SMITHSONIAN INSTITUTIONUNITED STATES NATIONAL MUSEUMBulletin 162
LIFE HISTORIES OFNORTH AMERICAN GALLINACEOUSBIRDS
ORDERS GALLIFORMES and COLUMBIFORMES
BYARTHUR CLEVELAND BENTTaunton, Massachusetts
UNITED STATESGOVERNMENT PRINTING OFFICEWASHINGTON : 1932
For sale by the Superintendent of Documents, Washington, D. C. Price ?1.00 (Paper cover)
ADVERTISEMENTThe scientific publications of the National Museum include twoseries, known, respectively, as Proceedings and Bulletin.The Proceedings series, begun in 1878, is intended primarily as amedium for the publication of original papers, based on the collec-tions of the National Museum, that set forth newly acquired factsin biology, anthropology, and geology, with descriptions of new formsand revisions of limited groups. Copies of each paper, in pamphletform, are distributed as published to libraries and scientific organi-zations and to specialists and others interested in the different sub-jects. The dates at which these separate papers are published arerecorded in the table of contents of each of the volumes.The series of Bulletins, the first of which was issued in 1875, con-tains separate publications comprising monographs of large zoologi-cal groups and other general systematic treatises (occasionally in sev-eral volumes), faunal works, reports of expeditions, catalogues oftype specimens and special collections, and other material of similarnature. The majority of the volumes are octavo in size, but a quartosize has been adopted in a few instances in which large plates wereregarded as indispensable. In the Bulletin series appear volumesunder the heading Contributions from, the United States NationalHerbarium, in octavo form, published by the National' Museum since1902, which contain papers relating to the botanical collections ofthe Museum.The present work forms No. 162 of the Bulletin series.Alexander Wetmore,Assistant Secretary, Smithsonian Institution.Washington, D. C, February 12, 1932.n
CONTENTS
PageIntroduction jxOrder GalliformesFamily PerdicidaePerdix perdix perdixEuropean partridgeHabitsDistribution 9Colinus virginianus virginianus 9Eastern bobwhite 9Habits 9Distribution 39Colinus virginianus floridanus 32Florida bobwhite 32Habits. 32Colinus virginianus texanus 34Texas bobwhite 34Habits. 34Colinus ridgwayi 3gMasked bobwhite 3gHabits 36Distribution 49Oreortyx picta palmeri 49Mountain quail 49Habits 49Distribution 43Oreortyx picta picta. 43Plumed quail 43Habits. 43Oreortyx picta confinis 51San Pedro quail 51Habits 51Callipepla squamata pallida 51Arizona scaled quail. 51Habits 51Distribution 57Callipepla squamata castanogastris 58Chestnut-bellied scaled quail 58Habits 58Lophortyx californica californica 59California quail 59Habits 59Distribution 60Lophortyx californica vallicola 62Valley quail 62Habits 62Lophortyx californica catalinensis 79Catalina quail 79Habits 79Lophortyx californica plumbea 71ni
IV BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMOrder Galliformes?Continued.Family Perdicidae?Continued. pageSan Quintin quail 71Habits 71Lophortyx californica achrustera 72San Lucas quail 72Habits 72Lophortyx gambeli gambeli 73Gambel's quail 73Habits 73Distribution 82Lophortyx gambeli sanus . 84Olathe quail 84Habits 84Cyrtonyx montezumae mearnsi 84Mearns's quail 84Habits 84Distribution 90Family Tetraonidae 91Dendragapus obscurus obscurus 91Dusky grouse 91Habits 91Distribution 95Dendragapus obscurus richardsoni 96Richardson's grouse - 96Habits 96Dendragapus obscurus flemingi 102Fleming's grouse 102Habits 102Dendragapus fuliginosus fuliginosus 103Sooty grouse 103Habits 103Distribution 113Dendragapus fuliginosus sierrae 114Sierra grouse 114Habits 114Dendragapus fuliginosus howardi 117Mount Pinos grouse 117Habits 117Dendragapus fuliginosus sitkensis 119Sitka grouse 119Habits 119Canachites canadensis canadensis 120Hudsonian spruce grouse 120Habits 120Distribution 128Canachites canadensis osgoodi 129Alaska spruce grouse 129Habits 129Canachites canadensis canace 131Canada spruce grouse 131Habits 131Canachites canadensis atratus 135Valdez spruce grouse 135Habits 135
CONTENTS VOrder Galliformes?Continued.Family Perdicidae?Continued. pageCanachites franklini 136Franklin's grouse 136Habits 136Distribution 141Bonasa umbellus umbellus 141Ruffed grouse _ 141Habits .- 141Distribution 165Bonasa umbellus togata 166Canada ruffed grouse 166Habits 166Bonasa umbellus umbelloides 171Gray ruffed grouse 171Habits 171Bonasa umbellus sabini 174Oregon ruffed grouse 174Habits 174Bonasa umbellus thayeri 177Nova Scotia ruffed grouse . 177Habits 177Bonasa umbellus yukonensis 177Yukon ruffed grouse 177Habits 177Lagopus lagopus albus 178Willow ptarmigan 178Habits 178Distribution 189Lagopus lagopus alleni 191Allen's ptarmigan 191Habits 191Lagopus lagopus alexandrae 194Alexander's ptarmigan 194Habits 194Lagopus lagopus ungavus 197Ungava ptarmigan 197Habits 197Lagopus lagopus alascensis 200Alaska ptarmigan _ 200Habits 200Lagopus rupestris rupestris 202Rock ptarmigan 202Habits 202Distribution 211Lagopus rupestris reinhardi ___. 213Reinhardt's ptarmigan 213Habits 213Lagopus rupestris nelsoni 215Nelson's ptarmigan 215Habits 215Lagopus rupestris atkhensis 218Turner's ptarmigan 218Habits 218Lagopus rupestris townsendi 220
VI BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMOrder Galliformes?Continued.Family Perdicidae?Continued. PageTownsend's ptarmigan 220Habits 220Lagopus rupestris chamberlaini 221Chamberlain's ptarmigan 221Habits 221Lagopus rupestris dixoni 223Dixon's ptarmigan 223Habits 223Lagopus rupestris sanfordi 225Sanford's ptarmigan 225Habits 225Lagopus rupestris kelloggae 227Kellogg's ptarmigan 227Habits 227Lagopus rupestris evermanni 230Evermann's ptarmigan 230Habits 230Lagopus leucurus leucurus 232Northern white-tailed ptarmigan 232Habits 232Distribution 233Lagopus leucurus peninsularis 234Kenai white-tailed ptarmigan 234Habits 234Lagopus leucurus altipetens 234Southern white-tailed ptarmigan 234Habits 234Lagopus leucurus rainierensis 240Rainier white-tailed ptarmigan 240Habits 240Tympanuchus cupido americanus 242Greater prairie chicken 242Habits 242Distribution 262Tympanuchus cupido attwateri 263Attwater's prairie chicken 263Habits 263Tympanuchus cupido cupido 264Heath hen 264Habits 264Tympanuchus pallidicinctus 280Lesser prairie chicken ._ 280Habits 280Distribution 284Pedioecetes phasianellus phasianellus 285Northern sharp-tailed grouse 285Habits 285Distribution 286Pedioecetes phasianellus columbianus 288Columbian sharp-tailed grouse 288Habits 288Pedioecetes phasianellus campestris 291
CONTENTS VIIOrder Galliformes?Continued.Family Perdicidae?Continued. PagePrairie sharp-tailed grouse 291Habits 291Centrocercus urophasianus 300Sage hen 300Habits 300Distribution 309Family Phasianidae 310Phasianus colchicus torquatus 310Ring-necked pheasant 310Habits 310Distribution 322Family Meleagrididae 323Meleagris gallopavo merriami 323Merriam's turkey 323Habits 223Distribution 325Meleagris gallopavo silvestris 326Eastern turkey 326Habits 326Meleagris gallopavo osceola 340Florida turkey 340Habits . 340Meleagris gallopavo intermedia 342Rio Grande turkey 342Habits 342Family Cracidae 345Ortalis vetula vetula 345Chachalaca 345Habits 345Distribution 351Order Columbiformes 353Family Columbidae 353Columba fasciata fasciata 353Band-tailed pigeon 353Habits 353Distribution 361Columba fasciata vioscae 363Viosca's pigeon 363Habits 363Columba flavirostris flavirostris 365Red-billed pigeon 365Habits 365Distribution 368Columba leucocephala 369White-crowned pigeon 369Habits 369Distribution 375Columba squamosa 376Scaled pigeon 376Habits 376Distribution 379Ectopistes migratorius 379
VIII BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMOrder Columbiformes?Continued.Family Columbidae?Continued. PagePassenger pigeon 379Habits 379Distribution 399Zenaidura macroura carolinensis 402Eastern mourning dove 402Habits 402Distribution 412Zenaidura macroura marginella 416Western mourning dove 416Habits 416Zenaida zenaida zenaida 417Zenaida dove 417Habits 417Distribution 421Leptotila fulviventris angelica 422White-fronted dove 422Habits 422Distribution 425Melopelia asiatica asiatica 425Eastern white-winged dove 425Habits 425Distribution . . 426Melopelia asiatica mearnsi 428Western white-winged dove 428Habits 428Columbigallina passerina passerina 435Eastern ground dove 435Habits 435Distribution 439Columbigallina passerina pallescens 440Mexican ground dove 440Habits 440Columbigallina passerina bahamensis 443Bahama gurond dove 443Habits 443Scardafella inca inca 444Inca dove 444Habits 444Distribution 449Oreopeleia chrysia 450Key West quail dove. _ 450Habits 450Distribution. . .. 452Oreopeleia montana. . 453Ruddy quail dove. - 453Habits 453Distribution 455Starnoenas cyanocephala 456Blue-headed quail dove. . 456Habits... 456Distribution.. ? 458Literature cited 459Index 479
INTRODUCTIONThis is the ninth in a series of bulletins of the United StatesNational Museum on the life histories of North American birds.Previous numbers have been issued as follows
:
107. Life Histories of North American Diving Birds, August1, 1919.113. Life Histories of North American Gulls and Terns, August27, 1921.121. Life Histories of North American Petrels and Pelicans andtheir Allies, October 19, 1922.126. Life Histories of North American Wild Fowl (part), May25, 1923.130. Life Histories of North American Wild Fowl (part), June27, 1925.135. Life Histories of North American Marsh Birds, March 11,1927.142. Life Histories of North American Shore Birds (pt. 1), De-cember 31, 1927.146. Life Histories of North American Shore Birds (pt. 2), March24, 1929.The same general plan has been followed, as explained in previousbulletins, and the same sources of information have been utilized.The nomenclature of the new Check List of the American Ornithol-ogists' Union has been followed, but it has seemed best to continue inthe same order of arrangement of families and species as given in theold (1910) check list.This is the first group in which any considerable number ofsubspecies have had to be treated. An attempt has been made togive as full a life history as possible of the best-known subspeciesand to avoid duplication by writing briefly of the others and giv-ing only the characters of the subspecies, its range, and any habitspeculiar to it. In many cases certain habits, probably commonto the species as a whole, have been recorded for only one subspecies
;
such habits are mentioned under the subspecies on which the obser-vations were made. The distribution gives the range of the speciesas a whole, with only rough outlines of the ranges of the subspecies,which can not be accurately defined in many cases.The egg dates are the condensed results of a mass of recordstaken from the data in a large number of the best egg collectionsin the country, as well as from contributed field notes and from afew published sources. They indicate the dates on which eggs have
X BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbeen actually found in various parts of the country, showing theearliest and latest dates and the limits between which half the datesfall, the height of the season.The plumages are described in only enough detail to enable thereader to trace the sequence of molts and plumages from birth tomaturity and to recognize the birds in the different stages and atthe different seasons. No attempt has been made fully to describeadult plumages; this has been already well done in the many man-uals. The names of colors, when in quotation marks, are takenfrom Ridgway's Color Standards and Nomenclature (1912) and theterms used to describe the shapes of eggs are taken from his Nomen-clature of Colors (1886 edition). The bold-faced type in the meas-urements of eggs indicates the four extremes of measurements.Many of those who contributed material for previous bulletinshave continued to cooperate. Receipt of material from more than320 contributors has been acknowledged previously. In additionto these, our thanks are due to the following new contributors:Clinton G. Abbott, W. C. Adams, M. C. Badger, J. H. Baker, PaulBartsch, Glenn Berner, E. J. Booth, O. M. Bryens, R. L. Coffin, L. V.Compton, J. V. Crone, P. D. Dalke, Ben East, F. L. Farley, J. A.Gillespie, F. J. Herman, R. B. Horsfall, F. N. Irving, Miss A. M.Keen, E. A. Kitchin, T. T. McCabe, Norman McClintock, E. A.Mcllhenny, G. W. Monson, G. W. Morse, J. J. Murray, L. T. S.Norris-Elye, W. W. Perrett, G. B. Pickwell, Gower Rabbitts, W. M.Rosen, W. B. Savary, E. W. Schmidt, W. E. Sherwood, F. H. Shoe-maker, W. A. Squires, J. W. Sugden, J. G. Suthard, L. M. Terrill,C. W. Tindall, I. R. Tomkins, Miss F. May Tuttle, J. H. Wales,N. A. Wood, and Miss M. W. Wythe.Through the courtesy of the Bureau of Biological Survey, the serv-ices of Frederick C. Lincoln were again obtained to compile thedistribution paragraphs. With the matchless reference files of theBiological Survey at his disposal, his many hours of careful andthorough work have produced results far more satisfactory thancould have been attained by the author, who claims no credit andassumes no responsibility for this part of the work.Dr. Charles W. Townsend and Dr. Winsor M. Tyler renderedvaluable assistance in reading and indexing, for this group, thegreater part of the literature on North American birds, which savedthe author many hours of tedious work. Doctor Townsend alsocontributed the entire life histories of four species and DoctorTyler also contributed one. Dr. Alfred O. Gross contributed twolife histories and wrote up the diseases of the ruffed grouse. Dr.Arthur A. Allen contributed the courtship of the ruffed grouse.E. A. Mcllhenny and the Rev. P. B. Peabody loaned the author
INTRODUCTION XI
valuable negatives and Maj. Allan Brooks loaned two beautifuldrawings. Thanks are due to the late Frank C. Willard for manyhours of careful work in collecting, arranging, and figuring a greatmass of data on egg dates and measurements; the author wouldbe glad to have some one volunteer to undertake this work infuture. The author is much indebted to Dr. Charles W. Richmond,of the United States National Museum, for many hours of carefuland sympathetic work in reading the proofs and correcting errorsin this and all previous volumes; his expert knowledge has beenof great value.The manuscript for this volume was completed in January, 1931.Contributions received since then will be acknowledged later. Onlyinformation of great importance could be added. The reader isreminded again that this is a cooperative work; if he fails to findin these volumes anything that he knows about the birds, he canblame himself for not having sent the information toThe Author.

LIFE HISTORIES OF NORTH AMERICAN GALLINA-CEOUS BIRDS (ORDERS GALLIFORMES ANDCOLUMBIFORMES)By Arthur Cleveland BentTaunton, MassachusettsOrder GALLIFORMESFamily PERDICIDAE, QuailsPERDIX PERDIX PERDIX (Linnaeus)EUROPEAN PARTRIDGEHABITSAttempts to introduce the European gray partridge into NorthAmerica have met with marked success in certain favorable locali-ties and with many dismal failures in other places less congenial toit. Dr. John C. Phillips (1928) has summarized the whole historyof these attempts. Of the earlier unsuccessful importations hesays
:
The earliest attempt at introduction, which so far as known was made byRichard Bache, son-in-law of Benjamin Franklin, who stocked his plantationon the Delaware River near what is now the town of Beverly, N. J., withHungarian partridges, dates back to the latter part of the eighteenth century.There were subsequent attempts in Virginia and New Jersey, most importantof which was Pierre Lorillard's effort in 1879 at Jobstown, N. J. Later at-tempts commenced in a small way in 1899, but the real fever of importationalong the Atlantic coast began about 1905 and has lasted up to the present,although the period 1907 and 1914 saw the height of the industry. In EasternStates importations of these hardy little birds have been put down all the wayfrom Portland, Me., and northern New York to South Carolina, Georgia, Flor-ida, and Mississippi. In Connecticut, Pennsylvania, and New Jersey, the workwas done on a large scale and, at first, with encouraging results. In a fewplaces the birds undoubtedly bred the first season, and in other places as inthe Connecticut Valley, they persisted for 8 or 10 years in considerable num-bers ; eventually they vanished, however, between 1915 and 1920.The results on the western plains and prairies have been quitesuccessful, of which he writes
:
The results in the far Western States and in western and central Canadamay be briefly summarized. The most remarkable success followed immedi-1
2 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
ately upon the first introductions into Alberta, near Calgary, in 1908-9. OnApril 20, November 16, and December 10, 1908, Calgary sportsmen liberatedabout 70 pairs over a small area mostly south and west of Calgary. Morecame on April 20, 21, and 22, 1909, and in all some 207 pairs seem to haveformed the basis for this wonderful result. The first birds were placed some15 miles south of Calgary, and after the first large plantings, 40 pairs inone place and 30 not far away (High River and west of that place), the restwere planted mostly in lots of 10 pairs. This stock came from Hungary.Some time later the Northern Alberta Game and Fish Protection League liber-ated a fresh importation of 230 birds in Alberta near Edmonton, but the stockfrom Calgary had in the meantime spread north to that city. The gain interritory from this nucleus has been little short of marvelous. The birdshave now spread at least 60 miles northwest of Edmonton (Pembina River)and breed there. There has been an open season on them in Alberta for years,and they are now by far the commonest of imported game birds in westernCanada. The spread from this initial plant has carried the Hungarian partridgeinto Saskatchewan and all over its western part as far north as township 60 andsouth to the international boundary. All this happened within only five yearsfrom the time the bird was first recorded in the Province.A. G. Lawrence writes to me
:
First liberated in Manitoba in April, 1924, when the Game Protective Leaguereleased at Warren, Manitoba, 40 pairs imported direct from Czechoslovakia.A second shipment was received in January, 1925, 17 pairs being later releasedat Neepawa, Manitoba, and 26 pairs at Warren. These birds are apparentlywell adapted to the prairies and seem to be establishing themselves in theareas in which they were liberated.The experience with this partridge in the State of Washingtonwell illustrates the fact that it will flourish, increase, and spreadin the type of open country that it prefers, but will barely hold itsown or will die out entirely in less favorable regions. D. J. Lef-fingwell says in his notes
:
We find the introduction of the partridge has been most successful in thedry nonforested areas with an elevation of 1,000 or more feet above sea leveland where the game enemies are rare. The lack of vermin and the largeopen fields in which the birds may feed are probably the most importantfactors. The birds should not be introduced outside of the Temperate orTransition Zone.In the comparatively humid regions of western "Washington at-tempts to introduce Hungarian partridges have not been very suc-cessful. S. F. Rathbun says of this section
:
Western Washington is a picturesque region of mountains, hills, valleys,and streams. Originally it was clothed with a dense and luxuriant forestmostly coniferous, but now a great change in this respect is apparent. Ashas been so often the case in the past in a new country, the development ofthe region began along the lines of least resistance?in this instance it beingwhere land and water met?and now to a large extent the tall forests havebeen replaced by broad cultivated areas that steadily encroach upon the stillundeveloped ones.
EUROPEAN PARTRIDGE 6On the other hand the birds have prospered and spread in theeastern part of the State, of which he writes
:
Eastern Washington, on the contrary, is a section quite devoid of forests ex-cept along many of its streams and some of the more rugged parts, and eventhen this growth lacks the luxuriance of that of the west side ; in fact, beingscanty by comparison. And many parts of eastern Washington are more orless elevated and open, wide-sweeping plateaus rolling in turn to the water-courses.In Oregon the story is much the same. William L. Finley writesto me:During the years 1913-14 we liberated 1,522 of these birds in various coun-ties throughout the State. In the Willamette Valley and places in southernOregon the climate is mild, and the country is varied with patches of timber,fields, and gardens, which from all reports is very similar to the Europeanhome of these birds. In the eastern part of Oregon where the partridges wereliberated the altitude is a little higher ; it is colder in wiuter ; the hills arecovered with broad grain fields with quite a lot of wild sagebrush country sur-rounding, also more or less trees and brush in the canyons. It came ratheras a surprise to find that the partridges did not increase and thrive in theWilliamette Valley and southern Oregon, but they multiplied quite rapidlyall through the northeastern part of Oregon, and especially in the southeasternpart of Washington, where quite a number of these birds were imported andreleased.According to Charles J. Spiker (1929) Hungarian partridges havebeen introduced successfully in northern Iowa, where they havespread into six counties, as well as three counties in southern Minne-sota. He says of this bird:There is no more charming bird on the Iowa landscape than the HungarianPartridge, nor one which better deserves protection at the hands of thosewho have brought it from its native haunts to become acclimated and ad-justed to new environments. While it is not highly colored, like the Ring-necked Pheasant, yet it is a beautiful bird and merits a great deal of en-thusiasm from an aesthetic point of view as well as the more mercenary pointof view of the sportsman. In size it is somewhat larger than the Bob-white,and has some of the characteristics of this species. Seen as it flies directlyaway from the observer, especially as it first takes off from the ground orspreads its tail in alighting, it presents its very distinguishing field mark. Thisis the rich russet of the tail feathers, visible only in flight, and concealed bythe upper coverts when at rest, but greatly resembling the sheen of that ofthe Red-tailed Hawk. If one be so fortunate as to behold the bird on a bankabout on a level with his eyes or slightly above him, as it has upon two orthree occasions occurred with me, he will note the black crescent just belowthe breast, practically in the middle of the belly, but so located that the birdmust be in just the exact position for this mark to show itself.Courtship.?The Rev. F. C. R. Jourdain has sent me the follow-ing quotation from F. Menteith Ogilvie
:
In March courtship proper will have begun. In the great majority of cases,the birds will have definitely selected their partners. Here and there, where
4 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthe males are in excess, constant fights will take place, often resulting in theelder male ousting the younger from the possession of the female, a mostundesirable occurrence when it happens, looked at from the breeding point ofview. The old males are not only more pugnacious and stronger birds, butthey are also either infertile or much less fertile than the young male andthe result of the union is likely to be a small laying, a still smaller hatching,and a large percentage of rotten eggs. Throughout March, while pairing isgoing on fighting is generally continuous and severe. Tbese fights are veryamusing to watch?the two males, bristling with fury, feathers raised andwattles showing, rush at each other striking and buffeting with their wings,generally jumping a few inches from the ground. The " round " may last 3 or4 minutes; the lady, close by picking up a seed here and there and preeningherself, is apparently unconscious of the furious rivalry she is exciting. Thefighters now separate a little distance and recommence feeding and peace seemsto be declared, till one or other approaches too near the female, when war isinstantly declared again. So the battle continues with intervals over a con-siderable period, possibly a week or more, until one of the two is finally van-quished and the happy pair are left to their honeymoon. I have often watchedfights of this kind, and I never could see that the Partridges inflicted any realdamage on each other ; their principal offensive weapon seemed to be theirwings. Their bills they rarely used, and their feet they didn't appear to useat all. The studied inattention of the female is most amusing to watch, andI conclude she exercises no choice in the matter at all, beyond promisingher hand to the better man.Nesting.?The nest of this partridge is a very simple affair, aslight depression in the ground, lined with a few dead leaves, drygrass, or straw. It is usually placed among bushes, or in long grass,fields of clover, or in standing grain. Mr. Jourdain says in his notes
:
It should be noted that during the time of laying (which may last for threeweeks) the eggs are carefully covered up by the hen bird with grass or deadleaves. When she comes to the nest to lay she scratches away the covering,deposits an egg, and then replaces it again. Until the clutch is complete theeggs are laid anyhow. When the hen is about to incubate she arranges themwith the greatest care and for a single day curiously enough, leaves the eggsuncovered and then begins to incubate. She is a good mother and sits veryclosely, especially after the first few days. The male bird takes no part inbrooding but remains close at hand for defense if necessary.Eggs.?Of the eggs Mr. Jourdain says thatnormally the clutch ranges from 8 or 9 to about 20. I have known cases ofas many as 21 and 22, which may have been the produce of one hen but thehigher numbers which occasionally are met with, 26 to 40 ( !), are undoubtedlydue to two hens laying together in one nest. In color they are uniformlyolive, sometimes darker, sometimes lighter, but occasionally clutches havebeen found with almost white eggs, while a bluish type has also been recorded.One hundred British eggs measured by myself averaged 36.8 by 27.4 millimeters.The eggs showing the greatest extremes measured 38.9 by 28.4 and 37.7 by29.4, 33.8 by 26.3, and 37.5 by 25.7 millimeters.Young.?Mr. Jourdain writes:Incubation lasts not less than 24 full days, as a rule, though Hanroth gives23% as the period in Germany. In England most birds hatch out on the 25th
EUROPEAN PARTRIDGE 5day. When the young are hatched both parents take charge and are mostactive and courageous in defense of the young. On one occasion I heard apair on the far side of a hedge, and looking over the top I was surprised tofind that the bold little cock flew straight at my head with loud outcry whilethe hen busied herself in getting the young under cover as soon as possible.Several of the early British writers have referred to an incident,related by Yarrell (1871) as follows:A person engaged in a field, not far from my residence, had his attentionarrested by some objects on the ground, which, upon approaching, he foundto be two Partridges, a male and female, engaged in battle with a CarrionCrow ; so successful and so absorbed were they in the issue of the contest, thatthey actually held the Crow till it was seized and taken from them bythe spectator of the scene. Upon search, young birds, very lately hatched,were found concealed amongst the grass. It would appear, therefore, that theCrow, a mortal enemy to all kinds of young game, in attempting to carry offone of these, had been attacked by the parent birds, and with this singularresult.Plumages.?In Witherby's handbook (1920) the downy young isdescribed as follows
:
Crown chestnut with a few small black spots sometimes extending to lines;back of neck with a wide black line down centre, at sides pale buff markedblack ; rest of upper -parts pale buff with some rufous and black blotches or ill-defined lines, at base of wings a spot, and on rump a patch, of chestnut ; fore-head and sides of head pale yellow-buff (sometimes tinged rufous) with spots,small blotches, and lines of black; chin and throat uniform pale yellow-buff;rest of under-parts slightly yellower, bases of down sooty.And the juvenal plumage, in which the sexes are alike, is thusdescribed
:
Crown black-brown finely streaked buff, each feather having buff shaft-streak ; back of neck, mantle, back, rump and upper tail-coverts buff-brown,with whitish to pale buff shaft-streaks inconspicuously margined blackish
;
lores and sides of head dark brown streaked whitish ; chin, throat and centre ofbelly whitish to pale buff; breast, sides and flanks and under tail-covertsbrown-buff slightly paler than mantle and with whiter shaft-streaks, faintlymargined brown on flanks ; tail much like adult but feathers tipped buff andwith subterminal dusky bar and spots and central ones speckled and barreddusky ; primaries brown with pale buff tips and widely spaced bars on outerwebs; secondaries with pale buff bars extending across both webs and ver-miculated brown, shafts pale buff; scapulars, inner secondaries and wing-coverts brown-buff with wide brown-black bars and mottlings and pale shaft-streaks widening to white spots at tips of feathers.A postjuvenal molt, which is complete except for the outer twoprimaries, produces a first winter plumage. The sexes are nowdifferentiated and resemble the two adults, except for the morepointed tips of the outer primaries. This molt begins when theyoung bird is about half grown and is sometimes prolonged throughDecember.74564?32 2
6 BULLETIN 162, UNITED STATES NATIONAL MUSEUMAdults have a partial prenuptial molt in May and June, some-times in April, and a complete molt from July to November orDecember. Several observers have experienced some difficulty indistinguishing the sexes among adults, chiefly because many femaleshave the dark chestnut patch on the belly more or less well de-veloped. There seems to be some difference of opinion as to whetherthis character is more pronounced in old or in young females. Butthe sexes can always be distinguished by two characters; the lightchestnut on the sides of the head is lighter and more restricted in thefemale; the median wing coverts of the female are dark brown orblack, with widely spaced, pale-buff bars; whereas these coverts inthe male have no transverse bars, but only a pale-buff shaft streak.Food.?In Witherby's handbook (1920) the food of the partridgeis summarized by Mr. Jourdain, as follows
:
Chiefly shoots and leaves of grass and clover as well as seeds of many speciesincluding Polygonum, Trifolium, Alchemilla, Galium, Spergula, Persicaria, Poa,etc. Turnip leaves, young shoots of heather, bramble and blaeberry, hawthornberries, and corn also eaten. In spring and summer insects are also taken,including diptera (Tipulidae and larvae), coleoptera and hymenoptera (nntsand their pupae being very favourite food). Also aphides. Once recorded aseating pears on tree
!
Crops and stomachs of American birds also contained wheat,barley, and oats, mainly waste grain, seeds of wild buckwheat, pig-weed, and other weeds, and grasshoppers. It is said that thesebirds do not pull up sprouting corn as the pheasants do. Theirfood habits seem to be wholly beneficial.Behavior.?Macgillivray (1837) says, of the gray partridge, tnat itis fond of rambling into waste or pasture grounds, which are covered withlong grass, furze, or broom ; but it does not often enter woods, and neverperches on trees. It runs with surprising speed, when alarmed or in pursuitof its companions, although in general, it squats under the apprehension ofdanger, or when nearly approached takes flight. Its mode of flying issimilar to that of the Brown Ptarmigan ; it rises obliquely to some height, andthen flies off in a direct course, rapidly flapping its wings, which producea whirring sound.Yarrell (1871) writes:During the day a covey of Partridges, keeping together, are seldom seenon the wing unless disturbed ; they frequent grass-fields, preferring thehedge-sides, some of them picking up insects, and occasionally the greenleaves of plants ; others dusting themselves in any dry spot where the soilis loose, and this would seem to be a constant practice with them in dryweather, if we may judge by the numerous dusting places, with the marks andfeathers to be found about their haunts; and sportsmen find, in the earlypart of the shooting-season, that young and weak birds are frequently infestedwith numerous parasites. In the afternoon the covey repair to some neigh-Douring field of standing corn, or, if that be cut, to the stubble, for the seconddaily meal of grain; and, this completed, the call-note may be heard, accord-
EUROPEAN PARTRIDGE 7ing to White, as soon as the heetles begin to buzz, and the whole move awaytogether to some spot where they jug, as it is called?that is, squat andnestle close together for the night ; and from the appearance of the mutings,or droppings, which are generally deposited in a circle of only a few inchesin diameter, it would appear that the birds arrange themselves also in acircle, of which their tails form the centre, all the heads being outwards
?
a disposition whicK instinct has suggested as the best for observing theapproach of any of their numerous enemies, whatever may be the direction,and thus increase their security by enabling them to avoid a surprise. Inthe morning early they again visit the stubble for a breakfast, and pass therest of the days as before. Fields of clover or turnips are very favourite placesof resort during the day. Mr. Harvie-Brown informs the editor that whenthe snow lay upon the ground he has known a covey to roost regularly on alimb of a large tree ; and he has also seen Partridges " treed " by a dog.Considerable discussion has appeared in print on the effect, on ournative game birds, of introducing Hungarian partridges. Someclaim that where the partridges are increasing the native grouse aredisappearing. Most of our grouse are subject to periodic fluctuationin numbers from other causes; and it does not seem to have beendefinitety proved that the partridges are the cause of any local de-crease in grouse. There are certainly plenty of suitable nesting sitesfor all these ground-nesting species ; there is no proof that any short-age of food supply has led to any disastrous competition betweenthem ; and there is no evidence that the smaller partridges ever attackthe larger grouse, which should be more than a match for them.Though there is always danger in introducing a foreign species, itwould seem that the little gray partridge is more likely to prove acomplementary than a competitive species.Enemies.?Partridges, like all other ground-nesting species, arepreyed upon by the whole long list of furred and feathered enemies,but they are such prolific breeders that their natural enemies are notlikely seriously to reduce their numbers. Their habits of feedingin the open during the day and roosting in the open at night, makethem especially exposed to the attacks of hawks and owls. The ring-necked pheasant may have to be reckoned with, as an enemy of thepartridge. Mr. Spiker (1929) writes:Northwestern Iowa has not until fairly recently been afflicted with this per-nicious bird, but they are on the increase, and farmers have told me that withthe coming in of the Ring-necked Pheasant, the partridges are departing. Per-haps a concrete example would be admissible here. Mr. Raymond Rove, afarmer living a few miles northwest of Sibley, while plowing late last fall(1927), observed something of a commotion in a little swale a short distancefrom his plowing. Prompted by curiosity he walked over to the place andflushed half a dozen partridges and three Ring-necked Pheasants. On theground before him lay the bleeding bodies of three partridges newly killed. Itwas just dusk, and doubtless the smaller species had crept into the long grassto spend the night and had been fallen upon by the pheasants who were alreadythere. Stories are also told of the destruction of the nests of the HungarianPartridge by pheasants.
8 BULLETIN 162, UNITED STATES NATIONAL MUSEUMVoice.?Mr. Spiker (1929) says that the voice isnot unmusical, and yet not conspicuous unless listened for, it is especiallynoticeable on a still spring evening, when there is little or no breeze, and theshadows of dusk follow the disappearance of the sun. There is a single two-syllabled chuckling note which may be represented somewhat by the syllables
" kee-uck," the second syllable being rather raspy or throaty as compared tothe first, which is high pitched and nasal. Upon being flushed, the bird takesoff with the startling whirr of wings characteristic of this family, utteringthe while a rapid cackling which diminishes to the above given notes repeatedseveral times and with a gradually increasing interval between them. Inthe immediate vicinity of Ashton it is not unusual to hear from four to eightof these birds calling at the same time and from as many different directions.Walter H. Rich (1909) writes of some birds in captivity:In their coop they used a great variety of language ; they clucked like aGrouse ; they chattered like a Blackbird ; they snapped their bills like an Owl
;
they " jawed " like a Parrot ; they made a guttural note of alarm like the
" br-r-r-r " of a startled Pigeon ; they hissed like a Black Duck guarding hernest, or like a Thomas cat whose dignity is ruffled not quite enough for anger
;
and, in addition, they are said to " crow " at evening.Game.?Provided that the Hungarian partridge does not seriouslyinterfere with the welfare of our native species, it seems to be a wiseand valuable addition to our list of game birds. I have never huntedit, but those who have speak very highly of it. It is a strong, swiftflier, smart and sagacious, well fitted to test the skill of the bestsportsmen. It is a fine bird for the table. Unfortunately it willsurvive and flourish only in certain favorable sections, mainly thenorthwestern grainfields and grassy plains. There it can probablysurvive much more intensive hunting than either the prairie chickenor the sharp-tailed grouse.One of the men who helped to introduce the partridge in Wash-ington wrote to Mr. Rathbun as follows
:
From the standpoint of a game bird I believe them to be the gamest ofthem all. The law of the covey is very strong, and when they flush all of themgo at the same time. There seems to be less than a fraction of a second be-tween the time the first one and the last one makes his get-away. They willalways be able to take care of themselves, since they become very wild whenmuch shooting is done. When one is winged or slightly wounded so that hecan not fly he will run a mile sometimes before a hunter's dog overtakes him.During the winter months they come right into the towns and eat at the backdoors of the residences. They will help themselves to strawstacks, haystacks,and anything edible. At night they burrow in the snow, sometimes makinglittle tunnels 4 or 5 feet long under 2 feet of snow. I have hunted upland gamebirds in the West covering a period of 37 years, but I believe the Hungarianpartridge, considered from every standpoint as a game bird, is the premierone of the Pacific coast.Winter.?Mr. Spiker (1929) says:The species is gregarious during the winter, beginning to flock in Octoberand continuing till the last of February. During this season they frequent the
EASTEUN BOBWHITE y
stalk fields left after the picking of the corn. When the gregarious spiritis upon them they are exceedingly wary and are up and away almost as theysee the hunter enter the field. The startling noise with which they take flightand their extremely rapid coursing across the field make them a very difficulttarget, and, although many attempts are made by poachers, few birds fall asvictims. By the latter part of February, however, there comes a change whenthey begin breaking up and pairing off, and at this time they appear to losesome of their wariness. DISTRIBUTIONIntroduced more or less unsuccessfully in the Eastern States, fromMaine and New York southward to Florida and Mississippi, also inCalifornia. In the Central States, from Minnesota and Michigan toKansas and Arkansas, most attempts at introduction have failed,except in extreme northwestern Kansas, in parts of Iowa (Osceolaand Lyon Counties), and in southeastern Wisconsin (WaukeshaCounty). Introduced birds have done well in southern BritishColumbia (Fraser Valley) and in eastern Washington and Oregon(east of the Cascade Mountains). The most remarkable success hasbeen attained in Alberta, Saskatchewan, and Manitoba, where thebirds have flourished and spread over a wide territory.Egg dates.?Washington : 4 records, May 25 to June 10.COLINUS VIRGINIANUS VIRGINIANUS (Linnaeus)EASTERN BOBWHITEHABITSIn the springtime and early in summer bobwhite deserves his name,which he loudly proclaims in no uncertain terms and in a decidedlycheering tone from some favorite perch on a fence post or the lowbranch of some small tree. But at other seasons I prefer to call hima quail, the name most familiar to northern sportsmen, or a partridge,as he is even more appropriately called in the South. But Europeansportsmen would say that neither of these names is strictly accurate,so we may as well call him bobwhite, which is at least distinctive.By whatever name we call him, he is one of our most popular andbest beloved birds. From a wide distribution in the East, he hasfollowed the plow westward with the clearing of the forests and thecultivation of the fertile lands of the Middle West ; and more recentlyhe has been successfully introduced into many far-western States.Bobwhite is one of the farmer's best friends ; his economic status iswholly beneficial; he is not known to be injurious to any of our crops,as what grain he eats is mostly waste grain, picked up in the stubblefields after the crops are harvested. It seems to me, however, thattoo much stress has been laid on his services as a destroyer of weedseeds. Nature has provided so lavishly in the distribution of weed
10 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
seeds that only a very small fraction of them can find room to germi-nate, and the seeds picked up by birds, which never glean thoroughly,only leave room for others to grow. I doubt if even a square foot ofground has ever been kept clear of weeds by birds. The hoe and thecultivator will always have to be used. But bobwhite has a finescore to his credit as a destroyer of grasshoppers, locusts, potatobeetles, plant lice, and other injurious insects.It has been suggested by some bird protectionists that the bobwhiteshould be removed from the game-bird list and be rigidly protectedat all seasons as a song bird and an insectivorous bird. But we mustnot lose sight of its economic value as a game bird and the pleasureand healthful exercise that it gives to thousands of sportsmen. Thereare hundreds of other birds that bring joy to the hearts of amateurbird admirers and many others that are nearly or quite as useful asinsect destroyers, so why should we deprive the sportsmen of theirmost popular upland game bird, when they have not more than twoor three species at best in any one section of the country? EdwardH. Forbush (1927) has summed up the matter very well, as follows:As a popular game bird of the open country Bob-white has no rival. Probablyabout 500,000 sportsmen now go out annually from cities east of the RockyMountains to hunt this bird. This necessitates a great annual expenditure forhunters' clothing, guns, ammunition, dogs, and guides. It adds to the revenueof farmers and country hostelries. In some of the southern states Bob-whitepays the taxes on many farms where the farmers sell their shooting rightsto sportsmen. Perhaps there is no bird to which the American people are moredeeply indebted for both aesthetic and material benefits. He is the mostdemocratic and ubiquitous of all our game birds. He is not a bird of desert,wilderness, or mountain peak which one must go far to find. He seeks thehome, farm, garden, and field ; he is the friend and companion of mankind
;
a much needed helper on the farm; a destroyer of insect pests and weeds;a swift flying game bird, lying well to a dog; and, last as well as least, goodfood, a savory morsel, nutritious and digestible.One does not have to go far afield to find the haunts of bobwhites,for they shun the deep forest areas, seldom resort to the woods exceptto escape from danger, and are rarely found on the wide openprairies. They seem to love the society of human beings and theircultivated fields. During spring and summer they are particularlydomestic and sociable, when it is no uncommon occurrence to heartheir loud, ringing calls almost under our windows, to see one perchedon a fence post near the house or on the low branch of an apple treein the orchard, or to find them running along the driveway or agarden path. They are very tame and confiding at that season andseem to know that they are safe. At other seasons they resort to moreopen country and seek more seclusion. In New England they preferthe vicinity of farms, where they find suitable feeding grounds in oldweed patches and stubble fields where crops of buckwheat, millet,
EASTERN BOBWHITE 11
rye, wheat, oats, or other grains have been harvested. But near athand they must have suitable cover, thick, swampy tangles or brierpatches in which to roost at night, or dense thickets or woodlots inwhich to seek refuge when pursued. In the South, according toM. P. Skinner, they " like weedy corners of cornfields nest to atangle of blackberry briars, cane, cat briars, and brush, into whichthey can retreat at a moment's notice. They also like cotton fields,especially if a corner be grown up to broom sedge and low brush."The cultivated fields of the South are usually well overgrown withweeds in the fall, where the partridges find both food and shelter inthe old fields of cowpeas, ground nuts, and other crops, overgrownwith crabgrass, foxtail grass, Japan clover, plume, and wild grasses.Courtship.?It is not until spring is well advanced that the coveys,which have kept together all winter, begin to break up and scatter.Then it is that the young cock, which has now acquired full maturityand vigor, begins to feel the urge of love and, separating from hiscompanions, sets about the important business of securing a mate.Dressed in his best springtime attire, his bosom swelling with pride,he selects his perch, a fence post, the low branch of a tree, or someconvenient stump, from which to send out his love call to his expectedbride. Bob-white/ Ah, bob-white bob-bob-white/ It rings out, loudand clear, repeated at frequent intervals, while he listens for a re-sponse, perhaps for half an hour or more in vain. At length he mayhear the coveted sound, the sweet, soft call of the demure little hen.With crest erected and eyes aglow, he flies to meet her and displayhis charms, fluttering and strutting about her and coaxing her withall the pomp and pride of a turkey gobbler. But she is shy and coy,and does not yield at first. Perhaps she runs away, and then ensuesa lively game of chase. Aretas A. Saunders tells in his notes of sucha chase that he saw under favorable circumstances. The hen keptabout 5 feet ahead of the cock, running rapidly, faster than he hadseen this species move at any other time, back and forth, in and out,around some clumps of grass. Though he watched for 15 minutes,the cock did not seem to gain an inch. Doubtless he did eventually.But bobwhite's road to happiness is not always so smooth. Ashis clarion call of defiance rings out across the fields an answeringcall, bob-bob-v:hite, reaches his jealous ears, the voice of an unknownrival. Back and forth the challenges are exchanged, as the bravelittle warrior advances to meet his foe. Louder, sharper, and angrierare their cries, as they dodge about, bursting with rage and eagerfor the fray, seeking a vantage point for the attack. At last theyclinch in furious combat, like small game cocks, savagely biting andtearing with sharp little beaks, scratching with claws, and buffetingwith strong little wings. The fighting is fast and furious for a timeuntil one gives up exhausted and slinks away. Finally the brave
12 BULLETIN 162, UNITED STATES NATIONAL MUSEUMiittle conqueror enjoys the spoils of victory, the acceptance of hissuit by the modest little hen, who now knows that she has picked awinner.Herbert L. Stoddard (1931), in his excellent and exhaustive mono-graph on the bobwhite, published by the committee on the Coopera-tive Quail Investigation, has added to our knowledge a vast fund ofinformation on the habits of this valuable species, its enemies, dis-eases, and means for preserving and increasing it, based on a fiveyears' study in cooperation with the Biological Survey. Anyoneinterested in this subject should study this voluminous report, asour space will permit only brief extracts from it.As to the breaking up of the coveys in the spring, which "areusually composed of the remnants of several hatchings," he saysthat " many of the birds are not closely, if at all, related." At thisseason, "cocks, which had been peaceable companions previously,became pugnacious," and frequent fights occurred. In the inclosuresthe fights were harmless, as a rule, but in the wild " an occasionalcombat no doubt proves fatal, for two dead cock quail that had beenpicked up afield were brought to us with the flesh bitten to the boneat the junction of head and neck."Referring to the courtship display, he writes
:
This display is a frontal one. The head is lowered and frequently turnedsideways to show the snowy-white head markings to the best advantage, thewings are extended until the primary tips touch the ground, while the elbowsare elevated over the back and thrown forward, forming a vertical featheredwall. The bird, otherwise puffed out to the utmost in addition to the spread,forward-thrust wings and lowered, side-turned head, now walks or advancesin short rushes toward the hen, and follows her at good speed in full displayin case she turns and runs.Some evidence was obtained to indicate that some mated pairsremain mated during winter and for at least two breeding seasons.As to the devotion of mated pairs, he says
:
Two weeks to a month may elapse, depending on the weather, between thetime of pairing and the beginning of nesting. During this period the pairsappear inseparable, the hen usually taking the lead in foraging expeditions,with the cock a devoted follower. He is very attentive at this time, as indeedhe is all during the breeding season, unless he takes up incubation duties,when he appears to lose interest in the opposite sex. It is amusing to seehim catch a grasshopper or other large insect after a lively chase. He puffshimself up and, holding the insect out in a stiff, wooden manner, starts asoft, rapidly repeated cu-cu-cu-cu to attract his mate, who rushes to him andeats the dismembered insect. This common habit may be frequently observedall during the breeding season, the hen usually being the one to get the insectscaught by the cock, even when the pair are rearing a brood.Nesting.?The bobwhite's nest is a very simple affair, but artfullyconcealed and seldom found, except by accident, as the bird is a very
EASTERN BOBWHITE 13
close sitter and usually does not leave the nest until almost troddenupon. The favorite nesting sites seem to be along old fence rows,where the grass grows long and thick or is mixed with tangles ofvines or briers, in neglected brushy corners of old fields, under dis-carded piles of brush, or in the tangled underbrush that, mixed withgrass, grows on the edges of woods, thickets, or swamps. The nestis often placed in open fields of tall grass, where the hay cutter some-times destroys it, in cultivated fields of grain or alfalfa, or at thebase of a tree in the farmer's orchard, if the grass is long enoughto conceal it. A nest is often found in an unexpected place. Once,at my cottage on Cape Cod, I worked for two days weeding mygarden within 3 feet of a boundary fence and was surprised the nextday, on cutting the grass along the fence, to uncover a quail's nest,with 15 eggs, from which the bird had never stirred. I was toldone day that there was a quail's nest under a brush pile at our golfclub and went up to photograph it. I found a pile of pine boughsthat had been cast aside just off the edge of an elevated putting green.I walked around it carefully several times trying to see the bird,but I never found it until I lifted the right bough and flushed her.I saw her several times afterwards and believe she raised her broodsuccessfully.George Finlay Simmons (1915) tells of a nest found by him inTexas " under the edge of a bale of hay in an old shed on the prairie,"which he discovered by flushing the bird. Charles R. Stockard(1905) writes from Mississippi:In fields of sedge grass or oats many pairs will often nest very close together.June, 1895, I found in a thirty acre field of sedge grass sixteen nests of the Bob-white, all containing large sets, ranging from twelve to twenty-two eggs, and thetotal number of eggs in this field must have been about three hundred.Out of 602 nests, studied by Stoddard (1931) and his associates,97 were in woodland, 336 in broom-sedge fields, 88 in fallow fields,and " about 4 per cent in cultivated fields, mostly in the grassy growtharound stumps in corn or cotton fields, but occasionally under trashcast aside by plows or cultivators." In the few cases where nestconstruction was under observation the work was done entirely bythe male under the supervision of his mate.The construction of a typical nest is very simple. Having se-lected a suitable spot, where the vegetation is thick enough to af-ford effective concealment, a hollow is scooped out and lined withdead grass or other convenient material; after that the dead andgrowing grass or other vegetation is woven into an arch over thenest, often completely concealing it, and leaving only a small open-ing on the side, just large enough for the bird to enter or leave thenest; while incubating, the bird looks out through this opening; ifthere are any vines or briers growing about the nest, these are also
14 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMwoven into the arch to make it firmer and more impenetrable.F. W. Rapp describes in his notes a more elaborate nest, resemblinga marsh wren's nest in construction and shape and very firmlybuilt; it was located in a fence row and was made of oak leaves andJune grass, neatly woven together into a ball, flattened on the bot-tom, with a hole on one side. Often the simplest nest is made byentering a thick clump of grass and flattening down a hollow in thecenter, without disturbing the grass tops at all.Major Bendire (1892) quotes Judge John N. Clark as havingseen a male bobwhite building a nest, as follows
:
In May, 1887, while on a hill hack of my house one morning, I heard a Quailwhistle, but the note, which was continually repeated, had a smothered sound.Tracking the notes to their source, I found a male Bob White building a nestin a little patch of dewberry vines. He was busy carrying in the grasses andweaving a roof, as well as whistling at his work. The dome was very expertlyfashioned, and fitted into its place without changing the surroundings, so thatI believe I would never have observed it. had he kept quiet.He also speaks of a nest, found in Louisiana, which "was entirelyconstructed of pine needles, arched over, and the entrance probablya foot or more from the nest proper."Eggs.?The bobwhite ordinarily lays from 12 to 20 eggs, 14 to 16being perhaps the commonest numbers ; as few as 7 or 8 and as manyas 30, 32, and even 37 eggs have been found in a nest; but theselarge numbers are probably the product of more than one femaleand are deposited in layers. The eggs are mainly subpyriform inshape, sometimes quite pointed or again more rounded. The shellis smooth, with very little gloss, and decidedly hard and tough.The color is dull white or creamy white, rarely "light buff" or
"pale ochraceous-buff." They are never spotted, but are usuallymore or less nest stained. The measurements of 55 eggs in theUnited States National Museum average 30 by 24 millimeters; theeggs showing the four extremes measure 32.5 by 24, 31 by 26, and 26by 22.5 millimeters.Bobwhites occasionally lay their eggs in other birds' nests. H. J.Giddings (1897) reports the finding of a quail's egg in a towhee'snest ; and the editor in a footnote refers to one laying in a domesticturkey's nest. E. B. Payne (1897) adds that he "found in a mea-dowlark's nest five of the meadowlark's eggs and four of the quail's."Mr. Rapp mentions in his notes a quail's nest shown to him thatcontained 12 eggs of the quail and 2 of the domestic hen. HerbertL. Stoddard has a photograph of a bantam's egg in a quail's nest.Young.?It is generally supposed that at least two broods ofyoung are raised in a season, perhaps three in the southern part ofthe quail's range, as very early and very late broods are of common
EASTERN BOBWHITE 15
occurrence. But, as the quail has many enemies and many nestsare broken up or deserted, it may be that the late broods are merelybelated attempts to raise a family ; in which case, perhaps one broodin the North and two in the South is more nearly the average.Most authorities agree that the period of incubation is about 23or 24 days. Both sexes share this duty. In the study of 276 nestsby Mr. Stoddard (1931), in southern Georgia and northern Florida,so far as could be ascertained 73 were entirely in charge of the cockand 175 in charge of the hen. If any fatal accident befall the hen,as too often happens, then the cock assumes full charge of the eggsand afterwards takes care of the young. It is said, too, that after theyoung are two or three weeks old the mother hands the brood overto the care of their father and starts to lay a second set of eggs ; butI doubt if this has been definitely proved.Young quail leave the nest almost as soon as they are hatched,and the eggshells are generally left in the nest, although occasionallya chick is seen rumiing away with part of the shell on its back. Theyare carefully tended by their devoted parents, who use every knownartifice to distract an enemy. Dr. T. M. Brewer (Baird, Brewer, andRidgway, 1905) relates the following to illustrate an extreme caseof parental boldness
:
Once as I was rapidly descending a path on the side of a hill, among a lowgrowth of scrub-oak I came suddenly upon a covey of young Quail, feeding onblueberries, and directly in the path. They did not see me until I was closeupon them, when the old bird, a fine old male, flew directly towards me andtumbled at my feet as if in a dying condition, giving at the same time a shrillwhistle, expressive of intense alarm. I stooped and put my hand upon hisextended wings, and could easily have caught him. The young birds, at thecry of the parent, flew in all directions; and their devoted father soon fol-lowed them, and began calling to them in a low cluck, like the cry of the BrownThresher. The young at this time were hardly more than a week old, andseemed to fly perfectly well to a short distance.Their ability to fly at such an early age is due to the fact that theirwings begin to sprout almost as soon as they are hatched; I haveseen young chicks not more than 2 inches long with wings reachingto their tails; they are very active and vigorous and grow veryrapidly. They are experts at hiding; a warning note from thewatchful parent, who previously has kept the brood together byfrequent gentle twitterings, sends them to cover instantly; instinc-tively they dart under some fallen leaf, beneath a tuft of grass, intosome thick vegetation or little hollow, where they remain motionlessuntil told by their parent that danger has passed. Edwyn Sandys(1904) has described this so well that I quote the following:If those who may stumble upon a brood of quail will take a sportsman-naturalist's advice, they will promptly back away for a few yards, sit down,and remain silently watchful. No search should be attempted, for the searcher
16 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
is more likely to trample the life out of the youngsters than to catch one.But if he hide in patience, he may see the old hen return, mark her cautiouslystealing to the spot, and hear her low musical twitter which tells that the perilhas passed. Then from the scant tuft here, from the drooping leaf yonder,apparently from the bare ground over which his eyes have roved a dozen times,will arise active balls of pretty down until the spot appears to swarm withthem. And the devoted mother will whisper soft greetings to each, and in somemysterious manner will make the correct count, and then with nervous careshepherd them forward to where there is safer cover. And they will troopafter her in perfect confidence, to resume their bug-hunting and botanicalresearches as though nothing important had transpired.Young quail are busy foragers, and they grow rapidly. Within a few daysafter leaving the nest they are capable of a flight of several yards. A broodflushed by a dog will buzz up like so many overgrown grasshoppers, fly a shortdistance, then dive into cover in a comical imitation of the tactics of theirseniors. As insect catchers they are unrivalled, their keen eyes and tirelesslittle legs being a most efficient equipment even for a sustained chase. Theparents scratch for them and call them to some dainty after the manner ofbantam fowls, and the shrewd chicks speedily grasp the idea and set to work forthemselves. A tiny quail scratching in a dusty spot is a most amusing sight.The wee legs twinkle through the various movements at a rate which theeye can scarcely follow, and the sturdy feet kick the dust for inches around.When a prey is uncovered it is pounced upon with amazing speed and accuracy,while a flying insect may call forth an electric leap and a clean catch a footor more above the ground. As the season advances grain, seeds of variousweeds, berries, wild grapes, and mast are added to the menu, in which insectsstill remain prominent. After the wheat has been cut the broad stubbles becomefavorite resorts, especially when they are crowded with ragweed. Patchesof standing corn now furnish attractive shelter and the suitable dusting placesso necessary to gallinaceous birds.Plumages.?Only in the smallest chicks can the pure natal downbe seen. In a typical chick the forehead and sides of the head arefrom " ochraeeous-tawny " to " ochraceous-buff," with a stripe ofbrownish black from the eye to the nape; a broad band from thehind neck to the crown, terminating in a point above the forehead,is "chestnut," deepening to "bay" on the edges; there is a similarbroad band of the same colors from the upper back to the rump ; therest of the upper parts is mottled with " chestnut," dusky, and buff
;
the chin and lower parts are pale buff or buffy white. In somespecimens from the South the back and rump are almost wholly
" chestnut," mixed with some black.The juvenal plumage begins to appear on the wings and scapularsat a very early age, even before the chick has increased perceptiblyin size ; I have seen chicks 2 or 3 inches long that had wings extend-ing beyond the tail and that would soon be able to fly. In this plum-age the sexes are alike, except that, according to Dr. JonathanDwight (1900), "the males are apt to be richer colored than arefemales, with grayer tails, whiter chins, blacker throat bands, andoften a slight dusky barring on the breast." The first feathers to
EASTERN BOBWHITE 17appear on the back and scapulars are black on the inner web, broadlytipped with white, and mottled with brown (" russet " to " tawny")and dusky on the outer web, with white shaft stripes, broadeningat the tip ; as these feathers grow out longer the black appears onlyas a large subterminal spot. In full juvenal plumage the crown iscentrally dusky, laterally gray (" hair brown " to " drab "), mottledor variegated with black; the throat is white in the male and buffywhite in the female ; the breast and flanks are " drab " to " lightdrab," with whitish shaft streaks; the belly is paler or white; thetail is gray, mottled with white ; and the primaries are mottled withpale buff on the edges.Even before the juvenal plumage is fully acquired the postjuvenalmolt into the first winter plumage begins. This molt is completeexcept for the outer pair of primaries on each wing, which areretained all through the first year; and it takes place at any timefrom late in summer until November, depending on the time at whichthe young were hatched. The first winter plumage is scarcely dis-tinguishable from that of the adult, and the sexes are widely differ-entiated ; but the colors above are duller with paler edgings, and theunderparts are more buffy and somewhat less barred. Young birdscan be distinguished from adults all through the first winter andspring by the outer pair of primaries, the first and second, on eachwing, which are still juvenal (pointed).The first prenuptial molt, as well as all subsequent prenuptialmolts, amounts to the renewal of only a few feathers about thehead and throat. The first postnuptial molt, the following summerand fall, chiefly in September, is complete and produces the adultwinter plumage. Adults then continue to have similar molts eachyear, a very limited head molt in spring and a complete postnuptialmolt from August to October. The slight seasonal difference betweenspring and fall plumages is mainly due to wear and fading.Among the thousands of quail shot and the large series preservedin collections, some odd types of plumage are to be found, such asmales with black or buff throats, very dark or melanistic types, othersin which the browns are replaced with buff or the buffs with white,producing a pallid type; partial albinos are occasionally seen andvery rarely one that is wholly pure white. Erythrism is reportedand illustrated by Stoddard (1931).Food.?Quail are very regular in their feeding habits. Everysportsman knows this and takes advantage of it, for he knows whenand where to look for them. They do not leave their roosting placevery early in the morning, as they prefer to wait until the rising sunhas, at least partially, dried the dew off the grass ; in winter or latein fall, when every blade of grass, twig, or spray of vegetation is
18 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
white with hoarfrost, and when the feeble rays of the sun are latein rising, they are slow to venture out. But usually by an hour aftersunrise they are afoot toward some convenient weed patch, stubblefield, berry patch, or cultivated field. Here they feed for an houror two, filling their crops, and then retire to some sheltered spot fora midday siesta, digesting their food, dusting or preening theirplumage, or merely basking in the sun or dozing. About two hoursbefore sunset they return to their feeding grounds again for an-other feast before going to roost at dusk.The food of the bobwhite has been exhaustively studied, and a massof material has been published on it. Space will not permit anydetailed account of it here; I can give only a general idea of it.The most complete account of it that I have seen is given by SylvesterD. Judd (1905) of the Biological Survey, to which the reader is re-ferred. He says that the bobwhite is "one of our most nearly om-nivorous species. In addition to seeds, fruit, leaves, buds, tubers, andinsects, it has been known to eat spiders, myriapods, crustaceans,mollusks, and even batrachians." In analysis of 918 stomachs, col-lected during every month in the year, in 21 States and in Canada,the food for the year as a whole consisted of vegetable matter, 83.59per cent, and animal matter, 16.41 per cent, mixed with some sandand gravel. Of the vegetable food, grain constituted 17.38 per cent,seed 52.83 per cent and fruit 9.57 per cent; the grain was probablymostly waste kernels, and the seeds were mainly weed seeds; not asingle kernel of sprouting grain was found in any of the crops orstomachs ; and there is no evidence that quail ever do any damage tostanding crops. The fruits eaten were practically all wild fruits.The animal matter was distributed among beetles, 6.92 per cent;grasshoppers, 3.71 per cent; bugs, 2.77 per cent; caterpillars, 0.95per cent; and other things, 2.06 per cent. From October to Marchthe food is almost entirely vegetable matter, but late in spring andin summer it is made up largely of insects, August showing 44.1 percent of insect food. The insects eaten are mostly injurious species,many of which are avoided by other insectivorous birds, such as
" the potato beetle, twelve-spotted cucumber beetle, striped cucumberbeetle, squash lady-bird beetle, various cutworms, the tobacco worm,army worm, cotton worm, cotton bollworm, the clover weevil, cottonboll weevil, imbricated snout beetle, May beetle, click beetle, the red-legged grasshopper, Rocky Mountain locust, and chinch bug."Since the above was written, the author has seen Stoddard's (1931)much more elaborate account of the food and feeding habits of quailin the Southeastern States, contributed by C. O. Handley. DoctorJudd's report covered a wider territory, and the stomachs wereobtained for each month of the year, but most of them were taken
EASTERN BOBWHITE 19late in fall and in winter, and there were no stomachs of young birdsexamined. Mr. Handley's report is based on the examination ofthe food of 1,625 adult and 42 young bobwhites; it covers 53 pagesand is far too voluminous and too elaborate for me even to attemptto quote from it. It should be carefully studied. A condensed tablegives the monthly and yearly percentages of the various items in thefood. The total yearly averages show 85.59 per cent of vegetableand 14.41 per cent of animal food. The principal items in thevegetable food are: Fruits, 19.41; legumes, 15.17; mast, 13.42; grassseeds, 10.65; and miscellaneous seeds, 10.24 per cent; and in theanimal food: Orthoptera, 7.43; Coleoptera, 2.98; Hemiptera, 1.96;and other insects, 1.06 per cent. For all the interesting details thereader is referred to this exhaustive report.E. L. Moseley (1928) gives a striking illustration of the value ofbobwhites as destroyers of potato beetles in Ohio, where these birdshave increased enormously under 10 years of rigid protection. Hesays:For several years past potatoes have been raised successfully on many farmsin Ohio without spraying for beetles, or taking any measures to combat theinsects. In fact many patches have been practically free from the " bugs."Bob-whites have been observed to spend much of the time among the potatovines. They have been seen to follow a row, picking off the potato beetles.When the potato patch was located near woodland there was no trouble withthe beetles ; but when the patch was near the highway or buildings, even onthe same farm, the insects were troublesome. On farms where the Bob Whitefound nesting sites and protection, the potato vines, if not too near the buildings,were kept free from the insects. A patch of potatoes surrounded by openfields, without bushes, tall weeds, or crops that might shelter the Bob White,was likely to be infested with beetles. A farmer living eight miles south ofDefiance raised about fifty Bob Whites on his place. During the two years thatthese birds were there he had no trouble with insects on either potatoes or cab-bage. The following autumn a number of the birds were killed by hunters, whileothers were frightened away. The next summer the potato beetles were backin numbers. The farmer is again raising Bob Whites and protecting them fromhunters.Mrs. Margaret M. Nice (1910) found that a captive bobwhite ate568 mosquitoes in two hours, another 5,000 plant lice in a day, andanother 1,000 grasshoppers and 532 other insects in a day; also thatit ate from 600 to 30,000 weed seeds each day, according to the size ofthe seeds and the bird's capacity. I can not give here a complete listof the food of the bobwhite, as given by Doctor Judd (1905), but afew of the most important seeds are those of various grasses, rushes,sedges, sorrel, smartweed, bindweed, duckweed , lupine, clover,vetches, spurges, maples, ashes, oaks, pines, violets, morning-glory,ragweed, sunflower, beggarticks, and foxtail and witch grass.Among the fruits are waxmyrtle, barberry, bayberry, mulberry,
20 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthimble berries, blackberries, wild strawberries, rose hips, wild apples,cherries, poison ivy, sumacs, holly, black alder, bittersweet, frostgrapes, blueberries, huckleberries, elderberries, viburnums, honey-suckle, partridgeberry, and woodbine. Wherever the foregoingplants are cultivated or allowed to grow in profusion, bobwhites willfind abundant food all through the year and will be encouraged toremain, with profit to the farmer and joy to the sportsman.The more important items of insect food have been mentionedabove. From 35 to 46 per cent of the summer food of adults con-sists of insects, but the young chicks eat a much larger proportion ofthis food. Small beetles of various kinds, weevils, small grasshop-pers, caterpillars, ants, stink bugs, spiders, and thousand legs havebeen identified in the food of small chicks.Behavior.?When a flock of quail suddenly bursts into the air fromalmost underfoot the effect is startling and gives the impression ofgreat strength and speed. They have been referred to as featheredbombshells. Such sudden flights of a whole bevy in unison are due tothe fact that they have crouched, trusting to their wonderful powersof concealment, until the very last moment, when they are forced tomake a quick get-away. From their crouching attitude they are inposition to make a strong spring into the air, giving them a good start,which their short but powerful wings continue in a burst of speed.Such bombshell flights are the rule when the birds are feeding inclose formation, or when suddenly disturbed in their roosting circles.At other times their flight is much less startling but often quite asswift. I have often seen a single quail, or a pair or two, rise and flyaway as softly and as silently as any other bird, when not alarmed.Their flight is not long protracted and generally ends by scaling downon set wings into the nearest cover. In settling, a flock usuallyscatters, to be joined together later by the gather call. Often singlebirds and sometimes a whole flock will alight in a tree, if alarmed.When leaving the tree their flight is silent and usually scaling down-ward. That they are not capable of long flights is shown by the factthat they become very much exhausted in flying across wide riversand have even been known to drop into the water in attempting suchflights.Stoddard (1931) made a number of tests to determine the speed ofbobwhites in flight. " These showed a speed for mature birds rang-ing from 28 to 38 miles an hour. It seems fair to estimate that thesportsman's hurtling mark sometimes exceeds 40 miles an hour, andbirds just ahead of ' blue darters ' are believed to go even faster forshort distances."Quail do much of their traveling on foot, and they are greattravelers. They cover considerable ground in a day's routine, and
EASTERN BOBWHITE 21
a bevy may be found in any one of several feeding places. In somesections they are said to make seasonal migrations from one typeof country to another, the journeys being made largely on foot.It is no uncommon occurrence to see a pair in spring, or a flock in fall,running along or across a country road. They make a very smartand trim appearance, with bodies held erect and heads held high,as they run swiftly along on their strong little legs. If too hardpressed they rise, flit gently over a fence or wall, and disappear.One can not help admiring their graceful carriage and their effi-ciency as runners. I believe they prefer to escape from their enemiesby running, until too hard pressed; a bird dog will often trail arunning bevy for a long distance.Their characteristic method of roosting in a close circle, withbodies closely packed and heads facing out, is well known. Forthis they select some sheltered spot under an evergreen tree or thickbush, or in some dense tangle of briers or underbrush. Sometimesthey select a small island in a river or a pond for a roosting place.If not disturbed they will occupy the same spot for many nightsin succession, as evidenced by an increasing circle of droppings.Miss Althea R. Sherman told me that she had seen young quail, onthe day they were hatched, assume the circular arrangement of aroosting covey, heads outward and tails in the center of the circle.An interesting account of how this circle is formed is given by Dr.Lynds Jones (1903) based on an observation by Robert J. Sim underespecially favorable circumstances:First one stepped around over the spot selected, then another joined him,the two standing pressed close together, forming the first arc of the circle.Another and another joined themselves to this nucleus, always with headspointing out, tails touching, until the circle was complete. But two were leftout ! One stepped up to the group, made an opening, then crowded himselfin, with much ruffling of feathers. One remained outside, with no room any-where to get in. He, too, ran up to the circle of heads, then round and round,trying here and there in vain ; it was a solid mass. Nothing daunted, henimbly jumped upon the line of backs pressed into a nearly smooth surface,felt here and there for a yielding spot, began wedging himself between twobrothers, slipped lower and lower, and finally became one of the bristlingheads. In this defensive body against frost and living enemy we may leavethem.But quail do not always roost on the ground. Mr. Sandys (1904)says thatit is no uncommon thing to find them regularly roosting in such places as a massof wild grape vines attached to a fence or a tree, in some thick, bushy tree, in anapple tree near the poultry, sometimes in the fowl-house, barn, or stable, on thelower rails of a weedy fence, on top of logs, and occasionally on the bare railsof a fence.74564?32 3
22 BULLETIN- 162, UNITED STATES NATIONAL MUSEUMThe ability of quail to hide and escape detection under the mostscanty protection is truly remarkable. One is often surprised to seea bird or a whole covey arise from a spot that seems to offer no chancefor concealment. Their ability to withhold their scent under suchcircumstances will be referred to later. Mr. Forbush (1927) relatessome interesting observations on a bobwhite that spent a winter inhis yard and became quite tame. He escaped the notice of a wan-dering dog by squatting on bare ground. A slow, quiet settling ofhis whole body was followed by the widening of the shoulders and anindrawing of the head, and, shaking out his feathers, be squatted onthe snowy ground " as flat as a pancake." The white markings ofthe throat and head were cunningly concealed, the top of the headprojecting barely enough beyond the general outline to allow him acomprehensive view of his surroundings. Once he effaced himselffrom sight in a little hollow at the foot of a tree, where he wasinvisible even through a glass at 40 feet away, until he " grew " outof the ground and walked away. Again he faded from view in acleft in a stump less than 3 inches deep. Where there are dry leavesor grass concealment is easy.Voice.?The most characteristic and best-known note of the bob-white is the spring call, or challenge note, of the male, from whichits name is derived. It is heard all through the breeding season insummer. It is subject to considerable individual variation and hasbeen variously interpreted as bob-white, more-wet, no-more-wet, peas-most-ripe, buck-wheat-ripe, wha-whoi, sow-more-wheat, and others.This call is subject to considerable variation; the number of thepreliminary bobs varies from one to two or rarely three; sometimethese first syllables are entirely omitted and we hear only the loudwhite, which again may be shortened to whit. Aretas A. Saunders,who has made a study of the voice of the bobwhite, has sent me someelaborate notes on it. He says that the pitch in this call, countingall his records, varies from G" to F" ', one tone less than an octave.One 3-note call covered this whole range, but the 2-note calls generallybegin on A", most commonly have the white note begin a tone higher,and slur up a single tone or a minor third. Sometimes the second notegives more accent and time to the first part of the slur, and sometimesthe lower note of the slur is on the same pitch as the first note. Theleast range of pitch is shown in a 2-note call beginning on C, startingthe slur on C#, and ending on D. What he calls the slur comes, ofcourse, in the last, or white, note.The bobwhite note is almost invariably given while the bird isstanding on some favorite perch, but R. Bruce Horsfall writes to methat while visiting in Virginia, on August 2, he saw a male bobwhitefly across an old orchard, with few remaining trees but much uncut
EASTERN BOBWHITE 23grass, uttering this note in flight, fully a dozen calls in rapid suc-cession, ceasing only with the termination of the flight.Although many writers refer to the " bobwhite " note as the callof the cock bird to his sitting mate, Stoddard (1931) says:We respectfully express our belief, based upon all tbe data we bave been ableto obtain personally, tbat tbe " bobwhite " call note is largely tbe call of theunmated cocks ; ardent fellows eager to mate, but doomed to a summer ofloneliness, from lack of physical prowess or an insufficient number of hensto go around.The sweetest and loveliest call, entirely different from the fore-going or the following, is the 4-syllable whistle of the female, usedto answer the male in spring and to call the young later in theseason. My father, who was an expert whistler as well as a keensportsman, could imitate this note to perfection. He often amusedhimself, when bobwhites were whistling in spring, by concealinghimself in some thick brush and answering the bobwhite call ofthe male with this enticing note. It was amusing to see the effecton the cock bird, as he came nearer at each repetition of the answerto his call, looking in vain for his expected mate, and sometimescoming within 20 feet before detecting the deception. Once twococks came to look for the anticipated hen ; then a lively fight ensued,all on account of an imaginary bride. This call consists of fournotes, the first and third short, soft, and on a low key, and thesecond and fourth longer, louder, richer, and on a much higher key.I have seen it written je-hoi-a-cMn, or whoooeee-che, but to me itsounds more like a-loie-a-hee. It is a beautiful, soft, rich note,with a decided emphasis on the second syllable, of a liquid qualitywith no harsh sounds.The third whistling note is the well-known gather call, so oftenheard during the fall when the flock has become scattered and thebirds are trying to get together again, particularly toward nightwhen they are gathering to go to roost. It has also been called thescatter call. It is a loud, emphatic whistle of two parts, slurredtogether, with an emphasis on the first. It has a human quality andto my mind is much like the whistle that I use to call my dog. Itsounds to me like quoi-hee. To Mr. Sandys (1904)it sounds very like ka-loi-hee, ka-loi-hee, especially when the old hen is doingthe calling. There are many variations of it, too, ichoil-kee representing acommon one. It is an open question if the cock utters this call, although someaccomplished sportsmen have claimed that he does. The writer has been aclose observer of quail and would think nothing of calling young birds almostto his feet, yet he has never been able to trace this call to the old male ; thatis, as a rallying call to the brood. He is well aware that young males use it inreplying to the mother, but he has yet to see a male of more than one seasonutter it.
24 BULLETIN 162, UNITED STATES NATIONAL MUSEUMMr. Saunders has three records of this call, which he describes inhis notes as a " repeated, slurred whistle, with usually an 1-like soundbetween the notes, so it sounds like coolee." His records show rangesin pitch from Atf to C", or from Bb to D*.In addition to these three very distinct and striking calls, there isoften heard a subdued, conversational chatter while the birds arerunning and feeding. Doctor Judd (1905) heard, as a part of thecourtship performance, " a series of queer responsive ' caterwalings,'more unbirdlike than those of the yellow-breasted chat, suggestingnow the call of a cat to its kittens, now the scolding of a caged graysquirrel, now the alarm notes of a mother grouse, blended with thestrident cry of the guinea hen. As a finale sometimes came a loudrasping noise, not unlike the effort of a broken-voiced whip-poor-will."Sandys (1904) says:A winged bird running, or an uninjured one running from under brush,preparatory to taking wing, frequently voices a musical tick-tick-tick-a-voy.A bird closely chased by a hawk emits a sharp cackling, expressive of extremeterror. Quite frequently a bevy just before taking wing passes round a low,purring note?presumably a warning to spring all together. When the hen iscalling to scattered young, she sometimes varies the cry to an abrupt Ko-lang,after which she remains silent for some time. This the writer believes to be ahint to the young to cease calling?that the danger still threatens, and isprompted by her catching a glimpse of dog or man. A bevy travelling afootkeeps up what may be termed a twittering conversation, and there is a lowalarm note, like a whispered imitation of the cry of a hen when a hawk appears.Stoddard (1931) describes the above-mentioned notes more elabo-rately, with slightly different interpretations. He also describesseveral others. The crowing or caterwauling note, a rasping call thatvaries considerably, is uttered habitually by the cocks at all seasons.He mentions several variations of the scatter call, used to bringtogether scattered birds or as a morning awakening call, and says:One of the most interesting features of the " scatter " call and its variationsis that it evolves by imperceptible degrees from the shrill, piping " lost call
"
of the baby chicks. This starts out with the newly hatched chicks as ananxious piping hti-hii-hu-hu-ich-e-ivhc-icJw-whee-ioheee with rising inflectionlike do-re-mi of the musical scale.Of the decoy ruse call, he writes
:
One of the strangest calls of bobwhites, and a very important one from thestandpoint of their preservation, is the fine cheeping p-s-i-e-u, p-s-i-e-u, p-s-i-e-ucall, uttered by adults and their baby chicks in unison as the brood is stumbledupon by man or beast. This note, proceeding alike from both the franticparents as they beat about in the dust trying to lure the enemy away, and bythe fleeing chicks as they scatter and hide, proves most confusing to the senses,and is a real quail " sleight of hand " that is apt to leave the confused disturberin such a frame of mind that he questions whether he saw fleeing chicks, orwhether it was all just a trick of the eye. Deciding it was the latter, mostenemies pursue the seemingly wounded parents, which sail away on perfect
EASTEBN BOBWHITE 25
wing after the enemy has been decoyed from the vicinity of the brood. Thou-sands of chicks must be saved yearly by this cleverly executed ruse, in whichparents and chicks display perfect teamwork, even before the latter are a dayold.The alarm note is started " as soon as the chicks have scatteredand hidden or the parents have failed to decoy an intruder away.It consists of a monotonous t-o-i-l?ick, ick, ick, ick; t-o-i-l?ick, ick,ick, t-o-i-l-i-c, t-o-i-l-i-c, t-u-e-l-i-c-k; or t-o-i-l-i-c, ip, ip, ip, tic, tic,tic, t-u-e-l-i-c, t-u-e-l-i-c, ick, ip, etc., uttered with machinelike regu-larity for a time, or as long as danger appears to be imminent."He also mentions a distress call, a " piteous whistled c-i-e-u, c-i-e-u,uttered loudly and as rapidly as the mouth can open and close,"given as old or young birds are captured ; also a " cheeping " orcackling call of the developing chick, referred to as the " flicker call."Then there is the " battle cry," of the unmated cocks, a harsh, scream-ing note, uttered in flight; the food call, " a soft, clucking cu, cu, cu,cu-, and a variety of soft conversational notes."Fall.?When fall comes the bobwhite becomes a quail. Its habitschange entirely, as it forsakes the haunts of man and becomes awild bird. It is no longer a sociable and trusting friend of humanbeings, so it resorts to the fields and woods, where it can find shelterin the brushy tangles. It travels now in coveys made up of familyparties or in larger flocks of more than one family.Quail are not supposed to be migratory, in the usual sense of theword, and in many sections, New England, for instance, I believethat they are practically sedentary throughout the year. In somesections, however, they seem to perform short migrations to betterfeeding grounds, or perhaps to escape adverse winter conditions.Audubon (1840) writes:This species performs occasional migrations from the north-west to the south-east, usually in the beginning of October, and somewhat in the manner of theWild Turkey. For a few weeks at this season, the north-western shores of theOhio are covered with flocks of Partridges. They ramble through the woodsalong the margin of the stream, and generally fly across towards evening.Like the Turkeys, many of the weaker Partridges often fall into the water,while thus attempting to cross, and generally perish ; for although they swimsurprisingly, they have not muscular power sufficient to keep up a protractedstruggle, although, when they have fallen within a few yards of the shore, theyeasily escape being drowned. As soon as the Partridges have crossed the prin-cipal streams in their way, they disperse in flocks over the country, and returnto their ordinary mode of life.This habit is also mentioned by Amos W. Butler (1898), who saysthat in Indiana they desert the uplands in fall and congregate inlarge numbers in the Ohio River bottoms; many attempt to crossthe river into Kentucky ; some perish in the attempt and others reach
26 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthe farther shore in an exhausted condition. H. D. Minot (1877)says:In Delaware and Maryland, however, coveys of Quail often appear, whoare distinctively called by the sportsmen there " runners." On the westernside of the Chesapeake, an old sportsman assured me that covey after coveypassed through the country, where food and shelter were abundant, crossingthe peninsula on foot, but often perishing by the wholesale in attemptingto pass the wider inlets, and he added in proof of this that he had takenas many as forty at a time from the middle of the river near his house.But everywhere quail become very restless in fall and are muchgiven to erratic wandering from no apparent cause. They are lesscrazy in this respect than ruffed grouse ; I have never known them tofly against buildings and be killed; but I have frequently seen themin my yard and garden in the center of the city. Mr. Butler (1898)says that " they are found in trees and among the shrubbery in gar-dens, in outbuildings, and among lumber piles. I have seen them inthe cellar window-boxes and over the transoms of the front doors ofthe houses." These wanderings may be due to a latent migratoryinstinct.Game.?Everything taken into consideration, the quail, partridge,or bobwhite is undoubtedly the most universally popular of all NorthAmerican game birds, in spite of the fact that man}' sportsmen con-sider the ruffed grouse the prince of game birds. The sophisticatedgrouse may be the more difficult bird to bag, but the quail, with itssouthern subspecies, has a much wider distribution, nearer to thehaunts of man, is generally more numerous and more prolific, liesbetter to the dog, flies swiftly enough to make good marksmanshipnecessary, and is an equally delicious morsel for the table.One who has never tried it can hardly appreciate the joy and thethrills of a day in the field, with a congenial companion and a braceof well-trained bird dogs, in pursuit of this wonderful game bird.The keen, sparkling October air and the vigorous exercise stimulateboth body and mind. The tired business man breathes more freelyas he starts out from the old farmhouse across the fields for his holi-day with the birds. On a frosty morning, when the grass and herb-age are sparkling white with hoarfrost, it is well not to start tooearly, as quail are not early risers and do not like to get their feetand plumage wet. But when the sun is well up it is time to lookfor them, for they may be traveling along some brushy old fencetoward their favorite buckwheat stubble, one of the best places tofind them. When you reach the field where the birds are expected tobe found, the most interesting part of the sport begins; the intelli-gent dogs have learned to quarter the ground thoroughly and huntin every likely spot where their bird sense leads them; excitementbecomes intense, as they show by their careful movements that they
EASTERN BOBWHITE 27have scented game; and, finally, the sudden stop and the rigid pose,with nose pointed toward the birds, bring the climax, as the sports-men step up and the covey bursts into the air with a whir of wings.A good shot may bag two or even three birds on the first rise ; I haveseen men that boasted of stopping as many as five with an automaticrepeater, but I have never seen one do it ; and I have seen many cleanmisses. The rest of the covey have flown straight to the nearestcover, perhaps scattered in several directions, some into a patch ofscrub oaks on a hillside, some into the tangled underbrush in aswampy hollow, and others into the nearest woodlot. The menshould mark them down, but had better leave them for a while untilthey begin to run about and leave a little scent ; otherwise they willbe very hard to find. Picking up these scattered singles is hardenough at best; it requires good work on the part of the dogs andgives the hunter many difficult shots in unexpected places. The manthat can put two quail in his pocket for every four shells fired is agood shot.Perhaps the birds have not been found in the buckwheat stubble.Each covey has several feeding places and it is necessary to coverconsiderable ground, hunting the wheat, rye, oat, and corn stubbles,especially if overgrown with ragweed or other weeds, as well as anyother old neglected fields and weed patches where the birds can findfood and shelter. Sometimes the dogs will show signs of game in alikely spot but fail to find the birds; quail often make short flightsfrom one field to another, thus breaking the scent. Sometimes aflushed covey will be marked down very carefully in a fairly openfield and be immediately followed up; but a careful search by ex-perienced men and good dogs will fail to reveal the presence of asingle bird. This has caused much controversy as to the power ofquail to withhold their scent. The explanation probably is that therapid passage through the air dissipates most of the scent from theplumage ; the birds, being frightened, crouch low on the ground withfeathers closely pressed against the body, shutting in body odors;and as they have not run any there is no foot scent. It has oftenhappened that, in a later search over the same ground, after thebirds have begun to run about, they have been readily found. Therehas been no willful or even conscious withholding of scent.For about four hours during the middle of the day, quail retirefrom their feeding grounds for their noonday rest. The huntersmay as well do likewise, until the birds come out to feed againabout two hours before sunset. The hours of waning daylight oftenfurnish some of the best and most interesting shooting ; the scatteredcovey is anxious to get together before roosting time; and thehunters get the final thrills of the day as they hear the sweet,
28 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMgentle gather call, quoi-hee, quoi-hee, from a distant patch of scruboaks, an answering call from the brier patch in the swale, and an-other from the edge of the near-by woods. They are content to callit a day and leave the gentle birds to settle down for the night.Enemies.?Quail have numerous enemies, furred, feathered, andscaled, but fortunately they are such persistent and prolific breedersthat they can stand the strain from natural enemies if man will givethem half a chance.Stray cats, or domestic cats run wild, are doubtless the mostdestructive enemies of quail. They catch and devour enormous num-bers of both young and old birds, as they hunt them day and night.Mr. Forbush (1927) gives some striking illustrations of this andspeaks of one big cat that is said to have killed more than 200 bob-whites. Dogs that are allowed to run loose and hunt independentlykill a great many old and young birds. Stray cats in the woodsand fields should be shot on sight. Domestic cats and dogs shouldbe restrained during the nesting season.In Jamaica the mongoose is said to have virtually exterminatedthe introduced quail. Foxes, minks, and weasels kill some birds, butthey probably find rabbits easier to catch and more to their liking.Raccoons, opossums, skunks, and rats destroy a great many eggs.Among bird enemies the crow is one of the worst. Crows arevery clever in hunting up nests and destroy a great many eggs;they have even been known to kill the adult birds in winter. Crows,in my opinion, should be shot whenever possible, for they can notbe much reduced at best. Cooper's hawk is probably the worst ofthe hawks. The goshawk and the sharp-shinned hawk are almostas bad. Red-tailed hawks have been known to kill quail, but theyare too slow to catch very many, and they are useful as rodent de-stroyers. Great horned and other owls must be reckoned with, butthe former is very fond of skunks, and all the owls keep the destruc-tive rodents in check. Quail have learned that brier patches andthick tangles offer good protection against their enemies in the air.Any of the larger snakes will eat the eggs and probably destroya great many, but here again we must give them credit for livinglargely on the rodent enemies of the bobwhite. Major Bendire (1892)speaks of a large rattlesnake, killed in Texas, that had swallowedfive adult quail at one meal, and another that had taken four bob-whites and a scaled quail.In his chapter on mortality, Stoddard (1931) states that of 602nests studied about " 36 per cent were more or less successful andabout 64 per cent unsuccessful." The failures were due to nest de-sertion, destruction by natural enemies, destruction by the elements,rains, floods, or droughts, and disturbance by human beings, by farm
EASTERN BOBWHITE 29
work, or by poultry and cattle. Among the destroyers of eggs hementions, in addition to the enemies named above, blue jays, turkeys,and red ants; the ants enter the egg as soon as the membrane ispunctured by the emerging chick, which is literally eaten alive; outof 278 nests studied by Louis Campbell in 1928, 34 were taken overby ants.After hatching, young quail are preyed upon by most of the moreactive enemies named above, to which must be added turkeys, guineafowl, pheasants, and shrikes. The chief winged enemies of the olderyoung and adults are Cooper's and sharp-shinned hawks, and in theNorth the goshawk. The Buteos are mainly, or wholly, beneficial.Mr. Stoddard (1931) exempts the sparrow hawk from blame andsays : " In several instances individuals took up quarters temporarilyon the fence posts of propagating enclosures and made forays againstthe large grasshoppers on the ground beneath, without harming thequail chicks in the least." In favor of the marsh hawk, he writes:In view of the fact that not more than 4 quail were discovered in approxi-mately 1,100 pellets, marsh hawks can hardly be accused of making any seriousinroads on the number of quail in the region. On the other hand, one or morecotton rats were found in 925 of these pellets. Since cotton rats destroy theeggs of quail, the marsh hawk is probably the best benefactor the quail has inthe area, for it is actively engaged in reducing the numbers of these rodents.Remains of at least 14 snakes, most of which were colubrines, were discovered.These also are probably eaters of quail eggs.Diseases.?The chapters on parasites and diseases, in Stoddard's(1931) report, were contributed by Dr. Eloise B. Cram, Myrna F.Jones, and Ena A. Allen, of the Bureau of Animal Industry.They are well worth careful study, but are too long (110 pages) andtoo technical for any adequate presentation here. Suffice it to saythat bobwhites are attacked by many of the same parasites andsuffer from many of the same diseases as ruffed grouse. Amongthe Protozoa the most important are those which cause malaria,coccidiosis, and blackhead. Nematodes, or roundworms, were found
" in a high percentage of the birds examined " ; 16 species were iden-tified, and their life histories explained. In the intestines fivespecies of tapeworms were found and similarly described. As ex-ternal parasites, lice, ticks, mites, and fleas are mentioned. Amongthe nonparasitic diseases the following are fully described: Footdisease, bird pox, dry gangrene, chicken pox, " nutritional roup,"aspergillosis, " quail disease," and tularemia. This brief summaryand other references to Stoddard's (1931) work give a very inade-quate idea of the wealth of material that his exhaustive report con-tains; it must be read to be appreciated; some of the interestingchapters can not even be summarized here.
30 BULLETIN 162, UNITED STATES NATIONAL MUSEUMWinter.?In the southern portions of their range, where quail en-joy open winters, their habits and haunts are about the same asduring fall ; but in the northern regions of ice and snow they havea hard struggle for existence and many perish from hunger and coldin severe winters. Quail have been known to dive into soft snow-drifts for protection from severe cold; Sandys (1904) says hehas caught them in such situations. More often, at the approach ofa snowstorm, they huddle together in some sheltered spot and letthe snow cover them. This gives them good protection from thewind and cold; but if the snow turns to rain, followed by a severefreeze the birds are imprisoned and often perish from hunger be-fore they can escape. Birds seldom freeze to death, if they can getplenty of food, but cold combined with hunger they can not stand.Mr. Forbush (1927) tells an interesting story of a man who hadbeen feeding a covey of quail ; for 10 days after a heavy snowstorm,followed by a thaw and freeze, they failed to come to their usualfeeding place; believing them to be imprisoned under the snow hewent to the place where they were accustomed to sleep and brokethe crust ; the next day they came to feed and a search showed thatthey had found the place where he had broken the crust for them.Quail often find more or less open situations where they get someshelter, under logs or fallen trees, under thick evergreens, in tanglesof briers, in brush piles, or under banks with southern exposure; insuch places they find bare ground and can pick up some food, as wellas the gravel or grit that they need. They avoid open places and donot like to travel on snow, where they are so conspicuous; but theyhave to go out to forage for food, such as the seeds of weeds, project-ing above the snow, rose hips, dried berries, seeds of sumac, bay-berries, and other plants. When hard pressed they often visit thebarnyard to feed with the poultry. Farmers, sportsmen, boy scouts,and many other persons make a practice of feeding quail regularly inwinter. They should have a shelter, made of brush, evergreenboughs, or corn stalks, open at both ends so that the birds can escapeat either end. The ground under this should be kept bare and wellsupplied with almost any kind of grain and plenty of grit. Quailwill come regularly to such places and the lives of many will be saved.DISTRIBUTIONRange.?Chiefly the Eastern United States, ranging west to east-ern Texas, eastern Colorado, and the Dakotas. The great interestshown by sportsmen in the bobwhite has resulted in introductions overthe entire country. Though many of these experiments have resultedin failure, others have been notably successful, and in some regionsthe introduced birds have spread out and met those that are indige-
EASTERN BOBWHITE 31
nous, thus causing an actual extension of range. The problem is fur-ther complicated by the fact that in addition to true virginianus, in-troductions of so-called Mexican quail {Colinus v. texanus) have beenmade in several States. As these have readily crossed with thenative birds, and as individuals show a great deal of variation, it isfrequently difficult to determine the natural limits of the differentraces. The following summary, however, presents a reasonablyaccurate picture of the natural range of the species
:
North to southeastern Wyoming (Horse Shoe Creek) ; South Da-kota (Faulkton) ; North Dakota (Bartlett and Larimore) ; Wisconsin(Danbury and Menominee) ; Michigan (Douglas Lake and AlconaCounty) ; southern Ontario (Mount Forest, Listowel, Toronto, andPort Hope) ; Vermont (Londonderry) ; and Maine (West Gardiner).East to Maine (West Gardiner and West Fryeburg) ; rarely easternNew Hampshire (Hampton) ; Massachusetts (Gloucester, Boston,and Cape Cod) ; Rhode Island (Newport) ; New York (ShelterIsland and Roslyn) ; New Jersey (Red Bank, Vineland, and Sea IsleCity) ; Delaware (Lincoln) ; Virginia (Belle Haven, Eastville, CapeCharles, Norfolk, and Dismal Swamp) ; North Carolina (CurrituckSound and Raleigh) ; South Carolina (Waverly Mills, St. HelenaIsland, and Frogmore) ; Georgia (Savannah, Riceboro, Jekyl Island,and Okefenokee Swamp); and southern Florida (Miami). Southto southern Florida (Coconut Grove, Indian Key, and Key West)
;
Alabama (Bon Secour and Mobile) ; Mississippi (Biloxi and BaySt. Louis) ; Louisiana (New Orleans, Houma, Abbeville, Mermentau,and Iowa Station) ; southeastern Texas (Galveston, Corpus Christi.and Brownsville) ; Tamaulipas (Tampico, Altamira, and Victoria)and southern Nuevo Leon (Mier y Noriega). West to Nuevo Leon(Mier y Noriega and Montemorelos) ; Coahuila (Sabinas) ; westernTexas (Langtry, Lozier, Fort Stockton, and Pecos) ; probably south-eastern New Mexico (Carlsbad) ; eastern Colorado (Monon, Beloit,Yuma, and Crook) ; and southeastern Wyoming (Uva and HorseShoe Creek).Introductions.?Introductions have been made in Colorado (Up-per Arkansas Valley, Wet Mountains, Pueblo, Denver, Estes Park,Loveland, Fort Collins, Greeley, Saguache, and Grand Junction)Utah (Salt Lake Valley) ; Montana (Sappington, Anaconda, Flat-head Lake, and Kalispell) ; Idaho (Boise, Nampa, Coeur d'Alene,and Rathdrum) ; Oregon (Lake Alvord, Snake River Valley, Pen-dleton, Scio, Dayton, and Portland) ; Washington (Starbuck, Whid-bey Island, Walla Walla, Cheney, Spokane, Osoyoos Lake, Olympia,Tacoma, Seattle, and Blaine) ; and California (San Felipe andGilroy)
.
32 BULLETIN 162, UNITED STATES NATIONAL MUSEUMThis species also has been transplanted to southwestern Canada(British Columbia and Manitoba) : the West Indies (Cuba, Jamaica.New Providence, Haiti, Porto Rico, Barbados, St. Kitts, Antigua,St. Croix, and Guadeloupe); New Zealand; England (Norfolk);Sweden; Germany (Hanover) ; France; and China (Kashing). Theintroductions in the West Indies (in some instances the Cuban form,C. v. cubanensis) have been more or less successful, but so far asknown all European and Asiatic attempts to acclimatize this birdhave been total failures.The foregoing distribution covers the range of the entire species,which is subdivided into three subspecies. C. v. floridanus occupiesthe whole of Florida from the vicinity of Gainesville, Palatka, andTarpon Springs southward. C. v. texanus occupies central andsouthern Texas, from southeastern New Mexico to northeasternMexico. Typical virginianus occupies the rest of the range in East-ern and Central United States.Egg dates.?Massachusetts, Rhode Island, and Connecticut: 22records, May 20 to October 10; 11 records, June 5 to July 28. NewYork, New Jersey, and Pennsylvania : 18 records, May 21 to August31 ; 9 records, June 7 to July 16. Virginia, Kentucky, and Kansas
:
18 records, May 14 to September; 9 records, June 2 to July 23.South Carolina and Georgia : 15 records, April 24 to September 16
;
8 records, May 21 to June 18. Indiana, Illinois, and Iowa: 31 rec-ords, April 28 to October 16 ; 16 records, June 2 to July 13.Florida {floridanus) : 46 records, April 19 to July 26; 23 records.May 13 to June 8.Texas {texanus) : 50 records, March 18 to August 19; 25 records,May 11 to June 2.COLINUS VIRGINIANUS FLORIDANUS (Coues)FLORIDA BOBWHITEHABITSThe Florida bird is merely a small, dark variety of the common,northern bobwhite. It is confined entirely to the peninsula of Flor-ida, where it is universally common and generally distributed in allthe drier portions of the State, except in the extreme north and theextreme south; in the north it intergrades with the northern formand in the south with the still smaller and darker Cuban form.If due allowance be made for the different environment in whichthey live, the Florida birds will be found in similar localities to thosechosen by their northern relatives and their habits are essentiallythe same. Nearly everything I have written about the northern birdsapplies equally well to the southern. They are equally fond of the
FLORIDA BOBWHITE 33
society of human beings, where they probably feel more secure fromtheir wild enemies and perhaps find more food. Major Bendire(1892) quotes Doctor Ralph, as saying that " localities they like bestare open woods grown up with saw palmettos or low bushes, orfields with woods near them, and they are particularly fond ofslovenly cultivated grounds that have bushes and weeds growingthickly along their borders." I have frequently seen them in smallvillages, in gardens, and about houses ; but I have more often foundthem in the open flat pine woods where there are extensive patchesof saw palmettos; these thick clumps of low palmettos are oftenalmost impenetrable and afford them excellent protection fromtheir enemies.Nesting.?Donald J. Nicholson has sent me the following notes onthis subject:These quail begin to pair off by February- and by March most of themhave chosen their mates, but still some will be found in coveys into March.In March the bobwhites begin to think of domestic duties, and the woodsand cultivated fields resound with their cheery bob-bob-ichite. This continuesuntil late June, when the calls become very much less frequent. A few pairsbreed as early as March or early April, but the height of the nesting seasonis late May or early June. They also nest up into August, and one nest wasfound in December by a hunter, which is quite unusual. From 9 to 16eggs are laid, which are deposited in arched nests of dead grass, in oldfields, in pine woods, or on the edges of grassy ponds. The female does notcommence to sit until the last egg is deposited. How long it takes to incu-bate is unknown to me. The birds are quite suspicious if a nest is found andwill generally leave it, but if not touched sometimes continue their duties.When a nest is found the bird sits until almost trodden upon, then eitherflies directly off with great speed, or more often flits off and cackles excitedly,running about close by and feigning lameness.Eggs.?The number of eggs laid averages less, but the eggs arequite indistinguishable from those of the northern bird. They arenot even appreciably smaller. The measurements of 51 eggs aver-age 30.6 by 23.8 millimeters; the eggs showing the four extremesmeasure 32.6 by 24.1, 31.5 by 24.5, and 28.9 by 22.8 millimeters.Young.?Mr. Nicholson says in his notes:Both parents attend the young and are very courageous in the defense ofthe little fellows, running about close to the intruder, jumping up to attractattention, and uttering strange notes. When the hiding spot of the young isdiscovered they scurry for other shelter, cheeping as they run, and if pressedtoo closely will take flight, when very young. I once saw a bobwhite makea mad rush at the wheels of a passing auto, and after the machine passed themate also came out. Undoubtedly there were young near by.Behavior.?In general behavior and habits the Florida bird doesnot differ materially from its northern relative. Its food does notseem to have been separately studied and is included in reports onthe species as a whole. It feeds on such of the seeds, fruits, and
34 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMinsects, mentioned in such reports, as are to be found within itsrange. Its voice is essentially the same. Once exceedingly abundantin the open country in Florida, it has been greatly reduced in num-bers and mainly by its human enemies. It has long been a favoritewith sportsmen; its long open season, with little or no protectionin many places, has offered them attractive opportunities long afterthe season in the North has closed. A steadily increasing numberof winter visitors have been tempted to spend part of their idletime in hunting, where the shooting is mostly open and the birdseasily killed. The quail have shown the effect of this slaughter.They have also suffered greatly from illegal trapping and nettingand from shooting out of season, mostly by poor whites and negroes.Fortunately for the future of the Florida bobwhite the laws arenow better enforced and there are many large areas where no shoot-ing is allowed and where public sentiment is protecting this andall other desirable birds.COLINUS VIRGINIANUS TEXANUS (Lawrence)TEXAS BOBWHITEHABITSGeorge N. Lawrence (1853) first called attention to the charactersthat separate the bobwhite of Texas from the two eastern forms. Ingeneral appearance it is decidedly grayer than either of the easternforms. Mr. Lawrence described it more in detail, as follows
:
This somewhat resembles O. virginianus, but is smaller, and differs also inhaving the lores white, in being without the conspicuous dark markings on theback and wings, and the bright chestnut red so prevalent in the upper plumageof that species; the bill is proportionately longer and narrower, the legs moreslender, and the black markings on the abdomen and breast are fully twiceas broad.The Texas bobwhite closely resembles the eastern form in behavior,habits, and haunts. George Finlay Simmons (1925) gives us thefollowing long list of places in which it may be found
:
More or less open country, particularly in mesquite-chaparral-cactus pas-tures ; old plantations, clearings, and cultivated fields ; open, semi-arid tree-dotted, bushy pastures in farming country, particularly where such pastures areinterspersed with small bodies of woodland ; hay, grain, brown stubble, corn,cotton, and open weedy fields ; clearings and brushy edges of woodlands
;
thickets, brush, and briar patches along edges of meadows and creeks; weedyroadsides and fencerows; wooded hillsides; cedar brakes; cultivated fields.Never in bottom woods or open praries. In fall and winter, among dead stalksin cotton and corn fields.
TEXAS BOBWHITE 35Nesting.?Major Bendire (1892) says:The favorite nesting site of the Texan Bob White is a bunch of sedge grass.A slight cavity is made in the center, this is lined with a few straws andarched over with similar material. Sometimes a covered way or tunnel leadsto the entrance of the nest. Occasionally a nest is placed under a bush andnot covered or arched. Two broods are usually raised in a season, and eventhree at times.Mr. Simmons (1925) mentions a number of other nesting sites asfollows
:
Nest on ground along fencerows, in small dewberry thickets, in prickly-pearbeds, in brushy mesquite lands or bushy, grassy pastures, in thick tussocks orclumps of big grass, along weedy fence lines or roadsides, or in the middle ofcotton or corn fields on the rolling prairies ; usually well hidden by a small bushor by weeds and prairie grasses ; occasionally placed in a fencerow with over-hanging vines or beside a stone wall or a log.Eggs.?The eggs of the Texas bobwhite are indistinguishable fromthose of the eastern bird. From 10 to 15 eggs constitute the usualset, but 18 or 19 are occasionally laid, and Bendire (1892) states thatH. P. Attwater once found 33 eggs in a nest. Probably this largeset was the product of two females. According to Bendire (1892),
" two broods are usually raised in a season, and even three at times."The measurements of 59 eggs, in the United States National Museum,average 30 by 24 millimeters; the eggs showing the four extremesmeasure 32 by 25, 30 by 25.5, and 27.5 by 22 millimeters.Plumages.?The molts and plumages correspond to those of theeastern bobwhite, but Dr. Jonathan Dwight (1900) says that "thejuvenal plumage is browner than in virginianus."Food.?Mrs. Florence Merriam Bailey (1928) writes:The Bob White is of special agricultural value because it destroys a largeamount of weed seed and a considerable number of insects. Half of its foodis weed seed, only a fourth grain?mainly from the stubble fields?and abouta tenth wild fruits. Fifteen per cent is composed of insects, including severalof the most serious pests of agriculture. It feeds freely upon Colorado potatobeetles and chinch bugs, and eats also grasshoppers, cucumber beetles, wire-worms, billbugs, clover-leaf weevils, the Mexican cotton-boll weevils, armyworms, cotton worms, cutworms, and Rocky Mountain locusts.Behavior.?Mr. Simmons (1925) says:Observed singly, in pairs in summer, or by threes and fours ; in winter,from middle fall to early spring, in coveys or bevies of from 10 to 30, breakingup for the breeding season and reassembling as soon as it is over. Rather shyand difficult to find after the hunting season has opened. During summer daysthe birds seek shelter under the bushes which dot the pastures ; winter days,in scattered brush heaps and tiny hollows. On spring and summer nights theygenerally roost in the open fields ; on fall and winter nights, roost under cover,usually in lowlands. When frightened, or when preparing to sleep and keepwarm, a covey arranges itself in a elose-huddled circle, heads out from center
;
when approach is too close, the birds burst in all directions, making it very
36 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMdifficult to shoot more than one at a time; were they to follow each other, asmost gregarious birds do, the hunter would have a better opportunity toexterminate the whole covey. Occasionally, but rarely, takes to the trees whenflushed, aud remains squatting close to limb and practically invisible.Voice.?The Texas bobwhite has a range of varied calls similarto those of its eastern relative, but the notes are said to be less loud,clear, and ringing.Enemies.?Major Bendire (1892) quotes William Lloyd, as follows:They are very insuspicious, and their low notes, uttered while feeding, attracta good many enemies. I have seen foxes on the watch, and tbe Marsh Harrierperched on a clump of grass on the lookout, waiting for them to pass. But themany large rattlesnakes found here are their worst enemies. One killed inMay had swallowed five of these birds at one meal; another, a female evidentlycaught on her nest and a half dozen of her eggs ; a third, four Bob Whites anda Scaled Partridge. Tbe young are also greatly affected and many killed byheavy rains in June and July ; numbers perish then from cold and protractedwet weather. When alarmed by a Hawk sailing overhead they run under themother for protection, as domestic chickens do.COLINUS BIDGWAYI BrewsterMASKED BOBWHITEHABITSThis well-marked species once inhabited a narrow strip of countryin southern Arizona, where its range extended for not more than40 or 50 miles north of the Mexican boundary. It has long sincebeen exterminated in Arizona and has now perhaps disappeared en-tirely from its more extensive range in Sonora. Herbert Brown wasthe first to discover the presence of this quail in Arizona, and we areindebted to him for practically all we know of its former distributionand habits. He first met the bird in Sonora, hearing its note andsupposing it to be our eastern bobwhite. He says of this incident(1904) :It is not easy to describe the feelings of myself and American companionswhen we first heard the call bob white. It was startling and unexpected, andthat night nearly every man in camp had some reminiscence to tell of Bob Whiteand his boyhood days. Just that simple call made many a hardy man heart-sick and homesick. It was to us Americans the one homelike thing in allSonora, and we felt thousands of miles nearer to our dear old homes in thethen far distant States.In the spring of 1884 a pair of these quail was taken on the easternslope of the Baboquivari Mountains, Arizona, and brought to Brownas specimens; but because of his absence they were allowed to spoiland were thrown away. Meantime he had reported to a local paperthat a pair of bobwhite quail had been taken in Arizona, and the notewas republished in Forest and Stream. This aroused the interest of
MASKED BOBWHITE 37Robert Ridgway, to whom fragments of the two birds were sent foridentification. He pronounced them Ortyx {Colinus) graysoni, aMexican species.During that same year Frank Stephens collected in Sonora, Mexico,a male masked bobwhite for William Brewster, which was describedby Mr. Brewster (1885) as the type of a new species, which he namedColinus ridgwayi. This whole interesting history is given in moredetail, with reference to several published articles on the subject, inan excellent paper by Dr. J. A. Allen (1886), to which the readeris referred.Brown (1904) was told that "in early days they were plentiful inRamsey's Canyon in the Huachucas, and also on the Babacomori, avalley intervening between the Huachuca and Harahaw ranges."Speaking of conditions prior to 1870, he says
:
At that time the valley was heavily grassed and the Apache Indians notori-ously bad, a combinrtion that prevented the most sanguine naturalist fromgetting too close to the ground without taking big chances of permanentlyslipping under it. For many years Indians, grass, and birds have been gone.The Santa Cruz, to the south and west of the Souoite, is wider and wasmore heavily brushed. Those conditions gave the birds a better chance forlife and for years they held tenaciously on. Six or seven years ago I was toldby a ranchman living near Calabasas, that a small bunch of Bob-white Quailhad shortly before entered his barnyard and that he had killed six of them atone shot. It was a grievous thing to do, but the man did not know that hewas wiping out of existence the last remnant of a native Arizona game bird.Later I heard of the remaining few haviug been occasionally seen, but forseveral years now no word has come of them.I never found them west of the Baboquivari Mountains, and from my knowl-edge of the country thereabouts I am inclined to fix the eastern slope of thatrange as their western limit. Between that and Ramsey's Canon, in theHuachucas, is a distance of nearly one hundred miles. Their deepest point ofpenetration into the Territory was probably not more than fifty miles, and thatwas down the Baboquivari or Altar valley.As to the causes of the bird's disappearance, he writes:The causes leading to the extermination of the Arizona Masked Bob-white(Colinus ridgwayi) are due to the overstocking of the country with cattle,supplemented by several rainless years. This combination practically strippedthe country bare of vegetation. Of their range the Colinus occupied onlycertain restricted portions, and when their food and shelter had been troddenout of existence by thousands of hunger-dying stock, there was nothing leftfor poor little Bob-white to do but go out with them. As the conditions inSonora were similar to those in Arizona, birds and cattle suffered in com-mon. The Arizona Bob-white would have thriven well in an agriculturalcountry, in brushy fence corners, tangled thickets and weed-covered fields,but such things were not to be had in their habitat. Unless a few can stillbe found on the upper Santa Cruz we can, in truth, bid them a final good-bye.Nesting.?Verj7 little is known about the nesting habits of themasked bobwhite. Brown (1904) offered a reward of $1 an egg74564?32 1
38 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMfor the first nest found for him. A nest containing six eggs wasfound on the mesa on the eastern side of the Baboquivari Mountains." Unfortunately these precious things were lost through the cupidityof the finders, whose expectations ran to more eggs, but while wait-ing for the increase the nest was robbed of the eggs that were thenin it. I was, however, notified of the find, but when I reached thereI found only an empty nest, a bowl-shaped depression in a bunch ofmountain grass."Major Bendire (1892) quotes from a letter received from Otho C.Poling, relating his fruitless attempt to find a nest and reportingthe finding of an egg in the oviduct of a female, which he shot onMay 24, 1890.Col. John E. Thayer has a set of seven eggs in his collection, pre-sented to him by Miss Engel, of Cleveland, Ohio. It was collectedin Sonora, Mexico, on May 4, 1903. The nest was " placed in sandunder a bunch of dry grass." The female was closely observed,but as the females of this and some other Mexican species are prac-tically indistinguishable, there may be some doubt about theidentification.Eggs.?The eggs of the masked bobwhite are indistinguishablefrom those of the eastern bobwhite. The single egg in the UnitedStates National Museum is pure white and unstained, as it was takenfrom the oviduct of a bird by Herbert Brown. This egg measures32.5 b}r 25 millimeters. The egg recorded by Major Bendire, pos-sibly the Poling egg, measured 31 by 24 millimeters ; I do not knowjust what has become of this egg, or how Major Bendire got themeasurements.Colonel Thayer's eggs are slightly smaller; they average 29.6by 23.4 millimeters; the largest egg measures 30.5 by 23.6 and thesmallest 28.3 by 23.1 millimeters.Plumages.?The downy young of the masked bobwhite is un-known. In the juvenal plumage, which is worn for only a shorttime in summer and early in fall, the sexes are alike and closelyresemble the similar stage in the Texas bobwhite. Doctor Allen(1889) describes a young male, taken on October 10 and still partlyin first plumage, as follows.The top of the head is blackish, with each feather narrowly bordered withashy brown. The hind neck, sides of the neck, and jugulum are yellowishwhite, with each feather barred at the tip with black. The scapulars arebrownish, each feather with a rather broad whitish shaft stripe, and barredwith yellowish white and black, and the wing coverts have much the same pat-tern, but the barring is pale cinnamon and brown. The throat is purewhite, with new black feathers appear ng irregularly along the sides of the chinand upper throat. Breast pale browi . with light shaft stripes and faintlybarred with blackish, passing into brownish white with more distinct barson the upper abdomen. The new featheps ulong the sides of the breast and
MASKED BOBWHITE 39flanks are chestnut, tipped with a spot of clear white, which is borderedbehind with a more or less V-shaped bar of deep black. The broad yellowishwhite superciliary stripes extend to the nostrils.The sexes differentiate during the fall, and young males show con-tinuous progress toward maturity during their first winter. Thejuvenal plumage does not wholly disappear until December orlater ; there is more or less white in the black throat and more or lessblack barring on the breast; but the throat becomes clearer blackand the breast and underparts purer " tawny " as the season advances.Food.?Major Bendire (1892) quotes Brown on this subject, asfollows
:
Of three stomachs of this species examined, one contained a species ofmustard seed, a few chaparral berries, and some six or eight beetles and otherinsects, ranging in length from a half inch down to the size of a pin head.The second was similarly provided, but contained, in lieu of mustard seed, agrasshopper fully an inch in length. These two were taken on the mesa. Thethird, from a bird taken in the valley, contained about 20 medium-sized redants, several crescent-shaped seeds, and a large number of small, fleshy, greenleaves.He also says that Lieut. H. C. Benson, who secured a number ofspecimens in Sonora, in 1886, wrote him " that they only frequentedcultivated fields there, where wheat and barley had been raised."John C. Cahoon, who collected in the same section of Sonora, foundthese quail abundant there ; " several large coveys were seen and 8specimens shot in one day " ; he sent 10 specimens, taken February5-8, 1887, to William Brewster (1887) ; they were " haunting patchesof weeds in gardens and barren sand wastes, where they fed on theseeds of a plant called red-root."Behavior.?Brown (1885) says of the habits of the maskedbobwhite
:
They appear to resemble very closely those of the common quail (C. vir-ginianus), only slightly modified by the conditions of their environment. Theyutter the characteristic call, " Bob White," with bold, full notes, and perch onrocks and bushes when calling. They do not appear to be at all a mountainbird, but live on the mesa, in the valleys, and possibly in the foothills * * *.In addition to their " Bob White " they have a second call of hoo-ice, articulatedand as clean cut as their Bob White. This call of hoo-ice they use when scat-tered, and more especially do they use it when separated toward nightfall. Atthis hour I noted that, although they occasionally called "Bob White" theynever repeated the first syllable, as in the daytime they now and then attemptedto do * * *. I will venture to say that when frightened and scattered theyare a hard bird to get. Hear one call, locate it as you may, see one fly andmark it down, and without a dog it is virtually impossible to flush it.Griffing Bancroft wrote to me in 1928, as follows:The masked bobwhite, Collnus rldgwayi, is virtually extinct. Its formerbreeding range was confined to a transition zone somewhat oval in shape and
40 BULLETIN 162, UNITED STATES NATIONAL MUSEUMlying at an elevation of 3,500 to 4,000 feet and over. The western axial centeris situated in Sonora, about 20 miles due south of Nogales. The city ofMagdalena appears to mark the southern boundary, while the northern line liesabout halfway between Tucson and Nogales. The western swing of the circlenaturally is broken and irregular; nevertheless it can be traced quite clearly,as the land falls in every direction but east. As soon as desert conditionsbegin to intergrade with grass and oak, environments no longer seem favorablefor the masked bobwhite.The eastern limits of the bird's range are also determined by altitude. Justwhere the mountains begin to get too high for them it is hard to say, thoughit is not very far east of Nogales. I believe it is about 50 miles and feel quiteconfident it is not more than 100, though I have not traced that line personallyas I have the other.There have been seen within the past two years two small bands of thesequail north and west of Magdalena. Except for this no hunter, sportsman, orobserver with whom I have been able to establish contacts has ever seen thebird or heard of its existence. As my inquiries have been carried on for thepast six years and have covered a large stretch of country and been thorough,my conclusion that the bird is almost extinct does not seem open to question.The reasons for his disappearance are not so clear. The greatest contributingcause appears to be the habit the Mexican wood cutters have of burning thehillsides in order to get better firewood. The rise in the cattle industry, in theprosperity and population of the country, and civilization in general seemsto have wiped out the bobwhite and the turkeys, but not Gambel's or Mearns'squail. Colinus ridgwayi can not maintain itself against civilization. The rea-sons why it can not do so are not wholly obvious.DISTRIBUTIONRange.?Formerly north to southern Arizona (Baboquivari Moun-tains and Huachuca Mountains) . South to northern Sonora (Sesabeand Magdalena). The eastern and western limits determined bythe extent of grassy plains at altitudes of from 3,500 to 4,000 feet.This bobwhite has now disappeared entirely from Arizona and isnearly, or quite, extinct in Sonora.Egg dates.?Arizona : 3 records, May 4 to 12.OREORTYX PICTA PALMERI OberholserMOUNTAIN QUAILHABITSThe common name, which is fortunately more stable than thescientific name, of this quail remains as we have always known it.But Dr. H. C. Oberholser (1923) has discovered that the bird whichDouglas described as Ortyx picta was really the lighter-colored birdof the interior, rather than the darker bird of the humid coastalstrip, to which we have always heretofore applied the name picta.Therefore, the paler bird of the interior must take the name Oreortyxpicta picta, plumed quail. This relegates the name plwnifera to
MOUNTAIN QUAIL. 41synonymy and leaves the bird of the humid coast belt without a name,for which Doctor Oberholser has proposed the name Oreortyx pictapalmeri, in honor of Dr. T. S. Palmer, who had reached the sameconclusion some years ago. The range of this race is restricted tothe humid Transition Zone of the Pacific coast, from southwesternWashington south to Monterey County, Calif.J. H. Bowles (Dawson and Bowles, 19-09) says of the status of thisquail in the State of Washington:The Mountain Quail, as it is generally called, and its close relative, thePlumed Quail, are neither of them native to Washington, several crates ofliving birds having been imported from California between the years 1880and 1890. So kindly did they take to the conditions they found here, that, atthe end of a long season of protection imposed by law, they fairly swarmedin suitable localities. But what a change a few years of persecution havewrought ! Where formerly a dozen large coveys could be found within a smallarea, only an occasional solitary bird, rarely a pair, is now left of this gemamong our upland birds. The entire blame cannot be laid at the door of the
.sportsman, altho modern rapid-fire guns have played their part. By far theworst havoc has been wrought by the treacherous nets, snares, and traps ofall descriptions, which unscrupulous persons set in defiance of law. Too lazyto hunt, these human vermin catch the poor birds alive and wring their necks.Before close association with mankind had proved so fatal a mistake, thesepartridges were among the tamest and most confiding of birds. Utterly un-suspicious of danger, they would run into the yard and eat with the farmer'shens, paying little attention to any passing human being. When flushed fromtheir haunts in the woods, the whole covey would merely fly into the nearestbushes and trees. Now all is changed, for the " fittest " survivors have inheritedthe knowledge that mankind is their deadliest enemy.He described its haunts as follows:Somewhat inclined to high altitudes, as their name implies, the favoritelocalities for these birds are the large areas in our forests that have beencleared of standing timber. In the course of a year or two these " burns," asthey are called, become over-grown with huckle-berry, salal, and occasionally adense growth of the wild sweet pea. Here is food in abundance at all sea-sons; for in summer the decayed mold of the fallen trees contains grubs andinsects galore as change from fall and winter diet of berries and seeds. Tell-tale hollows in the soft dry earth, sprinkled with a feather or two, speak ofluxurious dust-baths, and a net-work of three-toed tracks in a neighboring wood-road shows where the band has taken its morning constitutional.Nesting.?Major Benclire (1892) gives the following account of thenesting habits of this subspecies:Nidiflcation commences about the middle of May, and ordinarily but onebrood is raised. The nest is placed on the ground, alongside or under anold log, or on side hills under thick bushes and clumps of ferns, occasionallyalong the edges of clearings, grain fields, or meadows. A nest found May 27,1877, near Coquille, Oregon, containing six fresh eggs, was well concealedunder a bunch of tall ferns, in a tract of timber killed by a forest fire.Another, taken in Ukiah Valley, Mendocino County, California, June 2, 1883,by Mr. C. Purdy, contained twelve fresh eggs. This nest was found under
42 BULLETIN" 162, UNITED STATES NATIONAL MUSEUM
a bush of poison oak among a lot of dry leaves on a steep hillside. Theaverage number of eggs laid by this Partridge is about ten, most of the setscontaining from eight to twelve. An occasional nest contains as many assixteen, but such large sets are rare.Eggs.?The eggs of the mountain quail are indistinguishable fromthose of the plumed quail. The measurements of 61 eggs average34.1 by 26.7 millimeters ; the eggs showing the four extremes measure35.6 by 27.7, 33.5 by 27.9, 30.7 by 25.2, and 34 by 25 millimeters.The plumages, food, and general habits of this quail are so muchlike those of the plumed quail that I shall not repeat them here.Louise Kellogg (1916) relates the following incident, which illus-trates the sagacity of the weasel rather than that of the quail
:
On July 8, on north fork of Coffee Creek, the writer caught sight of aweasel in pursuit of a mountain quail. The bird was clucking in a distressedmaimer and evidently leading the enemy away from where her chicks were.When the weasel got her to a safe distance he ran back, jumped over a log,and was seen to make off with a small victim in his mouth. The wholeepisode did not occupy two minutes and occurred in a clearing in broaddaylight. DISTRIBUTIONRange.?The Western United States and Lower California; suc-cessfully introduced at points in Washington, British Columbia,Montana, and Idaho.The range of the mountain quail extends north to northwesternOregon (Astoria) ; southern Washington (Kalama) ; and north-eastern Oregon (Ironside). East to northeastern Oregon (Iron-side and Vale) ; western Nevada (Big Creek, Granite Creek,Truckee, Carson City, and Mount Magruder) ; southeastern California(Willow Creek, Coso, Little Owens Lake, and Thomas Mountain)
;
and Lower California (Laguna Hanson and Mision San Pedro Mar-tir). South to Lower California (Mision San Pedro Martir and Val-ladares). West to Lower California (Valladares, La Grulla, LasCruces, and Los Pozos) ; western California (San Diego, FrazierMountain, Mansfield, Big Creek, Monterey, Camp Meeker, Cahto,Stuarts Fork, and Fort Jones) ; and western Oregon (Port Orford,Coquille, Newport, Netarts, Tillamook, Batterson, and Astoria).The species has been reported as occurring south to Cape St. Lucas,Lower California, but the record is not considered satisfactory.The range as outlined is for the entire species. Oreortyx pictapalmeri is confined to the humid northwestern coast region south tothe coast ranges of Monterey County, Calif. (Big Creek). Oreortyxp. picta occupies the arid and semiarid regions east of the coastranges and south to southern San Diego County (Campo, MountainSpring, Cuyamaca, and Volcan Mountains). A third race, O. p.
PLUMED QUAIL 43
confinis, has been described from the San Pedro Martir Mountains,Lower California.Migration.?A slight vertical migration down the slopes of themountains is performed in the autumn. The journey is made al-most entirely on foot, the birds following railroads, wagon roads,and trails, sometimes passing close to human dwellings. The startfor lower altitudes is made about the first of September, and byOctober 1 the flocks have entirely abandoned those parts of thesummer range above 5,000 feet. The movement has been knownto start as early as August 28 (Webber Lake, Calif.). The returntrip is made early in spring, but exact dates are not available.These birds have been introduced with fair success at points inWashington (Whidbey Island, San Juan Island, and others) ; Brit-ish Columbia (Vancouver Island and Fraser Valley) ; Idaho(Nampa, Silver City, and Shoshone) ; and southeastern and westernMontana. Attempts to acclimatize them in the Eastern States andin New Zealand have been failures.Egg dates.?California (palmeri) : 8 records, March 10 to June20. Oregon: 24 records, May 1 to June 15; 12 records, May 29 toJune 6. Washington and British Columbia: 4 records, June 10to 21.California (picta) : 50 records, April 7 to August 15; 25 records,May 25 to June 20.Lower California (confinis) : 5 records, March 29 to May 28.OREORTYX PICTA PICTA (Douglas)PLUMED QUAILHABITSAs explained under the preceding subspecies, the scientific namesof the two California forms have been changed, but fortunately theEnglish names have shown more stability and will probably standas they always have. The plumed quail of the semiarid interiorranges was formerly called Oreortyx picta plumifera, but will nowstand on our new check list as given above. This is the most widelydistributed and best known form in California, where it is com-monly known as the mountain quail and is so named by Grinnell,Bryant, and Storer (1918).It is the largest and handsomest of the North American quail, isfrequently called " partridge," and has been thought by some toresemble the European partridge, although it is decidedly smaller.It is a shy, retiring species, more often heard than seen. I havehunted for it, where it was common and where it could frequentlybe heard calling, and been favored with only an occasional glimpse
44 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
of one walking stealthily away among the underbrush. It prefersto steal away quietly rather than show itself by flying. All theearlier writers speak of it as uncommon or comparatively rare; itwas doubtless often overlooked because of its secretive habits. It iseven more of a mountain bird than the preceding, ranging up to10,000 feet in summer. W. Leon Dawson (1923) says of its haunts:Save in the extreme northwestern and southeastern portions of its range,the Mountain Quail is to be found in summertime somewhere between 2,000 or3,000 and 9,000 feet elevation, according to local conditions of cover. It inhabitsthe pine chaparral of the lesser and coastal ranges, but its preference is formixed cover, a scattering congeries of buck-brush, wild currant, service berry,Symphori-carpus, or what not, with a few overshadowing oaks or pines. In thenorthwestern portion of its range the bird comes down nearly to sea-level andaccepts dense cover. In the southeastern portion, namely, on the eastern slopesof the desert ranges overlooking the Colorado Desert, the Mountain Quail,according to Mr. Frank Stephens, ventures down and nests at an altitude ofonly 500 feet. It is closely dependent here upon certain mountain springs,which it visits in common with L. c. vallicola and L. gambeli. Under certainconditions, therefore, its breeding range overlaps that of the Valley Quail.There are several instances on record of nests containing eggs of both species,and at least one hybrid has been found, conjectured to be between O. p. confinisand L. c. califoiiiica.Gourtshijy.?On this subject I can find the brief statement fromBendire (1892) that " the mating season begins in the latter part ofMarch and the beginning of April, according to latitude and altitude.The call note of the male is a clear whistle, like whu-ie-whu-ie,usually uttered from an old stump, the top of a rock, or a bush."Grinnell and Storer (1924) write:With the coming of the warm days of late spring, and on into early summer,the males perch on fallen logs, open spaces on the ground, or even on branchesof black oaks, and announce their amatory feelings by giving utterance totheir loud calls with such force and vigor that these resound through theforests for a half-mile or more, commanding the attention of all within hearing.One type of call consists of but a single note, quee-ark, and this is repeatedat rather long and irregular intervals. One bird timed by the watch, June 3,1915, gave his calls at intervals of 7, 6, 8, 5, 8, 6, 7, 5, 7, 9, and 9 seconds,respectively, and continued at about the same rate for a long time afterward.This intermittent utterance lends to the call a distinctiveness and attractivenesswhich would be lost if it were given in quicker time.Nesting.?Of the nest Bendire (1892) says:The nest, simply a slight depression in the ground scratched out by the bird,and lined perhaps with a few dry leaves, pine needles, grasses, and usuallya few feathers lost by the hen while incubating, is sometimes placed alongsidean old log, at other times under low bushes or tufts of weeds, ferns, and,when nesting in the vicinity of a logging camp, a favorite site is under thefallen tops of pine trees that have been left by wood-choppers, the boughs ofwhich afford excellent cover for the nest.Mr. L. Belding found a deserted nest of this species in a cavity of thetrunk of a standing tree near Big Trees, California, but in this locality they nest
PLUMED QUAIL. 45
oftener in thickets of the rock rose or the tar-weed, and according to his obser-vations they do not desert their nests for slight cause, like the Bob White orthe California Quail.H. W. Carriger writes to me:On June 20, 1914, I found four nests of the Mountain Quail. What struckme as unusual was how close the birds remain on the nest; one was undersome brush that a camper had cut the previous year, the leaves were all offand the bunch was a mere handful and, though I stood looking down at thebird, she paid no attention to me and did not get off till I had hit the brushpile several times with a stick. The one with 19 eggs was also a very closesitting bird; I flushed a male and then began a search about, in a radius ofabout 15 feet from where he flew up ; I used a stick and beat all the sur-rounding bushes and vines (mountain misery), but could not flush the female,which I figured must be in the vicinity; after covering all the near-by ground1 sat down near a large tree and accidentally saw a bird move its eye and there,about 6 feet away in a patch of " misery," was a sitting bird ; she allowed meto practically touch her before flying. I am sure that I passed this tree andbeat this bush before, but not a movement from the bird. The third nest waslocated without seeing the bird and was at the base of a small tree. The fourthhad but one egg and was also at the base of a very large tree.Grinnell, Bryant, and Storer (1918) mention two cases wherenests of this species have been found containing eggs of the Cali-fornia, or valle}7 , quail.Chester Barlow (1899) describes as follows some nests found inEldorado County, at an elevation of 3,500 feet in the pine belt:Three nests of the Plumed Quail were found by us, all built in the tar-weedor "mountain misery" (ChamaebaUa foliolosa), and all near paths or roads.The one shown in the illustration was built at the foot of a large cedar tree,and was nicely concealed and shaded by the foliage of the weeds. The nestingcavity was about six inches across and three inches deep, lined with feathersfrom the parent bird. It held ten eggs, in which incubation was well advanced.Several times the bird was flushed in order that we might observe the nest,but she was persistent and always returned. Another nest containing 11incubated eggs was found on the same day, placed amongst the tar-weed in theshade of large cedars. This nesting cavity was about six inches in depth, andcomposed of dry leaves from the tar-weed and lined with feathers. From thenests observed it seems certain that the Plumed Quail makes a nest of its own,for the one last mentioned was substantial enough to bring home.Charles K. Keyes (1905) found six nests of the plumed quail inthe heavily timbered portion of the Sierra Nevada at an altitudeof 3,000 feet. One nest was " protected under the outer edge of amass of deer brush {Ceanothus velutinus) r' ; another was "neatlytucked away along the northwest side of a small boulder and partlyconcealed by dwarf manzanita"; still another "was in rather anopen situation under a Murray pine and five feet away from thetrunk " ; it was " composed entirely of pine needles " and was " par-tially concealed by low sprigs of manzanita." Of the fifth nest hesays:
46 BULLETIN 162, UNITED STATES NATIONAL MUSEUMThe fifth nest was found on June 20 by tramping through deer brush nearthe place where a male had been heard calling for several days. It was thebest concealed of any, being under quite a thick mass of ceanothus, though Ihardly think I should have overlooked it, even though the female had not flushedwith a great whirr of wings when I was three or four feet away from her. Thenest was quite well constructed of coarse dry grass, a few small twigs, and manybreast feathers from the bird. The measurements were the same as those ofthe last nest described and the eggs were twenty-two in number, laid in twolayers, the lower of the nineteen eggs with three on top in the center.Eggs.?The plumed quail does not lay so large sets of eggs as thevalley quail. Probably the average is not more than 10 or 12, butas many as 19 or even 22 have been found in a nest, probably theproduct of two females. In shape they vary from ovate to subpyri-form ; some eggs are quite pointed ; the shell is smooth and somewhatglossy. The color varies from pale cream to a reddish buff, or from
" pinkish buff " to " pale ochraceous-salmon." They are entirely un-spotted. The measurements of 61 eggs average 34.7 by 27 milli-meters; the eggs showing the four extremes measure 38 by 28, 35 by29, and 33 by 25 millimeters.Young.?Bendire (1892) says that incubation lasts about 21 daysand that " in the higher mountains but a single brood is raised ; butin the lower foothills they rear two broods occasionally, the malecaring for the first one while the female is busy hatching the second."Probably both sexes share the duties of incubation. Mrs. Irene G.Wheelock (1904) gives the following account of the hatching process
:
I stole back alone for a last peep at them, and two had pipped the shells whilea third was cuddled down in the split halves of his erstwhile covering. Thedistress of the mother was pitiful, and I had not the heart to torture the beauti-ful creature needlessly ; so going off a little way, I lay down flat along the
" misery," regardless of the discomfort, and awaited developments. Before Icould focus my glasses she was on the nest, her anxious little eyes still regard-ing me suspiciously. In less time than it takes to tell it, the two were out andthe mother cuddled them in her fluffed-out feathers. This was too interestingto be left. Even at the risk of being too late to reach my destination, I mustsee the outcome. Two hours later every egg had hatched and a row of tinyheads poked out from beneath the mother's breast. I started toward her andshe flew almost into my face, so closely did she pass me. Then by many wilesshe tried in vain to coax me to go another way. I was curious and thereforemerciless. Moreover, I had come all the way from the East for just such hoursas this. But once more a surprise awaited me. There was the nest, there werethe broken shells; but where were the young partridges? Only one of all thatten could I find. For so closely did they blend in coloring with the shadowson the pine needles under the leaves of the " misery " that although I knewthey were there, and dared not step for fear of crushing them, I was not sharpenough to discover them.Bendire (1892) writes:I met with a brood of young birds, perhaps a week or ten days old, nearJacksonville, Oregon, on June 17, 1883. The male, which had them in charge,
PLUMED QUAIL 47performed the usual tactics of feigning lameness, and tried his very best todraw my attention away from the young, uttering in the mean time a shrillsound resembling Quaih-quaih, and showed a great deal of distress, seeing Ipaid no attention to him. The young, already handsome and active littlecreatures, scattered promptly in all directions, and the majority were mosteffectually hidden in an instant. As nearly as I was able to judge they num-bered eleven. I caught one, but after examining it turned it loose again.The feathers of the crest already showed very plainly.Dawson (1923) says:Not less uncanny nor less fascinating are the vocal accompaniments withwhich a scattered covey of youngsters is coached or reassembled. If the littleones are of a tender age and the need is great, the parent will fling herself downat your feet and go through the familiar decoy motions; but if the retreathas been more orderly, the parents clamber about, instead, over the rocks andbrush in wild concern. Once out of sight, the old bird says querk querk querkquerk, evidently an assembly call, for the youngsters begin scrambling in thatdirection ; while another old bird, presumably the cock, shouts quee yatok,with an emphasis which is nothing less than ludicrous.Plumages.?In the downy young a broad band of deep " chestnut,"mixed with and bordered by black, extends the whole length of theupper parts, terminating in a point in the middle of the crown;the rest of the upper parts, including the cheeks, are buffy or buffywhite, with large blotches of " chestnut " on the wings, thighs, andflanks and with a dusky line behind the eye; the underparts aregrayish white or yellowish white, palest on the chin.The wings begin to grow almost at once, and the juvenal plumagecomes on very fast, while the chick is still small, 2 or 3 inches long.In this plumage the scapulars, which appear with the wings, are
" clay color " or " cinnamon-buff," peppered, edged, and partiallybarred with brownish black ; the crown and upper back are " hairbrown," the crown barred with dusky and the back mottled withdusky and spotted with white; the crest is brownish black, barredwith brown; the wing coverts, tertials, and tail are pale buff, con-spicuously patterned with black, washed with bright browns onthe wing coverts and tail, and peppered and barred with dusky onthe tertials ; the breast is " light Quaker drab," with white edgings,becoming whitish on the throat and belly, and brownish on thethighs and crissum.The juvenal plumage is worn but a short time before it is replacedby the first winter plumage, which is acquired by a complete molt,except that the outer pairs of primaries are retained for a full year.The time of this molt varies greatly with the date of hatching. Ihave seen birds in this transition stage at various dates from July 10to September 27, and from less than half grown to fully grown.In first winter plumage young birds are practically indistinguish-able from adults, except for the outer juvenal primaries.
48 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMAdults have a very limited prenuptial molt in spring, confined tothe head and neck, and a complete postnuptial molt late in summer.This species has been known to hybridize with the valley quail,where their ranges meet in the foothills.Food.?Concerning the food of the plumed quail, Dr. Sylvester D.Judd (1905) says:Their feeding hours are early in the morning and just before sundown inthe evening, when they go to roost in the thick tops of the scrub live oaks.Their feeding habits are similar to those of the domestic hen. They arevigorous scratchers, and will jump a foot or more from the ground to nip offleaves. This bird is especially fond of the leaves of clover and other legumi-nous plants. It feeds also on flowers, being known to select tbose of Compositaeand blue-eyed grass (Sisyrinchium). Flowers, leaves, buds, and other kinds ofvegetable matter form the 24.08 per cent marked miscellaneous. The birdsprobably eat more fruit than these stomach examinations indicate. LymanBelding says that this quail feeds on service berries, and that during certainseasons it lives almost entirely on grass bulbs (Melica bullosa), which it getsby scratching, for which its large, powerful feet are well adapted. The fruitin its bill of fare includes gooseberries, service berries (Amelanchier alnifolia),and grapes (Vitis californica) . The bird is probably fond also of manzanitaberries, for it is often seen among these shrubs. The food of the mountainquail of the arid regions has been studied in the laboratory of the BiologicalSurvey. The stomachs examined, 23 in number, were collected in California.Five were collected in January, 2 in May, 6 in June, 3 in July, 3 in August,and 6 in November. The food consisted of animal matter, 3 per cent, andvegetable matter, 97 per cent. The animal food was made up of grasshoppers,0.05 per cent; beetles, 0.23 per cent; miscellaneous insects, including ants andlepidopterous pupae, 1.90 per cent; and centipedes and harvest spiders(Phalangidae), 0.82 per cent. The vegetable food consisted of grain, 18.20 percent ; seeds, practically all of weeds or other worthless plants, 46.61 per cent
;
fruit, 8.11 per cent; and miscellaneous vegetable matter, 24.08 per cent. Thegrain included wheat, corn, barley, and oats. The legume seeds include seedsof alfalfa, cassia, bush clover, vetch, and lupine. The miscellaneous seeds comefrom wild carrot {Daucus carota), tar-weed (Madia saliva), Collomia sp.,Anmnckia sp., labiate plants, dwarf oak, snowbush {Ceanothus cordulatus)
,
and thistle.Behavior.?H. W. Henshaw (1874) writes:It seems nowhere to be an abundant species. * * * The bevies are verysmall, and I do not remember to have ever seen more than fifteen together, of-tener less. It is a wild, timid bird, haunting the thick chaparral-thickets, andrarely coming into the opening. When a band is surprised they are not easilyforced on the wing, but will endeavor to find safety by running and taking refugein the thickness and impenetrability of their favorite thickets. If forced, how-ever, tbey rise vigorously and fly swiftly and well, and sometimes to a con-siderable distance, and then make good their escape by running. During theheat of midday, they will be found reposing under the thick shade of thechaparral, and there they remain till the cooler hours invite them to continuetheir quest for food.
PLUMED QUAIL 49Grinnell, Bryant, and Storer (1918) say:When alarmed the Mountain Quail carries its crest feathers erect, bowingbackwards towards the tip but not tilted forward as in the case of the ValleyQuail. This action gives the bird an alert attitude?consistent with its evidentanxiety in case there are young about. Although habitually occupying brushyand forested areas, this quail but seldom perches in trees, and as far as weknow the adults never roost in one at night. They stick close to the groundand usually seek safety by running beneath cover rather than by flight. Forthis reason the Mountain Quail is considered an unsatisfactory bird to hunt.When hunted in the brush they generally run some distance before flying, scat-tering and finally taking wing like as not behind a bush so as to preclude theprobability of a successful shot.Voice.?Dawson (1923) describes the varied notes of this quailvery well, as follows
:
The Mountain Quail's is the authentic voice of the foothills, as well as thedominant note of Sierran valleys and of bush-covered ridges. Spring andsummer alike, and sometimes in early autumn, one may hear that brooding,mellow, slightly melancholy too' wook, sounding forth at intervals of five orsix seconds. Now and then it is repeated from a distant hillside where a rivalis sounding. This note is easily whistled, and a little practice will enablethe bird-student to join in, or else to start a rivalry where all has beensilent before. And quite as frequently, in springtime, a sharper note issounded, although this, I believe, is strictly a mating or a questing call, queelkor queelp. This has alike a liquid and a penetrating quality which defies imita-tion, so that the unfeathered suitor is not likely to get very far in milady'saffections. Thus, also, I have " witnessed " the progress of courtship and itsimpending climax in the depths of a bed of ceanothus where not a featherwas visible. The quilk of the preceding days had evidently taken effect. Thelady was there, somewhere. The mate was still quilking, but his efforts werehurried, breathless. Between the major utterances, ecstatic took notes wereinterjected. As the argument progressed I heard a low-pitched musical series,rapidly uttered, look look look look look. (But there was no use in looking.)This series, employed six or eight times, was suddenly terminated by half adozen quilks in swift succession, indicative of an indescribable degree ofexcitement.Leslie L. Haskin writes to me
:
Their call when quietly feeding is not greatly unlike that of young turkeypoults when following their mother. Another call often given is a simple,rapid, chirring thrill, t-r-r-r-r-r-r-r-r, often long continued. Their alarm notewhen startled but not badly frightened is an exact reproduction in accent,though not in tone, of a hen's cackle. In the partridge the cut-cut-ca-do of thebarnyard fowl is charmingly altered into a shrill t-t-t-t-t-tr-r-r-r-r-r-t ortut-tut-tut-tr-r-r-r-r-tut all very rapidly delivered, and in sharp crescendo. Be-cause of the difference in tone, and the rapidity of the partridges' delivery, fewhave noticed this resemblance, but once the ear has grasped the accent thesimilarity can never be forgotten.Game.?Although the mountain partridge is a fine, large, plumpbird and makes a delicious morsel on the table, it is not highly re-garded as a game bird and not fully appreciated by sportsmen. It is
50 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
a difficult bird to hunt in the dense mountain thickets it frequents.Edwyn Sandys (1904) says:This comparatively large and exceedingly handsome species is not highlyesteemed by sportsmen in general, owing to its true value not being well un-derstood. In certain portions of California, and notably in the WillametteValley, Oregon, when abundant it affords capital sport, while upon the tableit is a delicacy not to be forgotten. As a rule, one, or at most two, broods arefound on a favorite ground, the birds seldom, if ever, flocking like some oftheir relatives. 0. pictus prefers moist districts and a generous rainfall. Itis a runner, and in comparison with Bob-white, by no means so satisfactory abird for dogs to work on. After the first flush the covey is apt to scatterwidely and the beating up of single birds is a slow and frequently a wearyingtask. On the wing, its size and moderate speed render it a rather easy mark.Grinnell, Bryant, and Storer (1918) say:Its flesh is excellent, being declared juicier and more finely flavored thanthat of the Valley Quail. But its comparatively small numbers, even undernormal conditions, the difficulty attendant upon reaching its habitat, and thefact that it does not lie well to dogs, deter many sportsmen from hunting thespecies. Except when the birds may be out of their natural habitat, as duringtheir fall migration, it takes stiff, hard climbing and a deal of patience toget a limit of ten. In former years Mountain Quail were commonly sold onthe markets of San Francisco. In some instances they were trapped alongthe western flanks of the Sierras and sent to the markets alive. Mr. A. E.Skelton, of El Portal, has reported to us that while shooting for the marketnear Raymond, Madera County, many years ago he averaged about a dozenand a half Mountain Quail a day. The birds then brought from $2.50 to$4.00 a dozen. At the present time it is illegal to sell quail of any sort,except for propagation and then under permit only.Enemies.?Predatory animals and birds help to account for thehigh rate of mortality among the young birds, so that, in spite of thelarge broods hatched, only a few ever reach maturity. Wildcatsand gray foxes seem to be their greatest enemies. These animals arealso sufficiently agile to capture the old birds as well, for bunchesof their feathers are often found.Fall.?From the nature of its summer haunts in the mountains,which must be abandoned when the winter snow comes, the plumedquail has developed an interesting migratory habit. Barlow andPrice (1901) say:By the first of September the quail are restless and are beginning theirpeculiar vertical migration to the west slope of the mountains. Sometimesfour to six adults with their young will form a covey of ten to thirty individualsand pursue their way, almost wholly on foot along the ridges to a more con-genial winter climate. By October 1 the quail have almost abandoned the ele-vations above 5,000 feet. In the fall the woodland is full of the disconsolatepeeps and whistling call notes of the young who have strayed from theircoveys.Lyman Belding (1903) adds:The fall migration of the mountain quail (Oreortyx pictus plumiferus)appears to be influenced but little by the food supply or temperature in its
SAN PEDRO QUAIL 51summer habitat in the Sierras, which it appears to leave because the propertime has arrived for its annual tramp down the west slope. The first flocksstart about the first of September, or sometimes two or three days sooner.At Webber Lake after three cold cloudy days, they began to move westwardAugust 28, 1900. When they are migrating their whistle is frequently heard,and they do not seek cover for protection but follow a wagon road, railroad,travel in snow sheds, pass near dwelling, and seem to care but little forself preservation.Winter.?Belding says further (1892) :The mountain quail (Oreortyx pictus plumiferus) , which are so plentiful inthe high mountains in summer, are only summer residents there. They usu-ally spend the winter below the snow line, but as it is not possible to tell justwhere that is, or rather where it is going to be, they are sometimes caughtin snow storms, but I have been astonished at the correctness of their apparentforecast of different winters. A few birds winter high in the mountains, butI think they are parts of flocks which were nearly annihilated, or young birdswhich got scattered and lost, and a few that were wounded and survived.OREORTYX PICTA CONFINIS AnthonySAN PEDRO QUAILHABITSIn the mountain ranges of southern California and northernLower California a grayer race of the mountain quail occurs. Thisrace was discovered and named by A. W. Anthony (1889) fromspecimens collected in the San Pedro Martir Mountains in LowerCalifornia, which he describes as " differing from Oreortyx pictaplumifera in grayer upper parts and thicker bill."He says of its haunts and habits
:
From an elevation of 6,000 to 10,000 feet above the sea, in the San PedroMountains, I found this quail abundant, occurring wherever water and timberafforded it drink and shelter, and only leaving the higher elevations when thefrosts of winter make life in the lower valleys desirable. A few pairs bredabout my camp at Valladores, 6 miles from the base of the range and 2,500feet above the sea ; but nearly all of the flocks that wintered along the creek atthis point were gone in March, leaving only an occasional pair, which sought theshelter of the manzanitas high tip on the hill-sides, from whence their clear,mellow notes were heard morning and evening, so suggestive of cool brooks andrustling pines, but so out of place in the hot, barren hills of that region.CALLIPEPLA SQUAMATA PALLIDA BrewsterARIZONA SCALED QUAILHABITSThe Mexican plateau, with its elevated and arid, or semiarid,plains, extends northward into southern Arizona, New Mexico, andwestern Texas. Much of this region is dry and barren, except for ascattered growth of creosote bushes, dwarf sagebush, stunted mesquite,
52 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
catclaw, mimosa, various cactuses, and a few yuccas. In the mesasbetween the mountain ranges and at the mouths of canyons, whereunderground streams supply some scanty moisture, there are grassyplains, with a scattered row of sycamores or cottonwoods marking thecourse of an unseen stream. Such are the haunts of the scaled quail,blue quail, or white topknot, as it is appropriately called in Arizona.Here, as one drives along over the winding desert trails, dodging thethorny shrubs or still more forbidding clumps of cactus, he may sur-prise a pair or perhaps a little bevy of these birds, invisible at firstin their somber gray dress, which matches their surroundings so well.They do not attempt to escape by flight, but scatter in different direc-tions, running with remarkable speed, with their necks upstretchedand their white crests erected, dodging in and out among the desertvegetation, like so many rabbits scurrying off to the nearest brierpatch. They are soon lost to sight, for they can run faster than wecan and will not flush.George Finlay Simmons (1925) says that, in Texas, it "shunstimbered country," but is " characteristic of the barren plateaus inthe mountainous districts of western Texas, usually where the soil isfine, loose, and sandy; broad, dry, arid washes, gulches, and semi-barren plateaus of the hills where hard ground is covered with a fewthorny bushes, scattered scrub oak, chaparral, mesquite, sagebrush,and different species of cactus; chaparral and mesquite country, gen-erally in the vicinity of water, but sometimes miles from any streamor pond."Mrs. Florence Merriam Bailey (1928) says that in parts of NewMexico scaled quail collect about the ranches and huts of the settlers,picking up grain left from feeding the horses, or feeding with thechickens.Frank C. Willard says in his notes
:
Of the three species of quail found in Cochise County, Ariz., the scaled quailis the one most commonly met with on the dry, brush-covered mesas and valleys.Here it frequents the dry washes, or arroyos, with their fringe of mesquites,small desert willows, and an occasional flat area covered with bear grass. Thenear proximity of living water does not seem to be at all necessary for theirexistence, as I have frequently found them 7 or S miles from any water at all.Such of these quail as do live in the vicinity of water make regular daily tripsto it and they congregate more thickly around ranches and water holes than theydo away from them.Along the San Pedro River and the Barbacomari River (a branch of theformer), there is a mingling of the scaled quail and Gambel's quail. These twospecies are also found inhabiting the foothills of the mountains, and the lowranges of hills like those around Tombstone and in the Sulphur Spring Valley.In such localities the two quails occasionally lay their eggs in the same nest.Nesting.?While going to and from our camp in Ramsey Canyon,in the Huachuca Mountains, Ariz., we frequently saw two or three
ARIZONA SCALED QUAIL 53pairs of scaled quail on a grassy flat where a few sycamores and smallbushes were scattered along the course of a dry wash, the bed of anunderground stream running out from the canyon. On May 25, 1922,while hunting this flat we flushed a quail from the only nest of thisspecies that I have ever seen. The nest was well concealed under atuft of grass surrounding a tiny mesquite ; the grass was well archedover it, and the hollow in the ground was lined with dry grass and afew feathers. It held 14 fresh eggs.Nests found by others have been similarly located under the shel-ter of some low bush, sagebush, creosote bush, mesquite, catclaw,cactus, or yucca, rarely in an open situation among rocks or undera fallen bush. Simmons (1925) says that in Texas the nest is rarelyplaced in a meadow or grainfield. The hollow is lined with what-ever kind of dry grass is available. Willard says in his notes
:
Most of the nesting occurs during the months of June and July. I am in-clined to believe that this is because the rainy season in Arizona commences,under normal conditions, early in June. Thereafter there are more or lessheavy showers nearly every day. This assures a supply of drinking waterwithin easy reach of the newly hatched young. The nests are usually placedunder some tussock of mixed dry and green grass. In the vicinity of gardens,they sometimes build under tomato vines. Where a haystack is available, theyare quite likely to work out a hollow near the bottom and lay their eggs theremuch after the manner of the domestic hen. It is not at all unusual for twoscaled quail to lay their eggs in the same nest, if the presence of two distincttypes of eggs in the same nest can be considered as evidence. In several in-stances I have had nests under observation (which did not yet hold completeclutches) and in three of these instances eggs were deposited at the rate of twoper day, quite positive proof that two birds were using the same nest.Eggs.?The scaled quail lays from 9 to 16 eggs, rarely more, andusually from 12 to 14. They are ovate or short ovate in shape andusually quite pointed. The shell is thick and smooth, with little ifany gloss. The ground color varies from dull white to creamywhite. Some few eggs are thickly, or even heavily speckled withvery small spots or minute dots of dull, light browns, " sayal brown "to pale " cinnamon-buff." Most of the eggs are sparingly markedwith similar spots. Some are nearly or quite immaculate. MajorBendire (1892) says that "occasionally a set is marked with some-what more irregular, as well as larger, spots or blotches, resemblingcertain types of eggs of Callipepla ga??ibeli, but these markings arealways paler colored and not so pronounced." The measurementsof 57 eggs average 32.6 by 25.2 millimeters; the eggs showing thefour extremes measure 35.8 by 26, 34 by 27, 30 by 24.5, and 31.5 by23.5 millimeters.Young.?Major Bendire (1892) believed that "two and even threebroods are occasionally raised in a season, the male assisting in the74564?32 5
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care of the young, but not in incubation. This lasts about 21 days."Mrs. Bailey (1928) writes:That the downy young are also obliteratively colored is well illustrated byan experience of Major Goldman when climbing the Florida Mountains. At5,300 feet, among the oaks and junipers, he reports, " I came suddenly on anadult bird and a brood of recently hatched young. The old bird disappearedafter giving several sharp cries of alarm, and the young also disappearedin an open patch of short grass. On reaching the place I began looking aboutcarefully and soon saw one young bird flattened down, with not only its littlebody but its head and neck also pressed close against the ground, its downyplumage blending in well with the color of the ground and the dead grass stems."There it lay, pressed close to the ground until approached within three feet when,
" it suddenly started up with sharp peeping cries, and the entire brood whichhad scattered and hidden in an area about fifteen feet across, half ran, half flewinto some thick bushes where they were more securely hidden."Plumages.?In the downy young scaled quail the forehead, thefront half of the crown, in front of a little gray topknot, and thesides of the head are " cinnamon-buff " or " pinkish buff " ; there is abroad band of " chestnut " from the middle of the crown, back ofthe topknot, down to the hind neck, bordered narrowly with blackand with broad stripes of buffy white; the auricular spots are dark
" chestnut " ; the chin and throat are buffy white and the rest ofthe underparts are pale grayish buff; the back is mottled with palebuff and " russet."The juvenile plumage starts to grow at an early age, beginningwith the wings; on a small downy chick, less than 2 inches long,the wings are well sprouted; the wings grow so fast that the youngcan fly long before they are half grown. The sexes are alike inthe juvenal plumage. In this the crown is " buffy brown " to " woodbrown " and the crest is " vinaceous-buff " ; the rest of the head, neck,and shoulders shades off gradually to shades of drab; the feathersof the back, scapulars, and wing coverts are from drab to "sayalbrown " or " tawny-olive," barred with " sepia " or brownish black,finely sprinkled or peppered with brownish black, with conspicuousmedian stripes of buffy white and with buffy edgings on the scapu-lars ; the tail is mainly dark drab, but more buffy near the tip, barredand peppered with brownish black; the underparts are buffy whiteor grayish white, spotted or barred with dusky, most distinctly onthe breast, where many feathers are tipped with white arrowheads.The molt into the first winter plumage takes place during Sep-tember and October. This is a complete molt, except for the twoouter primaries on each wing, which are retained all through thefirst year. The molt begins on the back, breast, and flanks. Youngbirds are practically indistinguishable from adults during the firstwinter and spring except for the retained outer primaries. Bothfirst year and adult birds have a partial prenuptial molt early in
ARIZONA SCALED QUAIL 55
spring, restricted mainly to the head and throat, and a complete post-nuptial molt in August and September. Albinism occurs occasion-ally, and this species has been known to hybridize with Gambel'squail and with the bobwhite.Food.?Mrs. Bailey (1928) sums up the food of this species verywell, as follows:The Scaled Quail apparently eats more insect food than any of the otherquails, or more than 29 per cent, as against 70 per cent of vegetable matter.Of this vegetable matter over 50 per cent is weed seeds, among which arethistle, pigweed, and bindweed, a troublesome weed that often throttles otherplants. Dasylirion seeds almost entirely filled six stomachs examined. Wildfruit, such as prickly pear and the succulent parts of desert plants, togetherwith its larger per cent of insect food, doubtless help it to live with a minimumamount of water. Its insect food includes grasshoppers, ants, and beetles
?
among them leaf chafers and cucumber beetles?weevils, such as the cloverpest and scale insects (several hundred in one stomach) that feed on theroots of plants.Sylvester D. Judd (1905) says of its vegetable food:The species resembles the ruffed grouse in its habit of feeding on greenleaves and tender shoots. It feeds upon budded twigs, but more often limitsits choice to chlorophyll-bearing tissue, often picking green seed pods ofvarious plants. Like domestic fowls, it eats grass blades. Fruit was eatenby only 6 of the 47 birds, and none was taken from cultivated varieties. Asmight be expected from inhabitants of arid plains, these birds like the fruitof cacti, and have been found feeding on the prickly pear {Opuntia lindheimeri)
.
The fruit of Ibervillea lindheimeri also is eaten. The blue berries of Adeliaangustifolia, which furnish many desert birds and mammals with food, areoften eaten by the scaled quail. Different kinds of Ruius fruits are relished,and the berries of Koeberlinia spinosa and Monisia pallida also are eaten. Thefruit and succulent parts of plants no doubt serve in part in the parcheddesert as a substitute for water.Behavior.?The scaled quail is a decidedly terrestrial bird withvery powerful legs, which it uses to advantage in the rather opendesert growth in which it lives and where it can run very fast inthe smooth open spaces among the desert plants. It prefers toescape by running rather than flying; but, if come upon suddenlyand surprised, it rises with a whir of wings, flies a short distance,and scales down into cover again, much after the manner of thebobwhite ; it then starts running and can not be easily flushed again.If in a flock, they sometimes follow a leader in Indian file, but moreoften they scatter in several directions and are soon lost to sight.Major Bendire (1892) quotes Dr. E. W. Nelson, as follows:In many instances I have found them far from water, but they make regularvisits to the watering places. On the Jornada del Muerto and on Santa FeCreek I found them frequenting the open plains, away from the water inthe middle of the day, and in the vicinity of the water late in the afternoon.At this time they are often seen in company with Gambel's Quail amongstthe bushes and coarse grass or weeds bordering the water courses.
56 BULLETIN 162, UNITED STATES NATIONAL MUSEUMBendire (1892) quotes William Lloyd as saying:The Blue Quail loves a sandy table land, where they spend considerable timein taking sand baths. I have often watched them doing so, pecking andchasing each other like a brood of young chickens. Good clear water is anecessity to them. They are local, but travel at least 3 miles for water.In the evenings they retire to the smaller ridges or hillocks and their callsare heard on all sides as the scattered covey collects. Several times I haveseen packs numbering sixty to eighty, but coveys from twenty-five to thirty aremuch oftener noticed. During the middle of the day they frequently alight intrees, usually large oaks, but they roost on the ground at night.Voice.?Major Bendire (1892) says:According to Mr. Lloyd their call note sounds something like a lengthenedchip-ohurr, chip-churr; the same, only more rapidly repeated, is also givenwhen alarmed, and a guttural oom-oom-oom is uttered when worried or chasedby a Hawk. The young utter a plaintive peep-peep, very much like youngchickens. Like the rest of the partridge tribe they are able to run about assoon as hatched.Mr. Simmons (1925) refers to their notes as: "A single low, long-drawn whistle ; a nasal, musical, friendly pe-cos', pe-cos'."Enemies.?Mrs. Bailey (1928) writes:Although protective coloration and attitudes partly serve their purposes,protective cover is still vitally important, for as Mr. Ligon has found, " PrairieFalcons, Cooper Hawks, Roadrunners, snakes, skunks, wildcats, and coyotes alltake their toll of these birds or their eggs" ; in the northern part of their range,Magpies destroy both eggs and young; and over much of their range hail,cold rains, and winter storms deplete their numbers.Mr. Willard says in his notes
:
The Gila monster, rattlesnake, and skunk are natural enemies which takea large toll from the nests of the scaled quail. I once observed a femalequail fluttering excitedly over a clump of grass and making dashes downat it. On investigating I found a rattlesnake and nine quail eggs in thenest. I dispatched the snake and on opening it found three whole eggs inside.A Gila monster, which I caught and caged, evidently disgorged two scaledquail eggs, as there were two eggs in the box a short time later, and I amsure no one had been near it but myself. In passing, it may be of interestto say that this great lizard will devour a hen's egg by gradually workingit far enough into its mouth to be able to clamp down on it with its powerfuljaws, crushing it, and then sucking out the contents. They are large enoughto swallow easily a quail's egg whole. We occasionally found a mass of loosefeathers of this quail scattered on the ground and clinging to near-by bushes.The presence of cat tracks told what was responsible for the tragedy thefeathers betrayed. On at least four occasions I have surprised a long-leggedMexican lynx stalking the same game I was after, and was able to collect acat as well as a quail.Fall?Mrs. Bailey (1928) says further:The entire life of the Scaled Quail is spent in the environment to whichit is so well adapted, but in the fall it is sometimes found a few hundredfeet higher than in the nesting season. When the young are raised these
ARIZONA SCALED QUAIL 57delightful little Cotton-tops go about in small flocks, visiting water holes andriver bottoms. Picking up insects, seeds, and berries as they go, they wanderthrough brushy arroyas, over juniper-clad foothills, cactus flats, and sage-brush or mesquite plains, calling to each other with a nasal pay-cos, pay-cos,which by long association comes to take on the charm attaching both to thegentle-eyed birds themselves and to the fascinating arid land in which theymake their homes.Game?Hunting the blue quail will never figure as one of themajor sports, although it is a gamy bird and makes a delicious andplump morsel for the table. The birds are widely scattered over avast expanse of rough country, on desert plains covered with thornyunderbrush, or on stony or rocky foothills where walking is difficultand slow. The hunter must be prepared to do some long, hardtramping, for he is more likely to count the number of miles to a birdthan the number of birds to a mile. A dog is useless, for these quailhave not yet acquired the habit of lying to a dog. Eastern quailhave learned to lie close, a good way to hide from human enemiesbut a very poor way to escape from the many predatory animals inthe West. Scaled quail are shier than Gambel's quail and are gen-erally first seen in the distance running rapidly and dodging aroundamong the bushes. They run faster than a man can walk, and thehunter must make fast progress over the rough ground to catch upwith them. By the time he gets within range he will be nearly outof breath and will have to take a quick snap shot at a fleeting glimpseof a small gray bird dodging between bushes. This is far moredifficult, under the circumstances, than wing shooting and can not beconsidered pot shooting. Sometimes, when a large covey has beenscattered and rattled, the hunter may surprise single birds and getan occasional wing shot; but they are apt to jump from most unex-pected places, ahead of or behind the hunter, and give him a difficultshot. Late in the season they are often found in large packs of100 or 200 birds, when the chances for good sport are better. Eventhen the hunter may well feel proud of a hard-earned bag.DISTRIBUTIONRange.?The Southwestern United States and northern Mexico.Nonmigratory.The scaled quail is found north, to southern Arizona (Picacho, Rice,and Clifton) ; northern New Mexico (Haynes and the Taos Moun-tains) ; east-central Colorado (Mattison and Holly) ; and northeast-ern Texas (Lipscomb). East to Texas (Lipscomb, Mobeetie, Colo-rado, San Angelo, Fredericksburg, San Diego, Fulfurrias, andBrownsville) ; Tamaulipas (San Fernando) ; and San Luis Potosi(Ahualulco). South to San Luis Potosi (Ahualulco and Ramos);
58 BULLETIN" 162, UNITED STATES NATIONAL MUSEUMDurango (Kancho Baillon) ; Chihuahua (San Diego) ; and Sonora(San Pedro and Sesabe). West to Sonora (Sesabe) ; and southernArizona (Arrivaca, Sierrieta Mountains, and Picacho). It has beendetected casually in eastern Texas (Gainesville and Bonham). OnAugust 19, 1926, three specimens were collected at Elkhart, MortonCounty, Kans. It is a common species across the State line in south-eastern Colorado.The range above outlined is for the entire species. By recognitionof the Arizona, New Mexico, and Colorado bird (C. s. pallida), thetypical subspecies (C. s. squamata) is restricted to northwestern Mex-ico. Another race, the chestnut-bellied scaled quail (C. s. castano-gastris), inhabits the lower Rio Grande Valley in Texas and adjacentregions in northeastern Mexico.Attempts to transplant the scaled quail to other regions have gen-erally resulted in failure. Among these may be mentioned introduc-tions in Louisiana, Florida, Georgia, and Washington. So far as isknown, the only successful transplantation was made in Colorado atColorado Springs and probably also at Canyon City. From thesepoints the birds have spread and increased until they are now com-mon in the Arkansas Valley from Pueblo east to the State line, andit appears that the introduced stock has met and blended with thenative birds working northward along the Las Animas River.Egg dates.?Texas {'pallida) : 11 records, May 7 to June 22. Ari-zona and New Mexico: 37 records, April 16 to September 22; 19records, June 11 to July 7.Texas and Mexico (castanogastris) : 44 records, March 7 to June28 ; 22 records, May 3 to June 2.CALLIPEPLA SQUAMATA CASTANOGASTRIS BrewsterCHESTNUT-BELLIED SCALED QUAILHABITSThe scaled quail of the lower Rio Grande Valley in Texas andeastern Mexico is more richly and darker colored than the quail foundfarther west, and, as its name implies, it has a well-marked patch ofdark chestnut in the center of its belly, which is more prominent inthe male than in the female. George B. Sennett (1879) thus de-scribes its habitat
:
The foothills of the Rio Grande, about 100 miles back from the coast, are theeastern limits of this bird, as well as of the Cactus Wren and the Yellow-headedTitmouse. The first rise of ground in going up the river occurs at LomitaRanch, and here we often saw these beautiful birds running about ; but althoughwe frequently collected a mile or two below the hill, there we never saw them,and not even in the fertile and heavily wooded lowlands in the vicinity of thishill did we observe them. A few miles up from Lomita and back from the river,
CALIFORNIA QUAIL 59
near the water-holes, rises are numerous, covered with thin, poor soil, wherecactuses and scrubby, thorny bushes grow, and here the blue quail abounds.Nesting.?Sennett evidently found only one nest, of which he says
:
On the 22d of May, near the buildings of the ranch at San Jose Lake, Mr.Sanford shot a fine male, which was on the brush fence forming the enclosure.In searching among the weeds where the bird fell, we found a nest and 16 fresheggs. The nest was under the edge of the fence, and was simply a saucer-likedepression in the ground, with leaves for lining.Three sets of eggs in my collection were taken from hollows in theground, under cactus plants or bushes, lined with grass, weeds, ortrash. Major Bendire (1892) says that " their nests are always placedon the ground; a slight hollow in the sand is scratched out by thebird, usually under a clump of weeds or grass, or a prickly-pearbush. They are very slightly lined with dry grasses."Eggs.?The eggs of the chestnut-bellied scaled quail are practicallyindistinguishable from those of the Arizona form, though they mayaverage a little more richly colored or more heavily spotted and atrifle smaller. Major Bendire (1892) says:Full sets of eggs have been taken near Rio Grande City, and at Camargoon the Mexican side of the river opposite, as early as March 11, and fromthat time up to July 10. Two broods are unquestionably raised in a season.Mr. Thomas H. Jackson, of West Chester, Pennsylvania, gives the averagenumber of eggs laid by this species as fifteen, based on data taken fromtwenty-seven sets. The largest number found in one nest was twenty-three.The measurements of 77 eggs in the United States NationalMuseum average 31 by 24 millimeters; the eggs showing the fourextremes measure 34 by 24.5, 33 by 25.5, 28.5 by 23.5, and 30 by 22millimeters.Behavior.?In its general habits this quail does not differ ma-terially from its western relative. Its plumage changes, its food,voice, and behavior are all similar. Both forms are resident in theirrespective ranges, moving about only as the food supply demands.LOPHORTYX CALIFORNICA CALIFORNICA (Shaw)CALIFORNIA QUAILHABITSThe type race of this species originally inhabited the narrow stripof humid coast region from southwestern Oregon south to south-ern Monterey County, Calif. It has been introduced on VancouverIsland and in Washington, where it has become well established.It differs from the more widely known valley quail in having theupper parts olive-brown, rather than grayish brown, and the innermargins of the tertials deeper buff. It does not differ materially
60 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMfrom the other races in its nesting, food, or general habits. A fullaccount of the habits of the species is given under L. c. vallicola,the next form treated. J. Hooper Bowles (Dawson and Bowles,1909) writes:This bird and its near relative, the Valley Partridge, are not natives ofWashington ; but, like the Mountain and the Plumed Partridge, were introducedhere from the State of California. Dr. Suckley, one of our pioneer naturalists,tells us that as early as 1857 two shipments of birds were turned out in thevicinity of Puget Sound by Gov. Charles H. Mason and a Mr. Goldsborough.Conditions seem to have proved most suitable for them, since, in the faceof constant persecution, they continued to increase in numbers, spreadingtheir ranks over new territory every year. Although ofteu found in dry,bushy uplands, they are much more inclined to damp localities than theMountain Partridge, their favorite haunts being the low ground of the rivervalleys. Here they may be found searching for seeds in the weed-patchesof the open fields, or gleaning amongst the growing cabbages, beans, andother vegetables of the farmer's garden. Indeed, few birds are so much thefriends of the farmer as our partridges, for their food consists almost en-tirely of weed-seeds, worms, beetles, grass-hoppers, and other insects. Whatlittle of the newly-sown crops they may eat is repaid a thousand fold bythe vast amount of good they accomplish.Nesting.?Mr. Bowles, in his notes sent to me, says
:
Like most species of introduced game birds these quail lay their eggs in thenests of other varieties of birds. I have in my collection a set of nine eggs ofthe sooty grouse and three eggs of this quail, personally taken here at Tacoma.All the eggs were heavily and evenly incubated.I have also a nest of the Nuttall's sparrow containing four eggs of the sparrowand two eggs of this quail, which was taken near the city of Seattle, Wash., onMay 8, 1918. Incubation was slight. This set was collected and presented tome by D. E. Brown, of Seattle. His notes say that one of the quail eggs wason end in the nest, the other on top of two of the sparrow eggs. The sparrowwas on the nest and showed much anxiety.Eggs.?The eggs of this quail are indistinguishable from those ofthe valley quail, but they average slightly larger. The measurementsof 60 eggs average 32 by 25 millimeters; the eggs showing the fourextremes measure 35.5 by 24.5, 35 by 26, 30 by 24, and 31 by 23millimeters. DISTRIBUTIONRange.?Southwestern Oregon, California, extreme western Neva-da, and Lower California. Successfully introduced in Washington,Idaho, Utah, Nevada, and British Columbia, as well as Hawaii, NewZealand, Chile, and probably locally in France.Because of the many attempts to extend the range of this speciesin the Western States, it is difficult to outline the area to which theyare indigenous. It appears, however, that the natural range extendsnorth to southwestern Oregon (Anchor and Algoma). East toOregon (Algoma and Klamath Falls) ; western Nevada (Anaho
OALIFOKNIA QUAIL 61Island and Stillwater) ; south-central California (Fresno, Tehachapi,and San Bernardino) ; and southeastern Lower California (Pin-chalinque Bay, Triunfo, and San Jose del Cabo). South, to southernLower California (San Jose del Cabo and Cape St. Lucas). West toLower California (Cape St. Lucas, San Javier, Rosarito, San Andres,San Quintin Bay, and Los Coronados Islands) ; western California(San Diego, Santa Catalina Island, Santa Barbara, Monterey, Wat-sonville, Alameda, San Francisco, Marysville, Chico, and Baird)
;
and southwestern Oregon (Grants Pass and Anchor).The range as above outlined is for the entire species. Truecalifomica is restricted to the humid coast region from southwesternOregon south to Monterey County, Calif. The valley quail (L. c.vallicola) occupies the rest of the range south to the northwesterncorner of Lower California (about latitude 32? N.), including LosCoronados Islands. The birds found on Catalina Island have beendescribed as a distinct race, LopJiortyx c. catalinensis.In Lower California, in addition to vallicola, which occurs in thenorthwestern part, the species has been divided into two subspecies.The San Quintin quail (L. c. plumbea) is distributed over most ofthe territory between latitudes 30? and 32? N., while the San Lucasquail (Z. c. aclirusteva) is found from latitude 30? N. south to CapeSan Lucas. There is more or less intergradation in the areas wherethese races meet.As previously indicated, the California quail has been a favoritein attempts at transplantation. The birds on Los Coronados Islandsare considered by some to be introduced, although there also is evi-dence that they fly back and forth to the mainland. They have beensuccessfully introduced into Nevada (Virginia City, McDermitt,Quinn River Valley, Paradise Valley, Lovelock, and probably alsoCarson City and Reno) ; Oregon (Willamette Valley and Jacksonand Josephine Counties) ; Utah (Salt Lake County and Ogden)
;
Washington (Olympia, Garfield County, Walla Walla County, Ya-kima County, and probably many other points, as they are now welldistributed over the western part of the State, including the islandsin Puget Sound and Bellingham Bay) ; British Columbia (Van-couver Island, Denman Island, and the Okanagan Valley). Attemptsto introduce this species in Illinois, Maryland, Massachusetts, NewYork, Delaware, Mississippi, and Missouri have not been successful,although in a few instances the birds seemed to thrive during thefirst season.California quail also have been successfully transplanted to theHawaiian Islands (Hawaii, Maui, and Molokai), New Zealand, andChile. Other foreign experiments apparently have failed.
62 BULLETIN 162, UNITED STATES NATIONAL MUSEUMEgg dates.?California (californica) : 92 records, January 12 toJuly 21 ; 46 records, May 8 to June 8. Washington and British Co-lumbia : 10 records, May 11 to July 3.California (vallicola) : 125 records, February 9 to October 29; 63records, May 1 to June 5.LOPHORTYX CALIFORNICA VALLICOLA (Ridgway)VALLEY QUAILHABITSThe valley quail being the most widely distributed form, it shallhave the most complete life history of the species, as it is the bestknown of the various subspecies. In Pasadena and vicinity, southernCalifornia, it is a common dooryard bird, coming regularly into thecity to feed on the lawns and to roost in the trees and shrubbery. OnDr. Louis B. Bishop's lawn, in the thickly settled part of Pasadena,one might see from 10 to 20 of the pretty birds almost any afternoonafter 4 o'clock. Although rigidly protected and regularly fed, theyseemed very nervous and shy ; if they saw us moving, even at a win-dow, they would run or fly into the shrubbery. J. Eugene Law hasa flock of 100 to 200 birds, which he feeds every morning duringwinter on his driveway in Altadena. I was able to photograph someof these birds one morning from a blind, but I found them verynervous ; at the slightest noise or movement they would all fly off butwould soon return. Mr. Law told me that these quail all bred in thevicinity, nesting commonly in the old abandoned vineyards over-grown with rank grasses and weeds. They travel around in flocksduring winter but begin to break up into pairs during March. Thelatest flocks I saw were two small flocks on April 1. Outside of thecities and towns we saw these quail on the brush-covered hillsides, onthe grassy plains in the wider canyons, in cultivated fields and in thefruit orchards, or almost anywhere that they can find a little cover.Claude T. Barnes writes to me:In northern Utah the favorite habitat of the valley quail (L. c. vallicola) isthe patches of scrub oak (Quercus gamoellii), which grow upon the foothills ofthe Wasatch Mountains and along the deeper stream-gullies of the valleys. Itis very fond, also, of fences along which, in early days, the golden currant(Ribes aureum) was planted and permitted to spread along irrigation ditches;in fact, any dense covert adjacent to grainflelds and near one of the manycrystal streams for which the region is noted suits very well this semidomesti-cated bird. Many farmers, convinced of not only the aesthetic but also theeconomic value of the quail, habitually in winter sprinkle grain upon the snowabout their barnyards for the quickly responding coveys.Courtship.?W. Leon Dawson (1923) writes:The Quail's year begins some time in March or early April, when the coveysbegin to break up and, not without some heart-burnings and fierce passages at
VALLEY QTJAIL 63
arms between the cocks, individual preferences begin to hold sway. It is thenthat the so-called " assembly call," ku kwak' up, ku kioak' up, ku kicak' u k-k o, is heard at its best ; for this is also a mating call ; and if not always directedtoward a single listener, it is a notice to all and sundry that the owner is veryhappy, and may be found at the old stand. Although belonging to a polygamousfamily, the Valley Quail is very particular in his affections; and indeed, fromall that we may learn, is at all times a very perfect model of a husband andfather. Even in domestication, with evil examples all about and temptressesin abundance, the male quail is declared to be as devoted to a single mate as inthe chaparral, where broad acres may separate him from a rival.Nesting.?The valley quail is not at all particular about the choiceof a nesting site and is not much of a nest builder. A slight hollowin the ground lined with grass or leaves may be well hidden undera bush, hedge, or brush pile, beside a log or rock, in some thickclumps of grass or weeds in an orchard or vineyard, in a clump ofcactus or pricklypear, under the base of a haystack in an open field,or even in a cranny in a rock. W. Leon Dawson (1923) shows aphotograph of a nest in the latter type of location. Often the nestis near a house, in a garden, or close to a much-traveled path or road.This quail often lays its eggs in other birds' nests. M. L. Wicks(1897) tells an interesting story of a partnership nesting with a long-tailed chat; the quail had laid two eggs in the chat's nest, in whichthe chat laid four eggs ; both birds took turns at incubating. HaroldM. Holland (1917) twice found a quail occupying a road runner'snest. John G. Tyler (1913) speaks of a curious habit this quail hasof dropping its eggs at random anywhere ; this happens early in theseason, and he thinks it is due to the fact that the vines under whichit wants to nest have not yet developed enough foliage for conceal-ment. Grinnell, Bryant, and Storer (1918) mention a few more oddnesting sites, as follows:H. R. Taylor records the finding of ten fresh California Quail's eggs in aSpurred Towhee's nest in a cypress hedge about four feet from the ground, andalso two eggs of this quail in a Spurred Towhee's nest on the ground, both inAlameda. Near Los Angeles, Wicks found two eggs of the Valley Quail in aLong-tailed Chat's nest. Several cases of tree-nesting of the California Quailcame to the attention of W. E. Bryant. The sites which had been chosen werethe upright ends of broken or decayed limbs, or the intersections of two largebranches. The same observer found a nest in a vine-covered trellis over amuch-used doorway, from which the young later successfully reached theground. Howell found a nest with three fresh eggs four feet above the groundon top of a bale of hay in the shade of an orange tree at Covina, Los AngelesCounty.Mr. Dawson (1923) tells of a nest placed " on a horizontal stretchof dense wistaria covering an arbor, at a height of 10 feet from theground " ; when the young were hatched the parent birds calledthem, and they came tumbling down, stunned at first but not seriouslyinjured. He mentions another nest on the roof of a house.
64 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMEggs.?The valley quail lays ordinarily from 12 to 16 eggs; largesets of more than 20 eggs are sometimes found, but these may be theproduct of two hens. They are short ovate in shape, and sometimesrather pointed; the shell is thick and hard, with little or no gloss.The ground colors vary from " cream buff " to " ivory yellow " or,rarely, dull white. They are usually heavily marked and show con-siderable variation. Some eggs are well covered with large blotches,irregularly scattered; others are evenly covered with minute dots;but there are many intermediate variations, and there is generally amixture of both kinds of markings on the same egg and several typesof eggs in the same set. An occasional egg is entirely unmarked.The colors of the markings are dull browns, varying from " snuffbrown " or " cinnamon-brown " to " Isabella color." The measure-ments of 77 eggs average 31 by 24 millimeters; the eggs showingthe four extremes measure 34 by 25, 32 by 26, and 28 by 23 millimeters.Young.?Mrs. Irene G. TVheelock (1904) writes:Incubation requires three weeks, and usually the hen alone broods the eggs,after the young are hatched they are kept in the underbrush or heavy stubbleand can rarely be discovered, so expert at hiding are they. Like the Californiapartridge they run to cover rather than fly, and they are so swift-footed thatit is almost impossible to flush them. When the young are feeding, the adultmales constantly call them, either to keep the covey together or to give warningof danger, and they answer each call with a faint piping note. This is notunlike the scatter call of the Eastern Bob White, but consists of two syllables inone tone, or one longer note. It is not unusual to come upon a covey of thesewhen driving through the foothills and valleys of Southern California, but thesensation is simply of something scampering into the brush rather than a definitesight of any bird, unless the cock comes out into view for a moment to soundhis warning and draw your attention from the brood to his handsome self.Bendire (1892) quotes William Proud as saying that only onebrood is raised in a season, that incubation lasts about 18 days, andthat " as soon as the young are hatched, they immediately leave thenest, keeping under cover as much as possible. Should the broodbe disturbed, the old birds will run and flutter along the ground todraw the attention of the dog, or whatever may have frightenedthem, to themselves and away from the young. In about 10 daysthese can fly a short distance."F. X. Holzner (1896) says:I walked unsuspectingly upon a bevy of Valley Partridges (Oallipepla cali-fomica vallicola), consisting of an old male and female with about 15 youngones. They were in a crevice of a fallen cottonwood-tree. On my steppingalmost upon them, the male bird ran out a few feet and raised a loud call ofca-ra-ho; while the female uttered short calls, addressed to her brood. Seeingme, she picked up a young one between her legs, beat the ground sharply withher wings, and made towards the bush, in short jumps, holding the little onetightly between her legs, the remainder of the brood following her.
VALLEY QUAIL 65Plumages.?In the small chick of this species the front half ofthe crown and sides of the head are " ochraceous-tawny " ; a broadband of " russet," bordered with black, extends from the center ofthe crown to the hind neck, and there is an auricular stripe of thesame color; the rest of the upper parts are from " ochraceous-buff "to " warm buff," striped, banded, or blotched with black ; the chinand throat are white, and the rest of the underparts are grayishwhite, suffused with buff on the breast.As with all young quail and grouse, the juvenal plumage comes inwhile the chick is still very small, the wings and scapulars sproutingfirst, so that the young birds can fly before they are half grown.In the full juvenal plumage, the forehead is " hair brown," thecrown and hind neck " wood brown," and the chin and throat " drabgray " ; the feathers of the back, wing coverts, and scapulars arefrom " hair brown " to " clay color," with median stripes of buffywhite, broadest on the scapulars, peppered with black and tipped orbanded near the end with black ; the tertials are from " sayal brown "to " cinnamon," peppered with black, and bordered on the inneredge with a broad band of black and a broad edge of " pinkish buff "
;
the rump is grayish buff, barred with dusky and whitish ; the tail isfrom " drab " to grayish buff, tipped with " cinnamon," and pep-pered and barred with blackish brown; the underparts are grayishwhite, barred with dusky, more buffy, and marked with triangularwhitish spots on the chest ; the head crest is " warm sepia." Thesexes are alike.A complete postjuvenal molt, except for the primary coverts andthe outer pairs of primaries, begins before the young bird has at-tained its full growth. The time varies, of course, with the date ofhatching, but it takes place between August and October. The lastof the juvenal plumage is seen on the head and neck. This molt pro-duces the first winter plumage, which is practically indistinguish-able from the adult, except for the outer pairs of juvenal primariesand primary coverts, which are retained until the next postnuptialmolt. The sexes are now different.Young birds and adults have a partial prenuptial molt, confined tothe head and neck, early in spring, and a complete molt late insummer and early in fall. Hybrids between this species and gam-oeli and between this and picta have been recorded, where the rangesof the species come together.Food.?These quail are very regular in their feeding habits. Whenthey have found a good feeding place they resort to it day after day,often traveling long distances on foot and not flying unless forcedto. They travel in flocks at all times except during the nesting sea-son, when they are paired. Formerly they came to the watering
66 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMplaces in immense flocks of hundreds, but now in flocks of 30 or 40,or aggregations of two or three families. Their feeding hours arefor an hour or two after sunrise and an hour or two before sun-set. During the middle of the day they congregate near the drink-ing places or rest in the shade of trees or bushes. While feeding, onebird acts as sentinel or guard until relieved by others in turn.Grinnell, Bryant, and Storer (1918) write:The Valley and California quails are believed to be more exclusively vege-tarian than any other of our game birds, save those of the pigeon family.The United States Bureau of Biological Survey, in an examination of 619stomachs (representing both subspecies), found that only about 3 per cent ofthe food consisted of animal matter. The remaining 97 per cent was vegetablematerial and consisted of 2.3 per cent fruit, 6.4 per cent grain, about 25 percent grass and other foliage, and 62.5 per cent seeds. The animal food com-prised chiefly insects, and of these, ants were most frequently present. Somebeetles, bugs, caterpillars, grasshoppers, flies, spiders, " thousand-leggers," andsnails were also found in the stomachs examined. A case is cited by Beal ofa brood of young quail feeding extensively on black scale.Fruit evidently does not form any important part of the food of the quail, asit was found in only about one-sixth of the stomachs and then only in verysmall quantities. Damage is sometimes done to grapes, but this is not shownclearly by examination of stomach contents. Beal mentions two cases where1,000 and 5,000 quail, respectively, had been seen feeding upon grapes in vine-yards. Under such circumstances severe loss was undoubtedly sustained;but these are exceptional instances. Florence A. Merriam states that on theranch of Major Merriam at Twin Oaks, San Diego County, quail were in 1889so abundant as to be a severe pest. For several years previously great flocksof them came down the canons to the vineyard, " where they destroyedannually from 20 to 30 tons of fruit." A report comes from the Fresno dis-trict to the effect that grape growers are occasionally troubled by the birdsscattering the drying raisins from the trays.Behavior.?The movements and actions of valley quail seem to mestrikingly like those of our eastern bobwhites, except that the}' areless inclined to fly or to hide and more inclined to run. Whenalarmed or forced to fly they jump into the air with a similar whirof wings and dash away with an equal burst of speed, scaling downinto the nearest cover on stiff, down-curved wings. If they alighton the ground, they do not stop, but continue running at terriiicspeed, their long, strong legs fairly twinkling in a hazy blur; itseems as if they continued to fly along the ground almost as fast asthey flew in the air. On the ground their movements are quick,alert, and graceful; their trim and pretty little bodies are held in asemierect attitude, leaning forward a little as they run, with thecrest held forward. They are most attractive in appearance andmost winning in their confiding ways.John J. Williams (1903) made some very interesting observationson the use of sentinels by valley quail ; his article is well worth read-
VALLEY QUAIL 67ing in detail, but I prefer to quote Grinnell, Bryant, and Storer's(1918) summary of it, as follows:A flock was heard calling and moving about on a brushy hillside somedistance from the observer, but before coming into view a single individualpreceded the rest and took his station in the branches of an apple tree, whencehe could survey the region round about. After carefully scrutinizing hissurroundings for several minutes the kayrk note was uttered several timesin a low guttural tone. Soon members of the flock were seen coming downthe hill in the same direction as taken by the sentinel, but their manner ofapproach was entirely different; he had exercised great caution and care-fully examined the surroundings for possible danger, while they came withtheir plumed heads held low, searching among the clover roots for seeds andother articles of food. Some preened and fluffed out their feathers; otherstook dust baths. While so occupied they all kept up a succession of lowconversational notes. Meanwhile the sentinel remained on his perch and con-tinued on the alert even after the flock had moved some distance beyond him.Then a second bird mounted a vantage point and took up the sentinel dutyand after a few minutes the first relinquished his post. While the flock wasstill in view, yet a third bird relieved the second. It would seem that by thispractice, of establishing sentinels on a basis of divided labor, the flock hadincreased its individual efficiency in foraging. The same observer also statesthat he had seen sentinels used when a flock was crossing a road, or when
" bathing " in the roadside dust, and that the practice is made general useof in open areas ; but he had never observed the habit when the birds werein tree-covered localities. During the breeding season it is known thatthe male mounts guard while the female is searching for a nesting site, andagain when she is incubating the eggs. Sometimes he also performs thisguard function after the chicks are out but not fully grown.Unlike our eastern bobwhite, which roosts on the ground, thevalley quail roosts at night in safer places, in bushes or in low,thick-foliaged trees. In the treeless region of Lower California,Laurence M. Huey (1927) found quail roosting in the centers ofcactus patches. Dawson (1923) says he has "seen a wounded birdswim and dive with great aplomb."Voice.?Some of the notes of this quail suggest the familiar bob-white of our eastern quail. Grinnell, Bryant, and Storer (1918) havedescribed them very well, as follows:The Valley Quail has a variety of notes which are used under different con-ditions and to express various meanings. When anxious or disturbed themembers of a flock utter a soft pit, pit, pit, or whit, whit, whit, in rapidsuccession, as they run about under the brush or when about to take wing.Then there is a loud call used by the males to assemble the flock when scat-tered. This has been variously interpreted as ca-loi'-o, o-hi'-o, tuck-a-hoe'
,
k-woik'-uh, ki-ka-kce', ca-ra'-ho, tuck-ke-teu' , or more simply as who-are'-yow-ah. However, the easiest and by far the most usual interpretation is come-right' -here, or come-right-home, with the accent on the second syllable. Some-times when excited a bird calls come-right, come-right, come-right-here. In atleast one instance a female bird has been observed to utter this call. The notesof the Valley Quail are less elaborate than those of the Desert Quail, the
68 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
"crow " lacking the two additional notes which the latter gives at the end ; alsothe Valley Quail lacks much of the conversational twitter of its desert relative.Enemies.?Quail have numerous enemies; the eggs and young ofthese and other ground-nesting birds are preyed' upon by crows,ravens, jaj's, snakes, raccoons, weasels, skunks, squirrels, and badg-ers; the adults also are pursued and killed by hawks, owls, coyotes,foxes, bobcats, and domestic cats and dogs. Gopher snakes areparticularly fond of quail's eggs. Joseph Dixon (1930) tells astriking story of a brood of 19 young quail that was entirely de-stroyed b}' a pair of California jays, which he says are one of thequail's worst enemies; he saw four chicks carried off by one jaywithin 15 minutes.Game.?The California quail, in its two forms, has often beenreferred to as the game bird of California, has been hunted bymore sportsmen and market hunters than any other bird, and hasbeen killed in enormous numbers. Its great abundance in formeryears seems almost unbelievable to-day. Dr. A. K. Fisher (1893)wrote
:
Throughout the San Joaquin Valley, Mr. Nelson found it common ahoutranches, along water courses or near springs. It was excessively abundant atsome of the springs in the hills about the Temploa Mountains and CarrizoPlain. In the week following the expiration of the close season, two men,pot hunting for the market, were reported to have killed 8,400 quail at asolitary spring in the Temploa Mountains. The men built a brush blind nearthe spring, which was the only water within a distance of 20 miles, and asevening approached the quails came to it by thousands. One of Mr. Nelson'sinformants who saw the birds at this place stated that the ground all aboutthe water was covered by a compact body of quails, so that the huntersmowed them down by the score at every discharge.Grinnell, Bryant, and Storer (1918) say:In the days when the Valley Quail was plentiful far beyond its conditionto-day, it was a common bird on the markets and could be obtained at prac-tically every hotel and restaurant. Records show that during the season1895-96 as many as 70,370 quail (mostly Valley Quail) were sold on themarkets of San Francisco and Los Angeles ; 'while an earlier report statesthat full 100,000 were disposed of in a single year in the markets of SanFrancisco. W. T. Martin, of Pomona, states that in 1S81-S4 he and a partnerhunted Valley Quail in Los Angeles and San Bernardino Counties for theSan Francisco markets. Eight to fourteen dozen were secured daily, andin the fall of 1S83 the two men secured 300 dozen in 17 days. Martin himselfsecured 114 birds in one day's hunt. In 18S1 and 18S2 over 32,000 dozen quailwere shipped to San Francisco from Los Angeles and San Bernardino Counties,and brought to the hunters engaged in the business one dollar a dozen. In thosedays restaurants charged thirty cents for quail-on-toast. By 1885 hunting hadbecome unprofitable because of the reduction in the numbers of quail.Quoting from T. S. Van Dyke (1892), they say:At your first advance into the place where the quail last settled in confusion,a dozen or more rise in front of you and as many more on each side anywhere
VALLEY QUAIL 69from 5 to 50 yards away. They burst from the brush with rapid flight andwhizzing wing, most of them with a sharp, clear, pit, pit, pit, which apprizestheir comrades of the danger and the course of escape taken. Some dartstraight away in a dark blue line, making none too plain a mark against thedull background of brush, and vanish in handsome style, unless you are veryquick with the gun. Others wheel off on either side, the scaling of their breastsshowing in the sunlight as they turn, and making an altogether beautiful markas they mount above the skyline. Some swing about and pass almost over yourhead, so that you can plainly see the black and white around their heads andthroats, and the cinnamon shading of their under surfaces.Although this quail is a splendid game bird and as good on thetable as our eastern quail, all sportsmen who have shot both seemto agree that our bobwhite is a far more satisfactory bird to hunt.The valley quail will not lie to a dog, unless thoroughly frightened
;
it has a most exasperating habit of running, which is quite dis-heartening to both man and dog. Dwight W. Huntington (1903)referring to Mr. Van Dyke's comments on the former abundanceand habits of this quail, says
:
He said that when he first came to California, in 1875, quail in flocks nowquite incredible soared out of almost every cactus patch, shook almost everyhillside with the thunder of a thousand wings, trotted in strings along theroads, wheeled in platoons over the grassy slopes and burst from around almostevery spring in a thousand curling lines. The same writer says that thepartridges have already deserted many of the valleys and are now more oftenfound in the hills, ready always to ruu and fly from one hillside to another,and " their leg power, always respectable enough to relieve you from anyquestion of propriety about shooting at one running, they have cultivated tosuch a fine point that sometimes they never rise at all, and you may chase andchase and chase them and get never a rise." Writing at another time Mr.Van Dyke advises the shooter not to attempt to bag anything at first, but tospend all the time in breaking and scattering the coveys, racing and chasingafter them and firing broadsides over their heads and in front of them, untilthey are in " a state of such alarm that they will trust to hiding." He thenadvises that the dog (which I presume has been used in coursing the birds)be tied to a shady bush and that the coat be laid aside, that the sportsman maytravel fast after the scattered birds.Occasionally they may behave differently and offer good sport, asin the following account by Henshaw (1874) :As a rule, their ways are not such as to endear them to the sportsman ; forthey are apt to be wary, and unless under specially favorable circumstances,are not wont to lie closely. I have, however, flushed a large bevy contiguousto a bushy pasture where the scrub was about knee-deep, with cattle-pathsthrough it, and have had glorious sport. The birds lay so close as to enableme to walk almost over them, when they got up by twos and threes, and wentoff in fine style. The sportsman may now and then stumble upon such chances,but they do not come often. A bevy once up, off they go, scattering but littleunless badly scared, the main body keeping well together ; and having flowna safe distance they drop, but not to hide and be flushed one after another atthe leisure of the sportsman. The moment their feet touch firm ground, off74564?32 6
70 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthey go like frightened deer, and if, as is often the case, they have been flushednear some rocky hill, they will pause not a moment till they have gained itssteep sides, up which it would be worse than useless to follow. Should they,however, be put up hard by trees, they will dive in among the foliage andhide, and there standing perfectly motionless will sometimes permit one toapproach to the foot of the tree they are lodged in ere taking wing.Winter.?What birds are left in the big flocks, after the sportsmenhave taken their toll, remain together during fall and winter, for-merly in great droves of hundreds, but now more often in flocks of40 or 60. They are not migratory to any extent. A. C. Lowell, oneof Major Bendire's (1892) correspondents in Nevada, told him thatthey were not able to stand the severe cold, accompanied by a heavyfall of snow in the Warner Valley; 2 feet of snow and 3 nights of28? below zero killed most of the birds. On the other hand, MajorBendire (1892) tells of a flock that spent the winter successfullynear Fort Klamath, Oreg., where the snowfall is quite heavy andthe thermometer fell " more than once considerably below zero."LOPHORTYX CALIFORNICA CATALINENSIS GrinnellCATALINA QUAILHABITSBased on a series of six specimens Dr. Joseph Grinnell (1906)gave the name Lophortyx califomica catalinensis to the Californiaquail inhabiting Santa Catalina Island. He characterized it assimilar to the valley quail, " but about 9 per cent larger throughout,and coloration somewhat darker; similar to L. c. californicus, butlarger and much less deeply brownish dorsally." He says further
:
The bulkiness of catalinensis is at once apparent when one sees it amongspecimens of the mainland vallicola. The tail is particularly long, the rectricesbeing proportionately broader. The bill is heavier, and the toes and tarsidecidedly stouter These characters hold equally in the males and females.In coloration catalinensis shows a deepening of shades especially on the lowersurface. In both sexes the flanks and lower tail-coverts are more broadlystreaked with brown; the terminal black edgings of the lower breast feathersare broader, and the light markings beneath are suffused with deeper ochraceous.Especially in the female of catalinensis is the lower surface darker than invallicola, due to the encroachment of the dark portions of each parti-coloredfeather upon the light part. The dorsal surface is not however much brownerthan in vallicola?it is decidedly slaty as compared with the deep bright vandykebrown of californicus from the vicinity of San Francisco Bay.It was thought at first that these quail had been introduced fromthe mainland, but more recent evidence shows that they were prob-ably native on the island, which perhaps was once connected withthe mainland. Doctor Grinnell (1906) was assured that they werethere at least as early as 1859.
SAN QUINTIN QUAIL 71After examining a series of 16 skins, at a later date, Doctor Grin-nell (1908) writes:When compared with a series of the mainland vallicola the island birds aredistinguished by larger size, especially of the feet, broadness of terminalbarring on the posterior lower surface, and broadness of shaft-streaks on lowertail coverts and flanks. An additional character which shows up in the largerseries is the averaging more intense and extensive chestnut patch on the hindchest, in the male, of course. This does not seem to be due to the different
" make " of the skins. An examination of individual variation in the twoseries shows that any one character alone is not diagnostic of every singleindividual. For instance, a small-footed island bird can be duplicated in thatrespect by an extra large-footed mainland bird. But at the same time thebarring and streaking of the former renders it easily recognizable. Then in thematter of barring on the lower surface, a mainland female appears as heavilymarked as the average island female. But at the same time the former has adecidedly shorter wing and weaker foot. It is therefore evident that there is amergence of separate characters thru individual variation ; and according tothe criterion now apparently most popular, the island form would be given atrinomial appellation. The binomial, however, appears to me most useful, as itsignifies complete isolation because of the intervention of a barrier.LOPHORTYX CALIFORNICA PLUMBEA GrinnellSAN QUINTIN QUAILHABITSUnder the name Lophortyx californica plumbea Dr. JosephGrinnell (1926) has separated the quail found in certain parts ofnorthern Lower California from the subspecies found on either sideof it. He describes the new form as " in general characters similarto Lophortyx californica vallicola and L. c. ac7irustera, but tone ofcoloration clearer, less buffy or brownish; gray or lead-color ondorsum, foreparts and sides, and remiges, more slaty than in either."He says further:It should go without saying that in quail fresh fall plumages should be reliedupon chiefly, if not altogether, in seeking color values. When this is done, thequail of the " San Quintin district " show themselves to differ in mass effectappreciably from Valley Quail from anywhere north of the Mexican line. SanDiego County birds, even, and those from Riverside and Inyo counties, welleast of the desert divides, all are markedly browner dorsally, the remigesbrowner, the chest less clearly ashy gray, and the " ground " tone of the hinderflanks and crissum more brightly tan. This holds for both sexes. The creamyarea on the lower chest of male plumbea, while not so pale as in achrustera,is not so deep-toned as in average vallicola. In females the grayness about thehead and on the chest in plumbea is almost constantly diagnostic ; and in bothsexes, the plumbeous tone of the remiges is as a rule strikingly different fromthe brown tone in vallicola. In the dried specimens, the feet and legs ofplumbea average blacker than in vallicola.
72 BULLETIN 162, UNITED STATES NATIONAL MUSEUMKeferring to its distribution and haunts, he says in a later publi-cation (1928) :Abundant resident of the northwestern portion of the territory, roughlybetween latitudes 30? and 32??practically as comprised in the San Quintinsubfaunal district. While the metropolis of the subspecies lies on the Pacificslope of the peninsula, colonies or pairs occur also to the eastward, in canyonsat the east base of the Sierra San Pedro Martir, and even at San Felipe, onthe Gulf coast (Mas. Vert. Zool.). As regards life-zone, inhabits the LowerSonoran, Upper Sonoran, and Transition, without any seeming choice. Asso-ciationally, adheres to an open or interrupted type of chaparral, especially asadjacent to springs or water-courses. Altitudinally, extends from sea level upto as high as 8,800 feet, on the Sierra San Pedro Martir.Eggs.?The eggs are probably indistinguishable from those ofother races of California quail. Griffing Bancroft (1930) says thatthe measurements of 150 eggs average 30.6 by 23.3 millimeters.LOPHORTYX CALIFORNICA ACHRUSTERA PetersSAN LUCAS QUAILHABITSJames L. Peters (1923) is responsible for the name L. c. achrustera,which he has applied to the quail of this species inhabiting southernLower California. Based on the examination of a series of 27 malesand 15 females, he says that it is " similar to Lophortyx califomicavatticola (Ridgway), but slightly paler above; band across breastgrayer; the buffy patch on the lower breast of the male much paler;dark feather-edgings on the lower breast, middle and sides of abdo-men, narrower; flanks paler."William Brewster (1902), with much of the same material, noticedthat the Lower California specimens were "slightly paler" thanCalifornia birds and their bills averaged " a little heavier," but hedid not consider these differences well marked or constant. ButPeters (1923) says that "while the bill character is of no diagnosticvalue, the color characters are constant and serve to distinguish thevalley quail of southern Lower California almost at a glance."Griffing Bancroft (1930) says of their haunts: "They insist onriparian associations, but they follow these without regard either toaltitude or to the character of the country adjoining the streambeds. They definitely do not require the presence of water."Nesting.?-Of the nesting of the San Lucas quail, Bancroft (1930)says:Our experience with the breeding of these quail was limited to San Ignacio.That was because the nests were too well hidden to be found, except acci-dentally, and those we saw were shown to us by the natives. Three ofthe sites were in damp ground in rank grass; one of them, to our surprise,on a tiny islet in a swamp. Two nests were in vineyards, two in natural
gambel's quail 73
cavities among the sucker growths of date palms, and one was under a lavarock on the mesa. In all but the last three cases the birds had excavateda cup nearly as deep as it was broad and had lined it with materials broughtin, grass, leaves, and feathers. The breeding season commences about thefirst of June and is hardly well under way until after the middle of thatmonth. The number of eggs in a clutch is rather consistently ten or eleven,sixteen being the most we found in any one nest.Eggs.?He gives the average measurements of 80 eggs as 32.3by 24.7 millimeters. The measurements of 15 eggs in P. B. Philipp'scollection average 31.8 by 24.3 millimeters; the eggs showing thefour extremes measure 33.6 by 25.4, 29.7 by 23.4, and 31.3 by 23.1millimeters. LOPHORTYX GAMBELI GAMBELI GambelGAMBEL'S QUAILHABITSGambel's quail is also very appropriately called the desert quail,for its natural habitat is the hot, dry desert regions of the South-western States and a corner of northwestern Mexico. Its center ofabundance is in Arizona, but it ranges east to southwestern NewMexico and El Paso, Tex., and west to the Colorado and MohaveDeserts in southeastern California. On the western border of itsrange it is often associated with the valley quail and has been knownto hybridize with it.This beautiful species was discovered by Dr. William Gambel " onthe eastern side of the Californian range of mountains in 1841 " andnamed in his honor, according to John Cassin (1856), who gives usthe first account of its distribution and habits, based largely on notesfurnished by Col. George A. McCall. He did not meet with itwest of the Colorado Desert barrier in California or east of the PecosRiver in Texas.We found Gambel's quail very common in southern Arizona,especially in the lower river valleys, where the dense growth ofmesquite (Acacia glandulosa) afforded scanty shade, or where theycould find shelter under the spreading green branches of the palo-verdes, which in springtime presented great masses of yellow blos-soms. They were even more abundant in the thickets of willowsalong the streams or in the denser forests of mesquites, hackberries,and various other thorny trees and shrubs. We occasionally flusheda pair as we drove along the narrow trails, but more often we sawthem running off on foot, dodging in and out among the desertunderbrush until out of sight. My companion on this trip, Francis
74 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMC. Willard, has sent me the following notes, based on his longexperience in Arizona
:
Gambel's quail is essentially a bird of the areas in southern Arizona wherethe mesquite abounds. Unlike their neighbor, the scaled quail, they seem torequire the close proximity of a water supply. They are, therefore, foundprincipally along the few living streams and close to permanent water holes.I found them swarming in the mesquite forest along tbe Santa Cruz Riversouth of Tucson and almost as plentiful along the Rillito between Tucson andthe mountains. In the valley of the San Pedro River they were also presentin large numbers. Between the valley of this last-named river and the variousranges of mountains fringing it are long sloping mesas from 5 to 20 mileswide where the "black topknot" is rarely seen except close to the infrequentwater holes. In the foothills of the Dragoon, Huachucas, Whetstones, Chirica-huas, and other less well-known ranges this quail again appears in somenumbers but nothing like those in the lower valleys.Dr. Elliott Coues (1874) has given us the best account of thisquail, which I shall quote from quite freely. He says of its haunts
:
Gambel's Quail may be looked for in every kind of cover. Where they aboundit is almost impossible to miss them, and coveys may often he seen on exposedsand-heaps, along open roads, or in the cleared patches around settlers' cabins.If they have any aversion, it is for thick high pine-woods, without any under-growth ; there they only casually stray. They are particularly fond of thelow, tangled brush along creeks, the dense groves of young willows that growin similar places, and the close-set chaparral of hillocks or mountain ravines.I have often found them, also, among huge granitic boulders and masses oflava, where there was little or no vegetation, except some straggling weeds;and have flushed them from the dryer knolls in the midst of a reedy swamp.Along the Gila and Colorado they live in such brakes as I described in speakingof Abert's Finch ; and they frequent the groves of mezquite and mimosa, thatform so conspicuous a feature of the scenery in those places. These scrubbytrees form dense interlacing copses, only to be penetrated with the utmostdifficulty, but beneath their spreading scrawny branches are open intersectingways, along which the Quail roams at will, enjoying the slight shade. In themost sterile regions they are apt to come together in numbers about the fewwater-holes or moist spots that may be found and remain in the vicinity, sothat they become almost as good indication of the presence of water as theDoves themselves. A noteworthy fact in their history, is their ability to bear,without apparent inconvenience, great extremes of temperature. They areseemingly at ease among the burning sands of the desert, where, for months,the thermometer daily marks a hundred, and may reach a hundred and forty,
" in the best shade that could be procured," as Colonel McCall says ; and theyare equnlly at home the year round among the mountains, where snow lieson the ground in winter.
. In New Mexico, according to Mrs. Florence M. Bailey (1928),it is foundin the Lower Sonoran Zone in quail brush (Atriplex lentiformte) and creosote,and in hot mesquite valleys or their brushy slopes, in screw bean and paloverde thickets and among patches of prickly pear. It is not generally foundso far from water as the Scaled Quail, which eats more juicy insect food,but at times both are seen in the same landscape.
GAMBEl/s QUAIL 75In inhabited regions, in places where cattle trails lead to water, the Gambel'spretty foot prints call up pleasant pictures of morning procession of thirsty little" black-helnieted " pedestrians, talking cheerfully as they go. For it seemsmost at home about small farms, such as those cultivated by the Spanish-Americans, which dot the narrow canyons and river valleys.Courtship.?Springtime in Arizona is most charming as the desertplants burst into bloom with their profusion of many colors. Thenew fernlike foliage of the mesquite mingles with dangling yellowtassels. The long slender stems of the ocotillo are tipped with ver-milion spikes. Even the lowly creosote bush is clouded with yellowhaze. The various chollas and the pricklypears are studded withpink, yellow, or crimson flowers, and the little rainbow cactus bloomsby the roadside with a wealth of large magenta and yellow blos-soms. Even the giant cactus supports a crown of white, and thepaloverde is the showiest of all, a great bouquet of brilliant yellow.Then we may look for the trim figure of the cock quail, perchedon some low tree, bush, or stump, and listen to his challenging lovecall.Major Bendire (1892) has described his courtship very well, asfollows
:
During the mating and breeding season, the former commencing usuallyin the latter part of February, the latter about the first week in April andoccasionally later, according to the season, the male frequently utters acall like yuk-kae-ja, yuk-kae-ja, each syllable distinctly articulated and thelast two somewhat drawn out. A trim, handsome, and proud-looking cock,whose more somber-colored mate had a nest close by, used an old mesquitestump, about 4 feet high, and not more than 20 feet from my tent, as hisfavorite perch, and I had many excellent opportunities to watch him closely.Standing perfectly erect, with his beak straight up in the air r his tail slightlyspread and wings somewhat drooping, he uttered this call in a clear strongvoice every few minutes for half an hour or so, or until disturbed by some-thing, and this he repeated several times a day. I consider it a call ofchallenge or of exultation, and it was taken up usually by any other malein the vicinity at the time. During the mating season the males fight eachother persistently, and the victor defends his chosen home against intrusionwith much valor. It is a pleasing and interesting sight to watch the malecourting his mate, uttering at the time some low cooing notes, and struttingaround the coy female in the most stately manner possible, bowing his headand making his obeisance to her. While a handsome bird at all times, hecertainly looks his best during this love-making period.Nesting.?My experience with the nesting habits of Gambel'squail is limited to three nests found near Tucson, Ariz., in 1922.On May 19 we were hunting through the mesquite forest, a largetract of once heavy timber that had been much depleted by theraids of Papago Indian woodchoppers. There were only a fewlarge trees left, some very large hackberry trees, which were moreor less scattered with many open spaces; but there was plenty of
76 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
cover left in the extensive thickets of small mesquite and thornyundergrowth, or in the patches of large mesquite, oaks, and hack-berry. While walking along the edge of a dry ditch, we flusheda Gambel's quail from its nest under a tiny thorn bush. The nestwas a rather deep hollow lined with sticks, straws, leaves, andfeathers; it held 10 eggs.Two days later, while hunting among the giant cactuses, whichhere were scattered over an open plain, scantily overgrown withlow mesquite and greasewood bushes in dry stony soil, we flushedanother quail from under a large mesquite bush ; way under it, at thebase of the trunk and almost beyond reach in the thorny tangle, wasits nest with 16 eggs. The next day while investigating a Palmer'sthrasher's nest, five feet above the ground in a cholla, we were sur-prised to find in it three eggs of this quail. I have since learnedthat it is not unusual for this quail to use old nests of thrashers orcactus wrens. Perhaps the birds have learned by sad experiencethat ground nests are less safe.My companion, Frank Willard, who has had much wider experi-ence than I, has sent me the following notes
:
From early in May well into July and sometimes even into August nestswith eggs may be found. The last week of May and the first week in Juneseem to be the height of the egg-laying season. The eggs of gambelii are laidin more exposed situations than those of squamata. The most frequentlychosen site is at the foot of a small mesquite or other bush where a slighthollow is scratched in the dry ground. There is one protection, however, whichthe quail seems to find necessary. There must be something to shade them fromthe hot midday sun. The scanty shade afforded by the fern leaf of themesquite is sufficient but there must be some at least.Two or more females lay their eggs in the same nest very frequently. Ihave had nests under observation where two or more eggs were added dailyto the complement therein. It has occurred to me that this is a wise provisionof nature to secure a nest full of eggs with as little delay as possible so thatincubation could be undertaken promptly and an even hatching take placewithout the eggs being exposed to the dry desert heat until one bird could laya full set, which averages a dozen eggs or more. A few days of exposure tothe dry air without the moisture from the body of the sitting bird would makemany of the eggs sterile. Nearly every egg in nests where incubation hadcommenced was fertile and I seldom found more than one unhatched eggamong the debris of a nest from which the young had hatched and gone.On May 14, 1908, I went to collect a set of Gambel's quail eggs which I hadbeen watching for some time. The previous day there had been IS eggs in thenest, some of them those of the scaled quail. As I looked around the largerock behind which the nest was concealed I found the female quail flutteringabove her nest in which was coiled a large rattlesnake. With head upliftedit was striking at the bird which deftly avoided the blows. On my appearancethe bird flew away. I prodded the snake, driving it from the nest, and thenkilled it. Eleven rattles adorned its tail. There were 16 eggs left, all of whichwere fresh. I foolishly neglected to open the snake and look for the twomissing eggs.
gambel's quail 77
I once found a Gambel quail sitting on 16 eggs laid in a Palmer thrasher'snest 5 feet up in a cholla sheltered by a large sycamore. The bird sat veryclose. On another occasion I found several eggs of this quail in the nestof a Palmer thrasher and the thrasher sitting on them and her own three eggs.Major Bendire (1892) writes:The nest of Gambel's Partridge is simply a slight oval-shaped hollow,scratched out in the sandy soil of the bottom lands, usually alongside of abunch of " sacaton," a species of tall rye grass, the dry stems and blades oflast year's growth hanging down on all sides of the new growth and hidingthe nest well from view. Others are placed under, or in a pile of, brush ordrift brought down from the mountains by freshets and lodged against someold stump, the roots of trees, or other obstructions on some of the numerousislands in the now dry creek beds, refreshing green spots amid a dreary wasteof sand. According to my observations only a comparatively small numberresort to the cactus and yucca covered foothills and mesas some distance back,where the nests are usually placed under the spreading leaves of one of thelatter named plants. If grain fields are near by they nest sometimes amidstthe growing grain in these, and should the latter be surrounded by brushfences, these also furnish favorite nesting sites.Among the nests observed by me two were placed in situations aboveground. One of these was found June 2 on top of a good-sized rotten willowstump, about 2*A feet from the ground, in a slight decayed depression in itscenter, which had, perhaps, been enlarged by the bird. The eggs werelaid on a few dry Cottonwood leaves, and were partly covered by these.Another pair appropriated an old Road-runner's nest, Oeococcyx caU/ornianus,in a mesquite tree, about 5 feet from the ground, to which apparently a littleadditional lining had been added by the bird. The nest contained 10 fresheggs when found on June 27, 1872.M. French Gilman (1915) found this quail quite tame and confid-ing, nesting in much-frequented localities, for he says
:
Two nests were in the school woodpile, containing 19 and 13 eggs, respec-tively. Another, in a pile of short boards and kindling about 10 feet fromthe school woodshed, had 7 eggs in it. The nest out in the fields had 9 eggs,and was at the base of a Lycium bush. About the middle of June I put somestraw in an old nail keg, open at one end, and placed it on its side in theforks of a mesquite tree about two feet from the ground. The mesquite hadsome saplings starting from the trunk that sheltered the keg. June 24, Ifound that a quail had moved in and had laid two eggs. Later she completedthe set, only eight eggs, and successfully hatched all but one. She was quitetame on the nest, and would not be scared off by any mild measures. Itried hammering on the rear of the keg, rolling it gently and talking to her,requesting her to get off and let me count the eggs, but unless I put my handat the front of the keg she sat pat.Eggs.?Ten or a dozen eggs constitute the average set, but setsof 18, 19, and 20 have been recorded. These large sets are doubtlessthe product of two hens, as indicated by Mr. Willard's observationsabove and by the fact that these sets usually contain two types ofeggs. The eggs are short ovate in shape and sometimes somewhatpointed; they are smooth and slightly glossy. The ground colors
78 BULLETIN 162, UNITED STATES NATIONAL MUSEUMvary from dull white to " cartridge buff " or " pale pinkish buff."They are irregularly spotted and blotched with a mixture of largeblotches, small spots, and fine dots; sometimes the ground color iswell covered, but more often not. The markings are in shades ofdark or purplish browns, from " warm sepia " or " chocolate " to
" snuff brown." One of my sets has 5 eggs of the ordinary typeand 11 beautiful eggs with a " pinkish buff " ground color, wellcovered with small blotches, spots, and fine dots of " deep grayishlavender " and " deep heliotrope gray." This set shows the purplishbloom referred to by Bendire (1892), which turns dark brown, whenwashed, and dries out to purple again. The measurements of 99eggs in the United States National Museum average 31.5 by 24 mil-limeters; the eggs showing the four extremes measure 34 by 26, 28.5by 24, and 30 by 23 millimeters.Young.?Incubation seems to be performed by the female aloneand is said to require from 21 to 24 days. Both sexes share in thecare of the young. Very early and very late dates indicate that atleast two broods are raised in a season. Coues (1874) says of theyoung
:
They run about as soon as they are hatched, though probably not " with haltshell on their backs," as some one has said. In a few days they become verynimble, and so expert in hiding that it is difficult either to see or catch them.When the mother bird is surprised with her young brood, she gives a sharpwarning cry, that is well understood to mean danger, and then generally flies alittle distance to some concealed spot, where she crouches, anxiously watching.The fledglings, by an instinct that seems strange when we consider how shorta time they have had any ideas at all, instantly scatter in all directions, andsquat to hide as soon as they think they have found a safe place, remainingmotionless until the reassuring notes of the mother call them together again,with an intimation that the alarm is over. Then they huddle close aroundher, and she carefully leads them off to some other spot, where she looks forgreater security in the enjoyment of her hopes and pleasing cares. As longas they require the parent's attention they keep close together and are averseto flying. Even after becoming able to use their wings well, they prefer torun and hide, or squat where they may be, when alarmed. If then forced up,the young covey flies off, without separating, to a little distance, often re-alighting on the lower limbs of trees or in bushes, rather than on the ground.As they grow older and stronger of wing, they fly further, separate more readily,and more rarely take to trees; and sometimes, before they are fully grown,they are found to have already become wary and difficult of approach. Asone draws near where a covey is feeding, a quick, sharp cry from the birdwho first notices the approach alarms the whole, and is quickly repeated by therest, as they start to run, betraying their course by the rustling of driedleaves. Let him step nearer, and they rise wiLh a whirr, scattering in everydirection.Plumages.?In natal down the young Gambel's quail is much likethe California quail at the same age, but the colors are paler andduller. The front half of the crown and the sides of the head are
gambel's quail 79from "clay color" to "pinkish buff"; there is a broad band of
"russet," bordered with black, from the tiny topknot to the hindneck, and a dark brownish auricular patch; the rest of the upperparts is light " pinkish buff," banded lengthwise and blotched with
" warm sepia " ; the underparts are pale grayish buff.As with all other quail and grouse, the juvenal plumage begins togrow soon after the chick is hatched, appearing first on the wingsand scapulars ; the topknot, or crest, appears at once, " hazel " at firstand then dull brown. In fresh juvenal plumage the feathers of themantle are variegated with " cinnamon," " pinkish buff," and black,each with a broad, median, white stripe ; later these fade to gray, palebuff, and dull brown; the scapulars have buffy edgings; the tertialsare at first " pinkish buff," later grayish, barred with dusky andtipped with white ; the tail is grayish, barred near the tip with dusky,dull whitish, and dull buffy ; the underparts are grayish white, faintlybarred with dusky; on the head, which is the last part to be feathered,the forehead is dusky and the crown " mikado brown." In thisplumage the sexes are alike, and the birds closely resemble youngCalifornia quail of the same age.The birds are hardly fully grown and the juvenal plumage ishardly complete before the change into the first winter plumagebegins on the back and wing coverts. This change is accomplishedby a complete postjuvenal molt, except that the two outer juvenalprimaries, and their coverts, on each wing are retained for a fullyear. Otherwise the young birds are practically indistinguishablefrom adults after the molt is completed in October, or later.Subsequent molts for old and young birds consist of a very limitedprenuptial molt in April and May, involving only the head and neck,and a complete postnuptial molt in August and September.Food.?In the stomachs, collected from January to June, Dr.Sylvester D. Judd (1905) found that less than half of 1 per centof the food consisted of insects, which included ants, beetles, grass-hoppers, leaf hoppers, and stink bugs. Vegetable matter made up99.52 per cent of the food; 3.89 per cent of this was grain, 31.89per cent seeds, and the balance, 63.74 per cent, was made up of leavesand shoots of various plants. The grain included corn, wheat, andoats, much of which was probably picked up among the grain shocks,where large flocks have been seen feeding with domestic poultry.The seeds were largely those of leguminous plants such as alfalfa,bur clover, and mesquite, and also of alfilaria, mustard, chickweed,peppergrass, and atriplex. Succulent foliage and shoots form byfar the larger percentage of the food. Of this, alfalfa, bur clover,and the foliage of other legumes constitute the greater part. Boththe green leaves and pods of alfalfa are freely eaten. In spring this
80 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMquail shows a fondness for buds, and in some localities its flesh has adistinctly bitter taste due to a diet of willow buds. Certain kindsof fruit also are eaten.Baird, Brewer, and Ridgway (1905) say that during the summerGambel's quail feeds extensively on the berries of the nightshade.Grinnell, Bryant, and Storer (1918) say:Evidence is also at hand that this quail, like many other desert animals,feeds upon the fruit and seeds of certain kinds of cactus. Stomachs of Gam-bel Quail collected along the Colorado River in the spring of 1910 containedmasses of mistletoe berries, and, at the time the mesquites were first cominginto leaf, quantities of the tender green foliage of this plant.It has been said that these quail can not exist very far from water,to which they have to resort twice a day to drink, but Dr. RobertC. Murphy (1917) found them abundant at all hours about his campin the Colorado Desert, which " was upwards of 20 miles from theriver, 7 miles from the miserable hole of the Tres Pozos, 10 milesfrom the Laguna Salada, and an equal distance from the nearestmountain 'tinaja.' The soil was ever}'where sandy and porous; nota suggestion of moisture was to be detected even in the beds of thedeepest barrancas." He also says : " The crops of the specimenstaken early in April were mostly crammed with caterpillars of thegenus Hemileuca, assorted sizes of which were at that time march-ing in legions across the desert."Behavior.?Gambel's quail is not so persistent a runner as thescaled quail, but it is quite reluctant to fly and prefers to escape byrunning very swiftly away among the underbrush. It does not oftensquat and hide, as our bobwhite does, for this would be a poor wayto escape from its natural enemies. When it is forced to fly its flightis swift and strong and often protracted to a long distance in theopen; in thick brush, which it largely frequents, it flutters ratherawkwardly away for a short distance; flocks separated in this waysoon begin to call and gather again. We found it not particularlyshy, especially among the mesquite thickets, where it probably feltthat it was not observed. About the ranches and farmhouses itbecomes very tame, often feeding with the domestic poultry. Dur-ing the heat of the day it rests quietly in the scanty shade of themesquites, or under denser thickets, or even on the branches of leafytrees. It comes out to feed and drink early in the morning andtoward night. It takes to the trees readily at any time and prob-ably roosts in trees at night. When a flock of Gambel's quail isfeeding there is usually a sentinel on guard.Voice.?Cassin (1856), quoting from Colonel McCall's notes, saysof the voice of the male in June
:
A very good idea may be formed of his cry by slowly pronouncing, in a lowtone, the syllables " kaa-wale," " kaa-wale." These notes when uttered close
gambel's quail 81
at hand, are by no means loud ; yet it is perfectly astonishing to what a distancethey may be heard when the day is calm and still. There was to me somethingextremely plaintive in this simple love-song, which I heard for the first timeduring a day of burning heat passed upon the desert.Again he writes:Later in the season, when a covey is dispersed, the cry for assembling is
" qua-el," " qua-el." The voice at all seasons bears much resemblance to thatof the California Partridge?having, in its intonation, no similarity to thewhistle of the Virginia or common partridge.Bendire (1892) gives the mating call of the male as " yuk-kae-ja,yuk-kae-ja, each syllable distinctly articulated and the last two some-what drawn out." Another note, given while moving about incoveys, "resembled the grunting of a sucking pig more than any-thing else, and it is rather difficult to reproduce the exact sound inprint. Any of the following syllables resembles it, quoit, oit, woet,uttered rapidly but in a low tone. The alarm note is a sharp, dis-cordant craer, craer, several times rapidly repeated, and is usuallyuttered by the entire covey almost simultaneously."Dr. Joseph Grinnell (1904) quotes from Joseph Mailliard's notes,as follows:The notes of the desert quail differ from those of the valley quail in variety,and to a certain extent in character, though they have some notes in common.The " crow " of the latter consists of three notes, varying in length and accentaccording to the call given, in one case the last note being a falling one. The
" crow " of the desert quail, while rather similar to the other, has two addi-tional notes at the end, rendered in a softer tone. Besides the alarm calls thevalley quail has a few twittering or conversational notes, while the other speciesbas a lot of these, quite varied and often given in a way that seems remarkablyloud to one accustomed only to the notes of the former. Another peculiarity ofthe desert quail is the queer sound that it makes as it rises from the ground onbeing surprised into flight?the sort of screeching cackle, on a small scale, thata hen makes when frightened from her nest.Game.?Although Gambel's quail is a plump and delicious morselfor the table, it is an exasperating bird to hunt. It loves the thickestand thorniest cover and frequents the roughest and hardest country,through which it runs, and keeps on running, faster than a man canfollow; often it will take refuge in a rocky creek bed or canyon,where it is hopeless to follow. What few I have shot have requiredmore vigorous leg exercise than they were worth and usually had tobe shot on the run. When flushed in the open it flies swiftly andrequires good shooting. The birds will not lie to a dog, so the bestbird dogs are utterly useless in hunting them, except as retrievers.In past years these quail were an important item in the markethunter's game bag. Herbert Brown (1900) was informed by an
82 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
express agent that 3,000 dozen quail were shipped out of Salt RiverValley in 1889 and 1890. He says further
:
The Mohawk valley, in Yuma county, is probably the most prolific breedingspot in the territory. It was, at one time, a favorite place for trappers andpot-hunters, and it was not until the game law had been amended that theirnefarious practices were broken up. In six weeks, in the fall of 1894, no lessthan 1,300 dozens were shipped to San Francisco and other California markets.The price at first realized, so I was told by the shippers, was $1.12% per dozen,but later GO cents only were realized. The Quail were trapped, their throats cut,then sacked and shipped by express. I was told by one of the parties soengaged that he and his partner caught 77 dozens in one day. They used eighttraps and baited with barley. Their largest catch in one trap, at one time, was11 dozens. At the meeting of the next legislature the game law was againamended, and it was made a misdemeanor to trap, snare, or ship Quail orPartridges from the Territory. This effectually stopped the merciless slaughterof the gamiest bird in Arizona?Gambel's Partridge.Enemies.?Coues (1874) -writes:Man is, I suppose, the Quail's worst enemy; what the White does with dogand gun the Red accomplishes with ingenious snares. The Indians take greatnumbers alive in this way, for food or to trade with the whites along theColorado ; and they use the crests for a variety of purposes that they considerornamental. I saw a squaw once who had at least a hundred of them strung ona piece of rope-yarn for a necklace. But the birds have other foes ; the largerHawks prey upon them, so also do the wolves, as I have had good evidence uponone occasion, when hunting in a precipitous, rocky place near Fort Whipple.I heard a covey whispering about me as they started to run off in the weeds, andfollowed them up to get a shot. They passed around a huge boulder thatprojected from the hill-side, and then, to my surprise, suddenly scattered onwing in every direction, some flying almost in my face. At the same instanta wolf leaped up from the grass, where he had been hiding, a few feet off,intending to waylay the covey, and looking very much disappointed, not to saydisgusted, at the sudden flight.The quail have numerous other enemies. Coyotes, foxes, wildcats,and various hawks and owls kill the old birds and young; even thelittle pigmy owl has been known to kill an adult Gambel's quail.Skunks, rats, rock squirrels, snakes, Gila monsters, and even landterrapins eat the eggs. Fortunately these quail are prolific breeders,so they are not exterminated.DISTRIBUTIONRange.?Southwestern United States and northern Mexico. Therange of Gambel's quail extends north to southern California (rarelyLos Angeles, Hesperia, and Daggett) ; southern Nevada (AshMeadows and Pahranagat Valley) ; southern Utah (Hamblin, Har-mony, and Fruita) ; Arizona (Cedar Ridge, Roosevelt, NantanPlateau, and Blue); and central Texas (Eagle Springs). Eastto southwestern New Mexico (Socorro and Las Cruces) ; central
gambel's quail 83Texas (Eagle Springs and Fort Clark) ; and southeastern Sonora(Camoa). South to southern Sonora (Camoa and Guaymas) ; andnorthern Lower California (Laguna Salada). West to northernLower California (Laguna Salada and Signal Mountain) ; andsouthern California (Calexico, Pelican Lake, Agua Dulce, PalmSprings, and rarely Los Angeles). Some of the northern localities,such as those in Utah, may possibly be the result of introductionsin contiguous areas. Bryant (1889) states that " a few pairs withsmall young were seen " on the western side of Lower Californiaat about latitude 30? N., but it is likely that the birds seen wereSan Quintin quail (L. c. plumbea). There is one record of the oc-currence of gambeli at San Diego, Calif., where a female was found(1924) mated with a male valley quail (L. c. vallicola).It is difficult to outline the natural range of this quail, since it hasbeen transplanted extensively. For example, they were found rathercommonly at Furnace Creek, Death Valley, having been introducedthere from Resting Springs, Calif., by the borax company. Theyare easily trapped and for this reason probably have been favoritesin many ill-advised projects. In the early nineties they were intro-duced into Massachusetts (Marthas Vineyard). Other eastern ex-periments were made in Pennsylvania and Kentucky. Naturally,all these were failures, as were also introductions in northern Cali-fornia and Washington.They were, however, successfully acclimated on San ClementeIsland, Calif., and there have been several successful introductionswell outside the normal range in Arizona (Snowflake, Holbrook,Vernon, and Colfax Counties) and New Mexico (Cortez, Gallup,San Juan Valley, and Huntington). A successful introduction alsois reported from Montana (Blue Creek, near Billings). The mostremarkable achievement, however, is the transplantation of thesebirds to western Colorado. About 1,000 birds from southern Cali-fornia (which locality gave rise to the name " California quail,"under which they have appeared in the literature on Colorado birds)were liberated at Montrose in 1885 or 1889. From that point thebirds have increased and spread over the Uncompahgre Plateauand the valleys of the Grand and Gunnison Rivers. Because ofplumage changes that have taken place since the introduction, theyhave been described as a subspecies (L. g. sanus). There is no defi-nite record of the natural occurrence of Gambel's quail anywherein Colorado.Egg dates.?Arizona: 68 records, March 19 to September 20; 34records, April 20 to May 29. California: 8 records, March 26 toJune 2.
84 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMLOPHORTYX GAMBELI SANUS MearnsOLATHE QUAILHABITSThe Gambel's quail of southwestern Colorado has been describedby Dr. Edgar A. Mearns (1914) as "rather larger than the average"of Arizona birds, from which "it differs in coloration as follows:Adult male with upper parts neutral gray (Ridgway, 1912), un-washed with olive; crown chestnut-brown instead of hazel; chest-patch cartridge buff instead of warm buff or chamois. Adult femalewith upper parts as in the male, differing from gambelii and fulvi-pectus in having the crown darker (sepia instead of cinnamondrab) ; chin and throat darker and more grayish; chest and abdomenpale olive-buff instead of cream color."The type specimen came from Montrose County, but it seems tobe a rare bird even there. Its habits are probably no different fromthose of the species elsewhere.CYRTONYX MONTEZUMAE MEARNSI NelsonMEARNS'S QUAILHABITSOne of the handsomest and certainly the most oddly marked of theNorth American quails presents a bizarre appearance when closelyexamined ; one look at its conspicuously marked face would brand itas a clown among birds; its dark-colored breast is contrary to thelaws of protective coloration and would make it very conspicuouson open ground. But when one tries to find it in its native haunts,squatting close to the ground among thick underbrush, weeds, andgrass, one realizes that its dark belly and spotted flanks are completelyconcealed, that the grotesquely painted face becomes obliteratedamong the sharp lights and shadows, and that the prettily markedback matches its surroundings so well that the bird is nearly invisible.Mearns's quail has been called the " fool quail " because it haslearned to trust this wonderful protective coloration and lie close,rather than trust to its legs or its wings to escape. How successfulit has been nobody knows. The only two I ever saw I almost steppedon before they flew, and I wonder how man}' more I walked nearwithout knowing it. I question whether it is as much of a fool as itis said to be. It lives in different haunts and has developed differenthabits from its neighbors, the scaled and Gambel's quails. Its shapeand carriage, its white-spotted sides, and its habit of clucking as itwalks or feeds have suggested a possible relationship with the guineafowls.
MEARNS'S QUAIL 85The haunts of Mearns's quail are generally far removed from thehabitations of man. Major Bendire (1892) quotes William Lloyd assaying that, in Texas, " the favorite resorts of the Massena Partridgeare the rocky ravines or arroyas that head well up in the mountains.They quickly, however, adapt themselves to changed conditions oflife and are now to be seen around the ranches picking up grain andscratching in the fields. In the vicinity of Fort Davis, Texas, theyhave been exceptionally numerous and ma}7 frequently be seen sittingon the stone walls surrounding grainfields in Limpia Canon."In Arizona we found them in the lower parts of the canyons andin the foothills of the Huachucas and the Chiricahua Mountains,where the ground was rough and more or less rocky, with tall tuftsof grass, low bushes, scattered mescals, and small oaks. They rangeup the sides of rocky ravines and into the mountains up to 9,000feet in summer and are seldom found below 4,000 feet. In ApacheCounty, according to Major Bendire (1892), "the favorite localitiesfrequented by this species during the breeding season are thicklive-oak scrub and patches of rank grass, at an altitude of from7,000 to 9,000 feet. Here they are summer residents only, descend-ing to much lower altitudes in winter." Henry W. Henshaw (1874)writes : " This beautiful partridge is quite a common resident inthe White Mountains, near Apache, Ariz., where, in summer, it seemsto shun the open valleys, and keeps in the open pine-woods, evincinga strong preference for the roughest, rockiest localities, where itsstout feet and long, curved, strong claws are admirably adapted toenable it to move with ease."Nesting.?Major Bendire (1892) refers to two nests described asslight hollows, one under a small shin-oak bush, the other alongsidea sotol plant. He quotes descriptions of two other nests quite fully.Otho C. Poling wrote
:
I was climbing up a steep mountain side on the northeast of the HuachucaMountains, some 10 miles north of the border, when, at an elevation of aboutS,000 feet, I flushed the female almost directly under my feet and shot it.The hillside was covered in places with patches of pines and aspens, as wellas with low bushes and grasses. The nest was directly under a dead limbwhich was grown over with dead grass, and so completely hidden that untilI had removed the limb and some of the grass it was not discernible at all.The nest was sunken in the ground, and composed of small grass stems, archedover, and the bird could only enter it by a long tunnel leading to it from underthe limb and the grass growing around it. The eggs were eight in numberand naturally white, but they were badly stained by the damp ground, theircolor being now a brownish white. They were almost hatched. The femalemust have remained on them all the time to have caused such uniform incuba-tion and preserved the eggs from spoiling by the excessive dampness.74564?32 7
86 BULLETIN 162, UNITED STATES NATIONAL MUSEUMG. W. Todd's notes state
:
The only nest of this species I have ever seen was situated under the edgeof a big bunch of a coarse species of grass, known as "hickory grass." Thisgrass grows out from the center and hangs over on all sides until the bladestouch the ground. It is a round, hard-stemmed grass, and only grows on themost sterile soil. According to my observations the Massena partridge is seldomseen in other localities than where this grass grows. I was riding at a walkup the slope of a barren hill when my horse almost stepped on a nest, touchingjust the rim of it. The bird gave a startled flutter, alighting again within 3feet of the nest and not over 6 feet from me ; thence she walked away withher crest slightly erected, uttering a low chuckling whistle until lost to viewbehind a Spanish bayonet plant (yucca), about 30 feet off. I was riding arather unruly horse and had to return about 30 yards to tie him to a yuccabefore I could examine the nest. This was placed in a slight depression,possibly dug out by some animal, the top of the nest being on a level withthe earth around it. It was well lined with fine stalks of wire grass almostexclusively, the cavity being about 5 inches in diameter and 2 inches deep.At the back, next to the grass, it was slightly arched over, and the overhangingblades of grass hid it entirely from sight. The nest was more carefullymade than the average bobwhite's nest and very nicely concealed.There is a set of 13 eggs in Col. John E. Thayer's collection, takenby Virgil W. Owen in Cochise County, Ariz., June 18, 1905. Thenest is described as " a slight depression under a bunch of saw-grass,which was growing on a hillside. It was quite compactly built ofstraws, grass, and leaves, with a tubular entrance extending out aboutsix inches from the nest. The whole nest was roofed over and wellconcealed by overhanging grass."Eggs.?Mearns's quail is known to lay from 8 to 14 eggs. Thesevary in shape from ovate to short ovate or ovate pyriform, usuallymore elongated than those of the bobwhite. The shell is smooth andsomewhat glossy. They are pure white, dull white, or creamy whiteand unmarked, but often much nest stained. The measurements of39 eggs average 31.9 by 24.7 millimeters; the eggs showing thefour extremes measure 33 by 24.4, 32.3 by 27.3, 30.5 by 24.6, and31.5 by 23 millimeters.Young.?The period of incubation seems to be unknown. Bothsexes share in the incubation and in the care of the young. FrankC. Willard, who has seen many nests, tells me thatin about half of the nests examined the male was on the eggs. In two in-stances both birds were at the nest. In one which I went to see on August 17,1913, the male was sitting at the entrance with a newly hatched chick pokingits head out from under his wing. The female was in the nest, which was wellarched over. Around her was a row of little ones, and one was sitting on herback. This charming picture lasted but a few seconds. The two old birdsfluttered away, pretending disability to fly. The scarcely dry young could notwalk, but crawled away with astonishing rapidity.
MEARNS'S QUAIL 87Henshaw (1874) describes the following exhibition of parentalsolicitude
:
August 10, while riding with a party through a tract of piny woods, a broodof 8 or 10 young, accompanied by the female, was discovered. The young,though but about a week old, rose up almost from between the feet of the fore-most mule, and after flying a few yards dropped down, and in a twinkling werehidden beneath the herbage. At the moment of discovery, the parent bird roseup, and then, tumbling back helplessly to the ground, imitated so successfullythe actions of a wounded and disabled bird that, for a moment, I thought shemust have been trodden upon by one of the mules. Several of the men, com-pletely deceived, attempted to catch her, when she gradually fluttered off,keeping all the time just beyond the reach of their hands, till she had enticedthem a dozen yards away, when she rose and was off like a bullet, much totheir amazement.Plumages.?In the newly hatched chick the upper parts are varie-gated with " cinnamon-buff," " hazel," " chestnut," and black, mostlyhazel and chestnut, fading out to white on the chin and throat andto grayish white on the belly ; there is a broad band or patch on thecrown, of " hazel," bordered with chestnut, and an auricular stripeof blackish brown.The juvenal plumage is acquired at an early stage. A smallyoung bird, only about 2!/2 inches long, is nearly covered above withjuvenal plumage; the wings reach beyond the tail, which has notyet started ; the sides of the breast and flanks are feathered ; but thehead, neck, and center of the breast are still downy. This youngbird could probably fly.In this plumage the sexes are much alike, except that in the youngmale the crissum, lower belly, and flanks are black, and the centerof the breast is suffused with brown, whereas in the young femalethese parts are whitish; these characters are conspicuous in flight.In both sexes the crown is " hazel " or " russet," spotted with blackand with some whitish shaft streaks ; the sides of the head are buffywhite, mottled with black and with a dark brown auricular patch;the mantle is " tawny " to " ochraceous-tawny," barred heavily withblack and with broad buffy median stripes; the wing coverts areashy with rounded black spots; the primaries and secondaries arebanded with white spots; the throat is white, and the rest of theunderparts are pale buff, or grayish white, barred or spotted withblack and white.The juvenal plumage is worn for only a short time, for the post-juvenal molt begins early in September and is prolonged in someindividuals through November. This molt is complete except forthe outer two primaries on each wing, which are retained during thefirst year. The sexes now begin to differentiate rapidly, the brilliantbody plumage of the male and the " vinaceous-pink " breast of thefemale replacing the juvenal plumage.
88 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe molt starts on the upper breast, flanks, and shoulders, workinggradually downward ; the rich chestnut in the center of the breastof the male is the last of the body plumage to be acquired; but theentire body molt is completed before any change takes place on thehead; the conspicuous head markings are not assumed in the youngmale until December or later. A very limited prenuptial molt takesplace in spring and a complete postnuptial molt late in summer andin fall, after which young birds are fully adult.Food.?Mrs. Bailey (1928) summarizes the food of Mearns's quailas follows:As far as known, lily bulbs
?
% of tbe food in 5 specimens and to judgefrom their large strong digging feet provided with sharp claws perhaps theprincipal article of their diet?also great numbers of acorns and pinyon nuts,and in addition seeds and spines of prickly pear, acacia, seeds of legumes andspurges, grass blades, berries of mountain laurel, arbutus, and cedar, and suchinsects as weevils, caterpillars, bugs, crickets, and grasshoppers.A pair that Mr. Bailey started at the head of the Mimbres at about 8,000feet had been scratching under the pine trees. " In the freshly scratchedground," he says, " I found a quantity of membranacious shells of a little bulb
?
probably Cyperus?and several of the bulbs. I ate one of these and found itgood, starchy, juicy, crisp, and of a nutty flavor. The Quail had dug two orthree inches deep in the hard ground and seemed to find plenty of bulbs, but Icould not find one by digging new ground, nor could I find the plant whichbore them.Behavior.?Mearns's quail is a gentle, retiring bird of rather sed-entary habits. It prefers to walk about slowly and quietly amongthe rocks, bushes, and clumps of grass on the rough hillsides whereit lives. If alarmed, it squats and freezes, immovable, until almosttrodden upon or touched, when it rises from almost under foot, fliesa short distance, drops into cover and squats again. When greattyalarmed it sometimes flies to a great distance in a very swift anddirect flight. Several observers have mentioned coming upon oneor more of these birds in the mountain roads, where they are fondof dusting; they showed no alarm, either walking away quietly orsquatting and freezing. Captain William L. Carpenter says in hisnotes
:
I once slopped my horse when about to step on one and watched it for sometime without creating alarm. After admiring it for several moments squattingclose to the ground within a yard of the horse, watching me intently, butapparently without fear, I dismounted and almost caught it with my hat, fromunder which it fluttered away.Henshaw witnessed a remarkable exhibition of the confidence thatthis bird shows in its protective coloration, for he says in some notessent to Mrs. Bailey (1928) :Of the several quail known to me the " fool quail " of New Mexico andArizona seems to depend for his safety upon his protective coloration more
MEARNS'S QUAIL 89than any other. As an example I recall one that squatted on a log near thetrail our pack train was following, and so closely did the colors of his backand sides harmonize with his surroundings that 12 or 15 pack mules and horse-men passed by him without seeing him or disturbing his equanimity in theleast. He seemed so completely petrified by astonishment at the novel sightas to be incapable of motion, and he was so close to vis that one might havetouched him with a riding whip. While the bird was no doubt actuated to someextent by curiosity, be depended for his safety, I am sure, upon the nice wayin which his plumage matched his surroundings and upon his absolute immo-bility. No one saw the bird but myself, and when the train had passed I hadto almost poke him off his perch before he consented to fly. Whoso calls thisthe " Fool Quail " writes himself down a bigger fool than the bird, who hasbeen taught his lesson of concealment by Mother Nature herself.Louis A. Fuertes (1903) thus describes his first impression of aMearns's quail:I awoke in the cool, just before sun-up, and was lazily dressing, half out ofmy sleeping bag, when my sleepy eye caught a slight motion in the grass about20 feet away. I looked and became aware that I was staring at my firstMearns quail. Even as I took in the fact, he apparently framed up his ideasas to his vision, and telling himself in a quiet little quail voice that it wereperhaps as well to move on and look from a safer distance, he slimmed downhis trim little form and ran a few steps. Meanwhile I was clumsily trying toget my gun cut from under my sleeping bag, where I had put it to keep it outof the dew. The quail, getting wiser every second, doubled his trot, and withhead erect and body trim ran like a plover for a few yards through the shortdesert grass, and with a true quail f-r-r-r-r-r-r-r-r burst into flight and droppedinto the thick brush across the arroyo. The most noticeable thing about himas I watched him running was the curious use of his queer little crest. Insteadof elevating it as the mountain quail does his, he raised his painted head on slimneck and spread his flowing crest laterally, till it looked like half a mushroom,giving him the most curious appearance imaginable. When he flew I markedhim clown carefully, hastily drew my boots half on, grabbed my gun andstumped after him with all speed. I got to his point within a short time, butthrash and kick around as I might, I never succeeded in making him flush asecond time.Mr. Willard writes to me
:
One morning, as I arrived in front of our store in Tombstone, I found aflock of a dozen or more of these birds running around in the street. Most ofthem flew up onto the roof of the building, but one male ran into the doorway,stuck his head down into a corner, and waited for me to pick him up.Voice.?Major Bendire (1892) quotes Mr. Todd as saying:When scared they utter a kind of whistling sound, a curious combinationbetween a chuckle and a whistle, and while flying they make a noise a gooddeal like a Prairie Hen, though softer and less loud, like " chuc-chuc-chuc,"rapidly repeated.H. S. Swarth (1909) writes:Their call consists of a series of notes slowly descending the scale, and endingin a long, low trill, the whole being ventriloquial in effect and most difficult to
90 BULLETIN 162, UNITED STATES NATIONAL MUSEUMlocate. It is easily imitated, however, and the birds readily answer when onewhistles ; when the flock is scattered they will sometimes even return, calling;it intervals as they approach. The only other note I have heard is a quaveringwhistle uttered as they take flight.Mrs. Bailey (1928) says thatthe low call of the Mearns Quail, suggestive of the quavering cry of a ScreechOwl, adds to the fascination of the pursuit of this illusory bird, for it is ventrilo-quial in quality and leads you such a fruitless chase that you return to campwith an exaggerated interest in this feathered Will-o'-the-wisp.Fall.?In some notes sent to Major Bendire (1892) Doctor Nelsonstates thatthe birds breeding along the northern limit of their habitat migrate southwardin October*. In southern Arizona the same result of a warmer winter climateis obtained by descending the flanks of the mountains. The summer range ofthis species is just above and bordering that of Gambel's Quail in parts ofArizona and New Mexico. The fact that Gambel's Quail changes its range butlittle in winter results in these birds being found very frequently occupyingthe same ground at this season. I have never seen the Massena Partridgein coveys larger than would be attributed to a pair of adults with a small broodof young. Frequently a pair raise but three or four, and I do not rememberhaving ever seen more than six or seven of these birds in a covey.DISTRIBUTIONRange.?Southwestern United States (except California) andnorthern Mexico. The range of Mearns's quail is extremely circum-scribed. It extends north to Arizona (Fort Whipple, Camp Verde,Mogollon Ridge, Wilcox, and Marsh Lake) ; New Mexico (ZuniMountains, San Mateo Mountains, White Mountains, and GuadalupeMountains) ; and central Texas (San Angelo and Mason). East toTexas (Mason, Kerrville, Bandera Hills, and San Antonio)
;
Durango (Ramos, El Salto, and Huasamota) ; and Nayarit (SanBlasito). South to Nayarit (San Blasito). West to Nayarit (SanBlasito) ; western Chihuahua (50 miles northeast of Choix, Sinaloa)central Sonora (La Chumata and Patagonia Mountains) ; and Ari-zona (Baboquivari Mountains, Rincon Mountains, Mount Turnbull,and Fort Whipple).The species has a vertical range from 4,000 to 9,000 feet, the birdsmoving to the higher altitudes early in fall and retiring upon theapproach of winter. They apparently do not, however, descendbelow the lowest parts of the breeding range.Egg dates.?New Mexico, Arizona, and Mexico : 29 records, April24 to September 5; 15 records, June 23 to August 1G.
DUSKY GROUSE 91Family TETRAONIDAE, GrouseDENDRAGAPUS OBSCURUS OBSCURUS (Say)DUSKY GROUSEHABITSThe big " blue grouse " of the southern Rocky Mountains rangesfrom southern Idaho and northern Colorado to Arizona and NewMexico. Two other subspecies of obscurus occupy the more northernparts of the Rocky Mountains. Its home is in the mountains fromabout 2,000 feet in the foothills, or as low as quaking aspens grow,up to timber line at 11,000 or 12,000 feet. It was formerly verycommon in the coniferous forests, but it has now disappeared frommany of its former haunts and is becoming scarce in others. Coues(1874) quotes from Mr. Trippe's notes, regarding its haunts inColorado, as follows:The " Gray Grouse," as this species is universally called, is a rather commonbird throughout the mountains, from the foot-hills up to timberline, aud, duringsummer, wanders at times above the woods as high as the summit of the range.Excepting for a brief period in August and September, it rarely approaches thevicinity of clearings, frequenting the dense pine forests, and showing a pref-erence for the tops of rocky and inaccessible mountains. In its nature, in short,it is the exact counterpart of the Ruffed Grouse, having the same roving, restlessdisposition ; living upon the same diet of buds and berries ; frequenting thesame rugged, craggy mountain haunts; and, like that bird, is more or lesssolitary in its habits, and constantly moving from place to place on foot.Mrs. Florence M. Bailey (1928) writes of it in New Mexico:In the depths of the coniferous forest you may suddenly discover a DuskyGrouse with its small pointed head and henlike body sitting quietly on a logfacing you, as if secure in its disguise?a dusky bird in the dusky woods sur-rounded by shadowy tree trunks. From a forest trail you may flush one thathas been dusting itself in the soft earth, or hearing a muffled ventriloquialhooting may creep up within sight of the lordly cock at the foot of a conifer,with purplish red neck pouches dilated. Again in the open, you may bestartled by a loud whir and look up to see great dark forms with a wide spreadof wing disappearing over your head ; or, on a steep mountain side, catch sightof a big Grouse sailing off below you with stiff outstretched wings and a spreadtail, whose gray band makes a striking mark to follow among the branches.If still more fortunate, you may surprise a family in a mountain meadow, forstrawberries are evidently one of their favorite summer foods.Courtship.?Bendire (1892) quotes from some notes received fromDenis Gale, as follows
:
During the mating season if you are anywhere near the haunts of a pair youwill surely hear the male and most likely see him. He may interview you onfoot, strutting along before you, in short hurried tacks alternating from rightto left, with widespread tail tipped forward, head drawn in and back andwings dragging along the ground, much in the style of a turkey gobbler. At
92 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
other times you may hear his mimic thunder overhead again and again in hisflight from tree to tree. As you walk along he leads, and this reeonnoiteringon his part, if you are not familiar with it, may cause you to suppose that thetrees are alive with these Grouse. He then takes his stand upon a rock, stump,or log, and in the manner already described distends the lower part of his neck,opens his frill of white, edged with the darker feather tips, showing in itscenter a pink narrow line describing somewhat the segment of a circle, thenwith very little apparent motion he performs his growling or groaning, I don'tknow which to call it, having the strange peculiarity of seeming quite distantwhen quite near, and near when distant ; in fact, appearing to come from everydirection but the true one. The first time I heard the sound I concluded itwas the distant laboring of one of our small mountain sawmills wrestling inagony with some cross-grained saw-log.Nesting.?I have never seen a nest of this grouse, and very littleseems to have been published about it. W. L. Sclater (1912) says:
" The nest, which has been described by Henshaw and Burnett, isplaced on the ground, generally in an open glade but sheltered orsomewhat concealed; that found by Burnett was placed in a hollowunder two old logs, it being a simple structure of dried grass or pineneedles." Mrs. Bailey (1928) says that the nest is "a shallow de-pression beside a log or under grass or bushes, slightly lined with afew pine needles or a little grass."Bendire (1892) describes a nest found by Henshaw, as follows:A nest found June 16 contained seven eggs on the point of hatching. Thenesting site was a peculiar one, being in an open glade, where the grass hadbeen recently burned off. The nest proper was a slight collection of driedgrass placed in a depression between two tussocks, there apparently havingbeen no attempt made at concealment.Eggs.?From 7 to 10 eggs are usually laid by the dusky grouse,and as many as 12 have been found in a nest. Bendire (1892) saysthat " an egg is deposited daily, and incubation does not commencetill the set is completed." The eggs are ovate to elongate ovate inshape. The shell is smooth, with little or no gloss. The groundcolors vary from " pinkish buff " to " pale pinkish buff " or to " car-tridge buff." They are usually evenly covered with very smallspots or minute dots, generally quite thickly; some eggs are lessthickly and more irregularly covered with somewhat larger spots;very rarely an egg is immaculate. The colors of the spots are " sayalbrown," " clay color," or " cinnamon-brown," rarely darker. Themeasurements of 54 eggs average 49.7 by 34.9 millimeters; the eggsshowing the four extremes measure 52.3 by 34.4, 50.6 by 37.1, 46.2by 34, and 50 by 31.8 millimeters.Young.?Bendire (1892) quotes Denis Gale as follows:Upon one occasion I met with a covey which had just been hatched ; theywere quite nimble, and with the exception of one which I caught they hid them-selves with great address. Until I released the little prisoner the femaleshowed great distress, clucking in the most beseeching manner, accompanied
DUSKY GROUSE 93
with suitable gestures, similar to but more tender and graceful than those ofour domestic hen. She stood within 6 or 7 feet of me pleading her cause andeasily won it. In her beautiful summer dress of brown, handsomely plumedas she was, she looked very interesting.In a single instance only, with a brood about ten days old, have I noted thepresence of both parents. Perched upon a fallen tree the male seemed to be onthe lookout, while the female and young were feeding close by. This seemingindifference of the male while the brood is very young, allowing his mate toprotect them, if he really is always near at hand, looks very strange, and yetit may be the care, since he is generally with the covey when the young are wellgrown. Directly the young are able to travel, the hen Grouse leads them tosome desirable opening, skirting the timber or gulch, where bearberries, wildraspberries, gooseberries, and currants, as well as grasshoppers, worms, andgrubs are abundant, managing them just as the domestic hen does her brood.The young grow rapidly, and when about two weeks old can do a little withtheir wings ; then, instead of hiding on the ground, they flush and endeavor toconceal themselves in the standing timber. Until almost fully grown they arevery foolish ; flushed, they will tree at once, in the silly belief that they are outof danger, and will quietly suffer themselves to be pelted with clubs and stonestill they are struck down one after another. With a shotgun, of course thewhole covey is bagged without much trouble, and as they are, in my opinion,the most delicious of all Grouse for the table, they are gathered up unsparingly.Plumages.?The sequence of molts and plumages is the same as inthe sooty grouse, of which I have found more material for study.The racial characters are apparent in immature birds. Kichardson'sgrouse has been known to hybridize with the Columbian sharp -tailedgrouse (Allan Brooks, 1907).Food.?Mrs. Bailey (1928) summarizes the food of the duskygrouse very well, as follows
:
In 45 crops and stomachs examined, the food consists of 6.73 per cent animalmatter?5.73 per cent grasshoppers, and the rest beetles, ants, and caterpillars
?
and 93.27 per cent vegetable matter?seeds, fruits, and leaves, coniferous foliageamounting to 54.02 per cent, fruits 20.09 per cent, including manzanita berries,mountain ash, service-berry, currant, gooseberry, etc. One cock shot at 11,600feet on Pecos Baldy in a strawberry patch had both crop and gizzard filledmainly with strawberries. The crop of another shot between 8,000 and 9,000feet contained 27 strawberries, 28 bear-berries, 12 Canadian buffalo-berries,flowers of Indian paint brush and milk vetch, leaves of vetch and buffalo-berry,and a few ants and caterpillars, while its gizzard was filled with seeds of bear-berry, Canadian buffalo-berry, and strawberry, a few green leaves, and a numberof ants, beetles, and other insects. Grasshoppers, the green leaves of blue-berryand vetch, salal, and other berry seeds, needles of Douglas spruce and fir,together with gravel and hard quartz grinding stones were among the itemsthat the field examination of other specimens revealed. The quartz grindingstones were found in gizzards apparently filled with hard coniferous needles.These needles seem to be the regular winter food as under a winter roostingtree on Pecos Baldy the winter dung was composed entirely of spruce needles.(Three birds taken in September near Golden, Colorado, had their " cropscrammed with the berries of kinnikinick.")
94 BULLETIN 162, UNITED STATES NATIONAL MUSEUMBehavior.?Coues (1874) quotes from Mr. Trippe's notes, as fol-lows:On being suddenly startled, this bird takes wing with great rapidity, some-times uttering a loud cackling note, very much like that of the Prairie Hen onsimilar occasions, frequently alighting on the lower limb of a tree after flying alittle way, and watching the intruder with outstretched neck. Sometimes theywill fly up to the top of a tall pine and remain hidden in the thick foliage for along time ; nor will they move or betray their position, although sticks andstones are thrown into the tree, or even a shot fired. Late in summer manyof them ascend to the upper woods to feed upon the multitudes of grasshoppersthat swarm there in August and September, in the pursuit of which they wanderabove timber-line, and may sometimes be met in great numbers among thecopses of willows and juniper that lie above the forests.Edwyn Sandys (1904) says that this grouseis most difficult to locate even when perched upon a limb only a few yards away.In its native woods the light is baffling and there is a confusion of shade, amidwhich the general slaty tone of the plumage is barely distinguishable. A coatof feathers especially designed with a view to protective coloration could notbetter serve the purpose, and the bird appears to be perfectly aware of this.Indeed, its habit of trusting to its trick of treeing and remaining motionlesshas earned for it the names of " fool-grouse," which I believe should be appliedonly to young birds. These unquestionably will tree and foolishly maintaintheir positions while their comrades are being shot or clubbed down, but theolder birds, except in seldom disturbed localities, are wiser.But fool grouse or no, when once the bird concludes to start there is no morefoolishness. With a nerve-shaking whirring it promptly gets to top speed, andusually darts downhill, a maneuver which greatly adds to the difficulty of theshot. When taking wing it cackles like a scared fowl.Game.?Referring to it as game bird, Sandys (1904) writes:Among western sportsmen it is termed the " blue," or " gray," grouse, andthose who have enjoyed the pleasure of shooting and later eating it have yetto be heard from in the line of adverse criticism. Its sole fault as a game birdconsists in its seldom being found in cover which affords a fair chance to thegun. In fact, it is such an inveterate lover of trees that it takes to the branchesas naturally as a duck takes to water. Like the ruffed grouse, it will tree, andremain motionless until it fancies it has been observed ; then it at once departswith a sounding rush, which may only be stopped by the quickest and mostskilled of shots. I have flushed it when it seemed to do hardly anything morethan leap from the ground to a convenient limb, and more than once, whileseeking to trim off its head, it has left the perch so suddenly that the gun couldnot be shifted in time to prevent the wasting of a shell?and this little joke atthe expense of a notoriously quick shot.Only those familiar with the western cover can understand how easy it isto fail to bag at short range a bird about as large as a common barnyard hen
?
to be accurate, of between three and three and one-half pounds' weight. Thetenderfoot would imagine such a bird, rising close at hand, to be an easy, per-haps too easy, mark. Let the tenderfoot climb the steeps and try a few bluegrouse as they leave the trees, and his song may take on an undertone sugges-tive of blasted hopes and trust betrayed. In the first place, the cover usuallyis standing timber big enough to stop a locomotive, to say nothing of small shot.This timber, as I found it, is about as close as it can stand, thereby forming
DUSKY GROUSE 95
something closely akin to a gigantic stockade with extremely narrow inter-spaces.In spite of the bird's penchant for timber it frequently is found in theopen and in grain fields. In such places the sportsman may enjoy " bluegrouse " shooting as it should be, and sport of a very high order. Then thefull strength and speed of the game becomes apparent, and the man whomakes uniformly good scores has no reason to fear any ordinary company.Fall.?Prof. Wells W. Cooke (1897) says:In August they begin to gather into flocks of 10 to 15 individuals andvisit the grain fields or the more open gulches and foothills for berries. InSeptember they wander above timber-line to feed on grasshoppers, reaching32,500 feet. In winter they come down into the thick woods during the severestweather, but many remain the whole year close to timber-line.DISTRIBUTIONRange.?The Rocky Mountain region of the United States andCanada.The range of the dusky grouse extends north to east-centralYukon (probably the south fork of the Macmillan River) ; andsouthern Mackenzie (Mount Tha-on-tha). East to southernMackenzie (Mount Tha-on-tha) ; Alberta (Stony Plain and 60 mileswest of Calgary) ; Montana (Zortman, Judith Mountains, BigSnowy Mountains, and Fort Custer) ; Wyoming (Sheridan, Trap-pers Creek, Guernsey, Wheatland, Pole Mountain, and Sherman)
;
Colorado (Estes Park, Boulder, Golden, Jefferson, Wet Mountains,Sangre de Cristo Mountains, and Fort Garland) ; and New Mexico(Culebra Mountains, Halls Peak, Pecos Baldy, San Mateo Moun-tains, Eagle Peak, and Mogollon Mountains). South to south-western New Mexico (Mogollon Mountains and San FranciscoMountains) ; Arizona (Blue Mountains, Mount Thomas, Mount Ord,and the Kaibab Plateau) ; and central Nevada (Arc Dome). Westto central Nevada (Arc Dome, Toiyabe Mountains, Monitor Moun-tains, Ruby Valley, Clover Mountains, and East Humboldt Moun-tains) ; eastern Oregon (Fort Harney, Strawberry Mountain, andTurtle Cove) ; eastern Washington (Butte Creek, Hompeg Falls,Okanogan, and Haig Creek) ; eastern British Columbia (Princeton,Nicola, Ashcroft, Bonaparte, Babine Mountains, Nine-mile Moun-tain, Fort Connolly, Groundhog Mountain, Tset-ee-yeh River, Sec-ond South Fork, Doch-da-on Creek, and Telegraph Creek) ; andYukon (Teslin Lake and probably the south fork of the MacmillanRiver).Three subspecies of Dendragapus obscurus are recognized. Trueobscurus is found from central Arizona and southwestern New Mex-ico north to northern Colorado, northern Utah, and southeasternIdaho, extending west to central Nevada. D. o. richardsoni occu-
96 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMpies the range from Wyoming and south-central Idaho north towestern Alberta and central British Columbia and west to easternOregon and Washington. D. o. flemingi is found in northernBritish Columbia, southern Yukon, and southwestern Mackenzie.Although technically nonmigratory, both the dusky and the sootygrouse have the curious habit of performing a postbreeding verticalmigration. The cocks will sometimes move to the higher elevationswhile the hens are still incubating their eggs, usually in July orearly in August. The females and half-grown young follow, so thatby the last of September the species has deserted the breedinggrounds. The winter months are usually spent in the upper spruceforests, the birds living entirely in the trees. By May or June theyhave descended again to their breeding areas.Egg dates.?Colorado and Utah (ohscurus) : 18 records, April17 to July 1 ; 9 records, May 21 to June 13. British Columbia andAlberta to Idaho and Wyoming (richardsoni) : 15 records, May 3to August 20 ; 8 records, May 13 to June 23.DENDRAGAPUS OBSCURUS RICHARDSONI (Douglas)RICHARDSON'S GROUSEHABITSThe name D. o. richardsoni was applied to all the " blue grouse "of the northern Kocky Mountain region until, in 1914, the bird ofthe extreme north was separated under the name flemingi, the nextform. Richardson's grouse is now restricted to the intermediateterritory, from central British Columbia and western Alberta toeastern Oregon, south-central Idaho, and Wyoming. It differs fromthe southern bird, ohscurus, in having no very distinct, terminal,gray band on the tail, and from the northern bird, -flemingi, in be-ing lighter colored. It does not differ materially from either inits habits, except as hereinafter mentioned.M. P. Skinner (1927) writes:Here in the Yellowstone National Park, we are in the borderland occupiedby both the southern and northern forms. It is very difficult to place thesePark birds under either ooscurus or richardsoni, for there are birds of eachform present and all degrees of graduation between them. I once found abird dead near the Buffalo Ranch that had no band at all on the end of itstail. Twice I have seen very dark birds with only a slight amount of graytipping their tails, so that they were more typical of the Sierra Mountainform. Another time, I found a bird so tame that I could approach closeenough to determine positively that he had more than three-quarters of aninch of gray on the ends of the middle tail-feathers. At other times, Ihave found feathers with terminal bands more than a half inch wide. Theredoes not seem to be any segregation of the two forms within the Park, but,
richabdson's grouse 97
so far as I know, the different forms are found in all sections. Both obscurusand richardsoni are larger than Ruffed Grouse, but the Yellowstone specimensare even larger than the average.Richardson's Grouse live in all kinds of brush and forest from the willowsas low as 5,500 feet altitude, through the service-berry (Amelanchier alnifoliaNutt.) areas, aspen groves, Douglas firs, limber pines, lodgepole pines, spruceand white-bark pines to the stunted spruces at timber-line (about 9,500 feet).1 have seen them out on the open plateaus and high meadows above timber-line, almost up to the 10,000-foot level. These timber-line birds were seenfrom July to October, but they apparently went down into the forests duringthe winter. I have seen these grouse in thick-growing lodgepole saplings wherethe small trees were so thick it appeared impossible for the grouse to walk orfly through them ; and I have also seen them out in burned forests, especiallywhere there were berries to be had.While Richardson's Grouse prefer the forests, I have found them out in thesage-brush (Artemisia tridentata Nutt.), sometimes as much as five hundredyards from the nearest tree. Usually, they are wild in such localities, andsoon fly to the nearest trees for protection ; but on May 31, 1921, one wasfound hiding under a two-foot sage-bush in the open. I have also seen singlebirds in open grasslands without brush and as much as two hundred yardsfrom the nearest tree ; and once I rode my horse past one in full view in thegrass and within twenty feet of me. No doubt this tameness is largely dueto the fact that they are absolutely protected and have nothing to fear fromman within the Park. They often come boldly about the buildings of the littlevillage at Mammoth, and even walk across the lawns, both in winter, andwhen the tourists are numerous in summertime. I have sometimes thoughtthe birds liked the open in winter, because of the sunny warmth there. Still,they are also out in the open on the warmest of summer days.Spring.?A. W. Anthony (1903) gives a very interesting accountof the extensive spring migration of Richardson's grouse in easternOregon. The flight was mainly southward from the higher ridges,heavily timbered with pine and fir, to the sage-covered benches andridges where they nest. He writes
:
On the first of March, 1902, when the first of the migrating grouse made theirappearance along the edge of the timber north of Sparta, the snow was from2 to 4 feet in depth, though the lower slopes near Powder River were bareand had begun to show the first signs of sprouting grass. Snow squalls andrough weather seemed to check the southward flight until about the 10th,although a few birds were passing over daily. The tracks on the snow boreample testimony as to the manner in which the migration was made.From the higher slopes north of Eagle Canon, the birds sailed until therising ground brought them to the surface of the snow on the south side ofthe creek, usually well above the canon. From this time until the highestpoint of the ridge south was reached the journey was performed on foot.Immediately north of Sparta lies a conical peak known as Baldy, some 700feet above camp, the highest point in the ridge south of Eagle Creek. Fromthe top of Baldy, and in an area not to exceed 100 feet square, 1 think fully85 per cent of the grouse passing over Sparta take their departure. From east,north and west up the steep, snowy slopes hundreds of trails led toward thetop and not one could be found leading downward. The flight from the topof the peak was almost invariably undertaken at about sunrise or sunset. It
98 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
is only when birds are disturbed and driven from the peak that they will at-tempt to cross to the southern ridge during the middle of the day. Through-out the day grouse are arriving along the upper slopes of Baldy, singly, in pairs,and small flocks that have perhaps formed since the southward march began,as I think they do not winter in company, but the flight from the peak is usuallyin flocks of from a dozen to a hundred birds. Though the ridge south of Spartais 400 feet or more lower than the top of Baldy, it is fully a mile and a halfdistant in an air line, and the flight is seldom sustained to carry the birdsto the top. Usually they alight on the snow half way up the slope, and aftera few moments' rest, continue the journey on foot ; those passing over in theevening spend the night, I think, in the pines, the last of which are seen alongthis divide ; but those arriving in the morning soon pass on, walking down anyof the small ridges leading toward Powder River.Courtship.?Maj. Allan Brooks (1926) has published a full account,with illustrations, of the courtship display of this grouse, from whichI quote, as follows:His first position was a crouching one, the tail spread to extreme extensioncocked right over the back and a little to one side, the neck feathers showingas a snowy mass, with the red gular sac looking like a small oyster on alarge shell.He maintained this attitude for several minutes, then the head was raised,the neck swelled, and he turned towards me and commenced to nod his head
;
the gular sacs were a deep purple-red, the " combs " over each eye changed fromyellow to a dusky orange and were inflated to the extent that they almost meton the crown, and the inversion of the neck feathers showed as a huge blaze ofwhite on each side. After six or eight nods the head was lowered to withintwo inches of the ground and with the neck inflated until the sacs showed adiameter of three inches, the tail still elevated and spread to its full extent, thefeathers of the lower back standing on end, the wings trailing on the ground,the bird made a short quick run of six or eight steps curving to the right andemitted the deep Oop!Brooks compares the hooting of Richardson's grouse with that ofthe sooty grouse, as follows
:
Often in later years a male Richardson's Grouse has been seen uttering hislow hooting, similar to the resonant hooting of the Sooty Grouse (Dendragapusobscurns fuliginosus) but with only a small fraction of its volume. In myexperience Richardson's Grouse always utters this hooting from the ground.The tempo is the same as in the Sooty Grouse, five or rarely six deliberate evenlyspaced hoots or grunts
?
Humph-humpJi-humphr-ma-h.umph-humph?but thesound is barely audible up to 75 yards or so. In the Sooty and Sierra Grousethis chant assumes a dominant character, ventriloquial to a degree it soundsfar off when quite near and yet has a carrying power of at least 2 miles. Inneither species is the tail very widely spread when uttering, nor is there anyspecial posture or action excepting a crouching attitude, high up in a coniferoustree in the Sooty Grouse, and on the ground, usually on some rocky ridge, InRichardson's. The Sooty Grouse also has the single Oop! although I have rarelyheard it and then only late in the breeding season.A very pronounced distinction in the two species, however, is the nature ofthe gular air sacs. In the Sooty Grouse and allied races these in the breedingseason become cellular, gelatinous masses, capable of great distention, and theexterior surface is velvety, deeply corrugated, and of a deep yellow color. In
richaedson's grouse 99the fall this specialized character is largely lost, the skin loses most of itscorrugations but still retains a yellow color.In Richardson's Grouse and its sub-species flemingi very little change fromnormal in the character of the neck-skin occurs in the breeding season, theexterior is flesh-colored tinged with purple, deepening to purple-red whentemporarily surcharged with blood.Charles de B. Green (1928) has noticed a curious fluttering habitof the male grouse, which he describes as follows
:
Years afterward I saw it done for the first time in my experience. I was lyingin bed looking out of the door at a female grouse walking on the ground some20 yards away. Near her was a small pine, the lowest branch of which wasabout 10 feet from the ground. On this branch was a male bird and, just asI caught a glimpse of him, he fell as straight as a string to the ground,tumbling over as he came, in what looked like a ball of feathers. This tookonly a second or two, but as he fell the fluttering sound was made. Then hemade short runs at the female, stopping abruptly each time he got near, andgiving one hoot. This running and hooting went on until they were lost toview, but there was no more fluttering.In the last few years, while in the sheep business, it has been my customin March to be with the flock before dawn. Every day for a month this verysame fluttering is the first sound to be heard in the dusk, before things are atall clearly seen at 50 yards away. In fact, before the faintest sign of daylight,from all around comes in fluttering sound. Only once was I able to see thebird : I was watching a grouse about 30 or 40 yards away in the growing light,when he sprang about 3 feet into the air, turned over, and came down withthe fluttering sound on the same spot from which he sprang.Nesting.?Anthony (1903) writes:A few birds undoubtedly remain and nest throughout the timbered region ofPowder River Mountains, but the percentage is small indeed compared withthose that nest on the bare sage plains along Powder and Burnt Rivers. Manyof the nests are placed in the shelter of the scattered growth of chokecherry,aspen, or Cottonwood that fringes the water courses tributary to the river ; anda few of these nests may produce young that reach maturity, but fully as manybirds lay in the shelter of a bare rock, or scanty sage brush in the open plain,in company with Sage Grouse ; and fortunate indeed is the bird, nesting in suchlocation, that raises its young. In a circuit of not over 6 miles from my campon Powder River the past May, were ranged not less than 20,000 sheep whichtramped out the nests so completely, that, while finding dozens of broken nests,I saw not one that had not been destroyed, of either Richardson's or SageGrouse, and only one young bird. Nevertheless, many of them do escape, astheir numbers testify, although I am told, on good authority, that there are veryfew in comparison with their former numbers.Skinner (1927) says that, in Yellowstone Park, "nesting takesplace in May, and the nests are usually placed at the foot of foresttrees at any altitude from 6,300 to 8,000 feet, and perhaps evenabove the last-named height. The Richardson's nests are shallowdepressions lined with grass, pine needles and leaves, and containfrom seven to ten creamy eggs speckled and blotched with brown,
100 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMFresh eggs are laid from May 10 until almost the first of July inbelated cases."Eggs.?The eggs of Richardson's grouse are similar to those of thedusky grouse. The measurements of 32 eggs average 47.9 by 33.3millimeters; the eggs showing the four extremes measure 51.7 by 34,47.5 by 36, 43 by 33.5, and 45 by 32 millimeters.Young.?Bendire (1892) mentions, in some notes from Robert S.Williams, the behavior of some young grouse that were probablytoo young to run away or hide ; he writes
:
On June 21, 1885, while crossing over the almost bare summit of a smallknoll in the foothills of the Belt Mountains, I suddenly almost ran into a broodof young Richardson's Grouse, which had evidently been hatched out but avery short time. The young, about ten in number, were closely huddled to-gether, the old bird standing by their side, with head up, and eyes fairlyblazing at the unexpected intruder. I was almost within reach of them,but neither old nor young made a single motion or uttered a sound while Istood watching them for several moments; and I left them in the same position.Mrs. Florence M. Bailey (1918) writes:As we rode out of the dark woods the peeping voices of young were heard,and as the first horse shied a big mother grouse flew conspicuously into thetop of a low evergreen, while her brood, circling out on widespread curvingwings like young quail, disappeared under cover. Early in August, on theSwiftcurrent, an old grouse and seven half-grown young, finding our campnearly deserted, walked calmly past the tents and under the kitchen awningon their way to the creek. On reaching it the mother flew across, callingthe brood till they followed, when they all walked off toward the blueberrypatch in the pine woods.Food.?In Yellowstone Park, according to Mr. Skinner (1927),their food consists largely of berries, such as bear berries (Arctostaphylosuva-ursi L.), huckleberries [VaccinUim scorparium Leiberg), high bush blue-berries (Vaccinium membranaceum Dough), service-berries (Amelanchier abii-foJia Nutt.), false buffalo berries (Shepherdia canadensis L.), raspberries(Rubus striyosvs Michx. ), gooseberries (Ribes saxosum Hook, and Ribesparvulum Gray), and strawberries (Fragaria amcricana Porter). In additionthey eat many insects, especially grasshoppers. When other food becomesscarce they eat fir, pine, and spruce needles.Aretas A. Saunders has sent me the following interesting notes:In August and September the food is mainly grasshoppers obtained in themountain meadows. A method of obtaining them is interesting. A smallflock of 8 or 10 birds, apparently a female and her fully grown young, standin a circle some 30 feet or more in diameter, the birds 10 or 12 feet apart.Each bird faces toward the center of the circle, and they slowly move in-ward, scaring up the grasshoppers as they go. As the circle grows smaller thegrasshoppers are more and more concentrated in the center, and the birdscapture a great many. A grasshopper jumps and flies from a bird on oneside only to fall victim of one on the opposite side. I have watched thisperformance once in the Sun River country, and flushed birds that were inthis formation several other times, so that I believe it is a common habit.
Richardson's grousb 101Enemies.?The same observer says:Although not here hunted by man, these grouse have plenty of other enemies,such as wolves, coyotes, mink, weasels, and all the other predacious fur bearers,as well as the duck hawks, Cooper's hawks, and western horned owls. Thebig western red-tailed and Swainson's hawks do not bother the grouse evenwhen they are small. I remember once watching a redtail circling overhead.I could not see what it was hunting, but apparently it was not Richardson'sgrouse, for almost at the same time an undisturbed grouse and seven littlefluffy youngsters were noted in the grass under the circling hawk. Soon, themother grouse flew up into a small sapling and from there clucked loudly toher brood, while the little grouse tried their best to fly up to her, one at a time,on very shaky little wings. Sometimes I see a Richardson's grouse with manytail-feathers missing, as if it had just escaped an enemy by the sacrifice of afew feathers.Fall.?The fall migration to the higher elevations is thus describedby Anthony (1903)
:
The return migration is less pronounced in its beginning, and more gradualin its progress. Toward the last of July the broods of well grown young, at-tended by the adults, begin to appear along the ridges, returning as they cameby walking invariably up to the tops of the hills and ridges and as invariablyflying as near to the top of the next as their gradually descending flight willcarry them. Before the middle of August, the migration is in full swing, andflocks are seen each evening, passing over Sparta. Frequently they alight inthe streets and on the house-tops. I recall with a smile the memory of a flockof a dozen or more which lit one evening in front of the hotel. For a tiinqpistol bullets and bird shot made an accident policy in some safe companya thing to be desired, but strange to relate none of the regular residents ofthe town were injured. The same may be said of most of the grouse, thoughone, in the confusion, ran into the livery stable and took refuge in a stall,where it was killed with a stick.Straggling flocks from south of Powder River prolong the tall migrationuntil near the first of October, after which none are seen below the highelevations north of Eagle Creek.Winter.?Skinner (1927) says of the winter habits of these grousein Yellowstone National Park:Normally in summer, these grouse are on the ground, or on low logs andboulders ; and they live mainly in the evergreen trees while the snow coversthe ground. But, when skiing through the winter forests in December andJanuary, I have had roosting birds burst out from under snow drifts. Atother times, most of these birds roost in heavy coniferous trees. If not dis-turbed, they may stay in a small grove of trees, and not descend to the groundfor several successive days. At such times, they eat needles for food and usethe snow instead of water.Mrs. Bailey (1918) was told that in Glacier National Park wherethe snow is from 1 to 25 feet deep, " they roost in holes in the snow."74564?32 8
102 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMDENDRAGAPUS OBSCURUS FLEMINGI (Taverner)FLEMING'S GROUSEHABITSThe range of Fleming's grouse, as given in the new A. O. U.Check List, includes northern British Columbia, southern Yukon,and southwestern Mackenzie. The bird was described by P. A.Taverner (1914), based on a small series of " specimens taken within30 miles of Teslin Lake, on the boundary between British Columbiaand Yukon Territory, longitude 130? 30', at the west base of theCassiar Mountains. The range of the form cannot therefore bedefined."It was named after J. H. Fleming, of Toronto, Canada, and issaid to be " like Dendragapus obscurus richardsoni, without termi-nal tail band, but darker in general coloration even than D. o.fuliginosusPThe subspecies is based on the characters of the male, and Mr.Taverner (1914) says:The characters of the female are less marked than of the male and withoutseries for comparison may be difficult of recognition. However, they averagein the same directions as the male, being bluer underneath than richardsoniand darker dorsally, with the rufous or rusty markings bolder and moredecided in character.Maj. Allan Brooks (1927) adds the following suggestion:The best character for separating flemingi from typical ricluirdsoni is notthe darker coloration of the males ; any large series will show dark coloredrichardsoni and, as in Swarth's Teslin Lake bird, light colored flemingi; butthe blacker under tail coverts of the last named form with small white tips,instead of the white tip covering almost the whole exposed portion of thefeather as in richardsoni. But birds from Revelstoke, Selkirk range, insouthern British Columbia show the extreme of blackness of the lower tailcoverts. It is possible that a wedge running southward to or near the Brit-ish Columbian southern boundary splits the range of richardsoni.Harry S. Swarth (1926) says of the haunts and habits of Fleming'sgrouse
:
In the Atlin region the " blue grouse " is resident and fairly common at highaltitudes. It is a favorite game bird of the region, both from its large sizeand from the excellent quality of its flesh. Its habitat is about timber line,where there is open country interspersed with clumps of balsam firs. The densethickets of these stunted trees, with their gnarled and spreading branches,afford shelter from enemies and from inclement weather, and in the foliagefood also is furnished when other sources fail.The broods are cared for solely by the hen. The old cock is usually soli-tary during the summer, though males of the previous year sometimes formbmall coveys, together with non-breeding females. Such gatherings were en-countered on several occasions. The hen with a brood is sometimes tame to theverge of stupidity; I found several that were, literally, as indifferent to
SOOTY GROUSE 103
approach as any barn-yard fowl. I have, however, seen an occasional covey ofyoung birds that was extremely hard to approach. The broods often feed overopen meadows, where they are exposed to attack by hawks and other enemies,and there must be a heavy mortality from such causes. That this is so is borneout by the small size of most of the broods encountered, and by the numberof hens seen with no broods at all.DENDRAGAPUS FULIGINOSUS FULIGINOSUS (Ridgway)SOOTY GROUSEHABITSIn the heavily timbered, humid coast ranges, from southern Alaskato extreme northern California, we find one of the most widely dis-tributed and best-known races of the " blue grouse " group. It isthe darkest of all the races, except the extremely dark race of theclosely related species D. dbscurus fle?}iingi, which is found in north-ern British Columbia. All the " blue grouse " (genus Dendraga-pus) have until recently been regarded as subspecies of Dendragapusobscurus, but the studies and suggestions of Allan Brooks (1912,1926, and 1929) and of Harry S. Swarth (1926) have resulted inseparating the various races into two groups as subspecies of twodistinct species. The new A. O. U. Check List, therefore, givesthree races of the Rocky Mountain, or eastern, group, obscurus,richardsoni, and flemingi, all as subspecies of obscurus; and thisleaves the four coastal, or western, races, fuliginosus, sierrae, sit-kensis, and hoivardi, as subspecies of a new species, fuliginosus.Major Brooks (1926) gives the following distinctive characters ofthe two groups
:
Group I, including richardsoni and fiemingi.1. Air sacs. Skin not conspicuously thickened or corrugated even in themating season, color flesh, changing to purple red under the influence ofexcitement.2. Voice. " Hooting of five or six notes audible for less than 100 yards,uttered from the ground. Note : The single hoot when in full displayis alike and common to both groups.3. Tail. In adult males, squarer, the feathers truncate at the tips; terminalband of gray darker, sometimes (rarely) absent or but faintly indicated.Group II, including fuliginosus, sitkensis, sierrae and hoivardi.1. Air sacs. Skin highly specialized in the mating season, thick, gelatinous,the surface deeply corrugated into a series of tubercles of a velvetytexture and of a deep yellow color. This condition is reduced when themating period is over.2. "Voice. " Hooting " of five or six notes of great power, audible for severalmiles. Always (?) uttered from high up in a tree.3. Tail. In adult males rounded, the feathers rounded at the tips ; terminalband of light gray averaging narrower than in group No. I.For a further, detailed discussion of the question the reader isreferred to Mr. Swarth's (1926) remarks under fiemingi, with a map
104 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMshowing the distribution of the seven forms and a study of theplumage changes.Major Bendire (1892), who had a wide experience with this species,writes
:
The favorite locations to look for the Sooty Grouse during the spring andsummer are the sunny, upper parts of the foothills, bordering on the heaviertimbered portions of the mountains, among the scattered pines and the variousberry-bearing bushes found in such situations, and along the sides of canons.According to my observations these birds are scarcely ever found any distancewithin the really heavy timber. In the middle of the day they can usually belooked for with success amongst the deciduous trees and shrubbery found alongthe mountain streams in canons, especially if there is an occasional pine orfir tree mixed amongst the former.In the vicinity of Seattle, Wash., in 1911, we found sooty grousestill quite common almost within the city limits. Where much ofthe virgin forest had been cut off, where some of the land had beencultivated, and where scattered houses were being built, the grousestill clung to the remnants of the coniferous forests and the brushyclearings near them. W. Leon Dawson (1909) has expressed it verywell, as follows
:
Indeed, the Blue Grouse and the Douglas fir are nearly inseparable. In thesheltering branches of this tree the bird takes refuge in time of danger ; fromits commanding elevation he most frequently sends forth the challenges ofspringtime; and in its somber depths he hides himself thruout the winter season.So great is this devotion on the part of the bird that it is found indifferentlyat sea level or at the timber line of the highest mountains ; and it will notwillingly quit a favorite piece of woodland even tho the supporting forests becut away on every side.Keferring to its summer haunts on Mount Rainier, Wash., R. A.Johnson (1929) says:The summer range of this species is restricted to the alpine meadow zone, ahabitat extending around the mountain approximately between the elevationsof 5,000 and 6,500 feet. The ground cover of this habitat consists largely ofheather meadows, small mountain ash, and mountain willow thickets inter-spersed with clumps and individual trees of alpine fir (Abies lasiocarpa) andmountain hemlock (Tsuga mertensiana) , the branches of which usually droopso that the lowest ones touch the ground.Courtship.?Early in March the grouse begin to migrate downwardfrom their winter resorts in the heavily forested mountains to theirsummer homes in the more open foothills and valleys. From thattime until the last of May their haunts resound with the loud, deep-toned hootings of the male, his challenge to his rivals, or his courtshiplove notes. I once had a good opportunity to watch a fine old cockgrouse hooting and displaying under very favorable circumstances.It was on Mercer Island in Lake Washington, within the city limits
SOOTY GROUSE 105
of Seattle, where Samuel F. Rathbun had taken us to see some ofthese grouse. After some difficulty in following up the ventriloquialnotes, we located the performer on a horizontal limb, close to tiietrunk of an enormous Douglas fir, fully 50 feet up in the densest partof the tree, but in plain sight ; we watched him for some time throughpowerful glasses and could plainly see every detail. He turned aboutoccasionally on the branch, facing first one way and then the other,with drooping wings, lifting and spreading his tail. When ready tohoot he stretched out his neck, on which two large, white, rosettesappeared, swelling open and showing the naked sacks of dull yellowskin, which puffed out to semiglobular shape with each of the fouror five hoots; the plumage of the whole neck and throat swelled outwith each note and the bill opened slightly. The hooting notes weremuch like the soft, low notes of the great horned owl ; when near athand they seemed to be softer and less powerful, but they reallyhave great carrying power and can be heard for a long distance.They have been likened to the noise made by blowing into the bung-hole of an empty barrel or by swiftly swinging a rattan cane. The}7were given in groups of four to six notes each ; I wrote down the fullgroup as hoooo, hoot, hoot, hoot, a-hoot, hoot, or sometimes as four,five, or six straight hoots, or as different combinations of the abovenotes. I recorded the intervals between the hootings as varying from12 to 36 seconds ; the following series was noted : 12, 18, 22, 23, 35, 14,22, 22, 16, IT, 19, 14, and 32 seconds. During the intervals of silencethe bird assumed a normal pose, or strutted about. Dawson (1909)says : " The hooting, or grunting notes, of this Grouse are among thelowest tones of Nature's thorobase, being usually about C of the FirstOctave, but ranging from E Flat down to B Flat of the ContraOctave."Leslie L. Haskin has sent me the following notes
:
Early in March the males begin to manifest their presence by their muffledhooting, which proceeds from high up in the trees. At this time the birdswill usually be found sitting close to the bole of the fir, their bodies hard todiscern against the general grays and browns of the surrounding limbs. The
"hooting" of the grouse is one of the most distinctive and peculiar bird notesof the Douglas-fir region, as characteristic in its way as the drumming of itsrelative, the ruffed grouse. This call can be best imitated by closing the lipstightly, puffing out the cheeks, and then articulating the sound oo in a deeptone, low in the throat. The hooting of the grouse has a muffled, ventriloquialquality that makes it exceedingly hard to locate?now seeming far, now near,now high, now low?and as a dozen birds may be calling at the same timeit becomes very confusing.As March advances the hooting becomes more pronounced, and the birdsbegin to move about more freely, and when mating takes place they descendto the ground. Now, where no grouse were visible only a few days previous,the whole forest may be alive with them. The females have a cackling call to
106 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
which, early in spring, the males respond very readily. An experienced hunter,by imitating this cackle, can call the birds to him and thus pot them with theleast possible effort.Nesting.?My first and only nest of this species was found onMay 7, 1911, on the island in Lake Washington referred to above.Mr. Rathbun guided us to an old clearing, which he said was thebreeding ground of these grouse; it was an open, sunny hillside, sur-rounded by large fir trees and covered with large fallen logs, clumpsof brakes, shrubs, and small trees. We had been hunting here lessthan 10 minutes when a big grouse flushed from under a small firtree just ahead of me, and there was the nest with seven fresh eggs.The nest was on a little knoll under the small fir, which was onlyabout 7 feet high, and was only partially concealed; the nest cavitywas about 7 inches in diameter and 3 inches deep, with a well-builtrim of green mosses and ferns, and was lined with dry leaves, ferns,bits of rubbish, and plenty of grouse feathers.F. Seymour Hersey collected a set of seven eggs for me, nearTacoma, Wash., on May 3, 1914. The nest was in an open situationin a fir forest and was plainly visible at a distance of 25 feet ; it wasunder the end of a fallen log, a hollow 4 inches deep and 7 inchesin diameter, well shaped, and lined with twigs and moss, with whichthe ground was carpeted, and with a few feathers of the bird.J. Hooper Bowles has sent me his records of some 23 nests foundnear Tacoma between April 30 and May 22. He says in his notes
:
In the selection of a nesting site this grouse, as a rule, prefers a very dry,well-wooded locality, where it scratches out a considerable depression in theground at the base of a tree, under a fallen branch or other shelter. I havefound nests under old boxes, a large tangle of wire netting from a henyard, etc.However, at times they will nest as far as a hundred yards from trees of anykind and with very little concealment. The nesting material is the same asa barnyard hen would use under similar conditions, being mostly what it canreach when the bird is on the nest and what falls into it in the course ofIncubation. A goodly number of feathers also come out of the bird, so thatthe nests are often objects of decided beauty.The period of deposition of the eggs is extremely variable, and it is doubt-ful if they ever lay an egg a day regularly. To give what may be consideredan extreme case was a nest found on May 3, 1928, when it contained two eggs.This nest was built in the side of a huge ant hill about 4 feet in diameter andagainst a large fir tree, the whole place being swarming with large andferocious red ants. On Mny 9, the nest contained only five eggs, and on the12th it still had only five eggs and had them half buried with hill material,so I concluded that the ants had made the bird desert. However, on passingthe place on May 22, I walked over to see if they were all covered over andwas astounded to see the female grouse sitting peacefully on her treasures.She was completely surrounded with ant hill and hundreds of ants wererunning all around her busy with their duties, but not an ant went on her,although a friend and I watched her for some 20 minutes. We flushed herand found that she had cleaned out the nest and lined it with green fir sprays.
SOOTY GROUSE 107Why the ants did not molest her is beyond comprehension, as they certainlynearly ate us alive while we were examining and photographing. The eggsvaried from addled to slightly advanced in incubation, so it would be difficultto judge when the last one was laid.Bendire (1892) gives an interesting account of a "most exposednest, without any attempt at concealment whatever," at an eleva-tion of 6,800 feet in Oregon ; it was in " a beautiful oval-shapedmountain meadow of about an acre in extent, near the summit ofwhich stood a solitary young fir tree." There were no other treeswithin 30 yards. He lay down to rest under the shade of this tree,when his setter dog came running up and pointed a grouse sittingon her nest within 3 feet of him. She allowed him almost to touchher before she fluttered off ; the nest held two chicks and seven eggson the point of hatching.Eggs.?The sooty grouse lays from 6 to 10 eggs, but 7 or 8 arethe commonest numbers; as many as 16 have been found in a nest,but evidently the product of two females. The eggs are indis-tinguishable from those of the dusky grouse, which I have alreadydescribed. Bendire (1892) describes them as follows:The eggs are ovate in shape, and the ground color varies from pale creamto a cream-buff, the latter being more common. In a single set before meit is a pale cinnamon. The eggs are more or less spotted over their entiresurface with fine dots of chocolate or chestnut brown ; these spots vary con-siderably in size in different sets, ranging from the size of No. 3 shot to thatof mustard seed. These markings are generally well rounded, regular inshape, and pretty evenly distributed over the entire egg. An egg is usuallydeposited daily and incubation does not begin until the set is completed, themale taking apparently no part in this duty nor in the care of the young afterthey are hatched.The measurements of 92 eggs average 48.5 by 35 millimeters; theeggs showing the four extremes measure 53 by 34.5, 52 by 37, and 45by 32.5 millimeters.Young.?But one brood is raised in a season. Various observershave reported the incubation period, as from 18 to 24 days ; this dutyis performed by the female alone. Bendire (1892) writes:The cocks separate from the hens after incubation has commenced, I believe,and keep in little companies, of from four to six, by themselves, joining the youngbroods again in the early fall. At any rate, I have more than once come uponseveral cocks in June and July without seeing a single hen amongst them. Highrocky points near the edges of the main timber, amongst juniper and mountainmahogany thickets, are their favorite abiding places at that time of the year.The young chicks are kept by the hen for the first week or two in close prox-imity to the place where they were hatched, and not until they have attainedtwo weeks* growth will they be found along the willows and thickets borderingthe mountain streams. Their food consists at first principally of grasshoppers,insects, and tender plant tops, and later in the season of various species ofberries found then in abundance everywhere, as well as the seeds of a species
108 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
of wild sunflower, of which they seem to be very fond. It is astonishing howsoon the young chicks learn to fly, and well, too, and how quickly they canhide and scatter at the first alarm note of the mother bird, which invariablytries by various devices to draw the attention of the intruder to herself andaway from her young. A comparatively small leaf, a bunch of grass, anythingin fact will answer their purpose; you will scarcely be able to notice thembefore they are all securely hidden, and unless you should have a well-traineddog to assist you, the chances are that you will fail to find a single one, evenwhen the immediate surroundings are comparatively open. After the youngbroods are about half grown, they spend the greater portion of the day, and Ibelieve the night as well, among the shrubbery in the creek bottoms, feeding alongthe side hills in the early hours of the morning and evening. During the heatof the day they keep close to the water, in shady trees and the heavy under-growth. They walk to their feeding grounds, but in going to water theyusually fly down from the side hills.Plumages.?In the downy chick the head and underparts vary from
" cream color " to " ivory yellow " ; the crown is mottled with blackand a little " hazel," and the auriculars are spotted with black ; theupperparts are variegated with " hazel," " chestnut," dusky, and palebuff. The wings begin to grow soon after the chick is hatched; ina chick 3 inches long they already reach beyond the tail. These firstwing feathers and their greater coverts are broadly tipped withwhite and have white shaft streaks.The juvenal plumage comes on very rapidly and is fully acquiredbefore the young bird is half grown, the last of the natal down dis-appearing on the belly and head. In the full juvenal plumage, inwhich the sexes are alike but in which the racial characteristics beginto show, the crown is mottled with " amber brown " or " hazel " andblack; the feathers of the mantle are variously patterned with dullbrowns, " hazel " or " tawny," and black or dusky, with conspicuouswhite shaft streaks, broadening at the tip ; the primaries, secondaries,and tertials are barred, notched, or mottled with pale buff on theouter web; the chin and throat are yellowish white and the bellydull white; the breast and flanks are spotted with dusky and palebuff; the rectrices are narrow and pointed, banded, and mottled,much like the plumage of the back. The juvenal flight feathers aremolted during July and August ; the molt begins as soon as the lastof these feathers are fully grown, or even before that ; and the bodymolt into the first winter plumage is continuous from August toOctober. The postjuvenal molt is complete, except that the outerpairs of primaries are retained for a full year. A. J. van Rossem(1925) says of the tail molt:The juvenal rectrices are shed at a very early age. The lateral pairs gofirst, followed soon after by the central pairs. The chicks can be scarcelymore than two or three weeks old when the tail-feathers are dropped, andthe characteristic post-juvenal tail begins to appear beyond the tips of thecoverts. These new tail-feathers are comparatively slow in growing, and reachmaturity when the first winter plumage is fully acquired. Their shape and
SOOTY GROUSE 109
size, as well as the shape of the tail itself, is diagnostic of birds of the year. Ascompared to that of the adult, the tail is much rounder and more fan-shaped,because of the greater proportionate shortness of the lateral feathers. Theindividual feathers are shorter, narrower, and have rounder tips. Most ofthese rectrices are replaced during the following winter, spring, and summer,but some (usually the outer pairs) are apparently always retained until thefollowing fall. The replacement-feathers are similar to those of the adult.However, the whole tail (replacement and first winter feathers alike) is castat the second fall moult, at which time the longer and broader tail-feathersof maturity are acquired. Replacement is so invariable as to preclude the pos-sibility of accidental or fortuitous moult. These differences are most easily seenin males. They are present in females in lesser degree.The first winter plumage is much like the adult and the sexes arequite unlike; but young birds, during the first year, can alwaysbe recognized by the outer primaries, by the shape of the tail, andby the narrow tail feathers. There is little evidence of even a verylimited prenuptial molt, but a complete postnuptial molt takesplace the next summer, which produces the fully adult plumage,when the bird is more than a year old. Of this Harry S. Swarth(1926) says:The first post-nuptial molt begins about the middle of July of the secondyear and lasts until about the middle of September. The change in characterof rectrices is the one conspicuous feature of the mature plumage. I cannotfind that there is any renewal of rectrices (except sporadically, presumably asthe result of accidental feather loss) until this molt regularly begins. This,I believe, is the only point in which I disagree with van Rossem in the con-clusions drawn by him regarding molt in this genus. The fully adult tail, nowacquired, is square ended, the feathers broad and truncate. Minor color dif-ferences are a clearer gray coloration below and less white spotting on breastand sides, while the mottling on dorsal surface of wings and on interscapularsis less in extent, and gray instead of brown.According to A. W. Anthony (1899) the sooty grouse has beenknown to hybridize with the ring-necked pheasant.Food.?Dr. Sylvester D. Judd (1905a) sums up the food of the
"blue grouse" (various races), as follows:The food consisted of 6.73 percent animal matter?insects, with an occa-sional spider?and 93.27 percent of vegetable matter?seeds, fruit, and leaves.Grasshoppers constitute the bulk of the animal food, amounting to 5.73 per-cent. Beetles, ants, and caterpillars form the rest of the insect food. Onestomach contained the common land snail (Polijgyra sp.).Of the vegetable food, he says
:
Browse is eaten by the blue grouse to the extent of 68.19 percent of itsannual food, and is distributed as follows : Buds and twigs, 5.2S percent
;
coniferous foliage, 54.02 percent; other leaves S.89 percent. The speciesspends most of its time in pine forests feeding on needles, buds, and flowers.The yellow pine (Pinus ponderosa)?male flowers, the white fir (Abiesconcolor), Abies magnified, the Douglas fir (Pseudotsuga mucronata), thewestern hemlock (Tsuga heterophylla), and the black hemlock (Tsuga mer-
110 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMtensiana) are among the trees that afford it subsistence. Plants other thanconifers furnish 14.17 percent of the annual food of the species. This materialincludes red clover leaves, willow leaves, blueberry leaves, miterwort (Mitellabreweri), birch shoots, and poplar flower buds. During July, in Montana andUtah, field agents of the Biological Survey have seen the bird feeding on theleaves, buds, and flowers of the Mariposa lily (Calochortus) . It eats also theblossoms of lupine, columbine, and the Indian paint brush (Castilleja)
.
At times it visits fields for oats and other grain. It feeds also on pine seeds(Pinus fiexilis and other species). It picks up polygonum seeds (P. poly-morphum and others), is fond of wild sunflower seeds, and has been known tosample false sunflower (Wyethia mollis), caraway (Glycosma occidentalis) ,and the capsules of Pentstemon gracilis. It picks up also the seeds of variousspecies of lupine, and is fond of acorns, including those of the canyon live oak(Quercits chrysolepis)
.
Among the berries eaten he includes manzanita berries (13.48 percent of the whole), bear berries, gooseberries, huckleberries, service-berries, salmon berries, and the fruits of red elder, honeysuckle,cherries, mountain-ash, salal, and currants.Mr. Bowles writes to me
:
In the examination of a great many stomachs of this grouse I have invariablyfound the contents to be 100 per cent vegetable, with no signs of animal foodof any kind. Green leaves of different kinds form at least 75 per cent of theirdiet, such as grass, ferns, kinnikinnick, etc., with different small berries next,while one was packed with dry green peas. This diet continues from about thefirst of April until the end of October, when the only food of any kind that Ihave found consists of the tips of green fir sprouts of the Douglas fir. As withthe rest of the family, small gravel is swallowed to aid in grinding up the food.Behavior.?When rising from the ground in the open clearings inits summer haunts the sooty grouse flushes with a loud whirring ofwings, like other grouse, and flies directly away across the clearing tothe nearest timber, with a strong steady flight. It usually alightsin a tall, thick, coniferous tree, where it stands so still that it is noteasily seen ; it crouches down on a limb lengthwise or huddles againstthe trunk, where its colors match its surroundings so perfectly thatit is easily overlooked ; only the keenest, practiced eyes can discover it.If flushed from a tree on the mountainside, where it is usually foundin fall and winter, it sails downward on silent, extended wings anddisappears in the forest below.In regions where these grouse have not been hunted much they aretame, almost to stupidity, the young birds particularly; it has beensaid that, when a number are perched in a tree, several of them maybe killed, if the lowest one is shot first, before the others take flight
;
this must be unusual, however. Even where they have been hunted,they are not nearly so wary or so resourceful as the ruffed grouse.They seem to feel particularly safe in the tops of the tall fir trees,where it is sometimes difficult to flush them even by shooting at them.The males during the mating season are quite unconcerned, even in
SOOTY GROUSE 111the presence of quite an audience ; they seem wholly absorbed in theirhooting and keep up their display in plain sight. How different fromthe behavior of the shy ruffed grouse
!
K.A.Johnson (1929) writes:On September 6, I observed a brood of Sooty Grouse going to roost. I locatedthem by their low cackling noise which is often heard at such times. Thiscackle is a subdued one and is somewhat similar to that of the domestic fowl,except for the absence of the outbursting notes with which the domestic henends her performance. In going to roost each bird settled upon a thick mat ofbranches about eight feet from the trunk of the tree and from twelve to thirtyfeet from the ground. In this position their dusky color blended somewhatwith the moss and lichen-covered branches. The birds settle singly, usuallywith the head turned toward the tip of the broad frondlike branch upon whichthey perch, yet concealed for the most part by overhanging branches.Voice.?In addition to what has been written above, I quote thefollowing observations by John M. Edson (1925), which describethe hooting of this grouse, as well as its tameness
:
The voice of the grouse has an almost ventriloquial carrying power. Al-though in reality his notes are not loud, they often may be heard for very con-siderable distances. Still, it happens as frequently that the supposedly distanthooter is in fact close at hand. The hooting appeared to commence about 4A. M., or perhaps a bit earlier, and was heard off and on till after 9 P. M.,and sometimes even as late as 9 :45. The notes of the hooting Sooty Grousemay perhaps be described as deep bass, but soft in quality, expressed as:Oot, oot, oot, oot, t-oot, the second and fifth notes being noticeably subdued.Different individuals vary this slightly. Frequently the last note is omitted, andoccasionally but three notes are given. The watch was held on one bird thathad been vociferating steadily for some time, one afternoon about 3 o'clock,and it was found that his performances numbered just seven per minute foreach of four minutes, following which he became silent for a time. This birdgave only four notes.The most interesting incident in connection with our grouse acquaintanceshipoccurred on the morning of the 11th. The weather was pleasant, with a fewfleecy clouds and a southwest breeze. Somewhere down the rocky ridge thatpitched to the north from our camp, a grouse had been hooting since 4 o'clock.As we sat by our camp-fire at breakfast the hooting seemed to draw nearer, andeventually as if the bird might be out in the open space that commenced notfar below the camp. I rose to peep over a near-by rock that obstructed my view,wondering if the bird might not be in sight. And indeed he was. Standingupon another small, flat rock immediately behind the first and just eight pacesfrom where I stood, was our performer all posed for his act. I moved outcautiously till he was in full view, then stood motionless. The bird seemed notthe least disturbed by my presence, and after giving me an inquisitive glance,soon started the hooting ceremony once more. Standing with his side towardme, his body pitched at an angle of about 45 degrees, the tail slightly drooping,head well up and neck and breast feathers somewhat puffed out, he began bydrawing down his head and further inflating his feathers till the bill and head,except from the eyes up, were concealed. Then throwing open the pocket of hisneck feathers he showed a hoilzontally elongated patch of white lining, in themidst of which was distinguishable the yellowish air-sacs peculiar to the genus.
112 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
It was not greatly distended or conspicuous, being partially concealed by thefeathers. With his effort of giving voice to his feelings, the bird's whole formpulsated with each note, the half-spread tail vibrating vertically. After theconcluding note, he would raise his head, close the neck pocket and calmly lookabout. He repeated this numerous times for six or seven minutes. Occasionallya whiff of smoke from the camp-fire would sweep past, and this would take hisInstant attention but caused no serious alarm.Fall.?Writing of conditions on Vancouver Island, where themigratory habits of the sooty grouse are evidently the same as else-where, Swarth (1912) says:The species is locally migratory, descending into the valleys during breed-ing season, and retreating into the higher mountains at the end of the summer.The old males go first, beginning to leave about the time the females are bring-ing their young from the nest. At Beaver Creek a few still lingered throughJune and could occasionally be heard hooting. In the mountains south ofAlberni, in July, no old males were seen at the bottoms of the basins, or in thecanons, where females with young were frequently met with, but on the higherslopes and the summits of the surrounding ridges they were quite abundant.At the top of Mount Douglas (altitude about 4,200 feet) several were heardhooting July 14 to 16.At Errington, early in September, sooty grouse were abundant and gatheredin flocks, usually of from six to ten individuals, though as many as fifteen wereseen in one gathering. At this time there were no males in the lowlands,these flocks being in all probability composed usually each of a female with herbrood; but a trip to the summit of Mount Arrowsmith, September 6 to 8,disclosed the presence of the cock birds in numbers everywhere on the higherslopes of the mountains. About the second week in September the others beganto follow, and they soon became quite scarce in the lowlands. By the end ofthe month but very few remained.Game.?The " blue grouse," or " gray grouse," or " mountaingrouse," as it is variously called, has been the finest game bird of thenorthwest coast region; but it is becoming very scarce near thecenters of population. It is a fine, large, heavy bird, weighing 3or 4 pounds. When the bird is feeding on berries in summer or fall,its flesh has a delicious flavor and it is often very fat ; but in winter,when it feeds almost exclusively on coniferous browse, it has aresinous flavor which is not so good. It lies well to a dog and, whenflushed in open clearings or in the mountain pastures or berrypatches, it flies away with a straight, steady flight, making very prettyshooting. In the heavily timbered mountains it is not so easj' amark, as it glides silently out of the top of some tall fir tree and goesscaling downhill at a swift pace. It is a difficult mark too, as itflies awTay among the maze of tree trunks in a heavy stand of Douglasfir, where many a charge of shot finds a tree trunk instead of thebird.Winter.?Keversing the habit of most other birds, the "bluegrouse " spends the summer in the lowlands and retires to the higher
SOOTY GROUSE 113mountain forests to spend the winter. Mr. Haskin says in hisnotes
:
It is a remarkable bird, with habits all its own. Throughout the wintermonths the birds are seldom seen, even in places where they are most abundant,for at this season they retire to the heavy fir timber, and spend their timevery quietly high up in the trees. During this season they feed almostexclusively on fir buds, and do not even descend to the ground to drink, for theabundant rainfall makes it possible for them to quench their thirst in thetreetops. Personally, I do not think that the grouse ever voluntarily comes tothe ground during these months of retirement. Only when by accident theyare disturbed, as when woodsmen ave felling trees, are they likely to be seenat all. In sections where the grouse are very abundant you may pass throughthe woods day after day, and unless you understand their ways, never suspectthat such a bird is present. DISTRIBUTIONRange.?Pacific coast mountain ranges of the United States, Can-ada, and southeastern Alaska.The range of the sooty grouse extends north to southeasternAlaska (Glacier Bay and Portage) and northwestern British Colum-bia (Wilson Creek). East to British Columbia (Wilson Creek,Hastings Arm, and Westminster) ; central Washington (Barron,Buck Creek Pass, Cascade Tunnel, Ellensburg, and Signal Peak) ;central Oregon (Wapinita, Fort Harney, and Drews Creek) ; north-eastern California (Warner Mountains and Honey Lake) ; westernNevada (Truckee and Marlette Lake) ; and southeastern California(Bishop Creek, Mount Whitney, and Piute Mountains). South tosouthern California (Piute Mountains and Mount Pinos). Westto California (Mount Pinos, Dunlap, Yosemite Valley, Big Trees,Mount Sanhedrin, summit of the Yolla Bolly Mountains, Summer-ville, and the White Mountains) ; western Oregon (Glendale, Rose-burg, Spencers Butte, Newport, and Tillamook) ; western Washing-ton (Cape Disappointment, Grays Harbor, Hoh River, and CrescentLake) ; western British Columbia (Victoria, Mount Douglas, prob-ably Delia Lake, Haida Mountain, and Massett) ; and the westernpart of southeastern Alaska (Coronation Island, St. Lazaria Island,Sitka, Cross Sound, Skagway, and Glacier Bay).The sooty grouse, formerly considered as a subspecies of the duskygrouse {Dendragapus o. obscurus), is raised to full specific rank inthe 1931 edition of the A. O. U. Check List. Three additional sub-species are recognized, the distribution as above outlined being forthe entire species. True fuliginosus is confined to the mountains ofthe northwest coast from northwestern California and Oregon, northto Alaska (Skagway) and northwestern British Columbia. D. f.sitkensis occupies the islands of the southeastern Alaskan coast (ex-
114 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
eept Prince of Wales Island), Queen Charlotte Islands, and PorcherIsland ; D. f. sierrae is found from central southern Washington andFort Klamath, Oreg., south on the inner side of the coast range toMount Sanhedrin, Calif.; and D. f. howardi is confined to an areain southern California from Mount Pinos east through the TehachapiRange and north in the Sierra Nevada to about latitude 36? N.As stated under Dendragapus o. obscurus, strictly speaking thesooty grouse is nonmigratory, but locally it has a curious verticalmigration as it descends into the valleys during the breeding seasonand retreats to the higher mountains at the end of summer.Egg dates.?Washington and Oregon (fuliginosus) : 60 records,April 16 to July 12; 30 records, April 30 to May 22. California(sierrae) : 5 records, April 27 to June 9. California (howardi) :1 record, May 21. Alaska (sitkensis) : 1 record, June 2.DENDRAGAPUS FULIGINOSUS SIERRAE ChapmanSIERRA GROUSEHABITSThe sooty, or " blue," grouse of the Sierra Nevada and the innerside of the coast ranges, from southern Washington to about latitude31? N. in California, was described by Dr. Frank M. Chapman(1904) as "most nearly related to Dendragapus obscurus, but thenuchal region often browner and usually vermiculated with black,the whole dorsal region less black and more heavily vermiculatedwith brown and gray ; terminal tail band narrower and more speckledwith blackish; the median tail-feathers more heavily marked withgray or brownish ; the scapulars and tertials with the terminal whitewedge less developed or entirely wanting; the basally white neck-tufts practically absent; the throat averaging duskier and the feathersof the sides, flanks, and under tail-coverts with much less white.Differs from Dendragapus obscurus fuliginosus in much palercoloration above, in the heavier vermiculation of the entire uppersurface, practical absence of neck-tufts, whiter throat, and palerunderparts." He says further:In spite of the fact that the Sierra Grouse more nearly resembles obscurusthan it does fuliginosus it apparently has been derived from the latter ratherthan from the former. That is, it represents a southern extension of thenorthwest coast form and not a westward extension of the Rocky Mountainform.This theory is supported by the apparent continuity of range of sierrae andfuliginosus and by their evident intergradation in the vicinity of Klamath,Oregon. Several of the specimens, in an admirable series collected by MajorBendire, at Fort Klamath, are referable to sierrae rather than to fuliginosus,though not typical of the former. Other examples in this series, however,
SIEREA GROUSE 115
are much nearer to fuliginosus. On the other hand, I have no material prov-ing continuity of range in sierrae and obscurus, and the character of thecountry intervening between the nearest known portions of their respectiveranges would lead one to suppose that they do not intergrade geographically.It lives chiefly in the Canadian Zone and only locally in the UpperTransition, ranging " upward into the Hudsonian Zone during latesummer," according to Grinnell and Storer (1924), who write:Acquaintance with the Sierra Grouse may begin in several ways, but rarelydoes it come in the conventional manner through which we learn to know mostbirds. Upon entering the Jeffrey pine and red fir forests of the Canadian Zonein spring and early summer, one may often hear a very un-bird-like, dull soddenseries of booming notes that have a ventriloquial quality. These are the court-ing notes of the male grouse. Less often, whatever the time of the year, theintroduction may come suddenly and much more impressively when, close athand, a heavy-bodied " blue grouse " rises quickly from the ground and makesoff through the forest on loudly whirring wings, and showing an expanse ofsquare-ended gray-banded tail. When a small flock of the birds get up, asthey often do, in rapid succession, or even simultaneously, the aggregate effectis bewildering, to say the least.The Sierra Grouse lives in the high country throughout the year, never migrat-ing to lower levels as does the Mountain Quail. The thick heavy plumage andlegs feathered clear down to the toes enable the grouse to withstand the coldof the midwinter months ; while their ability to subsist on pine and fir needlesassures them at any season an abundance of food to be easily obtained withoutseeking the ground.A. B. Howell (1917) says of its haunts:Although most of the published information pertaining to the Sierra Grousegives one the impression that these birds haunt the pines and associations ofscant undergrowth, my experience has been that they seldom resort to thelarger conifers except to roost, and to escape their enemies by remaining mo-tionless in the upper branches. At least in the locality under consideration,their favorite habitat is in the vicinity of dense aspen thickets, and the tanglesof manzanita, hazel and other brush on the dry hillsides and benches of thehigh Transition Zone, from which they flush to the timbered ravines.Courtship.?Grinnell and Storer (1924) have described the boom-ing courtship of this grouse very well, as follows :During the spring and early summer, the males are in the habit of takingsolitary positions near the tops of pines or firs, sixty or more feet above theground, where they stand on horizontal limbs close to the trunk. They holdsuch positions continuously for hours, one day after another, and send forth atintervals their reverberant booming. With different birds the series of notescomprising this booming consists of from five to seven syllables, six on anaverage. The quality of the sound can be likened to that produced by beating ona water-logged tub, boont, boont, boont', boont', boont, boont, crescendo at thefirst, diminuendo toward the end of the series. As each note is uttered thetail of the bird is depressed an inch or two?perhaps an index to the effortinvolved. The separate series of notes in two instances were uttered at in-tervals of 40, 20, 25, 45, 12, 21, and 29 seconds, and again 10, 10, 20, 26, 14, 15,17, 12, 11, 15, 13, 28, 17, and 11 seconds, respectively. These two birds hadbeen heard booming for a long time before we began to pay special attention
116 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUM
to them, and they continued long after we finished this record. The ventrilo-quial quality is discovered when one attempts to locate the producer, a difficultfeat as a rule. The observer may succeed in locating the proper tree, but islikely to circle it many times, peering upward with painfully aching neck, andstill utterly failing to locate the avian performer amid the foliage high over-head. The notes are commonly supposed to be produced by the bird's inflatingand exhausting the glandular air sacs on the sides of the neck. These sacs arecovered by unfeathered yellow skin, and we think it more likely that theyserve only as resonators, being kept continually inflated, while the air actuallyproducing the sound passes to and from the lungs along the regular air pass-age. It rests with some one gifted with patience for long-continued observa-tion to determine exactly how the notes are produced.W. Leon Dawson (1923) also says:As the hooter becomes vehement he struts like a turkey-cock, spreading thetail in fan-shape, dropping the wings till they scrape the ground, and inflatinghis throat to such an extent as to disclose a considerable space of bare orange-colored skin on either side of the neck. This last certainly makes a stunningfeature of the gallant's attire, for Nature has contrived that the feathersimmediately surrounding the bald area should have white bases beneath theirsooty tips. During excitement, then, as the concealing feathers are raised andreversed, a brilliant white circlet, some five inches in diameter, suddenly flaresforth on each side of the neck, to the great admiration, no doubt, of theobservant hen.These more emphatic demonstrations are probably reserved for such timeas the hen is known to be close at hand, for I have never frightened a strut-ting cock without finding a female hard by, at least at no greater distance thanthe lower branches of a neighboring tree. She has responded to the earliercalls of the male by a single musical toot note, uttered at intervals of ap-proach ; but once arrived at the trysting place she has become very shy, andwill take no part in the celebration, save by a few tell-tale clucks and many coyevasions. On these occasions, also, the cock works himself up into such atransport that he becomes oblivious to danger, so that he may be narrowlyobserved or even captured by a sudden rush.Nesting.?The nesting habits of the Sierra grouse are similar tothose of other forms of the species. Mrs. Irene G. Wheelock (1904)writes
:
In May or June, according to location, the wooing begins, and soon the motheris brooding on her eight buffy eggs in the shade of a fern tangle, near a log,or in a clump of manzanita. No part does the father take in the three weeksof patient incubation, but the mother can seldom be surprised away from thenest. It would be far easier to discover the eggs were she not covering them,for so protective is her coloring that you may be looking directly at her andnever suspect it, although at that very moment you are searching for a nest.Her food is all about her?buds, berries, and insects. If she leaves the eggs,it is only to stretch her tired little legs and pick up a few dainties close by.Eggs.?The eggs of this grouse are indistinguishable from thoseof other races of the species. The measurements of 23 eggs average48.7 by 35.2 millimeters ; the eggs showing the four extremes measure45.1 by 34.3, 49 by 37.5, 51.2 by 34.6, and 46.1 by 32.8 millimeters.
MOUNT PINOS GROUSE 117Young.?Mrs. Wheelock (1904) says that as soon as the young areout of the shell and dry, away goes the mother,proud as a peacock, with them at her heels. And now the father is intro-duced to family cares, and he scratches for bugs, calling the young with impera-tive little chucks to come. He is the drill-master of the little flock, teachingthem with infinite patience all that they need to know of wood lore. He standson guard at every suspicious noise, and whistles his warning when dangerthreatens. When their wing-feathers have developed and they can flutter upto a low branch in the bush, they roost there instead of cuddling under themother's broad wings at night. But they remain with the parents and evi-dently uniler discipline throughout the first six or eight months of their exist-ence. In the wintry weather, when their mountain homes are covered deepwith snow, they often sleep huddled together deep in a drift, waking to feedupon the buds of the coniferous trees, but seldom seeking a lower level. Theyare the hardy mountaineers, the children of the forest ranges.Grinnell and Storer (1921) write:By early July the new broods of grouse are to be looked for in the brush-bordered glades of the forests. When the chicks have been partly reared themales desert their mates, and, forming in flocks of 6 or 8, work higher inthe mountains. The females remain with, and continue to care for, their off-spring, these family units remaining separate for the time being. Finally, asthe summer wanes, they, too, work up into the Hudsonian Zone. Thus, whilethe Mountain Quail go down-hill in the fall, the grouse go up-hill.Food.?The same observers say on this subject
:
One of the above-mentioned male birds was shot, and its crop was found tocontain 1,520 needle tips of the lodgepole pine. The bitten-off ends of needlesvaried from one-fourth to one inch in length. The crop also contained a fewfragments of very young pistillate cones. The bill of this bird was smearedwith pitch. The crop of an adult female grouse obtained at Walker Lake heldeleven ripe rose hips, and the gizzard was filled with the hard seeds of the rose,together with grains of quartz which of course had served to grind theresistant portions of the bird's food.Grinnell, Dixon, and Linsdale (1930), in their report on theLassen Peak region, say:The crop of a female trapped on July 6, 1924, in a rolled-oat baited steeltrap contained a quantity of unripe manzanita berries. On September 16, 1923,at 8,200 feet on Warner Creek, a bird was discovered as it was eating manzanitaberries. At the same station a grouse was watched as it stalked grasshoppers.The bird would stretch out its neck and slowly approach the insect until withinone-half meter when it would make a quick rush forward and capture thehopper in its bill.DENDRAGAPUS FULIGINOSUS HOWARDI Dickey and van RossemMOUNT PINOS GROUSEHABITSThe southernmost race of the fuliginosus group of " blue grouse "has been recently discovered on Mount Pinos in Kern County, Calif.74564?32 9
118 BULLETIN 162, UNITED STATES NATIONAL MUSEUMIt was described by Dickey and van Rossem (1923) and named inhonor of O. W. Howard. The characters given are
:
Nearest to Dendragapus obscurus sierrae, but differing from that form inpaler dorsal coloration, and in coarser and more conspicuous vermiculation andbarring. Underparts darker, a brownish suffusion replacing the clearer grayof sierrae. The white median shafting and terminal pattern of the feathers offlanks and sides reduced in area and entirely lacking on anterior part of body,whereas in sierrae traces of this pattern extend forward to the shoulders.Wing slightly longer; tail decidedly longer and much more graduated, withterminal band averaging wider. Culmen, tarsus, and middle toe averagingslightly longer and decidedly heavier.Throughout the range of Dendragapus obscurus in California there is agradual geographic variation which particularly affects the length and gradu-ation of the tail. These characters increase steadily from north to south.Birds from Mount Pinos express in ultra-typical form this lengthening of thetail itself, as well as the greater ratio between the length of the lateral andmedian rectrices, a truly striking character which the writers have termed
" graduation " in the above description. In this same region, the variationin color and pattern from typical sierrae is also most pronounced.J. R. Pemberton (1928) says of the haunts of this grouse:These birds inhabit the crests of some of the higher mountains from thesouthern extremities of the Sierra Nevada through the Tehachapi Range toMount Pinos in Southern Kern County, California. Mount Pinos is the southern-most recorded station. This high peak reaches an altitude of 8,826 feet and isbeautifully wooded with several species of pines and the silver fir. The grouselive only on the higher portions of the mountain and I believe have not beenobserved below 7,800 feet, which is the elevation of the old sawmill. In asense their range coincides with the areas where the silver fir reaches its bestdevelopment. The upper part of Mount Pinos consists of a rather gentiyrolling table-land. The automobile road ends at an altitude of 7,800 feet, andin a walk of two miles the summit, 8,826 feet, is reached. The mountain isreally a broad ridge with an exceedingly steep north slope which falls 3,800feet in a distance of three miles to San Emigdio Creek. This creek runs in aneast and west valley paralleling the longer diameter of Mount Pinos.Nesting.?Mr. Pemberton (1928) was the first, and so far as Iknow is the only one, to find the nest of this grouse. Describinghis experience with it, he writes
:
This year, Dudley DeGroot and the writer spent May 21 looking for eggs,being unable to make the trip earlier. The interesting discovery was madethat at that time no birds could be located on the higher part of the mountain,while well down on the cliff-like north slope many hooters could be heard. Webelieved that the hooters were near the sitting females, so we spent our timeclambering about on this steep slope. Many tons of rocks were rolled downbut no birds could be flushed. Finally, as I was about ready to give it up andabout 200 feet below the rim of the steep slope at about the 8,200 foot level, Iflushed a female at a distance of about 50 feet and immediately saw the egg*.The bird left with a great whirr, lit on the lowest branch of a large pine about100 feet distant, clucked a few times as she walked to the end of the limb, andthen flew noiselessly downhill. The location was near a point where a hooterhad been circling all day and, although he moved his location many times, it was
SITKA GROUSE 119now evident that he had been in sight of the sitting female all the time. Thenest was in clear open ground and without the slightest cover for the eggs.A depression less than an inch in depth seemed to have been scratched out of thedry, sandy soil and lined rudely with bits of pine bark, a few needles andvegetable trash. Many feathers lay loosely with the eggs. It was a poorexcuse for a built nest and was rather a simple resting place for the eggs.I believe the following generalizations can be made. Howard's Grouse neston Mount Pinos during the first week in May, and full sets will be found beforethe 15th. The nests are fairly well down on the steep north slope and placed inentirely open ground in sunny spots well covered from the distance by observa-tion trees. Nests ought to be found by search near where hooters are active.In early May the snow banks will eliminate all unlikely ground. As soon asthe young are able to walk they are led to the flatter upper slopes of the moun-tain where there is good cover and more food. It is obviously unsafe to at-tempt a statement concerning the number of birds which live on Mount Pinos,but one can say that there are not many and I believe that the number is lessthan one hundred.Eggs.?I saw these eggs in Mr. Pemberton's collection and, as Iremember them, they are like certain types of sooty-grouse eggs. He(1928) describes them as follows:The five eggs were nearly ready to hatch and the embryos had feathers aninch long. They resemble miniature turkey eggs but with larger spots. Theground color is light buff while the spots are auburn, using Ridgway's ColorStandards and Nomenclature (1912). The more prominent spots are 2 and 3millimeters in diameter and one egg has two spots 8 and 10 millimeters in diam-eter. The measurements are 49 by 36, 49 by 37, 50 by 36, 51 by 36, 51 by 37
;
the average is 50 by 36.5.Behavior.?The summit and steep slope of Mount Pinos haveproved difficult country to hunt on, and late snowstorms, cold rains,and high winds make it very uncomfortable for the collector, evenlate in spring. Several good collectors have made repeated attemptsto get specimens of this grouse without success. The birds seem tobe very wild and unusually crafty in avoiding capture. Dickey andvan Kossem (1923) say:The birds of Mount Pinos display a sagacity in eluding capture that is utterlybeyond anything observed by the authors in birds from the central or northernSierra Nevada. One "hooting" site, in a Jeffrey Pine, was carefully watchedon several different occasions during a period of two years, before the bird waslocated and secured. By contrast, the species in like season in the Sierras isoften lacking in suspicion to the point of actual stupidity.DENDRAGAPUS FULIGINOSUS SITKENSIS SwarthSITKA GROUSEHABITSOn some of the islands of southeastern Alaska there occurs a raceof the sooty grouse in which the females are conspicuously reddish in
120 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
color. Harry S. Swarth (1921) has described this race under thename sitkensis, characterizing it asmost nearly like Dendragapus obsrurus fitliginosiix. Adult male not appreciablydifferent from the male of D. o. fuligino&uz. Adult female aud immature ofboth sexes, as compared with those of fuliginosiis, much more reddish in generalcoloration. This color feature affects practically all the plumage except somelimited areas, as the slaty colored abdominal tract, the chin and throat, andthe unmarked and generally concealed portions of the remiges and rectrices.The predominant color dorsally is close to pecan brown. Individual feathersare barred with black and brown, and are brown tipped. On head and neckbrown predominates, the narrow black bands being almost entirely hidden.Upper tail coverts and central rectrices are conspicuously of this reddish browncolor. Breast and sides are mostly pecan brown and black. There are con-spicuous white spots on sides of breast and flanks. Tarsus brown.Female fuliginosus, in comparison, is colored as follows : The upper partsare a duller brown with a great deal of black showing through and with thebrown everywhere sprinkled with black or gray. There are no pure reddishbrown areas as in sitkensis. The neck above is predominantly grayish ; uppertail coverts and remiges are mostly grayish. Breast and sides are mostly grayand black, with very little reddish. Feathers on sides of breast are dullbrownish, mottled with black and tipped with white. Flanks are mostly grayish.Tarsus gi'ay.There are four eggs of the Sitka grouse in the collection of P. B.Phillip, which measure 52.7 by 35.3, 52.7 by 35.1, 52.7 by 35.1, and49.7 by 35.1 millimeters.Alfred M. Bailey (1927) says of their habits:The birds are often so high they will not flush with the discharge of a gun ; infact, I have seen a Grouse sit within twenty feet of a gunner, who tired a dozenshots with a high power rifle in an attempt to shoot the bird in the head, with-out the bird seeming the least alarmed. In the early fall, many repair tothe mountain tops with their broods, where they find ample cover among thedwarfed pines and dense alder thickets; then they drop to sea level during thecold winter months, and one will often see them below snow line, where thetide has cleaned the beaches. They feed on the hillsides, among the dead devil-club and berry bushes, and rarely fly when one passes. Although stronglytainted with the spruce, which makes up a great part of their food, the flesh ofthese birds affords a welcome addition to the camp-fare when one is afield.CANACHITES CANADENSIS CANADENSIS (Linnaeus)HUDSONIAN SPRUCE GROUSEHABITS
? !ON TRIBUTE!) BY CHARLES WENDELL TOWNSENDThe Hudsonian spruce grouse thrives best in regions where manis absent. In fact it remains so woefully ignorant of the destructivenature of the human animal that, unlike its cousin, the ruffed grouse,it rarely learns to run or fly away, but allows itself to be shot,clubbed, or noosed, and, in consequence, has earned for itself the
HUDSONIAN SPRUCE GROUSE 121proud title of " fool hen." As a result, wherever man appears, thespruce grouse rapidly diminishes in numbers, and, in the vicinity ofvillages or outlying posts, is not to be found. It is a bird of thenorthern wilderness, of thick and tangled swamps, and of spruceforests, where the ground is deep in moss and where the delicatevines of the snowberry and twinflower clamber over moss-coveredstubs and fallen, long-decayed tree trunks.Although spruce grouse are resident wherever found even to thenorthern limit of their range, a certain extent of movement occursamong them in winter, dependent probably on the food supply andnot on the severity of the cold.Courtship.?As with the ruffed grouse, " drumming " by the wingsis an important feature of the courtship, but in this the spruce grousehas not reached so high a degree of evolution. It appears to be ata stage midway between the bird that in courtship flies with rapidlyand noisily fluttering wings and the bird that stands still and fluttersor " drums " with its wings.J. L. Devany (1921), writing of the courtship of the spruce grouse,says:His favorite location at such a time is between two trees standing apart some20 or 30 feet, and with the lower branches large and horizontal. Perched onone of these branches he pitches downward, pausing midway to beat and flutterhis wings, and ascend to a branch of the opposite tree. After a short intervalthis manoeuver is repeated and so continued by the hour, swinging back andforth from tree to tree, the time between each swing being as exact as ifmeasured by a watch. If such an ideal situation is not at hand, the fact doesnot prevent the " fool ben " from giving vent to his exuberance. Selecting asmall open space among the bushes, he takes his stand in the center and, likea jack-in-the-box, pops up a few feet in the air and, giving his triumphant flut-tering, drops again to earth * * *. The sound produced by the drummingof the Canada grouse can in no-wise compare with that of the ruffed grouse;it has neither the roll nor the volume. It is in fact little more than a flutter,such as might be made by birds forcing their way through thick branches afterbuds or berries. Unlike the ruffed grouse, however, he seems to have no verystrong objections to an audience. The performance of a ruffed grouse canonly be witnessed by the exercise of stealth and caution. Our little sprucepartridge on the other hand will peer and look at the intruder, and then, as ifsuddenly remembering, go through his evolutions with a gusto that excitesour startled amusement. Though the drumming of the grouse is peculiar tothe male, its practice is not confined to the nesting season alone, but may beheard in any month of the year, and occasionally at any hour of the day ornight.Everett Smith (1883) thus describes the performance:The Canada Grouse performs its " drumming " upon the trunk of a standingtree of rather small size, preferably one that is inclined from the perpendicular,and in the following manner : Commencing near the base of the tree selected,the bird flutters upward with somewhat slow progress, but rapidly beating
122 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
wings, which produce the drumming sound. Having thus ascended 15 or 20feet it glides quietly on the wing to the ground and repeats the manoeuvre.Favorite places are resorted to habitually, and these " drumming trees " arewell known to observant woodsmen. I have seen one that was so well wornupon the bark as to lead to the belief that it had been used for this purposefor many years. This tree was a spruce of 6 inches in diameter, with aninclination of about 15 degrees from the perpendicular, and was known tohave been used as a " drumming tree " for several seasons. The upper surfaceand sides of the trunk were so worn by the feet and wings of the bird or birdsusing it for drumming, that for a distance of 12 or 15 feet the bark hadbecome quite smooth and red as if rubbed.Bendire (1892) quotes another description of the drumming byJames Lingley:After strutting back and forth for a few minutes, the male flew straightup, as high as the surrounding trees, about 14 feet; here he remained sta-tionary an instant, and while on suspended wing did the drumming with thewings, resembling distant thunder, meanwhile dropping down slowly to thespot from where he started, to repeat the same thing over and over again.Bendire also quotes this from Manly Hardy : " My father, whohas had opportunities to see them drum, told me they drummed inthe air while descending from a tree."Nesting.?The nest of the spruce grouse is difficult to find, as itis generally placed under the low protecting branch of a spruce orin deep moss and concealed in a tangle of bushes. As the motherbird so perfectly matches the dead leaves and twigs of the forestfloor, and as she does not move except in imminent danger of beingstepped upon, the difficulty of discover}' is increased, and it not in-frequently happens that the intruder steps in the nest and breakssome of the eggs before he realizes that it is there. This method offinding a nest occurred when Mr. Bent and I were cruising alongthe Canadian Labrador coast. Mr. Bent had offered a reward toanyone who would bring him a set of the eggs of this bird, as ourown search had hitherto been fruitless. While we were anchoredbehind Little St. Charles Island, a fisherman came on board witheight of these beautiful eggs in his hat. He explained somewhatruefully that he had stepped into the nest almost on the sittingbird, and crushed four of a set of 12 eggs before he knew they werethere.The nest is generally a slight depression in the moss, lined withdead grass and leaves. Lucien M. Turner describes a nest foundin the neighborhood of Fort Chimo, Ungava, as " merely a fewgrass stalks and blades loosely arranged among the moss of a higherspot under the drooping limbs of a spruce situated in a swamp. Afew feathers of the parent bird were also in the nest."A. D. Henderson describes a nest as follows : " It was in a muskeg,a slight hollow in the moss scantily lined with a few twigs and leaves
HUDSONIAN SPRUCE GROUSE 123
of the Labrador tea. It was under a moss-covered, dead, fallen sprucebranch beneath a low-branching green spruce. The sitting bird wasvery reluctant to leave the nest." Another " nest was in a hollowlined with dry leaves and spruce needles under a small spruce bush,about 2 feet high, on the edge of a muskeg."Eggs.?[Author's note: The spruce grouse and its near relative,Franklin's grouse, lay the handsomest eggs of any of the grouse.Ten or a dozen eggs usually make up the set, but as many as 14 oreven 16 have been found in a nest. Sets of less than eight are proba-bly incomplete. The eggs vary in shape from ovate to elliptical ovate.The shell is smooth with a very slight gloss. The ground colorsvary from " cinnamon " to " pinkish buff," or from " cream-buff "to "cartridge buff." They are usually boldly and handsomelymarked with large spots and blotches of rich browns, sometimesmore sparingly marked and sometimes thickly and evenly coveredwith small spots and dots. The colors of the markings vary from
" chestnut-brown " or " chocolate " to " hazel " or " russet." One oddset in my collection has a " cartridge buff " ground ; one egg is nearlyimmaculate and the others are sparingly, or only slightly, spottedwith " bone brown." The measurements of 54 eggs average 43.5 by31.7 millimeters; the eggs showing the four extremes measure 47.1 by34, 40 by 30.4, and 40.1 by 29.8 millimeters.]Young.?The duration of incubation is about 17 days, accordingto Lucien M. Turner. Incubation is performed by the female andshe alone looks after the young. Bendire (1892) states that "anegg is deposited every other day, and incubation does not begin untilthe clutch is completed." Turner, however, states that " layingbegins about the fifth of June and incubation about the twelfth " inUngava.The young are able to run about and follow the mother almost assoon as their feathers dry, and they are able to fly vigorously at anearly age, when they appear about a quarter of the size of the adults.One of these young, a good flyer, that I collected at Shecatica Inlet,Canadian Labrador, on July 23, 1915, measured 5 inches in length,and its wing measured 3.5. The adult's length is about 13 inches andthe wing 6.5. On this occasion the brood of young startled me byflying up with a slight whirring sound almost from under my feet.They flew to the branches of a low spruce, while the mother appearedmost conspicuously, standing in a bed of curlew-berry vines andreindeer lichen, with head up and tail erected. As a rule the young-fly off and conceal themselves so thoroughly that it is difficult to flushthem again, while the mother, clucking and ruffling her feathers,flies to a spruce tree or remains on the ground, in both cases allowingan approach to within a few feet. On one occasion, when I was in
124 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthe marshes about the mouth of the St. Paul River, Canadian Labra-dor, the female stood on a small rock and crooned, while the young,one after the other, until seven in all, flew and joined her on the rock.Mr. Bent contributes the following note on the behavior of motherand young when the latter are still unable to fly; this was nearHopedale, Labrador:We caught one of the young and had an interesting time watching the motherbird in her solicitude. Her boldness was remarkable and she showed no fearwhatever ; we could walk right up to within a few feet of her, on the groundor in small trees. We tied the young one with a string and, as soon as themother heard its peeping notes, she came right up to it, clucking and scolding,with her feathers all ruffled up and her tail spread like a turkey's ; she struttedaround over the logs and rocks near us ; her soft clucking notes sounded likekruk, kruk, kruk, with an occasional rolling note like krrrrruk, soft and low.The young evidently understood it as a danger signal, for they remained sowell hidden that we found only two of them. While I was photographing thecaptive little one, the mother came almost near enough to touch and even ranbetween the legs of the tripod.Plumages.?[Author's note : In the small, downy chick the generalground color is yellowish buff, varying from " chamois " above to
" colonial buff " below ; there is a black spot at the base of the culmen,a larger one in the middle of the forehead, one on each of the lores,and a broken stripe on the auriculars; there is a large patch of
''hazel," bordered with black, on the crown and occiput; the backand rump are washed with " hazel " and " tawny " and indistinctlyspotted with black; the underparts are unmarked.The juvenal plumage appears at an early age, beginning with thewings ; these start to grow within the first 5 daj^s and, at the age of10 or 12 days, the wings reach beyond the tail and the young bird canmake short flights. By the time the young bird is half grown it isfully feathered. In this juvenal plumage the sexes are alike andresemble the adult female, but are browner above, rustier on the headand neck, and whiter on the chin. The crown is " cinnamon-rufous "or "' hazel," spotted with black; the back, scapulars, and wing covertsare " ochraceous-tawny " or " tawny," boldly patterned with blackblotches or bars, and with broad, median, buffy stripes with triangu-lar white tips ; the remiges are sepia, the primaries narrowly notchedwith buff, the secondaries edged with buff, and the tertials barredand spotted with " ochraceous-tawny " ; the pointed rectrices are" ochraceous-tawny," heavily barred and peppered with black ; thebreast is " ochraceous-tawny," spotted with black, the belly grayishor yellowish white, faintly spotted with dusky, and the chin andthroat yellowish white.Beginning early in August and lasting through September, thepostjuvenal molt takes place, during which the sexes begin to differ-entiate, the young males showing patches of black feathers in the
HUDSONIAN SPRUCE GROUSE 125breast. This molt is complete except for the two outer primaries oneach wing. It begins on the breast, extends to the flanks and back,and is finally completed on the throat and crown. When this moltis completed in October, young birds can hardly be distinguishedfrom adults, though there is more white in young birds and the outerprimaries are diagnostic.Adults probably have a very limited prenuptial molt about thehead and neck in spring; and they have a complete postnuptial moltin August and September. The Hudsonian spruce grouse is a grayerbird than the Canadian, with rather more white and purer gray in themale ; the difference is even more pronounced in the female, which ismuch more purely black and gray, with much less buffy or ochraceous.]Food.?The spruce grouse not only lives in spruce woods butdepends upon the buds, tips, and needles of the spruce, as well asof the fir and larch, for a considerable part of its diet. This isparticularly the case in winter when snow and ice cover the ground,concealing many berries, which it enjoys eating in summer. In thelatter season, I have found in Canadian Labrador the followingstomach contents of this bird : A young able to fty had eaten 5 redspiders, 10 green snowberries, and 75 achenes of a bulrush (Scirpuscaespitosus) ; an adult had eaten 25 snowberries, 20 crowberries(Empetrum nigrum), and many leaf tips of a dwarf bilbern^{Vaccinium ovafolia). Another adult had been feeding entirely oncrowberries and a third had "been eating the leather woodfern(Dryopteris marginalis). Bearberries (Arctostaphylos uva-ursi),as well as grass and weed seeds and various insects, including grass-hoppers, are also eaten in summer, although the regular diet ofspruce is not entirely given up.E. A. Preble (1908) found nothing but spruce needles in thestomachs of four spruce grouse. Another, taken on the shore of aninlet had in its crop several mollusks (Lymnoea palustris). Thecrops of a number taken late in fall and in winter contained onlythe needles of the jack pine {Pinus banksiana). A young birdjust ready to fly had eaten bits of the American rockbrake fern{Cryptogramma acrostichoides) , blueberries {Vaccinium uligi-nosum), and mountain cranberries (F. vitisidaea) .Lucien M. Turner says
:
The food of the spruce partridge consists of the tender, terminal buds ofspruce ; and this, in winter, seems to be their only food * * * mixed with,at times, an astonishing quantity of gravel. I was surprised to find thesestones of such uniformity of size and material. Crystallized quartz fragments,in certain instances, formed alone the triturating substance.If a bird be opened when just killed the contents of the gizzard has apowerful terebinthine odor which quickly pervades the flesh and renders ituneatable to a white person. In the spring and summer months these birds
126 BULLETIN 162, UNITED STATES NATIONAL MUSEUMconsume quantities of berries of Empetrum and Yaccinium and in the fall theflesh of the young * * * has a fine flavor, and, as the meat is white, it isvery acceptable.In some instances the flesh of adults in summer is also free from anytaste of turpentine. According to A. L. Adams (1873) it is saidthat the flesh is sometimes poisonous when the birds have been eatingmountain-laurel berries.Behavior.?The chief characteristic of the Hudsonian sprucegrouse is its unsuspicious character, which amounts, indeed, to stupid-ity. This is illustrated by an experience of Lucien M. Turner, whosays:I once shot 11 and did not move a yard in distance to do so. The people ofLabrador employ a method which they term " slipping," i. e., a slip noose on along pole which enables the holder to slip the noose over the heads of the birdsand jerk them down. One who is expert in this method rarely fails to obtainall the birds within reach.D. G. Elliot (1897) says: " I have seen birds push this noose asidewith their bills without changing their position, when throughawkwardness, or unsteadiness of hand on account of a long reach,the noose had touched the bird's head but had not slipped over it."I have known a botanist to kill an adult grouse by throwing hisshort-handled collecting pick at it.The plumage of spruce grouse often makes them difficult to dis-tinguish from their surroundings, and if their tameness depends onthis protective coloration, they are overconfident, for, in a setting ofreindeer lichen or snow, or an open branch of a spruce, they arevery conspicuous. When flushed they generally fly only a few yardsor even feet, and, alighting in trees, they continually thrust thehead and neck now this way, now that, and appear to be blindlytrying to discover what has disturbed them. As a rule the flightis noiseless, or a slight sound only is heard, but at times they rise witha loud whir of wing beats.I have already mentioned the tameness or boldness of the femalebird with her brood. This boldness is also shown at the nest contain-ing eggs. J. Fletcher Street writes:The nest was somewhat hidden under a dense spruce shrub, and while Iwas cutting away some of the inclosing branches to obtain a better view, thebird left and at first charged toward me. Then she withdrew and kept retreat-ing as I approached toward her, keeping about 10 feet between us. Sheexhibited but little concern after having left the nest but would not returnto it while I remained nearby. I left the locality for three definite periodsand upon each return found the bird sitting upon the eggs, yet becoming morewary at each successive disturbance.Referring to this bird, Joseph Grinnell (1900) says: "After thesnow came, grouse were seldom found for they remained continually
HUDSONIAN SPRUCE GROUSE 127in the trees. I saw but few tracks on the snow all winter, thoughin the fall their tracks were numerous on the sand-dunes and amongwillows along the river." According to A. Leith Adams (1873),they do not dive under the snow like the ruffed grouse. Lucien M.Turner says: "I have reason to suspect that some of these birdsretain their mates for more than one season as I have frequentlyfound a pair together in the depth of winter and these two beingthe only ones of the kind to be found in the vicinity." Audubon(1840) says that "the males leave the females whenever incubationhas commenced, and do not join them again until late in autumn;indeed they remove to different woods, where they are more shyand wary than during the love season or in winter." He also im-parts the following curious information
:
All the species of this genus indicate the approach of rainy weather or asnow storm, with far more precision than the best barometer; for on theafternoon previous to such weather, they all resort to their roosting placesearlier by several hours than they do during a continuation of fine weather.I have seen groups of grouse flying up to their roosts at mid-day, or as soonas the weather felt heavy, and have observed that it generally rained in thecourse of the afternoon. When, on the contrary, the same flock would remainbusily engaged in search of food until sunset, I found the night and the follow-ing morning fresh and clear.D. G.Elliot (1897) says:The spruce grouse is found usually in small flocks consisting generally of onefamily, but also old males are met with alone, and I have always regarded itas a bird that was rather fond of solitude. Frequently, even in autumn, whenthe nights were becoming frosty, and snow flurries would hide the sun by day,heralding the coming winter, I have seen an old male, in the recesses of aswamp, strut about with ruffled feathers and trailing wings, as if the air werebalmy and mild and spring were at hand.Tales are told of immense numbers of these birds collecting in greatflocks, or " packs," but such collections have probably not occurredfor many years.Voice.?The spruce grouse is a silent bird except when disturbed.His courtship " song " is instrumental, made by the rapid " drum-ming " of the wings striking the air as already described. I haveheard slight clucking sounds from }roung birds and somewhat similarduckings from adult males. Adult females when disturbed cluckincessantly, a sound described by Mr. Bent as " kruk, kruk, kruk,with an occasional rolling note like km^rrruk, soft and low." Streetsays : " The only note that the grouse uttered at any time was a lowchuck chuck upon the occasion of her first leaving the nest." For-bush (1927) records the voice of the immature male as " a lowwailing whistle, weeo-weeo-weeo?
128 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMField marks.?The male is a handsome bird distinguished by itscompact form, its jet-black breast contrasting sharpW with whito,its red combs over the eyes, and its yellow-tipped tail. The femaleis a plain brown bird barred with black above, in this way differingfrom the ruffed grouse, which is spotted. It is smaller than theruffed grouse and has a shorter tail.DISTRIBUTIONRange.?Northeastern United States, Canada, and Alaska.The spruce partridge is nonmigratory. Its range extends northto Alaska (Noatak River, Coldfoot, Fort Yukon, and Circle) ; Yu-kon (latitude G6? 40' N.) ; Mackenzie (Mackenzie River, Fort Frank-lin, Lake Hardisty, Gros Cape, and Fort Simpson) ; northern Sas-katchewan (Cochrane River) ; northern Manitoba (Lac du Brochet,Fort Churchill, and York Factory) ; northern Ontario (Fort Sev-ern) ; northern Quebec (Fort George, Great Whale River, FortChimo, and Whale River) ; and Labrador (Okkak). East to Lab-rador (Okkak) ; eastern Quebec (Rigolet, Groswater Bay, head ofthe Magdalen River, and Mount Albert) ; northeastern New Bruns-wick (Bathhurst) ; Nova Scotia (Baddeck, Canso, Halifax, andShelburne) ; eastern Maine (Fort Fairfield, Mount Katahdin, Houl-ton, Kingman, Calais, Orono, and North Livermore) ; and south-ern New Hampshire (Dublin). South to southern New Hampshire(Dublin) ; northern New York (Raquette Lake) ; southern Ontario(Kingston, Peterboro, and Bradford) ; northern Michigan (Au Sa-ble River, Vans Harbor, and Palmer) ; northern Wisconsin (MamieLake) ; northern Minnesota (Northern Pacific Junction, Leech Lake,and Hallock) ; southern Saskatchewan (Fort Pelly and Osier)
;
central Alberta (Mundare, Blueberry Hills, and Simpson Pass)southeastern British Columbia (Goat Mountain) ; and northernWashington (Chopaka Mountain and Barron). West to northwest-ern Washington (Barron) ; northwestern British Columbia (FloodGlacier, Glenora, and Atlin) ; and Alaska (Chilcat, Kodiac, Nusha-gak Lake. Aleknagik, Bethel. Russian Mission, Nulato, Kowak River,Kotzebue, and Noatak River).Spruce partridges are of casual occurrence in Massachusetts(Gloucester, in 1851, and Roxbury, about 1865).The 1931 edition of the American Ornithologists' Union CheckList of North American Birds recognizes four races of Canachitescanadensis, all of which are included in the foregoing ranges.True canadensis is found from the Labrador Peninsula west to theeastern base of the Rocky Mountains west of Edmonton, Alberta.Canachites c. canace ranges over the Maritime Provinces of Canada(New Brunswick and Nova Scotia) ; northern New England and
ALASKA SPEUCE GROUSE 129New York; southern Quebec, Ontario, and Manitoba; and northernMichigan, Wisconsin, and Minnesota. It is now largely extinct inthe southern part of its range. Canachites c. osgoodi is found fromGreat Slave Lake and Athabaska Lake west to the Yukon regionand the Mount McKinley Range of Alaska, while Canachites c.atratus occupies the coast region of southeastern Alaska.Egg dates.?Central Canada (canadensis) : 8 records, May 23 toJune 14. Labrador Peninsula : 11 records, June 1 to July 4.Alaska (osgoodi) : 9 records, May 11 to June 25.Quebec to Nova Scotia and Maine (can-ace) : 21 records, May 5 toJune 24; 11 records, May 24 to June 2.CANACHITES CANADENSIS OSGOODI BishopALASKA SPRUCE GROUSEHABITSThe Alaskan form of the spruce grouse, or spruce partridge, wasdiscovered and described by Dr. Louis B. Bishop (1900) and namedin honor of his companion on the Yukon River trip, Dr. Wilfred H.Osgood. He described it as " similar to Canachites canadensis butwith the ochraceous buff bars replaced everywhere by cream-buffand grayish white. On the upper parts the gray tips are paler, theochraceous buff replaced by cream-buff and whitish, and the palebars of the cervix grayish white instead of buff ; below the white tipsare larger, the pale bars whitish and cream color instead of buff,becoming cream-buff only on the jugulum."Doctor Osgood (1904) says of its habitat:The range of the spruce grouse is practically coextensive with that of thespruce tree. We traveled much of the time near the western limit of thetimber, and found grouse fairly common, even up to the edge of the tundra,where the spruce was considerably scattered. The last one seen was a finecock, which was started very early on the morning of September 10, from asmall beach on the Nushagak River about 25 miles above its mouth. Thegrouse are said to occur within a very few miles of Nushagak, however.Herbert W. Brandt contributes the following notes on his experi-ence with this grouse in Alaska
:
The Alaska spruce grouse proved to be a common bird throughout the woodedarea that we traversed while en route to Hooper Bay. We first met with itabout 60 miles west of Nenana, and from that time thereafter, when we werein the spruce areas, we were continually coming upon it. This noble fowl wascommon in the spruce timber right up to the highest pine growth in the BeaverMountains, but its apparent preference is for the densely grown spruce riverbottoms. The " fool hen's " noted lack of fear was often in evidence, and itsretreating from ahead of our caravan often quickened the pace of the chase-loving dogs. It proved to be much more arboreal in habits than the otherAlaskan gallinaceous birds we encountered, for we seldom saw it on the ground,
130 BULLETIN 162, UNITED STATES NATIONAL MUSEUMand its snow tracks are rarely observed, which contrasts markedly with thenetwork of telling ptarmigan trails that everywhere enliven the barren snowwastes. In the heart of the dense spruces, such as it frequents, the beautifuldark plumage pattern gives it almost complete coloration protection, and ifit did not reveal itself by movement, this bird would seldom be observed.Nesting.?According to A. H. Twitchell the Alaskan spruce grouse nestsregularly in the vicinity of the Beaver Mountains at the head of the DistnaRiver, and here he has noted nests containing from five to eight eggs. Onenest found by him on June 10, 1924, was placed out in the open in winter-driedgrass near a small, dead spruce, which afforded but scant concealment. Thisbird chose a site about 100 yards from the reindeer corral in a scattered growthof small spruce, and in an area where there was often considerable activity.The male bird was frequently seen flying about, but the female was very wary.The nest was sunken 5 inches in the moss and made of circular-formed drygrass and a few dead spruce twigs, and contained a number of feathers of thesitting bird. The nest was found on June 1, when it contained one egg, and onJune 10 the clutch numbered eight eggs, all of which proved to be fresh. Duringthe egg-laying period, when the bird was off the nest the eggs were hiddenbeneath a covering of surrounding dead vegetation, artfully arranged beforethe bird departed.Eggs.?In shape, the egg of the Alaskan spruce grouse is elongate-ovate, andthe surface reflects a noticeable luster. The shell is somewhat greasy and quitesturdy. The ground color is prominent, as the spots occupy less than one-thirdof the surface, and the egg, on account of the bold richness of its markings, isquite handsome. The ground color varies from " salmon color " to " chamois "and " cream buff." The spots are flecked over the entire surface, but are sparseston the larger end. In size these markings range from dots to those the size of apea. There are only a few of the larger spots on each egg and these are wellscattered over the surface, seldom exhibiting confluence. In contour the spotstend to be circular, with their rims well defined. When the pigment is thin thecolor is " chestnut " to " chestnut-brown," but the usual shade is " haematitered." When the egg is newly laid it is evident that both the ground color andmarkings are soft and moist, like those on the egg of the willow ptarmigan, aseach egg somewhere on the surface is streaked with feather scratches, whichoften show distinctly the individual feather barbs. Occasionally a considerablearea is so rubbed, exposing the ground color. When the egg dries, however, themarkings are very durable.Food.?Dr. Frank M. Chapman (1902) quotes from J. D. Figgins'snotes, as follows
:
In all the timber region I visited, the Canada Grouse was found common andbreeding. Their chief food during early summer is the leaves of various decidu-ous bushes and spruce needles. About the 1st of August they repair to theedge of the barren grounds for berries which are then ripening. These aretheir food until September, when they return to the timber where raspberriesand currants are abundant. During winter and spring their food consists en-tirely of spruce needles. Both adults and young appreciate their protectivecoloration, and when approached remain perfectly motionless until the dangeris past. During the winter their color is to their disadvantage, and they becomevery shy and will not allow a close approach.
CANADA SPRUCE GROUSE 131Near the base of the Alaska Peninsula, Doctor Osgood (1904)found these grouse in abundance about Lake Clark, " more commonthere than " he had " ever found them elsewhere in Alaska." He says
:
They feed largely cm berries in the summer time, being particularly fond ofthose of Vaccinium vitis-idaea, which they eat almost exclusively from the timethe little green berry first begins to swell until it is dead ripe. At this time theflesh of the birds is sweeter than in the early winter, when a diet of spruceneedles has made them fatter but less palatable. In the spruce forest, which istheir ordinary habitat, they are unable to obtain on the moss-covered groundthe grit necessary for a gallinaceous bird, so they make daily excursions to theshores of the rivers and lakes where fine gravel is to be had in abundance.Early morning before sunrise is the time for this ; then they may often be seenon the beaches, singly, in pairs, or in small flocks. Doubtless they also come tothe rivers to drink, though pools are common enough in the swampy openingsin the timber. On the Chulitna River one was caught in a steel trap which hadbeen set for a possible mink or weasel in the marsh grass at the water's edge.CANACHITES CANADENSIS CANACE (Linnaeus)CANADA SPRUCE GROUSEHABITSThis is the form of the spruce grouse found in extreme southernCanada and the extreme Northern States, east of the Eocky Moun-tains. The male is practically indistinguishable from the male ofthe Hudsonian spruce grouse, but the female is decidedly morerufous or rusty, both above and below. The haunts and habits ofthe two are practically identical.Edward H. Forbush (1927) has given us the following attractivedescription of its haunts
:
In the dense spruce, fir, cedar and tamarack swamps of the great Mainewoods the Spruce Grouse dwells. Where giant, moss grown logs and stumps ofthe virgin forest of long ago cumber the ground, where tall, blasted stubs ofothers still project far above the tree-tops of to-day, where the thick carpet ofgreen sphagnum moss deadens every footfall, where tiny-leaved vinelets radiateover their mossy beds, there we may find this wild bird as tame as a barn-yardfowl. In the uplands round about, there still remain some tall primeval woodsof birch and beech and rock maple where the moose and bear have set theirmarks upon the trees. In winter the deer gather in the swamps, and there theirmany trails wind hither and yon. Gnarled, stunted trees of arbor vitae, somedead or dying, defy the blasts of winter, while the long, bearded Usnea droopsstreaming from their branches.An equally satisfactory account comes from the facile pen ofWilliam Brewster (1925) :For the most part the birds frequent dense, matted growths of cedar (i. e.arbor vitae), black spruce, and hackmatack (American larch), overspreading,low-lying, flat, and more or less swampy lands bordering on sluggish streamsor on semiopen bogs similar to those known as Muskegs in the far North. Fromsuch coverts they wander not infrequently up neighboring hillsides to evergreen
132 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMforests on still higher ground beyond, or perhaps into neglected pastures chokedwith intermingling young balsams, red spruces, and white spruces no morethan eight or ten feet tall. Nor are they unknown to appear well out in ratherwide upland clearings, where the only available cover consists of thickets ofraspberry bushes, or even in river?or brook?meadows, where it is furnishedsolely by rank grass. Ramblings, thus venturesome, are exceptional, of course,and undertaken, I believe, at no seasons other than late summer and earlyautumn, when the lowly vegetation that clothes such perfectly treeless groundis most luxuriant, and also best supplied with berries or insects of variouskinds ; these Spruce Partridges devour eagerly whenever, and wherever, theycan obtain them readily, although subsisting during the greater part of theyear on a nearly unmixed diet of spruce and balsam spills (leaves), pluckedmostly from branches at least fifteen or twenty feet above the ground.Courtship.?William Brewster (1925) gives a slightly differentaccount of this from what others have given; he relates LumanSargent's experience with it, as follows:Many years ago he was skirting a dense swamp, when his attention wasattracted by a peculiar whirring sound that came from it. Advancing cau-tiously he soon perceived two Spruce Partridges, cock and hen, together on theground. The cock left it presently, and vibrating his wings with great rapiditybegan mounting upward in a spiral course around the trunk of a large balsam,producing all the while a continuous drumming sound. After rising to a heightof about 20 feet, and making three or four complete turns around the stem ofthe tree, he alighted on one of its branches where he rested for a moment ortwo and then flew down just as he had risen, that is by circling spirally aroundthe trunk, with the same uninterrupted sound of wings. On reaching the spotwhere he had left his mate, he strutted about her like a Turkey cock, with wide-spread tail. Luman saw all this repeated fifteen or twenty times. For thefirst 10 feet above its base the trunk of the balsam was smooth and bare, butabove that the Partridge had to conduct his drumming flights, both upward anddownward, through numerous stiff branches. The sound of his drumming wasdistinctly audible at least 50 yards away.Watson L. Bishop's account, quoted by Bendire (1892), of thedisplay of a male bird in captivity is as follows
:
The tail stands almost erect, the wings are slightly raised from the bodyand a little drooped, the head is still well up, and the feathers of the breastand throat are raised and standing out in regular rows which press the feath-ers of the nape and hind neck well back, forming a smooth kind of cape onthe back of the neck. This smooth cape contrasts beautifully with the ruffledblack and white feathers of the throat and fore breast. The red comb overeach eye is enlarged until the two nearly meet over the top of the head. Thiscomb the bird is able to enlarge or reduce at will, and while he is struttingthe expanded tail is moved from side to side. The two center feathers do notmove, but each side expands and contracts alternately with each step asthe bird walks. This movement of the tail produces a peculiar rustling, likethat of silk. This attitude gives him a very dignified and even conceitedair. He tries to attract attention in every possible way, by flying from theground up on a perch, and back to the ground, making all the noise he canin doing so. Then he will thump some hard substance with his bill. I havehad him fly up on my shoulder and thump my collar. At this season he isvery bold, and will scarcely keep enough out of the way to avoid being stepped
CANADA SPRUCE GROUSE 133
on. He will sometimes sit with bis breast almost touching the earth, hisfeathers erect as in strutting, and making peculiar nodding and circularmotions of the head from side to side ; he Will remain in this position two orthree minutes at a time. He is a most beautiful bird, and he shows by hisfictions that he is perfectly aware of the fact.Nesting.?The nesting habits of this grouse are not essentiallydifferent from those of its more northern relative. Brewster (1925)records a nest found by Aldana Brooks near Richardson Lake, Me.,
" where the land was low and wooded with black ash, birch, alder,and a few larches. It was sunk, he said, in the top of a little moundwith no rock, log, or even tree-trunk very near it. There were nineeggs. The bird did not leave them until almost stepped on, whenshe fluttered off over the ground for a few yards, and then stoppedto watch Brooks who finally continued on his way without molest-ing her, or taking any of the eggs, w7hich he never saw again.''Watson L. Bishop (1890), who was succeeded in domesticatingthis bird, says
:
As the nesting season approaches I prepare suitable places for them byplacing spruce boughs in such a way as to form cozy little shelters, wherethe birds will be pretty well concealed from view. I then gather up someold dry leaves and grass and scatter it about on the ground near where Ihave prepared a place for the nest. She will then select one of these places,and, after scratching a deep, cup-shaped place in the ground, deposit in it hereggs ; * * * if there should be sufficient material within easy reach ofthe nest the bird will sometimes cover the eggs up, but not in all cases.No nesting material is taken to the nest until after three or four eggs arelaid. After this number has been deposited, the hen, after laying an egg,and while leaving the nest, will pick up straw, grass, and leaves, or whateversuitable material is at hand, and will throw it backward over her back asshe leaves the nest, and by the time the set is complete, quite a quantity ofthis litter is collected about the nest. She will then sit in her nest and reachout and gather in the nesting material and place it about her, and when com-pleted the nest is very deep and nicely bordered with grass and leaves.So strong is the habit, or instinct, of throwing the nesting materials overthe back, that they will frequently throw it away from the nest, instead oftoward it, as the hen will sometimes follow a trail of material that will turnher " right about " so that her head is toward the nest, but all the time she willcontinue to throw what she picks up over her back. This, of course, is throw-ing the material away from the nest. Discovering her mistake, she will then
" right about face " and pick up the same material that an instant before wasbeing thrown away, and throw it over her back again toward the nest.E99s -?The eggs of the Canada spruce grouse are indistinguishablefrom those of the Hudsonian race, already described. The measure-ments of 53 eggs average 43.2 by 31.1 millimeters ; the eggs showingthe four extremes measure 47.5 by 31.5, 46.5 by 32.5, 39.9 by 31, and40.4 by 29 millimeters.Food.?On dissecting some young birds shot on a meadow, on Sep-tember 11, Brewster (1925) " found in their crops very many grass-74564?32 10
134 BULLETIN 162, UNITED STATES NATIONAL. MUSEUMhoppers of various kinds and sizes, numerous ripe raspberries, a fewleaves of Spiraea fomentosa and (in one crop only) a few larchspills." Evidently these birds had wandered out of their usual hauntsfor a change of diet. He says further
:
In this connection it may be well to add that in the crops of two youngSpruce Grouse only about half-grown and killed in the Tyler Bog on August 13,1873, I found raspberries, blueberries, checkerberries, and balsam buds as wellas needles ; that from the crop of an adult female shot near MollidgewaukStream on September 28, 1890, I took 51 berries of Viburnum lentago?, somefragments of small mushrooms, and a few spills of the black spruce ; and thata young male found and killed in company with the old female just men-tioned had in his crop 13 Viburnum berries, uncounted pieces of mushrooms,and a few larch spills. Hence it will appear that food of various kinds otherthan that supplied by the foliage of coniferous trees is partaken of ratherfreely by Spruce Grouse in late summer and early autumn.Behavior.?Although they usually flutter awkwardly away orsilently fly for very short distances in the thick woods, when flushedin the open by Brewster (1925) they behaved quite differently, "ris-ing all at once like Quail, from within a space no more than two yardssquare, with what seemed a deafening roar of wings, they spedstraight for the woods, flying precisely like Ruffed Grouse and quiteas swiftly.''Forbush (1927), while walking on a trail, almost stumbled overa male spruce grouse. He says
:
The bird was somewhat startled and flew heavily up into a near-by spruce,alighting near the tip of a little limb about 20 feet from the ground. As thelimb drooped under his weight, he walked up it to the trunk, hopped up abranch or two higher, and immediately began to feed on the foliage. After afew minutes of this, he moved a little into another tree and continued feeding.Pounding on the trunk with an axe did not alarm him, and it was only afterseveral sticks had been thrown and one had hit the very limb on which he satthat he was induced to fly.Edwyn Sandys (1904) writes:The writer has twice caught mature specimens with his bare hands, and itis a common trick of woodsmen to decapitate a bird with a switch, or noose itwith a bit of twine. Once the writer came precious near hooking one with atrout fly, at which the grouse had pecked. Only a dislike to needless cruelty, anda respect for a fine rod, saved this particular bird. Quite often the brood is metwith in the trail, when they will sedately step aside about sufficiently far tomake room for the intruder's boots, meanwhile regarding him with a laughableair of affectionate interest. No doubt this grouse could fly rapidly should itchoose to exert its powers, but it is content with more leisurely movements.Game.?The spruce grouse is not much esteemed as a game bird, asit lacks most of the qualities that appeal to the sportsman. Its hauntsare usually too difficult to hunt in, it is too tame and stupid to makeits pursuit interesting, and, except when on rare occasions it is foundin open clearings or on meadows, it seldom offers a flying shot. It is
VALDEZ SPRUCE GROUSE 135
not highly regarded as a table bird, for its flesh is said to be unpalat-able. This is probably so in winter, when it has been feeding onspruce and balsam leaves; its flesh is then dark and decidedly resin-ous in flavor. But during fall, when it feeds largely on berries, greenherbage, and insects, its flesh has a very different color and flavor.Brewster (1925) says that some young birds, shot in September,
" proved delicious eating, their flesh being much sweeter and finerflavoured than that of any Kuffed Grouse. Both before and aftercooking it was nearly as white as the Ruffed Grouse's, where as thefully-matured Spruce Grouse has invariably dull reddish flesh some-what too redolent of spruce foliage to be relished by everyone,although I do not dislike it. The flesh of at least some of the youngbecomes, almost, if not quite, as dark as that of the adults, by thelast of September."These birds are killed for food all through the fall and winter byhunters of large and small game, by lumbermen, and by trappers andothers. They are so easily killed that they are disappearing veryrapidly and are now very scarce in northern New England in anybut the most inaccessible regions.Winter.?Walter H. Rich (1907) writes:During a snowstorm the Spruce Grouse usually flies up into the densestclump of spruce or fir trees in the neighborhood, and, under their thick, archingbranches, snow-laden and bending, he finds shelter from the weather and foodin abundance. He may not leave the tree for several days if undisturbed andthe storm continues. The question of temperature troubles him little, andwith his wants all provided for, the Spruce Grouse is more independent in hismode of life than any of his feathered neighbors, for when other birds arescurrying about for something to eat and perhaps going hungry, this gentlemanfinds plenty of food in his shelter, and sits in comfort, " at ease in his own inn."CANACHITES CANADENSIS ATRATUS GrinnellVALDEZ SPRUCE GROUSEHABITSDr. Joseph Grinnell (1910), in describing the spruce grouse of thecoast region of southeastern Alaska, says that it " resembles Gana-chites canadensis osgoodi of the interior of Alaska (Yukon andKowak Vallej's), but general tone of coloration darker: white mark-ings less in extent; black areas more extended; and grays less ashy,more olivaceous."Of its distribution he says : " The indications are that this formis generally distributed in the humid coast belt from the easternside of the Kenai Peninsula southwestwardly at least as far asHawkins Island, and probably beyond."
136 BULLETIN 162, UNITED STATES NATIONAL MUSEUMReferring to its haunts and food, he writes
:
Spruce grouse were not abundant in the Prince William Sound region, butappeared to be generally distributed. Two shot on Hawkins Island were bothin heavy timber near the beach. Their crops were filled with the fresh greenleaf-buds of spruce. On Hinchinbrook Island a male was shot on the moun-tain side near timber-line. The example secured by Heller on Hoodoo Islandwas flushed from a rank growth of salmonberry bushes ; its crop containedberries, some fern fronds and a few seed pods of the devils-club. Grouse signwas noted on Chenega Island ; and, as previously noted, skins were secured atKnight Island and at the head of Port Nell Juan.CANACHITES FRANKLINI (Douglas)FRANKLIN'S GROUSEHABITSThis handsome species might well have been named the westernspruce grouse, for it is the western counterpart of the well-knownspruce grouse of eastern and northern Canada. It lives in similarhaunts, has similar habits, and is so closely related to the sprucegrouse that it may eventualty be shown to intergrade with some ofthe western races of canadensis and be reduced to subspecific rank.It occupies a comparatively limited range in the mountainous interiorof the Northwestern States and southwestern Canada.While stationed in Idaho, Major Bendire (1892) found these grousequite commonalong the edges of wet or swampy mountain valleys, the so-called " Camasprairies," or the borders of the numerous little streams found in such regionsamong groves or thickets of spruce and tamarack. Few naturalists have asyet been sufficiently interested to invade their favorite haunts. In the sum-mer of 1881 I found a single covey, numbering about ten birds, in the lowflat and densely timbered region between the southern end of Pend d'OreilleLake (the old steamboat landing) and Lake Coeur d'Alene, Idaho, at an alti-tude not exceeding 3,500 feet, I should think. I bagged three of these birds,and was quite surprised to find them in such a locality. As far as I have beenable to learn, they usually occurred only at altitudes from 5,000 to 9,000 feet,and scarcely ever left the higher mountains. They were scratching in thedust on the trail I was following, and simply ran into the thick underbrushon each side, where they were quickly hidden.Courtship.?Thomas T. McCabe has sent me the following interest-ing notes on the courtship of Franklin's grouse, as observed by himand Mrs. McCabe in British Columbia on May 28, 1929:In the course of a morning's nest hunting in second-growth spruce andbalsam, carpeted with deep green moss, we had found the cock in his usuallocality, sitting quietly on a tussock, showing no erection of the crimson combs,no inclination to display, and typically indifferent to us. An hour later anda little before noon we were about 200 yards from this point, when he ap-peared above us, flying through the tree tops, and lit in a spruce about 15
franklin's grouse 137feet from the ground and close to a pack trail. He still displayed no unusualexcitement, and we left him again for 15 minutes. When we came back thehen had appeared and was squatting flat on the ground in the center of thebeaten trail. Her appearance was normal and remained so through the ensuingepisode. Perhaps 20 feet away the cock was walking down the trail towardher in a typical attitude of display?head drawn up and back, tail spreadthrough two-thirds of a circle and vertical (not bent forward over the back),the fine undercoverts falling back from it like the sticks of a fan, the wingpoints slightly dropped, the combs bulged upward into elongate crimson rolls,which met in the center of the cranium. In this guise he strutted very slowly,with a statuesque pause of about 8 seconds every 2 or 3 feet.At a distance of about 10 feet the whole bird was transformed with thesuddenness of a conjuring trick, and the similarity to the courting ruffedgrouse disappeared. The tail snapped together and sank nearly to the ground.The head was lowered and extended far forward. The plumage was flattenedso that a hard sleekness replaced the fluffy rotundity, and the size of the birddiminished by half. The attitude was like that often assumed by Bonasa orLagopus when luring an intruder away from nest or young, but in place oftheir whining note a low guttural was produced, vibrant and threatening, infrom 5 to 7 periods, the first two distinct and slow, the remainder losing intervaland less sonorous, trailing off to silence. This was accompanied by a slightmovement of the tail in the vertical plane, rather a periodic trembling than asnapping like that of the courting Bonasa. One of us thought that with thismovement the rectrices were slightly opened. Between these utterances thebird moved 3 to 6 feet, very slowly, and not in short runs after the manner ofBonasa. The movements were in various directions, a few perhaps directlytoward the hen, but for the most part oblique and keeping 5 or 6 feet away.After about 4 minutes of this the hen took wing, silently, but with amazingsuddenness and speed for so phlegmatic a bird, flashed down the trail, and madea quick turn into the woods. Her sudden start scarcely gained a foot on theeager male, and both disappeared together. Perhaps a final scene was enactednear by. Perhaps many more were set, with many variations, before the subtleinterplay of impulse and reaction rose to its climax. The details of the primi-tive drama, with its suggestions of threatening, beseeching, lamenting, liebeyond our power of interpretation, but, except in the opening movement, theelement of simple and lavish display, so widespread among other genera, wasabsent.Nesting.?The nesting habits of Franklin's grouse are similar tothose of the spruce grouse. The eggs are very rare in collections, asthe nests have seldom been found, and very little has been publishedabout them. Major Bendire (1892) writes:Through the kindness of Mr. W. E. Traill, in charge of one of the HudsonBay Company posts in British Columbia, parts of three sets of these rare eggs,fifteen in number, were collected during the season of 1890 ; taken on May 20,27, and 30, respectively. The nests were shallow depressions in the moss-coveredground, lined with bits of dry grass, and were placed at the borders of sprucethickets. The eggs were fresh when found.A set of six eggs, fresh when taken on May 27, 1906, is in my col-lection; it was taken by E. C. Bryant in Flathead County, Mont.The nest was at the end of an uprooted tree among some lodgepole
138 BULLETIN 162, UNITED STATES NATIONAL MUSEUMpines; the hollow in the ground was lined with pine needles, weeds,and other material that came handy.William L. Dawson (1896) describes a nest that he found inOkanogan County, Wash., as follows
:
On the 2Sth of April, 1S96, I found a nest of this bird at an altitude of abouta thousand feet above Lake Chelan. It was placed in the tall grass, whichclothed the side of an inconspicuous "draw" bottom, and although the ploughhad recently turned up the soil within five feet of her, the mother bird clungto her post. I took several " snap shots " of her at close range, and she allowedme to advance my hand to within a foot of her, when she stepped quietly offthe eggs and stood looking back at me over her shoulder. The nest was adepression in the gravel-filled soil, lined with grass and dry corn leaves, besidesa few stray feathers ; depth 3 inches, width 7 inches.Eggs.?The eggs of Franklin's grouse are similar to those of thespruce grouse; what few I have seen average more finely and moreevenly spotted with smaller spots, but practically all types can bematched in a series of either. They are beautiful eggs and greatly indemand by collectors. The measurements of 33 eggs average 42.7 by31.2 millimeters; the eggs showing the four extremes measure 45.1 by30.2, 44.5 by 33, and 39 by 30 millimeters.Young.?Mrs. Florence M. Bailey (1918) gives the following in-teresting account of her experience with a brood of young in GlacierNational Park
:
A brood of three half-grown buffy-breasted and tailless young were seen inthe Waterton Valley about the middle of August, wandering around enjoyiuc,'themselves in deep, soft-carpeted woods of spruce and fir, where they jumpedup to pick black honeysuckle berries from the low bushes, or answered theirmother's call to come and eat thimbleberries. One of them, which flew up on abranch, also passed the time eating fir needles. When surprised by our appear-ance the little fellows ran crouching down the trail showing a keen hidinginstinct, but their mother had little sense of danger. When the young wereapproached she merely turned her head over and called mildly in soft remon-strance. She was the genuine fool hen of Montana, we were told, whom theFlatheads and the mountain Indians never kill except when in great need offood, as the birds are so tame they can be snared at will, without ammunition :as the Indians say, with string from a moccasin.The same brood, we supposed, was met with a few days later on the sametrail. One of the young was in the trail and the mother was sitting on a logwhen we came up, but on seeing us she called the little ones into the bushes.When driven out for a better view she climbed a bank adorned with bear grass.dwarf brake, and linnaea carpet, and, stopping under a long drooping spray ofStreptopus?under whose light-green leaves hung beautiful bright red berries
?
she jumped up again and again to pick off the berries. Then, flying up on afallen tree trunk almost over my head, she sat there looking very plump andmatronly and entirely self-possessed, while I admired the white and tawnypattern of her plumage. She sat there calmly overlooking the brushy coverwhere the young were hidden and showed no disapproval when the three cameout and walked a log by the trail. She called to them in soft, soothing tonesand they answered back in sprightly fashion. It would have been so easy to
franklin's grouse 139win their confidence completely and to watch their engaging ways that it wastrying to have to leave them and pass on up the trail.Plumages.?The Franklin's grouse chick is beautifully colored.The central crown patch, which is bordered with black, and theupper parts in general are rich brown, from " Sanford's brown " to
" amber brown " ; the colors of the forehead, sides of the head, andunderparts vary from " mustard yellow " to " Naples yellow," deep-est and tinged with brownish on the forehead and flanks, and paleston the sides of the head and belly; there are black spots below theeyes, on the lores and auriculars, on the lower forehead, and on therump ; and there is a black ring around the neck.The juvenal plumage comes in first on the wings, when the chickis only a few days old, then on the scapulars, back, flanks, and breast,in that order; the tail appears next and the head and neck are thelast to be feathered. By the time the young bird is half grownit is fully clothed in juvenal plumage. The feathers of the upperparts are beautifully patterned in rich browns, black, and white;those of the crown are barred with black and white; those of theback, scapulars, wing coverts, and flanks are barred and patternedwith " tawny " and " ochraceous-tawny," and have narrow bars orlarge areas of black separated by bars of creamy white; many ofthese have central shaft stripes of creamy white, broadening onsome into a white tip ; the breast is " cinnamon-buff " or yellowishwhite, with large black spots; the chin and throat are white andthe belly grayish white; the tail is barred with sepia and grayishbuff and tipped with white. During the latter part of August themolt begins from the juvenal plumage, in which the sexes are alike,into the first winter plumage. This is a complete molt, except thatthe two outer primaries on each wing are retained for a year. Thesexes now differentiate and look much like adults, though the blackareas are less purely black and there is more white spotting.Adults may have a very limited prenuptial molt in spring, butthey have a complete postnuptial molt in summer and early in fall.J. H. Riley (1912), who collected a fine series of these grouse inBritish Columbia, says
:
The males were never found with the females and young, but always bythemselves and in full molt, July 18th to 21st, while at this time it had barelybegun in the females. All the males taken had molted the tail and the newfeathers were just appearing, while the only female taken that the molt hadprogressed so far was shot August 27th. This seems to show that while thefemale is brooding and bringing up the young, which she does unaided by herspouse, he goes on by himself and moults, while the process in the femaleis delas'ed until her young are able to shift for themselves.In the series of females collected tnere are two phases of plumage; one ofwhich I shall call the red phase and the other the gray phase. In the redphase the lower parts, down to the abdomen, are tawny ochraceous with the
140 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
sub-terminal black bars on the leathers oiten interrupted, giving to theseparts a beautiful "spangled" appearance; in the upper parts the tawny ochra-ceous barring is very prominent ; the middle tail feathers and upper tail-coverts are black crossed by .irregular narrow ochraceous bars and tippedwith white. In the gray phase the lower parts only as far as the upperbreast are ochraceous-buff, the feather of the breast being broadly tippedwith white, and the black bars are not interrupted, giving to the breast andabdomen the appearance of being black and white, entirely different from thered phase; the light bars on the neck and upper back are ochraceous-buffand the tawny ochraceous of the red phase in the rest of the plumage ofthe upper parts is replaced by wood or hair brown ; the central tail feathersare black, barred irregularly or stippled with wood brown and tipped withwhite ; the upper tail-coverts lack the white tips. Both phases were takenat the same locality with young; the gray phase is the rarer and in our seriesthere are intermediate stages.Food.?The same writer says of the summer food : " The food con-tents of the crops of the adults was either spruce leaves or the greenberries of a low-growing plant, while that of the young was theblossoms of the red heather Ph/yllodoce empetriformis, and a fewinsects."Aretas A. Saunders has sent me the following notes
:
Late in summer this bird seems to feed mainly on berries, particularly the
"huckleberries" (Yaccinium) . Up to the middle of October their crops con-tained a blue-colored berry of this genus that grows along the edges of sprue;'forests. There were sometimes the smaller red-berried ones, another Vacci-mum that forms undergrowth in the lodgepole-pine forest. All these are called
"huckleberry" in Montana, though allied to the blueberries. I do not thinkthe true huckleberry (Gaylussacia) ever grows there. The only other foodI have noted is spruce and balsam needles, and late in the season these aloneare found in the crops. Apparently these needles are the main winter food,though I have never shot and examined a bird later than late November.Behavior.?Mr. Saunders says in his notes:All through Montana this bird is known as "fool hen" because of its lackof fear of man. It will sit still, even when close to the ground and allow oneto approach very near. They are often killed with sticks or stones. Whena dog approaches they fly up into the trees, and sit there. By shooting thelowest one first, I have shot several in a flock, the others sitting and waitingtheir turns. In Jefferson County, Montana, we had a small brown spanielthat would put them up a tree, and then stand beneath and yelp till we came.In Lewis and Clark Counties, on the upper waters of the Sun River, I onceclimbed a small pine, and grasped a cock Franklin grouse by the foot, just tosee if I could do it. The bird moved to a higher limb when I let go, butdid not fly away. The male, even in fall, is fond of puffing out its blackbreast, and opening and shutting the red "comb" over its eve, apparentlyby a sort of lifting of its " eyebrows."John O. Snyder (1900) says that " one sat sedately on a limb whilea revolver was emptied at her. The shots having missed, roots andstones were thrown, which she avoided by stiff bows or occasionalsteps to the side."
BUFFED GROUSE 141Fall.?Mr. McCabe writes to me
:
The male, of course, takes zio part in rearing the young and is never seen nearthe broods in the summer. Yet in the late summer or early autumn the birdsgather into mixed groups, young males and females and adult females, to thenumber of six or seven, under the leadership of a single adult male. The latterwhen they are disturbed, assumes his display attitude, just as we have describedit at courting time, utters a rapid clucking sound, and approaches the intruder,a magnificent creature, while the rest either draw off quietly or squat insupposed concealment. There would be nothing peculiar about this were it notfor the fact that these groups only hold together for the two or three autumnmonths and then dissolve. The birds are very hard to find in winter as theyremain sluggishly in the hearts of the big balsams, eating the needles (wecounted 5,500 in one crop) or sometimes on the ground in dense masses of smallbalsams whose lower branches are weighed down into the snow, but when found(we collected three females last January) they are invariably single.Mr. Saunders tells me that though he has traveled on snowshoesmany times in midwinter, through forests where Franklin's grouseare known to occur, he has never seen one at that season. Evidentlythey remain well hidden. DISTRIBUTIONRange.?Northwestern United States and southwestern Canada.It is nonmigratory.The range of Franklin's grouse extends north to southeasternAlaska (Kasaan Bay) ; northern British Columbia (Tatletuey Lake,Thudade Lake, Ingenika River, and Hudsons Hope) ; and central Al-berta (Edmonton). East to central Alberta (Edmonton, PipestoneRiver and Banff) ; western Montana (St. Marys Lake, Belton, Paola,Mount McDonald, and Belt Mountains) ; and central Idaho (BakerCreek). South to central Idaho (Baker Creek, Sawtooth City, andResort); and northern Oregon (Mount Hood). "West to northernOregon (Mount Hood) ; Washington (Cowlitz Pass, Bumping Lake,Yakima Pass, Lake Chelan, and Pasaytens River) ; British Columbia(probably Chilliwack, Alpha Lake, Fort George, Fort St. James,Stewart Lake, Babine Lake, and Nine-mile Mountain) ; and south-eastern Alaska (Kasaan Bay).Egg dates.?British Columbia, Alberta, and Montana : 13 records,May 18 to July 29 ; 7 records, May 27 to June 9.BONASA UMBELLUS UMBELLUS (Linneaus)RUFFED GROUSEHABITSSpring.?During the first warm days of early spring the wandererin our New England woods is gladdened and thrilled by one of thesweetest sounds of that delightful season, the throbbing heart, as it
142 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUM
were, of awakening spring. On the soft, warm, still air there comesto his eager ears the sound of distant, muffled drumming, slow anddeliberate at first, but accelerating gradually until it ends in a pro-longed, rolling hum. The sun is shining with all its genial warmththrough the leafless woods, thawing out the woodland pools, wherethe hylas are already peeping, and warming the carpet of fallenleaves, from which the mourning cloak butterflies are rising fromtheir winter sleep. Other insects are awing, the early spring flowersare lifting up their heads, and all nature is awakening. The breastof the sturdy ruffed grouse swells with the springtime urge, as heseeks some moss-covered log, a fallen monarch of the forest, or per-haps a rock on which to mount and drum out his challenge to allrivals and his love call to his prospective mate. If we are fortunateenough to find his throne, on which he has left many a sign of previ-ous occupancy, we may see the monarch of all he surveys in all hisproud glory.Courtship.?Dr. Arthur A. Allen, who has made some carefulstudies of the display and drumming of the ruffed grouse and shownsome wonderful photographs of them, has contributed, at my request,a very full account of the whole performance, with some quotationsfrom other, earlier observers. I have had to condense it somewhat,but it is substantially as follows
:
" In a species as well known as this familiar game bird, which hasclaimed the attention of naturalists and sportsmen for nearly 200years, and whose courtship performances have been watched anddescribed by many observers, one would not expect many discrepan-cies in the accounts?at least among those recent observers who havehad the benefit of the arguments of the earlier naturalists. Such isnot the case, however, and it seems worth while, therefore, to sum-marize, here, the descriptions of the plumage displaj^ and the variedexplanations of the drumming performance, before concluding withthe writer's personal experience. Published records of the court-ship performances of the ruffed grouse date back to the year 1755when a communication from George Edwards on the pheasant ofPennsylvania was printed in the Philosophical Transactions of theRoyal Society of London (Edwards, 1755). In this classic com-munication Edwards quotes largely from a letter received from thatfamous naturalist, John Bartram. He says in part
:
When living, they erect their tails like turkey-cocks, and raise a ring offeathers round their necks, and walk very stately, making a noise a little likea turkey, when the hunter must fire. They thump in a very remarkable man-ner, by clapping their wings against their sides, as is supposed, standing on afallen tree. They begin their strokes at about two seconds of time distantfrom each other, and repeat them quicker and quicker, until they sound likethunder at a distance, which lasts about a minute, then ceases for 6 or 8
RUFFED GROUSE 143
minutes, and begins again. They may be beard near balf a mile, by which thehunters find them. They exercise their thumping in a morning and evening inthe spring and fall of the year.Edwards likewise quotes a Mr. Brooke, surgeon of Maryland, whosays:The beating of the pheasant, as we term it, is a noise chiefly made in thespring by the cock birds. It may be distinctly heard a mile in calm weather.They swell their breasts, like a pouting pigeon, and beat with their wings,which sounds not unlike a drum.Edwards then goes on to quote from La Hontan (1703), who inhis New Voyages to North America, vol. 1, p. 67, in speaking aboutthe grouse says
:
By flapping one Wing against the other, they mean to call their Mates, andthe humming noise that issues thereupon, may be heard half a quarter of aLeague off.
" There is the argument in a nut shell ; it is a problem of longstanding. Bartram says that the grouse beats its body with itswings; Brooke intimates that it merely fans the air; La Hontanreports that it hits one wing against the other. A further com-plication is advanced by Hodge (1905) when he tells us that thegrouse was called ' the carpenter bird ' by the Indians because theybelieved that it beat upon a log with its wings to produce the drum-ming sound.
" Let me quote from some of the apparently more authentic descrip-tions and explanations of the act. Audubon (1840) states that
?
* * * the drumming is performed in the following maimer. The malebird, standing erect on a prostrate decayed trunk, raises the feathers of itsbody, in the manner of a Turkey-cock, draws its head towards its tail, erectingthe feathers of the latter at the same time, and raising its ruff around the neck,suffers its wings to droop, and struts about on the log. A few moments elapse,when the bird draws the whole of its feathers close to its body, and stretchingitself out, beats its sides with its wings in the manner of the domestic Cock,but more loudly, and with such rapidity of motion, after a few of the firststrokes, as to cause a tremor in the air not unlike the rumbling of distantthunder.
" Between 1842 and 1874 one finds numerous references to thedrumming of the grouse, but which of the four beliefs the respectiveauthors hold as to the method of its production seems to dependupon whom they quote. William Brewster (1874), however, writ-ing in the American Sportsman, describes the drumming of a grouseas actually watched by him from a distance of 12 feet
:
Suddenly he paused, and sitting down on his rump and tarsi, crossways onthe log, with tail slightly expanded and hanging down loosely over the edgebehind, with body exactly perpendicular, neck stretched to its full length andfeathers drawn closely to the body, he stretched out his wings stiffly at nearlyright angles with the body. In this attitude he remained several seconds,
144 BULLETIN 162, UNITED STATES NATIONAL MUSEUMand I was instantly reminded most forcibly of the pictures one sees of thatsingular family of birds, the penguins. Now the wings were drawn slightlyback, a quick stroke given forward, at the air, and a pulsating throb entirelydifferent from any sound I have ever heard, struck my ear, producing at suchshort range an almost painful sensation on the drum ; the wings were immedi-ately recovered, and another stroke, a trifle quicker than the first, was succeededby another still quicker, until the wings vibrated too fast to be followed bythe eye, producing the well-known terminal "roll of muffled thunder," andnot till then the " semicircular haze." I say not till then, for the first two orthree strokes could be distinctly followed by the eye. This over, the birdimmediately rose to its feet, shook its feathers with an air of relief, andresumed its attitude of repose. * * *I think the drumming of the ruffed grouse is produced by the forward beats ofthe stiffened wings on the air, the planes of their motion being nearly horizontal,about four inches in length, with the initial ends represented by the pointsof a wire passed through the center of the erect body from side to side.
" Brewster's explanation of the drumming seems to have beenrather generally accepted until 1905. At that time Prof. C. F.Hodge, of Clark University, was experimenting in the rearing ofgrouse in captivity and enjoyed exceptional opportunities forwatching the drumming performance at a distance of but a few feetand made 40 different photographs of the drummer. He publishedquite an extended account of this drumming with his conclusions anda number of his photographs in the Country Calendar (Hodge,1905). He states:As to the matter of interpretation, I can not entirely agree with ProfessorBrewster. The appearance to the eye, however, supports his theory that thewing strikes nothing but air. But I am convinced that at just the criticalmoment, when the sound is produced, the wing moves with too lightning-likerapidity, even in the first slow sti-okes, for the eye to follow it. The wing,consequently, disappears from sight as it approaches the contour surfaces of thefeathers of the sides. We must defer here to the eye of the camera, and someof the photographs certainly show the blur of the rapidly vibrating wings comingup and touching the tips of the feathers along the sides. It is the impact of thestiffly held concave wing on the feather cushions of the sides that causes all thesound. In fact, the sound, so far as quality goes, can be best imitated bystriking with a wing properly stretched, or even a concave fan, on an extremelylight eider-down cushion.
" The next account which should be included is that of FrederickK. Vreeland reporting in Forest and Stream, for April, 1918, andreprinted in the Bulletin of the American Game Protective Associa-tion. After watching a grouse drum at a distance of 6 feet and aftertaking a remarkable series of photographs of the drumming bird, hecame to the conclusion that the thumping sound was produced by thewings striking behind the back, and he introduces a photograph thaihe says ' will prove to the most skeptical that they (the wings) didactually strike behind the drummer's back.'
RUFFED GROUSE 145
" E. J. Sawyer, after watching from a distance of a dozen feet thebeginning, progress, and ending of at least a hundred drumming.-during the spring of 1921, concludes in the Roosevelt Wild LifeBulletin for March, 1923
:
1. Tlie outward and upward motion [of the wings] is chiefly responsible forthe drumming sound, particularly during the first half of the performance, thoinward and forward motion of the wings being for the most part silent ornearly so.2. The striking of the air alone with the wings is practically the sole causeof the sound.
"With this framework to build upon, the writer here submitspictures (pis. 26, 27) of a grouse drumming, taken from the rear,which are made from enlarged motion pictures. Unfortunately it isnot practicable to reproduce all the 123 frames making up the drumbut a careful analysis of the pictures printed should satisfy the mostskeptical as to how the sound is produced. The film was made inthe spring of 1929 on Connecticut Hill, 17 miles southwest of Ithaca,N. Y., after four nights spent in a blind about a dozen feet from thedrumming log. The first night, April 26, the bird came to the log at1.40 a. m. and drummed every 5 minutes until 5 o'clock. He thenmoved to another log about 100 feet away and drummed until6 o'clock.
" The night of April 27 was cloudy and windy but warmer andthe grouse did not arrive until 4 a. m. He drummed every 3 min-utes until 5 a. m. and then every 5 minutes until 5.30, and thenwalked off the log not to return that morning.
" Stormy weather prohibited returning to the blind at the drum-ming log until May 4, when we resolved to try an experiment thatmight hold the grouse on the log later in the morning and at thesame time throw some light on the object of the drumming as wellas the method. We hoped it might also give us an indication asto the polygamous proclivities of the grouse. Accordingly we tookwith us a captive female grouse in a crate with 2-inch wire nettingover the top. This we set in front of the log, concealing the sideswith boughs so that the female would be most visible from thelog. The experiment was entirely a success.
" The night was cloudy, and it rained intermittently with occa-sional snowflakes. At 4.45 we were awakened by a fluttering in thecrate and peering through the peephole beheld the male bird infull display?tail up, ruffs extended, wings drooping but pressed tothe body and not touching the log. He moved along the log everso slowly until near the female. Then he lowered his head, ex-tending his neck and shaking out the ruffs still more, and madea few pecks at the log in front of him, though not always hittingit, with his bill. Next he started shaking his head and ruffs with
146 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
a rotating motion and commenced a series of short hisses, each onesounding like drawing the palm of one's hand rapidly backward,forward, and backward again over the sleeve or the trouser leg. Heshook his head more and more rapidly, the hisses corresponding, andfinally with a quick little run forward and a prolonged hiss, hestruck a pose and held it for several seconds. In this pose the tailwas swung over more to one side, the rump feathers on the offsidelifted, and the extended head and ruff turned toward the female.The male continued this strutting and posing, usually on the logbut sometimes in front of the crate, until 5.30, when he attemptedto mate with the female reaching his head far down through thewire and apparently seizing hold of her, for he pulled out at leastone feather. He went through all the motions of actual matingthough the wire separated them by several inches. He then pro-ceeded down the log to his accustomed place and drummed. Thisseems rather significant as indicating that the drum is probably achallenge to other males even more than an announcement of hispresence to the female."Audubon states that ' the female, which never drums, fliesdirectly to the place where the male is thus engaged,' but so far asI know, no one has ever seen a female grouse come to the male on ornear his drumming log, although a great many hours have been spentby different observers watching the drumming bird. I think we arejustified in concluding that while she may do so occasionally, she doesnot do so with any regularity and that the male must find her some-time during the day when he is not drumming. The drumming,therefore, resolves itself primarily into a challenge to other malesto keep out of the drummer's territory.
" Let us next analyze how the sound is produced. The ordinarydrum, such as the one filmed, requires almost exactly 8 seconds fromthe first wing beat until the last. With the motion-picture camerataking 16 pictures per second, the performance is registered on 123frames. The first one or two wing beats are almost silent and aregiven while the bird is in a nearly normal horizontal position, thewings striking downward and inward. The bird's tail is beinglowered against the log during this preliminary beat or beats. Thenabruptly he stands erect with his tail against the log, wings droopingat his sides and appears to throw his ' shoulders ' back. This mightgive the impression that the wings were struck behind the back,because the forward stroke of the wing follows so instantaneouslythat the eye scarcely perceives it, and it is given with such force andthe wings come back to the normal position so quickly that theentire action registers on only one frame of the motion-picture filmhaving an exposure of approximately one-fiftieth of a second. Be-
RUFFED GROUSE 147tween the ' thumps ' the wings of the bird register on the film withscarcely a blur representing the intervals between thumps. Thevarying tempo of the intervals between thumps has been noticedby all observers and as registered on the film is as follows, eachnumber being the number of pictures or the number of sixteenthsof a second between thumps
:
5_6_8-S-fr-5-5-4-^3-3-3-2r-2-l-2-l-l-l-l-l-l-000000000000000000000-l.
"If one now examines the series of pictures he will see that notonce is the back blurred, as it would be if the wings struck behindthe back, and that wherever the wings have moved with sufficientrapidity to cause a compression of the air and resulting sound, theyare registered forward and upward. This then is the effective sound-producing stroke of the wing
?
forward and upward?not outwardand upward as stated by Sawyer?more like his inward and forward,which he says is silent or nearly so.
" Moreover, if one watches the tail of the grouse during the drum-ming performance, he will see it become more and more flattenedagainst the log, for 'action and reaction are equal and opposite indirection ' and the forward-upward stroke of the wings tends to drivethe bird backward and downward on its tail. The reaction thatfollows cessation of drumming is even more clear to the observer,for always, upon the completion of the drum, the bird pitches slightlyforward and the tail lifts from the log as if it were a spring undercompression ; when the pressure is suddenly released by the cessationof drumming, the tail throws the bird forward and upward and isitself carried upward by the impetus given the bird."A single weak thump heard at the conclusion of the drum registerson the film in frame 122 after an interval of one frame where thewings are quiet. It corresponds to the beginning ' thump ' and isgiven as the bird pitches forward and is in a more horizontal position.This stroke is forward and downward rather than upward andperhaps helps the bird to regain its balance.
" During the four nights and mornings spent in the blind thegrouse drummed approximately one hundred times. I am frank toconfess that I did not watch every performance, for the strain ofkeeping one's eyes at a peephole is considerable in the small hoursof the morning. I did watch most of the performances, however,until I was absolutely convinced that the sound was produced ashere set forth."Nesting.?The ruffed grouse is a woodland bird, and its nest isalmost always in thick woods or under dense cover, though I oncefound a nest in a fairly open situation; it was placed at the baseof a small white birch in a clearing, with only a few small treesand bushes near it. Most of the nests I have seen in Massachusetts
148 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMhave been found by flushing the bird while hunting through heavy-woods in search of hawk's nests. The commonest location is at thebase of a tree ; this may be a large oak among heavy deciduous tim-ber, or a birch or other small tree in lighter, mixed woods; severalnests have been in dense white-pine groves at the base of a largeor a small pine. One nest was beside a rock in mixed woods, onewas partially hidden under a corner of a woodpile in open pinewoods, and others have been well concealed under fallen dead pineboughs or under old piles of brush. Several have been within 50or 100 yards of a red-shouldered hawk's nest. One that I was watch-ing was near a crow's nest; all the eggs but one were taken fromthis nest, probably by the crows. Edward H. Forbush (1927) tellsof a nest, found by J. A. Farley, that was directly under a sharp-shinned hawk's nest. The nests are merely deep hollows in theground, lined with whatever material is at hand, usually oak orother hardwood leaves; I have seen nests in pine groves that werelined with nothing but pine needles. There are usually a fewfeathers of the grouse mixed with the leaves. The female is aclose sitter and often does not leave the nest until the intruder isclose at hand; but, when approached on a second visit, she is moreapt to flush wild. In leaving she flies directly from the nest, ifclose pressed, with a great whir of wings, which makes the leavesfly and thus partially covers the eggs. Perhaps, on a more leisurelydeparture, she may cover the eggs more carefully.All the nests that I have seen have been on the ground in perfectlydry situations. But Major Bendire (1892) says: k ' Mr. Lynds Jones,of Grinnell, Iowa, found a nest of the Ruffed Grouse in a hollowstump, and Mr. C. M. Jones, of Eastford, Connecticut, found one ina swamp, on a little cradle knoll, surrounded by water. Mr. WilliamN. Colton, of Biddeford, Maine, records a nest found between thestems of three young birches, fully 8 inches from the ground."George M. Sutton (1928) reports two nests found in a sphagnum bognear Hartstown, Pa., one of which was " sheltered by leaves of skunkcabbage." E. A. Samuels (1883) records two instances where thisgrouse has nested in an abandoned crow's nest in a tree.Eggs.?From nine to a dozen eggs constitute the usual set, occas-ionally fewer and often as many as 14. Lester W. Smith sent mea photograph of a set of 23 eggs, found in Connecticut, and hetold me that every egg hatched. This was perhaps the product oftwo females. I believe that ordinarily an egg is laid each day untilthe set is completed ; but often, especially if bad weather occurs, aninterval of a day or two may intervene. E. P. Warner (1911)reported that, in a nest he had under observation, he found 3 eggson April 17, 4 on the 20th, 6 on the 24th, 10 on the 30th, and 14 onMay 7, all of which hatched.
RUFFED GROUSE 149The eggs are ovate in shape, with variations toward short-ovate orelongate-ovate. The shell is smooth with a very slight gloss. Theground colors vary from " chamois " to " cream-buff " or " cartridgebuff," or, more rarely, from " pinkish buff " to " cinnamon-buff."About half of the eggs, perhaps more, are entirely immaculate ; othersare more or less spotted with a few small spots or dots of " sayalbrown," " clay color," or duller buffs. The measurements of 73 eggsaverage 38.9 by 29.6 millimeters ; the eggs showing the four extremesmeasure 42.7 by 28.3, 40 by 32, and 33 by 25 millimeters.Young.?The incubation period has been variously recorded bydifferent observers at 21, 24, or 28 days; probably 21 days is thenormal period under faAorable circumstances, which may be length-ened by cold or wet weather or by interrupted incubation. The femalealone performs this duty and raises only one brood in a season. Theyoung leave the nest soon after they are hatched, or as soon as thedown is dry, leaving the empty eggshells in the nest neatly split intohalves.The female ruffed grouse is a model mother, assuming full care ofthe young, leading them away from the nest, teaching them to scratchand hunt for insects and seeds among the fallen leaves, and showingthem where the best berries are to be found and what green food isgood to eat. The young are at first very sensitive to dampness andmust not be allowed to wander in wet grass or herbage; she broodsthem under her wings, keeping them dry and warm during wetweather, and she broods them also at night. When they are oldershe leads them to bare places in the woodland roads, where they aretaught to dust themselves and free their plumage of vermin. Aspecies of wood tick causes the death of many very young chicks byburrowing into the skin of the neck or back?a situation with whichthe mother seems unable to cope. William Brewster (1925) ob-tained some evidence that the ticks kill the small chicks by boringthrough the tender skull into the brain.But her devotion is shown to the best advantage in her heroic de-fense of the young against their enemies. While walking quietlythrough the woods we may be startled by a shrill, whining cry andsee the infuriated bird, bristling with rage, rushing toward us, hertail spread and all her plumage extended; she seems twice her nat-ural size and imposing enough to cause any enemy to pause. Some-times she is less aggressive and merely flutters away, feigning lame-ness, or skulks away, crouching close to the ground and utteringthe same whining cry, which is the signal to the young to hide.When the young are older a clucking note is given as a signal to fly.The behavior of the mother is so startling that we have lost sight of74564?32 11
150 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthe young ; they have disappeared completely ; and search as we may,our chances of finding any are small. We had better not hunt forthem, as we may step on them. But, if we conceal ourselves andwait patiently, we shall see a pretty sight, which is well described byEdmund J. Sawyer (1923) as follows:There follows perhaps ten minutes of silence. Then conies a low, mewingnote, pe-e-e-e-u-u-r-r-r. The note can be imitated by trying to pronounce theword " pure " in a strained, tremulous way with the mouth nearly closed.Soon there is an entirely different note like the low clucking of a hen or tur-key ; this grows louder and more confident and I catch a glimpse now and thenof the watchful hen picking her cautious way back among the low plants.Tsee?tsee?tsee-e-e-e, answers a chick here and there about me, all unseen.Puck-puk-puk, from the mother; tsee-tsee-tsee-e-e-e, from the chicks, and one ofthe latter comes flying down from some leafy lower branch ; tsee-tsee-tsee?andanother appears from around a stump or log. There follows more callingback and forth, more chicks come out of hiding and already the puk-puk-pukshave begun to grow faint in the distance as the mother quickly leads the broodoff under cover of the ferns. I have on two or more occasions discovered oneof the chicks in his hiding place on the leafy ground. In each case he wasmerely squatting there, his coat of mottled down perfectly matching the brownsand grays of the forest floor.The wings start to grow soon after the young are hatched, andbefore they are half grown they are able to fly, or, at least, toflutter up into the lower branches of a tree. They are zealouslyguarded by their mother all through the period of growth, and in thefall their father joins the family group, which keeps together duringwinter in a loose flock. Edwyn Sandys (1904) had an interestingexperience with a pointer pup, which was attacked by a hen grouse,the guardian of a brood about as large as quail. He describes theincident as follows:A sudden tremendous uproar attracted my attention, and, to my astonishment, I saw an old hen grouse vigorously belaboring the bewildered pup withher wings and giving him a piece of her miud in a torrent of cacklings, suchas I had never dreamed a grouse capable of uttering. The poor pup, afterfirst trying to make a point, and then to grab her, finally bolted in dismay.She followed him for about a dozen yards, beating him about the rump withher wings, which kept up a thunderous whirring. She acted exactly like awrathful old fowl, and the pup like a condemned fool.Edward H. Forbush (1927) relates the following incident:Once I saw a fracas between the ordinary inoffensive rabbit and a grousehen, defending her chicks. She " bristled up " and struck at bunny, but heapparently tried to leap upon her. In the ensuing running fight he drove herabout a rod. Her chicks having hidden in the meantime, she then flew away.Very rarely, when the young are in danger, the male bird appears and takeshis turn at running toward and strutting near the intruder, and he has beenknown to care for a brood after the death of the mother bird.
RUFFED GROUSE 151The following extract is quoted from the journal of Dr. EdgarA. Mearns:We surprised an old hen pheasant (Bonasa umbellus). She gave a loudsquall, and whirred loudly as she beat her wings upon the ground in frontof us. The little chicks, only 2 or 3 days old and about 15 in number, atfirst piped out most lustily in their first surprise and bewilderment ; butsoon they recovered, and piled over each others' backs pell-mell in their effortsto escape and hide. Several of them rushed directly into the water of thebrook close beside which we discovered them. One swam clear across,another was carried a distance down the stream and then crawled under astone beside the bank. Others hid under stones and debris in the brook ; andone fellow was actually drowned in its effort to hide. We found it floatingdead upon the water when we returned to the spot sometime afterward. Theold bird dragged herself over the ground with a great clucking; but whenI ran rapidly after her, she took to wing and flew a little way off, and for afew minutes remained silent. We had captured three of the little chicks, andnow examined to see what sort of hiding places the balance of the flock hadchosen. One was packed like a sardine between two stones in the brook,with its head and streak of its back visible; another was wedged tightly be-tween a stone and some herbage growing beside it. Two cute little fellowswere found under shelving stones in the brook, running imminent risk ofdrowning. The drops of water were trickling off of one bird's head in rapidsuccession, and it was found fairly drenched. I verily believe that, had I notrescued it from this perilous situation it would surely have died rather thanbe discovered.Plumages.?In the ruffed grouse chick the entire crown and backare " tawny " or " russet," darkest on the back and rump, shadingoff to " pale ochraceous-buff " on the sides of the head, chest, andfianks; the underparts are pale yellow, shading off to yellowishwhite on the chin and belly ; there is a black auricular patch, but noother spotting on the head. The wings begin to grow soon afterthe chick is hatched and reach beyond the sprouting tail before thechick has grown much. The juvenal wings are fully developedand the young bird has reached the flight stage before it is halfgrown. The juvenal remiges are " light vinaceous-cinnamon," un-marked except for a very fine sprinkling of a slightly darker shade,somewhat lighter tips, and darker bases.The juvenal plumage is at its height when the bird is about three-quarters grown, for soon after that the remiges are molted and thefirst winter plumage begins to appear. In full juvenal plumagethe sexes are alike and closely resemble the adult female, but theyare much browner above and below, less distinctly barred on theunderparts, and more distinctly barred on the tails, which showboth red and gray phases. The mantle is variegated with brightand dull browns, heavily barred and finely sprinkled with black,and has many broad buff or whitish shaft streaks ; the chin is whitishinstead of buff ; and the chest is washed with " hazel."
152 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe first winter plumage is acquired by a complete postjuvenalmolt, except that the two outer juvenal primaries on each wing areretained throughout the first year. This molt begins in summerbefore the bird is fully grown and is generally completed beforeOctober. This plumage is practically adult and the sexes are nowdistinguishable. The new ruffs are a duller, more brownish blackin the young male and are at first tipped with " hazel."Adults have a very limited prenuptial molt, confined to the headand chin, and a complete postnuptial molt in August and September.The red and gray phases, most conspicuous in the tails, are presentin this and in all other races of this grouse, though one or the otherphase is supposed to predominate in each one of the races. In thenorthern races gray tails predominate; from Pennsylvania south-ward " silver tails," as they are called, are rare ; in the western andnorthwestern races red tails are rare; New England birds, as a whole,are about halfway between the extremes. " Red ruffs," birds withbrownish ruffs, tinged with a coppery red sheen, are occasionallyseen in many of the races.Food.?Forbush (1927) has published the most complete and con-densed list of the vegetable food of the ruffed grouse that I have seen,based largely on Dr. Sylvester D. Judd's report (1905a). Followingis his list in full
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Nuts or Seeds : Hazelnuts, beaclmuts, chestnuts, acorns. Seeds of tick trefoil,hornbeam, vetch, hemlock, pitch pine, maple, blackberry, lily, beggar's ticks,chickweed, sheep sorrel, sedges, violet, witch-hazel, beech drops, avens, persica-ria, frost weed, jewel weed. Buds, Blossoms or Foliage: Poplar, birch, willow,apple, pear, peach, alder, hazel, beech, ironwood, hornbeam, blackberry, blue-berry, spruce, arbor vitae, Mayflower, laurel, maple, spicebush, partridgeberry, sheep sorrel, aster, green ovary of bloodroot, clover, purslane, woodsorrel, yellow sorrel, heuchera, chickweed, catnip, cinquefoil, buttercup, speed-well, saxifrage, liveforever, meadow rue, smilax, horsetail rush, azalea, falsegoat's beard, dandelion, cudweed. Fruit : Rose hips, grapes, smooth sumac,dwarf sumac, staghorn sumac, scarlet sumac, poison ivy, partridge berry, thornapple, cockspur thorn, scarlet thorn, mountain ash, wintergreen, bayberry,blackberry, huckleberry, blueberry, cranberry, sarsaparilla berries, greenbrier,hairy Solomon's seal, smooth Solomon's seal, black raspberry, raspberry, domesticcherry, cultivated plum, wild black cherry, wild red cherry, elder, red elder,black haw, nannyberry, withe rod, maple-leaved arrow wood, high-bush cran-berry, mountain cranberry, snowberry, feverwort, black huckleberry, blackaider, flowering dogwood, bunchberry, cornel, silky cornel, pepperidge, mulberry,bittersweet, manzanita, barberry, Virginia creeper.Doctor Judd's analysis (1905a) showed 89.08 per cent of vegetablematter and 10.92 per cent of animal matter in the crops and stom-achs of 208 grouse, collected in every month of the year in Canadaand in 14 States. He says
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The animal food is almost all insects. The vegetable food consists of seeds,11.79 percent; fruit, 28.32 percent; leaves and buds, 48.11 percept, and miscel-
RUFFED GROUSE 153laneous vegetable matter, 0.86 percent. The insect food proper includes grass-hoppers, 0.78 percent ; caterpillars, 1.15 percent ; beetles, 4.57 percent, and mis-cellaneous insects, 3.86 percent. Some miscellaneous animal matter, made upof spiders and snails, is also eaten. The ruffed grouse eats a somewhat smallerproportion of insects than the bobwhite, but, like it, feeds on them to a largeextent in the breeding season.Judd lists among the animal food mainly insects, various grass-hoppers, crickets, various caterpillars, cutworms, army worms, cot-ton worms, apple worms, various beetles and their larvae, cloverweevil, potato beetle, various flies, bugs, ants, spiders, oak gallsmade by insects, snails, and slugs.The foregoing lists are probably not complete, for the grouse willeat, at different seasons, a great variety of food. In spring they arefond of the catkins, blossoms, and tender leaves of many of theplants named above, the fresh blades of new grass, and the wildstrawberries, when they come. Forbush (1927) adds:Perhaps the plant most sought after in the New England coastal region isthe cow-wheat, a low growing plant with small white blossoms which thrivesalmost everywhere that this bird is found. Ruffed Grouse in confinement areso fond of it that they eagerly eat quantities of it, consuming the entireplant, root and branch. Edible mushrooms are taken eagerly. Fern leaveswhich remain green in swamps under the snow of winter are eaten then aswell as at other seasons.During summer, when the birds find their food on or near theground, insects begin to form an important part of their food, about30 per cent of the adult food, according to Doctor Judd (1905a) . Hesays that the newly hatched chicks are nearly, or wholly, insectivo-rous, feeding on cutworms, grasshoppers, beetles, ants, wasps, spiders,and caterpillars. The old birds, too, like to wander out into thefields and meadows near the woods in search of grasshoppers andcrickets and to scratch among the woodland leaves for other insectsand grubs. All kinds of berries and fruits claim their attentionduring summer and fall; I have found them frequenting regularlythe edges of cranberry bogs near the woods, as well as wild-appletrees in secluded spots.In winter, when their ground food is buried under the snow, theyhave to resort to trees and bushes for what fruits and berries areleft, for leaves that remain green, and for dried catkins and buds.They are said to feed largely on leaves of sheep-laurel and mountain-laurel; and people have been poisoned by eating birds that had fedon such diet. I wonder if the poisoning was not due to berries ofpoison sumac and poison ivy, which are easily obtained in winter.The ruffed grouse has a bad habit of budding cultivated appletrees, quite extensively when other food crops fail. Forbush (1927)gives us the following surprising figures
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154 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMMr. Charles Hayward reports that he found in the crop of a grouse 140apple buds, 134 pieces of laurel leaves, 28 wintergreen leaves, G9 birch buds,205 blueberry buds, 201 cherry buds and 109 blueberry stems. Another birdhad G10 apple buds in its crop and a third had more than 300. Weed and Dear-born found in the crop of a female ruffed grouse 347 apple buds, S8 maplebuds and 12 leaves of sheep laurel.This damage may be serious during certain winters, especiallyin orchards close to woods where the grouse are numerous; he men-tions a case where a tree has been denuded of buds and killed. But,if not overdone, budding may be more beneficial than injurious,amounting to merely healthful pruning, for he says
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For twenty years one or two birds customarily " budded " on an apple treenear my farmhouse window. This tree seemed to be their favorite, but not-withstanding the " budding " or because of it, the tree bore a good crop oflarge apples nearly every year, while other trees not " budded " by the grouseoften bore none. Apparently the thinning of the buds by the birds was abenefit to the crop.Doctor Judd (1905a) quotes from a letter from Miss M. E. Paine,as follows:The ruffed grouse eats the buds of apple trees, but it is a help rather thana damage. Last year a wild apple tree on top of a hill, between pasture andmowing, was almost entirely budded. I thought entirely at first, but theterminal buds were almost always left uninjured, also many minute budson each limb. The result was the terminal buds were pushed out and grewrapidly and strongly. The tree blossomed abundantly and the fruit hung inclusters toward the ends of the branches. The tree is of medium size andthe branches droop to the ground. In the fall the golden apples occupiedfully as much room as the green leaves, and as one looked at the tree a fewrods away?a perfect picture, barrels of apples on it, all nearly perfect andfair, just the result of a vigorous trimming.William Brewster (1925) describes their method of budding asfollows
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At sis o'clock this morning my assistant, R. A. Gilbert, called me to see someRuffed Grouse budding in a large wild apple tree that stands within sixty yardsof our old farm-house, from one of the eastern windows of which I was ableto watch them very satisfactorily through my field-glass. Five or six werenoticed at once and before many minutes had elapsed I counted no less thannine scattered all over the tree, a few being low down on stout limbs close to itsmain trunk and hence inconspicuous, but the greater number near the ends ofits longer upper branches, where they could be plainly seen, while one or twowere perched on the very topmost twigs, boldly outlined against the grey skyand looking as big as Hen-hawks. They were busily engaged in budding, an op-eration which I have never before witnessed to such good advantage. It wasnot less surprising than interesting to see birds ordinarily so shy and retiring,and so very stately and dignified of bearing, assembled thus numerously in anisolated, leafless tree not far from a house, hopping and fluttering almost cease-lessly amid its branches, thereby displaying unwonted activity and sprightli-ness, as well as apparent fearlessness. At times, however, they would all standerect and motionless for a few moments, evidently looking and listening intently.
BUFFED GROUSE 155Those feeding near the ends of long and slender branches had some difficultyin keeping their foothold and were constantly obliged to jerk up their tails, andflutter their wings in order to preserve their balance, especially when as oftenhappened, they stretched forward or even for a moment bent almost straightdownward after the manner of Redpolls or Pine Siskins similarly engaged.They picked off and swallowed the buds in rapid succession, with much thesame quick, bobbing motion of the head as that of a domestic fowl feasting oncorn. The supply of such buds as they chose, within reach of the most favour-ing percb, seldom lasted more than a minute or two. When it became exhaustedthe partridge either moved still farther out among the terminal twigs, or flewto another part of the tree. Birds at work not far from the trunk behavedsomewhat differently, and with decidedly more dignity and deliberation, doubt-less because the buds they were obtaining grew on short twigs within easyreach of thick and perfectly rigid branches on which they could stand or workas easily as on level ground.Behavior.?There is a striking difference in behavior between theunsophisticated grouse of the primitive wilderness and that of thewise and wary birds of thickly settled regions. Birds that havenever heard the roar of a gun and have not learned to know theirdangerous human enemies are often absurdly, almost stupidly, tame
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whereas the birds that have been persistently hunted have developedsuch a high degree of wariness and strategy as to make it difficult tooutwit them. Formerly in much of New England and eastern Canadathe ruffed grouse well deserved the name of " fool hen," and was oneof the easiest of birds to shoot. It would either walk quietly awayor fly up into the branches of a tree and stare stupidly at theintruder. It was an easy matter for a good shot to pick off itshead with a rifle, and it was considered unsportsmanlike to shootgrouse in any other way. It has often been said that, when a numberof grouse are perched in one tree, if the lowest one is shot first andthen the next lowest one, the others will remain until the last one iskilled. I doubt, however, if this has often happened ; it hardly seemscredible; and even in Audubon's time it was doubted. Even now,in the wilder portions of Canada, in the southern Alleghenies, and insome of the Western States, the grouse are absurdly tame and takebut little notice of human beings.The normal behavior of sophisticated ruffed grouse will be referredto later, but, while we are on the subject of tameness, we must con-sider the numerous cases recorded in print of abnormal tameness ofindividual grouse in regions where their fellows are the wildest.Space will not permit reference to all of more than a dozen suchcases of peculiar behavior that I have heard or read about; one ortwo samples must suffice. In all these cases an individual grouse,sometimes a male and sometimes a female, showed a strong attach-ment for, or a decided interest in, one or more human beings, withthe element of fear entirely eliminated. Carleton D. Howe (1904)published a full account of the behavior of a hen grouse that devel-
156 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMoped a strong friendship for a farmer and even allowed herself tobe handled by other people. The friendship lasted through at leasttwo seasons. " When Mr. Rand called 'Chickee,' 'Chickee,' the birdwould come out of the woods and sit upon his knee. From his kneeshe would fly to his shoulder, and then to the ground. The birdwould repeat this performance a half dozen times, clucking con-tentedly the while."Howard H. Cleaves (1920) tells an interesting story of a bellig-erent cock grouse called " Billy " that " went forth to battle " witha motor tractor; he was evidently attracted by the noise it made,was not in the least afraid of it, and would even ride on it inmotion. When first seen he was25 yards up the road, his ruff extended and his head lowered and jerkin?nervously, after the manner of a rooster about to make battle with his foe.Billy took the middle of the lane and, following a peculiar, sinuous course,came steadily on to meet us with reckless abandon. The contrast was absurd.On the one side was a wild bird not larger than a bantam, and on the otherwere five adult humans led by a mobile mass of several thousand pounds ofsteel from which emanated a loud noise : a feathered David and a mechanicalGoliath.At the instant when it seemed that further advance by either side wouldmean annihilation for the eccentric Grouse, the pilot brought his tractorto a stop and descended to the ground, whereupon began one of the mostremarkable of exhibitions. Billy darted toward Mr. Armstrong's feet andpecked at his trousers, and when Mr. Armstrong walked away the bird ranafter him with the greatest agility, striking with wing or beak on comingwithin range. If a hand were extended toward him, Billy would peck it alsoand, most extraordinary of all, he would permit himself to be picked up andfreely handled, perching on finger, wrist, or shoulder. When on the latter hewas invariably prompted to investigate one's eyes and nose with his sharpbeak!A number of other published accounts illustrate similar traits.These abnormal birds are usually resident in some restricted areawhere they can generally be called by the human voice or whistle orcome to the sound of a moving vehicle, a woodchopper's ax, or a stickrustling among the leaves. They follow their human friends aboutlike pet dogs, can be coaxed to eat out of human hands, will oftenpeck at them in a possibly playful manner, and will eventually allowthemselves to be handled. Rae T. Hadzor (1923) tells of a hengrouse that flew into the yard one fall, possibly to escape from someenemy, and lived there about a year, mingling with the chickens butroosting by herself in an open shed. She became tame enough to eatout of the hand and even laid a set of eggs in the orchard. Of course,they did not hatch.When a ruffed grouse is suddenly flushed it springs into the airwith a loud whirring of wings, which is quite startling and discon-certing to a novice, and goes hurtling off through the trees or bushes
RUFFED GROUSE 157
at terrific speed, gaining momentum very quickly. Evidently itdepends largely on its feet for the initial spring from solid ground,for it has difficulty in rising from soft snow, where it leaves theimprint of its whole body and wings in its struggle to rise. But itdoes not always make a noisy " getaway " ; I have often seen one flitsoftly and silently up and over a stone wall, fence, or bush when itwas not frightened or thought it was not observed. Again, whenflying from a tree, it usually launches downward and flies awayalmost silently. The roar of a rising grouse, often too far away to beseen, is a common sound in regions where the birds are wild. Itsflight is strong, exceedingly swift, and usually quite direct, but not,as a rule, prolonged for more than 150 or 200 yards, unless the bird iscrossing a river or an open space between tracts of woods. A commonhabit is to fly low and straightaway along a woodland road or path,but its usual method is to rise above entangling undergrowth and thenrly away through the trees, soon setting its wings and scaling downinto thick cover. I have always thought it particularly skillful indodging the branches of trees in its swift flight through thick woods,but evidently it is not always successful in this, for Forbush (1927)says
:
It does not, like the Wood Duck, so control its movements as to avoid thetwigs and branches of trees, but dashes through them. I have seen one in sucha case strike bodily against a limb and fall to the ground. This bird had beentired at in a neighboring wood, and had crossed the open with tremendousspeed to another wood where it struck the limb. Aside from the shock the birdwas unhurt. Mr. Albert A. Cross of Huntington sent me a Ruffed Grouse thatin full flight had collided with the forked and broken end of a dead limb,driving one of the prongs three inches into its breast and the other into itsvitals, and tearing the head and neck from the body.Forbush also speaks of a habit I have never noted:A hard-pressed bird has been known to go into shoal water, apparently forconcealment. Mr. L. Barber tells us that a grouse that was startled by hisdog alighted in the water. She was entirely under water except her head whichwas covered by a projecting bush. Mr. W. L. Bishop writes that he killed aGoshawk near a brook, and afterward discovered by traces on the snow that thehawk had been pursuing a Ruffed Grouse. He found the frightened bird in thebrook entirely submerged with the exception of its head. Though the RuffedGrouse seems to drink mostly the dew and raindrops from the leaves, it is notafraid of water, and if winged over water can swim fairly well.Grouse are much given to dusting themselves in soft, dusty placesin woods roads, in country highways or on old rotted logs or stumps.They have favorite dusting places in which a few telltale feathersmay be found.Voice.?The vocal accomplishments of the ruffed grouse are quitesimple. The commonest note heard when the grouse is slightlyalarmed is a sharp quit-quit, usually given while walking on the
158 BULLETIN 162, UNITED STATES NATIONAL, MUSEUMground and indicating nervousness. The squealing or whining noteuttered while defending its young is probably also a signal to themto hide, as a clucking note is a signal to older young to fly. Thenthere is the call of the female to her young, crut-crut, car-r-r, andvarious soft cooing notes and chatterings.Enemies.?Besides its archenemy, man, who has shot and snaredit almost to extinction in many places, the ruffed grouse has manynatural enemies and is subject to many diseases. It always managedto survive, however, until man came on the scene; its large broodshave helped it to come back to normal numbers after periods ofscarcity. Foxes destroy large numbers of grouse, as well as theireggs and young; feathers scattered about their burrows and tracksin the snow tell the story. Forbush (1927) says:Mr. C. E. Ingalls, writing of an experience at Templeton, stated that he sawa fox approaching the nest of a Ruffed Grouse near the edge of the woods. "Abig ball of feathers," writes Mr. Ingalls, " flew out at that fox and drove himsome distance into the grassland." The fox, nevertheless, returned to theattack only to die in his tracks by a well-directed bullet from the rifle of thewatcher, not, however, until the brute had filled both mouth and throat withegg contents from the nest of the devoted mother.Wandering dogs, stray cats (of which we have too many in ourwoods), lynxes, and perhaps raccoons and weasels kill many oldand young grouse, the former being probably mostly caught on theirnests. Skunks, opossums, raccoons, and squirrels undoubtedly robthe nests. The goshawk, also called " partridge hawk," levies heavytoll during periods of its abundance ; it is often named as one of thechief causes of the periodic scarcity of grouse. The Cooper's, red-tailed, and red-shouldered hawks probably kill a few. Great hornedowls pounce on them in their night roosts and are very destructive.The evidence against the screech owl and the long-eared owl, bothof which have been seen eating grouse, does not seem conclusive.Crows are canny nest hunters and doubtless break up many nests; Ifeel confident that a nest that I was watching was robbed by a familyof crows that had a nest near by. Dr. Charles W. Townsend (1912)reports a crow seen flying off with a freshly killed grouse in itsclaws; examination of the body of the grouse, which the crow wasseen to drop, led to the conclusion that the crow had killed it.Fall.?Audubon (1840) refers to short migratory flights of grousein October across the Ohio and Susquehanna Rivers. These areprobably nothing more than autumn wanderings in search of food.But there is much evidence of an incipient or suppressed migratoryinstinct in the erratic short flights of ruffed grouse during the so-called " crazy season " in fall. At such times they certainly do be-have queerly. I have repeatedly known them to appear in my yardin the center of the city, or to kill themselves by flying against build-
RUFFED GROUSE 159ings or through windows. Once one flew through a window intoour machine shop, where the machinery was running and scores ofmen were working. One of my neighbors once found one inside herhouse with no visible means of entrance except through a chimney.Forbush (1927) says tha<>-Some have been known to go through the glass of moving motor cars ortrolley cars and even into locomotive headlights. So careless are they of ob-structions that a high wire fence around a covert is likely to kill all the RuffedGrouse within its confines. Dr. A. O. Gross found that three birds whichhad been killed by flying against obstructions were infected by internal para-sites, and he suggests the possibility that the irritation caused by such para-sites may be the initial cause of the " crazy " behavior commonly observed.Game.?Although the bobwhite may be more universally popular,for the reasons stated elsewhere, I think most sportsmen will agreethat the ruffed grouse, known in the South as " pheasant," in theNorth as " partridge " or " patridge," and in Canada as " birch par-tridge," is the unrivaled king of North American upland game birds.Shooting into a flock of whirring quail gives a thrill, but it is com-paratively easy, and shooting the straight-flying prairie chicken inthe open is child's play compared with stopping the swift rush ofthe wily grouse through the treetops. The thundering roar of therising bird, the flash of nitro at a vanishing glimpse of brownfeathers, the dull thud of a plump partridge falling to earth, andthe whir of wings among dry leaves as it beats its final tattoo, com-bine to produce the thrill of thrills for the successful sportsman.And with the freshly killed monarch of the woods in hand comeadmiring thoughts, so well expressed by Charles B. Morss (1923) :In no other game bird do the tones of gray, black, cinnamon, and whiteshade and blend with such quiet harmony. Child of the wilderness that he is,in the full dark pupil of that eye surrounded by an iris of October's own brown,seem always to dwell the brooding shadows of the great forest he loves sowell. And in the moulding of him Nature seems to have embodied all of thebeauty, all of the charm, all of the inexplicable strangeness and romance ofthe autumnal woods and produced her feathered masterpiece?the perfect gamebird. * * * And wherever you chance to find him?in the still shadow ofravine and glen where the climbing bittersweet twines its orange offeringabout old stumps and windfalls?on rocky hillsides clad with second growthwhere the wild barberry fruits in crimson racemes and berries of the winter-green flash among the leaves?or in the grass-grown tangles of birch meadowand maple swamp where glows the steady flame of the black alder?always ishe the woodland's pride, alert, instinct with life, and filled with a spirit anddash that furnishes, when in such mixed cover as we were hunting this day, thevery climax of shooting with the shotgun.A good partridge dog adds much to the pleasure and success of thehunt, but good partridge dogs are scarce, and a poor one is worsethan none. I once had one that showed real " bird sense," knewwhere to hunt for the birds, would not run too far away, was careful
160 BULLETIN 162, UNITED STATES NATIONAL MUSEUMabout flushing birds, and was a fine retriever. But I love to huntalone, with nothing to distract my attention from the beauties of theautumn woods, to watch and study the interesting habits of the otherwild creatures, to learn the haunts of the wily grouse and match mywits against his. Then, if I can, unaided, outwit this wizard of thewoodland glades, learn to beat him at his many clever tricks, I feelthat I have earned my prize. Well he knows the trick of puttingthe trunk of a big tree or a thick tangle of leaves and branches be-tween the hunter and himself in his headlong flight; or running offto one side, he will rise behind the gunner and get away safely ; per-haps he will alight in the thick top of a pine tree and slip away onthe farther side of it on silent wings, giving the hunter an unexpectedand difficult shot. Usually, if we miss, we can watch his distant flightto mark him clown and flush him again ; but he may run a long dis-tance or fly out across an opening to another bit of cover and escape.Occasionally we get a pleasant surprise by killing a bird we couldnot see, shooting in the direction it has taken behind thick brush. Orwe may think we have made a clean miss, as we see the bird keep onand on rising up and up into the sky until it appears as a small speck
;
but if we watch it, we may see it drop like a stone, shot in the headthen we need a good dog to find it. There is no bird that so tests theskill, patience, and endurance of a good wing-shot as the ruffedgrouse and no shooting that calls for so much experience and intelli-gent study. There are very few who can make a respectable ratio ofbirds killed to shots fired ; if one takes his shots as they come, thereare very few that can put a bird in the pocket for every three emptyshells. One must know where to look for his birds and study theirfood habits in the localities he hunts. One of the surest places tofind them in my section is where an old apple orchard has been aban-doned and overgrown with grapevines, briers, and birches, or whereold apple trees grow along the edges of the woods, or where tangledthickets of berry-bearing shrubs, junipers, cedars, pines, and otherforest growths are encroaching on deserted pasture lands. When thebeechnut crop is good they may be found in sheltered spots on thesunny side of the woods or brushy hillsides. In the mountains far-ther south they frequent the rhododendron clumps, the ravines linedwith laurel thickets, or the dense undergrowth along the streams,where shooting is difficult. In closing this chapter I am tempted toquote the appreciative words of one who has substituted the camerafor the gun; Edmund J. Sawyer (1923) says:And now there is little enough satisfaction in the reflection that that gun shotmany a grouse, albeit all of them on the wing and not one over a dog. I have,after all, never taken a Grouse except through the immense advantage of myinfernal powder and lead. I never outwitted him fairly ; I have never held hisJimp form in my hand without feeling the rebuke of his matchless wings. I
RUFFED GROUSE 161found no just ground to glory over the dead body of that perfect product of thewild outdoors, that past master of woodcraft with his wings, which so immeas-urably outmatched the best my limbs could do ; those wings with their damning,rebuking evidence?a drop of lead-tinctured blood. The triumph was all his.Winter.?If not too much disturbed the young birds remain in thefamily group with the female during fall and winter; and the malejoins them late in fall. They fatten on the abundant crop ofberries, fruits, and nuts in preparation for winter, their legs are nowwarmly feathered, their thick new plumage protects them againstthe increasing cold, and the comblike scales on their feet grow outto help them walk on the snow. They now seek their winter quartersin thick woods, where they can find shelter from cold winds and agood food supply within easy reach. They lead comparatively in-active lives, spending much of the daytime roosting in evergreen trees,in vine-clad thickets, on sunny borders, or even under the snow.They feed early in the morning and at dusk, mainly on the buds ofpoplars, birches, and apple trees. Their tracks in the snow are easilyrecognized, as the toes are widely spread and the tracks are in adirect line, one directly in front of another. In severe weather, whenthe snow is deep, they often dive into the soft drifts and find asnug, well-protected bed a foot or two below the surface. Unlessthe snow becomes very heavily frosted they can easily burst a wayout in the morning, or unless an enemy finds their hiding place. Agood shelter is sometimes found under a low branch or pile of brushcovered with snow, or under a log banked with snow. Such placesserve as either day or night roosts. When there is little or no snowthe birds gather for the night in the thickest groves of pines orother conifers, roosting on the branches near the trunk, often anumber together in the same tree, where the cover is dense enoughto protect them.Mortality.?The periodic scarcity of ruffed grouse, with subse-quent recovery to more normal numbers again, has long been a fruit-ful subject for discussion and study. Their natural enemies haveprobably served only to keep their increase within check. Theirdecrease is due to many other causes, mostly chargeable to man.Trapping, snaring, and smoking out roosting places have all beenstopped, and shooting has been reduced and periodically stopped.But clearing and cultivating land still goes on, and sportsmen arestill increasing. Severe winters and unfavorable breeding seasonshave their temporary effects, as do the periodic scarcity of certaininsects, on which the young depend for food, and the occasionalinroads of heavy flights of goshawks or great horned owls.But the principal causes of decrease and of excessive mortalityare the various diseases to which grouse are subject and the numer-ous parasites that attack them. At my request, Dr. Alfred O. Gross,
162 BULLETIN" 162, UNITED STATES NATIONAL MUSEUMwho has been making an extensive study of this subject for severalyears, has contributed a condensed, but quite complete, report onthe diseases and parasites of the ruffed grouse, which I considerimportant enough to quote almost verbatim. At least 6 infectiousdiseases and about 25 parasites have been discovered in the examina-tion of more than 2,000 birds received from a wide range extendingfrom Quebec to Virginia and westward to Minnesota.Among the infectious diseases he mentions enlargement of thespleen, enteritis (unknown etiology), hepatitis (diseased liver, un-known etiology), enterohepatitis (blackhead), aspergillosis (a fungusdisease chiefly of the respiratory system), and bird pox (an integu-mentary disease, producing tumorlike growths, generally in theregion of the head and mouth). Following is Doctor Gross's reporton external parasites
:
" Bird lice are wingless ectoparasitic insects with mouth partsadapted to biting, not sucking. They feed on parts of the feathersand epidermal scales and when present in large numbers causesevere irritation and considerable annoyance to the host. Liceof the genus Gonicotes have been taken most frequently from NewEngland specimens. Grouse collected during the summer monthsin southeastern New York were heavily infested with the louseEsthiopterum perplexum.
" Ticks of the species Haemaphysails leporis-pallustris have beentaken from birds collected throughout the entire range of the ruffedgrouse. The heaviest infestations of the parasite were found amonggrouse living in the heavily wooded sections of northern Maine andalong the Canadian Labrador coast.
" The ticks generally attach themselves in the region of the head,usually on the chin and skin around the eyes or on the nape, allplaces where it is difficult for the host to dislodge them. The ticksvary in size, depending on the relative quantity of sucked bloodthey contain. Even a few ticks are a discomfort to the birds, and ineases of heavy infestation they become a serious menace, decreasingthe vitality and resistance of the grouse and in certain rare casesresulting in the death of the bird. Observers have reported thatentire broods of young grouse have been destroyed by ticks.
" It has been demonstrated that the tick may serve as a carrier ofsuch diseases as tularemia, an infectious disease of rabbits, and sincethe tick HaemapTiysails leporis-pallustris is a common ectoparasite ofboth the rabbit and the grouse it may thus eventually prove to be ofconsiderable importance in the life of the grouse. The ticks Haema-physalis cinnabarlna and H. punctata punctata have also been takenfrom ruffed grouse.
" Live birds received from Canada and others collected in northernMaine were found to be heavily infested with the northern fowl mite
RUFFED GROUSE 163Lyponyssus sylvarium. Several individuals of a flock of ruffedgrouse kept in captivity at Brunswick, Me., were killed by an infes-tation of myriads of these blood-sucking mites.
" The hippoboscid fly Omithoponus amerlcanus is light brownand about the size of a common house fly but with longer wings andwith a radically different life history and behavior. This fly is veryactive and difficult to secure since it is quick to disappear among thefeathers or it may leave the bird entirely. The flies attack the birdsin the region of the head, where they suck the blood of the host andare thus capable of bearing blood parasites and diseases."The internal parasites are more numerous, of greater importance,and apparently more often prove fatal. Of these Doctor Grosswrites
:
" The internal parasites are chiefly in the alimentary tract, wherethey are introduced along with the food, chiefly in the form of eggsand larvae. The life histories of some of the parasites involve asecondary host.
" The crop worm Thominx annulata is an extremely slender wormfound beneath the epithelial lining of the crop and gullet. It isfrom 2 to 3 inches long, but less than the diameter of a hair. Thewalls of the crop and gullet are very much thickened and throwninto folds and ridges by the presence of this parasite.
" The stomach worm Dispharynx spiralis is one of the most im-portant of the internal parasites. In southern New England andNew York State a large percentage of the birds found dead andexamined, died from the effects of this nematode. Dispharynx spir-alis is between one-fourth and one-half of an inch in length, pointedat both ends, and always rolled up in a characteristic spiral form.It becomes established in the glandular walls of the proventriculusand in advance cases of infection this region is so excessively swollenas to equal the size of the gizzard. The worms become sexuallymature in the grouse, and great numbers of eggs are passed throughthe alimentary tract and eliminated with the droppings. The eggswill not develop if taken directly into the body of another grouse,but the life cycle involves an intermediate host, which Dr. EloiseB. Cram, of the United States Bureau of Animal Industry, workingwith the New England ruffed grouse investigation, has found tobe the pillbug.
" The gizzard worm Gheilospirura spinosa is distributed over awide range from Maine to Pennsylvania and west to Wisconsin. InWisconsin it is second only to Ascaridia lineata in the percentages ofcases of infection. Gizzard worms are slender, the female obtain-ing a length of iy2 inches. They are usually found between thechitinous lining and muscular walls of the gizzard. Like Dis-
164 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMpharynx the life cycle of this nematode worm involves a secondaryhost, which Doctor Cram has discovered to be a grasshopper.
" The intestinal worm Ascaridia lineata is the commonest of all theinternal parasites and in certain sections of the range of the ruffedgrouse as many as 75 per cent of the birds examined were found tobe infected. This worm is comparatively large, ranging from 2 to 4inches in length. It is yellowish white and pointed at both ends.Because of its size and abundance it is reported by sportsmen morefrequently than all other parasites combined. Ascarids normallyreside in the intestine and while there do comparatively little harmaside from absorbing a certain quantity of food. A number ofcases have been found, however, where the worms have been outsidethe intestine among the vital organs and in the body cavity wherethere was no evidence of shot wounds or scars to indicate that theworms had made their way through artificial openings."The hatching of the eggs occurs normally after the eggs areswallowed, and the life cycle does not involve a secondary host. Inthe larval stages this parasite is capable of doing serious injury tothe mucous lining of the alimentary tract, especially in young birds.Some of the larvae may even penetrate the mucous lining and thusbe transported by the blood stream to other parts of the body.Furthermore, wounds made by the larvae may open the way toinfections of a serious nature.
" The cecal worm Heterakis gallinae is a small nematode parasiteusually found in the ceca. Rarely it occurs in the small intestine,colon, and rectum. It is white and very rigid, and the head is bentdorsally, a character that serves to distinguish it from young Asca-ridia. The eggs of Heterakis pass in the feces and under favorableconditions of temperature and moisture develop in about 10 days tothe point where the eggs contain infective embryos. When the eggsare swallowed by the grouse the embryos are released and then de-velop into adult worms in the region of the ceca. This parasite isknown to have an important relation to the dreaded disease entero-hepatitis commonly known among poultrymen and game breeders asblackhead. The eggs of Heterakis may carry for great lengths oftime the blackhead germs and thus facilitate the spread of this di-sease from bird to bird. A species of Contracaecunv has been takenfrom the ruffed grouse.
" Tapeworms, designated also as cestodes, constitute importantparasites in the intestines of certain groups of birds. Three speciesof minor importance have thus far been found in the ruffed grouse."A large number of the ruffed grouse examined have been foundto be infected with flagellate protozoan parasites. Thus far six-species have been identified, of which Histomona<$ meleagridi*
RUFFED GROUSE 165(blackhead) is of the greatest pathological importance. Blackheadis a common and serious disease of poultry and of captive game birdsand has also been found in a number of cases of ruffed grouse collectedin their natural covers.
" The group Sporozoa is represented by two species of Eimeria,which are responsible for the disease coccidiosis, which is not onlyimportant in grouse raised in captivity, but, as in the case of black-head, is also found in grouse killed in a natural habitat."DISTRIBUTIONRange.?United States, Canada, and Alaska; north to the limitof trees. The species is nonmigratory.The range of the ruffed grouse extends north, to Alaska (Nulato,Tanana, Kampart, Fort Yukon, and the Porcupine River) ; Mac-kenzie (Roche Trempe 1'Eau, Fort Simpson, Fort Providence, FortRae, and probably the Slave River Delta) ; Manitoba (Hairy Lake,Oxford House, Knee Lake, and York Factory) ; northern Ontario(Martin Falls, probably Fort Albany, and Moose Factory) : Quebec(Lake Mistassini) ; and Labrador (Hamilton Inlet). East toLabrador (Hamilton Inlet and Sandwich Bay) ; southeastern Quebec(Wolf Bay, Natashquan, and Perce) ; Nova Scotia (Baddeck, JamesRiver, Musquodoboit, and Halifax) ; Maine (Calais, Columbia Falls.Mount Desert, and Portland) ; Massachusetts (Boston and CapeCod) ; New York (Shelter Island) ; New Jersey (New Brunswickand Vineland) ; Maryland (Laurel) ; Virginia (Bush Hill and Sur-rey) ; western North Carolina (Roan Mountain, Mount Mitchell, andBlack Mountain) ; western South Carolina (Caesars Head) ; andnorthern Georgia (Brasstown Bald and Grassy Mountain). South,to northern Georgia (Grass}r Mountain and Cloudland) ; north-eastern Alabama (De Kalb County and Long Island) ; westernTennessee (Danville) ; southern Missouri (Holcombe and CurrentRiver) ; formerly northwestern Arkansas (Fa3^etteville) ; formerlyeastern Kansas (Manhattan) ; rarely Colorado (Estes Park andSweetwater Lake) ; southwestern Wyoming (Fort Bridger) ; north-ern Utah (Parleys Park and Barclay) ; central Idaho (Lardo) ; andnorthern California (Oak Bar and Eureka). West to northwest-ern California (Eureka) ; Oregon (Anchor, Empire, Eugene, andTillamook) ; Washington (Vancouver, Cougar, Olympia, ElkhornRanger Station, Elwha River, Ozette Lake, and Neah Bay) ; BritishColumbia (Alberni, Parksville, Nootka Sound, Fort Rupert, PortSimpson, and Hastings Arm) ; and Alaska (Juneau, Tanana Cross-ing, Lake Minchumina, Tacotna, Akiak, Russian Mission, andNulato)
.
74564?32 12
166 BULLETIN 162, UNITED STATES NATIONAL MUSEUMRuffed grouse have been extirpated from much of the south-cen-tral parts of their range, as in Iowa, Nebraska, Kansas, Missouri,and Arkansas. Casual occurrences outside the normal range are notnumerous, probably the most unusual being a specimen that wastaken near Camden, S. C, on December 27, 1904.The range as above described is for the entire species, which has,however, been divided into six subspecies. Bonasa u. umbettus oc-cupies the territory from Massachusetts, Virginia, and Georgia westto southern Minnesota, eastern Kansas, and northern Arkansas;B. u. togata is found north of umbettus, meeting it on the south inMassachusetts, central New York, northern Michigan, northern Min-nesota, and eastern North Dakota, while northward its range ex-tends to Maine, northern Quebec, and northern Ontario. The raceB. u. thayeri is confined to Nova Scotia and probably eastern NewBrunswick.The gray ruffed grouse {Bonasa u. umbelloides) is found fromwestern South Dakota west to the coast and Cascade Ranges inBritish Columbia, north to west-central Mackenzie, and south tonorthern Colorado and northern Utah; the Oregon ruffed grouse(B. u. sabini) occupies the Pacific coast region from HumboldtCounty, Calif., north through Vancouver Island and the adjacentmainland of British Columbia; and the Yukon ruffed grouse (B. u.yukonensis) is found in the interior of Yukon and in Alaska.Egg dates.?Southern New England and New York {umbettus) :72 records, April 5 to June 10 ; 36 records, May 9 to 24. New Jerseyand Pennsylvania : 15 records, April 20 to June 12 ; 8 records, May9 to 27. Maryland to Iowa: 13 records, April 28 to June 4; 7records, May 6 to 27. Michigan to Minnesota: 11 records, May 2to June 10.Northern States and southern Canada {togata and thayeH) : 52records, April 14 to June 26 ; 26 records, May 15 to 28.Mountain regions of Western States and Canada {umbelloides) :45 records, May 1 to July 10 ; 23 records, May 18 to 29.Oregon and Washington {sabini) : 40 records, April 19 to June 18;20 records, May 5 to 14.BONASA UMBELLUS TOGATA (Linnaeus)CANADA RUFFED GROUSEHABITSThe ruffed grouse of northern New England and eastern Canadais a grayer bird, with more conspicuous and darker markings below,than typical umbettus. Its range is roughly the same as the spruce
CANADA BUFFED GKOUSE 167grouse, or "spruce partridge," from which it is commonly distin-guished as the " birch partridge."William Brewster (1925) well describes its haunts in Maine, asfollows
:
Ranging from the lowest levels to the crests of the higher mountains it occurspractically everywhere, although seen comparatively seldom among heavy,unmixed spruce timber, and still more rarely in the larch and arbor vitaeswamps, so beloved by Spruce Grouse. It likes best to dwell in woodscomposed of intermingling evergreen and deciduous trees. Far back in theseit is found oftenest about deserted lumber camps, and along old logging roads,where enough sunlight has been let in to stimulate a vigorous growth ofunderbrush ; or along the courses of alder-fringed brooks or runs, where fernsflourish in rich, moist soil ; or on river banks freely exposed to the sun, butdensely overgrown with cornels, Viburnums, and other berry-bearing shrubs.Near permanent settlements it is given to frequenting wood edges, neglectedpastures, and the outskirts of crudely tilled farms, where young spruces,balsams, birches, maples, and alders have been permitted to spring up incrowded thickets about sunny little openings filled with tall bracken. Thebirds are here reasonably safe from human molestation, except in autumnwhen everyone possessed of a gun bags as many of them as he possibly can.Nesting.?The nesting habits of this grouse are similar to thoseof other ruffed grouse. Brewster (1925) describes three nestsfound in May, 1896, near Lake Umbagog, Me. Of one found onMay 14, he says : " This nest was directly under the main stem of afallen poplar, on a dry knoll wooded with second-growth poplarsand birches among which were interspersed a few balsams andspruces. It was 30 yards back from a public road and within 10yards of an open pasture." The next day he found one " at theedge of a thicket of alders covering rather wet ground, betweentwo large, buttressed roots of an old stump. Overspreading branchesof a small arbor vitae and Viburnum, growing close beside it,screened it so perfectly that the brooding bird could be seen onlyfrom the direction whence we happened to approach." On May16, another " was in a very exposed situation, quite outside of theborder of wild cherry, mountain maple, and other undergrowththat fringed an extensive forest, half-encircling an upland mowingfield, and in the field rather than in the forest, although but a yardor two from where the latter ended. Here it was sheltered fromobservation and from blazing sunlight by only a few dead sticks,the remains of a disintegrated brush fence."J. W. Banks, of St. John, New Brunswick, wrote to Major Ben-dire (1892)
:
Here with us a very common nesting place is what is called a fallow. Thisis a piece of woods chopped down in the fall, to be burned when sufficientlydry, usually in the latter part of May or early in June. Being composedchiefly of spruce and fir, it burns very rapidly. I found two nests (or ratherthe remains, for the eggs were badly scorched) in one of these burnt fallows,
168 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMand a few feet from each nest the bones of the mother Grouse. A farmeracquaintance told me of rinding a nest of this bird, which contained ten eggs,in a fallow he was about to burn, and knowing of another nest with an equalnumber of eggs, the thought occurred to him to put the eggs in the nest ofthe otiier bird that would not be endangered by the fire, and watch develop-ments. He had the satisfaction of knowing that the eggs were hatched.Eggs.?The eggs of the Canada ruffed grouse are similar to andindistinguishable from the eggs of the more southern race. Themeasurements of 71 eggs average 39.2 by 30.3 millimeters ; the eggsshowing the four extremes measure 44 by 31, 42.2 by 31.5, 38.6 by28.7, and 38.6 by 28.2 millimeters.Young.?Bendire (1892) quotes the following from Ernest Thomp-son Seton's notes
:
Every field man must be acquainted with the simulation of lameness, bywhich many birds decoy or try to decoy intruders from their nests. Thisis an invariable device of the Partridge, and I have no doubt that it is quitesuccessful with the natural foes of the bird ; indeed, it is often so with man.A dog, as I have often seen, is certain to be misled and duped, and there islittle doubt that a mink, skunk, raccoon, fox, coyote, or wolf, would fare nobetter. Imagine the effect of the bird's tactics on a prowling fox ; he has scentedher as she sets, he is almost upon her, but she has been watching him, andsuddenly with a loud " whirr " she springs up and tumbles a few yards beforehim. The suddenness and noise with which the bird appears causes the foxto be totally carried away ; he forgets all his former experience, he neverthinks of the eggs, his mind is filled with the thought of the wounded birdalmost within his reach; a few more bounds and his meal will be secured.So he springs and springs, and very nearly catches her, and in his excitementhe is led on, and away, till finally the bird flies off, leaving him a quarterof a mile or more from the nest.Plumages.?The molts and plumages are the same as in the otherraces. There are also both color phases, gray and red. As men-tioned elsewhere, these two color phases occur in all the races, butin this and in other gray races the red phase is rarer and less pro-nounced, the reverse being the case in the red races.Food.?The long list of food given for the ruffed grouse (umbellus)would apply equally well for this race, with due allowance for thedifferent species of plants and insects available. Manly Hardy, insome notes sent to Major Bendire (1892), says that it feeds not onlyon the poplar buds but also on the hard old leaves. He writes
:
I have killed one with its crop filled with such leaves on the 20th of August,and they eat them continuously, until the last have fallen in late October.They do this when other food is abundant. Buds of willow, yellow and whitebirch, hophornbeam, thorn plums, rosehips, leaves of tame sorrel, of the rockpolypod, fungus from birch trees, the seeds of touch-me-nots (Impatiens fitlra),wild raisins, and highland cranberries (both species of Viburnum) form alsoa part of their bill of fare. They seem to be especially fond of beechnuts. Ihave a record of finding seventy-six in one bird's crop and over sixty in another.
CANADA RUFFED GROUSE 169Ora W. Knight (1908) says:In the winter they " burl " seeming to prefer the yellow and white hirches,and the poplar, but also eating spruce, fir, pine, maple and in fact many otherbuds.Brewster (1925) writes:To the best of my knowledge the Birch Partridges of the Umbagog Regionnever eat the spills of coniferous trees, although subsisting almost wholly onthe buds of deciduous ones during rather more than half of the year. In latespring and early summer their food is gleaned mostly from the surface offorest-shaded ground and consists largely of insects and low-growing herba-ceous foliage of various kinds. Even where it is most plentiful the birds seldomlinger anywhere to seek it, but continue to advance, picking up a leaf or grubnow here, next there, so daintily and infrequently that they often ramble onslowly for a quarter of a mile or more before filling their crops. They aresomewhat less dainty and fastidious when feasting in late August on the fruitof low blueberry bushes, while in September I have often seen them alightat sunset in cornel or Viburnum (especially V. opulus) bushes on river-banksand literally gorge themselves in the course of a few minutes, almost withoutchange of foothold, on the berries which these shrubs commonly bear in suchprofuse and crowded clusters. Later in the season the pale orange fruit of themountain ash is similarly dealt with whenever it can be had plentifully, whichis not oftener than every other year. The glowing red berries of the blackalder are also eaten freely in late autumn. The birds seem to have littleor no liking for oats, but are exceedingly fond of buckwheat, and to obtainit will venture out fifty yards or more from neighboring coverts into stubblefields where it has recently been grown and harvested.Behavior.?Illustrating the tameness of ruffed grouse under prim-itive conditions, Brewster (1925) relates his experience with themin the Umbagog region early in the seventies, as follows:At first I undertook to hunt them with a setter, and to shoot them only onwing, as had been my practice in coverts nearer home. My good dog found andpointed them readily, but was evidently not a little puzzled to comprehendwhy they should stand conspicuously upright in open ground, or on mossylogs, regarding him with seeming indifference from a distance of only a fewyards, instead of rising far in advance, or crouching unseen in dense brush,as had been the unvarying habit of all birds of their kind with which he hadhad previous experience. When I stepped in ahead of the staunch setter withthe intention of flushing the Grouse, their behavior was still more surprisingespecially if, as often happened, there were as many as five or six together.For instead of rising promptly on wing as I wished, and expected them todo, they would begin a snickering outcry almost precisely like that of a RedSquirrel, nod their heads slightly a few times, and then start off at a slow walkwith crests erect and perhaps also widespread tails, shaking their heads andnecks, and twitching their expanded ruffs at each deliberate step, and con-tinuing unceasingly to utter their derisive and unseemly snickering. Thiswas most likely to happen in a narrow cart-path tunneled through the forest,or on the outskirts of some woodland opening. In either case the birds hadseldom far to go before reaching fallen tree-tops, or dense evergreen thickets,from which it was difficult if not impossible to dislodge them, at least by theaid of a dog, who would never flush his birds. Into such sanctuaries they com-
170 BULLETIN 162, UNITED STATES NATIONAL, MUSEUMmonly skulked on foot, if not too closely pressed. Even when I forced themto take to wing by running after them, they rarely went more than a fewyards before alighting in a tree, or dropping again to the ground, over whichthey might continue to hasten, if much alarmed, until it was useless to followthem farther.After this and several similar experiences, he learned that the onlyway to secure any of these unsophisticated grouse was to " pot
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them in the " time-honored fashion of local hunters " by shootingoff their heads as they sat on trees, bushes, or logs, or on the ground.More than once he was able to bag all the birds in a flock, shootingone at a time, while the survivors refused to fly. While hunting forducks one day in a boat, he saw a grouse sitting on a log. " When Ifinally shot her," he says, " the report of the gun started six others,hitherto unseen, although close about her. Flying only a few yards,they alighted in trees and bushes within plain view, and remainedthere gazing at us while, sitting in the bow of the boat, I loaded andfired until the last bird had fallen."Such shooting was unsportsmanlike, of course, unless the huntergave the birds a chance by using a rifle and shooting at the head. Itwas the only method in those days, but is no longer necessary nowin any but the more remote regions, for the birds have become edu-cated and have learned to give the sportsman a chance to show hisskill. Good sport may now be had with dog and gun in these samewoods.Fall.?Brewster's (1925) very full and exceedingly interesting ac-count of the Canada ruffed grouse contains several references to amigratory movement or an erratic autumn wandering, similar, per-haps, to their queer behavior during the restless " crazy season " thatwe have noted elsewhere.. I quote a story, one of three similarincidents, told him by Luman Sargent
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After spending an October night (before 1870) in the old log-camp of MiddleDam and leaving it at about sunrise the next morning, he had gone only ashort distance into neighbouring woods when he saw a Partridge on a log andshot it. Startled by the report of his gun so many birds of the same speciesrose far and near on wing, one after another, that for half a minute the airseemed full of them as far as the eye could range through trees and brushwood.There could not have been less than one hundred of them, he said. When fol-lowed they all kept moving on by successive flights in the same direction, and tothe southward. They were so restless and shy that he had difficulty in gettingnear them and was able to kill only a few more before losing track of themaltogether.Winter.?Ernest Thompson Seton (Ernest E. Thompson, 1890),referring to the habits of the ruffed grouse in Manitoba, says:It seems to be the normal habit of this bird to roost in a snow drift during thecoldest weather. The wonderful non-conductivity of the snow is well known,but may be forcibly illustrated by the fact that although the thermometer
GRAY RUFFED GROUSE 171
registers 35? below zero, the 10 inches of snow which fell before the severefrost came, has effectually kept the wet earth in the woods from freezing,although the temperature has been at or below zero for over a week. Inview of these facts it is easy to understand that the grouse in the snow driftare quite comfortable during the coldest nights. In general the bird will befound to run about before burrowing into the drift; each makes its own bed,usually 10 or 20 feet from its neighbor ; they usually go down a foot or so andalong 2 feet ; they pass the whole night in one bed if undisturbed, as the largeamount of dung left behind would indicate. They do not come out at theingress, but burst through the roof of their cot at one side ; they do not usuallygo straight ahead and out, because their breath during the whole night hasbeen freezing into an icy wall just before their nostrils.BONASA UMBELLUS UMBELLOIDES (Douglas)GRAY RUFFED GROUSEHABITSThe name gray ruffed grouse describes very well the characters onwhich this race of the ruffed grouse is based; it is a decidedly graybird. Its range extends from Mackenzie south through the RockyMountain valleys and east to the western edge of the Great Plains.The birds found in Manitoba, of which we collected a good series,are intermediate between this and togata. It intergrades withsabini where their ranges meet.Harry S. Swarth (1924) found it " abundant throughout the pop-lar woods of the lowlands " in the Skeena River region of northernBritish Columbia, where the birds are " less grayish, more brownish "than typical unibelloides. In the Stikine River region he (1922)found the birds " relatively gray colored, but not so ashy " as thebirds from the Yukon region (yukonensis) ; they are practically thesame as unibelloides from Alberta.In the Glacier National Park, Mont., Mrs. Florence M. Bailey(1918) found this grouse " in the pines and aspen thickets of theeastern slope and also in the dense hemlock woods of the westernslope of the mountains." M. P. Skinner has sent me the followingnotes on its haunts in the Yellowstone National Park, Wyo.
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This grouse is more likely to be in the aspen groves than Richardson'sgrouse is. In fact, it is quite apt to prefer the aspen groves, but it is alsofound in forests of fir, mixed aspen and fir, lodgepole pine, mixed aspen andlodgepole, and in the spruce forests. Although this grouse is rather scarcein Yellowstone National Park, it frequents the forest in all sections between6,500 and 8,500 feet elevation, but I do not find them in the timber-line forests.It seems to be in heavy, thick standing forests, open forests, and thick saplinggrowth. Occasionally it is found out on the grasslands, but not so much asRichardson's grouse. Still, I think the gray ruffed grouse rather prefers anopen stand of trees, especially if berries are present.Courtship.?Skinner says that " as early as March 20 the malesbegin to strut and court the females. One day I found a pair near
172 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMTower Falls. The female sat as if roosting on a horizontal pinebranch, 4 feet from the ground, in a clump of thick-growing sap-lings; a male, with ruffs spread and tail spread in an upright fan,strutted about below. He may have been drumming, although Idid not actually hear him."J. H. Riley (1912) heard it drumming in the fall in Alberta; hesays : k - When we first made camp, September 5th, and for about aweek thereafter, we seldom heard this bird drum, but before wedeparted, September 22nd, one used to drum at intervals throughoutthe day, though the weather was cloudy with rain and later snow."Thomas T. McCabe writes to me that at Indian Point Lake, nearBarkerville, British Columbia, he heard one drumming for man}'weeks, not on the usual log, but on the large sloping root of a bigspruce on the lake shore, 175 yards away from a nest he had found
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he watched him drum at the same spot 10 days after the nest hadbeen robbed.Nesting.?Mr. McCabe very kindly collected and sent to me thisnest and the 10 very handsome eggs it contained. He took it onMay 27, 1929, at Barkerville, British Columbia, and he writes tome thatthe nest was in heavy, open, mixed, second-growth spruce and balsam {Abieslasiocarjja) , which had nearly the dignity of primeval forest, and was car-peted with our universal green moss. It was wedged between two roots of a9-inch balsam, and sunken to the extent that the top was about level with themoss. There was absolutely no bushy or herbaceous concealment. The brood-ing bird left in each case when we were from 15 to 20 feet away, but it didnot approach us. It kept at a distance of about 40 feet, running, crouched low,whining like an eager hound, and then flew away. The leaves lining the nestare Populus tirenwiloides, seldom far to seek here, even in the deep timber.Major Bendire (1892) mentions a nest found in Montana, "underthe trunk of a fallen cottonwood tree, which rested about a footfrom the ground. Otherwise the nest was not concealed in anyway." Another nest near Nulato, Alaska, was " found in an oldwillow stump." In Yellowstone National Park, Mr. Skinner recordstwo nests in his notes; one was in a grove of quaking aspens, undera fallen tree ; the other was at the foot of a lodgepole pine, a hollowin the pine needles.Eggs.?The eggs of the gray ruffed grouse are practically in-distinguishable from those of its eastern relatives. The measure-ments of 55 eggs average 40.3 by 29.8 millimeters; the eggs show-ing the four extremes measure 43 by 31.5, 38 by 30. and 40.7 by 28.7millimeters.Young.?Swarth(1922) found two broods of young in the StikineRiver region, of which he writes
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The young of one brood were still unable to fly. Our first knowledgeof their presence was derived from the mother bird, who burst forth from
GRAY RUFFED GROUSE 173the bushes and charged us furiously. She kept tail and ruff widely spread,the head crest depressed. She was mewling in a very catlike fashion, andalso hissing from time to time. There was an occasional faint peep from thegrass nearby, and once I caught a glimpse of a yellow chick slipping awaythrough the shubbery, but the young were too agile to be captured.The young of the second brood were somewhat larger and able to fly. Thissecond mother tried to toll us away from the chicks by feigning a brokenwing; the noise she made was not unlike the whining of a small puppy.Her actions, all together, gave the impression that she was frightened ratherthan angry. However, if frightened, she still did not desert her trust, but re-mained nearby, dragging herself back and forth across the road, with wingsdropping and all her feathers pressed closely against her body. Her tail wasnot spread nor were her ruffs displayed at any time, all in striking contrastto the behavior of the first bird that morning.Plumages.?The molts and plumages are similar to those of theeastern bird. It has a red phase, which apparently occurs about asoften as the gray phase occurs in the eastern birds. Swarth (1924)collected two red-tailed birds in a series of 14 in the Skeena Riverregion. Ernest Thompson Seton (1885) has described the red phase,as it occurs in Manitoba, as follows : " In general appearance thisbird differs but little from the well known Bonasa umbeUus umbel-loides, but it is distinguished by being more decidedly marked?thusthe bars on the belly are complete and nearly black?and by havingcopper-colored touches on the back, the subterminal tail-band andthe ruff a rich, iridescent, coppery red." He says that about 10per cent of the birds have copper ruffs and only about 20 per cent arepure umbelloides.Food.?Skinner says of its food in the Yellowstone : " In spring,they eat the buds of aspens. In summer, they frequent berry patchesand sometimes visit small openings for grasshoppers. In winter,they are said to eat mistletoe berries." In the Stikine region Swarth(1922) found that the food was practically all vegetable matter,consisting of leaves and stems of Populus, Galium, Artemisia, andViburnum, with a few berries.Behavior.?Skinner contributes the following notes on the behaviorof the gray ruffed grouse
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These are resident birds, remaining throughout the year in one locality, andI see no evidence that they even move up and down the mountains, as otherresident species do. But in winter they live mostly in the treetops, and wedo not see them on the ground much before March 15 or April 1. Althoughthese birds have the same habits and probably the same disposition towardtheir environment as the eastern ruffed grouse originally did, they have retainedtheir comparative tameness toward man under the protection accorded inYellowstone National Park. My notes are full of references to them as " quitetame " and " very tame," but occasionally I find one that is wild. The tameones are recorded in all places, at every elevation within the bird's range, and atall seasons. This unsuspicious attitude extends also to men on horseback, thegrouse sitting or remaining in their tracks while the horseman rides by, although
174 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUM
sometimes a grouse will move around to the opposite side of a tree fromthe intruder. I have even been permitted to ride past a grouse within 5 feet.At times, especially in autumn when they frequent the roads, the grouse oftenrun across in front of a horse and even apparently under his nose. I have seenmy horse almost step on a grouse. Sometimes they run ahead of my horsedown the trail. Even a mother with young is often extraordinarily tame, andshe does not make use of the broken-wing tactics nearly so often as unprotectedgrouse do. Often a bird springs directly from the ground to a tree overheadand allows me to pass directly under. Still, the gray ruffed grouse is not so aptto come about houses and hotels as Richardson's grouse.BONASA UMBELLUS SABINI (Douglas)OREGON RUFFED GROUSEHABITSIn the humid coast belt, west of the Cascades, in Oregon, Wash-ington, and British Columbia, we have the darkest, most richlycolored, and one of the handsomest races of the ruffed grouse. Thegrouse of the southern Alleghenies are quite richly colored, but theywill not compare in this respect with these western birds, whichBaird, Brewer, and Ridgway (1905) describe as follows:The upper parts are dark orange chestnut, mottled with black, the cordatelight spots very distinct. The feathers of the breast are strongly tinged withreddish-yellow; those of the sides marked with broad and conspicuous bars ofblack, instead of the obsolete brown. The under tail-coverts are orange-chest-nut, within distinct bars of black, and an angular terminal blotch of white. Allthe light brown blotches and edgings of the eastern variety are here dark brownor black. The jugular band between the ruffles is very conspicuously black.They say of its haunts, quoting J. K. Lord
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Dr. Cooper also speaks of this grouse as very abundant everywhere aboutthe borders of woods and clearings. It was common near the forests east of theCascade Mountains up to the 49th degree. In the spring their favorite hauntis in the vicinity of stagnant pools, or in the brush around a marsh in whichthe wild swamp-crab, the black birch, and the alder grow.William L. Finley (1896) writes:They are generally found on low land, a river bottom or along some smallcreek, but in times of high water, they will go to higher ground. I have oftenseen them when the water is high, in some small tree or bush, when the waterwas several feet deep under them, and around them for a half mile. In a boat,at such times, one can row right under the bird, or within a few feet of it.A great many are killed along the river bottoms in this way by hunters.Courtship.?The drumming performance of this grouse is appar-ently the same as with the eastern birds, but "Mr. Lord also statesthat he has seen the males of this species fighting furiously duringthe pairing season. Ruffing up their necks, with their heads andoacks almost in a straight line, and with wings dropped, they circleround and round each other, striking and pecking until the van-
OREGON RUFFED GROUSE 175quished gives in, and the victor mounts upon a log and proceeds todrum furiously."Nesting.?W. Leon Dawson (1909) says: "At the foot of a maplein some swampy thicket, or close beside a fallen log, the femalescrapes a slight depression in the earth, lining it roughly with deadleaves and a few small twigs."In some notes sent to me by D. E. Brown, he describes four Wash-ington nests. One was at the foot of an elder tree in an old riverbed, in the dead leaves, with no cover at all. Two nests, found April19, were in open woods, under the ends of logs. And one, found May6, was at the side of a log at the edge of large woods. In all cases theeggs were laid on dead leaves. Bendire (1892) describes a nest foundon Vancouver Island as " a slight hollow in the ground scratchedout by the bird, placed under the fallen branches of a spruce tree.The cavity was lined with dead leaves and spruce needles, as well asa few feathers. This nest was found close to a small creek and waswell concealed."William L. Finley (1896) reports a nest found by G. D. Peck inOregon that contained eight eggs of the sooty grouse and seveneggs of the ruffed grouse; the sooty grouse was flushed from thenest. He has also sent me a photograph of a nest containing eighteggs of the ruffed grouse and three eggs of the ring-necked pheasant.Eggs.?The eggs are indistinguishable from those of other ruffedgrouse, though they may average slightly darker in color and some-what more often spotted. The sets will average smaller in number.The measurements of 58 eggs average 41 by 30.2 millimeters ; the eggsshowing the four extremes measure 44 by 31.5, 43.5 by 32, 38 by 29,and 40.9 by 28.2 millimeters.Plumages.?Confirming what I have said elsewhere about thepresence of both color phases in all races of the ruffed grouse, H. S.Swarth (1912), referring to a series collected on Vancouver Island,says:The dichromatisru of the species is very apparent in the series collected,the gray and the red-colored birds being conspicuously different. Those inthe gray phase are quite uniform in color and markings, but the reddish birdsshow considerable variation. The former all have black ruffs, and gray tailswith a black subterminal band. Of the reddish birds some have red ruffs,some black, and others are variously intermediate. Some have a gray tailwith a red band, some a red tail with a dark band, and one a red tail with adarker red band. These different styles of coloration are not indicative ofage, sex, or season, for both phases are represented among adults and immaturesof both sexes.Food.?The food of the Oregon ruffed grouse is similar to thatof its eastern relatives, differing only with the available supply ofberries, seeds, leaves, fruits, and insects. William H. Kobbe (1900)writes: "They are extremely fond of the small wild crab apples
176 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM(Pyrus rivularls) which grow in the low, damp woods. The birds-visit these trees very early in the morning and late in the evening,at which times they may be found silently perched upon thebranches."Mr. Dawson (1909) adds: "They are fond of the fruit of theCascara, which they gather from the ground; and wild crab-applesare favorites in season. These last ripen about the middle ofOctober, and from that time until the alders bud again these Grouseare often to be found in evergreen trees."Behavior.?In general the habits of the Oregon ruffed grouse donot differ from those of the rest of the species, except so far as theyare affected by differences in environment. In the wilder sectionsthey are quite unsophisticated.Enemies.?J. H. Bowles (1901) tells the following remarkablestory
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The carnivorous habits of chipmunks as related in the recent issues of TheCondor were very interesting to me, though I believe mice are far more guilty.Mice are a perfect pest to ground-builders in this country, as they burrow intothe ground several yards away from the nest and then tunnel until they reachthe bottom of the nest.They then dig upward into the nest and carry the eggs into their tunnel toeat. I have often found broken and unbroken eggs several feet from the nestin a burrow. I have never actually seen mice do this, but the tunnels are muchloo small for anything else. The Ruffed Grouse (Bonasa umbellus sabini) arethe worst sufferers that I have yet found, and their eggs are the largest that Ihave seen destroyed in (his manner. All the small ground-builders suffer moreor less.Game.?Dawson (1909) writes:From the point of view of the sportsman, this bird is not to be compared withthe Ruffed Grouse of the Eastern States. Its cover is too abundant, and it doesnot take the discipline which has educated the wily " partridge." It seldomallows the dog to come to a correct point, usually flushing into the nearest smalltree, where it sits peeping and perking like an overgrown chicken, regardingnow the dog and now the hunter. Pot-shooting the birds under these circum-stances can hardly be called sport, but their fondness for dense thickets oftenmakes it the only way in which they can be obtained.Edwyn Sandys (1904) evidently agrees with Dawson, for he says:In British Columbia the sport, as found, could not compare with that of theEast. Those who know the wonderful western province will readily guess why.In many places the trees almost rival the famous big conifers of California,and they are crowded together as thickly as it is possible for such mightytrunks to stand. Frequently the lower spaces are filled with ferns of such sizeand luxuriance as to suggest semitropic lands rather than a portion of Canada.In such cover the keenest of guns can do little or nothing. The writer is oversix feet tall, but in that cover he felt like a veritable babe in the wood. Thesize of the firs was almost oppressive?but the ferns?ye gods ! such ferns.In places they grow like the big western corn, close and rank, towering a yard
NOVA SCOTIA BUFFED GROUSE 177
or more above one's head. Among them, grouse after grouse can buzz awayunseen, while, in addition, the tremendous fronds combine to form a mostbaffling light. BONASA UMBELLUS THAYERI BangsNOVA SCOTIA RUFFED GROUSEHABITSThe ruffed grouse of Nova Scotia had long been recognized andwas finally described by Outram Bangs (1912) under the name B. u.thayeri, given in honor of Col. John E. Thayer. He designates it assimilar to Bonasa umbellus togata (Linn.) but general color of upper partsdarker, more dusky or sooty, less grayish; the whole underparts (except throat)heavily and regularly banded with dusky, the dark bauds much blacker andmuch more boldly contrasted against the ground color?less blended.B. umbellus thayeri presents two phases of coloration, which are both verydark, and not very different ; a phase in which there is much dull chestnut orburnt sienna in the upper parts and tail and another in which the tail iswholly dull gray and black and the upper parts are but little varied with darkchestnut markings. The color and markings of the underparts is not differentin the two phases, except that very reddish birds sometimes have the bases ofthe feathers of the upper chest dull chestnut instead of dusky.Bangs says further in regard to it:Some years ago I was accustomed to go shooting every autumn in NovaScotia, and each season I was more and more impressed by the very darkcoloration of the Ruffed Grouse killed there. I therefore made into skinsduring my last two shooting trips to this province a series of sixteen Grouse.It is probable that the new form is confined to the almost insular provinceof Nova Scotia, although I cannot be sure about the bird from the coast ofNew Brunswick as the specimens I have before me are in worn midsummerplumage, and not comparable with the Nova Scotia specimens, all of which weretaken in October.We have no reason to think that the habits of this grouse, or anyof the chapters in its life history, are essentially different from thoseof its neighbors in other parts of Canada where the environment issimilar.Eggs.?The eggs are indistinguishable from those of other ruffedgrouse. The measurements of 47 eggs average 40.5 by 30.5 milli-meters; the eggs showing the four extremes measure 44 by 33, 37.9by 30, and 40 by 29.2 millimeters.BONASA UMBELLUS YUKONENSIS GrinnellYUKON RUFFED GROUSEHABITSThe ruffed grouse of the interior of Alaska and Yukon is, accordingto Dr. Joseph Grinnell (1916), the " largest and palest of the races ofBonasa umbellus; nearest like B. u. umbelloides, but general colora-
178 BULLETIN 162, UNITED STATES NATIONAL MUSEUMtion of light-colored parts of plumage more ashy, and pattern ofdark markings finer." He says also that it
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occurs along the Yukon River Valley down nearly to its rnouth, as also inadjacent wooded areas west even into the Seward Peninsula.As with the other subspecies of the Ruffed Grouse, yukonensis shows two colorphases. Three out of the eleven specimens at hand have pale rusty tails; buteven in this " red " phase the race is distinguishable from the correspondingphase in the other subspecies by paler tone of coloration. Typical umbelloidesis still a gray bird, but its grayness is more leaden, and its browns and blacksare deeper. The extreme fineness of the intricate pattern of barring andmottling on the plumage is in yukonensis an appreciable character.Dr. E. W. Nelson (1887) writes of its haunts:Like the Spruce Grouse, and sharing with the latter its range in NorthernAlaska, this bird is found everywhere where wooded land occurs, reaching thehead of Norton Sound and vicinity of Bering Straits, following the belts oftimber as they approach the sea in this portion of the Territory. It is notuncommon in the vicinity of Nulato, where it frequents the deep spruce growths,and feeds exclusively upon the buds of these trees, its flesh being tainted inconsequence. Dall found it nesting there in May, and a set of eggs was foundin a willow stump. Like the Spruce Grouse, this bird is found whereverspruces occur, and both species range well into the Kaviak Peninsula, so thatthey are found within a very short distance of Bering Straits.LAGOPUS LAGOPUS ALBUS (Gmelin)WILLOW PTARMIGANHABITSThe willow ptarmigan, with its various so-called subspecies, is ofcircumpolar distribution, inhabiting the arctic or subarctic regions ofboth the North American and Eurasian Continents. European writ-ers know it as the willow grouse, applying the name ptarmigan toonly the rock ptarmigan and its races. It seems more logical toapply the name ptarmigan to both species that assume a white plum-age in winter. The British red grouse is the only species of thegenus Lagopus that does not have a white winter plumage; it does,however, have the feathered toes, one of the principal characters ofthe genus. The following account includes the so-called Alaskaptarmigan.Spring.?As soon as the first bare spaces appear on the sunny slopesof the tundra the ptarmigan begin their spring migration from theirwinter quarters, among the willows in the sheltered valleys of therivers and creeks of the interior, to their breeding grounds on theopen hills or tundra. H. B. Conover says in some notes he sent to me
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On our sled trip from Nenana to Hooper Bay, this grouse was not encountereduntil we reached the Kuskokwim Mountains, where we found it very plentiful.It was generally encountered in flocks of 15 to 100 feeding among the dwarf
WILLOW PTARMIGAN 179
willows. A few males shot on April 5 had an occasional brown feather showingon the head and neck. The morning of April 24 we left Mountain Village onthe Yukon and cut across the tundra for Hooper Bay, a journey which took usfour days. For the first two of these we encountered ptarmigan everywherealong the willow-bordered sloughs and creeks. The hens were still white, butthe heads and necks of the cocks were about a third into the red spring plumage.As we approached the coast and the willows became scarcer, these birds were nolonger seen. On May 9 the ground about Point Dall was beginning to show inspots through the snow and the first ptarmigan made their appearance. Twodays later they were common. Each male now took possession of a little spotof bare ground, whence he sent out his challenges, com-ere, com-ere, go-bec, go-bec. Between calls they would bob their heads as if they were pecking at theground, or, jumping about 6 feet into the air, glide down to the earth, cacklingas they descended. The hens seemed to have but one call, a cackle similar tothat of a tame chicken. Often two cocks were seen chasing each other aroundover the tundra, but only rarely would they seem to stand and fight it out.In the evenings and early mornings these birds were especially noisy, and oftenit was no great stretch of the imagination, what with the calls of the water-fowl, to imagine oneself in some great barnyard. About this time the Eskimoboys began to range the tundra with their bows and arrows, and many anunwary cock and sometimes a hen were killed by the blunt shafts of these 8 and10 year olds.Courtship.?Edwyn Sandys (1904) gives the following very goodaccount of this performance
:
The love-making of the ptarmigan is not unlike that of the Canada Grouse,or " spruce-partridge." The males, with their plumage changing from whiteto the handsome summer dress, strut with all the pomposity of their kind.The red combs over the eyes are swollen and very conspicuous, as the birdstruts with head thrown far back, tail raised and spread, and wings trailing.Presently he leaps into the air, raises himself higher and higher with a vigor-ous flapping, then sails on set wings through a descending spiral, which bringshim back to his starting-point. While thus a-wing, he utters a curt, gruffchallenge oft repeated, a defiance to all rivals. Again he struts, and again goesinto the air, frequently to see male after male arise from near-by stations.While so occupied the birds make considerable noise, the bark-like challenge ofother calls being heard for some distance. Meanwhile, the females loiter aboutin the cover, admiring the efforts of the males, and gradually acknowledgingtheir charms. The inevitable battles follow?spirited encounters, in whichmany hard knocks are given, and much pretty plumage marred, until the weakerhave been well whipped.Nesting.?Herbert W. Brandt has sent me some elaborate notes onthis species. He says of its nesting:The willow ptarmigan at Hooper Bay is not at all particular as to the locationof its home site, for it dwells impartially from the drift-strewn sea beach to thehigher altitudes on the mountains. Down under the protection of a drift log, aclump of grass, a small bush, a mossy hummock, or any screeny object, shescrapes out a cavity to fit her requirements. This she lines more or less withany material at hand, and here she deposits daily her rich crimson egg. Duringthe period of egg accumulation, we found the nest to be covered with surroundingmaterial, because the bird does not begin to incubate until the full complementis satisfied. When the first egg is laid there is but little form to the nest, but
180 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
as the set progresses, the birds mold it into the proper shape, and by the timeincubation has progressed, the eggs snuggle together in a well-cupped basin.The brooding female spreads her feathers as does the sitting domestic hen, andwhen her frail body is examined in hand, it seems almost incredible that she cancover so large a clutch of eggs. The extreme measurements of 12 nests exam-ined are : Total height, 5 to 7 inches ; inside diameter, C to 8 inches ; depth ofcavity, 3 to 6y2 inches.F. Seymour Herse}T collected some 10 sets of eggs for me betweenSt. Michael and the mouth of the Yukon. They were mostly infairly open situations on the tundra. One nest was a depression inthe tundra moss at the base of a small clump of grass; it was linedwith dry grass, leaves, and a few feathers ; it measured 7 inches acrossand 5 inches deep. Another was in a hollow under a dwarf willow ona raised mound on the tundra. Another was in a deep hollow in a wetplace on the border of a marsh, a very open situation; it had a heavylining of dry grass and a few feathers. Some of the hollows werevery shallow, not more than 2 inches deep. There were three sets of7, two of 8, one of 9, three of 10, and one of 11 eggs.George G. Cantwell writes to me of a nest he found on CopperRiver, Alaska, that was " placed on an open river bar among a lightgrowth of willows, close to a growth of mountain spruce/' StantonWaiburton. jr., tells me of a nest he found late at night, saying:To preserve them intact for a photo in the morning, I put my khaki coatover the nest and eggs, completely covering them. Then (as an experiment)1 wrinkled up the collar by the eggs so that it formed an opening not over2 inches high. This slight opening did not expose the eggs to view from anyangle, as they were still completely covered. Early the next morning when Iapproached, the female flushed from right beside my coat. All 11 eggs werenow in a new nest just outside my coat, not over a foot from the originallocation. During the night the female had made a new nest, moved all 11 eggsinto it, and recommenced incubation.MacFarlane mentions in his notes another case where the eggswere probably removed by the birds. He " had reason to know thatsome, at least, of the nests were used by ptarmigan several seasonsin succession."Joseph Dixon (1927) writes:The male ptarmigan spends the day hiding in little thickets, keeping within50 or 100 feet of the nest. He has a definite form or nest of his own which lieoccupies when roosting. One reason for his staying so close is the danger ofShort-billed Gulls finding the ptarmigan's nest. These egg thieves work inorganized gangs, usually three together. One will swoop down at. the female.trying to make her shift about on the nest so as to expose the eggs. The secondor third gull following tries to slip in and grab an egg. As soon as the gullsappear, the hen ptarmigan gives a peculiar call for help. Upon hearing tinsthe cock ptarmigan bursts forth like a rocket and charges the thieving gulls.He doesn't beat around the bush but Hies directly at the intruders, knockingthem down with the impact of his body. An average cock Willow Ptarmiganat this season weighs ;!07 grams, while one on the Short-billed Gulls which was
WILLOW PTARMIGAN 181
shot weighed 358.2 grams. In addition to being one-third heavier than thegull the cock ptarmigan flies much the faster of the two, and when he hits agull it is almost like a Duck Hawk striking a duck.Eggs.?The willow ptarmigan lays commonly 7 to 10 eggs; asmany as 17 have been found in a nest, and in late or second setsthere may be only 5 or 6. An egg is laid each day, and incubationdoes not begin until the last egg is laid. For a description of thebeautiful eggs, I can not do better than to quote Mr. Brandt'sremarks, as follows
:
Eggs in the same set follow the same type of ground color, the same styleof markings, and are nearly uniform in size ; and as each bird seems to lay atype of egg individually its own, it may be noted that scarcely any two sets ofeggs of this interesting species are exactly alike. In shape the egg is almostalways ovate, but in rare instances tend to short ovate with stubby ends, andnearly elongate ovate when more slender in shape. The surface of the shellis smooth and often somewhat greasy like that of a duck egg; the texture ishard ; and the egg sturdy like its parents. The luster is rather shiny and thisapparently increases somewhat as incubation progresses.A study of the coloration of the egg of the willow ptarmigan is of specialinterest because of its change of hue after it is laid. When the egg firstappears, the markings are from " ox-blood red " to " scarlet red," and the wholesurface is vivid and moist, and it appears as if it has been dipped in fresh,red paint. This undried pigment is very easily rubbed off, and in consequence,it is unusual that an egg does not show the sign somewhere on the surface ofbeing brushed by the feathers of the parent bird. In fact, these rubbed spotsmay show distinctly the individual barbs of the ptarmigan feather thatscratched it when it was wet. As the moisture from the egg dries, however, thepigment sets rapidly, and at the same time so darkens, like congealed blood,that by the time the eggs are a few hours old, the brilliant reds turn to blackishbrown. Once the pigment becomes dry and sets, it is very durable, and egg-shells that have lain out in the weather from the previous year still retain theirbold markings.The ground color is usually inconspicuous, because it is seldom that morethan half of it is visible, and often it is all practically hidden by the overlyingspots. The only place that the ground color is prominent is on those areaswhere there is an aforementioned rubbed spot. This ground color exhibitsmany variations of the pale creamy tints, such as " ivory yellow " and " sea-foam yellow," but a few sets are further decorated with paler reds, making theground color " orange-crimson " to " vinaceous-tawny " and " pecan brown,"while the ground color of one egg is even " ocher red."The markings are the richest of any egg we collected at Hooper Bay and areirregularly flecked in profusion all over the surface. In size, these spots rangefrom the finest pepperings to blotches thumb-nail in size, and are all more orless confluent, some so much so that they cover the surface and almost envelopethe ground color. When the spots are large, the ground color is often welldefined, and then the most handsome effects are produced. If the ground coloris distinctly reddish, the surface markings are usually not nearly so numerous,and because of their sparse distribution, the egg then approaches in appearancethat of the spruce grouse {Canachites canadensis). The markings show almostuniformly blackish brown with a reddish suggestion, yet where the pigmenthas been scratched very thin, the color is often as light as "maroon" and ev. u74564?32 13
182 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
" garnet brown," while the deeper colors are " warm blackish brown " and
"blackish brown (1)."The measurements of 250 eggs, in the United States National Mu-seum, average 43 by 31 millimeters; the eggs showing the four ex-tremes measure 48 by 31.5, 46.5 by 34, 39 by 30, and 39.5 by 28millimeters.Young.?Mr. Conover says in his notes
:
Newly hatched young were first found on June 22. The incubation periodseems to last about 22 days. A nest found on June 2 with 11 eggs had 12 onJune 3, and on being visited on June 25, it was found to be empty. The chicksare very precocious. One day a hen was flushed from a nest containing twoeggs and eight youngsters still damp. Hardly had she left when every downychick scrambled weakly from the nest and attempted to hide in the grass. Theminute they were replaced, out they would go again, until finally they becametired out and stayed in the nest. Toward the end of June broods were con-stantly encountered about the tundra. Both parents were always with themand the cock was especially combative, although discretion always got thebetter part of valor. The young after running a few feet would suddenly dis-appear, whereupon the hen would join the male in threats and attempt to leadone off. It was amusing to imitate the peeping of a chick and watch the cockgo into a frenzy, ruffling himself up, making short dashes here and there, andin unmistakable language telling you just what he was going to do if you didn'tget away from his children. After a few minutes of this, both birds would beworn out and would retire a short way to watch for your next move. By July22 the young were about a third grown and had begun to shed their first brownprimaries and grow their new white ones. The adults were then in the midst ofshedding their toenails.Dixon (1927) noticed that the family party traveled as follows:First came two or three chicks in the thick grass, then the mother surroundedby the other chicks ; the cock sometimes led and at other times brought up therear. I timed them and found that they covered a lineal distance of 45 feet infive minutes. Following this there came a period of rest of five minutes, duringwhich the mother hovered her brood of young. We never saw the cock hoverthe young; but when one of the chicks became entangled in a network of twigshe was right there and helped it get free. By noon the ptarmigan family hadwandered out in the low bushes 100 yards from where they had hatched. Thechicks were now nearly 24 hours old, and all of them were strong and lusty,each able to run about with agility and to secure food for itself. At CopperMountain, about 4 o'clock in the afternoon of July 12, a family of WillowPtarmigan came feeding along through the dwarf willows near camp. Therewere six young about the size of quail. The cock kept a lookout for enemiesfrom elevated positions while the hen herded the young along through thewillows. The hen kept up a running conversation with the young as did alsothe cock. This liaison note was a loud ke-onck, repeated at intervals of fromfive to ten seconds. The cock's call was somewhat coarser than that of the hen.I had difficulty in hearing the thin peeping of the chicks at a distance of fiftyfeet, but it served to keep them together. The young were very active, jumpingup into the willows and catching insects over a foot off the ground.Plumages.?In natal down the young willow ptarmigan has a largepatch of " burnt sienna," bordered with black, on the center of the
WILLOW PTAEM1GAN 183crown and occiput; the rest of the head and the underparts are" colonial buff "; the upper parts are variegated with " colonial buff,"black, and " cinnamon-rufous," the last mainly in the center of theback and rump, bordered by broad black bands ; a small spot on thelores, a larger auricular spot, and a narrow line behind the eye areblack.The juvenal plumage comes in first on the wings, then on thescapulars and back, while the chick is very small; the juvenal remigesare " sepia," bordered with " cinnamon-buff " or buffy white, andtheir coverts are tipped with buff ; the last of the down to disappearis on the chin, neck, and belly. The juvenal plumage is at its heightbefore the young bird is half grown, at which time the postjuvenalmolt begins, early in August. In full juvenal plumage the feathersof the mantle are black, edged, notched, or barred with " ochraceous-tawny," and with triangular white tips; the breast and flanks are
" ochraceous-tawny," heavily barred or spotted with black or dusky
;
the belly is buffy or grayish white ; the remiges are as stated above,except the outer two, which are the last to appear and are white;the rectrices are black, edged, spotted, or barred with " ochraceous-tawny."When the young bird is nearly half grown the postjuvenal moltbegins by shedding the juvenal remiges, which are replaced by whiteones. This is a more or less incomplete molt, involving most of thewings (except the two pairs of outer white primaries) , the tail, anda varying extent of the body plumage. In this intermediate, orpreliminary, winter plumage, the sexes are still practically alike.There may be only a few scattered, reddish brown, finely vermicu-lated, or mottled feathers, or the renewal may be nearly complete;the belly, flanks, and legs become white.This intermediate plumage is worn for a very short time whilethe young bird is getting its growth ; for, during the last of Augustand through September, a supplementary, partial molt takes place,which completes the change into the white winter plumage. Youngbirds can be distinguished during their first winter by the outerprimaries, which are more worn and often speckled at the tips.Both young and old birds have a partial prenuptial molt in spring,the date varying greatly with the latitude and season. The reddishspring plumage begins to appear on the head and neck of the maleat about the time that bare ground begins to appear in its summerhome, from the first to the middle of April in Alaska. The femalesmolt a month or more later. Dr. Jonathan Dwight (1900) says:Females may now be distinguished with certainty from males for the firsttime by plumage characters, the barring being coarser and extending to thehead, throat, and breast, the feathers of which in the male are reddish brown,
184 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
chiefly with narrow, dusky terminal bands, and often tipped, on the chinespecially, with white. It should be observed that parti-colored feathersbasally or terminally white, may be assumed at this moult on the internal bor-ders of the sternal baud, just as in juvenal dress, the abdominal wedge, flanks,legs, and feet, retaining as a rule the white feathers of the winter plumage.The white remiges and their coverts are always retained and often much ofthe rest of the wing plumage, the median rows of coverts being the ones re-newed if any are. The tail-coverts may be renewed, but the fourteen blackrectrices remain.Dwight says of the next molt
:
Even before the nuptial dress is fully acquired the postnuptial moult setsin, beginning a little prior to the postjuvenal and resulting in an intermediateplumage partly white and partly reddish brown which may hardly be toldfrom that of young birds at the same season. It should be observed thatthe moult of the remiges now includes the two distal primaries which are re-tained in young birds. Adults, however, seem to be somewhat grayer withfiner mottling or vermiculation, the throat being of a deeper red-brown withless barring than that of young birds. Practically young and old, both malesand females, are all indistinguishable except by inconstant differences whenclothed by the preliminary winter dress, but their age and sex may usuallybe told by the left over tell-tale feathers of an earlier plumage.A supplementary molt early in fall, September and October inAlaska, completes the change into adult winter plumage. Femalesare indistinguishable from first winter birds, the feathers of thecrown being basally gray, whereas in the adult male these are basallyblack. Winter adults in high plumage often have a decidedly rosytint, which soon fades in the dry skin.Food.?During summer ptarmigan feed on the tender leaves andflower buds of the willows, birches, and alders, with a fair per-centage of berries, such as mountain cranberry, crowberry, blue-berries, arbutus, and kinnikinnick. They also eat what insects theycan find. Turner (1886) writes:During the winter these birds subsist on the past year's twigs of the willowand alder or other bushes. I have cut open the crops of many of these winter-killed birds and found them to contain only pieces of twigs about one-thirdof an inch long, or just about the width of the gape of the posterior, hornypart of the bill, as though this has been the means of measurement in cuttingthem off. The flesh at this time is dry and of a peculiar taste. In the springthe Ptarmigans congregate in great numbers on the willow-bushes and eatthe tender, swelling buds. The flesh then acquires a bitter, but not unpleasant,taste. As open weather advances they find berries that have remained frozenthe entire winter, and tender grass shoots, and later, insects. The young areinsectivorous to a great degree in their youngest days. They consume greatnumbers of spiders that are to be found on the warm hillsides.Dixon (1927) sent the stomach of a 5-day-old chick to Washingtonfor examination; it contained 17 yellow caterpillars, 1 spider, 15
WILLOW PTARMIGAN 1852'hysanura, and 15 other insects and larvae, but no vegetable matter.He says:The eight chicks foraged in a loose flock covering an area about five feet wideand six feet long. They pursue small insects and mosquitoes which they rundown or reach up for and pick off the grass. I watched one chick catch acranefly and after hammering and pecking at it awhile he concluded that itwas too tough, gave it up as a bad job, and left it.Dr. Joseph Grinnell (1900) writes:Occasionally a few spruce needles were also found. The gizzards of thebirds obtained, invariably contained a quantity of small polished pieces ofclear quartz, this probably being the hardest substance for the purpose ob-tainable by the birds. A bare place on a sand-bar in the river, kept clear ofsnow by the wind, was wont to be frequently visited by the ptarmigan andI have seen them scratching over the gravel in such places, even in the coldestmid-winter weather.In addition to many of the above items, E. A. Preble (1903)lists mushrooms, tops and seeds of grasses, and leaves, seeds, andberries of various other plants.Behavior.?Ptarmigan rise from the ground with a loud whirof wings and harsh cackling notes. When scattered about feedingamong the willows they do not all rise at once but jump up singlyor a few at a time, and there are generally a few laggards. Whenwell under way their flight is strong, swift, direct, and often pro-longed for a long distance. They alight readily on trees and bushes,where they are skillful at balancing. In winter it is difficult tosee them on the snow-laden branches, where they look much likeballs of snow. On bright, sunny days their shadows show up plainlyon the snow-covered ground; but on dull, hazy days they are veryhard to see, unless the black bill or eye is in motion. The blacktail is entirely concealed, except in flight, when it serves as a verygood direction mark for other members of the flock to follow. Theyseem to understand the value of protective coloration, for, if theground remains bare after the white plumage is assumed, they arevery shy ; but, after the snow comes, they become very tame. GraceA. Hill (1922) calls attention to the fact that, while the ptarmiganare molting into the white winter plumage, they frequent the opentundra at the time the cottongrass {Eriophoruvi polystachion) isbearing its white cottony plumes, which aid the birds in theirprotective coloration.Ptarmigan are evidently monogamous and make quite devotedcouples. Dixon (1927) tells of a male, perched on the top of aspruce, standing guard over his mate while she was feeding. Mr.Dixon writes:The bird gave a couple of warning calls as I approached the tree, and then itdawned on me that he was probably standing guard while his nesting mate
186 BULLETIN 162, UNITED STATES NATIONAL MUSEUMfed. So I hunted around ; and sure enough, I found the female ptarmiganfeeding in some dwarf willows about twenty feet from where I stood. As soonas I started after the female the male ptarmigan flew down from the tree topand ran off ahead of me, trying in various ways to decoy me away from his mate.I had been told of an instance where a cock willow ptarmigan had attackedand routed a large grizzly bear tbat happened to stumble upon his nest. Buteven after seeing the ptarmigan drive off the gulls I did not fully appreciatethe furiousness of the attack until June 23, when I came across an old henptarmigan with her brood of small young which were just able to fly. I rushedafter the young, trying to catch one. Just as I was about to grab a chick, awillow bush in front of me exploded and the crick ptarmigan flew directlyinto my face, knocking my glasses to one side as he slapped my face with hisbeating wings. He then dropped to the ground, but instead of retreating flewdirectly into my lace again ; but this time I was ready for him and caughthim with my bare hand when he became mixed up with my mosquito headnet. The bird then tried to bite and to flap his way to freedom. As I startedoff with the cock under my arm the hen ptarmigan left her young and camerushing at me and then crawled feebly about at my feet as though in mortalagony. When I started away she rushed frantically about flapping my heelswith her wings at every step. Every time she rushed at me she hissed. Whenthe male found he could not escape he uttered a few croaking notes and thehen left me at once and went back to her chicks.Voice.?Dixon (1927) says that the warning cackle of the male
" sounds like running a nail over a stiff comb." He mentions threenotes of the female : A harsh ke-ouk, ke-ouk is a warning of danger
;
a soft purring keer-er-eerk is a hush-a-bye, when hovering young;and a clucking cuck-cuck is a note used to call the chicks to her.The downy chicks give a soft cheep-cheep-cheep when in distress orwhen separated.Enemies.?Gulls, jaegers, hawks, owls, foxes, wolverenes, and otherpredatory birds and animals levy heavy toll on the ptarmigan andtheir eggs and young. Ptarmigan are so plentiful that they furnishthe principal food supply for many of these creatures, as well as forthe human natives. Dixon (1927) writes:After the young ptarmigan are out of the shell they are menaced by Black-billed Magpies as well as by the foxes. Thus on June 24 a family of fouryoung and two adult magpies was found systematically working the willowsin the Savage River bottom for ptarmigan chicks. When these magpies locateda pair of adult ptarmigan they would retire stealthily and hide in the willowsnear by, until the ptarmigan chicks began to run about. Then the magpiesswooped down and grabbed the chicks before they could hide, and then car-ried them off and ate them. A cock ptarmigan that I watched put one magpieto flight, but where there were six and in another case nine magpies workingtogether against two adult ptarmigan the odds were overwhelming. As aresult of this persecution by the magpies we found that by July 10 many fam-ilies of young ptarmigan had been reduced to only one or two individuals.Gyrfalcons also levy continuous toll on ptarmigan ; and since these large falconsare relatively numerous in the Mount McKinley district, the aggregate numberof ptarmigan killed by them is considerable. It is thus easy to see why the henptarmigan lays from 6 to 12 eggs. If only one or two eggs were laid eachseason the species would soon become extinct.
WILLOW PTAEMIGAN 187Game.?Were its haunts not so far removed from the centers ofcivilization, the willow ptarmigan would be a popular game bird.Our experience with the Aleutian ptarmigan taught us that thesebirds possess excellent game qualities. Except during the breedingseason, when they are very tame, they are wild and wary enoughto give good sport. Their flight is strong and swift and sufficientlyprolonged to give one all the exercise he wants. Their thick winterplumage is somewhat shot resisting, so that they have to be hard hit,with a close-shooting gun; it often requires considerable chasing toget within effective range. Edwyn Sandys (1904) gives a thrillingaccount of a winter ptarmigan hunt, with its hardships and dangers.The flesh of the old birds in winter is apt to be dark and dry and tohave a bitter flavor, as a result of a steady diet of willow buds, butthat of 37oung birds in fall, fed on fresh foliage, berries, and insects,is lighter colored and very good to eat.Dr. E. W. Nelson (1887) writes:Among the Alaskan natives, both Eskimo and Indian, especially those inthe northern two-thirds of the Territory, this bird is one of the most importantsources of food supply, and through the entire winter it is snared and shot ingreat abundance, and many times it is the only defense they possess againstthe ever-recurring periods of scarcity and famine.The Eskimo of the Kaviak Peninsula have a curious way of taking advantageof the peculiarities of this bird in their migrating season. Taking a longand medium fine-meshed fishing-net they spread it by fastening cross-pieces toit at certain distances ; then taking their places just at sunset in early Novem-ber or the last of October, on a low open valley or " swale," extending northand south, they stretch the net across the middle of this highway, with a manand sometimes two at each cross-piece, while the women and children concealthemselves behind the neighboring clumps of bushes. As twilight advances thenot is raised and held upright. Ere long the flocks of Ptarmigan are seen ap-proaching skimming along close to the snow-covered earth in the dim twilight,and a moment later, as the first birds come in contact with the obstacle, themen press the net down upon the snow sometimes securing 50 to 60 birds.While the men throw themselves upon the net and hold it down, the womenand children rush forward and kill the birds by wringing their necks or bybiting their heads. On some evenings several flocks are thus intercepted, andthe party of natives return to their houses heavily laden with spoils. In winterthe birds are snared in their haunts by placing fine nooses attached to lowbushes close to the ground. Sometimes small brush fences are built withsnares at the passage-ways purposely left open. In spring, as the snow beginsto leave the mossy knolls here and there, the natives shoot a male bird andstuff it roughly with straw, and, mounting it on a small stake, place this effigyupon one of the bare knolls in a conspicuous position ; then they surround itwith a fine sinew net held in place by slender stakes. The hunter then con-ceals himself close by and imitates the challenge cry of the male. All aroundcan be heard the loud cries of the pugnacious birds, and attracted by the decoynotes of the native some of them are almost certain to bestow their attentionupon the decoy ; they approach swiftly, and either fly directly at their supposedrival or alight and run at him in blind rage. In either case their jealously isfatal, as they are at once hopelessly entangled in the net <>f the hunter, who
188 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMdisposes of them and repeats the maneuver indefinitely, generally returninghome well laden.Ptarmigan are subject to great fluctuation in numbers from yearto year, and during periods of scarcity they may be nearly or quiteabsent from regions where they were once abundant. Alfred M.Bailey (1926) says:In 1919, ptarmigan were very scarce throughout the territory ; in December,on a trip to within a short distance of the source of the Copper River, we sawbut one bird. In 1920 the birds began to return, and in 1921 they were reportedabundant at all points where they usually occur. I am unable to explain thecause of this scarcity at intervals, for, so far as I know, no disease has beenreported among them.Mr. Warburton, who spent the summer of 1929 about the mouthof the Yukon, writes to me
:
The scarcity of these birds was a great disappointment. I had expected tofind them as plentiful as I had seen them about Nome and Teller, Alaska, inthe summers of 1924 and 1925. At that time on the Seward Peninsula theywere most plentiful, together with many rock ptarmigan. This year the exactreverse was true, at least about the Yukon mouth.Fall.?Turner (1886) says that the ptarmigan migrate to the in-terior late in the fall. He writes
:
When the snow has pretty well covered the ground in late November thePtarmigans assemble in immense flocks, often numbering thousands. I wasonce out on the higher grounds just south of the Crooked Canal. I ascended aslight hill and came, unexpectedly, on one of these large flocks that covered acresDf ground. I was among them before either was aware of it. They flew, andmade both the air and earth tremble. There must have been over five thousandbirds in this one flock. They flew beyond a neigbboring hill-range. Approach-ing night and a heavy snow falling prevented me from following them.Winter.?When the snow covers the tundra so deeply that no foodor shelter can be found there, the ptarmigan are forced to migrateinto the interior valleys, river bottoms, and creek beds, where theycan find shelter among the willows, alders, or spruces and can feedon the bugs and twigs of these trees or such berries and fruit asstill remain above the snow. They often congregate at that season,and, particularly when migrating, in enormous numbers. Mr.Brandt says that, where the railroad crosses the Continental Dividein Alaska, he saw large flocks arising " like snow drifts in motion,alongside the snorting engines, and whirling away over the greatwhite hills."In winter the ptarmigan's feet are thickly covered with long,hairlike feathers, resembling the foot of a hare, which serve as snow-Bhoes and enable the bird to walk on soft snow. Sandys (1904)Writes
:
And Nature, as if realizing the perils of the ptarmigan asleep, has taught itto plunge beneath the cold drifts to escape the cold, and to fly at, not walk t?.
WILLOW PTARMIGAN 189the chosen drift, so that there will be no telltale trail for some keen nose tofollow to the sleeping-place. And this the bird invariably does, going at speedand butting its way into the snow, leaving never a print to betray its re-treat, from which it flies in the morning. The game of life and death is in-terestingly played up North?where the weak white snow-shoers are everhiding from the strong white snow-shoers forever searching over a field ofbaffling ice-bound white.Doctor Nelson (1887) describes another method of roosting:On November 25, 1877, they were numerous, in large and small flocks, alongthe bushy gullies and hill slopes on the shore of Norton Sound, but were shy.In many places where they had stopped the night before, their sleeping-placewas well marked. In each instance they had occupied a small clear spot inthe midst of a dense thicket, and in no case had the birds approached on foot,but had flown in over the top and plumped down into the soft snow, wherethey had remained during the night, each bird thus making a mold of itselfin the snow. In some instances there were fifteen to twenty of these molds inthe snow in an area of a few feet. In leaving their stopping-place the birds aroseand flew directly from their " forms," as was shown by the marks of the wingson each side as they touched the snow in rising, so there were no tell-tale tracksto or from these places ; the open places were undoubtedly chosen to allow thebirds an unobstructed escape in case they were surprised by prowling foxes,which hunt these thickets for food.DISTRIBUTIONRange.?The range of the willow ptarmigan as a species is circum-polar, extending in the Old World from Scandinavia, Russia, andSiberia south to Turkestan, and in North America from Greenland,Newfoundland, the Arctic Archipelago, and Alaska south throughCanada. The species is casual in winter in the Northern UnitedStates.Breeding range.?The North American breeding range extendsnorth, to Alaska (Cape Lisburne, Wainwright, Point Barrow, SmithBay, Delta of Colville River, Camden 33av, Humphrey Point, andDemarcation Point) ; Franklin (Bay of Mercy, Port Kennedy, FelixHarbor, and Igloolik) ; northern Quebec (Fort Chimo) ; and Labrador(Okkak). East to Labrador (Hamilton Inlet) and Newfoundland(Raleigh and St. Johns). South to Newfoundland (St. Johns andsouth coast) ; west-central Quebec (Carey Island) ; northern Ontario(40 miles south of Cape Henrietta Maria) ; northern Manitoba(50 miles north of York Factory and Fort Churchill) ; southernMackenzie (Artillery Lake and Fort Resolution) ; central BritishColumbia (Moose Pass, Icha Mountains, and Ninemile Mountain) ;and southeastern Alaska (San Juan Island). West to Alaska (SanJuan Island, Kruzof Island, Glacier Bay, Nushagak, Nelson Island,Igiak Bay, Askinuk Range, Pastolik, St. Michael, Nome, Mint River,Cape Blossom, and Cape Lisburne).
190 BULlLETIN 162, UNITED STATES NATIONAL MUSEUMWinter range.?Although willow ptarmigan perform a very defi-nite migration, individual birds are frequently found in winter almostat the northern limits of the breeding range. At this season theyhave been recorded north to Alaska (Nunivak Island, Nulato, KutukRiver, and Miller Creek) ; Mackenzie (mouth of the Dease River,Fort Eae, and Fort Reliance) ; northern Manitoba (Fort du Bro-chet) ; Franklin (Igloolik) ; Quebec (Great Whale River, and Pi-ashti) ; and Newfoundland. Normally they are found south tosouthern Quebec (Lake St. John) ; central Ontario (east of Cochraneand Martin Falls) ; central Manitoba (Norway House and GrandRapids) ; Saskatchewan (Cumberland House and Fort Carleton)
;
and southwestern Mackenzie (Fort Simpson).Migration.?Though the bulk of the willow ptarmigan move southeach fall or winter, and return north in spring, the movement has noregularity and is directly correlated with the food supply and, inconsequence, with the fall of snow.First fall arrivals are: Quebec, Lake Mistassini, October 25, andGodbout, November 2; Ontario, Martin Falls, about October 20;Manitoba, Lac du Brochet, November 4, Grand Rapids, November 12,and Winnipeg, January 12; Mackenzie, Fort Rae, October 1; andAlberta, Fort Chippewyan, October 11.In spring, late departures for the north are : Quebec, Lake Mistas-sini, about May 1 ; Ontario, Cochrane, March 20 ; Manitoba, Winni-peg, March 21; and southwestern Mackenzie, Fort Simpson, March12. They have been observed to arrive at Fort Resolution, Macken-zie, on June 28, and at Demarcation Point, Alaska, as early as April 6.Casual records.?The willow ptarmigan has been reported a fewtimes in the Northern United States. Among these records are:Maine, one at Kenduskeag, April 23, 1892 ; New York, one at Watson,May 22, 1876 ; Wisconsin, two taken near Racine in December, 1846Minnesota, one taken at S*?4y Island, Lake of the Woods, April 20,1914; and Montana, three taken in Glacier National Park in thewinter of 1913-14. Prof. W. B. Barrows believed that they occasion-ally occurred on Keweenaw Point, Mich.The range as above described is for the entire species, which in the1931 edition of the American Ornithologists 1 Union Check List is sub-divided into five subspecies. According to this authority the sub-species albus is found in North America from the eastern AleutianIslands, northern Mackenzie, northern Banks Island, and centralGreenland south to central British Columbia, central Alberta, cen-tral Ontario, and southern Quebec. The Ungava ptarmigan (L. I.ungavus) occurs from northern Quebec west probably to the easternshore of Hudson Bay. Allen's ptarmigan (L. I. alleni) is confined toNewfoundland. The Alaska ptarmigan (L. I. alascen-sis) is found
allen's ptarmigan 191on the Alaskan mainland (except the southeastern coast), northernYukon, and eastward for a distance not yet determined. Alexander'sptarmigan (L. I. alexandrae-) occurs on Baranof Island, Alaska, andadjacent islands, west to the Shumagin Islands, and south to PorcherIsland. This race also may occupy a narrow strip on the mainlandfrom Glacier Bay to central British Columbia.Egg dates.?Northern Alaska : 68 records, May 25 to July 10
;
34 records, June 6 to 25. Arctic Canada: 37 records, June 2 toJuly 7; 19 records, June 10 to 21.Labrador Peninsula : 18 records, June 1 to 30 ; 9 records, June6 to 23.Newfoundland (alleni) : 11 records, May 12 to June 30; 6 records,June 8 to 12.Southern Alaska and British Columbia {alexandrae) : 3 records,May 28, June 25 and 26.LAGOPUS LAGOPUS ALLENI StejnegerALLEN'S PTARMIGANHABITSThe willow ptarmigan of Newfoundland was originally describedby Dr. Leonhard Stejneger (1884) as similar to the common willowptarmigan, " but distinguished by having the shafts of both primar-ies and secondaries black, and by having the wing-feathers, even someof the coverts, marked and mottled with blackish." He examinedonly 14 specimens in all, all of which presented the above characters.In a large series (I have 75 in my own collection and have examineda great many in other collections) these characters appear to benone too constant in Newfoundland birds and to crop out occasion-ally in willow ptarmigan elsewhere, with a great range of individualvariation. In this connection it is interesting to note what HarryS. Swarth (1924) has to saj' about the willow ptarmigan of southernAlaska
:
It is of interest to note in alexandrae the frequent presence of black shaftson the primaries, sometimes on secondaries and greater coverts. This characterhas been considered an important feature of the Newfoundland subspecies(L. I. alleni), but obviously it can not be used as a feature characteristic of thatrace alone. In an immature female from Prince of Wales Island, which hasacquired the winter flight feathers, not only are primaries and secondarieswith distinct black shafts, but there are large, tear-shaped spots of black nearthe tips of all the primaries and most of the secondaries. Furthermore, theprimaries have a black " freckling " over much of their surface, and the greatercoverts are also marked with black, though to a lesser degree.Dr. Hart Merriam (1885) has called attention to the great varia-tion in the extent of black in a large series of wings sent to him
192 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
b}?- Napoleon A. Comeau from Godbout, Quebec. Perhaps the rangeof alleni should be extended to include eastern Quebec.Allen's ptarmigan are very abundant and widely distributed inNewfoundland, on the upland tundra in the central and northernparts of the island and on the mountains. J. It. Whitaker tellsme that from about the middle of April to September 20 they liveon the high tundras and rocky ridges; he has never seen a bird onthe lower ground during summer. F. S. Hersey, who went to New-foundland in September and October, 1913, to collect ptarmigan forme, found these birds more abundant than he ever found willowptarmigan anywhere in northern Alaska. He found them on thehills a few miles back from Cape Ray, and some were seen in theLewis Hills near the west coast ; but they were most numerous aboutGafftopsail in the interior of the island. This is a rough, barren,open region, tending to run into low, rocky ridges, separated bylower marshy areas, carpeted with thick mosses, often knee-deep,and dotted with small ponds. There is a little cover here, althougha few sheltered spots may have low shrubs and nearly prostrateblueberry bushes, and in a few places there are small areas ofexceedingly tough, thick, dwarf spruces.Courtship.?Referring to the courtship of this ptarmigan, Mr.Whitaker says in his notes
:
The cock bird nearly always mounts a rock to utter his challenge; havinggained this vantage point he looks all around, then with head erect and breastexpanded bleats out his notes ; they will start a little before sunrise andcontinue for a short time and are seldom heard during the day. After theyoung are hatched crowing appears to cease entirely. The crow of this birdis very like the British red grouse ; it is quite easy to imitate, but to me, verydifficult to write. The first part sounds like the low trembling bleat of ananny goat, or, if one can imagine it, a very coarse drum of a snipe ; thereare usually five quickly uttered tremulous notes run off er-er-cr-cr, followedby three or four gobcck-gobeek^gobeck ; these latter may be copied perfectlyby partly closing your throat and uttering gobeck harshly through the nose.Nesting.?Whitaker says thatnesting begins in May and the young are usually full grown by the end ofAugust. The nest is usually placed at the base of some bowlder and is wellhidden ; the hen bird sits very close indeed. Some years ago a nest wasfound close to a watershoot by the railway; several trains a week stoppedhere for water ; the train hands used to take passengers to see the birdsitting, which did not upset her in the least.A set of eight fresh eggs, collected for me near Gafftopsail onJune 16, 1912, was taken from a hollow in the tundra moss, undera little bush, but in plain sight, between two hummocks. Mr. Hersey
allen's ptarmigan 193found a nest from which the young had hatched, which he describesin his notes, as follows
:
The site was on the side of a ridge some 20 feet above the marshy groundat its base. A few bushes grew at this point, and under the overhangingbranch of one of these the nest had been made. Even at this season the hollowthat had been the nest was well defined. There was no indication of liningbeyond the accumulation of fine dry leaves, bits of sticks, moss, and othervegetation that made up the general ground cover on the whole ridge. Theeggshells from which the young had hatched were still in the nest.Eggs.?The eggs of Allen's ptarmigan are indistinguishable fromthose of its mainland relative. The measurements of 54 eggs aver-age 42.5 by 33 millimeters; the eggs showing the four extremesmeasure 45.6 by 32.3, 44.1 by 37.6, 40.1 by 30.2, and 43.4 by 28.7millimeters.Plumages.?The molts and plumages of this race are similar tothose of the willow ptarmigan. It is interesting to note, however,that many of the wings sent by Mr. Comeau to Doctor Merriam(1885) taken from birds killed in November, were "deeply tingedwith a delicate and ver}^ beautiful shade of rose-pink which is morepronounced than in a freshly killed Roseate Tern." Mr. Whitakerhas also noted this, as he says in his notes : " They are in full whitewinter plumage by the end of October; I say white, but there is abeautiful faint pink flush; I first noticed this when having shotseveral one winter day and laid them on a block of pale blue ice,this showed up the pink to advantage ; the pink quickly fades afterdeath."Food.?In his notes on food, Whitaker writes
:
As the snow melts off the high ground many berries that have lain snugall winter provide plenty of food; after these are shriveled up by the sunand before the new crop matures, the birds feed on the tips of a low-growingplant, which looks like a very dwarf heather. In August the blueberries andpartridgeberries (Mitchella rcpens) ripen and on these they feed almost ex-clusively. When the snow begins to pile up on the highlands during Octoberthe bulk of the birds move down to lower levels and begin feeding principallyon buds of scrub birch ; they also eat buds of blueberry and pussy willow.The birds shot by Mr. Kersey had their crops filled with blueberriesand their leaves, the small green berries from a dwarf evergreenresembling a cedar, or with small white seeds.Behavior.?Mr. Whitaker says in his notes
:
These ptarmigan are very tame in the wilder districts, where they are notdisturbed or shot, and seem loath to take wing ; they will run in front of youstopping every few yards and spread their tails with a quick jerky motion
;
they often utter a suppressed little grunting note, possibly a protest againstyour intrusion in their midst.
194 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMMr. Hersey's notes contain the following observations
:
The birds were usually found in parties of 3 or 4 to as many as 11 and 13and, as many were in a plumage that indicated they were birds of the year,it was assumed that these flocks usually represented a pair and their grown-upyoung. They were very erratic in the matter of taking flight, on some occa-sions not flying until nearly stepped on and again taking wing before a personwas within 100 yards of them. As a general thing they were wildest on sharpfrosty days or when rather windy and would lie closest when the day waswarm and calm. I do not believe the ptarmigan range over any large areaat this season. Birds were seen day after day on the same ridges and werebelieved to be the same flocks. On bright clear days a flock often spends severalhours crouched on the ground in the sunlight, particularly in rocky places,where one or two males will take up a position on the rocks. These sentinelsperform their duty poorly, as it is usually rather easy to approach and shootsuch birds and then secure one or more from the flock with the remainingbarrel as the birds take wing. At times the birds on the ground will notflush, even after one or more has been shot. I remember one very warm daywhen I came upon three birds crouched on the ground a few feet apart. Iwas unable to flush any of them and was obliged to back away and shootthem on the ground one at a time, the remaining birds paying no attentionto the sound of the discharge. When flushed they are frequently silent, butsome birds give voice to a harsh cack-cack-cack-cack-cack. On foggy morningsbirds, presumably males, may be heard " crowing," at least the notes soundvery similar to the spring call of the male. They may be a signal or flock call.Game.?Allen's ptarmigan, known also as " willow grouse " or
" partridge," is by far the most important game bird in Newfound-land. It is jealously guarded during the close season, only to be shotin enormous numbers during the open season. Many parties of gun-ners go out every fall from St. Johns, as well as visiting sportsmenfrom Canada and the United States, camp along the line of the rail-way, and kill hundreds of the birds. Fortunately for the birds,there are immense tracts of uninhabited country, inaccessible bybranch lines or by roads, where they are not disturbed, so that theydo not seem to be in any immediate danger of extermination.Winter.?Mr. Whitaker says that " quite a number perish duringthe winter; they make a shallow scratching in the snow in which toroost and are frequently buried by drifts and are imprisoned; Ihave often found them dead in the spring when the snows are melt-ing, and once saw seven within a few feet of each other that had metthis fate." LAGOPUS LAGOPUS ALEXANDRAE GrinnellALEXANDER'S PTARMIGANHABITSThe willow ptarmigan of the humid coast region of southernAlaska and northern British Columbia is darker and more richlycolored than birds from northern Alaska and Labrador, with a
Alexander's ptarmigan 195smaller and narrower bill, the general color above being deep, richchestnut thickly vermiculated with black. The most typical birdsof this race are found on the islands along the coast of southernAlaska; the type came from Baranof Island. It was described byDr. Joseph Grinnell (1909) and was named in honor of Miss AnnieM. Alexander. Alfred M. Bailey (1927) first saw them on Wil-loughby Island in Glacier Bay, of which he says
:
This island is about five miles in length, with scant vegetation, other thanalder and stunted spruce. The knobs are devoid of plant life, and it is only onthe terraces that soil can hold. The southern and eastern slopes are ratherdensely clothed with alders, however, and form ideal cover for Ptarmigan.Seven other birds were seen by Young, one of which was still in the winterwhite. The pair collected were breeding birds.He suggests a good reason why they should prefer to live on theislands
:
The lack of predatory animals was very noticeable, for with the exceptionof two Eagles, none was noted, and no signs of predatory animals were seenon the outer Beardslee Island, although those nearer shore must have somecarnivorous mammals. We saw a fox in June and August on the mainlandshore, and wolves are abundant. It is rather apparent then that the birds arefree from molestation, in direct contrast to their life upon the mountain slopesof the mainland. Ravaged by fur-bearers, it is possible the Ptarmigan first usedthe islands for protection, and having found both food and comparative safety,have continued to live under such conditions. On the other hand, the islandsmight be considered as being alpine in nature, with timber-line conditions, asthe spruce are small and willow and alder predominate, with the characteristicprofusion of small growths. The soil is scant, the glacial sands and morainedebris being exposed, while the windward shores of the outer Beardslee are pre-cipitous; the glacial winds sweep down channel, icebergs line the shores, andtaking all into consideration, the region is probably the coldest of south-eastern Alaska.Harry S. Swarth (1924) noted that the birds he collected in theSkeena River region of northern British Columbia " are intermediatein color between lagopus of the interior and alexandrae from theislands; the average is nearer alexandrae." He says of their hauntsthere
:
Ptarmigan are said to occur occasionally in the lowlands of the Hazeltonregion in midwinter, but during most of the year they are restricted to theAlpine-Arctic mountain tops. We found them in limited numbers on the timber-less summit of Nine-mile Mountain. There are miles of open country on thetwo converging ridges that form the top of this mountain, barren of trees savefor occasional thickets of dwarfed or prostrate Alpine conifers, and here, at longintervals, we encountered ptarmigan.Nesting.?We know very little about the nesting habits of Alexan-der's Ptarmigan. According to Doctor Grinnell (1909), JosephDixon " records that at Coppermine Cove, Glacier Bay, July 10 to 20,the feathers and bones of a ptarmigan were found near a nest ofbroken eggs on the summit of the mountain, 2,100 feet. The nest was
196 BULLETIN 162, UNITED STATES NATIONAL MUSEUMunder a stunted hemlock. All the feathers were white, so the ptar-migan must have laid early."There is a set of 11 eggs in Col. John E. Thayer's collection, takenby John Koren on Kodiak Island, Alaska, on June 25, 1911. Theeggs were fresh and " were placed on a bed of moss in a two-footgroove in an elevated part of the tundra "; both parents were present.The eggs are not distinguishable from those of the willow ptarmigan.The measurements of 20 eggs average 42.8 by 30.9 millimeters; theeggs showing the four extremes measure 45.9 by 31.5, 43.6 by 32,41.5 by 30.8, and 42 by 30.2 millimeters.Young.?Swarth (1924) writes:In all, ten broods of willow and rock ptarmigan were encountered (the specieswere not always to be differentiated) and about five or six single birds in addi-tion. The broods ranged from three to twelve in number ; the aggregate of youngbirds seen was about fifty. The chicks grew rapidly. Some seen on July 25,and a day or two later, were down-covered and unable to fly. At that timethey were accompanied by the female parent only, and the male birds wereflushed separately. By August 10 the young ptarmigan were the size of quailand larger, and were strong on the wing. The old males were then associatedwith the families. In some of the larger broods seen the difference in sizeamong the young was so marked as to suggest the junction of two families. Itmight happen that upon the death of a hen her offspring would seek the com-panionship of another family.Food.?These ptarmigan probably feed on as varied a diet as otherptarmigan, but the following note by Alfred M. Bailey (1927) is allthat I can find in print on the subject
:
A flock was flushed from a bed of wild strawberries, at an altitude of scarcelythirty feet, when I had expected to find them above timber line. There wereseveral pairs of adults, as well as many young, and a good series was taken.An examination of crop contents proved the birds had been feeding eutirely onstrawberries and pea-vine, no alder or willow buds being found. After findingptarmigan in such a low altitude, I searched all the points along the east main-land shore, and did not fail once to find them, where there were berries. Itwas noticeable they preferred the points where they could feed close to afringe of alder.Behavior.?Mr. Bailey (1927) says also:I returned to Glacier Bay again from October 10-14, and observed the Ptar-migan under still different conditions. A stop was first made on the outerBeardslee, upon which the young bird had been taken. I was interested to seeif the Ptarmigan had left the island, as it was now drab and dry looking, andthe birds were assuming their winter's white. I found them very abundant, andover forty were flushed from the dense alders, and a few taken. They wereextremely wild, as they should have been, for their changing plumage?entirelywhite below with many white feathers in head and neck, made them extremelyconspicuous among the leafless alders. I had little chance to observe them, du<*to their wildness. Stops were made at two other points, not visited in August,and ptarmigan were noted in both places. At Sandy Cove, October 14, I col-lected another series from the point they were first found in August. These
UNGAVA PTARMIGAN 197
were as conspicuous as the others, but when the flock was broken up, the birdsdid not flush so readily, and I had a chance to study them at close range. Theyran over the ground the same as in August, evidently believing themselvesinconspicuous, and when closely pressed, crouched quail-like and depended upontheir " protective coloration." When the birds became scattered several climbedinto alders, about a foot from the ground, and sat hunched in some convenientcrotch, where they were more evident than ever. I was surprised to find straw-berries still abundant here, hidden under the mosses in shady places, and eachof the specimens taken had its crop full.Doctor Grinnell (1909) quotes from Chase LittleJohn's notebookas follows
:
While searching for eggs of the glaucous-winged gull on one of the smallislands on the east side of Glacier Bay on July 14, I suddenly came upon aflock of ptarmigan in a little opening among some spruce, hemlock, andalders, which covered the ground in dense masses in spots; the remainderof the area supported a thick growth of grass interspersed with patches ofmoss and low-growing flowering plants. There were about eighteen birdsall told, young and old, and as near as I could determine there were fouror five old birds present. They would not fly after they were first flushed, butkept dodging about on the ground, sheltered by the thick cover ; several timesI saw them, but so near that a shot would have ruined them as specimens.LAGOPUS LAGOPUS UNGAVUS RileyUNGAVA PTARMIGANHABITSThe ungavus race of the willow ptarmigan was described byJ. H. Kiley (1911) as " like Lagopus lagopus albus, but with aheavier bill," based on a series of 20 birds collected at Fort Chimo,Ungava, in the northwestern portion of the Labrador Peninsula.He gives as its probable range " from Ungava and probably theeastern shore of Hudson Bay south." Birds that I have seen fromthe eastern and southern parts of the Labrador Peninsula do notseem to have appreciably large bills. If this form is worthy ofrecognition at all, which I very much doubt, it will probably proveto be a northern race and perhaps identical with alascensis Swarth,about which I have expressed my views elsewhere. But, pendingfurther investigation, we may as well consider all the willow ptar-migan of the Labrador Peninsula as referable to this form.In this connection it might be well to consider what Lucien M.Turner has to say, in his unpublished notes, concerning the greatdegree of individual variation that he noted in the hundreds ofUngava ptarmigan that he handled. He found a great " individualvariation in the size and shape of the head and beak." He collecteda large series of skulls and found them to vary from " 1.94 to 2.30inches in length from occiput to tip of bill." He says that " the74564?32 14
198 BULLETIN 162, UNITED STATES NATIONAL MUSEUMlarge beak is especially noticeable in some of the birds, more espe-cially in the males," but that " some only of the males alonehave a noticeably large beak." He concludes by saying that thischaracter should not, in his opinion, warrant the separation ofthis form.I found and collected a few specimens of willow ptarmigan nearHopedale and at Ukjuktok Bay, on the northeast coast of Labrador,in 1912. They were breeding there, and I was told that they alsobreed on some of the islands. They are said to migrate farther southin October and return in April and May. I have a set of eggs takennear Okkak in July. Dr. Harrison F. Lewis (1928) said that thisspecies " bred in the summer of 1928 on the large island at the BluffHarbor where they had nested every year since 1925. It seems prob-able that two pairs of willow ptarmigan nested on this islandin 1928."O. J. Murie has sent me the following interesting notes:Along the Nastopoka River, on the shore of Hudson Bay, lies a region thatis apparently an ideal nesting ground, for willow ptarmigan. On the morningof May 13, my Eskimo guide led me inland over the granite hills ahoutparallel with the river. The immediate coast iu this vicinity is bare and tree-less. Ten or 15 miles inland we found scattered patches of small stuntedspruces, covering some of the lower hills. Here we found a ptarmigan para-dise. The birds were everywhere, apparently all paired, the males thrilledwith the energy of the mating season. It was invariably the male that flewout first when we approached small clumps of willows. He went off like arocket, with a clattering racket of harsh notes and whir of wings. He wouldgenerally take a commanding position on a rocky point or top of a hill and,strutting pompously with head high and tail raised and spread, let out hisclattering crow.At this time of the year I was struck by the perfect blending of the plum-age with the surroundings, especially in the case of the female. Her plum-age at this time is speckled irregularly with various shades of brown andwhite, in varying stages of molt, which was difficult to distinguish on a groundof mosses, lichens, rocks, and willows, sprinkled with remnants of snow. Thefemales showed a tendency to keep well under cover. The males were not par-ticularly difficult to see, and I spied them readily at a considerable distance onoccasion, but sometimes I thought I could detect a principle of protective color-ing. The body was pure white, with a brown head and neck, apparently theusual plumage in the season of courtship, judged from this one spring. Usuallythe white plumage flashed out prominently, but the brown neck was lust on thebackground of rock and moss. Although the eye was attracted at once, therewas a tendency to pass it over without recognizing the headless body as abird. Similarly, on a snowbank, the brown neck was seemingly detached, andnot a part of a bird.Nesting.?Turner says in his notes
:
In the vicinity of Fort Chimo, nesting of this species begins during the lat-ter part of May. The nest is usually placed in a dry spot among the swamps oron the hillsides where straggling bushes grow. The nest is merely a depres-sion in the mosses and contains a few blades and stalks of grass together
UNGAVA PTARMIGAN 199
with a few feathers from the parent bird, which is now in the height of themolt from the winter to the pre-aestival plumage. The first eggs obtained weretwo on June 1, 1S84.Young.?Referring to the period of incubation, Turner writes:It is a rather difficult matter to determine, as the female is compelled,during a stress of severe weather, to sit upon her eggs to prevent thembeing lost by cold or rain. It is not unusual for severe snow and sleet accom-panied by cold rains and even a severe freeze to occur during the early halfof June at Fort Chimo. Some of the most dismal days of the year occur inearly June. The parent bird, during such weather, may be two or three dayson the nest after the first young bird has appeared and thus prevented fromgiving such attention to the young as these tender creatures require. It isnot rare to find a nest containing two or three eggs and near by to find oneor more young which have perished while the mother has perhaps wanderedoff with three or four young which were able to follow her.He says that "the Indians consider the downy young of theptarmigan a special delicacy. Even taken from the shell the birdserves in lieu of an oyster." He frequently saw them eating theembryos taken from eggs that they were blowing. He says thatthey make special excursions to collect the small chicks for food.One party that he saw returning from such a hunt had more than250 of these helpless young.At Ukjuktok Bay, on August 3, 1912, I surprised a family partyof willow ptarmigan in a boggy, grassy hollow. The young, whichwere about half grown, rose with a startling rush of wings andwent whirring off like a flock of quail. The old birds did not flush.The female feigned lameness, in spite of the fact that the young hadall flown; I could not make her fly, and she finally walked away.The male walked boldly out into the open marsh, looking at me,too close to shoot, then ran behind some spruces and flew away tojoin the young. I followed them up and flushed the male first;then three of the young rose singly. I could not find any more ofthe young, but a little later I found both old birds in the exactspot where I had first seen them.Voice.?O. J. Murie has sent me the following good description ofthe ptarmigan's notes:The call of the Ptarmigan is very striking. It consists of a rattling krrr-r-r-ruk-uk-uk-uk-uk, followed by a more deliberate, low-toned, throaty puk-que'-o,puk-que'-o, puk-que'-o. I thought it fitted well with the surrounding hills ofrough granite and the scant growth of ragged, twisted spruces. Sometimesthe female was heard responding with a peculiar whirring sound, a nasalnyek, nyek, somewhat similar to some notes of other members of the grousefamily. By imitating this note we frequently drew the male to us in a head-long flight. He would drop on a knoll near by and send out his startling call.The Eskimos take advantage of this trait and decoy the birds to be shot.Winter.?Ptarmigan are great wanderers in wTinter, but veryerratic in their movements, appearing in enormous numbers dur-
200 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMing some seasons on the eastern and southern coasts of the LabradorPeninsula, and being nearly or quite absent during other seasons.Dr. Harrison F. Lewis writes to me
:
This species often appears on the coast in great numbers in autumn, but thenumbers so appearing vary from year to year. When these birds come outfrom the interior to the coast they seem to come as far out as possible, andoften go in numbers to the very outermost islands which, of course, are oftenunited to the mainland by ice at that time of year. Large numbers are takenby the residents for food. No matter how plentiful the ptarmigan may bein the fall and the month of December, comparatively few are usuallyto be seen after the 1st of January, although a few may often be observedfrom time to time until spring.But vast numbers remain in northern Ungava, at least duringsome winters, according to Turner. One day he saw a gyrfalconflying across the Koksoak River, and writes
:
We had seen hundreds of ptarmigans on the left bank among the thickets.The hawk plunged among these birds which began to rise as soon as thehawk was sighted. I am certain that not less than 1,800 ptarmigans rosebefore that hawk; and, as the latter did not reappear, we suspected there wasat least one less ptarmigan. The air fairly trembled as these birds arose.In my notes I find the following for the locality of Fort Chimo and date ofDecember 7, 1882. Hundreds of this species of ptarmigan have made theirappearance in this vicinity during the past week where two weeks ago not adozen birds of the kind could be found during a tramp of an entire day.The traffic in ptarmigan feathers is very heavy and will give anidea of the enormous number of birds killed. The Indians saveonly the clean feathers from the breasts and backs of the birds,pack them in bags, and trade them to the Hudson's Bay Co. Theyare then packed in barrels and shipped to England. Mr. Turnerestimated that it required the feathers of 16 ptarmigan to make apound and says that 31 barrels of feathers were shipped from FortChimo during the year ending June 1, 1883. As the average weightof feathers in a barrel was 51 pounds, he calculated that it accountedfor 25,296 ptarmigan killed. And to this must be added a verylarge number of birds killed whose feathers were soiled or for someother reason were not saved. Fortunately the ptarmigan is a prolificbreeder; otherwise, the wholesale slaughter of chicks and old birdswould soon exterminate the species.LAGOPUS LAGOPUS ALASCENSIS SwarthALASKA PTARMIGANHABITSHarry S. Swarth (1926) gave the name alascensis to this supposedsubspecies, which he characterized as " slightly larger than albus.A large billed race; bill slightly smaller than in ungavus, much larger
ALASKA PTAKMIGAN 201than in alius. In summer plumage, generally more reddish-coloredthan either ungavus or albus, a difference that is most conspicuous infemales in the barred breeding plumage."He gives as its range " the Alaskan mainland except on the south-eastern coast, northern Yukon Territory (specimens from vicinity ofForty-mile), and eastward for an undetermined distance."At least three authors have attempted to subdivide the willowptarmigans of North America, with decidedly confusing and some-what unsatisfactory results. Austin H. Clark (1910) after studying115 specimens, 20 from Newfoundland, 60 from Labrador, 3 fromcentral arctic North America, 18 from the mainland of Alaska, 2from Kodiak Island, and 12 from the Shumagin Islands, says
:
All those from Labrador and central arctic America, with others fromPoint Barrow, Kotzebue Sound, Cape Lisbourne, Kowak River, Yukon River,and near St. Michaels, belong to a well-differentiated race, with the beak verylarge, high and stout, the culmen strongly arched, and usually with a prominentridge from the inferior corner of the maxilla to in front of the nostril. Theyare identical among themselves, it being impossible to tell from the examinationof any one specimen whether it was taken in Alaska or in Labrador.J. H. Riley (1911) evidently differed from him, for when he de-scribed and named the Ungava bird, from the Labrador Peninsula,he gave as the range of alius " from the west side of Hudson Bay,west through northern Alaska to eastern Siberia." Thus two investi-gators, with practically the same material for study, have arrived atquite different conclusions.Now Swarth (1926) comes along with a still different theory, basedon a study of a large series of western birds in California collections,together with 3 from the west coast of Hudson Bay and 10 from FortChimo, Ungava. He writes:Comparison of these birds with the series in this museum convinced me ofthe existence of the following recognizable subspecies of the willow ptarmiganon the North American mainland: (1) Lagopus lagopus ungavus from theregion east of Hudson Bay, as defined by Riley; (2) Lagopus lagopus aliusfrom the west shore of Hudson Bay westward to the coast, ranges of northernBritish Columbia, and for an undetermined distance northward; (3) an unde-scribed subspecies from the Alaskan mainland and extending for an undeter-mined distance eastward in the extreme north.Swarth (1926) then goes on to describe the northern bird, alascen-sis, as quoted above and says further
:
Conditions in these western races of willow ptarmigan parallel to some extentthose found in the rock ptarmigan. In each species the northern Alaskan sub-species is an extremely ruddy-colored bird compared with the others, and laeach the British Columbia subspecies seems to reach an extreme of grayness.In each species, too, the Labrador birds are much more grayish than are thosefrom Alaska. Thus the Labrador willow ptarmigan (ungavus) and the BritishColumbia bird (albus) are much alike as regards color but differ in size of bill.
202 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMThe Labrador bird and the northern Alaskan bird (alasoensis) are both large-billed forms, but differ in coloration.Perhaps, after studying the three papers referred to above, thereader may get a clear idea of the subject ; but I must confess that Ican not. In my opinion, all the willow ptarmigan of the NorthAmerican mainland (excluding alexandrae and perhaps alleni) areof one subspecies, Lagopus lagopus albus. The differences pointedout above are too slight and too variable to be worthy of recognitionin nomenclature.LAGOPUS RUPESTRIS RUPESTRIS (Gmclin)ROCK PTARMIGANHABITSMuch has yet to be learned about the relationships of the variousforms of the rock ptarmigan in North America and their distribu-tion. There have been three forms named in Greenland, one in New-foundland, three on the mainland, and six on the Aleutian Islands.With the possible exception of some of the Aleutian forms, which arequite distinct, all might well be regarded as subspecies of the Euro-pean Lagopus mutus. Of the three mainland forms Harry S. Swarth(1926) has made an extensive study, the results of which are quiteenlightening. The three forms are: A gray-colored bird, rupestris,a ruddy-colored form, kelloggae, and a dark-colored form, dixonl.He says of their ranges:First, there is a gray-colored bird that extends from Labrador westwardto the coast ranges of northern British Columbia. In the east it apparentlyextends northward into the Arctic regions ; it also occurs on islands north ofMackenzie, but elsewhere in the west it is restricted to the southern part ofthe region covered by the species Lagopus rupestris. Second, there is a ruddy-colored form that occupies almost the entire mainland of Alaska and ex-tends eastward along the Arctic coast about to the one hundredth meridian.Third, there is a dark-colored form with a rather limited range in the coastalregion of southeastern Alaska.The latest contribution on the subject is a paper by P. A. Taverner(1929) based on a careful study of 105 adults of both sexes andyoung. Even with this quantity of material, the results are notwholly satisfactory, because of lack of adequate series of breedingbirds in comparable plumages and because of a number of puzzlingspecimens that seem to have wandered far away from their normalbreeding ranges. Taverner, however, has recognized two wide-ranging mainland forms, a southern, generally grayish bird (rupes-tris), ranging from northern British Columbia to Newfoundland,and a northern, generally yellowish bird (kelloggae) . ranging from
ROCK PTARMIGAN 203the interior of Alaska to northwestern Greenland. I recognize thisrevision and include welchi in the grayish form {mpestris)
.
It is the wide-ranging gray rock ptarmigan that we are now con-sidering. Lucien M. Turner, in his unpublished notes, has given usthe best contribution to the life history of this species. Bendire(1892) quoted some of these notes under Reinhardt's ptarmigan,but, for reasons stated under that form, I prefer to use them here.As to the haunts of these birds he says
:
They prefer more open ground and rarely straggle even into the skirts ofthe wooded tracts. The hilltops and " harrens " (hence often called barrenground bird) are their favorite resorts. As these tracts are more extensivein the northern portions of Labrador and Ungava, these birds are there veryabundant. During the summer months they are quite scarce in the vicinityof Fort Chimo, retiring to the interior and the hills of George River for thatseason. In the month of May the nuptial season arrives and is continued untilabout June, when nesting and laying begin. The birds are by this time scat-tered, each pair now taking possession of a large tract of stunted vegetation,among which they make their nest and rear their young. I was never able toprocure the eggs of this species.As to the haunts of rock ptarmigan in Newfoundland, all recentobservers seem to agree that " Welch's ptarmigan " is confined, atpresent at least, to the barren tops of the highest hills along the westand south coasts. Dr. G. K. Noble (1919) says that "all of theWelch's Ptarmigan observed were found on the very highest rangesof the Lewis Hills. These are composed mostly of syenite, verymuch weathered or fragmented." Ludlow Griscom (1926) agreeswith him that this species " is found only on the highest diorite andsyenite rock barriers " ; he found it also " on the summit ofBlomidon." J. R. Whitaker, who lived in Newfoundland for manyyears, says in some notes he sent to me
:
These birds are distributed through the high tundras and hills of Newfound-land. In the middle section of the country they are not nearly so numerousas along the hills nearer the coast. They remain on the high barrens far morepersistently than Allen's ptarmigan, but it was quite a rare event to see anyon the lower level even during severe winters; this statement applies to centralNewfoundland. I am told on good authority that along the west coast, CapeRay especially, they come down to sea level in numbers during the fall andwinter months.From extensive information recently gathered, I should extendthe range of the rock ptarmigan, in Newfoundland, to include suit-able highlands along the south coast from Fortune Bay to Cape Ray,with several records for Gafftopsail and Kettys Brook on the rail-way line. Probably the whole of the northern peninsula, north ofBonne Bay and White Bay, should be included also. I am told that80 per cent of the ptarmigan that come into the market from thissouthwestern rejrion are " rockers."
204 BULLETIN 16 2, UNITED STATES NATIONAL, MUSEUMF. Seymour Hersey, who spent two months in Newfoundland inthe fall of 1913 and collected 13 specimens of " Welch's ptarmigan "on the Lewis Hills, contributed the following notes on its distributionand haunts :Welch's ptarmigan is much more local in its distribution than Allen's. I didnot find it at Cape Ray or Gafftopsail, and I confidently expected to find itthere, but notwithstanding repeated hunting over many miles of ground Ifound Allen's only. After I collected in the Lewis Hills I returned to Gaff-topsail and showed my birds to the few people who lived at this lonely place,and all agreed that they had never seen a ptarmigan like them. One man whohunted and trapped a great deal did state that they occasionally shot a smallptarmigan in winter which I believe is Welch's.The only birds I obtained were collected in the Lewis Hills, which, at theplace where my collecting was done, run parallel with the beach and only ashort way back from it. They have the form of a continuous ridge, ratherthan of individual hills, and rise abruptly at a steep angle, to a height ofseveral hundred feet. The sides, as well as the strip from the shore to theirbase, are well wooded with trees of fair size. The top is undulating andstretches away for long distances without descent, which makes huntingcomparatively easy. Individual hills are indicated by slightly increased eleva-tions of the general level, but there are no distinct and separate hills.The whole surface is devoid of trees, but the ground is well covered withlow-creeping vegetation, prostrate blueberry bushes, mosses, and grass. Inmore protected spots are patches of dwarf spruces. These are usually 1% to 3feet high and are so twisted and interlaced that it is often possible to walkacross a patch of them without breaking through.My first day in these hills was foggy and I got no birds. Once I heard aptarmigan call and following the sound come upon three Allen's. The nextday was clear and crisp with a good breeze and only scraps of fog. We hadbeen tramping for several hours with little success when we came to an eleva-tion thickly strewn with large granitelike rocks of a very dark gray color.My guide halted and whispered that if we went cautiously he thought wewould soon see some birds. It was not long before I made out a ptarmiganperched on a rock, and very soon after seeing the first one I discovered severalothers. Their dark colors blended with the gray rocks, but they were never-theless rather easy to see. As there was no concealment we walked straighttoward them and had no difficulty in getting within gunshot. At the first shotthey arose, flew a few hundred yards, and alighted again among the rocks.Leaving the guide to retrieve the birds I had shot, I followed the flock andsoon flushed them again. This was done several times until I had collecteda number of specimens, when the remainder flew out of sight.In the Flatbay Brook country where I later camped were some high roundedhills. My guide stated there were ptarmigan to be found well up towardtheir summits, but only in winter, when they had formed into large flocks,could they be successfully hunted. I was also told they were to be foundin the mountains about Fortune Bay. Judging from my experience this speciesis restricted to the tops of certain ranges of hills, mainly near the coast, andthey are partial to areas covered with dark gray rocks rather than the lightercolored rocks where Allen's ptarmigan is so abundant.
ROCK PTARMIGAN 205Spring.?Bernharcl Hantzsch (1929), referring to the spring mi-gration at Killinek, northern Labrador, writes
:
Suddenly in early spring, mostly in April, seldom sooner, at times not untilwell into May, the wanderers appear from the south. Usually at first rathersmall advance posts are established. A short time after that the whole throngof birds follows. As I was assured by the missionaries, Messrs. Waldmannand Perrett, who each have passed a year up to 1906 in Killinek, by Mr. J.Kane who lived there six or seven years, as well as the Eskimos of theneighborhood agreeing, countless large flocks of these birds appear at times,usually passing through rather high in the air. For hours they hasten inmany thousands through the sky, so that their numbers cause astonishment.Many flights of the kind are observed from the same place. The birds mostly flydirectly across Hudson Strait without delaying. This is almost always coveredwith ice in the spring and little to be distinguished from the land. The flight isso swift and high, that Missionary Perrett was in doubt whether the birdswere migrating to Greenland, which can, however, be safely denied, accordingto the unanimous reports from there. The Canadian Neptune Expedition 1903-1904 observed a great migration of these birds at Fullerton, northwest HudsonBay. Only a small percentage of the ptarmigan make a stop in our region inorder to rest up and hunt food. The forerunners and the stragglers stop morefrequently than the main swarm, the latter having perhaps not much farther togo to reach their breeding places. The birds which stop, halt mostly in flocks often to thirty, occasionally still more together, and they usually do notact particularly shyly. When contrary winds and hunger tire the creaturesout, they are so tame that they can be killed with the long dog-whip. Thecaptured birds form a much-preferred article of food for whites and Eskimos,indeed the latter devour even the entrails, especially when these are warm.Courtship.?Turner describes the courtship of the rock ptarmiganin his notes as follows
:
The mating season begins in May, and during this period the male acts inthe strangest manner to gain the affection of his chosen mate. He does notlaunch high in air and croak like the willow ptarmigan, but runs around hisprospective bride with tail spread, wings either dragging like those of thecommon turkey, or else his head and neck stretched out, and breast in con-tact with the ground, pushing himself in this manner by the feet, which areextended behind. The male at this time ruffles every feather of his body, twistshis neck in various positions, and the supraorbital processes are swollen anderect. Pie utters a most peculiar sound, something like a growling kurr-kurr;as the passion of the display increases the bird performs the most astonishingantics, such as leaping in the air without effort of wings, rolling over and over,acting withal as if beside himself with ardor.The males engage in most desperate battles ; the engagement lasts for hoursor until one is utterly exhausted, the feathers of head, neck, and breast strew-ing the ground. A maneuver is for the pursued bird to lead the other off agreat distance and suddenly fly back to the female, who sits or feeds as un-concerned as it is possible for a bird to do. She acts thoroughly the most heart-less coquette, while he is a most passionately devoted lover. He would ratherdie than forsake her side, and often places himself between the hunter andher, uttering notes of warning for her to escape, while attention is drawn tohim who is the more conspicuous.
206 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMNesting.?The nest of the rock ptarmigan is a very simple affair,a hollow in the ground or moss of the open tundra, lined with grasses,mosses, or other convenient material, and a few feathers of the bird.It may be partially sheltered beside a hummock, tuft of grass, or lowbush, but it is usually in plain sight. But Roderick MacFarlane(1908) says:It proved no easy matter, however, to find the nests of this species, as theplumage of the birds and the color of the eggs both strongly resembled theneighbouring vegetation. At the same time the female sat so very closely thatmore than one was caught on the nest, and I recollect an instance where theparent, on the very near approach of our party, must have crouched as muchas possible in the hope that she might not be noticed, which would have hap-pened had not one of the smartest of our Indian assistants caught a glance ofher eye.Eggs.?The eggs of the different races of rock ptarmigan are allmuch alike and are very well described b}r MacFarlane and Brandtunder Kellogg's ptarmigan, but the six sets in my collection showsome types different from those described by others. The groundcolors vary from " clay color " or " pinkish cinnamon " to " palepinkish buff," " cream-buff," or " cartridge buff." They averagemore heavily marked than willow ptarmigans' eggs; two sets arenearly covered with great blotches and splashes of very dark browns,
" chestnut-brown " to " bone brown," or nearly black ; others aremarked like willow ptarmigans' eggs with similar colors. I alsohave one set of 12 eggs. The measurements of 99 eggs in the UnitedStates National Museum average 42 by 30 millimeters; the eggsshowing the four extremes measure 45 by 31, 44 by 32.5, 39 by 29,and 41 by 28 millimeters, including eggs of both mainland forms.Young.?Turner says in his notes:When the young are with the parents they rely upon their color to hide them-selves among the nearly similar vegetation from which they procure their food.1 am certain I have walked directly over young birds that were well able tofly. If the parent birds are first shot, the entire number of young may besecured, as they will not fly until nearly trodden upon, and then only fora few yards, where they may easily be seen. I have found on two occasionsan adult female with a brood of 13 young. All the flocks were secured with-out trouble. At other times only three or four young would be found with bothparents. The young are very tender when first hatched ; no amount of mostcareful attention will induce them to eat, and after only a few hours' captivitythey die. I could never keep them alive more than 12 hours. The changeableweather, sudden squalls of snow or rain, must be the death of scores of thesedelicate creatures. Their note is a soft piping pe-pe-pe, uttered several times,and has the same sound as that of the young bobwhite, Colinus virginianus.Captain MacMillan tells me that ptarmigan occur in flocks aboutBowdoin Harbor. Baffin Island, all through the breeding season, andthe residents say that these are young birds, which do not breed untiltheir second spring.
ROCK PTARMIGAN 207Plumages.?The newly hatched chick of the rock ptarmigan ismuch like that of the willow ptarmigan but is usually somewhat palerand grayer. The crown patch is " chocolate," mixed with andheavily bordered by black; the rest of the head, neck, and breastis " cream-buff " or " chamois," shading off to " colonial buff " on thechin and underparts; a spot on the forehead, a rictal stripe, and abroad auricular stripe are black, the remaining upper parts areheavily blotched and banded with black, " chamois," " honey yellow,"and " tawny."The juvenal plumage begins to appear almost immediately, thewings coining first, in which the two outer primaries on each wing arewhite; these white, juvenal primaries are retained all through thefirst year. Young birds reach the flight stage before they are halfgrown. In the full juvenal plumage the young birds are darker,more heavily barred, and have less rufous than in the same stage ofthe willow ptarmigan. The sexes are alike. The entire upper partsare variegated with black, brownish black, " ochraceous-buff," " ochra-ceous-tawny," and white; the feathers are mainly black, tipped,barred, edged, or notched with the buffs ; many feathers in the mantlehave a terminal white spot ; the breast and flanks are from " cinnamon-buff " to " pinkish buff," fading out to whitish on the chin and belly,spotted on the chin, throat, and neck, and barred on the breast andflanks with sepia or dusky. About the end of July the molt into theautumn plumage begins ; this is also called the tutelar or preliminarywinter plumage; it is common to both young and adult birds and isa transition plumage between the summer and the white winterplumages. The molts overlap and feathers of all three plumages areoften seen in the same bird. During August and September thebirds are molting almost continuously. The colored body feathers ofboth young and adult birds in this autumn plumage are much alikeand are quite different from the feathers of the juvenal or summeradult plumages, being more finely vermiculated or sprinkled, lessheavily barred, and therefore lighter in effect. The black tail feathersare acquired at this postjuvenal molt, which is nearly complete, and,according to Dr. Jonathan Dwight (1900), "the wings (except themedian coverts and inner remiges) become white together with theabdominal wedge of the ventral tract and all posterior to it includingthe flanks, legs and feet; while the head, throat, breast, sides andback become more or less dusky7 according to the extent of the renewalin different individuals and probably according to the latitude."From this time on the colored feathers are gradually replaced bywhite feathers, until the full, first winter plumage is acquired bythis supplementary molt. This is wholly white except for the blackrectrices, and in some young males there are traces of black lores.
208 BULLETIN 162, UNITED STATES NATIONAL MUSEUMIn adult males the black lores are complete and in some old femalesthere are traces of them.In the far north, where the ground is not wholly free from snow,the prenuptial molt, in both young and old birds, is much delayedand is more or less incomplete, as the short arctic summer giveshardly time for the birds to acquire and shed the summer andautumn plumages. But farther south, where the birds live on awholly bare ground, the molt of the body plumage is more com-plete, and the sexes become distinguishable in plumage, as the birdsare now practically adult. In this and subsequent nuptial plumagesthe males assume a coarsely vermiculated and mottled dress of gray-ish buff and dusk}^, except that the wings, tail, abdomen, and feetremain as in winter. The females make a more extensive molt,becoming coarsely mottled and barred with buff and black, butretaining the white remiges, white belly, and black rectrices.Soon after this nuptial, or summer, plumage is acquired, the moltinto the autumn plumage begins, which helps to make the completemolt into the white winter plumage. The autumn plumage is muchlike that of the young bird and is very finely vermiculated in bothsexes, instead of coarsely vermiculated or barred, as in summer. Thesexes are now much alike again. These two plumages are wellillustrated in the colored plates published by A. L. V. Manniche(1910).Food.?In summer the food of the rock ptarmigan includes anumber of insects, but it is made up more largely of leaves, buds, andberries, such as crowberries, whortleberries, bearberries, and the ten-der leaves and buds of birches and willows. Numerous seeds areeaten, as well as sphagnum and other mosses, and the leaves of Lab-rador-tea. In winter, when food is scarce, they have to feed on budsand twigs and the seeds of such weeds as they find above the snow.Their long, strong claws, which are highly developed in winter,enable them to dig down through the snow to reach the mosses andsuch of the above plants as they can find.Behavior.?Turner says in his notes
:
Their flight is rapid, and when flying with a stiff wind they require thequickest shot to stop them. The beat of the wing is so rapid that it is scarcelydiscernible, and when the bird is sailing the somewhat decurved wings areheld almost motionless as it rolls from side to side. The direction of flightis always in a straight line, rising only sufficiently to clear a patch of trees orintervening ridge, the latter at times passed over only by a few inches in height,to the plain or valley beyond. Sometimes they will fly more than a milebefore alighting and at other times only a few rods, depending altogether onthe character of the weather. If it is a cold, blustering day with much snowdrifting or falling, these birds dislike to take flight, and by using a slightamount of discretion the birds of an entire flock may be all secured, but ifit is calm and cold, or warm, they take flight and at this time are rarely
ROCK PTARMIGAN 209
approached within shooting distance. Warm, damp, weather with gusty windis best suited for hunting this ptarmigan.I have often been amused at these birds' actions when descending from ahigh bluff to the level ground below. If there is a place that gradually slopesto the bottom, they seem to prefer to slide or tumble down rather than takeflight. Just back of my house, and only a hundred or so yards, there was abluff nearly 100 feet in height. This was the side of a level tract of groundabove, and to it great numbers of " rockers " came every morning either to sunthemselves or to descend to tbe lower ground nearer the houses and beyond.Their growling and " snoring " could be heard nearly every morning. I oftenwatched them descend. Some individuals would push their feet forward andwith outspread tail on the snow slide to the bottom, while others would rolland tumble over and over until they came to the level ground, where they ranas unconcernedly as birds could do.Voice.?Mr. Whitaker likens the call of this ptarmigan to "thetattoo of a woodpecker on an especially mellow tree." Mr. Herseydescribes it as " a low guttural croak reminding me of the springsong of the crow." O. J. Murie has sent me the following notes
:
When slightly alarmed or annoyed by too close approach, these birds uttereda strange sound, like a short interrupted purr
?
prrt! prrt! This usually indi-cated that they were about to fly. Later, in spring, I heard a more prolongedutterance, possibly the " crowing " of this quiet bird. On several occasions inMay, when I disturbed a group of these birds, some of them produced thissound, which might be described as a rolling " snore "
?
k-r-r-r-a-r-r-r-uk?r-r-r-a, plainly varied by a change in the middle which resolves it into threeparts. Once when I shot a male for a specimen, the female called with a
" whining " note.Doctor Noble (1919) writes:On August 24th, during a heavy rainstorm, while making my way across oneof these fields of grotesquely shaped stones, I came suddenly upon an old malebird. It had just emerged from between two great blocks, and stood lookingat me. After a few moments' hesitation, it stretched out its neck and gavea long cackle, unlike any call I had ever heard. It was a crescendo of clucks,somewhat pheasant-like in quality
?
kulc, kuk, kuk, kuk?each syllable strongerand of a higher pitch than the last.Ludlow Griscom (1926) had a similar experience
:
An old grey cock suddenly appeared on the top of a rock in a field of hugeboulders, not more than fifty feet from me. It stretched its neck, cackledloudly and long, and exhibited no fear at all as I drew nearer. It then dis-appeared, but for some seconds I could hear it clucking to itself in greatdissatisfaction as it threaded its way through the maze of its chosen home.Fall.?From the northern portions of its range the rock ptarmiganmakes quite extensive migrations. Capt. Moses Bartlett told me,and others have confirmed it, that during the last of Septemberbeginning with the first heavy snow squalls and lasting up to aboutthe middle of October, a heavy flight of ptarmigan occurs acrossHudson Straits to Cape Chidley, hundreds of birds being in sightat one time. Often they alight on ships and are easily caught. A
210 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
similar flight occurs from Ellesmere Island to Greenland. In north-ern Labrador, according to Bernhard Hantzsch (1929), the flight isirregular. He says:In the middle of September, 1905, countless flocks are said to have flownsouthward over Killinek. In 1906 the first ptarmigan were not observed until28th September. From 4th October they appeared somewhat more numerouslyin a heavy, driving snow and some cold, but rather large hunting parties didnot get many, particularly as they could not yet travel in the light fall ofsnow. Whether the unusually long and mild fall kept the birds in theirnorthern dwelling places, or induced them to choose another migration route,must remain unknown.Winter.?Many rock ptarmigan evidently spend the winter in thelowlands of Ungava, mainly near the coasts, for Turner says:During the summer months they are quite scarce in the vicinity of FortChimo, retiring to the interior and to the hills of Georges River vicinity forthat season, to return about the first of November. This month will beconsidered as a starting point for their wanderings. Then they appear in fullwinter plumage and in flocks of various sizes, often numbering over 200individuals. They resort to the open ground and rock ridges where the snowis more apt to drift from, or during severest weather they retreat to thesheltered places amongst the bushes of alder and willow along the streamsand gullies. These situations afford food of berries of various kinds, whichyet remain on the stems of prostrate shrubs, twigs and buds of dwarf birch,alder, and willow, together with a few blades of grass. These ptarmiganremain until the latter part of March and disappear as suddenly as they came.O. J. Murie writes to me as follows
:
During winter the rock ptarmigan along the eastern coast of Hudson Baywere very tame and easily approached, much more so than the willowptarmigan. The latter birds confined themselves generally to willows oralders or even in the edge of the woods. L. rupestris, on the other hand,were generally found in the open or rocky slopes, feeding on the willow tipsprotruding from the snow or the berries of Empetrum, in spots where theground was blown bare of snow. At times the feathers around the beakwere stained purple as a result of feeding on this fruit. Even when restingthese hardy birds did not always seek shelter. Once I found a group of twoor three crouching on the bare top of a little rise, all hunched up in a littleball and facing a persistent, cold north wind. One of them remained inthe same position while I approached within about 4 or 5 feet to photographit, finally walking off quietly when I reached out still nearer. Generally thisspecies was found in the shelter of a little bush, a single stunted tree, or smallclump, but not often in heavy cover.Enemies.?Hantzsch (1929) writes:The flights of ptarmigan are accompanied by birds of prey; especially theproud Gyrfalcon, the smaller Duck Hawk, and the beautiful Snowy Owl followthem. If the flight is smaller, as in the autumn of 1906. then these birdsare observed only in small numbers. In addition, foxes assemble, particularlyVulpes layopus (Arctic Fox), in places where there are many Ptarmigan, and allthe other beasts of prey, in like manner, probably take a share at the appearanceof our much desired bird.
ROCK PTARMIGAN 211Game.?Hantzsch describes the ptarmigan hunt as follows:The arrival of the first flocks of these birds is greeted as an event of the day,which controls all the conversation. Now everyone cleans his gun, and eventhe little eight or ten-year-old chap is happy, whenever a gun is occasionallyloaned him. If the ptarmigan appear in great numbers, an occurrence that variesmuch from year to year, then each one who has a gun and ammunition, from themissionary to the youngest Eskimo lad, betakes himself out into the wildmountainous landscapes. And the district is so large for the few people
?
at the most 15-20 men assemble near the Killinek mission station?thatno one is in another's way. They prefer to go alone or in pairs with the dog-sleds, with a young man along for assistance, in order to overtake the birdsmore quickly and be able to take the bag home more conveniently. In thefew days when the birds are present, a good hunter is often able to shootseveral hundred. To be sure the hunt is strenuous. They travel across thewide, snowy landscape until they see a flock flying up somewhere. Sittingdown they do not see the ptarmigan until rather near, as I convinced myself.The hunter now usually springs from the halting sled and approaches thebirds in order to get one or two goods shots at them. The unwounded birdsrise at once and fly away, and it is now a matter of paying attention to wherethey stop again. After the game has been put on the sleds, they journey far-ther, seeking either the part of the flock which has flown away, or new bands.Now and then they see several at the same time, at other times they have towait a long time before coming across a single one.O. J. Murie says in his notes
:
The Indians and Eskimos take advantage of a trait of these birds to trapthem. Ptarmigan have a tendency to gather on a conspicuous dark spot on thelevel white expanse of snow, such as an exposed sandbar. The natives stretchsome kind of netting over a frame, which is tilted up over an exposed plotof ground or a spot where sand or earth has been spread on the snow. As thebirds gather under the net a string is pulled, which allows the net to fall.The trapped birds are then killed by biting them in the neck.The rock ptarmigan is an important game bird in Newfoundland,as about 20 per cent of the ptarmigan that come into the St. Johnsmarket are " rockers." But evidently the hunters prefer the largerwillow ptarmigan, which are, perhaps, more easily obtained. Fromreliable parties who have hunted in the vicinity of Quarry andGafftopsail, as well as on certain sections between Fortune Bay andCape Ray, I have learned that, when they have been unsuccessfulin their hunt for willow ptarmigan, they turn their attention to thehilly sections of the same locality to find " rockers." Sometimes theywould go to hunt for willow ptarmigan and not bother about " rock-ers " unless their hunt for willow ptarmigan was unsuccessful ; thenthey would climb to hilly sections of the same localitj' where theycould always be sure of getting rock ptarmigan.DISTRIBUTIONRange.?Alaska (including the Aleutian Islands), northern Can-ada, Greenland, and Newfoundland.
212 BULLETIN 162, UNITED STATES NATIONAL MUSEUMThe range of the rock ptarmigan extends north to Alaska (CapePrince of Wales, Point Barrow, Humphrey Point, and DemarcationPoint) ; northwestern Mackenzie (Cape Bathurst, Franklin Bay, andPierce Point) ; northern Franklin (Winter Harbor, Gaesefiord, LakeHazen, and Floeburg Beach) ; and northern Greenland (Thank GodHarbor, Newman Bay, and Lockwood Island). East to Greenland(Lockwood Island, Sabine Island, Clavering Island, and Ivigtut) ;and Newfoundland (mountain ranges). South to Newfoundland(Fortune Bay to Cape Ray) ; Anticosti Island (Fox Bay) ; northernQuebec (Fort Chimo and Sorehead River) ; southern Franklin (CapeFullerton and Bernard Harbor) ; and central British Columbia(Ingenika River and Ninemile Mountain). West to British Colum-bia (Ninemile Mountain) ; and Alaska (Baranof Island, St. LazariaIsland, Mount Edgecumbe, Port Frederick, Hinchinbrook Island,Montague Island, English Bay, Unalaska Island, Atka Island, AdakIsland, Tanaga Island, Amchitka Island, Kiska Island, Attu IslandsAskinuk Mountains, Nome, Teller, and Cape Prince of Wales). Itis of casual occurrence in summer on Bonaventure Island, Quebec,where one was taken July 8, 1922 (Stoddard).Although the majority of the rock ptarmigan withdraw from theextreme northern part of their summer range upon the advent ofsevere winter weather, they do not appear to retreat beyond thesouthern limits of the breeding range. Instead, a concentration isnoticed at some of the more southern points, as at Ivigtut, Greenland.Among late departure dates in the northern part of the range are:Franklin, Floeburg Beach, September 29. and Winter Harbor, Octo-ber 15. The species has been observed to return to this general areaas follows: Franklin, Newman Bay, March 9, Floeburg Beach,March 11, Thank God Harbor, March 24, Discovery Harbor, April10, and Winter Harbor, May 12.The range as above described is for the entire species, underwhich 10 subspecies are recognized. True rupestris is found fromNewfoundland and the Ungava Peninsula west to northern BritishColumbia and southern Yukon; Reinhardt's ptarmigan {Lagopus r.reinhardi) is found in southwestern Greenland, north to the vicinityof Disco ; Nelson's ptarmigan {Lagopus r. nelsoni) is found on Una-laska, Akutan, and Unimak Islands, Alaska, and also on some of theother eastern Aleutian Islands; Turner's ptarmigan {Lagopus r.atkhensis) is confined to Atka Island, of the Aleutian group; Cham-berlain's ptarmigan {Lagopus r. chamberlaini) is found on Adak Is-land of the Aleutians; Sanford's ptarmigan {Lagopus r. sanfordi) isconfined to Tanaga Island of the Aleutians; Townsend's ptarmigan{Lagopus r. townsendi) is found on Kiska Island, of the Aleutianchain; Evermann's ptarmigan {Lagopus r. evermanni) is found on
reinhardt's ptarmigan 213Attn Island, of the Aleutians; Kellogg's ptarmigan (Lac/opus r.kelloggae) occupies northwestern Greenland, the Arctic Islands (ex-cept Baffin Island), northern Yukon, the interior of Alaska, and thewest Arctic coast to Coronation Gulf; and Dixon's ptarmigan (Lago-pus r. dixoni) is found on the alpine summits of Baranof , Chichagof
,
and Admiralty Islands, and the adjacent Alaskan mainland.Egg dates.?Northern Alaska : 15 records, May 28 to July 29 ; 8records, June 9 to July 1. Arctic Canada: 23 records, June 3 toJuly 9; 12 records, June 17 to 28. Iceland: 18 records, May 7 toJuly 21 ; 9 records, May 20 to June 19. Greenland : 17 records, May20 to July 6; 9 records, June 16 to 30. Labrador Peninsula: 12records, June 11 to July 7; 6 records, June 3 to 20. AleutianIslands : 7 records, June 10 to 26. Newfoundland : 11 records, June2 to 12. LAGOPUS RUPESTRIS REINHARDI (Brehm)REINHARDT'S PTARMIGANHABITSThe race reinhardi of the rock ptarmigan was described fromsouthern Greenland, where it undoubtedly forms a well-marked sub-species. A. L. V. Manniche (1910) treats of the rock ptarmiganof northeast Greenland under the name Lagopus mutus, that ofthe Old-World species, from which the Greenland birds differ onlyslightly. I am inclined to think that a careful study may showthat mutus is a circumpolar species, of which the American formsare only subspecies. The 1910 American Ornithologists' UnionCheck List includes in the range of reinhardi the northern extremityof Ungava and western Cumberland Sound. Unfortunately summerspecimens of rock ptarmigan from anywhere on the Labrador Penin-sula are too scarce for us to form any satisfactory opinion as to whatthe birds of that region really are. Until we know more definitelywhat the birds of Labrador and Ungava really are, it seems morelogical to refer them to rupestris and confine reinhardi to westernGreenland south of Disco. The Greenland birds have been sub-divided into three races, as fully explained in Dr. R. M. Ander-son's footnotes in his translation of Bernhard Hantzsch's (1929)Labrador paper. But for life-history purposes, I prefer to treatthe Greenland birds as all of one form.Nesting.?What little information we have on the nesting habitsof Reinhardt's ptarmigan indicates that it does not differ in thisrespect from other races of the rock ptarmigan. Capt. D. B. Mac-Millan told me that he once found a nest, on Baffin Island, that wasplaced on a nubble, not more than 10 feet long, surrounded by74564?32 15
214 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
water in a pond; it would be interesting to know how the mothertransported her j^oung to the shore.Eggs.?The eggs are indistinguishable from those of the rockptarmigan. The measurements of 51 eggs average 41.9 by 30 milli-meters; the eggs showing the four extremes measure 44.5 by 29.5,44 by 32, 39 by 30, and 40 by 29 millimeters.Food.?A. L. V. Manniche (1910) says:Their principal food consisted of buds and short bits of stalks of Salixarctica. According to Dr. Lindhard's analyses stomachs of Ptarmigans shotat this season also contained leaves of Dryas ootopetala and crowns of leavesof Saxifraga oppositifolia.In stomachs of shot young ones I found remnants of plants as well as ofinsects. The old birds in summer also partly feed on insects.Behavior.?The same observer writes:In fine weather these hardy birds did not seem inconvenienced by thetemperature frequently as low as some 40? below zero. But it was hard forthe ptarmigans to support their lives during severe snow-storms and whenthe earth was covered by thick, evenly lying crusted snow.At sunset they flew to the rocks and remained there over night. In theheavy snowmasses on the leeside of the rock they digged holes some 20 centi-meters deep, just large enough for the body of the birds, and here they spenttheir nights apparently without ever altering position judging from themanner in which the excrements were deposited. When several ptarmiganshad spent a night in company, their holes were always placed within a rathernarrow circumference sometimes nearer and sometimes at a longer distance,but never quite close to each other. The ptarmigans would also often spendtheir nights in narrow ravines in the rocks filled up with snow.Sometimes I found my old foot-prints taken possession of by the ptarmigansas night-quarters. They were by night not seldom frightened out of theirholes of polar foxes and erimines. which could be easily seen on new fallensnow. I found, however, in no case, signs that ptarmigans were caught inthis way.When a female ptarmigan was going to fly up, she would raise the featherson the back of her head to a pointed crest and lay the tips of her wings onthe upper rump uttering a suffocated clucking, that could best be comparedwith the call of FrlvgiUa montifringilla ; at the same time she would executesome courtesying movements with her head and the forepart of her body.Just after a heavy snow-storm, that, covered all the earth evenly with snow,the ptarmigans would prove extremely shy. For a few moments at a timethey would settle on summits of rocks or stones, that reach over the snow, andthen, by a rapid soundless flight, disappear around corners of rocks throughdeep ravines or out over extensive plains. When the ptarmigans after somehours had found places with food, they would again become tranquil.Winter.?It seems to be well established that this and other truerock ptarmigan are migratory. Mr. Manniche (1910) says:In the absolutely dark season ptarmigans or foot-prints of them were nowherefound in spite of numerous researches on different places, and there can beno doubt, that tins species for some three months leaves this part of North-East Greenland.
nelson's ptarmigan 215
It may be supposed, that the birds only migrate to somewhat more southerlylying parts of East Greenland, as they already begin to return in the beginningof February, when the sun has not yet appeared. (In 1907 the first ptarmiganwas seen at the ship's harbour February 4th and the next year 4 days later.)The migration lasted through February. March, and the larger part ofApril, and the number of ptarmigans within a certain place might differ agood deal in this time.Captain MacMillan (1918) recorded them as "common at Etahin spring and fall migration " ; and as " not seen in July and Au-gust." The migration is very early, as he shot some at Etah onFebruary 13 and saw them on the inland highlands of EllesmereIsland and Grant Land in March.Hagerup (1891), referring to the region about Ivigtut in south-ern Greenland, writes:During winter the number is considerably increased by the birds comingfrom the north, but the abundance is very variable. Thus the first winterI was at Ivigtut, an uncommonly cold season, comparatively few were seen,though about 400 were shot; but the following winter, which was muchmilder, the birds were much more numerous, and about twice as many werekilled. When snow covers the ground they are less frequent in the valleys thanon the mountain slopes and in the clefts; but on the high lands they are notso numerous. They usually resort to side hills, where there are largebowlders, and where some herbs are easily accessible. They change theirfeeding-ground very often, and sometimes in the course of a single night theyarrive in such numbers that on the following day the birds or their tracks maybe seen everywhere, while at other times one may travel for days withoutseeing any sign of one.LAGOPUS RUPESTRIS NELSONI StejnegerNELSON'S PTARMIGANHABITSNelson's ptarmigan is perhaps the best known of the half dozenforms of the rock ptarmigan found in the Aleutian Islands. It is apermanent resident on Unalaska Island, particularly on the easternand more mountainous end, on the Krenitzin Islands, east of Un-alaska, and on the Alaska Peninsula, at the base of which it appar-ently intergrades with the mainland rock ptarmigan. Dr. WilfredH. Osgood (1904), who collected specimens on Portage Mountain, atthe base of the peninsula, says: "With the material at hand I havebeen unable to satisfactorily distinguish the rock ptarmigan of theAlaska Peninsula from those of Unalaska Island."About Iliuliuk Village, on Unalaska Island, we found ptarmiganscarce and wild, even on the mountains back of the village, wherethey are persistently hunted by the natives. Dutch Harbor is onAmaknak Island, which is separated from Unalaska by only a narrow
216 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
channel. Here, between June 5 and 9, 1911, we found them quitecommon on a small mountain, locally known as " Ballyhoo," wherewe collected most of our birds, shooting nine one day and sevenanother. This mountain is only about 1,800 feet high, but is verysteep and is topped by a knife-edged ridge covered with crusted snow.It is a stiff climb up the south side where the steep slope is clothed insoft mosses, with a sparse growth of coarse grasses, cow-parsnip, andother small herbaceous plants and with bare soil or rocks here andthere. The north side breaks off suddenly into precipitous, rockycliffs, straight down to the bay below. One day I started out to huntptarmigan on this mountain in a cold driving rain typical of AleutianIslands weather; when halfway up the rain changed to snow, andwhen I reached the crest of the ridge the bay below was hidden in ablanket of fog, and across the bay a snow-capped mountain stood outin bright sunlight. Austin H. Clark (1910) found these birds at" the rugged northeastern end " of this island " on the mossy lowerslopes, and one or two on the seacoast itself about the mouth of snow-filled ravines." But all of our birds were taken well up on the sidesof the mountains.Courtship.?At the time of our visit the ptarmigan were busywith their courtships. The males were very noisy and conspicuous;usually several could be seen sitting on little hummocks, as we lookedup the mountainside; evidently each cock bird has his own specialhummock, which he defends against intruders, for it is well dec-orated with droppings and molted feathers. Here he sits or strutsabout, clucking and displaying his charms, with the flaming redcombs above his eyes fully extended, while his prospective mate,now inconspicuous in her mottled summer dress, walks about inthe vicinity quite unconcerned. At frequent intervals he rises intothe air 30 or 40 feet above the ground and floats or flutters down-ward, sometimes scaling on down-curved wings, uttering duringthe descent his loud clucking, or rattling, call, icvck, wuclc, ivuck,many times repeated. A pair of birds could often be located byseeing this song flight of the male, his white wings being quiteconspicuous at a long distance. It is a very pretty performanceand makes a striking display. Lucien M. Turner (1886) refersto a performance that is evidently part of the courtship : " In themale the neck is stretched along the ground, the tail spread andthrown over the back, the wings outstretched, while he utters arattling croak that may be heard for a long distance."Nesting.?We found no nests of this ptarmigan, and none of thefemales collected were anywhere nearly ready to lay. We assumedthat the}' would nest later in the season on the lower, more grassyslopes. Turner's (1886) remarks probably refer in part to this
nelson's ptarmigan 217ptarmigan, which he says is extremely abundant on some of theislands in the eastern part of the Aleutian Chain. He writes:The mating season begins in the early part of May, and is continued forabout three weeks, by which time a site for the nest is chosen, usuallyamidst the tall grasses at the mouth of a wide valley, or else on the opentundra among the moss and scanty grass.The nest of this bird is composed of a few stalks of grass and a few feath-ers that fall from the mother's breast. The nest is a very careless affair,and often near the completion of incubation the eggs will lie on the bareground surrounded by a slight circle of grass stalks that have apparentlybeen kicked aside by the mother impatient of her task. The number ofeggs varies from nine to seventeen, eleven being the usual number.Dr. Richard C. McGregor (1906) found a nest on a small island inthe Krenitzin Group, east of Unalaska, that unquestionably belongedto the Nelson's ptarmigan; it was taken on Egg Island on July 6;
" the nest was a mass of grass, leaves, and a few feathers " andcontained six eggs.Eggs.?The eggs referred to above are described by Doctor Mc-Gregor (1906) as follows:In color the eggs are dull creamy brown overlaid with irregular spots ofdark reddish brown, almost black. The larger markings tend to form a ringnear the large end of each egg, but this ring is rather poorly defined. Theeggs measure as follows, in millimeters and tenths: 42.3X30.1; 42.4X30.6;40.8X30.5; 41.7X31.3; 42.4X31.8; 45.2X31.4. Incubation was begun.There are three eggs in the National Museum collection that re-semble certain types of rock ptarmigan's eggs. They are ovate inshape with very little gloss. The ground colors are creamy whiteor pale buff. In one the ground color- is nearly covered with smallspots and fine sprinkles of very dark brown ; in another it is nearlycovered with large blotches and small spots of " claret brown " and
" liver brown." The measurements of nine eggs average 42.5 by 30.5millimeters; the eggs showing the four extremes measure 44.2 by30; 42.5 by 31.8, and 41.6 by 29.9 millimeters.Food.?Doctor Osgood (1904) reported that " an examination ofthe crops of 10 birds killed at Cold Bay showed a variety of food,but buds, particularly willow buds, predominated. Tiny buds andtwigs of some small species of Vaccinmm were found in large num-bers, which must have been secured by a very tedious process. Someof the craws contained nothing but buds, others had a few leavesof Dryas and Ledum, and occasionally one contained some brokenpieces of large aments of Alnus viridis.'"Behavior.?Turner (1886) writes:The young are able to follow the mother as soon as they are hatched. Asthis bird never collects into large flocks, I always supposed the flocks seen inwinter were the parents with the brood reared the previous summer. Thepower of flight of this bird is much stronger than its congener. It is sustained
218 BULLETIN 1(52, UNITED STATES NATIONAL MUSEUMfor a Longer period and much more rapid. The flesh of this .species is betterthan that of the Willow Ptarmigan and is much sought for as food. The besttime to hunt this bird is early in the morning when the wind is calm and amoist snow is falling. The birds are then sluggish and dislike to rise to thehill-tops.There is not much more to be said about the habits of this ptarmi-gan, which apparently do not differ materially from those of themainland rock ptarmigan. This form is much closer to the main-land form in appearance and habits than it is to the other Aleutianforms. It is essentially a bird of the mountains and foothills. It istame enough where it is not molested, but it soon becomes sophisti-cated where it is hunted persistently. On the mountains back ofIliuliuk Village on Unalaska Island, where we collected a few speci-mens, we found it a really sporty game bird. It usually flushed atlong range, with loud clucking notes, and flew very swiftly for a longdistance, often across some deep ravine, where a long hard walk wasnecessary to flush it again. The man who makes a good bag undersuch circumstances earns his birds.LAGOPUS RUPESTRIS ATKHENSIS TurnerTURNER'S PTARMIGANHABITSThe four forms of ptarmigan found on four of the more centralislands of the Aleutian Chain, atkhensis on Atka Island, townsendion Kiska Island, chamberlaim on Adak Island, and sanfordi onTanaga Island, seem to me to be sufficiently different from theirnearest neighbors, nelsoni on the east and everwan n i on the west,to warrant recognizing them as four subspecies of a species distinctfrom rupestris. They are all somewhat larger than the mainlandrock ptarmigan, with larger and heavier bills, and their eggs aredecidedly larger. They are birds of the lowlands, living on thelow, rolling hills, grassy plains, and sand hills near the coast; where-as the rock ptarmigan, as well as nelsoni and evemianni, is essen-tially a bird of the mountains and moss-covered foothills, comingdown to the lowlands only on the arctic tundra. The dark colorsof the mountain forms match their habitat, as well as the lightcolors of the lowland birds match theirs. The Aleutian Islandsappear to be the summits of a submerged mountain chain, whichat one time may have formed a land bridge between Asia and NorthAmerica. It seems likely that the central portion of the chain majhave subsided first, isolating the central islands long before theeastern and western islands were separated from the two continents.This might have given the birds on the central islands a much longertime to differentiate, while the birds on the two ends of the chain
turner's ptarmigan 219have remained more like the mainland birds. This might accountfor the presence of light-colored birds in the center and dark-coloredbirds at both ends, as they now exist.On Atka Island we found this form of ptarmigan very abundant
;
there seemed to be more ptarmigan here than on any island wevisited. There were comparatively few of them on the hillsides,but in the grassy hollows and among the low, rolling hills of thevalleys we were constantly flushing them. They were apparentlymated and breeding on June 13, the day of our arrival, but wefailed to find any nests during the next two or three days. Themale usually flushed first with loud clucking notes, and the femalewas sure to follow soon after him.Courtship.?They were very tame and always in pairs, so we hadplenty of chances to observe their courtships. On his song flightthe male rises 30 or 40 feet in the air and floats down again ondecurrent wings, giving a few rapid wing strokes before alighting;sometimes, after checking his descent by rapid wing strokes justabove the ground, he sails along and upward to repeat the sameperformance; during his descent, and particularly during the rapidwing strokes, he utters his loud croaking notes, kruk, kuk, Jmk, huk,kuh, or kTTU-ru-ru-ruy ru-ruk, a prolonged, rattling, nasal, cluckingsound of great carrying power. Once I saw a male chasing a femalein the nuptial pursuit flight; she led him a long chase up and downthe valley and over some low hills, until they finally settled nearme on the tundra, where they strutted about in plain sight. Themale carried his head high, with the bright orange-vermilion combover the eyes swollen and distended and with the tail erected andspread as he walked about in a slow and stately manner.Nesting.?All we know about the nesting habits of this ptarmiganis contained in the following brief statement by Lucien M. Turner(1886), who discovered and described the bird:The nest is built amongst the rank grasses at the bases of hills and thelowlands near the beach. The nest is carelessly arranged with few driedgrass stalks and other trash that may be near. The eggs vary from eleven toseventeen, and are darker in color than those of rupestris, and but slightly in-ferior in size to those of L. lagopits. A number of eggs of this species were pro-cured, but broken in transportation ; hence, can give no measurements ofthem.Eggs.?There are 24 eggs of this ptarmigan in the United StatesNational Museum, which do not differ in appearance from thoseof the other Aleutian races. The measurements average 44.8 by32.9 millimeters; the eggs showing the four extremes measure 48.2by 32.1, 45 by 34.5, 42.4 by 32.3, and 43.6 by 31.3 millimeters.Plumages.?Not much is known about the plumage changes ofTurner's ptarmigan. Our birds were all in full summer plumage in
220 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMmid-June. A series of 15 birds collected on April 4 by Hamilton M.Laing (1925) were just beginning to molt into the summer plumage;a few were still in full, white, winter plumage.Food.?Of the birds collected by Laing (1925), "the cocks hadlittle in their crops; the hens in most cases were stuffed with thefoliage of the crowberry."Behavior.?Laing (1925) first met this ptarmigan on Atka Islandon April 4, 1924, of which he writes
:
On a cold, windy morning, with snow squalls between periods of sunshine, aDuck Hawk posting along the shore was seen to rout some of these white chick-ens of the north almost from the water's edge and send them whirling overthe white hilltop. On going ashore it was found that about fifty ptarmiganwere in the vicinity. They refused to fly very far and during the hunt seventeenspecimens were secured. Rank grass in tussocks and crowberry patches grewon the hills and as there had been a good fall of snow, walking was difficult.Sometimes the birds were wild, again rather stupid. They were first foundcuddled in the sun against the sheltered wall of a small canyon enclosing abrawling stream. Afterwards they flew from one hilltop to another. They werevery speedy on the wing, usually flew downwind, and were extremely difficultto kill.We noticed that the flight of the male is particularly strong andvigorous ; he seems to delight in sailing against a strong wind, whenhe can soar for a long distance, rising and falling again and again,or even remaining perfectly stationary in the air like a poised falcon.When walking the head is carried high, and the motions are verydeliberate and stately, almost stealthy in appearance, with frequentnervous twitches of the tail.Voice.?The male utters his loud clucking notes, similar to thosementioned under courtship, while walking on the ground, whenstarting to fly, or when alighting. The female has a much softernote, like cook, which is very seldom heard, as she is usually a silentbird. Laing's (1925) impressions were as follows:The strangest thing about them was their purring snort like that of astartled horse. This seemed an alarm call. A sentinel stood on a hill andgave it again and again. It could be heard 300 or 400 yards, but was veryelusive and difficult to locate. One bird gave it in flight, with opened beak. Acock that was winged purred again and again when chased and stoppedpurring only when caught.LAGOPUS RUPESTRIS TOWNSENDI ElliotTOWNSEND'S PTARMIGANHABITSIn his original description of this subspecies, Dr. D. G. Elliot(189G) gave as its range both Kiska and Adak Islands, although heobserved that " there is a slight difference in the appearance of thebirds from the two localities, and this can be attributed possibly
chamberlain's ptarmigan 221somewhat to the difference of date in their capture, the Adak birdshaving been obtained one month later, but more to their geographicaldistribution, as Adak is several hundred miles east of Kyska, and thebirds' environment has produced a different result upon them but one,not yet sufficiently pronounced to establish even a subspecific form."It is now well known that townsendi is confined to Kiska Islandand is a well-marked race, being darker and more heavily barred thanany of the other races found on the central islands. The Adak birdhas since been separated by Austin H. Clark (1907), under the namechamberlaini.We were on Kiska Island from June 17 to 21 and on Adak Islandon June 26 and 27. We collected good series on both islands, inwhich the characters of both races are well marked, showing that thetwo forms are quite distinct. It is, therefore, clearly a geographicaldifference and not a seasonal change, as Doctor Elliot (1896)evidently thought it might be.At Kiska Island, on June 17, the ptarmigan of this form were stillin the uplands, were much wilder than the Atka birds, and not nearlyso abundant ; but during the few days that we were there we succeededin collecting a good series. No nests were found ; probably we weretoo early for complete sets, and nests would be found later on in thelong grass of the lowlands. So far as we could see, the habits andbehavior of the Kiska birds are similar to those of the other races onthe neighboring islands, as their environment is practically the same.These islands are all so widely separated that it seems very unlikelythat the ptarmigan ever fly from one to the other. Hence each islandhas produced its peculiar form, which is completely isolated andpermanently resident. All these ptarmigan proved to be very goodto eat, and we found their plump bodies very welcome additions tothe ship's stores of canned food.LAGOPUS RUPESTRIS CHAMBERLAINI ClarkCHAMBERLAIN'S PTARMIGANHABITSThe Adak ptarmigan was separated from the Kiska and Atkabirds and given the name chamberlaini by Austin H. Clark (1907).It differs from the Kiska bird " in its finer vermiculations above,which give the bird a grayer appearance?the whole plumage pre-senting a much more delicate pattern." He calls it intermediatebetween the Kiska and the Atka birds, " but in general colorationit is much grayer than either, being the grayest of all the Aleutianptarmigan."
222 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMWe spent June 26 and 27, 1911, on Adak Island, where we foundthis ptarmigan common but not abundant; we obtained all that weneeded for specimens. We found them mostly in a broad valley ofsmall, low, rolling hills, with a number of small ponds scatteredthrough it; on either side of the valley were mountainous peaks, withrocky summits and with plenty of snow on them. The valley wasmostly dry tundra carpeted with a dwarf species of reindeer moss,which the gray plumage of the ptarmigan matched very well. Thebirds were also found on the low hills and in the grassy hollowsand lowlands but not on the mountains. They were much wilderthan we had found them on other islands, which seemed strange,as this island is uninhabited.Nesting.?On June 26, 1911, I found a nest of seven fresh eggsin a little valley on a hillside; it was a deep hollow in the groundbetween a tuft of grass and a little cow-parsnip; it was carelesslylined with dry grass and a few feathers. Dr. Alexander Wetmorealso collected a similar set of seven fresh eggs on the same day.Evidently all the Aleutian ptarmigan are late breeders.Eggs.?The eggs we collected are ovate in shape, and the shellis smooth with little or no gloss. The ground colors vary from
" pinkish cinnamon " to pale " pinkish buff " ; some eggs are washedwith " cinnamon " at one end or the other, giving the egg a richlycolored effect. In one set the eggs are thickly covered with verysmall spots and fine dots of dark browns, which are sometimesconcentrated near the small end; two of the eggs have a few large,irregular blotches near the small end. The other set is markedlike the common types of ptarmigan's eggs. The markings vary incolor from " chestnut-brown " to " bone brown." The measurementsof these 14 eggs average 46.1 by 32.9 millimeters; the eggs showingthe four extremes measure 47.5 by 33.2, 45.6 by 33.6, 45 by 32.8,and 46.5 by 32.4 millimeters.Plumages.?Five males collected by Hamilton M. Laing (1925) onAdak Island, April 13, 1924, had already begun to molt into thesummer plumage.Food.?All the Aleutian ptarmigan that we collected had beenfeeding entirely on green food, principally the young, green leavesand buds of the dwarf willows, the tops of ground evergreens andmosses, and the flower buds and blossoms of herbaceous plants.Behavior.?We noted nothing peculiar in the habits of this ptarmi-gan, which were similar to those of its neighbors. Laing (1925) sa}Ts
:
Ptarmigan were even more numerous at Kuluk bay, Adak island, than onAtka island. On April 13 ptarmigan were purring everywhere and were allnoted in the grass at low levels. There was no time to ascertain whether theywere also numerous on the dark. cro\vberry-c< >vered hills above, but they werescattered over the flats near the lagoon and the nearby lower grass-covered
DIXON 'S PTARMIGAN" 223
nills. In the distance was seen what, apparently, was a lively fight betweentwo birds. There was only one round. Several times birds when routedwhirled aloft 50 feet or so and then settled again slowly, purring loudly andperhaps threateningly, though what this manoeuvre was for was not clear.Several were shot with the .22 rifle, but they were very tenacious of life and ashot through the body with a hollow-point seldom actually killed them. Someso hit flew a hundred yards before falling. Some of the birds were quite wild,others comparatively tame. They were wildest during the cold, blustery snowsqualls. LAGOPUS RUPESTRIS DIXONI (Grinnell)DIXON'S PTARMIGANHABITSDixon's ptarmigan is one of the many dark-colored races so charac-teristic of the humid coast belt of the northwest. It was discoveredby Joseph Dixon and was named for him by Dr. Joseph Grinnell(1909), who ascribed to it the following characters: "ResemblingLagopus rupestris nelsonl in corresponding plumage, but muchdarker; in extreme blackness of coloration nearly like Lagopus ever-mannh but feathers of chest and back more or less finely vermiculatedwith hazel."Dixon's account (Grinnell et al., 1909) of securing the first speci-mens is interesting and gives an idea of the inaccessible haunts of thisbird
:
I was crawling down a ledge on the north side of the rocky summit of amountain at 2700 feet altitude. About twenty-five feet below me a sharp rockjutted out, forming the crest of a hundred-foot cliff. I had glanced along theledge below but saw nothing, when suddenly a gray-backed ptarmigan rosefrom a bunch of heather on a narrow ledge and trotted out on a jutting rock,bobbing its head and watching me intently the while. I fired a light charge atthe bird which dropped over the cliff. At the report two other ptarmiganjumped up and started swiftly away. I dropped one with the remaining barrel.Then I began the descent to retrieve the birds. By going down to one side of thecliff I had almost reached its base when I came to a sheer drop ; so I had todig my fingers into the crevices and work my way back up again. By going along way around I finally reached a twenty-foot snow drift at the foot of thecliff and there I found my two birds dead. Both had their crops stuffed withheather buds.George Willett (1914) says of its haunts:During the summer months these birds keep well up toward the summits ofthe mountain ranges, above timber line, where they feed on heather buds andberries. Owing to the difficulties in ascending these mountains, specimens arehard to secure at this season. They apparently move in bodies from onesection of the mountains to another, and locating them is largely a matter ofluck. I have been in sections of the mountains where sign less than a week oldwas abundant, but the most diligent search failed to locate a single bird.Whether these changes of location are due to the weather or food supply I amunable to state.
224 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMNothing seems to have been published about the nesting habits ofthis ptarmigan. There is a set of six eggs in P. B. Phillipp's collec-tion, of which the average measurements are 42.2 by 29.7 millime-ters; the eggs showing the four extremes measure 43.9 by 29.9, 42.3by 30.4, and 40.8 by 29.4 millimeters.Doctor Grinnell (1909) quotes from Mr. Dixon's notes, as follows:Their flight was very swift, more like that of a falcon than a quail. Themales would fly out over the mountain side, hover for a moment and thenswoop down, and alight on a rock, uttering their loud, rasping call, whichsounds similar to the noise produced by running a lead pencil over a stiffrubber comb.Alfred M. Bailey (1927) gives us two good pictures of Dixon'sptarmigan in its autumn habitat
:
During October there appeared a wealth of small birds, and many Ptarmiganwere seen and collected. The vegetation was in the height of its " autumnglory," and a peculiar " lily pad," which flourishes abundantly, colored the hillsan intense yellow above timber line, while still higher, among the piledboulders, there was a small ever-green growth upon which the Ptarmigan werefeeding. A few were found in such a site, and some of them were extremelywild. The Ptarmigan in Granite Creek were taken among the boulders andslide rock on the summit of the highest mountains surrounding the valley,at an altitude of over 4.000 feet. There was absolutely no vegetation. Thatthey are well named "Rock Ptarmigan" there can be no doubt after notingtheir habit of sunning themselves upon the tops of large boulders; one restedupon a little overhanging ledge which left a sheer drop to the valley floor farbelow. Several small flocks were seen flying about like so many Doves.They raised from the mountain on which we were hunting and sailed across thevalley to the foot of a hanging glacier. One band flew over me and I tried todrop a bird on our narrow ridge, but the tumbling Ptarmigan sailed on intospace and dropped at least 1,500 feet to the valley floor.Again, on November 11, he wrote in his notes
:
When just above timber line I saw a Three-toed Woodpecker on a dwarfedhemlock, and, on the snow fields above, about thirty Ptarmigan. The tops wereicy, making creepers a necessity. The birds were in full winter plumage, won-derfully handsome fellows, the white of the males being relieved by the blackeye patch. Their call notes could be heard from all sides of the snow coveredmountains, and here and there cream-colored birds, gleaming in the sun-light,could be seen. Overhead an eagle circled, and soon the air was filled withflying Ptarmigan, although I did not see the Eagle make a swoop toward them.Of all the birds seen, only five were in one band, while the others were scat-tered in singles or pairs, and I wondered if they spread to feed among thelittle patches of grass sticking through the snow, or for the protection whichisolation sometimes brings. The call note of the males was constantly heard.This note has an individuality about it which can be mistaken for no other bird.The Ptarmigan were tame, and often allowed us within good photographingdistance, especially if we tried to imitate their note. They rise from the groundwith great speed, and usually their flight is direct, although when flying outover a valley, they often slant down as though to attain greater speed. A fewspecimens were taken, and we found it difficult to secure our birds, for immedi-
sanford's ptarmigan 225
ately they were hit, they started sliding down the slippery mountain side, anddid not stop until they reached the brush line far below. This particular hab-itat was picturesque, to say the least, and on this day was remarkably beau-tiful ; for the cloudless sky was a deep blue, the horizon was the serrated whiteline of the mountain tops, and the winding glacier?from its colorless snowfields at the summit to the seamed and rugged ice field below?wiih its char-acteristic shadows and high-lights of blue and white, made a wonderfulpanorama.Bailey (1927) says of its winter habits:After the winter snows have covered the mountains to a considerable depth,these birds drop to the valley floors where they feed among the alders andwillows. They were often encountered during the following winter days, some-times in large flocks, and many specimens were secured. The species may beconsidered a rather common bird in its proper habitat, near Juneau; it issimply a matter of looking in the proper place?and often involves some ratherstrenuous work. LAGOPUS RUPESTRIS SANFORDI BentSANFORD'S PTARMIGANHABITSAlthough I described and named this race myself (1912), in honorof my friend Dr. Leonard C. Sanford, who cooperated with me inorganizing our expedition to the Aleutian Islands, I must confess thatit is only slightly differentiated from the Adak ptarmigan. We allnoticed a difference when our birds were collected, and when we laidour series of about 40 specimens of sanfordi beside nearly as many ofchamberlaini, it was easy to see that the Tanaga birds were appreci-ably paler in color than the Adak birds. The Tanaga birds aretherefore the lightest in color of any of the Aleutian ptarmigan andhave the finest vermiculations.We landed on Tanaga Island on June 25, 1911, and spent only halfa day on shore; so far as I know, no one had ever collected birds onthis interesting island before ; we found it very rich in bird life, andit is a great pity that we were not able to spend more time there.Back of the sandy beach on which we landed was a series of sand hillsor dunes, covered with long grass, and beyond these was a flat, alluvialplain or tundra, with one large and several small streams flowingthrough it from the mountains farther inland, and dotted with anumber of small ponds and wet meadows. Northern phalaropeswere breeding commonly among the ponds and meadows, and Aleu-tian sandpipers were abundant, indulging in their flight songs andnesting on the little knolls and hummocks on the tundra, where abrood of downy young was found. At the base of a steep hillside acolony of fork-tailed petrels was beginning to breed. The ptarmiganwere tamer and more abundant here than on anv of the other islands
226 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthat we visited; we shot more than 40 in one afternoon. They werecommonest on the rolling, grassy hillocks and grassy hills on thetundra. They flushed at short range, did not fly far, and were easilyshot.Nesting.?Five nests were found, but only three sets of eggs werecollected; the other two were left to be photographed the next day,but we were forced to go away and leave them, as well as some nestsof Aleutian sandpiper and an eagle's nest found by some of the crew.One nest containing nine fresh eggs was on the side of the steep over-hanging bank of a stream; it was in a hollow between two large tuftsof grass and well hidden under one of them; the hollow measured7 by 8 inches and was lined with coarse grass and feathers. Anothernest, containing eight fresh eggs, was a hollow in the ground, meas-uring 7 by 6 inches and 3 inches deep, between two little mossy hum-mocks and under a scraggly cow-parsnip; it was on a little grassyhillock near the beach and was lined with coarse grass and feathers.Other nests were well hidden in the long grass and were found byflushing the birds.Eggs.?Judged from the three sets of eggs that we collected, con-sisting of eight or nine eggs each, the eggs of this ptarmigan are veryhandsome, in fact the prettiest ptarmigan's eggs I have ever seen.They are ovate in shape with a smooth and slightly glossy surface.The ground colors vary from " ochraceous-tawny " or " cinnamon "to " cream-buff " or " cartridge buff " ; some of the eggs are washedat the large end or at the small end with " tawny," giving them a veryrich appearance. They are boldly and heavily marked with largeirregular blothes and small spots of the colors usually seen on otherptarmigan's eggs, dark browns, " chestnut-brown " to " bone brown,"or nearly black. The measurements of 25 eggs average 46.5 by 33.9millimeters ; the eggs showing the four extremes measure 48.2 by 34,46.6 by 34.6, 44.2 by 34.6, and 45.5 by 32.8 millimeters.Plumages.?We know nothing about the plumage changes of San-ford's ptarmigan, except that a series of two adult males and sixadult females, collected by Donald H. Stevenson on September 18,1921, now in the Biological Survey collection, show a postbreedingplumage quite different from the June breeding plumage. In themale this is darker, browner, or redder than the breeding plumage;the prevailing color of the breast, head, neck, and flanks is from
" tawny " to " ochraceous-tawny," instead of " cinnamon-buff " mixedwith pale grayish buffs, as in the June birds; on the upper partsthe tawny shades are much more heavily peppered, variegated, orbarred with black, entirely unlike June birds. The differences aresimilar in the female; the "cinnamon-buff" feathers, barred withblack, of the June plumage are being replaced by white feathers on
KELLOGG 'S PTARMIGAN 227the belly, and by " tawny " or " ochraceous-tawny " feathers on thebreast and flanks, more finely barred or peppered with black ordusky; the upper parts are also more tawny with finer barring orpeppering and with more black than in June birds. Breedingfemales have no peppered feathers and no white on the belly. Ajuvenal bird, collected at the same time and place, is like the adultfemale, but the colors are duller.LAGOPUS RUPESTRIS KELLOGGAE GrinnellKELLOGG'S PTARMIGANHABITSThe name kelloggae. " Montague rock ptarmigan," was firstapplied by Dr. Joseph Grinnell (1910) to a bird that he describedas a new subspecies from Montague Island, Prince William Sound,Alaska. But a later and more extensive study by Harry S. Swarth(1926) has shown that the specimens on which Doctor Grinnell'sname is based are merety variants toward dixoni of a race, distinctfrom rupestris, that inhabits the whole of northern and westernAlaska. He says of the characters and distribution of the race henow calls kelloggae:The notable feature of this bird is its bright ruddy tone of coloration, acharacter that is evident in both sexes and in all stages of the summer plum-ages. As compared with rupestris, the general tone of color throughout isbrighter and more reddish, and there is notable restriction of the dark areason individual feathers.The extreme manifestation of this race is reached on the northwestern andnorthern coast of Alaska, it occupies practically the whole of the Alaskanmainland, and it extends eastward of Alaska along the Arctic coast for somedistance. In the latter region the duller color of specimens from BaillieIsland, Coronation Gulf, and Bathurst Inlet, is to be interpreted, to my mind.as indicative of intergradation with rupestris.Southeastward there is intergradation again with rupestris as occurring inBritish Columbia, about at the Alaska-Yukon boundary line. A series of seven-teen skins from the vicinity of Eagle (U. S. Biol. Surv. coll.), in the upper Yukonregion, demonstrates such intergradation satisfactorily. Certain selected skinsfrom this series and from the British Columbia aggregation are hardly to bedistinguished, and none of the Eagle specimens shows the extreme of ruddinessthat is seen in Alaskan birds from more northern points. The Eagle series asa whole, however, certainly belongs with the northern Alaska subspecies ratherthan with rupestris. On the southern coast there is apparent intergradationwith dixoni, as shown by skins from Kodiak Island, Seward, and Prince WilliamSound.According to the latest revision of this species by P. A. Taverner(1929), as recognized in the new American Ornithologists' UnionCheck List, the range of this form is extended eastward along thearctic coast and islands to western Greenland, north of Disco.
228 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMAs the habits of this ptarmigan do not differ materially fromthose of other rock ptarmigan, I shall not attempt to duplicate whatI have already written on the species.According to Roderick MacFarlane (1908) the summer home ofthe rock ptarmigan in northern Canadaconsists of vast plains or steppes of a flat or undulating character, diversifiedby some small lakes and gently sloping eminences, not dissimilar in appearanceto portions of the North-West prairies.The greater part of the Barren Grounds is every season covered with shortgrasses, mosses, and small flowering plants, while patches of sedgy or peaty soiloccur at longer or shorter distances. On these, as well as along the smallerrivulets, river and lake banks, Labrador tea, crow-berries, and a few other kindsof berries, dwarf birch, willows, etc., grow.Referring to his trip to Hooper Bay, Alaska, Herbert W. Brandtwrites to me
:
Our first acquaintance with the rock ptarmigan was made in the upper soli-tudes of the Beaver Mountains high above timber line on April 6. On thesebald snow-beaten hills we found a number of straggling flocks ; one that num-bered about 20 birds contained only males, while the others were evidently matedcouples, banded together. On the wind-swept slopes were numerous mossyhummocks, and in the leeward side of this scant protection, the bird scoops outa snug little snowy igloo. This is its only retreat and roosting place during thelong cold winter, for Mr. Twitchell advises me that this hardy species seldomdescends to the larger willows and spruces, which line the streams below. Therock ptarmigan is a rather common summer resident in the Askinuk Mountains,where it seems to prefer the sterile open ridges in the vicinity of 1,000 feetin altitude.Nesting.?Brandt says on this subject :The contents of the nest of this species range from 6 to 11 eggs, but the usualnumber found is 8. The nesting site is so chosen that protection is affordedby a hummock, a small tree, or even a growth of frost-dried grass, but occa-sionally no concealment whatever is present. The lining of the nest consistssolely of surrounding materials, such as grasses, lichens, and moss, together witha few feathers.Eggs.?The rock ptarmigan, according to MacFarlane (1908), laysfewer eggs than the willow ptarmigan, the usual number being sixor seven and rarely more than nine. He describes the eggs, basedon the very large series collected by him, as follows
:
The eggs are ovate or short ovate in form, resembling the eggs of Lagopuslagopus considerably, both in colour and markings, but they average smaller.The majority are readily distinguished from those of the latter, the markings,as a rule, being smaller and better defined, and seldom running into indistinctand irregular blotches, as is frequently the case in the eggs of that species.The ground colour ranges from a pale cream to a decided yellowish-buff, andin many specimens this is entirely hidden by a vinaceous rul'ous suffusion.The spots and blotches range from a dark clove-brown to a dark claret-red,with paler coloured edgings ; they are of various sizes, from the size of a buck-shot to that of No. 10 shot, and are irregularly distributed over the egg.
KELLOGG 's PTARMIGAN 229Brandt gives a very good description of the eggs as follows
:
The egg of the rock ptarmigan is ovate to elongate ovate in shape andhas slight to considerable luster, which apparently increases as the eggincubates. The surface is smooth and greasy, and the sturdy shell stronglyresists the drill. The vivid markings on this beautiful egg are so numerous thatthey often all but envelope the pale ground color and produce rich-mottleddecoration that gives the egg a noteworthy appearance. These spots are dis-tributed evenly over the surface, except that often a confluent cap intensifiesthe color at the larger end. The eggs laid by one bird are similar in shape,plan of markings, and coloration, but seldom are there two sets from differentparents exactly alike. The inconspicuous ground color follows the paler buffsand creams, often with a reddish suggestion ; shell pink, pinkish buff, or creamcolor is often observed, while many eggs are still lighter than these palecolors. The markings range in size from the smallest spots to those approach-ing thumb-nail in size and are more or less confluent. As a rule, the largerthe spots are in size, the more the ground is shown. These markings whendry are blackish brown, but where the pigment has been sufficiently thinned,the color ranges from walnut brown and maroon to blackish brown. Underly-ing spots apparently are not present.Behavior.?Doctor Grinnell (1900) says of his experience withthis ptarmigan in the Kotzebue Sound region
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I first met with this species on September 17, 1898, about the summitof the Jade Mountains on the north side of the Kowak Valley. On thatday I saw three flocks of 6, 7, and 20 birds, respectively. In each case theywere flushed from ridges at some distance, and were probably feeding onheath and blue-berries, which fairly covered the ground on favorable slopes.At. a distance the birds appeared to be entirely white, at this date, thoughno specimens were obtained. I rather think the summer plumage of theRock Ptarmigan is of much shorter duration than that of the Willow Ptarmiganin the lowlands. The Rock Ptarmigan, according to my experience, are con-fined exclusively to the higher hill-tops and mountains in summer, and at suchelevations the snow remains later in the spring and comes much earlier inthe fall than in the valley, leaving a very brief summer. No Rock Ptarmiganwere detected in the Kowak Valley until February 11th. On account of thelight snow-fall in the early part of the winter, they probably found sufficientforage on the mountain sides up to this date. However, during March andApril flocks of from a dozen to a hundred were often met with in the lowlands.These flocks could be traced up by following their tracks, especially if thesnow was freshly fallen or laid by the wind. Then tracks of a large flockof Rock Ptarmigan would form a broad swath and extend across the tundrafor miles, the individual lines of tracks zigzagging back and forth so as totake in every willow twig or bunch of grass sticking up through the snow,but all tending in the same general direction. The birds, when on thesefeeding marches, apparently seldom take flight unless disturbed, and I havefollowed these roads from one set of " forms " in the snow, where the birdshad passed the preceding night to the second set of " forms " of the succeedingnight, and then finally found them, doubtless on their second day's walk with-out taking flight; except occasional individuals left behind. The tracks ofthe Rock Ptarmigan are easily distinguishable from those of the Willow Ptarmi-gan by their much smaller size and the shorter strides; and they seem not to74564?32 16
230 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbe in the habit of dragging their middle toes over the ground at each step.as evidenced by the tracks in the case of the Willow Ptarmigan,W. Sprague Brooks (1915) says:The males are quite pugnacious, when in flocks, often pursuing each otherand going through antics suggesting the young males of domestic fowls.Rock Ptarmigan exhibit considerable curiosity at times, a trait I have notnoticed in the Willow Ptarmigan. When one of its kind is dead or woundedthe rest frequently show great concern and interest in the unfortunate one.Many times while walking over the tundra I would be startled by therattling call of a male Rock Ptarmigan, and turning about see him alight withina few yards of me with tail spread and eye wattles erect. After strutting aboutand "showing off" a moment he would busy himself searching for food asthough no man were in the country. In the winter plumage the males arevery beautiful.Voice.?Dr. E. W. Nelson (1887) came across a pair of these birdsnear St. Michael, of which he writes
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They allowed me to approach within 20 feet, and paid no attention beyondlooking curiously at me as I walked slowly along. The suspicion of the malebeing slightly excited, he uttered a low, rolling or whirring sound, like thatproduced by rolling the end of the tongue. The female answered with a low,clear yop-yop, with a peculiar intonation, strikingly like that of the female hen-turkey, except it was much lower. When we were about 15 feet from the birds,they stood looking at us for a moment with a pretty air of innocent curiosity,and then, without showing the slightest signs of alarm, arose and flew off to thehill-side, a hundred yards or more away.Migrations.?Nelson says of their migratory movements
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During the entire year these birds are resident north at least to BeringStraits, as I obtained specimens from that vicinity on one of my winter expe-ditions. In summer it extends still beyond this, to all portions of the countrycrossed by mountain chains and hills. In autumn, toward the last of Octoberand first of November, this bird unites with the common Ptarmigan in greatflocks, on the northern shore of Norton Sound, and migrates thence across thesound to Stuart's Island, thence reaching the mainland. The birds are fre-quently seen by the natives while they are passing Egg Island, on their way tothe island just mentioned. They are said to commence their flight just beforedark in the evening, and at this season, as mentioned under the preceding spe-cies, many are snared at the head of Norton Bay. In April the birds returnto the north, always traveling in the evening or night, as they do during theirautumnal migrations.LAGOPUS RUPESTRIS EVERMANNI ElliotEVERMANN'S PTARMIGANHABITSThe well-marked subspecies evermanni, the darkest colored of allthe ptarmigans, the males being almost black, is confined to AttuIsland, the westernmost of the Aleutian Islands, 1,400 miles west of
eveemann's ptarmigan 231Unalaska. Dr. D. G. Elliot (1896) in his original description of itobserves
:
The males of L. evermanni bear a certain resemblance to specimens of L.mutus, of the Eastern Hemisphere, where these have much black in theirplumage; but between Attu and the continent of Asia is found L. ridgwayi,a very distinct form from Bering Island, about 300 miles west of Attu. Thiswould seem to bar any possible relationship between L. evermanni and any con-tinental species, though it is a surprising fact, and one that can only betheorized upon and not thoroughly explained, that species which are closelyallied can be separated by many miles of sea and land, and yet retain theirspecific characteristics, though distinctly different species may be found occupy-ing interlying territory. This is one of the curiosities of geographical dis-tribution, ? the solution of which is probably beyond the power of man tofathom.Everyone who has visited Attu Island has remarked on the scarcityof this ptarmigan. Perhaps it may live so far up on the mountainsthat its favorite haunts have not been visited; this would fit mytheory that the dark-colored birds are mountain birds and thatthe light-colored birds are lowland birds. But perhaps the scarcitymay be due to persistent hunting by the inhabitants of ChichagofVillage or to the presence of blue foxes, which are very common here.Doctor Elliot had only seven specimens on which to base his de-scription. Austin H. Clark (1910) writes:I did not find this bird at all common on Attu, doubtless because I did notsucceed in locating its favorite haunts. During an entire day's trip over themountains on the right of the harbor, behind the town, and about the largelake at the summer encampment, only three were seen, one in the mountainsabove the lake and two in the lowlands between the town and the lake.All three were shot. On arriving at the ship one of the men told me hehad never seen ptarmigan so common as about the summit of the mountainsat the left of the harbor entrance. As he had had considerable experiencewith ptarmigan in seldom visited portions of Alaska, and was a reliable man,
'I arranged to visit the locality the next day with him as a guide in orderto obtain a series of this little known species. We started early and reachedthe place a little before noon, but, although the droppings of the birds wereextremely abundant everywhere, we saw none of the birds themselves. Just aswe were preparing to leave, after searching the whole district thoroughly, afine cock came flying over from one of the neighboring peaks and waspromptly secured. On our way back to the shore we saw one other whichwas chased for over a mile but without success.Hamilton M. Laing (1925) says:During our three days' stay at Attu Island only three ptarmigan were found.A single bird on April 20 bounded up from the shore and flew wildly away.Next day, which was warm and sunny, two single birds were seen sitting onthe brow of the bluff above the shore and both were secured. Even in lifethe difference between these birds and the previous forms was evident, thenew blackish feathering giving them a decided speckled appearance. A climbto one of the hilltops disclosed no evidence of the birds at higher elevations.
232 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMWe spent only a day and a half at Attu Island and collected eightptarmigan, one pair that I shot in the valley and six that Rollo H.Beck shot on the mountains; the bare and moss-covered rocky sidesof the mountains seem to be their favorite haunts; the eight birdssecured were all that we saw.Nothing seems to be known about the nesting habits or eggs ofEvermann's ptarmigan. All we know about molts and plumages isthat the male described by Doctor Elliot (1896) was just completingthe molt into the summer plumage on June 4, and that the males col-lected by Laing (1925) were just beginning this molt on April 21.Our birds, seven males and one female, were in full summer plumageon June 22 and 23, and the female had a bare patch on the belly,showing that she was incubating. We learned nothing further abouttheir habits. LAGOPUS LEUCURUS LEUCURUS (Richardson)NORTHERN WHITE-TAILED PTARMIGANHABITSThe name " northern " white-tailed ptarmigan, adopted in thenew American Ornithologists' Union Check List, seems hardly suit-able for this, the type race of the species, for the Kenai race, penin-sularis, ranges entirely north of it. Typical leucurus is the bird ofthe western Canadian mountains in British Columbia and Alberta.This form is darker, with more black in the plumage and has shorterwings and tail than the southern bird, altipetens. The southern birdbeing the best known race, the reader is referred to altipetens forthe principal life history of the species.Dr. Frank M. Chapman (1908), in his attractive account of thisbird in the Canadian Rockies, says
:
They are said not to descend below timberline during the summer, but wenoted a striking exception to this rule at Lake Louise, where numbers of themcame regularly to feed about the forest-surrounded stable. They were evidentlyattracted by the fallen grain and may have learned of this supply of foodduring the winter when the heavy snowfall drives them to lower levels.Referring to their behavior, he writes:The first evidence they gave of being aware of my presence, was to remainperfectly motionless, then, as I made no further advance, they attempted tocombine action with rigidity of pose and were almost successful in achievingthis impossible feat. With painful slowness, one foot was placed in advance ofthe other, at the rate of about three steps to the minute. If I drew so nearthat the birds seemed convinced that they were seen, the male assumed a morealert, bantamlike attitude, ducking his upraised head and flirting his tail asthough inviting me to conflict. The pose of the female was more henlike, andless aggressive. She showed virtually no concern when I was within three feetof her, feeding about the rocks, and even stopping to scratch her head. After an
NORTHERN WHITE-TAILED PTARMIGAN 233hour or two, the male became more accustomed to me, and seemed as much atease as his mate, uttering a low, crooning note suggesting that of a comfortablechicken on a sunny day. DISTRIBUTIONRange.?Alpine sections of southern Alaska, western Canada, andthe United States.The range of the white-tailed ptarmigan extends north to Alaska(Lake Clark, Savage Kiver, and Robertson River) ; Yukon (proba-bly La Pierre House) ; and southwestern Mackenzie (Nahanni Moun-tains). East to southwestern Mackenzie (Nahanni Mountains);western Alberta (Henry House, Laggan, and Sulphur Mountain)
;
western Montana (St. Marys Lake, Piegan Pass, and BeartoothMountain) ; Wyoming (Medicine Bow Mountains) ; Colorado(Mount Zirkel, Arapahoe Mountain, Longs Peak, Bald Mountain,James Peak, Breckenridge, St. Elmo, Cochetope Pass, and SummitPeak) ; and New Mexico (Costilla Peak, Taos Mountains, and MoraPass). South to northern New Mexico (Mora Pass) ; southwesternColorado (Dolores Mountain) ; and northwestern Oregon (MountJefferson). West to northwestern Oregon (Mount Jefferson andMount Hood) ; Washington (Mount St. Helena, Mount Rainier,Pyramid Peak, Cloudy Pass, Mount Sahale, and Mount Baker)British Columbia (Mount Arrowsmith, Delia Lake, NinemileMountain, Groundhog Mountain, head of the Iskut River, and Doch-da-on Creek) ; and Alaska (Admiralty Island, Hooniah, Glacier Bay,Valdez, Kenai Mountains, and Lake Clark).The white-tailed ptarmigan is confined entirely to mountainousregions, and it does not perform a migration comparable to that ofthe willow and rock ptarmigans. A slight vertical movementusually takes place in winter when the birds descend from the peaksand ridges to sheltered valleys in search of food.The range as described is for the entire species, which has beendivided into four subspecies. The " northern " white-tailed ptarmi-gan {Lagopus I. leucunis) is found from northern British Columbiaand central Alberta south to Vancouver Island; the Kenai white-tailed ptarmigan (Lagopus I. peninsularis) occurs from centralAlaska, northern Yukon, and Mackenzie south to the Cook Inletregion, Kenai Peninsula, and central Yukon ; the Washington white-tailed ptarmigan {Lagopus I. rainierensis) is found in the CascadeMountains of Washington ; and the southern white-tailed ptarmigan{Lagopus I. altipetens) occupies the Rocky Mountain region of theUnited States, from Montana to northern New Mexico.Egg dates.?Colorado : 14 records, June 19 to July 15 ; 7 records,June 26 to July 6.
234 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMLAGOPUS LEUCURUS PENINSULARIS ChapmanKENAI WHITE-TAILED PTARMIGANHABITSThe type of the local race of the white-tailed ptarmigan namedpeninsularis, one of a series of 26 specimens, was taken by J. D.Figgins in the Kenai Mountains, on the Kenai Peninsula, Alaska, onAugust 11, 1901. It is in the gray fall, or transition, plumage. Dr.Frank M. Chapman (1902) in his original description gives it thefollowing subspecific characters : " In nuptial plumage differs fromcorresponding phase of plumage of Lagopus leucurus in having theblack areas of great extent, the buff areas much paler. In fall,transition or ' preliminary ' plumage differs from similarly plum-aged specimens of Lagopus leucurus in being decidedly grayer."Chapman quotes from Mr. Figgins's notes, as follows
:
Reared far above all timber, tbese interesting birds must depend upon theircolor for protection at all times. Found only on the bleak barren grounds, noteven a blade of grass rises to offer them a retreat. Their color is an exactimitation of their rocky surroundings, and if the bird remains at rest it isimpossible to detect it though only a few feet distant. When approached theycrouch as closely to the ground as possible, usually near some small boulder,and remain thus while you are in motion, but if a stop is made they try tosteal away and in that way reveal themselves. As soon as a movement ismade they resume their former position. They are hard to flush, dependingrather upon their color for safety than their wings. A low cackling whentheir young are disturbed are the only notes I have heard. The food of thisptarmigan is berries and the leaves of small plants. The principal berryresembles our blueberry in appearance and remains fresh the year round,falling from the plant only when a new crop is grown.LAGOPUS LEUCURUS ALTIPETENS OsgoodSOUTHERN WHITE-TAILED PTARMIGANHABITSDr. Wilfred H. Osgood (1901) discovered that the white-tailedptarmigan of the Colorado mountains is subspecincally distinct fromthe bird of the mountains north of the United States and gave it thename altipetens. He gives as its characters : "Adult in fall plumagesimilar to Lagopus leucurus, but general color of upperparts buffinstead of gray; adult in summer plumage indistinguishable in colorfrom leucurus; wings and tail decidedly longer than in leucurus."The white-tailed ptarmigan is an alpine species, a permanent resi-dent in the high mountains, above timber line during most of the3rear at least. In the southern portion of its habitat it ranges from10,000 to 14,000 feet altitude and somewhat lower farther north. Itis the only ptarmigan known to breed within the limits of the United
SOUTHERN WHITE-TAILED PTARMIGAN 235States. Mrs. Florence M. Bailey (1918) gives the following attrac-tive description of its haunts in the Glacier National Park:Skirting an acre of snow, I zigzagged back and forth over the face of the
"ideal ptarmigan slope," open to swift-winged enemies, but by its broken sur-face and variety of colors affording a safe background for ptarmigan in themixed summer plumage. Even the wide expanse of slide rock was broken byoccasional dwarf evergreens and streaks of grass, and many of its red shaleswere patterned with lemon-yellow or curly-brown lichen covering deep ripplemarks. Above the main mass of slide was a wide grassy slope of soft yellow-Isti brown tones thai would soon match the brown of the ptarmigan. Abovethis the narrow outcropping ledges and stony slopes made a terraced Alpineflower garden, one of the gardens that are among the choicest of all nature'slavish gifts to man; this one, with its maturing ^vvd harvest, providing ver-itable grain fields for hungary bird and beast. Some of these Alpine terraceswere fairly white with the lovely low, wide-smiling Dryas octopetala. In otherplaces the beds of white were .-polled with the pink mossy cushions of Silencacaulis, while iii still others there were clumps of dwarf sedum, whose dark-red flowers and seed pods contrasting strikingly with their pale green leavesmight well attract the attention of furry vegetarians locating granaries, andmake good feeding grounds Cor the Arctic grouse.Nesting.?Bendire (1892) quotes A. W. Anthony as writing:In southern Colorado, where I have met with I his species, nesting must beginseme time from the first to the middle of June, as I have found young birds butan hour or so from the egg, from July 1 t<> the L8th. The nests I have seenwere located in the loose rocky debris of steep hillsides, a simple depression inthe short line grass which grows in small patches between the rocks above thetimber line. Although utterly devoid of protection from hush or shrub, sonearly does the sitting hint resemble the gray bowlders which surround heron every side that the discovery of the nest is due largely to accident. Whenincubating it is nearly impossible i'> flush the bird, according to my experi-ence. Twice have I escaped stepping upon a sitting Ptarmigan by only an inchor so, and once I reined in my horse at a time when another step would havecrushed out the life of a brood of nine chicks lint an hour or so from theegg. In this case the parent crouched at the horse's feet, and, though inmomentary danger of being stepped on, made no attempt to escape until I haddismounted and put out my hand to catch her. She then fluttered to the topof a rock a few feet distant, and watched me as I handled the young, con-stantly uttering low anxious protests. The chicks were still too young toescape, mere little awkward bunches of down that stumbled and fell over oneanother when they attempted to run.He quotes Doctor Coues's description of a nest as follows:The nest in its present state measures scarcely 5 inches in diameter by aboutan inch in depth. It thus seems rather small for the size of the bird, but isprobably somewhat compressed in transportation. The shape is saucer like,but with very little concavity of surface. The bottom is decidedly and reg-ularly convex in all directions, apparently fitting a considerable depression inthe ground. The outline is to all intents circular. The nest is rather closelymatted, the material interlacing it in all directions, and retains considerableconsistency. The material is chiefly fine dried-^rass stems; with these aremixed, however, a few small leaves and weed tops and quite a number offeathers. The latter, evidently those of the parent birds, are imbedded through-
236 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
out the substance of the nest, though more numerous upon its surface, where adozen or so are deposited ; there may have been some loose ones lost inhandling.Illustrating the perfectly concealing coloration of the bird on itsopen nest, Evan Lewis (1904) writes:On reaching timber-line a Junco was seen building, and a search was madefor a loose stone to mark the spot for a photograph when the set was com-plete. In the search I was just about to put my hand on a Ptarmigan whenI saw what it was. I then made two exposures with the small camera andleft the camera on top of a large rock to mark the spot, the nest being threesteps and one foot due south from the mark. I went to the cabin at the lakeand got the large camera and tripod. When I returned I took three rathershorter steps, as I supposed, and looked for the bird or its nest. For tenminutes I looked over the ground foot by foot. I could not believe my own eyesthat the bird was not there, yet I could not see her. At last I was about toreturn to the mark and step the ground over again, when a reflection from thebird's eye showed her to me just one foot from where I was standing.W. C. Bradbury (1915) had some similar experiences. Nests,which he had previously located and marked, he had difficulty in find-ing again even when standing within a few feet of them. One nestwas right beside a stone that his foot was on ; but it had been lightlycovered with grass when the bird left it. In one case he found thatonly three eggs had been deposited in a period of five days.Eggs.?The white-tailed ptarmigan has been credited with layingas many as 15 eggs and as few as 1 ; probably the usual numbers runfrom 6 to 8. In shape they vary from ovate to elongate-ovate, andthey have little or no gloss. They are quite unlike other ptarmigans'eggs and are colored more like small eggs of the dusky grouse. Theground color is usually " cartridge buff " or "pinkish buff " andrarely " cinnamon-buff." The lightest-colored eggs are sometimesnearly immaculate ; most of the eggs are more or less evenly coveredwith small spots or fine dots; some are more heavily marked withlarger spots or small blotches; but the ground is never largely con-cealed. The markings are in various shades of brown, usually verydark brown, " Vandyke brown " to " bone brown," but sometimes aslight as " snuff brown " or " tawny-olive." The markings are rarelyconcentrated into a ring. The measurements of 31 eggs average 42.9by 29.5 millimeters; the eggs showing the four extremes measure 49.3by 29.6, 44.7 by 32.3, 39 by 28.3, and 42.9 by 28.2 millimeters.Young.?Mrs. Bailey's (1918) account of the behavior of motherand young is worth quoting, as follows:Listening, I caught it again?the softest possible call of a mother ptarmigan
!
There she stood, only a few feet from me, hard to see except when in motion, sowell was she disguised by her buffy ground color finely streaked with gray.A round-bodied little grouse with a small head, she was surrounded by abrood of downy chicks, evidently just hatched, as their bills still held the sharpprojection for pipping the shell. Preoccupied with the task of looking after
SOUTHERN WHITE-TAILED PTARMIGAN 237
ner little family, as I talked reassuringly to her, she ignored my presence.Nothing must hurry the unaccustomed little feet, nothing must interfere withtheir needed rest. Talking softly she gradually drew the brood in underher motherly wings and sat there only a few yards from me, half closing oneeye in the sun and acting oblivious to all the world. Once the downy head ofa chick appeared between the fluffed-out feathers of her breast, and once shepreened her wing so she showed the white quills remaining from the whiteplumage of winter.Her bill opened and her throat palpitated as if she were thirsty, as shesat brooding the young, and I imagined that the last hours of hatching highabove water had been long and trying to the faithful mother. But thoughwater?clear cold mountain brooks?were below, no need of her own couldmake her careless of her little ones. Keeping up a motherly rhythmic cluck-uk-uk, cluck-uk-uk, interlarded with a variety of tender mother notes, sheled them down by almost imperceptible stages, slowly, gently, carefully, raisinga furry foot and sliding it along a little at a time, creeping low over theground with even tread, picking about as she went, while the little toddlersgradually learned the use of their feet. Like a brood of downy chickens,some were more yellowish, seme browner than others, but they all had darklines on head and body giving them a well-defined color pattern. Peepinglike little chickens, while their mother waited patiently for them they toddledaround, trying to hop over tiny stones and saving themselves from going ontheir bills by stretching out wee finny wings. As chickens just out of theshell instinctively pick up food from the ground, they gave little jabs atthe fuzzy anthers of the dryas, little knowing that pollen was the best foodthey could find, a rich protein food from which the bees make bee bread tofeed their larvae.Plumages.?In the small, downy chick of the white-tailed ptar-migan the crown, shoulders, central back, and central rump are
" tawny," bordered and sprinkled with black ; the rest of the headshades from buffy white on the forehead to dull white on the chinand throat, with black spots and bands on the front and sides of thehead in somewhat different patterns ; the rest of the upper parts arevariegated with pale buffs, grays, and black; the underparts aregrayish white, with a slight buffy tinge on the breast. As with allother grouse, the juvenal plumage appears very early, the wTings firstwhen the chick is very small; and the neck and head are the last tobe feathered.By the time that the young bird is half grown it is fully featheredin juvenal plumage. In this the crown is barred or mottled withblack, white, and pale buff ; the back and rump are mainly vermic-ulated or peppered with black on a grayish white to " cinnamon-buff " ground color, but some feathers show larger black areas; thescapulars, wing coverts, and tail coverts are similar, with more blackon the scapulars ; the chin and throat are grayish white, barred withdusky ; the breast and flanks are pale buff, barred with black, andthe belly is buffy white ; the two outer primaries are white and theothers dusky, the tail feathers are dusky, banded and mottled with
* l cinnamon-buff." This plumage is hardly complete in August,
238 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbefore the postjuvenal molt into the late summer plumage begins.This is similar to the preliminary winter, or tutelar, plumage ofadults. About a month later, in September, the molt into the pure-white winter plumage begins. These two molts effect a completeehange of plumage between August and October, except that the twoouter primaries are retained for a year, which serve to distinguishyoung birds from adults.What little material is available seems to indicate that the moltsand plumages of adults are similar to those of other ptarmigan. Apartial prenuptial molt of the contour plumage, head, neck, andback takes places from March to June. The late summer, or tutelar,plumage is assumed by a partial postnuptial molt beginning late inJuly; this is much grayer, more finely vermiculated, and with lessblack than the nuptial plumage, which is more heavily vermiculated,with more black spots, on a more " ochraceous-buff " ground color.Females are more buffy or ochraceous than males in both plumages.A supplementary molt produces, in September and October, thecomplete change into the pure-white winter plumage.Food.?Dr. Sylvester D. Judd (1905a) summarizes the food of thisptarmigan, as follows
:
During winter in Colorado, according to Professor Cooke, they subsist, like
'>ther ptarmigan, largely on willow buds. The stomachs of two birds collectedat Summitville, Colo., in January, 1891, at an altitude of 13,000 feet, were foundto contain bud twigs from one-third to one-half inch long, but the kind of bushfrom which they came could not be determined. Doctor Coues, quoting T. M.Trippe, states that the food of this bird is insects, leguminous flowers, and thebuds and leaves of pines and firs. According to Major Bendire, the flowers andleaves of marsh marigold (Caltha leptosepala) and the leaf buds and catkinsof the dwarf birch (Betula glantiulosa) are eaten. Dr. A. K. Fisher examinedthe stomachs of two downy chicks collected on Mount Rainier, Washington, andlound beetles and flowers of heather (Cassiope mertensiana) and those of asmall blueberry.Mrs. Bailey (1928) adds:The crop of one New Mexico specimen was filled mainly with leaves ofthe dwarf willow, and fruiting spikes of Polygonum viviparum, with one flowerof Geum rossii, while the gizzard held mainly Polygonum seeds, a few othersmall seeds, a few small grasshoppers, and other small insects.Behavior.?Quoting Trippe, Coues (1874) writes:In localities where it is seldom molested it is very tame, and I have beeninformed by persons whose word is worthy of belief, that they have fre-quently killed it with sticks. But when persistently persecuted, it soon be-comes wild, and leaves the range of a shot-gun with surprising quickness.After hunting several large flocks for three or four days, they grew so shythat it was difficult to approach within gunshot, although at first they hadbeen comparatively tame. Nimble of foot, the Ptarmigan frequently prefers torun away on the approach of danger, rather than take wing, running overthe rocks and leaping from point to point with great agility, stopping every
SOUTHERN WHITE-TAILED PTARMIGAN 239
little while to look at the object of alarm. I have sometimes chased themhalf a mile or more, over the rocky, craggy ridges of the main range, withoutbeing able to get within gunshot, or force them to take wing. The flight ofthe Ptarmigan is strong, rapid, and at times sustained for a considerabledistance, though usually they fly but a few hundred yards before alightingagain. It resembles that of the Prairie Hen, consisting of rapid flappings ofthe wings, alternating with the sailing flight of the latter bird. The noteis a loud cackle, somewhat like the Prairie Hen's, yet quite different; andwhen uttered by a large flock together, reminds one of the confused murmurand gabble of a flock of shore-birds about to take wing. It is a gregariousbird, associating in flocks throughout the year, except in the breeding season.The different broods gather together as soon as they are nearly grown, forminglarge flocks, sometimes of a hundred or more.Dr. D. G. Elliot (1897) says:They were not what may be called tame, unlike the Willow Grouse in thisrespect, but were always very uneasy at my presence, and ran about withuplifted tail as if uncertain which way to fly, but when they once got startedthere seemed to be no farther difficulty in their minds as to the proper direc-tion, which I noticed never led near where I stood. Sometimes I have seenthem light on the bare limbs of a stunted tree or large bush at the edgeof the timber line, where they stood perfectly motionless for quite a lengthof time, observing every movement I made, and then suddenly burst away withgreat speed, uttering a low cackle as they flew. They are very skillful inconcealing themselves, either squatting in the snow with only the head exposedto view, or else crouching behind some stone or large bowlder. In summertheir peculiar gray plumage assimilates so well to the hue of the groundand the moss-covered stones lying about in all directions that it is next toimpossible to perceive them, and at this period, especially during the breedingseason, they rarely move when approached, perhaps only going a few feeton one side to avoid being stepped upon.Denis Gale wrote to Major Bendire (1892) :Irrespective of season, as a general rule, a single bird will not flush unlessurged to it. During the summer months this is especially noticeable ; theywill only move out of your way when directly in your path, and close uponthem, by short tacks right and left, sidling off from you, at each tack chang-ing sides, moving quickest on the short run just before slowing up forthe turn. Two or more together are much more likely to flush, and if alarmedwhile flying will utter a quick repeated kock, kock, very like the note uttered byPediocaetes phasianellus campestris under similar circumstances.Game.?The white-tailed ptarmigan is a fine game bird for thosewho are hardy enough to stand the hard tramping necessary for itspursuit in the high mountains. Edwyn Sandys (1904) writes:Unlike many of its kin, this bird is not troubled with overconfidence in man,but is apt to fly smartly and present none too easy a mark. It is also quitea runner, and taken altogether, the " snow quail," as the miners call it, is a fitquarry for an expert, especially if he be a " tenderfoot," unused to Alpine workand the pure, thin air of the heights; for this ptarmigan is a lover of highaltitudes, seldom, if ever, being seen lower than five or six thousand feet.
240 BULLETIN 162, UNITED STATES NATIONAL MUSEUMFall.?Dr. George Bird Grinnell wrote to Major Bendire (1892)
:
In the autumn the birds are generally rather wild, and if nearly approachedbecome quite uneasy and run about, holding the tail elevated and looking verymuch like a white Fan-tail Pigeon. At this season the only cry that I haveheard is a sharp cackle like that of a frightened hen. This the bird begins toutter a short time before it takes wing, and continues it for quite a little whileafter having begun to fly.On the high plateaus where this bird is found the wind often blows witha tremendous sweep and is almost strong enough to throw down a man. Whensuch a wind is blowing the Ptarmigan dig out for themselves little nests orhollows in the snow banks, in which they lie with their heads toward the windand quite protected from it. Often on the rocky slopes where there is no snowthey may be seen lying crouched on the ground behind rocks or small stones,with their heads directed to the quarter from which the wind blows. Ifstartled from such a place they all take wing at once, looking like a flock ofwhite Pigeons, and fly for a short distance, but as soon as they touch the groundagain they throw themselves flat on it behind the most convenient shelter.Winter.?Sandj^s (1904) says:At the approach of winter the broods of a district frequently join forces ina packlike formation. I have seen 40 or 50 together, and heard the minersspeak of packs of several hundreds; this, however, is hearsay, and perhaps100 birds together would be a large pack. During rough weather the birdswill go under the snow ; in fact, they will hide in snow whenever it is available.During severe winters and when the snow is so deep that their foodsupply is covered, these ptarmigan desert their normal home abovetimber line and descend into the edges of the spruce timber on thehillsides or into the creek bottoms among the willows, where theycan find food and shelter.LAGOPUS LEUCURUS RAINIERENSIS TaylorRAINIER WHITE-TAILED PTARMIGANHABITSDr. Walter P. Taylor (1920) described and named this dark racefrom a series of eight adults and four young birds collected on MountEainier, Wash. He says that adults in nuptial plumage aresimilar to Lagopus leucurus leucurus, but dark areas more blackish ; buffy washover light areas not so consistently present, and when present paler.Comparison with specimens of Lagopus leucurus leucurus from Moose Pass,British Columbia, Moose Pass, Alberta, and Moose Branch of Smoky River,Alberta (one specimen from Henry House, Alberta), practically topotypes ofleucurus, all in nuptial plumage, indicates that the dark areas in rainierensisaverage more blackish than in leucurus. In the latter the shade is close tomummy brown (Ridgway, Color Standards, 1912), while in rainierensis theyapproximate one of the darker shades of blackish brown. The buffy portionsof the feathers in rainierensis are paler than in leucurus, being, in the former,near light ochraceous-buff, while closer in the latter to ochraceous or ochraceous-tawny.
RAINIER WHITE-TAILED PTARMIGAN 241There is no evidence that this race differs materially in habitsfrom other white-tailed ptarmigan. Doctor Taylor (1927) says thatit ranges in altitude between 6,000 and 8,000 feetin the Arctic-Alpine Zone all around the mountain, rarely dropping down intoupper Hudsonian, except in winter.The ptarmigan finds congenial surroundings on the pumice slopes at andabove timber line on Mount Rainier. Here the combination of bright light,freezing boreal blasts, dwarfed and wind-blown vegetation, and extensive snowand ice fields provide Arctic conditions in fact.Nesting.?Taylor describes a nest as follows
:
Consciously or unconsciously the ptarmigan had here selected a nest sitewhich for grandeur of outlook would be hard to equal. The nest was on theground on the south side of a rock on a southwest slope of Pyramid Peak, atan altitude of about 6,100 feet, where the hardy conifers, dwarfed and mattedin their unequal struggles with the elements, had at length given up com-pletely. At first glance the nest did not appear to have been specially con-structed ; but it was later found that a hollow had been excavated and filledwith dried vegetation. The nest itself was comfortably dry, though the soilbelow was damp, and doubtless usually frozen solid. A few feathers were scat-tered about the nest. Plants in the immediate vicinity were the red and whiteheathers and the Siberian juniper. There were five eggs, one infertile, oneaddled, and three in various stages up to approximately 10 days' development.Voice.?Taylor gives the best description of notes of this speciesthat I have seen. Referring to the female, he writes
:
While on the nest she several times uttered a hoot, hoot, hoot, hoot, hoot,a low, almost inaudible, soothing series of grouse-like notes. Another noteuttered by the ptarmigan as she turned the eggs was a cluck! cluck! muchresembling the call of a barnyard fowl to her little chickens. When awayfrom the nest she stalked about rather slowly, occasionally jerking back herhead in a characteristic manner, and regarded with evident anxiety the nestsite about which we were grouped. If we approached the nest too closely thegentle bird was not a little perturbed and warned us port! perrt! or some-times pit-prrrrt! prrrrtlOf the male he says
:
One of the birds, a cock, remained in the vicinity for upward of an hour,watching the observer and calling for his mate. His principal call was some-think like Su-squeek ! cluck-luck-a-luck, cluck-luck-a-luck! or sometimes Squeek!cluck! cluck! cluck! cluck! Cluck lucka-lucka-lucka-cluck ! Occasionally thecall is blurred at the end, Cluckrrrrrrrr ! The squeaking note, which is ofstaccato quality, high pitched and conspicuous, may be twice repeated, as fol-lows: Squeek! chuck chuck chuck chuck chuck chuck chuck Squeek! chuckchuck chuck! A call somewhat resembling that of the red-shafted flicker washeai-d yip! yip! yip! yip! Another combination Yip, yip squeech! yip! yip I Awarning note may be represented by the syllable chirrr chirr chirr chirr chirrchirr.
242 BULLETIN 162, UNITED STATES NATIONAL MUSEUMTYMPANUCHUS CUPIDO AMERICANUS (Rcichcnbach)GREATER PRAIRIE CHICKENHABITSContributed by Alfred Otto GrossThe prairie chicken ranks first among the game birds of the prairiesof our Middle West. It is to the prairie what the ruffed grouse isto the wooded sections of the country. As intensive agriculturepushed to all sections of the range of the prairie chicken and asinterest in hunting increased, this fine game bird at one time seemedin grave danger of following the course taken by the heath hen,to extinction as a game bird. In fact, it is gone from much of itsformer range, and its original numbers have been greatly reducedin practically the entire area of its distribution.Because market hunting has been made a thing of the past sincethe beginning of the twentieth century and also because of the in-creasing restrictions on hunting by State departments, as well asvarious effective conservation programs, the prairie chicken is nowholding its own and is increasing its numbers in many sections ofits present range. Another hopeful sign is the fact that it has beenextending its range to the northwest, and to-day the species is wellrepresented on the prairies of Manitoba and is gradually spreadingwestward through Saskatchewan and Alberta, where formerly itdid not exist.The State Department of Conservation of Wisconsin has under-taken a comprehensive investigation of the prairie chicken to ascer-tain all the facts that affect its life, with the expectation that the de-partment will be able to carrj? on a more effective program of con-servation. Until the fundamental facts in the biology of our gamebirds are clearly known, conservation commissions will be handi-capped in handling questions of game legislation and gamemanagement.Prairie chickens, in common with other grouse, go through definitecycles of numbers. The problem of fluctuations in numbers of vari-ous species of wild life is not yet definitely solved, but work on itin relation to the ruffed grouse is being undertaken by many institu-tions and individuals in different parts of the country; hence thereare excellent prospects of this work being brought to a successfulconclusion.The weather condition during the nesting season, especially duringthe height of the hatching period, is so important that it is frequentlythe determining factor in the number of young birds available forthe next hunting season. A series of torrential cloud-bursts fol-
GKEATER PRAIRIE CHICKEN 243lowed by long, cold, rainy spells during the first two weeks of Junewill cause hundreds of broods to perish.During severe winters, especially when deep snows cover theground, the birds are severely pressed to obtain enough food. Asuccessful attempt has been made in Wisconsin to relieve this con-dition by the establishment of winter feeding stations. Crops ofbuckwheat and other grains are planted and left in the field to pro-vide food to tide over the birds during these severe times.One of the major problems involved in the conservation of theprairie chicken is the menacing fires that have swept the prairieregions during the nesting season of the birds. A fire at this timewill destroy hundreds of nesting birds and their nests and eggs andin the course of a few hours undo the work of years of conservationwork. Fires in fall destroy quantities of prairie-chicken food andthe much-needed cover, without which the birds are left exposed topredators. The encroachment upon the breeding and feeding areaby agriculture has long been recognized as a factor that has affectedthe status of the prairie chicken in the Middle West. This unfavor-able situation is being relieved somewhat by the establishment oflarge State game preserves, on which the birds are given absoluteprotection and where conditions are systematically improved for thebirds. The maintenance of winter feeding stations has been espe-cially helpful in tiding the birds over the times when deep snowscover most of the normal food supply.Intensive hunting has done much toward decimating the numbersof prairie chickens. The automobile and the fine modern roads haveall been in favor of the hunter and against the birds.Predators in their relation to game birds are important, but thevalue of vermin control is frequently overestimated. The wholesalekilling of all hawks and owls, for example, should be rigidly avoided,for in the past this practice has actually acted as a boomerang to theobjective of conservation of game birds.Diseases and parasites of birds have not been well known in thepast, but they are now becoming to be recognized as importantfactors in the life of our game birds. Under ordinary conditions,diseases and parasites may be of minor importance, but just as soonas the vitality and normal resistance of the birds are lowered by aseries of adverse conditions, such as severe weather and scarcity offood, diseases and parasites manifest themselves and become ofprime importance. It is the exceptional bird that is not parasitized,and hence this menace is ever present. There is also danger ofinfectious diseases, such as blackhead, which has been found toaffect the prairie chicken and which figured in the decline of theheath hen. It is highly probable that the cycles in the grousepopulation are primarily dependent on some disease, either in itself
244 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
or in combination with other factors. The evidence points to theconclusion that the vast majority of the parasites and diseases ofour game birds have been introduced through poultry and exoticgame birds. It is apparent that the adaptation of the prairie chickento the conditions imposed by civilization is not a simple matter. Inthis adjustment, birds such as the prairie chicken will require muchorganized assistance on the part of conservation commissions,sportsmen, and bird lovers.Courtship.?The courtship of the prairie chicken generally beginsduring the first warm days that lay bare the open fields of thewinter's accumulation of snow. Though an early beginning may bemade, the courtship does not reach its maximum until the latter partof April or the first of May, when companies of prairie chickensmajr be seen collected in favorable, often traditional, spots of theopen fields throughout the prairie-chicken country.O. M. Bryens, of Luce County, Mich., reports the first booming,or " crowing," as it is generally termed in the Middle West, asMarch 22, 1925; March 13, 1926; April 17, 1928; and March 27,1929. According to Prof. W. W. Cooke, the booming of the prairiechicken was from March 7, at Caddo, Okla., to March 24, at Barton,N. Dak.The courtship season continues through the month of May, butthe vigor of its execution diminishes and the number of individualsthat take part decreases as the sets of eggs hidden in the grasses ofthe prairie are completed and the domestic duties of incubation onthe part of the female begin.A few .birds were still booming on the prairies near Hancock,Wis., when I arrived there the first week of June, 1929, and birdswere also booming the second week of June, 1930, in various partsof Wood and Waushara Counties, central Wisconsin. I heard nobooming and obtained no authentic accounts of birds booming afterthe second week of June.The courtship of the prairie chicken is similar to that describedfor the heath hen, but since these performances are such an import-ant part of the behavior of this bird a number of interpretations asmade by other observers are of great interest.Dr. Frank M. Chapman (1908) has given us a very vivid account ofthe prairie chicken as he observed it in the sandhills of Nebraska :At short range the bird's note suggested the mellow resonant tone of a ket-tledrum, and when bird after bird, all still unseen, uttered its truly startlingcall, the very earth echoed with a continuous roar. As a rule, each bird haditsi own stand separated by about ten yards from that of its neighbor. The boomis apparently a challenge. It is preceded by a little dance in which the bird'sfeet pat the ground so rapidly as to produce a rolling sound. This cannotbe heard for a greater distance than 30 yards. It is immediately followed by
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the inflation of the great orange air sacks at the side of the neck, which puffout as quickly as a child's toy balloon whistle; the tail is erect and widelyspread, the wings drooped, the neck tufts are raised straight upward, givingthe bird a singularly devilish look, then with a convulsive movement of thelowered head, the boom is jerked out and at its conclusion the air sacks havebecome deflated.One might imagine after so violent a performance the bird would feel acertain sense of exhaustion or at least quiescent relief, but his excess of vitalityseeks still other outlets; uttering hen-like calls and cacks he suddenly springsa foot or more straight into the air, whirling about as though he were sufferingfrom a combined attack of epilepsy and St. Vitus dance. But all this activityis only a prelude to the grand finale of actual combat. Like a strutting turkeycock, the neighboring birds go towards each other by short little runs, headdown, the orange eye-brow expanded and evident pouch inflated, neck tufts,and tail straight up, and looking like headless birds with two tails. Theirmating is followed by no make-believe duel but an actual clash of wings. Utter-ing a low, whining note they fight as viciously as game cocks ; and the number offeathers left on the ground testifies to effective use of bill and claws.First bird called at 4.40 and by seven o'clock the performance was pi-acticallyover.A prairie cock when in the lists is a strikingly conspicuous creature ; hewears no adornment which cannot be concealed at a moment's notice. Thesight of a passing hawk changes the grotesque beplumed, be-oranged bird intoan almost invisible squatting brownish lump, so quickly can the feathers bedropped and air sack deflated. With woodland birds so great a change isunnecessary, but the prairie hen can hide only under its own feathers.H. L. Stoddard (1922), in notes from southern Wisconsin, saysthat " the ' cooing ground ' at the sandy west end of Sauk Prairie hasbeen used each spring for over 30 years, the birds always using thesame knoll whether in rye, stubble, or grown to grass." Cooingstarted early in March and continued well into June. The birdsarrived early; some were on the grounds before daylight, but onother occasions the bulk came shortly after daylight. The cooingis a " resonant C-A-O-O-O-O-O, H-O-O, H-O-O, rising and in thesame tones as do re mi of the musical scale." This note carries along distance. " I have heard it over water when the nearest landwas nearly 2 miles away." Two cackling calls were like that ofroosters, " one a loud Ka-Ka-Ka-Ka-a-a-a-a and the other a long-drawn q-u-a-h."Alexander Sprunt, jr., says in his notes:I witnessed a dance one afternoon of five pairs of the birds which came toa sudden end in a strange manner, and one which would thrill the heart of anornithologist. Lying ensconced behind a log, I was reveling in the eaves-dropping act of witnessing the ludicrous antics of five males, who, with airsacs inflated, tails spread, and wings drooping, were bobbing up and down likecorn in a hopper, about an admiring group of hens. The booming was intenseand incessant, all having something to say at once. Suddenly, without amoment's warning, a huge snowy owl appeared from behind a low ridge at thefar edge of the dancing ground and on widespread wings shot low over the74564?32 17
246 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMdancers at a height of about 3 feet. Like so many feathered bombs, thechickens scattered to right and left, and in an instant the dancing ground wasdeserted. No attempt was made by the disturber to follow any of the revelers.Nesting.?The nest of the prairie chicken is invariably on theground, but the character of the vegetation in which it is built revealsconsiderable individual variation. Generally the nesting site isamong grasses and weeds or low shrubbery in very open situations,but sometimes it may be adjacent to trees and woodlands and in rareinstances may be surrounded by trees of considerable size. The vege-tation about the nest is usually very thick and effectively concealsthe eggs and the incubating bird from view. It also serves as aprotection from extremes of temperature. There are sometimeskilling frosts during the nesting season, in May, and there are manydays in June when the heat is great enough to kill the embryos if leftexposed to the direct rays of the sun for any great length of time.The nest is placed in a natural hollow of the ground, or a slightexcavation may be made by the bird by scratching out the looseearth and then molding the cavity to conform to the size and shapeof the body. In this cavity the bird places a scant quantity of nest-ing material, in some instances the nest lining being little more thanthe bent-over blades and weeds growing about the structure.The following descriptions serve to represent the character of thenesting site as well as the nature and construction of the nests builtin three different types and situations located in central Wisconsin
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A prairie-chicken nest containing 17 eggs was found 4 miles south-east of Bancroft, Portage County, on June 4, 1929, in a small clear-ing of a jack-pine grove, the trees of which ranged from 35 to 50feet in height. The trees of the clearing had been cut the yearbefore and piles of brush left in place. Some of the brush was moreor less hidden by the rank growth of grass and weeds which hadsprung up around it. The nest, built in a very shallow depression.4 centimeters deep and 18 by 20 centimeters in diameter, was nearone of the piles of brush. Some of the smaller branches were archedover the nest when found. It was protected by the brush on oneside, but on the other it wTas well exposed to view, a condition veryfavorable for observations and photography from the blind that waslater placed in position. Several of the pine trees were so near thatthey provided shade for the nest during certain hours of the day.Although this nest was not built in the usual surroundings, it isinteresting to note that there were extensive marshlands all aboutthe site.In the drainage area of Wisconsin there are isolated areas of highground locally called " islands," a name originating from the daysbefore the drainage ditches, when they were in reality " islands "
GREATER PRAIRIE CHICKEN" 247during the rainy period of the year. One such island of about 100acres near Bancroft, Wis., is called Prairie Chicken Island, becausethese birds have always lived and nested there in unusually largenumbers.Since agriculture has been encroaching on the original habitat ofthe prairie chicken, many of the birds have adapted themselves withmore or less success to conditions created by farming activities. Anest of the prairie chicken was found on June 24, 1929, on a farm10 miles northeast of Friendship, Adams County, Wis. The nestwas on rather high meadowland and was completely surroundedwith a luxuriant growth of clover, timothy, and other grasses. Theeggs were well concealed by a beautiful canopy of red-cloverblossoms. The bowl of the nest measured 15 by 19 centimeters indiameter but only 4 centimeters in depth. The lining of the nestconsisted of grasses and weed stems, all apparently picked up fromthe vicinity of the nest. The clover field was bordered on one sideby a low, wet marsh, in which various sedges and rushes prevailed
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on the other side was a cleared area being used as a potato field.Other individuals cling to the more remote prairie districts awayfrom farms, often in situations occupied in common with the sharp-tailed grouse. A nest containing 11 eggs was found in such a situa-tion in Portage County on June 17, 1929. It was placed amongtufts of sedge (Carex stricta). The nesting cavity was 18 by 21centimeters in diameter and 8 centimeters deep. The lining of thenest was made up entirely of sedge, among which were a fewfeathers of the incubating bird. Near the nesting site was a low,wet marshy area, and on the other side there were thickets of smallwillows and poplars. No farmland or farm buildings were within3 miles of this location.Eggs.?The background color of the eggs varies from a " darkolive-buff " to a " grayish olive " tint. Most of the eggs are dottedwith many fine and a few larger spots of " sepia." The spottingvaries considerably in different eggs from those with scarcely anymarks to those with many fine dots and 20 or more well-defined spotsranging from 1 to 2 millimeters in diameter. Sets of eggs found inopen prairie regions seemed to have less spotting and the color ofthe spots was a " vinaceous-buff " rather than the dark markings of
" sepia " present on eggs found in nests located in the woodedsections of the State. This difference in coloration, however, isprobably a mere coincidence and is not to be correlated with a con-sistent difference in habitat.Nine sets comprising 100 eggs, all found in central Wisconsinduring the summer of 1929, were weighed and measured. Theaverage long diameter was 44.86 millimeters; average short diameter,
248 BULLETIN 162, UNITED STATES NATIONAL MUSEUM33.59 millimeters. The eggs showing the four extremes measured49.1 by 34.1, 45 by 35.3, 40.5 by 32.9, and 40.8 by 30.2 millimeters.The laying of the eggs as determined by studies of captive birdsextends over a period of days equal to nearly twice the number ofeggs in the set. Apparently approximately the same ratio holds forbirds living under natural conditions. In a nest in Adams County,Wis., the first egg was found on May 5, 1929 ; on May 12 there were5 eggs; and the set of 11 eggs was completed on May 22. The timerequired to complete a set of eggs depends on a number of factors,such as the condition of the weather, the health of the bird, and theavailable food supply. The laying of the eggs is not necessarilyon alternate days, but more apt to be very irregular. Certain eggs ofthe set are laid on successive days to be followed by a lapse of twodays before the next egg is deposited.During the laying period of one individual under observationthe bird covered the rather exposed eggs with nesting material beforeshe left them. This instinctive habit may be for the purpose of con-cealment or for protection from extremes of temperature or forboth.The number of eggs layed by the prairie chicken based on studiesmade of 40 nests, in which the number of eggs was presumably com-plete, varied from 7 to 17. The average number was 11.5 eggs perset. A nest containing 21 eggs has been reported, but such unusualsets probably represent the eggs of two females using the same nest.The dates when the nests were found are of some interest to thosewho may desire to know the probable date when a nest of the prairiechicken may be found. Of 41 nests in which the date of findingwas recorded, 20 were found in May, 18 in June, and 3 in July. Theearliest date was of a nest found on May 5, when it contained a singleegg, and the latest was a nest found July 10 that still contained eggson July 15, the last time it was visited. The average date when the41 nests were found is June 3.Judging from these results we may say the last week of May andfirst week of June are the times when one may expect to find thelargest numbers of nests of the prairie chicken in central Wisconsin.Young.?The incubation of the eggs is performed by the female.Soon after the courtship season the male goes into retirement andundergoes the ordeal of molting. I have never seen the male birdnear the nest, nor have I ever observed him participating in the careof the young.Incubation begins soon after the set of eggs is completed, but incertain cases where we have found live embryos in eggs left in thenest after the rest of the brood had departed it was evident thatincubation had started before these eggs were laid. Sometimes an
GREATER PRAIRIE CHICKEN 249
unusual disturbance about the nest may delay the start of incuba-tion for several days, thus making the determination of the incuba-tion period under normal conditions in the field a difficult task. Thisaccounts for discrepancies in the determinations made by differentobservers, which vary from 21 to 28 days as the incubation period forthe prairie chicken. In the case of a nest under continuous observa-tion, incubation began on May 22 and the eggs hatched on June 14,establishing an incubation record of 23 days. The incubation periodfor the eggs of the closely related heath hen, as determined by Dr.George W. Field, is 24 days.After incubation is started the prairie chicken, under normal con-ditions, remains faithful to her duties through the vicissitudes ofweather, storms, and dangers of attacks from enemies. Unlessflushed from the nest she leaves only for very short intervals to feed,usually at dawn or late in the afternoon about sunset, times when theeggs will not be exposed to extreme heat. Excessive heat is verydestructive to the embryos, and great care must be exercised in flush-ing birds away from nests in which the protecting vegetation hasbeen removed for purposes of photography. The birds that left thenest normally slipped off it quietly and made no attempt to cover theeggs, as they did during the laying period when the nest was aban-doned for a longer time. One bird, after nervously surveying hersurroundings, sneaked off the nest and walked briskly in a crouchedposition until she had gone several yards from the nest. There shehesitated, elevated her head, and looked about as if to determinewhether her movements had attracted any attention. She then casu-ally nipped at the grasses as she walked along, and finally whenabout 25 yards away from the nest she arose with a loud whir ofwings and disappeared in the scrub pines, where she probably foundat least a part of her meal. After an absence of half an hour sheflew into view, circled the nest, and alighted in the tall grass 15 to30 yards away from the nest. At first she crouched in the grasscompletely concealed from view, but after being assured all was wellshe walked along stealthily though not directly toward the nest. Itwas not possible to keep her in view at all times, but now and thenshe would come to an open place and from this vantage point morecarefully scrutinize the surroundings. Sometimes she completely en-circled the nest and blind with a wide radius and frequently retracedher steps to make a careful inspection in order to satisfy herself thatno spying enemy was near. One could not be sure whether this be-havior was prompted by apprehension of harm to herself, fear ofrevealing the presence of the nest, or to both. After these maneu-vers, which usually took about 20 minutes, were completed, she ap-parently was assured, and then without hesitation she walked quickly
250 BULLETIN 10 2, UNITED STATES NATIONAL MUSEUMand directly to the nest. The position she assumed on the nestvaried, and one could never be sure whether she would be facingtoward the blind or away from it. The ruffed grouse, which buildsits nest at the base of a stump, a log, or tree, invariably faces awayfrom the side of the nest thus protected from the sneaking approachof some prowling enemy. Prairie chickens, which nest in thick veg-etation, will usually face toward the side that is opened up for pur-poses of photography, a very desirable position for the photographer.Many of the prairie chickens studied exhibited extreme restless-ness and made much more of a task of incubation than does theruffed grouse. One continuously shifted her position during thecourse of the day; at other times she would pick aimlessly at thenesting materials, and not infrequently she would reach far out ofthe nest at some unsuspecting grasshopper or other insect thatchanced to alight on the tall grass. Certain females revealed a sen-sitive, nervous temperament and quickly responded to any stimulus,whether it was the caw of a crow or the hum of a distant tractor;and even the shadow of a passing cloud was sometimes sufficient tomake her respond. When startled she would frequently elevate herhead to command a wider view of her surroundings. If she caughtsight of the source of her alarm, such as the passing of a dog, shewould retract her head, become perfectly motionless, and retaina " frozen " position until the source of danger passed. If theanimal ventured too near she would fly off with a violent " whirrrr "of her wings, which was sure to attract, if not startle, the intruder.She would drop into the grass a short distance away, utter a sharpdistressing cry, and feign a wounded bird. After several repetitionsof the performance, until she had attracted the enemy away to asafe distance from the nest, she would sail gracefully away, leav-ing the bewildered creature behind. This behavior is common tomany birds, but the deception is remarkably well executed by theprairie chicken. I have seldom seen a prairie chicken try thesetactics with a human being; apparently they have learned that itis best in such cases to get out and away without the least delay.That the leaving of the nest when flushed by a human being is a quickperformance is revealed by moving pictures. The usual speed is 16frames per second in the ordinary moving pictures. It requiresonly three or four frames of the picture to show the bird until sheis away from the scene. This means that it requires only three-sixteenth to one-fourth of a second for the bird to leave the nest.As it is not possible in most cases definitely to establish the causeof nest destruction, it is necessary to depend on circumstantial evi-dence. In 1929, out of 12 nests studied in Wisconsin only threereached the period of hatching. The adudt birds of two of the nests
GREATER PRAIRIE CHICKEN" 251
were killed and the eggs destroyed, presumably by coyotes, as thetracks of the animals were found in the sandy soil around the nests,and the mass of feathers left behind seemed to indicate the workof such animals. One incubating prairie chicken was killed by ahorned owl; the eggs of another located in an open situation weredestroyed by crows, and in two other cases the eggs disappearedwithout a trace of the intruder. One nest was accidently destroyedby a farmer while plowing, and the eggs of two others were de-serted where it was problematical whether the incubating birds weretaken by some predacious bird or mammal or were merely frightenedaway in some manner.During the summer of 1930, 28 prairie chicken nests were found,of which 17 reached the period of hatching and the others failedfor one reason or another. Indirect evidence indicates that crowswere responsible for the destruction of three nests of eggs. One wasprobably broken up by a dog, four were drowned out by floods, inone case the bird was killed, presumably by a mink, within a fewfeet of the nest, in another case the embryos in the eggs were killedby exposure to the heat of the sun, and one nest was deserted. Com-bining the records of ,L929 and 1930, we have 40 nests of which 3hatched in 1929 and 17 in 1930, 20 in all, or an average of 50 per centfor all the nests observed.Every egg hatched of the three nests that reached the stage ofhatching in 1929. The 17 nests that succeeded in reaching the hatch-ing stage in 1930 contained 208 eggs. Twenty-nine eggs, or approxi-mately 14 per cent, failed to hatch. Only 8 of the 17 birds suc-ceeded in hatching every egg, and in 9 nests there were one or moresterile eggs, eggs with dead embryos, or both. Of 29 eggs that failedto hatch, 6 were sterile and 23 contained dead embryos. The lat-ter were killed by excessive heat of the sun or by failure of the eggsto hatch in time before the old bird left the nest with her young.During the summers of 1929 and 1930, the date of hatching wasnoted for 23 sets of eggs in the field or by special incubation. Ofthese, 3 hatched in May, 17 in June, and 3 in July. The earliest dateof which we have a record was May 29, 1930, and the latest July 7,1929. Nest No. 12 contained eggs on July 15, 1929, but it was notpossible to record its hatching. The average date of hatching ofthe 23 nests was June 10. These records indicate that the majorityof the nests hatch during the first two weeks of June. The condi-tion of the weather at this time is a most important factor in thedetermination of the number of birds to be expected the followingseason. A long continuous cold rainy spell with cloud-bursts, such asis sometimes experienced in the Middle West during the first partof June, is certain to have a disastrous effect on the broods of youngbirds.
252 BULLETIN 162, UNITED STATES NATIONAL MUSEUMThe first eggs are pipped on about the twenty-second day afterincubation starts. The shell is slightly raised at the point on thecircumference between the blunt and pointed ends, but slightlynearer the larger end. The embryo inside the egg can be heard to
" peep " at this time, an event that greatly excites the mother andat once becomes a great stimulus in her behavior. It causes her toreach under her breast feathers to turn the eggs every few minutesand to exhibit a great deal of nervousness in her response to variousother stimuli.On the morning following the day on which the eggs are slightlypipped, all the eggs destined to hatch are pipped. In those eggswhere there was but a slight elevation in the shell the day before,there are now well-defined openings through which the tip of thebill of the embryo can be clearly seen. At this stage the mother birdmay frequently roll the eggs with her body in addition to turningthem with her bill. I have even observed the old bird pick out bitsof shell from a pipped egg, as if attempting to facilitate the processof hatching. After a few hours the calcareous shell is cracked forits entire circumference, but the shell membrane may remain intactfor a longer time. The struggles of the young, however, soon flingthe cap open with a part of the shell membrane on one side servingas a kind of hinge. After the embryo has kicked itself out of theprison shell, the tension of the drying shell membrane pulls the capback in place and thus prevents the youngster from being cupped byits own shell.The time of hatching of the various eggs of a set is remarkablyuniform, and in some instances the time elapsed from the time ofhatching of the first to the last egg is less than an hour. In caseswhere incubation started before the last one or two eggs were laid,the latter may be delayed, and in several cases under observation,during the summer of 1930, they failed to hatch in time, and the con-tained young were left behind when the brood left the nest.The precocious young of the prairie chicken are ready to leave thenest as soon as their down is dry, and the mother bird often hasdifficulty in preventing the first young from leaving the nest beforethe last to hatch are prepared to go. The brood may leave in a fewhours after hatching, but if the hatching takes place late in the af-ternoon the old bird, unless disturbed, will brood them on the nestduring the night, but leaves the next morning j ust as soon as the tem-perature and weather conditions are suitable for the chicks to move.The eggshells are never removed from the nest by the old bird.The chicks, as well as embryos in the eggs, are very sensitive to ex-tremes of temperature. Young left exposed to the cool damp nightair will quickly perish, and brooding at such times is absolutely
GREATER PRAIRIE CHICKEN 253
necessary. Though the young quickly perish if cold, especially ifwet, they will make rapid and remarkable recoveries from nearlyparalyzed conditions. Young that seem almost lifeless can bequickly restored to an active condition by merely holding them inyour hand and blowing your breath over them for 10 or 15 minutes.In one instance a mother bird was frightened away from the nestat night and failed to return. The young, still damp from the fluidsof the eggs, seemed destined to die. They were taken inside theblind, revived, and then kept alive and contented all night byplacing them against my body inside of a flannel shirt. At day-break the old bird appeared on the scene and claimed the youngstersone by one as they were released under the burlap of the blind.The language of the prairie chicken is readily understood by theyoung, even when first hatched. If the young are taken inside of theblind at the time of hatching they are indifferent to various soundsand notes of other birds calling outside of the blind, but as soon asthe adult prairie chicken appears and begins calling they respond atonce. If she gives her brirrrb-brirrrb call, they struggle all overone another trying to get out, but if the old bird becomes alarmedand suspicious she gives a sharp shrill call of caution and imme-diately each little chick cows down and " freezes " to a perfectlymotionless pose. Chicks set free at the edge of the blind made noeffort to go to the mother bird unless called. As soon as the call noteis given there seems to be an irresistible impulse on the part of theyoung to follow that call, although they can not see the old bird.When the chicks wander from the nest at hatching time that samecall brings them back, and hence this response is important in theirpreservation and doubtless is a matter of evolutionary development.Though the character of the call has a distinct meaning to the birds,one can with little effort imitate it and completely deceive the adultor young. I have often made use of this fact by inducing the birdto come near enough to the blind to obtain large portrait pictures ofher. One can so excite the bird that is nearing the hatching timethat she will exhibit an unusual behavior, such as turning the eggsover and over or twisting and squirming about the nest. She will,if the call, is well imitated and continued, leave the eggs to search forthe young. The bird at one nest under observation circled the blindagain and again and even attempted to get under the burlap toreach what she apparently supposed was a young in distress. Theold bird seems to have but little resourcefulness in aiding a youngin unusual situations. In one case where the bird was taking heryoung away from the nest, two of the young accidentally fell into adeep horse track. The young called desperately, but the motherseemed helpless. She raced around the opening several times and
254 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthen settled down in the grass to call for them. These two youngwould have perished without my intervention. An experiment ofplacing young in a hat near the nest also proved the bird's lack ofresourcefulness to cope with an unusual situation.Few things in nature have a greater human appeal than a familyof gallinaceous birds. The whole scene from the hatching of thefirst young to the departure of the brood is one brimming withthrilling incidents. The motherly interest of the old bird when thefirst youngster pokes its head through the breast feathers and givesa contented peep, as it picks at its mother's bill or her eye, is anevent never to be forgotten. Then the unexpected poking of a downyhead through the plumage, first at the side, then through a rearwindow, and perhaps two youngsters surprising each other as theyappear simultaneously, all are experiences that make a long vigilin the blind well worth the effort. As more of the young hatch theybecome more daring and may vigorously compete for a position onthe mother's back. They make repeated attempts to scale the slip-pery feathered dome, and finally when one does succeed he has anunmistakable look of triumph. All these things seem to have atruly human aspect, and surely the most skeptical can not help buttake an anthropomorphic attitude toward their behavior.Ordinarily a few hours after the young are hatched the old birdleaves the nest, allowing the young to follow after her. She gener-ally goes a few yards, settles in the grass and then continues callinguntil all the young are gone. This procedure is repeated until she iswell away from the nest. During this time the young are broodeda great deal, but before many hours, especially if the day is warm,they become active in searching for insects and other food.Plumages.?In a prairie chicken a few hours old the chin and loresare " primuline yellow," sides of head, including down on eyelids,
" naphthalene yellow," throat and breast " wax yellow," remainder ofunderparts " barium yellow," and down on tarsus " straw yellow."The lighter areas above are yellowish, strongly tinged with " cin-namon," which approaches " Mikado brown " on the rump, and theentire upper parts are marked with numerous irregular black spotsand patches. There is a small black patch back of the eye and threeirregular shaped black spots in the auricular region. The iris is
" dark Quaker drab," the base of the upper mandible is " pecanbrown " and tipped with a lighter color, and the upper surface of themandible is black, which extends for a distance equal to two-thirdsthe length of the bill. The lower mandible is pale "flesh color,"tipped with " straw yellow." The posterior part of the tarsus, notcovered with down, is " yellowish citrine." The upper surface of
GREATER PRAIRIE CHICKEN" 255the toes is " honey yellow," the undersurface " mustard yellow," andthe nails " flesh color."The young of the sharp-tailed grouse is similar to that of theprairie chicken, but with the following minor differences : The yellowof the unclerparts, not so deep or so bright as in the prairie-chickenyoung. Upper parts with much less black, especially in the regionof the back and rump. The brown of the rump is a paler shade.The black on the upper mandible extends down only one-half the dis-tance of the length of the bill. When the young of the prairiechicken and sharp-tailed grouse are compared with young of theruffed grouse, they are seen to be much yellower and much lessreddish brown, so characteristic of the day-old ruffed grouse.The average weight of 17 young prairie chickens that hatched ina nest near Bancroft, Portage County, Wis., was 15.9 grams. Theaverage weight of the eggs 2 days before hatching was 19.4 grams.[Author's note : As with all young grouse, the wings and scapu-lars begin to grow soon after the chick is hatched, and juvenal plum-age is rapidly acquired, long before the young bird attains its growth.In this plumage the crown is " hazel," spotted with black ; the feath-ers of the back, scapulars, tertials, and wing coverts are boldly pat-terned with " ochraceous-tawny," black, and " snuff brown," manyfeathers, especially the scapulars, having broad white shaft streaksspreading out into a white tip ; the primaries are spotted with palebuff on the outer web; the pointed tail feathers are barred or pat-terned with the colors of the back; the chin is white, and the under-pays are dull white, washed with buff and spotted with dusky onthe breast and flanks.Before the young bird is fully grown, in July, the postjuvenalmolt begins with the primaries. This is a complete molt, except thatthe outer pairs of juvenal primaries are retained for a whole year.Otherwise the first winter plumage is practically adult. Adults havea partial prenuptial molt, about the head, in March and April, anda complete postnuptial molt in August and September.]In addition to the normal plumages of the prairie chicken thereare unusual types that have attracted the attention of sportsmen andornithologists. In certain individuals of the prairie chicken thereis a prevalence of rufous or reddish brown, which is due to an excessof red pigment in the feathers, a condition known as erythrism. Thered phase of the plumage is a common occurrence among ruffedgrouse, but as yet it has been noted in comparatively few cases ofpinnated grouse. The following cases, which have come to my atten-tion, are of interest
:
George N. Lawrence (1889) described a specimen in which all ofthe light markings were tinged with light, bright rufous and the
256 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUM
entire underparts, throat, and neck tufts were deep rufous (red-dish brown).William Brewster (1882 and 1895) called attention to four speci-mens exhibiting the red phase in which there was but little variationwith respect to the depth and extent of the reddish brown or chest-nut coloring. The upper parts of the birds were strongly suffusedwith reddish brown, while most of the underparts were clear reddishor rusty chestnut and the usual blackish chestnut bars were nearlyor quite wanting on the sides. All four specimens examined byBrewster were males.Cases of albinism, in which there is a lack of pigment, resultingin a white plumage, have been frequently noted. Some of the albi-nos are not pure, but may have a little pigment in certain of thefeathers, giving them a dusky appearance.An albinistic specimen collected March 6, 1893, near the MissouriRiver, Iowa, is in the collection of the Museum of ComparativeZoology, Cambridge, Mass. This bird is white, with the exceptionof minor, pale, rusty brown crossbars and markings.J. A. Spurrell (1917) records a very interesting case of an albinoin Sac County, Iowa, which attained quite a local reputation becausethe bird was so clever that it eluded all attempts to trap it. Thehunters in the vicinity made a point to spare the " white chicken "in the hope that it might be captured alive.Hybrids between the prairie chicken and the sharp-tailed grousehave been noted by many observers. In Wisconsin, where the rangesof the two species overlap, it is a common experience to see themassociated together at all seasons of the year, and it is not at allsurprising that they frequently interbreed.J. H. Gurney (1884) described a male hybrid between the sharp-tailed grouse and the prairie chicken in which the pinnae werepresent, but only one-fourth of an inch long. The tail was a hybridgray between the brown of the sharp-tailed grouse and the white ofthe prairie chicken. The sides of the toes were only slightly feath-ered and the general coloration was intermediate between the twospecies.A hybrid between the prairie chicken and the sharp-tailed grousein which the elements of the prairie chicken predominated has beenreported by F. C. Lincoln (1918). The author has examined ahybrid specimen in the possession of Mrs. H. M. Hales, of Hancock,Waushara County, Wis., which also resembles the prairie chickenin most of the characters of its plumage.William Eowan (1926) figures and describes two female hybridsof the sharp-tailed grouse and the prairie chicken collected in Al-berta, Canada. One individual collected near Edmonton resembles
GREATER PRAIRIE CHICKEN 257the pinnated grouse more closely than the sharp-tailed grouse, whilethe other, shot at Gough Lake in the southern part of the Province,resembles more nearly the sharp-tailed grouse. The ovary of theEdmonton bird, according to Mr. Rowan, was normal. These are theonly Alberta hybrids known to him, but he states that hybrids be-tween the prairie chicken and the sharp-tailed grouse are frequentin Manitoba, where the pinnated grouse is more numerous than it isin Alberta.Glenn Berner, of Jamestown, N. Dak., writes that he killed ahybrid grouse in 1923 in which the back, head, and tail resembledthe prairie chicken, whereas the breast, legs, feet, and under tailparts were like those of the sharp-tailed grouse. The breast was notbarred as in the prairie chicken but spotted as in the sharp-tailedgrouse. The bird when flying had the characteristic cackle of thesharp-tailed grouse.According to O. A. Stevens, Fargo, N. Dak., there is a hybrid inthe collection at the North Dakota State Agricultural College.I have examined a female and three male hybrid specimens of theprairie chicken and sharp-tailed grouse in the Museum of Compara-tive Zoology, at Cambridge, Mass., which were obtained in the Bos-ton markets, March 24, 1873, February, 1887, December 29, 1899, andJanuary 24, 1893, respectively. So far as I know, these hybrids donot reproduce themselves, and in most cases this is probably due tothe sterility of the individuals.Food.?The prairie chicken, like other grouse, is adaptable in itsfood eating habits, varying its diet from season to season and sus-taining its life on the food that is most abundant and easilyobtained.Dr. Sylvester D. Judd (1905a) reported on the examination of 71stomachs of prairie chickens collected in the Middle West and rep-resenting all months of the year except July. The food consisted of14.11 per cent animal matter, chiefly grasshoppers, and 85.89 percent vegetable matter, made up of seeds, fruit, grain, leaves, flowers,and bud twigs.According to Judd's report the prairie hen is highly insectivorousfrom May to October inclusive, insects constituting one-third of thefood of the specimens shot during this period. The species is par-ticularly valuable as an enemy of the Rocky Mountain locust. Dur-ing an invasion by this pest in Nebraska, 16 out of 20 grouse killedby Prof. Samuel Aughey (1878) from May to October inclusive,had eaten 866 locusts. Beetles and miscellaneous insects were eatenin smaller numbers.From October to April, inclusive, according to the BiologicalSurvey report (Judd, 1905a), the prairie hen takes little but vege-
258 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMtable food, consisting of fruit, leaves, flowers, shoots, seeds, grain, andmiscellaneous vegetable material. It is especially fond of rose hips,which comprised 11.01 per cent of the food. When the deep snowcauses scarcity of other supplies, the sumac affords the prairie henwith abundant food. Seeds make up 14.87 per cent of the annualdiet, of which grass seeds form 1.03 per cent, seeds of various polyg-onums 8.49 per cent, and miscellaneous weed seeds 5.35 per cent.The prairie chicken eats more grain than any of the other nativegallinaceous birds; the food examined by the Biological Survey was31.06 per cent grain. The stomach of one bird shot in June inNebraska contained 100 kernels of corn and 500 grains of wheat.Buckwheat, barley, oats, and millet are relished, but corn appears tobe the favorite cereal, amounting to 19.45 per cent of the food.Wheat was next in order represented by 11.Gl per cent. Like othergallinaceous birds, it is fond of mast such as hazelnuts and acorns,though it obtains much less than the ruffed grouse. A bird shot inMinnesota in March had bolted 28 scarlet-oak acorns.An analysis of organic material in the food of IT prairie chickenscollected in Wisconsin for the Wisconsin Conservation Commission,chiefly in fall, revealed that about 28 per cent of the food was animaland 72 per cent vegetable matter. Gravel constituted 6 per cent ofthe combined organic and inorganic material of the crop and stomachcontents. The average weight of the crop contents was 25.7 grams
;
the maximum 83 grams. The average weight of the stomach con-tents was 14.3 grams, and the largest quantity found in any onestomach weighed 23.1 grams.There were 84 kinds of vegetable matter and 82 kinds of animalmatter represented in crops and stomachs of the prairie chickenscollected in Wisconsin. Arranged in order of the percentages ofthe entire food eaten by the birds, the 25 more important foodsfollow: Short-horned grasshoppers, 26.7; ragweed, 11; oats, 10.8;clover, 7.7; black bindweed, 6.2; acorns, 4.5; greenbrier, 3.6; dogwood,3.5; crickets, 3.3; buckwheat, 3.1; bramble, 3.1; blueberries, 2.4; rose,1.7 ; hawkweed, 1.4 ; chokeberry, 1 ; galls, 0.94 ; ants and wasps, 0.88
;
poison ivy, 0.80; birch, 0.80; pin cherry, 0.64; woody debris, 0.64;bunchberry, 0.53; wild black cherry, 0.53; smartweed, 0.47; pigeongrass, 0.47.It will be seen that the chief difference between the above list offoods and the results published by the Biological Survey in 1905 isthe absence of corn in the recent list. In Doctor Judd's list cornmade up 19.45 per cent of the entire contents of all the birdsexamined. This may be accounted for by the change in methods offarming. In the past, corn was husked in the field and much grainwas accidently left behind by the harvesters. To-day, in Wisconsin,
GREATER PRAIRIE CHICKEN 259
a dairy State, all the corn that is raised is cut and made into ensilagefor the cattle. Practically no corn is allowed to ripen in the field;hence it does not appear as a food for the prairie chicken. Both listsagree in the large percentage of grasshoppers comprising the food.The prairie chicken and sharp-tailed grouse are notable grasshopperconsumers, which fact, together with their fondness for weed seeds,makes their presence a great asset to the farmers.Except in the northern part of their range, where very severeweather and deep snows prevail, there is sufficient natural food forthe prairie chickens at all seasons of the year. The prairie chickenis there hard pressed for an existence, since it does not seem to beable to subsist on buds and other foods above the snow to the extentthat it is done by the ruffed grouse. In Wisconsin, experiments con-ducted with winter feeding stations by the conservation commissionhave proved a great success. Plots of ground ranging from a halfacre to two acres in extent are planted chiefly to buckwheat, with sor-ghum, sunflowers, broomcorn, and corn planted as accessory foods onmost of the plots. Half of the crop is left standing and the otherpart is cut and placed in covered shocks, which are opened up afterthe deep snows arrive. According to the reports of the wardens incharge, as many as 200 to 300 birds visited a single station at onetime, a strong testimonial for the practicability of such stations ingame management.Migration.?The prairie chicken is a permanent resident in muchof its range, but in the Northern States there is a regular annualmovement of the birds southward at the approach of winter weather.There are counties in Wisconsin where prairie chickens do not breed,or are present in very small numbers during the summer season,whereas they are represented by large numbers of individuals dur-ing the winter months, especially when deep snows and extremelycold weather exist in the more northerly sections of the range fromwhich the birds apparently come. Observers in Door County, Wis.,have reported seeing flights of prairie chickens approaching theland from Green Bay. The birds supposedly came from the oppo-site shore, a distance of 12 to 15 miles, which, if true, means thatthey sometimes take flights exceedingly long and continuous for abird of the type of the prairie chicken. A. E. Doolittle, superinten-dent of Peninsular Park, Door County, saw a flock of 300 prairiechickens headed northeast, up the shore of Green Bay, which hethought were en route for the Michigan side of the bay. WilliamFairchild, former keeper of Chambers Island (near the middle ofGreen Bay, a distance of about 7 miles from the mainland), sawtwo prairie chickens arrive from Marinette in April, 1927. Theyremained several weeks, then flew eastward to Door County proper.
260 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMK. M. Anderson (1907), writing of the prairie chicken in Iowa,states : " While a certain number remain throughout the winter, largeflocks pass southward early in the winter, returning in March."The migration was even more marked in the past when the birdswere abundant. J. A. Spurrell (1917) states that there was a markedmigration of birds away from Sac County, Iowa, until about 1875-1880. After that date, he says, corn became a common crop and birdswintered as well as nested abundantly in that section of the State.Prof. W. W. Cooke (1888), in writing of migrations of prairiechickens in Iowa, stated
:
In November and December large flocks of prairie cbickens come fromnorthern Iowa and southern Minnesota, to settle in northern Missouri andsouthern Iowa. This migration varies in bulk with the severity of the winter.During an early cold snap immense flocks come from the northern prairies tosouthern Iowa, while in mild, open winters the migration is much less pro-nounced. During a cold, wet spring the northward movement in March andApril is largely arrested on the arrival of the flocks in northern Iowa ; but anearly spring, with fair weather, finds them abundant in the southern tier ofcounties in Minnesota, and many flocks pass still farther north. The mostremarkable feature of this movement is found in the sex of the migrants. It isthe females that migrate, leaving the males to brave the winter's cold. Mr.Miller, of Heron Lake, Minn., fairly states the case when he says : " The femalesin this latitude migrate south in the fall and come back in the spring about oneor two days after the first ducks, and they keep coming in flocks of from ten tothirty for about three days, all flying north. The grouse that stay all winterare males."In the spring of 1884, at Iowa City, Iowa, the first flocks passed over March10, and the bulk March 22 ; at Newton, Iowa, the bulk was noted March 23.Glenn Berner, of Jamestown, N. Dak., says in his notes
:
During the spring of 1924 I witnessed a decided northward movement ofprairie chicken flocks numbering 10 to 100, some quite high in the air?late inthe afternoon?possibly 30 flocks being seen from one location in two hoursand very few of them alighting.O. A. Stevens, of Fargo, N. Dak., writes that he saw a flock ofprairie chickens fly high overhead on October 25, 1930, a time thatcoincided with a marked migratory movement of other birds. Mr.Stevens has noted for several years this annual movement taking-place during the latter part of October. He considers these flightsas a distinct migration, as the birds were always moving in the samedirection.Information on the migration of the prairie chicken supplied byF. C. Lincoln, of the Biological Survey, is as follows:Although not a true migrant in the strict sense of the word, the prairie chickenhas been known to make more or less regular flights north and south. Curi-ously, these movements appear to be confined chiefly to the females, the malesremaining in the breeding areas during winter. The flights apparently werenot infrequent a generation ago when the birds were much more abundant thanat present. The extent of the movement and the number of individuals partici-
GREATER PRAIRIE CHICKEN 261pating were dependent upon the severity of the winter in the northern part ofthe range. The exodus usually took place in November or December (Wisconsin,Madison, November 25; Iowa, Ogden, December 14, and Osage, December 30),while the return trip was made in February, March, or April (Iowa, Sioux City,February 4, Marshalltown, February 7, and Osage, February 13 ; Wisconsin,Unity, February 15, Elkhorn, March 2, and Whitewater, April 1 ; and Minnesota,Fort Snelling, March 21, and Minneapolis, April 8).Thus it is clear that the prairie chicken, at least in the NorthernStates, makes flights of considerable length, which we can consider ofa migratory character. In addition to these movements there areshifts and concentrations of the birds that are very local and mainlyconcerned with the food supply. At the feeding stations establishedin Wisconsin it was not unusual for 200 to 300 birds to feed at asingle station at one time, a total far in excess of the numbers breed-ing in the vicinity. Some of the birds may have migrated fromthe north, but it is probable that the mass of these flocks are merelyaggregations from a limited region of a few counties.Winter.?In regions where deep snows prevail, the prairie chickensoften dig themselves into the deep drifts to avoid the excessive cold.One observer at Green Bay, Wis., relates observing five prairiechickens alight on the surface of the snow, which was about 2%feet deep. The birds walked up to some weed stalks that projectedthrough the snow and then dug themselves in at places about 10 to12 yards apart. A day later the same observer flushed the birdsfrom the snow bank and found well-molded places on the groundamong the weeds. There was an accumulation of droppings ineach burrow, indicating that the birds had remained in the samespot during the night. The practice of digging into the snow hasproved disastrous at times when it becomes covered over with aresistant layer of ice.Gale W. Monson, of Argusville, N. Dak., says in his notes:In winter the prairie chickens are our most conspicuous birds. They spendthe nights in the tall grass of marshy meadows making small pockets forthemselves in the snow. At sunrise they leave their beds and fly to the nearestcornfield, there to eat their fill of that grain. In the afternoon they returnagain to their sleeping quarters. Their chief enemy at this time is the snowyowl, which sometimes depletes their numbers to a noticeable extent.Several observers in Wisconsin report that the tall marsh grass isfrequented by the prairie chickens as soon as the water of the swampsis solidly frozen over.John Worden, of Plainfield, Wis., states that during times of deepsnows the prairie chickens are often in a semistarved condition.At such times the birds showed little fear of man and often allowedhim to approach within a few yards before attempting to fly. Afarmer living near Babcock, Wis., stated that in collecting shocks74564?32 18
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of corn following a heavy snowstorm, he had virtually to drive thebirds away, and when flushed they flew but a few yards to the nextshock of corn. Such behavior is probably very unusual except un-der very extraordinary circumstances when the birds are sufferingwith extreme hunger. According to Mr. Worden, the birds fre-quently alight in trees during winter, but he says they invariablyroost on the ground at night.F. Hall, of Babcock, Wis., states that during the winter of 1928-29a flock of about a dozen prairie chickens came regularly to thepoplar trees of his back yard, but he was not certain to what extentthey fed upon the buds.
iDISTRIBUTIONRange.?South-central Canada and the United States east of theRocky Mountains, except the Southeastern States.The full range of the prairie chicken extended north to southernSaskatchewan (Quill Lake and Indian Head) ; southern Manitoba(Oak Lake, Carberry, Westbourne, Ossono, and Shoal Lake) ; north-ern Minnesota (Crooked Lake) ; central Wisconsin (Unity, WildRose, and West Depere) ; Michigan, (Chatham, McMillan, SaultSte. Marie, and Fourmile Lake) ; southern Ontario (Wallaceburgand Chatham) ; and Massachusetts (Springfield, Newton, and CapeAnn). East to Massachusetts (Cape Ann and Marthas Vineyard)
;
Long Island, N. Y. (Miller Place and Hempstead) ; New Jersey(Barnegat) ; and southern Maryland (Marshall Hall). South tosouthern Maryland (Marshall Hall) ; District of Columbia (nearWashington) ; probably Virginia and perhaps North Carolina; south-western Pennsylvania (Blairsville) ; central Ohio (near Columbus)southern Indiana (Bloomington, Marco, and Bickwell) ; northwesternKentucky (Henderson) ; southern Louisiana (Iowa Station and Cal-casieu Pass) ; and Texas (Beaumont, Richmond, Edna, Port Lavaca,St. Charles Bay, Austin, and Tascosa). West to northwestern Texas(Tascosa) ; Colorado (Barton and Barr) ; southeastern Wyoming(Chugwater) ; northwestern Nebraska (Chadron) ; South Dakota(Pine Ridge Reservation, Kadoka, and Short Pine Hills) ; NorthDakota (Bismarck, Charlson, and Crosby) ; and southern Saskat-chewan (Johnston Lake and Quill Lake).A specimen taken in the fall of 1917 near Huntley, Mont., is atpresent the only record for that State. Prairie chickens have beennoted as rare in winter near Fayetteville, Ark., and are said to occurin that season at De Witt, Ark.The prairie chicken and its eastern relative, the heath hen, havebeen extirpated over great areas in their former range. The heathhen is, in fact, extinct except for a single bird, which at the time
attwater's trairie chicken 263
of writing (November, 1930) was still living on Marthas Vineyard,Mass. Western Indiana marks the present eastern boundary of thespecies.Many attempts have been made to transplant the prairie chickeninto other parts of the country. These have all met with failure,except for an apparent introduction in northern Michigan. Theyare reported as thriving in the vicinity of Sault Ste. Marie andMcMillan.The range as described is for the entire species, which has, how-ever, been separated into three subspecies. True americanus, thegreater prairie chicken, formerly occurred from the southern partsof the prairie Provinces of Canada and eastern Colorado east tosouthwestern Ontario, northwestern Ohio, and western Pennsylvania.The range of the heath hen (T. c. cwpido) included New England(to southern New Hampshire), New York, and other States of theAtlantic seaboard, probably south to and including Maryland. Att-water's prairie chicken (T. c. attwaterl) is found in the coastal regionof Texas and southwestern Louisiana.Egg dates.?Manitoba : 6 records, May 24 to June 20. Minnesotaand Dakotas : 30 records, May 1 to June 18 ; 15 records, May 18 to29. Wisconsin: 37 records, May 5 to July 10; 19 records, May 28to June 20. Illinois and Iowa: 32 records, April 20 to June 6; 16records, May 6 to 25. Marthas Vineyard : 3 records, June 2 and 5and July 24. Texas : 4 records, April 3 to May 16.TYMPANUCHUS CUPIDO ATTWATERI (Bendire)ATTWATER'S PRAIRIE CHICKENHABITSThe small dark race of the prairie chicken named atttvateri is con-fined to southwestern Louisiana and eastern Texas, mainly in lowprairies in the coastal counties. It was described by Maj. CharlesE. Bendire (1894) and named by him in honor of Prof. H. P. Att-water. He gives as the subspecific characters:Smaller than T. americanus, darker in color, more tawny above, usually withmore pronounced chestnut on the neck; smaller and more tawny light coloredspots on wing coverts, and much more scantily feathered tarsus, the latter neverfeathered down to base of toes, even in front; a broad posterior strip of bareskin being always exposed, even in winter, while in summer much the greaterpart of the tarsus is naked.George Finlay Simmons (1925) describes its haunts as " rollingopen, grassy, fertile upland prairies, where the grass is from 1 to 3feet tall, old and thick and mixed with weeds ; wheat and corn fields
;
takes to timber only during snow and sleety storms."
264 BULLETIN 162, UNITED STATES NATIONAL MUSEUMReferring to its history, Simmons writes:Formerly abundant on the open prairies, these wonderful game-fowl becameextirpated in the Austin Region through two agencies ; civilization and hunting.They disappeared rapidly as the country was settled up and as cultivated fieldstook the place of the extensive, wild, unfenced prairies; and hunters quicklykilled the few remaining birds.Its courtship performances, nesting habits, eggs, plumages, andmolts are all similar to those of the common prairie chicken.Eggs.?The eggs are indistinguishable from those of the commonprairie chicken. The measurements of 27 eggs average 42.3 by 31.5millimeters ; the eggs showing the four extremes measure 44.9 by 32,42.4 by 33.5, 38.8 by 28.9, and 39.8 by 28.6 millimeters.Food.?Simmons (1925) says that itsometimes flies to treetops to inspect corn fields before alighting in them tofeed ; frequently feeds in the open in plain sight of observers several hundredyards away. During early breeding season, feeds largely on insects, such asgrasshoppers, crickets, potato bugs, and other beetles; in fall and winter, topsand seeds of leguminous plants, tender buds and green leaves of late winter,fruits, berries, and waste grain of stubble and corn fields.Behavior.?Simmons says that it isobserved singly and in pairs in spring ; in fall and winter, roamed about inflocks of from 10 to 12 up to a 100 or more, moving about over the prairies andgrain fields, generally keeping among bushes and tall grass inland, the openprairies and grassy knolls along the coast. Stately of bearing, but otherwisevery much like the domestic fowl in its actions. In spring, a " scratchingground " or smooth, open courtship ground is selected, where pairing takesplace.Voice.?The same observer describes the notes as " nondescriptcalls ; strange cackles, with a muffled booming love-call, uck-ah-umb-boo-oo-oo-oo-oo-oo; and a loud beating boom-ah-boom, perhaps pro-duced by a beating of the wings; when alarmed, a rapidly repeatedcluk-cluk-cluk-cluk; female, when flushed, utters a low kuk-kuk-kuk-kuk-kuk-7euk-kuk-kuk."TYMPANUCHUS CUPIDO CUPIDO (Linnaeus)HEATH HENContributed by Alfred Otto GrossHABITSThe heath hen and the prairie chicken are so closely related thatthey are now considered as geographical races and not as distinctspecies. In 1885 William Brewster (1885a) called our attention todifferences between the pinnated grouse of Marthas Vineyard (heathhen) and the western pinnated grouse, or prairie chicken. He named
HEATH HEN 265the western form as the new species, because Tetrao cupido of Lin-naeus was the eastern form from the fact that its habitat is givenas Virginia. The differences between the eastern and western birdsare so slight and the variations of the individuals so great thatornithologists now concede that Brewster was not well justified inthe establishment of a new species.In prehistoric times the common ancestors of the heath hen andthe prairie chicken probably ranged in an uninterrupted distribu-tion from the Atlantic seaboard to the plains east of the RockyMountains. Later the birds of the East became separated fromthose of the West, and as a result of this isolation and differencesin environment certain modifications arose that have resulted in theestablishment of the two geographical races
?
Tympanuchus cupidocupido, the heath hen, and Tympanuchus cupido americanus, theprairie chicken.The prairie chicken is still flourishing and in recent years hasbeen rapidly regaining its numbers in favorable sections of theMiddle West, but the heath hen has been unable to cope with thechanging conditions of its restricted environment, and to-day isrepresented, so far as we can ascertain, by a single male individual,which is living out its normal life on the scrub-oak plains of MarthasVineyard Island, Mass.The following account is made up primarily of modified excerptsof the contributor's monograph on the heath hen (Gross, 1928) andfrom subsequent annual census reports
:
Historical.?The heath hen is among the first of the Americanbirds to be mentioned in the writings of the early colonists who cameto our shores. There is, however, such a dearth of material concern-ing the heath hen during these early times that we know but littleconcerning the conditions under which it existed, and the records areso incomplete that we are unable to determine with any degree ofaccuracy its relative abundance and distribution prior to the nine-teenth century. Some of the earlier American writers designatedthe heath hen by the name " heathcocke," " pheysant," or " grous,"but their notes and descriptions are such that they can be clearlyreferred to this species. William Wood (1635) in his New EnglandProspect writes as follows : " Heathcockes and Partridges be com-mon : he that is husband, and will be stirring betime, may kill halfedozen in a morning. The Partridges be bigger than they be inEngland, the flesh of the Heathecocks is red, and the flesh of thePartridge white, their price is four pence a piece." Wood residedat what is now the city of Lynn in Massachusetts. His map includedCape Ann and the Merrimac River ; hence it is evident that the heathhen existed in northeastern Massachusetts in his day. Thomas
266 BULLETIN 162, UNITED STATES NATIONAL MUSEUMMorton (1637), writing concerning the heath hens, which he called" pheysants," stated that these birds were like the pheasant hen ofEngland in size but were rough footed and had " stareing " feathersabout the neck. The birds, according to Morton, were so commonthat they seldom wasted a shot upon them. The writings of manyothers who followed indicated that the birds were distributed alongthe Atlantic seaboard from Maine and Massachusetts southward toVirginia and possibly the Carolinas. They were by no means evenlydistributed over this region, but were restricted to certain areaswhose features and productions were suitable for their existence.There were large heavily timbered areas that probably were nevervisited by the heath hen. In favorable localities, such as the brushyplains of eastern Massachusetts, they were abundant. ThomasNuttall (1832) wrote as follows: "According to information I havereceived from Governor Winthrop, they were so common on theancient brushy site of Boston, that laboring people or servants stipu-lated with their employers not to have Heath Hen brought to tableoftener than a few times a week." No published statement has everbeen found that more impressively reveals to us the abundance ofthese birds in early colonial times. It was chiefly on the sandy scrub-oak plains of Massachusetts, Connecticut, Long Island, New York,New Jersey, and Pennsylvania that they existed in large numberswhen the white man first came to America. The birds served as avaluable source of food, and because they were easily tricked andkilled they were exterminated at an early date in the more accessibleareas, and soon after 1840 were entirely gone from the mainland ofMassachusetts and the State of Connecticut. The birds persisted fora longer time on Long Island, and a few continued to battle forexistence on the plains of New Jersey and favorable places among thepines and scrub oaks of the Pocono Mountains in NorthamptonCounty, Pa. Since 1870 the surviving members of this interestingrace have been restricted to Marthas Vineyard Island, off the south-eastern coast of Massachusetts.Because of conflicting reports and uncertain statements we cannot be positive whether the heath hen was native to Marthas Vine-yard or was introduced there from the mainland by man. In eithercase it is truly remarkable that the heath hen, after being so greatlydepleted in numbers, has persisted for more than half a century inthis very restricted area where excessive interbreeding has occurredand where the birds have been subjected to all the vicissitudes ofdiseases, enemies, and other adverse conditions.In 1890 William Brewster (1890) made a careful census and atthat time estimated that there were 200 birds on the entire island.Kentwood (1896) stated that there were less than 100 birds in 1896.
HEATH HEN" 267By the beginning of the twentieth century the birds had reached avery low ebb in their existence. The year 1908 witnessed one of themost notable steps taken in the history of the heath hen in an effortto preserve it from extinction, in the establishment of a reservationin the midst of the breeding range where the birds could be protectedfrom poachers and predators by competent wardens. Six hundredacres were purchased by private subscription, and an additionaltract of 1,000 acres was leased by the Commonwealth for a reser-vation, which was systematically improved to make it attractive forthe birds. There is no doubt that the prolongation of the life of theheath hen on Marthas Vineyard Island has been due to the interesttaken in it by the State of Massachusetts, conservation organizations,bird clubs, sportsmen, and bird lovers. The State Department ofConservation expended $70,000, and thousands more were contributedby individuals, in the unprecedented efforts to prevent the bird frombecoming extinct. Many attempts were made when the birds wereabundant to transplant them to other favorable places on the main-land and to other islands such as Long Island, one of their formerstrongholds. Furthermore, the most experienced sportsmen andgame breeders were unable to breed the birds in captivity, a factindicating that the heath hen was very sensitive to radical changesin its environment and that it would not yield to such methods ofconservation. All the many experiments of introducing the westernprairie chicken to the East have likewise proved unsuccessful.When the reservation was established in 1908 there were onlyabout 50 heath hens, but as a direct result of the efficient protectionthe birds increased very rapidly and by 1915-16 they were to befound in all parts of Marthas Vineyard with the exception of GayHead, the extreme western end of the island. It was then possibleto flush a flock of 300 or more birds almost any day from the cornand clover plots planted on the reservation for the birds. An esti-mate made by William Day, then superintendent of the reservation,indicates there were probably 2,000 birds on the island. This was agreat triumph for those who had encouraged and fostered the reser-vation, but unfortunately success was not long-lived.In spite of the unusual precautions taken to prevent the spreadof fire, a terrific conflagration broke out during a gale on May 12,191G, which swept the greater part of the interior of the island,destroying brooding birds and their nests and eggs, as well as thefood and cover of the birds on more than 20 square miles, rightin the heart of the breeding area of the heath hen. This fire undidin a few hours the accomplishment of many years of work. A hardwinter followed the fire, and in the midst of this came an unprece-dented flight of goshawks, which further decimated the number of
268 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbirds. The net result of this catastrophe was an amazing decreasein the number of heath hens, which according to official estimateswas reduced to less than 150 birds, most of which were males.There was a slight rally in numbers during the following fewyears, but the birds were too far gone to overcome the surmountinguncontrollable conditions of extensive interbreeding, declining sexualvigor, the condition of excess males, and, worst of all, disease.In 1920 many birds were found dead or in a weak and helplesscondition, indicating that disease was then exacting its toll. Theheath hen is very susceptible to poultry diseases, and when domesticturkeys were introduced to the island in large numbers the dreadeddisease blackhead came with them. The turkeys and heath hens fedon the same fields, and thus the disease was readily transmittedthrough droppings to the native birds.The heath hen continued to decrease in numbers, and by 1925it was apparent that they had reached their lowest ebb in history.The Federation of the Bird Clubs of New England then came tothe front and offered to raise $2,000 annually to support additionalwarden service. In spite of this splendid cooperation the birds, aftertwo years of effort on the part of all concerned, continued to de-crease. The 1927 spring census revealed but 13 birds, only 2 of whichwere females. During the fall of 1928 only two birds were seen andafter December 8 but one was reported. This bird was photographedfrom a blind on April 2, 1929, at the farm of James Green locatedon the State highway between Edgartown and West Tisbury. At thattime it was the common expectation that the bird would step out ofexistence before the end of another year. (See pi. 1, frontispiece.)It was seen regularly until May 11, 1929, but after that date it disap-peared among the scrub oaks to live a life of seclusion, as was cus-tomary for the heath hen to do in the past, during the summer months.After the molting season it again appeared at the Green farm toannounce to the world that it was still alive. It was seen at irregularintervals during the winter, and after the first warm days of Marchit appeared daily at the traditional booming field at the Green farm.The bird was studied and photographs were taken again at the timeof the annual census in March-April, 1930. The lone bird continuedto appear at the Green farm during April and May, where it wasobserved by many ornithologists and bird lovers who journej^d tothe island to get a glimpse of the famous last bird. The bird againdisappeared during the summer, and no reports were received untilit almost met a tragic death on September 15, when it was nearly runover by an automobile traveling one of the little-used roads leadingacross the scrub-oak plains. In October it resumed its daily visits tothe open field on the Green farm and at the time of this writing
HEATH HEN 269(November 15, 1930) was still alive. It is the first time in the historyof ornithology that a bird has been studied in its normal environmentdown to the very last individual. How long this bird will liveno one can safely predict ; its going is inevitable, and the death of thisindividual will mean the death of its race, and then another birdwill have taken its place among the endless array of extinct forms.Ornithologists, bird lovers, and sportsmen the world over, however,will have the satisfaction of knowing that all that could be done hasbeen done to save this bird from extinction. The State departmenthas assured us that the last bird will be allowed to live, and whendeath comes, whether it is due to old age, disease, or violence, we shallknow that the life of the last heath hen was not wilfully snuffed outby man. [During the fall of 1931, this lone survivor disappeared.]Courtship.?There was no part of the behavior of the heath henmore unique, more interesting, and more specialized than the ex-traordinary performances during the courtship season. I vividly re-member the thrill of hearing and seeing the heath hen's boom forthe first time on Marthas Vineyard Island during April, 1923. Atthat time the birds came regularly to a definite part of the meadowwest of the reservation house. A wooden blind, 4 by 6 by 6 feet,had been erected several years before for the convenience of thelarge numbers of ornithologists who journeyed to the island eachyear to get a glimpse of the heath hen. The blind had become apart of the environment of the drumming field around which thebirds came to enact their fantastic dances without any fear of beingharmed. At times one or two of the birds would even alight onthe flat top of the structure, offering unexcelled opportunities forstudy of the intimate details of their behavior.The following observations, as recorded in my notebook for April11, 1923, are typical of the many mornings spent inside the blind
:
" I left the reservation house at 3.30 a. m. It was very dark andonly the faint light of the stars illuminated the way. The cold,enervating air quickened my step, and as I walked along, the frostedgrass crunched under my feet with a metallic resonance. At thisearly hour all was quiet so far as voices of birds were concerned.I entered the blind, closed its door on creaky hinges, and preparedmyself to wait patiently for the first note of the heath hen. Aslight fog rolled in from the sea and for a time hid the stars.At 3.55 a. m. with the first dim light of dawn I heard the clearwhistled notes of a bobwhite perched somewhere among the scruboaks. At 4.05 the first robin was heard chirping, and five minuteslater a vesper sparrow was singing its awakening song. In a shortwhile a host of other birds were adding their notes to the morningchorus. At 4.21 the first " toot " of the heath hen was heard, a note
270 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthat has often been mistaken for a muffled blast of a tug boat or afog horn. Though I was well prepared for this deception, I mustadmit that I did not at first associate this curious note with theheath hen, for the light was yet dim and the fog obscured the viewof the bird. At 4.27 a heath hen appeared from the scrub oaks onthe south side of the meadow at a point relatively near the blind.The calls of this individual at once stimulated the birds on thewestern end of the field to greater activity in tooting. After 20minutes a second and then a third bird came out of the scrub oakson the south, and for a time all were busily engaged in feeding. Oneof the males flew to the roof of the blind, where he commanded asplendid view of the field and his companions. Later it was foundthat the resonant wooden roof proved to be an admirable place toconduct their stamping and courtship performances. At 4.45 thebirds without any warning interrupted their feeding and began
" booming or tooting " at a point only a few yards from the blind.The male on the roof joined his fellows on the ground. At thisclose range the call resembled whhoo-doo-doooh. The note variedsomewhat in subsequent renditions and was variously interpretedas whoo-oodul^doo-o-o-o-o, whoodle-dooh, or whoo-dooh-dooh.The sound was accented on the second syllable or the first part ofthe second and then gradually diminished in intensity. It requiredfrom iy2 to 3 seconds to render the different versions given above.The number of calls a minute varied greatly, according to weatherconditions, temperature, time of day, and the season. The boomingwas interspersed with henlike calls resembling cac, cac, cac, or oc, oc,oc, oc, goc, (/oc, goc, goc, occasionally ending with a queer call thatsounded like auk-ae-e-e-e-e-ek. The males frequently leaped intothe air to a height of 3 or 4 feet and so doing uttered a piercingrolling wrrrrrrrrb, followed by a curious indescribable laughterlikesound. In this wild demonstration the bird completely reversedits orientation in the air and landed on the ground, usually facingin the opposite direction. This leaping and screaming seemed tobe augmented by similar performances of the birds on the otherside of the field, and it was an evident challenge to their fellowantagonists.
"At 5 a. m. the sun pierced the screen of fog and appeared like agiant fiery ball above the eastern horizon. The morning chorus ofbirds then rapidly diminished in volume, but the heath hen nowprepared for real action. One male from each group ran rapidlytoward each other in a defiant warlike attitude. When near togetherthey hesitated, lowered and waved their heads, leaped at each other,and struck their wings vigorously as they lunged forward. A fewfeathers flew, but no real harm was done, and they settled back in
HEATH HEN 271
a comfortable position and occasionally uttered a long-drawn-outbut shrill cry, which fluctuated greatly in tone and intensity. Onebird arose after a few minutes, circled, paced a few steps, and wentthrough his repertoire of toots and calls without any interferencefrom his antagonist. Later one ambitious male insisted on chasingone opponent after another, following after them rapidly on footuntil they took wing. He flew after them for a distance of 30 to 50yards, then returned to repeat the performance with another weakermember of the group of a dozen birds. It reminded one of boys atplay after being pent up in school all day. These thrilling spectaclescontinued until 6.50 a. in., when with one common impulse all thebirds left the field to the seclusion of the scrub oaks to remain quietuntil the afternoon, when they again appeared on the drumming fieldduring the few hours preceding sunset. But when the last glow oftwilight faded into darkness the fantastic dance ceased to be resumedat the coming of dawn the next day."The first " tooting " calls of the year were usually heard the lastweek of February or the first week of March, the date varying fromyear to year and depending largely on the nature of the weather.In 1927 a series of warm days started the birds booming as early asFebruary 12. Though an early beginning was sometimes made it wasnot until the latter part of April or the first week of May that thecourtship reached its maximum intensity. It then gradually dimin-ished, and by the end of May the performance was generally over,but a few more persistent males often continued a few weeks longer.In 1923 the last " boom " for the year was recorded on June 11 ; andin 1920 a small group of males was still performing as late as June20. After the month of June the birds ceased their nuptial displaysuntil the mating season of the next year.The following details of the courtship performance were obtainedby repeated observations from blinds of the birds at close range andare supplemented by a study of captive birds and detailed labora-tory dissections: The tooting is usually prefaced by a short run,followed by a very rapid stamping of the feet, a part of the per-formance that is not readily detected unless the observer is verynear to the birds. The stamping is vigorous enough, however, to bedistinctly heard at a distance of 25 or 30 feet, and certain males,which did their stamping on the resonant roof of the blind, produceda noise second only to the tooting that followed. In preparationfor tooting the neck is outstretched forward; the pinnae (neck tufts)are usually directed upward or forward; the primaries are spreadand held firmly against the sides of the body and legs; and the tailis thrown upright at right angles to the axis of the body, thus dis-playing the white under tail coverts when viewed from the rear-
272 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMDuring this procedure the whole musculature of the body seems tobe in a strained state of contraction, as if it required great effort onthe part of the bird. As the inflation of the orange-colored sacs(it is really one large sac with two lateral areas devoid of feathers)begins, the tooting sound is heard. Sometimes there is a slightinflation before any sound is given. The inflation seems uniformand does not fluctuate with the inflections and accents of the tootingcall. At the end of the tooting the sacs collapse suddenly by therelease of air through the nares or more rarely through the openedmandibles. The sacs do not produce the notes, as was thought bysome of the earlier ornithologists, but have much to do with modi-fying the sounds produced by the syrinx (the vocal mechanism at thejunction of the bronchial tubes). The sounds are produced by theair forced from the lungs, which vibrates specialized membranes ofthe syrinx under control of a complex set of muscles. The soundwaves then issue through the trachea and glottis to the pharynx.In the production of such notes as the ordinary cackle the mandiblesare opened and the air accompanied by the sound waves issues outof the mouth. In the tooting performance the mandibles are tightlyclosed, the throat patch is elevated, and the tongue is forced againstthe roof of the mouth (palate) by the mylohyoides muscles, whichclose off the exit through the internal nares. The tongue is bentin such a way that it causes the glottis at the base of the tongueto open directly in front of the esophagus. The air now comingfrom the respiratory system is forced to fill the modified anterior endof the esophagus, or gullet, which becomes distended like a balloon. 1While the air sac is filling, the sound waves produced by the syrinxbeat against these tense drumlike membranes, which serve as resona-tors for the sounds and give them their great carrying power. Theordinary cackles and screams of these birds seem louder than thetooting or booming calls when one is near to the birds, but at a dis-tance of 200 yards or more you can scarcely hear these calls, whereasthe booming carries for long distances, often 2 miles or more underfavorable conditions.The female's part in the courtship is a passive one. She mindsher own business, and I have never heard her utter any calls or notesor show any concern in response to the ardent attentions of the males.When a male or pair of males came strutting and circling about afemale she kept on with her feeding. If the males came too near shemerely stepped to one side and continued with the serious businessof procuring food.
1 A number of detailed experiments were performed with both dead and living birds,which clearly demonstrate the nature of the vocal mechanism as described above. SeeGross (1928).
HEATH HEST 273The females gave frequent calls and notes when attending theiryoung. If the mother bird was suddenly surprised she gave a char-acteristic sharp call signal for her young to scatter and hide. If themembers of the brood were very young and unable to fly, she feigneda wounded bird and cried out as if in great distress as she flutteredalong the ground. If the young were older she usually sailed outover the scrub oaks and uttered a loud cackling call, which appar-ently was also for the purpose of attracting attention away from theyoung.Nesting.?The nest of the heath hen was built upon the groundand was usually composed of leaves, grasses, and twigs already inplace, to which were added materials found near the nesting site.The nests were concealed by the low dense vegetation of the scrub-oak plains. Indeed they were so well hidden from view and the eggsso well covered when the bird was away that few of the nests wereever found, in spite of the great efforts various observers have ex-pended to locate them.William Brewster (1890) states: " Only one person of the many Ihave questioned on the subject has ever found a heath hen's nest.It was in oak woods among sprouts at the base of a large stump andcontained either 12 or 13 eggs." There was a set of six eggs in theBrewster collection that were found in a nest in the woods near GayHead on July 24, 1885. This set was described and one egg wasfigured by Capen (1886). One of the eggs given to the UnitedStates National Museum is figured by Bendire (1892, pi. 3, fig. 2).Bendire stated that the six eggs referred to above were the only eggsin any collection known to him.In 1906, E. B. McCarta found a nest and nine eggs in a low butdense growth of scrub oak near the central part of Marthas Vine-yard Island. Dr. George W. Field photographed the nest on June2, and two days later the eggs were placed under a bantam hen. Oneof the eggs hatched on June 20, but unfortunately the chick waskilled by the hen, and the other eggs failed to hatch. This set ofeggs with the chick is now a part of a display group in the AmericanMuseum of Natural History, New York. On June 5, 1912, DeputyWarden Leonard, after a most prolonged and diligent search, founda nest and four eggs covered with leaves in a slight hollow sur-rounded by a dense mass of sweet ferns growing among the scruboaks. The oaks in the vicinity were 2 or 3 feet in height. When thenest was visited on June 12 the bird was incubating. On June 21,Doctor Field was able to determine that there were eight eggs, and aweek later he took an excellent series of photographs of the bird onthe nest. The bird sat so closely on the eggs that it was dislodgedonly by active effort. Deputy Leonard had no difficulty in approach-
274 BULLETIN 162, UNITED STATES NATIONAL MUSEUMing the bird, and she fought the approach of the hand in the samemanner as would a sitting hen, ruffling her feathers, opening herbeak, and striking viciously. The incubation period of the heathhen, according to Doctor Field, is 24 days.Eggs.?In addition to the eggs mentioned above in connection withthe account of the nests, there are the following: An egg in theBrewer collection of the Museum of Comparative Zoology markedTympanuchus cupido, Holmes Hole, Mass. (Holmes Hole is the oldname for Vineyard Haven). There is an egg in the John E. ThayerMuseum and another in the collection of the Boston Society ofNatural History. The latter was picked up on the plains of MarthasVineyard after the destructive fire of 1916. Bendire (1892) de-scribes the eggs as " creamy-buff in color with a slight greenish tint,ovate in form and unspotted." They are regularly oval in form,all specimens being quite uniform in this respect. The color is yel-lowish green of a peculiar shade. I have compared the colors ofthis set of eggs with Kidgway (1912) and find all of them to be a
" deep olive buff." One egg has a small spot of drab. All otherheath-hen eggs that I have examined are of this same deep olive-buff color.The measurement of the five eggs in the Brewster collection are asfollows, 43.5 by 32.5, 43.5 by 32.1, 43.8 by 32.5, 43.9 by 32.4, and46.2 by 32.9 millimeters, and the sixth egg, now in the United StatesNational Museum, measures, according to Bendire, 44 by 33 millime-ters. The egg in the Brewer collection collected at Holmes Holemeasures 44.2 by 32.6, and the egg at the Boston Society of NaturalHistory collected at Marthas Vineyard in 1916 measures only 40.3by 30.4, the smallest egg of the species I have examined.Young.?The first young of the heath hen for the season usuallymade their appearance during June. The earliest record is of abrood of 8 or 10 young seen near Edgartown on June 14, 1913, byDr. Charles W. Townsend. During the season of 1913, nine broods,with an average of four chicks to the brood, were seen. On July15, 1914, a dead chick was found that was estimated to be about fivedays old. In 1915, 14 broods were reported. The first brood, seenon June 19, consisted of six chicks about five days old. Allan Ken-iston saw seven broods with an average of five chicks each during thesummer of 1918, and in 1919 he reported broods on June 29, July 1,and July 11, the members of which were able to fly well at that time.For 1920 the following were noted: June 20, 10 young; June 24,6 young; July 4, 2 small broods, the numbers were not recorded;July 9, 5 or 6 young. During the summer of 1921 a brood of sevenwas seen on July 3, and a few days later a brood of eight was re-corded. On July 31 a brood of six, about two-thirds grown, was
HEATH HEN 275
seen. In 1922, between June 15 and June 30, there were five broodsreported containing four to eight chicks; the exact dates were notrecorded. In 1923 the writer made a continuous search for nestsand young throughout the summer, but the birds by that time wereso greatly reduced in numbers that only one brood of two chickswas seen, and that on July 3 during a downpour of rain, as we sud-denly surprised the brooding mother in the middle of one of thelittle-used crossroads. Since 1923 there have been no authenticrecords of broods of young birds, and I very much doubt if anyyoung have been reared since 1925.Plmnages.?The following description is based on a downy youngabout four days old : Underparts " cream-buff," the throat and mid-dle of the belly approaching " colonial buff." Sides of the head
" Marguerite yellow " with three small black spots back of the eye.Upper parts " tawny-olive " or " Isabella color," the region of therump " snuff brown " and " russet," variously marked with black.There is a conspicuous mark on the forehead. The remiges andcoverts marked with various tones of brownish drab and black, thefeathers tipped with dingy wThite. The measurements of this speci-men are as follows : Bill, 8 ; tarsus plus third toe, 39.8 ; wing, 42.5
;
length, 85 ; third toe, 23 millimeters. The natal down of a 2-day-oldspecimen hatched in captivity by Dr. John C. Phillips is similar tothe above description but with the following differences : The under-parts brighter yellow, the throat and sides of the head " amber yel-low." The bright yellows fade rapidly when exposed to air and light,and in chicks two weeks old the bright yellows of the underpartsare faded to a uniform " cream-buff." The measurements of the 2-day-old chick are : Bill, T.5 ; tarsus plus third toe, 37 ; wing, 28length, 79; third toe, 19 millimeters.I have been unable to obtain a specimen of the heath hen in thecompleted juvenal plumage, but it is reasonable to infer that thesequence of the molts and plumages are similar to those described forthe prairie chicken.The first winter plumage of the heath hen is acquired by a com-plete postjuvenal molt except on the two distal primaries. The firstwinter plumage is similar to the adult plumage, but the youngerbirds are readily distinguished from the adults by their smallersize, by the more rufescent color of the upper parts, and by the colora-tion of the throat, which is " cinnamon-buff " in contrast to the
" warm buff " or " cream color " of the adults and the white throatof the juvenals. The first plumages of the heath hen and the prairiechicken are so nearly alike that one can not readily distinguish themfrom each other. This ontogenetic resemblance indicates the closerelationship of the two races.
276 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe first nuptial plumage is acquired by a partial prenuptialmolt, which does not involve the wings and tail and body plumagebut is restricted to the head region. The nuptial plumage was usuallycompleted by April. The second or adult winter plumage is ac-quired by a complete postnuptial molt during August and Septem-ber. Specimens collected in October and November had the adultwinter plumage completed. From the adult winter plumage onwardthe plumages are repetitions of the nuptial and winter phases.Food.?The heath hen in its feeding habits was similar to othergallinaceous birds, such as the ruffed grouse, in being very adapt-able to the changing food supply throughout the seasons of the year.It was not dependent on any particular item but subsisted on whatwas most abundant and most easily procured.During the spring months the heath hen congregated in the openfields and meadows of the farms to feed upon the tender shoots ofgrasses, sorrel, and other plants, but when these became hardened andless palatable with the approach of summer the birds changed theirdiet to fruits and insects. In fall, berries and insects, such as grass-hoppers, were freely eaten, and in the winter months, acorns, seeds,and certain berries found throughout the range of the heath hen onMarthas Vineyard provided the birds with a livelihood. Com-paratively little snow falls on the island, and hence it was an excep-tional winter when the birds were not able to obtain sufficient foodfrom native plants. Even at times when the ground was coveredwith snow the scrub oaks held out an exhaustless supply of food inthe form of adorns.Our chief knowledge of the food of the heath hen is based uponthe meager notes of food included in the data with birds collectedand preserved as skins in various museums, upon field notes, andupon studies of birds kept in captivity. The following comprise theprincipal foods:The crops of three heath hens collected by C. E. Hoyle on January10, 1891, and two on December 28, 1895, contained bayberries{MyHca carolinensis) . A specimen of a heath hen killed by a snowyowl was reported by Allan Keniston to have had its crop gorgedwith bayberries. Birds kept in captivity at the reservation in 1915ate very freely of bayberries. During the winter of 1923-24 aflock of 15 birds frequented a large bayberry thicket near the southshore where they subsisted chiefly on these berries.The bearberry (Arctostaphylos uva-ursi), sometimes wronglynamed mountain cranberry or cranberry, is extremely abundantthroughout the central portion of the island and was freely eatenby the heath hen during the winter months when the trailing plantswere not covered with snow.
HEATH HEN 277Audubon (1840) stated that the barberry (Berberis vulgaris) wasthe chief food of the heath hen. This is also included in the listsof the food of the heath hen by other earlier writers, but there is noevidence that the birds in recent years depended very largely onthe barberry for a source of their food.The fleshy wild-rose berries, more frequently called rose applesor rose hips, were eaten by a bird collected March 7, 1896.The birds fed very freely on wild strawberries, which were abund-ant in the meadows and open areas of the reservation. They werealso fond of the cultivated varieties, as evidenced by their frequentdepredations in gardens grown near the haunts of the heath hen.The partridgeberry {Mitchella repens) was so frequently eaten bythe heath hen that the earlier settlers called this berry the heath-hen plum. Not only the berries but the leaves of this plant wereoften eaten by the birds during the early fall and winter months.The dryland blueberry (Vaccinium vacittans), the low-bush blue-berry (Y. pennsylvanicum) , and the black huckleberry (Gaylus-sacia baccata) were eaten during the berry season. On August 24,1913, William Day saw a flock of 51 heath hens feasting on blue-berries, and he also states that captive birds ate freely of blueberriesprovided for them.The acorns of the scrub oak {Quercus ilicifoHa) have been calledthe " bread " of the heath hen. No natural food is more abundant onMarthas Vineyard, and no food is more dependable during the win-ter months. The scrub-oak acorns are small and were swallowedwhole by the birds. The scrub oaks provide one of the reasons whythese birds have persisted in the scrubby plains.Leaves were found in the crops of many specimens examined byMr. Hoyle. In the spring months I found that the birds showeda great preference for the leaves of sheep sorrel (Rumex acetosella)over other plants such as clover and tender blades of grasses thatwere equally abundant in the same field. The distribution of thebirds on the field usually corresponded roughly with the distributionof this plant. One male specimen trapped on April 5, 1924, hadits crop gorged with leaves of sorrel. There were in this specimen,1,846 leaves and parts weighing 32.2 grams. Another bird trappedon May 20, 1924, had its crop completely filled with 38.6 grams ofseeds of the sheep sorrel. Although other leaves, such as those ofclover, alfalfa, and other herbaceous plants, were eaten, the birdsexhibited a decided preference for the leaves of the sorrel.Buds, including those of the scrub pine (Pinus virginiana) , wereeaten by certain birds during the winter months.The heath hen was especially fond of the grains of cultivated crops,such as corn, buckwheat, millet, and sunflowers, and in late years74564?32 19
278 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthese were planted on the reservation, especially for the use of theheath hen.The animal food of the heath hen consisted primarily of insects,chiefly grasshoppers, which were sometimes excessivety abundant onthe island late in summer and in autumn.There are very few data concerning the food of the young, but,judged from the food of the young prairie chickens, it probablyconsisted mainly of insects, especially in the case of the youngerbirds. The crop contents of one 5-day-old heath-hen chick acci-dentally killed on July 15, 1914, contained more than 80 insects repre-senting 10 species. The vegetable matter was merely incidental andnegligible in this specimen.Game.?The heath hen even when in its prime was never givena high place among game birds by the sportsmen. It was easily shot,because of its direct and laborious flight, and the habit of massingin flocks in the open fields made it too easy a victim for the pothunter and the market gunner. The ease with which the heath henwas tricked and killed readily accounts for the rapidity of its earlydisappearance from the mainland, whereas the clever ruffed grouseof the woodlands still holds its own and is ever ready to challengethe wits of the most skilled sportsman.The attitude of the sportsman in the past toward the pinnatedgrouse is well illustrated by the following excerpt from an articleby Elisha J. Lewis in his book The American Sportsman, 1885.So numerous were they a short time since in the barrens of Kentucky, andso contemptible were they as game birds, that few huntsmen would deign towaste powder and shot on them. In fact they were held in pretty much thesame estimation, or, rather abhorrence, that the crows are now, as they per-petrated quite as much mischief upon the tender buds of the orchards, as wellas the grain of the fields, and were so destructive to the crops, that it wasabsolutely necessary for the farmers to employ their young negroes to drivethem away by shooting of guns and springing loud rattles all around the planta-tion from morning till night. As for eating them, such a thing was hardlydreamed of, the negroes themselves preferring the coarsest food to this nowmuch admired bird.It is apparent that the heath hen was not considered an ideal birdfrom the point of view of the sportsman, and our efforts to save theheath hen were not made on the plea of its economic importance asa game bird. It is interesting to note, however, that it was thesportsman who took the initiative and who provided a large partof the funds to assist the State in the vain attempt to preserve thisinteresting race of birds.Enemies.?Man directly or indirectly is in part responsible forthe disappearance of the heath hen from most of its former range.Comparatively soon after the coining of the white man, it was drivenfrom one locality to the next, until it was forced to entrench on the
HEATH HEN 279
scrub-oak plains of Marthas Vineyard Island. It is a striking ex-ample of a bird that has not been able to adapt itself to the changingconditions brought about by civilization.Audubon (1840) wrote: "We frequently meet with the remains ofsuch [heath hen] as has been destroyed by the domestic cat whichprowls in the woods in a wild state." What was true in Audubon'sday was true in the more recent years of the heath hen, when catsranked high among the enemies of the bird. In addition to the catsreared on the island, large numbers were introduced by people whodumped them in the interior of the island when they left their sum-mer homes in autumn. A large part of the effort of the State De-partment and the special wardens in the control of vermin wasdirected toward the semiwild house cat.Large numbers of hawks are attracted to the island because ofthe abundance of mice and shrews, which live among the scrub oaks.Unfortunately, many of the hawks, for example the marsh hawk,which have a good reputation elsewhere, are frequently tempted, onMarthas Vineyard, to prey upon birds ; and when the heath hen wascommon, these birds were also numbered among their victims. Thegoshawks, notorious for their killing of game birds, played theirpart in the history of the heath hen. The most notable instance oftheir wholesale depredations was in the winter of 1916-17, followingthe destructive fire that swept over the island during the precedingspring. Other hawks, such as the red-shouldered, the rough-legged,the pigeon hawk, and others, as well as the different species of owls,were killed on sight by the wardens in charge of heath-hen protec-tion. It is probable that the wholesale killing of hawks and owls soupset the balance of nature that it acted as a boomerang to the heathhen.Disease was one of the most important factors in the recent de-cline of the heath hen and was one that man was unable to control.Blackhead is a disease common to poultry, but, so far as we knowat present, it is unusual in birds living in a free, wild state. Theheath hen, however, had the peculiar habit of congregating in theopen fields near farmhouses where poultry was kept. In most in-stances, chickens and turkeys had access to the fields visited by theheath hen, and thus the dreaded disease was readily transmittedto the native birds. Blackhead was found in the adult heath hen,and this is presumptive evidence that the disease was very destruc-tive to the young.Internal and external parasites were found on the few heath hensexamined, but these were all of minor importance as compared to thedisease blackhead.In the recent history of the heath hen it was well known that therewas a great excess of male birds. This abnormal ratio may have
280 BULLETIN 162, UNITED STATES NATIONAL MUSEUMbeen brought about in part by some hereditary influence, but it iscertain that this condition was aggravated by the fires that ravagedthe island during the breeding season. At such times the femaleswere destroyed on the nests, whereas the males escaped the con-flagration. Furthermore, a female with young was subject to moredanger of being killed than the male, which never cared for theyoung.Other factors that played their part were the excessive interbreed-ing, which was destined to occur after the heath hen was restrictedin range and to exceedingly small numbers of individuals. It wasalso found upon examination of dissected specimens that many ofthe birds were sterile.TYMPANUCHUS PALLIDICINCTUS (Ridgway)LESSER PRAIRIE CHICKENHABITSComparatively little seems to be known and still less has been pub-lished on the habits and distribution of the small, light-colored, lesserprairie chicken, which is found in the Upper Sonoran Zone of theGreat Plains from Kansas and Colorado to central Texas and easternNew Mexico. It has disappeared from many sections where it wasonce abundant; too much grazing on, and extensive cultivation of, thegrassy plains have driven it out. But it is still to be found in fairnumbers in its restricted range, where it is protected, or not disturbed.We are greatly indebted to Walter Colvin for the information thatfollows regarding this fine bird, which I have gathered from hispublished article (1914) and from the full notes and photographs hehas sent to me. Writing of its distribution and haunts in 1914, hesays:The natural habitat of this beautiful grouse is far remote from the habitatof its allied cousin, the heath hen, and still less remote from its nearer cousin,the common prairie heu of the Middle States. Its present confine is thesouthwestern counties in Kansas, extending west from Meade, through Seward,Stevens, and Morton and north into Stanton, Grant, and Haskell counties, cross-ing the line into Colorado some fifty miles, extending south through Beaver,Texas, and Cimarron counties of Oklahoma, into the panhandle of Texas, buthow far south and east I haven't sufficient data at hand to determine, althoughI believe it is safe to assert that they do not extend farther south into Texasthan two degrees by air line. In northwestern Oklahoma I have seen thechickens within a few miles of the New Mexico line.Formerly this variety of chickens was common in Woodward County, Okla-homa, and Captain Bendire, in his Life Histories, mentions securing their eggsnear Fort Cobb, Indian Territory, in 1S70. At that time reliable informationgoes to show that they were far more plentiful south of the great Indian high-way than north. The pan-handle is a typical bunch-grass country, and duringthe early eighties a great prairie fire broke out in its southern extremity, sweep-
LESSER PRAIRIE CHICKEN 281ing north to the narrow strip of short grass land in " No Man's Land," where itdied. The chickens that were driven north found an ideal home in the rolling,sandy bunch-grass country that abounded just across the line.Their range in its entirety would probably cover no greater area than a fourthof the State of Kansas, and the most abundant nucleus is in Stevens andMorton counties. Here they are quite plentiful in its sandhill and bunch-grassfastness, where, in the fall of the year they sometimes gather in flocks ofseveral hundred birds, roaming where they will, a typical bird of the long-grasscountry.Courtship.?Concerning the courtship of this species, Colvin says
:
The nuptial performances of the cocks are similar to those of the commonvariety, but the ventriloquial drumming sound does not appear to be quite sorolling or voluminous. In May, 1907, I put up in the heart of the nesting-ground, where I had an excellent opportunity to study their habits. The cocksgenerally select for drumming-ground a slight rise covered with buffalo grass,where they gather each spring for the nuptial performance. They are verypartial to their drumming-grounds, and even though disturbed will return totheir old haunts year after year. I saw one drumming-ground that had beenused for many years.Here the cocks would gather sometimes as high as fifty birds to perform theirantics. The drumming of so many cocks would be of such volume as tosound like distant thunder. Hens attracted by the drumming would causedisturbance. Cock fights and a general all-around rumpus would begin. Agreat deal of strutting and clucking would be done by the males. Finally,when, with lowered head and wings and air-sacks full, a successful cockwould drive his hen from the bunch, peace would reign again, and the drummingwould be resumed.Nesting.?A nest that Colvin found near Liberal, Kans., on May28, 1920, was " located in a bunch of sage, growing in a swale justbelow a brow of a hill ; " it consisted of " a hollow scooped out in thesand and lined with grasses. So well concealed was the nest thatone could observe only a small portion of the female as she sat uponthe nest. Disturbing the sage brush she left the nest, disappearingover the hill. The nest contained 12 eggs on the point of hatching."Another set of eggs, which he kindly presented to me, was takenin the same general region on June 2, 1920; these eggs were onlyslightly incubated. He says in his notes
:
On John Napier's farm I was shown a nest of the lesser prairie hen. Itwas placed under the south side of a bunch of sage, and was a mere hollowin the sand lined with grasses. When I saw the nest it had been exposed bya corn lister. The nest originally contained 12 eggs, but one had been brokenby the lister, leaving 11. Mr. Napier informed me that the team had passedover the nest and sitting hen twice before she was finally raked off by thedoubletree.Of a third nest he writes
:
Through the efforts of an old-time trapper, Ed. Ward, I was successful insecuring a set of 13 straw-buff-colored eggs. The nest, a mere hollow in thesand, was lined with a few grasses, and was situated under a tumbleweed,which had lodged between two tufts of grass on the north side of a sloping
282 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMhillock. The sitting hen allowed us to approach quite close before takingwing.Mr. Ward informed me that the nests were almost invariably placed on topof a rise, or on its sloping sides. The nests, though usually placed in opensituations, are extremely difficult to find, owing to the dichromatic arrange-ment of the feathers, which so harmoniously blends with the surroundingsof the sitting bird. A far greater protection to the sitting hens is their non-scent-giving powers during the nesting season, which was fully demonstratedthe following spring, when I again visited that vicinity in order to securea series of photographs.In company with one of the best-known chicken dogs, I thrashed over severalsections of bunch grass land where chickens were common and known tonest each year, but without success. I found no hens off the nests during theheat of the day, but quite frequently saw them flying to the feeding groundsafter twilight. Several times while hunting their nests I felt sure that I waswithin a few feet of the sitting birds, but was compelled to give up the search.The hens are close, hard sitters, and very few nests are found. Frairie firesexpose many nests and are the nesting hen's worst enemy.Eggs.?The full set seems to consist of 11 to 13 eggs, so far as weknow-. The eggs are ovate in shape, smooth, and rather glossy. Thecolors vary from " cream color " to " ivory yellow ' ; in my set ; mostof the eggs are sprinkled with very fine dots of pale brown or olive
;
but some are nearly or quite immaculate. Mr. Colvin calls them strawcolor or straw buff. Bendire (1892) says: "The ground color variesfrom pale creamy white to buff. The markings, which are all veryfine, not larger than pin-points, are lavender colored. More thantwo-thirds of the eggs are unspotted, and all look so till closelyexamined."The measurements of 47 eggs average 41.9 by 32 millimeters; theeggs showing the four extremes measure 43.5 by 33.5, 40.5 by 33, and40.7 by 30.4 millimeters.Plumages.?I have never seen a downy young of the lesser prairiechicken, but probably it is much like the chick of its northern rela-tive. The sequence of molts and plumages is doubtless similar tothose of the prairie chicken. A young bird, about one-third grownbut in full juvenal plumage, has the crown and occiput mottled with
" tawny " and black ; the chin and throat are white ; the feathersof the back and scapulars are variously patterned with transversebars of " ochraceous-tawny," " tawny-olive," " cinnamon-buff," andblack, with median white stripes or tips; the central tail feathersare similarly barred and tipped, the pattern diminishing on lateralrectrices; the underparts are dull whitish, heavily spotted or barredon the breast and flanks with black, sepia, and pale dusky, darkeston the chest, and more or less heavily tinged on the flanks andbreast with " ochraceous-buff."Food.?Colvin (1914) says that " during the summer months theyfeed largely on grasshoppers, but in the fall and winter they feed
LESSER PRAIRIE CHICKEN 283almost entirely on kaffir corn and maize, cane seed, and other va-rieties of semi-arid cereals. As to the palatableness of the meatI much prefer duck."Behavior.?The same observer says:In general characteristics and makeup the lesser prairie hen is of a sturdy,robust nature, being some two-thirds the size of the common prairie hen.They are veritable dynamos of " git up " and energy. Such vivacity andactivity I have never seen displayed in any other game bird. On a cold,snappy day they have the life and energy of half a dozen quails, and for speedthey put their first cousin to shame. A full-grown cock, well fatted, willweigh from a pound eleven ounces to a pound fourteen ounces.Game.?Judged from Mr. Colvin's published accounts (1914 and1927) the lesser prairie chicken must be a fine game bird, sufficientlywild and swift of wing to make sporty shooting, and large andplump enough to make a desirable table bird. These birds werewonderfully abundant in earlier days, as a few quotations from Mr.Colvin's writings will show. " In a cane field near the State Line,"he says, "we saw a flock of 500 or more, and when they arose itseemed that a hole had been rent in the earth." He wanted tostop and shoot a feAv, but his companion urged him on, saying,
" Those are only rovers. I'll show you some chickens when we getup in the State." Evidently he made good, for Mr. Colvin (1914)writes
:
Two miles farther along we came to Ed Ward's. He informed us thatthere were a " few " chickens in a cane and kaffir corn field a quarter of amile east. We flushed several birds from the tall bunch grass just beforewe reached the field, which were promptly despatched ; however, in the fieldthings became more lively. Such a sight I have never seen before nor since.Chickens were flushing everywhere, and droves of fifty to a hundred would takedown the corn rows, sounding like a moving avalanche as they touched theblades of corn. Still birds were quite wary, and the only good shots were to behad over the dog.As we thrashed back and forth across the grain field, the chickens arosein flocks of fifty to five hundred, and generally sixty to eighty yards distant,making shooting difficult. The majority of the birds, after being flushed, wouldfly back into the field, while some would go to the bunch-grass covered hills halfa mile away. Mr. Ward and I estimated that there were from thirty-fivehundred to four thousand chickens in this one field, a sight never to beforgotten.A few years later, hunting over the same ground, he found thechickens much diminished in numbers; his thoughts are expressedas follows:Gathering our duds together, we started for our long journey home. A fewclouds, fringed with gold, freckled the western sky, and over all a red mantlewas cast while the sun slowly lowered to the horizon. My mind went back tothe events of the day and to the time when the chickens were more plentiful,and I realized with a shudder that we were nearing the sunset life of the kingof upland game birds. But the decrease in their numbers is not due so much to
284 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthe gunners, as gunners are few per capita in those parts, but is due largely tothe cutting up of this vast wilderness into small farms. The bunch-grass landcan not be mowed for hay ; therefore, in such land the chickens have found anideal home in which to rear their young and harbor themselves during thewinter. Such land is soon destroyed by cultivation and small pastures. Withthe advancement of civilization the flocks scatter and become depleted.He wrote to me in December, 1927
:
The saving of the lesser prairie hen for future time is assured, as some recentlaws were enacted in Kansas that gave the game commission the power thatenables them to close and open seasons without any special legislation fromthe State. After visiting the nesting grounds this summer and noticing thatmany of the birds had been destroyed, I took it up with the State game warden,J. B. Doze, and Lee Larabee of the commission, and they established the closeseason this year. We are therefore assured of a good crop of birds nextyear.Winter.?Of their winter habits, Colvin says:Though naturally lovers of the free range, during the winter they rely largelyupon the farmer and rancher for their food. A large amount of grain is con-sumed by the flocks as they roam from one grain field to another. In theeighties a man by the name of Hatch settled in the sandhills just inside theKansas line in Seward County. Here he planted a grove of black locust treesand spread out his broad fields of maize and kaffir corn. The Texas bobwhites,mountain quail, and lesser prairie hens soon learned that this man was afriend of the birds, and straightway made it their rendezvous. Here, eachfall, the chickens gathered by the thousands, and each spring spread out overthe vast prairies, nesting and rearing their young. In the fall of 1904 mybrother estimated that he saw in a single day, 15,000 to 20,000 chickens in andaround this one grain field. Though timid if persecuted, if unmolested theybecome quite tame, coming to the barn lots to feed, and will put as much con-fidence in man as quails when protected.DISTRIBUTIONRange.?The Great Plains region, from southeastern Colorado andKansas south to west-central Texas and probably southeastern NewMexico.Breeding range.?The breeding range of the lesser prairie chickenextends north to southeastern Colorado (Gaumes Ranch and Holly)and southwestern Kansas (Cimarron). East to southwestern Kan-sas (Cimarron) and Oklahoma (Ivanhoe Lake, Fort Reno, and FortCobb). South to southwestern Oklahoma (Fort Cobb); northernTexas (Mobeetie and Alanreed) ; and east-central New Mexico(Portales). West to east-central New Mexico (Portales) ; andsoutheastern Colorado (Cimarron River and Gaumes Ranch).Winter range.?Confined chiefly to central Texas. North toMonahans, Midland, and Colorado City. East to Colorado City,Middle Concho River, and Bandera. South to Bandera, FortClark, and the Davis Mountains. West to the Davis Mountains
NORTHERN SHARP-TAILED GROUSE 285and Monahans. Casual in winter at Lipscomb in the Panhandlearea, and it appears probable that some winter in southeastern NewMexico (vicinity of Carlsbad).No information is available relative to the movements of this speciesbetween breeding and wintering areas.Casual records.?Widmann (1907) reports a specimen in the Hurtercollection said to have come from southwestern Missouri, and that inJanuary, 1877, large numbers were shipped to Fulton market, NewYork City, from Pierce County. Neff (1923) states the species wasnoted in Lawrence County, Mo., in 1887. A specimen in the collec-tion of the Academy of Natural Sciences of Philadelphia was takenbetween January 24 and 28, 1894, near Garneth, Kans., while anotherwas collected at Oakley, Kans., January 1, 1921. The exploringparty of Capt. John Pope collected two specimens (later made thetypes of the species) on the Staked Plains, N. Mex., on March 3 andMarch 11, 1854.Lesser prairie chickens also have been reported from Nebraska, butin the lack of specimen evidence it is thought that the records referto T. c. americanus. At the present time the Arkansas River appearsto be a very definite northern boundary to their range.Egg dates.?Colorado to Texas: 12 records, May 5 to June 12;6 records, May 20 to June 1.PEDIOECETES PHASIANELLUS PHASIANELLUS < Linnaeus)NORTHERN SHARP-TAILED GROUSEHABITSAs the specimen on which Linnaeus bestowed the type name of thespecies came from the Hudson Bay region, the name P. p. phasianel-lus is now restricted to the dark-colored race, which ranges throughthe forested regions of northern Canada to central Alaska. Its centerof abundance seems to be in the vicinity of Great Slave Lake, Macken-zie. Swainson and Richardson (1831) say that " it is found through-out the woody districts of the fur-countries, haunting open glades orlow thickets on the borders of lakes, particularly in the neighbourhoodof the trading-posts, where the forests have been partially cleared."According to Major Bendire (1892) it was found breeding at FortRae, in latitude 63? N., and at Fort Good Hope, in the MackenzieRiver Basin. MacFarlane (1908) found it breeding in the valley ofthe Lockhart and Anderson Rivers, where two nests were found, butthe eggs were afterwards lost. Herbert W. Brandt says in hisAlaska notes:The sharp-tailed grouse proved to be the most common gallinaceous bird weencountered during the early stages of our dog-sled trip to Hooper Bay. Wefirst collected it on March 22, when two handsome males were taken, but small
286 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMflocks were seen from time to time during the previous day. After we reachedthe Koskokwim River the birds became scarcer, and we did not see it at allalong the Yukon River. The center of abundance of this grand grouse appearsto be in the vicinity of Lake Minchumina, with its sheltered, scenic, birch-cladhills, that transcends in exquisite beauty any region I saw in glorious Alaska.Nesting.?The nesting habits of this grouse are apparently similarto those of its more southern relatives. The only published accountof it I can find is by Baird, Brewer, and Ridgway (1905), as follows
:
Mr. Kennicott found the nest of this bird at Fort Yukon, at the foot of aclump of dwarf willows. It was in dry ground, and in a region in whichthese willows abounded and were quite thickly interspersed with other trees,especially small spruces, but no large growth. The nest is said to have beensimilar to' that of Cupidonia cupido. Mr. Lockhart also found it breeding inthe same region. The nests seen by him were likewise built on a rising groundunder a few small willows.Eggs.?Major Bendire (1892) describes the eggs very well, asfollows
:
The number of eggs to a set varies from seven to fourteen and their groundcolor from a fawn color with a vinaceous rufous bloom to chocolate, tawny, andolive brown in different specimens. The majority of the eggs are finely markedwith small, well-defined spots of reddish brown and lavender, resembling themarkings found on the eggs of Tympanuchus americanus, only they are muchmore distinct. Compared with the eggs of the two southern subspecies,P. phasianellus columbianus and P. phasianellus campestris, they usually arevery much darker colored, even the palest specimens being darker than theheaviest marked eggs of either of the two subspecies. These markings areentirely superficial, and when removed leave the shell a creamy white in somecases and a very pale green in others. In shape they are usually ovate.The measurements of 27 eggs average 43.1 by 32.3 millimeters;the eggs showing the four extremes measure 48 by 33, 44.5 by 33.5,40.5 by 32, and 42 by 30 millimeters.Food.?Swainson and Richardson (1831) say: "They feed on thebuds and sprouts of Betula glandidosai, of various willows, and ofthe aspen and larch, and in autumn on berries." Macoun (1909)says : " These birds keep in pairs or small flocks and frequent thejuniper plains all the year. The buds of these shrubs are theirprincipal food in winter, as their berries are in summer."DISTRIBUTIONRange.?Alaska, western Canada, and the Western United States.The full range of the sharp-tailed grouse extends north to Alaska(Allakakat and Fort Yukon) ; Yukon (Ramparts) ; Mackenzie (FortGood Hope, Fort Norman, Grandin River, Fort Rae, Fort Resolu-tion, and Fort Smith) ; northeastern Alberta (Fort Chipewyan)
;
east-central Saskatchewan (Cumberland House) ; Manitoba (GrandRapids, Norway House, Oxford House, and York Factory) ; north-
NORTHERN SHARP-TAILED GROUSE 287ern Ontario (Severn House and Fort Albany) ; and Quebec (GreatWhale River). East to Quebec (Great Whale River and FortGeorge) ; eastern Ontario (Lake Abitibi, Lake Temiskaming, ParrySound, and Beaumaris) ; formerly northeastern Illinois (Wauke-gan) ; formerly east-central Iowa (Grinnell) ; and formerly centralKansas (Fort Hays). South to formerly Kansas (Fort Hays, Ellis,and Banner) ; New Mexico (Raton) ; Colorado (Pagosa Springs,Fort Lewis, and Cortez) ; southern Utah (Parawan Mountains) ; andformerly northern California (Canoe Creek and Fort Crook). Westto formerly northern California (Fort Crook and Camp Bidwell) ;Oregon (Fort Klamath, Caleb, and The Dalles) ; Washington (Top-penish and Dosewallops River) ; British Columbia (Nicola, Earn-loops, Cariboo Road, 158-mile House, Quesnelle, and Hudsons Hope) :southwestern Yukon (Tagish Lake, Alsek River, and Lake Kluane)
;
and Alaska (Kolmakof, Holy Cross, Tacotna, Lake Minchumina, andAllakakat).Sharptails are not now known to breed east of Ontario, Minne-sota, South Dakota, and Colorado, and apparently they have beenentirely extirpated from Iowa, Kansas, and California. The speciesoccurs only as a summer straggler in western Alaska (Allakakat,Holy Cross, Lake Minchumina, and Kolmakof), and as a winterstraggler in southeastern Ontario (Lake Abitibi and Temiskaming)and Quebec (Great Whale River). It has, however, occurred insummer at Fort George, Quebec. In some winters it is abundantat Vermilion and Watertown, S. Dak.Migration.?In common with some of the ptarmigans, there ap-pears to be a definite migration from the northern part of the range,governed by the severity of winter conditions and the available foodsupply. This exodus, however, does not extend south of the breedingrange. It has been observed, apparently, only by E. A. Preble, whonoted flocks moving southward at Fort Norman, Mackenzie, on Oc-tober 1, 1903, and who detected early spring arrivals at Fort Simp-son, Mackenzie, on March 12, 1904.The range as outlined is for the entire species, which has beenseparated into four subspecies. The northern sharp-tailed grouse(P. p. phasianellus) occupies the northern part of the range southto Lake Superior, the Parry Sound district (casually) of Ontario,and the Saguenay River, Quebec. The Columbian sharp-tailed grouse(P. p. columbianus) is found from the interior lowlands of BritishColumbia south (formerly) to northeastern California, Utah, Colo-rado, and northern New Mexico. The range of the prairie sharp-tailed grouse (P. p. campestris) extends from eastern Colorado.Kansas, northern Illinois, and Wisconsin north to southern Manitoba,Saskatchewan, and Alberta.
288 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMEgg dates.?Northern Canada: 12 records, May 1 to June 3; 6records, May 16 to 21.Washington and Oregon: 9 records, April 6 to June 18. Mon-tana: 5 records, May 19 to June 5. Alberta to Manitoba: 52records, May 2 to June 22 ; 26 records, May 30 to June 11. Dakotasand Minnesota: 27 records, May 2 to June 26; 14 records, May 19to June 8. Colorado : 2 records, April 1 and May 8.PEDIOECETES PHASIANELLUS COLUMBIANUS (Ord)COLUMBIAN SHARP-TAILED GROUSEHABITSThis is the grayest of the three races of sharp-tailed grouse; thenorthern race, typical phasianellus, is much darker, and the easternrace, campestris, is more buffy or rufous. This western race inhabitsthe lowlands of the Great Basin from the Rocky Mountains to theCascades and Sierra Nevadas. It was discovered by Lewis andClark on the plains of the Columbia River in 1805 and was namedby Ord in 1815. As it is not so widely distributed or so well knownas campestris, the reader is referred to the prairie sharp-tailed grousefor the life history of the species.Major Benclire (1892) says of this grouse:It is one of the most abundant and best known game birds of the Northwest,inhabiting the prairie country to be found along the foothills of the numerousmountain chains intersecting its range ; seldom venturing into the wooded por-tions for any distance, and then only during the winter months, when it ispartially migratory in certain sections. According to my own experience theColumbian Sharp-tail breeds more frequently on the sheltered and sunny slopesof the grass-covered foothills of the mountains than in the lower valleys andcreek bottoms.As to its past status in California, Grinnell, Bryant, and Storer(1918) say:When Newberry, Cooper, Henshaw, and other early naturalists were mak-ing observations upon the fauna of California, previous to 1880, they foundthis species numerous in the plateau region northeast of the crest of the Sierra-Cascade range. Since then, man's occupancy of that territory, and uncon-trolled levy upon its birds for food or sport, has resulted in the apparentlycomplete disappearance of this species. They say : Coming north from SanFrancisco, we first found it on a beautiful prairie near Canoe creek (nearCassel, Shasta County), about fifty miles northeast of Fort Reading; subse-quently, after passing the mountain chain which forms the upper canon ofPit River, we came into a level, grass-covered plain, through which the wil-low-bordered river flows in a sinuous course like a brook through a meadow(probably near Lookout, Modoc County). On this plain were great numbersof birds of various kinds, and so many of the sharp-tailed grouse, that, for two
COLUMBIAN SHABP-TAILED GROUSE 289
or three days, they afforded us fine sport and an abundance of excellent food.We found them again about the Klamath lakes.Grinnell, Bryant, and Storer (1918) say that it is " now almost orentirely extirpated " as a California bird, and that " the disappear-ance of this bird can be attributed to no other cause than to its in-cessant pursuit by man. As long as a single bird remained huntingpersisted. Moreover the fact that this grouse prefers grassy locali-ties, just such as are selected for ranch sites, indicates another ofthe factors that led to its extermination."The southern limit of its present range seems to be in northernNew Mexico, where Mrs. Florence M. Bailey (1928) says of it:Though naturally a bird of more northern country with abundant rank grassfor breeding places, the high altitude of the grassy, broken-rimmed mesas north-east of Raton, some 8,000-9,000 feet in elevation, " appears to create a littleworld suitable to it in New Mexico," which, the oldest settlers attest, has longbeen inhabited by it.Nesting.?Major Bendire (1892) says of its nesting habits:Nidification began usually from about April 15 to May 1, according to theseason. I found a set of fifteen eggs, which had been sat upon about a week orten days, on April 22, 1871. Some birds must have laid earlier still, as it wasno uncommon sight to find fully grown birds by July 10. All the nests of thisspecies which I examined were invariably well concealed and rather difficultto find. You might search daily for a couple of weeks and be unsuccessful infinding a nest, and again you might stumble on two or three on the same day.A bunch-grass covered hillside, with a southerly exposure, seemed to be afavorite nesting site with this Grouse at Fort Lapwai, while at Camp Harney,Oregon, they confined themselves during the breeding season to the sage brushcovered plains of the Harney Valley, interspersed here and there with a lowgrassy swale, nesting along the borders of these, where the grass attained aheavier growth. The nest, like that of all the Grouse, is always placed on theground, usually close alongside some tall bunch of coarse grass, which hidesit completely from view. Even if it did not, the female harmonizes in colorso thoroughly with her surroundings that she is not apt to be noticed, unlessshe should leave her nest, which she does not do very readily, as she is a veryclose sitter. A slight hollow, usually scratched out on the upper side of abunch of grass, if the nest is placed on a hillside, is fairly lined with dry grass,of which there is ordinarily an abundance to be found in the vicinity, and thisconstitutes the nest. A few feathers from the lower parts of the bird areusually mixed in among the eggs, each one of which is often imbedded abouttwo-thirds in its own mould and does not touch the others. Once only did Ifind the eggs placed on top of each other, eight in the lower and five in theupper layer.Eggs.?The eggs of the Columbian sharp-tailed grouse are indis-tinguishable from those of its prairie relative. The measurementsof 58 eggs average 43.3 by 32.1 millimeters; the eggs showing thefour extremes measure 46.5 by 34.5, 39 by 31, and 42.5 by 30.5millimeters.
290 BULLETIN 162, UNITED STATES NATIONAL MUSEUMYoung.?Major Bendire (1892) writes:The young are active, handsome little creatures, and able to use their legs atonce on leaving the shell. They are at first fed mostly on insects, young grass-hoppers and crickets forming the principal portion of their bill of fare. Theformer are always abundant and easily obtained; later, when the young areable to fly, the mother leads them to the creek bottoms, where they find anabundance of berries and browse. They are especially fond of the seeds of thewild sunflower, which grows very abundantly in some places, and when these areripe, many of these birds can be found in the vicinity where these plants grow.Coues (1874) says:The young, as usual among gallinaceous birds, run about almost as soon asthey are hatched ; and it is interesting to witness the watchful solicitude withwhich they are cherished by the parent when she first leads them from the nestin quest of food, glancing in every direction, in her intense anxiety, lest harmbefall them. She clucks matronly to bring them to brood under her wings orto call them together to scramble for a choice morsel of food she has found.Should danger threaten, a different note alarms them ; they scatter in everydirection, running, like little mice, through the grass till each finds a hidingplace; meanwhile, she exposes herself to attract attention, till, satisfied of thesafety of the brood, she whirrs away and awaits the time when she may reas-semble her family. In the region where I observed the birds in June and July,they almost invariably betook themselves to the dense, resistant underbrush,which extends for some distance outward from the wooded streams, seekingsafety in this all but impenetrable cover, where it was nearly impossible tocatch the young ones, or even to see them, until they began to top the bushesin their early short flights. The wing and tail-feathers sprout in a few daysand are quite well grown before feathers appear among the down of the body.The first coveys seen able to rise on wing were noticed early in July ; but bythe middle of this month most of them fly smartly for short distances, beingabout as large as Quails. Others, however, may be observed through August,little, if any, larger than this, showing a wide range of time of hatching, thoughscarcely warranting the inference of two broods in a season.Behavior.?The habits of this grouse are essentially the same asthose of the prairie sharp-tailed grouse, but Bendire (1892) observes
:
The habits of the Columbian Sharp-tailed Grouse vary very materiallyin different portions of the country where I have met with them. At FortKlamath, Oregon, where they are rather rare, I have found them inhabitingdecidedly marshy and swampy country, and keeping close to, if not in the edgesof, the pine timber throughout the year. At Fort Custer, Montana, this Grouse,during the winter, was much more arboreal than terrestrial in its habits, movingaround on the limbs of the large cottonwood trees as unconcernedly as on theground ; spending in this way almost all their time, except when feeding. AtHarney, Oregon, and Lapwai, Idaho, they might be frequently seen in small treesand bushes which grow along the creeks, but scarcely ever in large trees, ofwhich there was an abundance. Here, they uttered very few notes at any time,while at Fort Custer I have frequently heard them cackling in the tall cottonwoods which grew along the Big Horn River bottom, before I had approachedwithin several hundred yards of them, evidently giving notice to other birds inthe vicinity of my coming. This fine game bird is decreasing very rapidlythroughout its range. It does not seem to prosper in the vicinity of man, and
PRAIRIE SHARP-TAILED GPvOUSE 291
as the country is becoming more and more settled, it recedes before civilization.As it is not a particularly shy bird, it falls an easy victim to the gunner.PEDIOECETES PHASIANELLUS CAMPESTRIS RidgwayPRAIRIE SHARP-TAILED GROUSEHABITSOn my various trips to North Dakota, Manitoba, and Saskatche-wan, I became quite familiar with the sharp-tailed grouse of theeastern plains. It is not so much a bird of the open prairie as is theprairie chicken; but we found it very common in the sandhills,among the willow thickets, and on the low, rolling hills overgrownwith shrubbery. Its range is becoming more and more restricted asthe Central West becomes more thickly settled and more land comesunder cultivation. In some places it is decreasing in numbers whereprairie chickens are increasing. Edw}Ti Sandys (1904) writes:It has been claimed by more than one well-known' expert that the sharptailand pinnated grouse are bitter foes, but this I am inclined to doubt. I am wellaware of the belief among western sportsmen that the one species drives theother from its haunts, but believe that the true reason for the supplanting of onespecies by the other is nothing more than the closer settlement of what a fewyears ago were wild regions. In other words, one bird follows the farmer, whilethe other retreats before him.Courtship.?The courtship performance of the sharp-tailed grouseis no less interesting than that of other grouse. It is quite similar tothat of the prairie chicken and the heath hen; perhaps not quite sogrotesque but more animated. These birds have favorite spots,generally small knolls, to which they resort for this purpose everyspring ; these are known as " dancing hills." Frank L. Farley writesto me from Alberta
:
On my farm at Dried Meat Lake, an average of a dozen pairs nest every year.There are two dancing hills on this farm that the Indians told me had beenused as long as they could remember ; one is a little knoll right overlooking thelake, and the other is half a mile away. I have seen as many as 50 birdsdancing on each hill at a time ; that is, waiting until the ground was vacatedby the previous dancers. They would wait patiently for their turn. Thesedances take place every April and May, and often the grain, when up, is trampedentirely away. I can generally get up to within 25 feet of the dancers withmy car to watch them.Dr. D. G. Elliot (1897) gives a very good account of the " danc-ing," as follows:In the early spring, in the month of April, when perhaps in many parts oftheir habitat in the northern regions the snow still remains upon the ground,the birds, both males and females, assemble at some favorite place just as dayis breaking, to go through a performance as curious as it is eccentric. Themales, with ruffled feathers, spread tails, expanded air sacs on the neck, headsdrawn toward the back, and drooping wings (in fact, the whole body puffed
292 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
out as nearly as possible into the shape of a ball on two stunted supports),strut about in circles, not all going the same way, but passing and crossing eachother in various angles. As the " dance " proceeds the excitement of the birdsincreases, they stoop toward the ground, twist and turn, make sudden rushesforward, stamping the ground with short quick beats of the feet, leaping overeach other in their frenzy, then lowering their heads, exhaust the air in thesacs, producing a hollow sound that goes reverberating through the still air ofthe breaking day. Suddenly they become quiet, and walk about like creatureswhose sanity is unquestioned, when some male again becomes possessed andstarts off on a rampage, and the " attack " from which he suffers becomes in-fectious and all the other birds at once give evidences of having taken the samedisease, which then proceeds with a regular development to the usual con-clusion. As the sun gets well above the horizon, and night's shadows have allbeen hurried away, the antics of the birds cease, the booming no longer re-sounds over the prairie, and the Grouse scatter in search of food, and in pursuitof their daily avocation. While this performance is always to be seen in thespring, it is not unusually indulged in for a brief turn in the autumn, and whileit may be considered as essentially a custom of the breeding season, yet likethe drumming of the Ruffed Grouse, it may be regarded also as an exhibitionof the birds' vigor and vitality, indulged in at periods of the year even whenthe breeding season has long passed.Ernest Thompson Seton (Thompson, 1890) says
:
The whole performance reminds one so strongly of a Cree dance as to sug-gest the possibility of its being the prototype of the Indian exercise. Thespace occupied by the dancers is from 50 to 100 feet across, and as it is re-turned to year after year, the grass is usually worn off and the ground trampleddown hard and smooth.Hamilton M. Laing (1913) noticed that the birds danced in pairsand that each pair usually kept to a certain section of the hill. Hedescribes a lively fight he saw as follows
:
The fun had reached its frenzy pitch when suddenly I noted that somethingother than dancing was taking place. It very much resembled a fight; andsoon I realized that such it really was, though it had a most absurdly comicside to it. The fray was a three-cornered affair. The first fellow fled in circles
;
the second followed him ; and the third brought up the rear. I decided that itwas two cocks fighting, and that the cause at issue, and root of the trouble, wasmerely following the contestants. They whirled about the hill at lightningspeed, running on legs that fairly spun, or dashing short snatches on the wing,through the set or over the dancers. The second fellow had blood in his eye,and the first?evidently an interloper, who was not wanted?lacked the courageor fiber to turn and fight it out. Yet, when the pursuer caught him, they bitand held on with a grip like bull-dogs, and rolled over, and beat each otherwith their wings, and shed each other's feathers. The interloper always gotthumped, but not until he was properly mauled would he retreat.E. S. Cameron (1907) writes:At this date (April 18) the ball is opened by a single cock making a runacross the open space as fast as he can use his legs, the tail being inclinedstiffly over the back, while the wings are dragged, so that a large white area isexposed behind. The vivid yellow supraciliary fringe is erected, and, all thefeathers of the neck standing on end, a pink inflated sac is disclosed. At the
PEAIEIE SHARP-TAILED GROUSE 293same time the head is carried so low as almost to touch the ground, so that thebird is transformed in appearance and, as observed through binoculars at somedistance, looks to be running backwards. He then returns at full speed, whenanother cock comes forward toward him, both advancing slowly, with vibratingtails, to meet finally and stand drumming their quills in a trance with tightlyclosed eyes. After perhaps a minute one bird peeps at the other, and seeinghim still enraptured, resumes an upright graceful carriage, anon stealinggently away. His companion is thus left foolishly posing at nothing, butpresently he too awakes, and departs from the arena in a normal manner.Meanwhile the remaining cocks, one after another, take up the running tillall have participated, but the end of each figure seems to be the same. Twobirds squat flat on the ground with their beaks almost touching for about twentyminutes, and when they do this they are out of the dances for that day. Thedance appears to terminate by some bird, either a late starter or one morevigorous than the rest, being unable to find a partner to respond to his run.Having assured himself of this, he utters a disgusted clucking, and all thegrouse fly away at intervals as they complete their term of squatting.Nesting.?The sharp-tailed grouse is not very particular as to anesting site and is not much of a nest builder. The nest is a hollowin the ground, scantily lined with whatever loose material is avail-able. It is usually partially concealed under a thick tuft of tallgrass on the prairie or under bushes or thick herbage in a bushytract or near a stream. We found these grouse very common aroundCrane Lake in 1905. On one day, June 5, we discovered three nestson the bushy prairies near Bear Creek. One of the nests that Iphotographed was under a little rosebush; the hollow was 7 inchesacross and 2y2 inches deep ; it was lined with fine twigs, straws, andfeathers. Another similar nest was under a " grease bush," or
" silver willow." A nest found June 2, 1913, near Lake Winnipeg-osis, Manitoba, was a hollow in the ground 8 inches across and 4inches deep ; it was concealed in long grass on a grassy and bushydry place on a low, wet prairie near the lake. We found that thebirds were usually close sitters, flushing almost underfoot. In onecase, where we wanted to photograph the bird on a previouslylocated nest, we had difficulty in finding it again, although we hadmarked the spot with a tuft of cotton. After scanning the groundvery carefully, foot by foot, for a long time, wre finally discoveredthe bird sitting on her nest within less than 10 feet. She shows upplainly enough in the photograph, but in life her color patternmatched her surroundings so wTell that she was nearly invisible,though in plain sight.Dr. Alfred O. Gross has sent me his notes on five nests found by himin Wood County, Wis., June 1 to 4, 1930. Following are his notes onthree of them, containing 13, 11, and 12 eggs, respectively
:
Nest built among the grass and moss of a rounded knoll about 4 feet indiameter, located in a wet, swampy region remote from any farmed land.Growing throughout the swamp were small willows, poplars, and swamp grass.74564?32 20
294 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
It was lined chiefly with grass, much of which was the bent-over tips of thegrass growing about the nest. The nest and eggs were partially concealed bythe overhanging grasses.Nest lined with oak leaves, dry ferns, grass, and a few feathers of the nestingbird. About three-fourths of the lining consisted of oak leaves, which hadfallen from a scrub oak near by. About the nest was a thick growth of mossand checkerberry (Gaultheria). Very near to the nest was a thicket of pincherries, poplars, and willows, and here and there were masses of blueberrybushes about a foot in height. The country about the nest was a sand plain,with a few marshy sloughs, but most of the land was brushy, chiefly willows,cherry, hazel, black and scrub oaks, and poplars.This nest was located near the margin of a scrub-oak area that bordered anextensive grassy marsh. The edge of the marsh was only a few feet distant.The nest was very shallow and lined with sticks, grass, and a few leaves andgrouse feathers. Growing about the nest were vines and grass, and the wholewas well concealed by a mass of blueberry bushes.Gross saj^s further
:
Judged from the situations in which these five nests were found, the sharp-tailed grouse chooses a nesting site remote from farms and in places wherethere is considerable brushy growth. The prairie-chicken nests, however, arenearly all near farms or on the farmed fields, such as the meadows and alfalfaand clover fields. Though the nesting sites vary considerably, the birds arefrequently seen together during the hunting season, and Mr. Cole, local gamewarden at Wisconsin Rapids, states that on one occasion he killed a prairiechicken and a sharp-tailed grouse with one shot. Several times in traversing theprairies I have seen both species together, this summer (1930).Eggs.?From 10 to 13 eggs generally constitute the full set for thesharp-tailed grouse, but as many as 14 or 15 are sometimes found in anest. Thejr are ovate in shape, and the shell is smooth, with a slightgloss. They are quite dark colored when first laid and have a pur-plish bloom, but the bloom disappears vary quickly and the colorgradually fades. The colors vary from " buckthorn brown " or " oldgold " to " dark olive-buff " or " olive-buff." Many eggs are almost, orquite, immaculate, but more often they are speckled with very smallspots or minute dots of dark brown. The measurements of 58 eggsaverage 42.6 by 32 millimeters; the eggs showing the four extremesmeasure 45.2 by 33.8, 44.2 by 34.1, 40 by 31, and 41 by 30 millimeters.Young.?The period of incubation is about 21 clays, and this dutyis probably performed by the female alone. Only one brood israised in a season. Seton (Thompson, 1890) writes:A partial history of the young in a wild state is briefly as follows: At theage of weeks they are fully feathered and at 2 months fully grown, althoughstill under guidance of the mother at this time. There is usually not more thansix or seven young ones left out of the original average brood of fifteen, whichstatement shows the number of chicks which fall a prey to their natural enemies,while many sets of eggs also are destroyed by the fires which annually devastatethe prairies. As the fall advances they gather more and more into flocks andbecome regular visitors to the stubble fields, and, in consequence, regular
PRAIRIE SHARP-TAILED GROUSE 295
articles of diet with the farmers until the first fall of snow buries their foraginggrounds and drives them en masse to the woods.Doctor Gross says in his notes
:
So far as my observation goes only one bird incubates and cares for theyoung. I did not see the male about the nest at any time. The female is anervous, excitable creature and continually flits her tail when walking aboutor approaching the nest. The tail is usually widespread and displaying thepatches, and the long tail feathers are held up in an upright position, makinga striking picture.Plumages.?Downy young sharptails are decidedly yellowish ; thegeneral color varies from " mustard yellow " above to " straw yellow "below, washed on the crown and back with " ochraceous-tawny "
;
they are spotted on the crown and blotched or streaked on the backwith black; there is a black spot at the base of the culmen and ablack spot on the auriculars.As with all grouse, small chicks show the growth of juvenal plum-age, beginning with the wings; before the young birds are halfgrown they are fully feathered, and even before that they are ableto fly. In full juvenal plumage, in July, the crown and occiput are
" hazel," centrally black ; the feathers of the mantle and wing covertsare brownish black, broadly tipped, barred, or notched with " ochra-ceous-tawny " on the back and with " ochraceous-buff " on the coverts
;
the feathers of the mantle have a broad median white stripe; theunderparts are dull wThite, spotted or barred on the breast and flankswith " sepia " and " cinnamon-buff " ; the four central tail feathers(sometimes nearly all of them) are patterned with "ochraceous-buff " and black, with a broad median white stripe, and they are alldecidedly pointed; the chin and throat are "colonial buff."Within a month or so, during August, the postjuvenal molt takesplace. This is a complete molt, except that the outer primaries oneach wing are retained for a full year; it produces a first winterplumage, which is practically adult in September. Adults may havea partial prenuptial molt, about the head and neck. They have acomplete postnuptial molt, during July, August, and September.The sharp-tailed grouse has several times been known to hybridizewith the prairie chicken.Food.?Dr. Sylvester D. Judd (1905a), by his examination of 43stomachs of sharp-tailed grouse, showed thatanimal matter (insects) formed only 10.19 percent of the food, while vegetablematter (seeds, fruit, and ' Browse ') made 89.81 percent. The insect matter con-sists of bugs, 0.50 percent; grasshoppers, 4.62 percent; beetles, 2.86 percent,and miscellaneous insects, 2.21 percent in a total of 10.19 percent of thefood. Vernon Bailey, of the Biological Survey, found that three birds shot byhim in Idaho August 29 had eaten chiefly insects, including grasshoppers, smallbugs, and small caterpillars. The young of the sharp-tailed grouse, like thoseof other gallinaceous species, are highly insectivorous. A downy chick from 1
296 BULLETIN 162, UNITED STATES NATIONAL MUSEUMto 3 days old, collected on June 27, in Manitoba, by Ernest Thompson Seton,had eaten 95 percent of insects and 5 percent of wild strawberries. The sharp-tailed grouse is fond of grasshopper*. Vernon Bailey shot 3 birds at ElkRiver, Minn., September 17, 1S94, which had eaten, respectively, 7, 23, and 31grasshoppers. The species is a destroyer also of the Rocky Mountain locust.Of 9 birds collected by Professor Aughey from May to October, inclusive, 6had eaten 174 of these pests. The bird eats also a few crickets and, like othergallinaceous game birds, devours the Colorado potato beetle (Leptinotarsadecemlineata )
.
The vegetable food of the sharp-tailed grouse, so far as ascertained in thelaboratory, comprises weed seeds, 7.39 percent ; grain, 20.50 percent ; fruit.27.68 percent; leaves, buds, and flowers, 31.07 percent; and miscellaneousvegetable food, 3.06 percent ; making a total of 89.81 percent. Like many othergame birds, the species feeds on mast (largely acorns), including acorns of thescarlet oak (Quercus coccinea). Corn is eaten, but whf.at is the favoritegrain. It formed 17.21 percent of the food. A thousand kernels of wheatwere sometimes found in one stomach. The sharp-tailed grouse is a greatbrowser. It makes 31.07 percent of its food of leaves, buds, and flowers.Ernest Thompson Seton found it eating the buds of willow and birch. Itfeeds on the leaves ?f cottonwood, alder, blueberry, juniper, and larch; alsoleaves of quillwort (Isoctes), vetch, dandelion, grass, and rush (Juncus).Hearne says that in winter it eats the tops of the dwarf birch and the buds ofpoplars. Flowers form 19.90 percent of its diet, the species leading all otherbirds in this respect. A half pint of the showy, bluish blossoms of the pasqueflower (Pulsatilla hirsutissima) which brightens the western prairie are oftentaken at a meal, and those of the dandelion also are eaten. Inflorescence ofgrasses, alder, willow, maple, and canoe birch are plucked along with leafbuds. Like the prairie hen and the ruffed grouse, the sharp-tailed grouse isfrugivorous, and fruit forms 27.6S percent of its diet. Hips of wild rose aloneform 17.38 percent. Ernest Thompson Seton, who examined hundreds ofstomachs of the sharp-tailed grouse, says that he can not recollect an instancein which they did not contain the stony seeds of the wild rose. Mr. Setonstates that in places in Manitoba, where he has collected during the winter,gravel to pulverize the food is not to be had, and the stony rose seeds act inits stead. Rose hips appear difficult to digest, and, furthermore, are sometimesthickly set with bristles that would irritate the human stomach, but appear tocause no inconvenience to the grouse. The persistent bright-colored hips arereadily seen above the snow, and they are a boon to the birds in wintry north-ern regions where the struggle for existence is bitter. It feeds on blueberriesand cranberries and on the snowberry (Symphoricarpus racemosus) , variousspecies of manzanita, bearberry (Arctostaphylos uva-ursi), buffalo berry (Lep-argyrea argentea), juniper berries, huckleberries, and arbutus berries. It takesalso the partridge berry (Mitchella repens) , a favorite with the ruffed grouse,Like many other species, it eats with relish the fruit of cornel (Cornus stol-unifera) and poison ivy (both Rhus radicans and Rhus diversiloba)
.
Behavior.?The sharp-tailed grouse rises with a loud whir ofwings and flies away with considerable speed, usually in a straightline in the open, with rapid beats of the wings alternated with shortperiods of sailing on down-curved wings. While hunting for nests,where these grouse were common, we kept flushing them from theirroosting and dusting places among the bushes. As they rose, and
PRAIRIE SHARP-TAILED GROUSE 297for some distance afterwards, they uttered a peculiar clucking notesounding like whticker, ivhucker, whucker. They flew rapidly forsome distance and then set their wings and scaled downward intosome good cover or behind some little hill. Seton (Thompson, 1890)says:Their mode of flight is to flap and sail by turns every 40 or 50 yards, andso rapid and strong are they on the wing that I have seen a chicken save itselfby its swiftness from the first swoop of a peregrine Falcon, while another wasseen to escape by flight from a Snowy Owl.Aretas A. Saunders says in his notes:Though usually found on the ground, these birds sometimes perch in trees,either cottonwoods along a river valley or in yellow pines where hills andprairie come together. In southeastern Montana and northwestern SouthDakota they come into the pine hills when the snow gets deep and spend thenights there. In that region I saw them in larger flocks than elsewhere, often50 or 60 birds in a flock. In the pine hills I have seen as many as 30 birdsperched in a single yellow pine and have watched them fly from tree to tree,half a dozen birds leaving at a time, and then another group following tillthe whole flock had reached the new tree.Voice.?Besides the hollow booming sound made by the birds inthe courtship dance and the guttural clucking notes so commonlyheard when they are flushed, " the birds have several cackling notes,and the males a peculiar crowing or low call, that in tone soundssomewhat like the call of the turkey " (Goss, 1891).Enemies.?These grouse are subject to the attacks of the numerousenemies that beset all ground-nesting birds. Their eggs and youngare preyed upon by the smaller and slower predatory mammals andbirds, while the larger beasts and birds of prey attack the adults.While absorbed in their courtship dancing they are easy prey to thecrafty coyote. Laing (1913) tells of a family of coyotes that estab-lished their den, skilfully hidden, on one of the dancing hills; herehe found ample evidence that the wise old coyote and her pups hadlevied regular toll on the unsuspecting grouse. The larger falconsand the goshawk take their share of the grouse, though the latter arefast enough in flight to give even these swift-winged hawks a livelychase. C. L. Broley has sent me the following notes
:
One cold day this fall, while bird observing with Mrs. Broley on the banksof the Red River, north of Selkirk, we saw three sharp-tailed grouse fly outof the woods on the opposite side of the river as if they had urgent businesselsewhere, and a moment later a fourth one burst out with two goshawks inhot pursuit. The grouse flew directly out over the water, where one of thehawks, reluctant to follow in the open, dropped out of the chase. The grousefled upstream, keeping some 50 feet above the water and an equal distancefrom each shore, followed, some 30 feet behind, by the other hawk, which to oursurprise kept below the altitude of the thoroughly frightened chicken, which
298 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMwas taking advantage of a strong -north wind and making wonderful time,using his wings almost continually and sailing very little.Fortunately we had a clear view up the river for nearly 2 miles and whilethe birds were covering this distance their respective positions remained un-changed, but when the bend of the river was reached the grouse changed itscourse, going over the land. The goshawk immediately rose from below to wellabove its quarry, with the full intention, we thought, of making its swoop, butunfortunately intervening trees close to us hid the rest of the incident.Norman Cricldle (1930), in an interesting study of the fluctuationsin numbers of grouse in Manitoba, from 1895 to 1929, has shownon a graph " the remarkable regularity with which sharp-tailedgrouse fluctuate in company with grasshoppers." His theory is thatyoung grouse are fed to a large extent on grasshoppers and thatduring periods of abundance of these insects the birds rear largebroods. He says
:
Referring again to our graph it will be observed that all the high points ofSharp-tailed Grouse abundance were preceded and accompanied by grasshopperoutbreaks. Of these outbreaks the one of 1900-1903 was much the worst thatwe ever experienced in the Aweme district, and it was not finally subdued until1905. The peak of grouse abundance was also attained at this time, and we haveno records of Sharp-tailed Grouse ever being in greater numbers.Game.?My experience with sharp-tailed grouse was confined tothe breeding season, when they are very tame, flush near at hand,fly rather straight and evenly, and give one a good, open, easy shot.We considered them easy marks and had no difficulty in shootingall we needed for specimens. But in fall, after they have been shotat, they are evidently more gamy and seem to be popular as gamebirds. Sandys (1904) writes:The sport afforded by this grouse is of a very high order. At the openingof the season it lies well to the dog, and springs with the usual whirr of wings,at the same time uttering a vigorous clucking, which is repeated again andagain as the birds speed away, alternately flapping and sailing. When drivento brush, they very frequently behave not unlike quail, flushing close at hand,and offering the prettiest of single chances. The flesh is excellent, light-coloredin young birds, and darkening with age, but always worthy of a place on thehoard. Not seldom, as one nears the pointing dog, he will see the birdssquatted in the grass, and perhaps, have one after another turn and run afew yards before taking wing. When thus seen they are very handsome, thecrest is raised, and the white hinder feathers show like the flag of a deer, orthe scut of a cottontail rabbit. Almost invariably the flush is straggling, giv-ing a quick man a fine opportunity for scoring again and again. At the properseason, i. e., just before the broods begin to pack and become wary, this birdaffords sport to be long remembered. I have enjoyed it to the full, and knowof nothing better for a business-harassed man than a day on the sunny openwith the sharptails behaving well. Like all prairie-grouse, this bird, risingclose, is an easy mark for whoever has learned not to be hurried by the soundof wings. A good twelve-gauge, properly held, should stop its buzzing andclucking fully three-fourths of all reasonable chances.
PRAIRIE SHARP-TAILED GROUSE 299Winter.?Seton (Thompson, 1890) writes:During the summer the habits of the chickens are eminently terrestrial ; theylive, feed, and sleep almost exclusively on the ground ; but the first snowmakes a radical cL^inge. They now act more like a properly adapted perch-ing bird, for they spend a large part of their time in the highest trees, flyingfrom one to another and perching, browsing, or walking about among thebranches with perfect ease, and evidently at this time preferring an arborealto a terrestrial life. When thus aloft they are not at all possessed of that feel-ing of security which makes the similarly situated Ruffed Grouse so easy aprey to the pothunter. On the contrary, their perfect grasp of the situationusually renders them shy and induces them to fly long ere yet the sportsmanhas come near enough to be dangerous. Like most of the members of itsfamily, the Prairie Chicken spends the winter nights in the snow, which isalways soft and penetrable in the woods although out on the plains it is beatenby the wind into drifts of ice-like hardness. As the evening closes in the birdsfly down from the trees and either dive headlong into a drift or run about alittle and select a place before going under. The bed is generally about 6inches from the surface and a foot long from the entrance. Each individualprepares his own place, so that a flock of a dozen chickens may be scatteredover a space of 50 yards square. By the morning each bird's breath hasformed a solid wall of ice in front of it, so that it invariably goes out atone side. The great disadvantage of the snow bed is, that when there thebirds are more likely to become the prey of foxes and other predaceous animals,whose sagacious nostrils betray the very spots beneath which the unsuspectingbird is soundly slumbering. I am inclined to think this is the only chancea fox has of securing one of the old birds, so wary are they at all other times.Laing (1913) says:The question of winter food is never a big problem with these grouse.There is always an abundance of hawthorn hips, rose fruit, snowberries, orother winter-cured fruits; in addition to these, edible buds of many kinds,are in abundance. Best of all, they seem to relish the sweetish, frost-rip-ened berries of the dwarfish snowberry that peep above the snow just farenough to invite picking. Of the tree-buds, poplar, willow, and dwarf birchare winter staples, and these are consumed in great quantities. The questof wheat in winter frequently leads these grouse to come right into thetowns. They first took this bold step after finding wheat dropped along theroads and railway. As the clue led in the direction of the big red elevators,they followed it in at first, but soon needed no grain trail by way of invi-tation. They were quick to learn that they were not molested in the winter,and indeed received a ready welcome. The daily itinerary of this flock inmid-winter is about as follows : With the first peep of dawn they leave theirsnow beds and mount to the tops of the willows and poplars, close at hand.At sunrise or a little before it, they whizz off into town and scatter aroundthe various feeding grounds mentioned above. About ten o'clock they usuallytake a run out along the railway track, evidently to get a supply of gravel;then they return to spend the warm part of the day in the scrubby sand-hills. Here they sit about in the sun, and pick a few buds; or if the dayis very cold and the snow light and deep, they burrow for their noon-day nap.At three o'clock they return to their feeding-place of the morning, and thenshortly before sundown go back to the scrub to make their beds for the
300 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
night. Here while the cruel wind sweeps across the plains and the ther-mometer ofttimes registers in the minus forties, they remain cuddled snuglyaway from the bitter night-world.CENTROCERCUS UROPHASIANUS (Bonaparte)SAGE HENHABITSThe recent American Ornithologists' Union check lists, both oldand new, call this the sage hen, but I prefer to call it a grouse, asit really is and as it was called in earlier editions. I see no reasonfor calling it a hen, except that the cackling notes of the femaleremind one of that familiar domestic fowl. It is a true grouse anda grand one, by far the largest of our American species and thelargest in the world except the European capercaillie, which farexceeds it in size. A fully grown sage-grouse cock is said to weighas much as 8 pounds, but the hen will not weigh more than 5 pounds,probably both usually weigh much less.It was discovered by Lewris and Clark about the headwaters ofthe Missouri River and on the plains of the Columbia ; they namedit " cock of the plains " and gave the first account of it. The tech-nical description of it and the scientific name, urophasianus, weresupplied by Bonaparte in 1827.The range of the sage grouse is limited to the arid plains of theNorthwestern States and the southwestern Provinces, where the sage-brush (Artemisia triclentata and other species) grows; hence it iswell named sage grouse or cock of the plains. Its range stops wherethe sagebrush is replaced by greasewood in the more southern deserts.Like the prong-horned antelope, another child of the arid plains, ithas disappeared from much of its former range, as the countrybecame more thickly settled and these large birds were easily shot.It has been said that the sage was made for this grouse and thisgrouse for the sage, where it is thoroughly at home and where itscolors match its surroundings so well that it is nearly invisible whilesquatting among the lights and shades of the desert vegetation. Itseldom wanders far from the sagebrush, but may be found occa-sionally in the shade of the narrow line of trees that marks thecourse of some small stream. Dwight W. Huntington (1897) de-scribes its haunts very well, as follows
:
I found the Sage Grouse most abundant in the vicinity of Fort Bridger andsouth to the Uintah Mountains. Here the tufted fields of the gray-green sagesweep up to the sides and walls of the adjacent " bad lands," or buttes, devoidof vegetation but beautiful in color and fantastic in form. The buttes arestrangely fashioned by erosion, and are full of the fossil remains of animalsand fishes. Numerous domes, spires, and pinnacles surmount the buttes and theconglomerate layers running about them have been compared to Egyptian carv-
SAGE HEN 301ing. Towards the southwest are the blue Uintah Mountains, with snow flashingon their crests all summer, and towards the east the vast plain of sage extendsas far as the eye can reach, blending at the horizon into an azure sky. Thetrout streams which issue from the mountain side become the small rivers ofthe plains, flowing at long intervals and nourishing a narrow line of verdure ora yellow screen of cottonwood, which marks their course. It is along suchstreams that the sage grouse hunter must pitch his camp.Courtship.?Much has been written about the courtship of thesage grouse, which is the most spectacular performance indulged inby any of the grouse. It has been variously described by differentobservers. Frank Bond (1900) was one of the first to describe andillustrate this with a drawing ; he writes
:
During the months of April and May the Sage Cocks are usually found in smallflocks of a half dozen or more, stalking about with tails erect and spread afterthe manner of the strutting turkey cock, but I have never seen the Grouse drag-ging their wings upon the ground, turkey fashion, and in the manner describedby Dr. Newberry in the quotation from this author found on page 406 ofDr. Coues's " Birds of the Northwest," nor have I ever found a wing of aSage Cock in this or any other season, which exhibited the slightest wearingaway of the primaries. Instead of dragging its wings upon the ground theSage Cock will enormously inflate the air sacks of the neck until the whole neckand breast is balloon-like in appearance, then stooping forward, almost the entireweight of the body is thrown upon the distended portion and the bird slidesalong on the bare ground or short grass for some distance, the performancebeing concluded by the expulsion of the air from the sacks with a variety ofchuckling, cackling or rumbling sounds. This performance is continued prob-ably daily, during the pairing and nesting season, and of course the feathers areworn away by the constant friction.William L. Finley's account of it differs somewhat. He has sentme the following notes on the subject:On May 13 we rose at 3.15 and were in the blind a little after 4, still verydark. The birds at this time were already strutting. We could hear themand occasionally see a flash of white from the breasts. The birds were activebetween 4.30 and 6.30 a. m. and had left the strutting ground by 8 a. m.When the sage cock starts to strut, his tail spreads and the long pointed tailfeathers radiate out in a half arc. The air sacs are filled and extend nearlyto the ground, hiding the black breast feathers. This is the first movement.Then the bird takes one or two steps forward and throws up the pouch, appar-ently by drawing back the head and neck. The next movement is a repetitionof throwing the air sacs up and down and getting under headway for the lasttoss of the pouch, which is brought down with a jerk, as one would crack awhip, making a " plop " that on a quiet morning we easily heard for a distanceof 200 or 300 yards. The whole movement gives one the idea that the birdinflates the air sacs and then, by the rigid position of the body and throwingthe head and neck back, gives these air sacs a very vigorous shaking. In themovement when the pouch spreads, the bare yellow skin on the lower partof the pouch or chest shows clearly. As the pouch is thrown up and down,the wings are held rigid, the tips of the wing feathers sometimes touching theground. The white feathers that cover the chest are exceedingly stiff; thesegrate against the wing feathers, giving out a wheezy sound that at first I
302 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthought came from the inhaling and exhaling of air. I soon discovered thatthis rasping noise was made by the stiff feathers rubbing together. Thisrubbing of the breast feathers against the rigid wing feathers seems to accountfor the very worn appearance of the breast later in the season. If therewere any gurgling or chuckling noises I failed to catch them. As the struttingends, the air sacs are deflated and each time the bird goes through the motionas if gulping or swallowing something.There were 56 cocks scattered around in an area of 4 or 5 acres, each birdhaving a space for himself. Occasionally when one bird came too near anotherit resulted in a fight. Once I saw two fight with lowered heads. Occasionallythey would jump in the air, striking very much as an ordinary rooster strikes.Two or three times I saw a female feeding near by, where the males werestrutting. One passed by several cocks, but they paid no attention to her, orshe to them. This gave me the impression that the strutting was not so mucha courting performance or even a nuptial dance; it seemed to be a gatheringplace where the males came together and " showed off " among themselves.E. S. Cameron (1907) observed:By ruffling up all their feathers, spreading their tails, and dragging theirwings along the ground they looked much larger than they really we're, whilethey produced a rattling sound with their quills after the manner of turkey-cocks and peafowl.Some additional information is given by L. E. Burnett (1905),who writes:I have heard them drum as early as December. This performance is most,often observed where hundreds of males and females have congregated to-gether, a custom which they have in the fall of the year. By February, themales are all drumming, but this is not continued during bad weather whichcloses the session until fair weather returns. By the latter part of the monththe males are in full dress. Their protracted meetings last until the firstdays of May. After the violets and buttercups have come and the song of thesage thrush begins, their drumming is heard but occasionally. When drum-ming they stand very erect, holding the wings away from the sides and nearlyperpendicularly, while the large loose skin of the neck is worked up, and thehead drawn in and out until the white feathers are brought to the chin.At the same time the galls are filled with air until the birds look as if theywere carrying snowballs on their shoulders. Then the skin which lies be-tween the galls is drawn in with a sucking movement, thus bringing the gallstogether or nearly so. With this action the air is expelled from the throatproducing the noise, which is hard to mimic and which resembles that ofan old pump just within hearing distance. After the bird has accomplishedthis feat he walks away a few paces either in a straight line or a circle, withwings down, hanging loosely, but not grating on the ground. At times theydo drag the wings as they strut along with tail spread and erect, though notso perpendicular as that of a turkey. Again they will dance about with allthe pomp of a male pigeon.Maj. Allan Brooks (1930) says that "the feathers of the breastand neck of the male sage grouse are specialized feathers only " andare not worn away by rubbing on the ground during the display.
SAGE HEN 303Nesting.?Major Bendire (1892) says:The nest is always placed on the ground, in a slight depression, usuallyunder the shelter of a small sage bush. I have found several, however, somelittle distance from sage brush flats, alongside and sheltered by a bunch oftall rye grass (Elymus condensates?) , near the borders of small creeks. Thenest is usually very poorly lined, and in fact the eggs frequently lay on thebare ground without any lining whatever, and are often found in quite exposedsituations. I found such a one on May 11, 1875. My notes read as follows:
" I stumbled accidentally on this nest. It was placed within a yard of amuch-used Indian trail, in a very exposed position, so much so that I sawthe eggs while still 5 yards off. There really was no nest, simply a meredepression scratched out by the bird on the south side of a very small sagebush, which afforded no concealment or protection from rain whatever. Thebush itself was not over a foot and a half high, growing on a rocky plateauabout 3 miles east of Camp Harney. A few feathers were scattered amongthe eggs which laid on the bare ground, and were separated from each otherby bits of grass and dry leaves of the sage. One of the eggs was nearlycovered with dirt and almost buried out of sight. The set contained eighteggs, and these were nearly hatched. They were cold when found, and thenest had evidently been abandoned for some days.Illustrating the concealing colorations of the close-sitting bird,Bendire quotes Capt. William L. Carpenter as follows
:
I found a nest at Fort Bridger, Wyoming, where this species is numerous, June1, with nine fresh eggs. I was standing alongside a sage bush watching butter-flies ; several times looking down carelessly without seeing any thing unusual,when happening again to glance at the foot of the bush, in the very placebefore observed, I saw the winking of an eye. Looking more intently agrayish mass was discerned blending perfectly with the color of the bush,which outlined itself into the form of a Sage Hen not 2 feet from my foot.She certainly would have been overlooked had not the movement of her eye-lids attracted my attention. I stood there fully five minutes admiring thebeautiful bird, which could have been caught in my butterfly net, then walkedback and forth, and finally passed around the bush to observe it from behind.Not until then did it become frightened and fly away with a loud cackling.The nest was a depression at the foot of a sage bush, lined with dead grassand sage leaves. The spot was marked and visited several times, alwayspassing within a few feet without alarming the bird.D. E. Brown tells me of a nest found by a sheep herder. Thebird did not flush from the nest until the sheep were all aroundher; she then flushed with a great noise, scattering the sheep in alldirections. This habit may often prove very useful in preventingcattle from trampling on the eggs.Eggs.?The number of eggs laid by the sage grouse usually variesfrom 7 to 9, in some localities from 10 to 13 ; as many as 15, or even17, have been found in a nest. Bendire (1892) found but one set of10, and found more sets of 8 than any other number. They varyin shape from ovate to elongate ovate, and the shell is smooth withlittle or no gloss. The ground colors vary from pale " ecru-olive "or " deep olive-buff " to " yellowish glaucous," " olive-buff," or " green-
304 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMish white." They are generally quite evenly marked with smallspots and fine dots of dark brown, " bister," or " brownish olive "
;
in very light-colored eggs the spots are in very pale shades of brownor olive. The markings are very easily washed off when the eggsare fresh. The eggs in a set are seldom, if ever, uniform in type;there are usually two or more conspicuously different types in eachset. The measurements of 110 eggs average 55 by 38 millimeters;the eggs showing the four extremes measure 59.5 by 39.5, 58.5 by 40.5,51 by 37, and 56.5 by 35.5 millimeters.Young.?Bendire (1892) gives the period of incubation as 22 days.This duty is performed by the female alone, as the polygamousmales desert the females as soon as the eggs are laid and associate inflocks by themselves. Consequently the full care of the young restson the devoted mother. Mrs. Irene G. Wheelock (1904) writes:Like all grouse nestlings, they run about as soon as the down is dry, whichis about fifteen minutes after the shell breaks. They pick up food at herscratching all day, and at night they nestle on the ground under her wings, onlya row of little heads being visible. As soon as their own feathers are developed,they sleep every night in a circle about her, each one with head pointed tothe outside as before, and always on the ground ; for the Sage-Grouse nevertrees. It is not difficult to come upon a brood sleeping this way on a moon-light night ; but the only satisfaction will be to hear the sharp alarm of themother, a whirr as she runs by you, and a knowledge that though the young arehiding on the dust at your feet, you could not find them were your eyes ten-fold sharper. I have groped carefully on hands and knees among them, andactually touched one before I saw it at all. For the desert hides its secretswell, and the little grouse have learned to trust to it for safety.Grinnell, Bryant, and Storer (1918) say:After the young birds have learned to fly, they descend from the uplandsdown along the larger canons, often invading the meadow lands, where smalltender weeds are added to their diet. At such places the young birds maygather into large flocks. When approached they crane their necks and makea weak attempt at cackling. When closely pressed they run rather than fly.By the last of August or early September the young birds are joined by the oldmale birds, which come off the higher slopes and ridges where they have stayedduring the summer and large flocks become the rule.Plumages.?The sage-grouse chick is well colored to escape de-tection when crouching on the ground in the gray shadows of thedesert. The crown, back, and rump are mottled and marbled withblack, dull browns, pale buff, and dull white; the sides of the headand neck are boldly spotted and striped with black; there are twolarge spots of " sayal brown " bordered with black, on the fore neckor chest; underparts grayish white, suffused with buff on the chest.The juvenal plumage comes in first on the wings, while the chick isvery small, then on the scapulars, back, tail, sides of the breast, andflanks, lastly on the rump, head, neck, and belly. The juvenal plum-age is much like that of the adult female ; but the breast is more
SAGE HEN 305buffy and more spotted than barred ; the feathers of the black breastpatch are tipped with white; and the feathers of the mantle areconspicuously marked with a broad shaft streak of white. Youngmales seem to be much darker than young females.A nearly complete postjuvenal molt, including all but the outertwo primaries on each wing, produces the first winter plumage, whichis practically adult and in which the sexes are fully differentiated.There may be a partial prenuptial molt, but I have seen no evidenceof it in either young or old birds. Adults have a complete post-nuptial molt, mainly in August.Food.?Dr. Sylvester D. Judd (1905a) says:The feeding habits of the sage grouse are peculiar, and its organs of digestionare unlike those of other grouse. The stomach is not differentiated into apowerful grinding gizzard, but is a thin, weak, membranous bag, resemblingthe stomach of a raptorial bird. Such an organ is evidently designed for thedigestion of soft food, and we find that the bulk of the sage grouse's diet con-sists of leaves and tender shoots. A stomach collected September 7, 1890, inIdaho, by Dr. C. Hart Merriam, contained leaves of sage and other plants,seeds, and a ladybird beetle (Coccinellidae). Four birds shot in Wyomingduring May and September by Vernon Bailey had gorged themselves with theleaves of sagebrush (Artemisia tridentata). This and other sages, includingA. cana and A. frigida, furnish the bulk of the food of the sage grouse. Otherfood is taken, but it is comparatively insignificant. B. H. Dutcher, formerly ofthe Biological Survey, examined a stomach which, besides sagebrush leaves,contained seeds, flowers, buds of Rhus trilobata, and ants and grasshoppers.Three birds collected by Vernon Bailey on September 5, in Wyoming, had variedtheir sagebrush fare with ladybird beetles, ground beetles (Carabidae), flylarvae, ants, moths, grasshoppers (Melanvplus sp.) and the leaves of astersand yarrow. Of two birds killed in May, one had fed wholly on the leavesof sagebrush (Artemisia tridentata), while the other in addition had takeninsect galls from sagebrush and the flowers and flower buds of a phlox (Phloxdouglasii), together with some undetermined seed capsules, pieces of moss, andseveral ants. A third bird, killed in July, had eaten a few plant stems andnumerous grasshoppers.During the winter the sage grouse feeds almost entirely on theleaves of the sage, but "in summer," according to Bendire (1892)," its principal food (in Wyoming and Colorado) is the leaves, blos-soms, and pods of the different species of plants belonging to thegenus Astragalus, and Vicia, commonly called wild pease, which arealways eagerly sought for and consumed in great quantities."Eobert S. Williams reported to Bendire from Montana that hescared up a flock among tall grass in a mountain meadow; one ofthese birds had its crop full of the blossoms of a species of golden-rod. Bendire also quotes George H. Wyman as stating that a sagegrouse will go a long way for food in a wheat field; some that heexamined had traveled at least 8 miles to till their crops with ripewheat.
306 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMBehavior.?Dr. D. G. Elliot (1897) says:It is not always easy to flush these birds, as they will run long distances beforetaking wing, and skulk and hide at every opportunity. But when forced torise, they flush with a great fluttering of the wings and utter a loud kek-kek-kek,which kind of cackle is kept up for quite a considerable time. They seem tohave difficulty in getting well on the wing, and rise heavily, wabbling from sideto side as if trying to gain an equilibrium, but once started they go far andfast enough, with intermittent quick beats of the pinions and easy sailing onmotionless wings.Mr. Finley refers in his notes to their morning flights for water,as follows
:
September 22, Horsfall and I were up at 4.40 a. m., left the cabin at 5.40, justas it was getting light. As we walked down the long draw, through whichWarner Creek winds back and forth, we saw sage grouse coming in to water.They came from the rimrocks and higher plateaus, perhaps from several milesaway, as some of them sailed down from high over the rimrock, showing thatthey had come from a second high rimrock about a mile back. They came insingles, in small flocks of from 8 to 10, occasionally larger flocks numbering30 or 40. They lit out in the open at 100 or 200 yards back from the creek, andthen walked down to water. In a short distance I counted 150 birds. Fromthe cabin down to a small reservoir, where the water was backed up coveringseveral acres of ground, there must have been 1,500 to 2,000 birds. Aroundthe reservoir site there was also a larger number. It was the same alongthe creek above the cabin, where it winds through meadow and sagebrush. Thegrouse do not frequent the water at all during the middle of the day, andwe found very few toward evening. They seem to come in before daybreak,and a little after many have departed for the high plateau, by 6 o'clockor before sunrise. By 8 o'clock all had departed.Bendire (1892) quotes Dr. George Bird Grinnell as stating:On a very few occasions I have seen the Sage Grouse standing on thebranches of a sage bush, sometimes 2 or 3 feet from the ground, but I imaginethat this is quite an unusual position for the bird. This species, commonly, Ithink, goes to water twice a day, flying down to the springs and creek bottomsto drink in the evening, then feeding away a short distance, but roostingnear at hand. In the morning they drink again and spend the middle of theday on the upland. The young birds, when feeding together, constantly callto one another with a low peeping cry, which is audible only for a short distance.This habit I have noticed in several other species of our Grouse, notably inthe Dusky Grouse and the Sharp-tail.Coues (1874) quotes from Doctor Newberry's account, as follows:A very fine male which I killed there was passed by nearly the whole party,within thirty feet, in open ground. I noticed him as soon, perhaps, as he saw us,and waited to watch his movements. As the train approached he sank down onthe ground, depressing his head, and lying as motionless as a stick or root,which he greatly resembled. After the party had passed I moved toward him,when he depressed his head till it rested on the ground, and evidently madehimself as small as possible. He did not move till I had approached within15 feet of him, when he arose and I shot him.
SAGE HEN 307And from Mr. Holden, Coues quotes
:
They roost in circles on the ground. I have seen a patch of ground fifteenfeet in diameter completely covered with their excrement. I think they resortto the same place many nights in succession, unless disturbed.Voice.?The vocal efforts of the sage grouse seem to be limitedto a deep guttural clucking note, kuk, kuk, kuk, slowly repeated asthe bird flushes, a rapidly repeated scolding note, tuk-a-tuk, and acackling note of the female, like the cackle of a domestic hen. Thechicks call to one another with faint peeping notes.Enemies.?Sage grouse have been found to be infested with tape-worms and probably they are also infected with some of the otherparasites and diseases to which other grouse are subject. The eggsand young are preyed upon by various predatory animals and birds,mainly crows and magpies, but their worst enemy is man. Naturalenemies of all wild creatures have merely checked their increase, butwhen man comes on the scene it means extermination. So it is withthis fine large grouse, an easy mark for the gun. It has been extir-pated from much of its former range and is disappearing verysteadily in many other places.Sandys (1904) relates the following incident:One day I was watching an old male which had taken up a position upon analmost bare knoll. It was before the open season, a very idle period on theplains ; so, partly to pass away time, and partly in the hope of discoveringsomething, the field-glass was brought into play. Before the bird had beenthoroughly scrutinized, some falcon, which looked like a male peregrine, shotinto the field of vision, and made a vicious stoop at the huge quarry. Whetheror no the grouse had been watching the hawk is impossible to say, but in anyevent he was ready. As the hawk was almost upon him, up went the long tail,down went the head, and the wings were a trifle raised. Most readers, probably,have seen a man hump his back and get his shoulders about his ears when heexpected to be struck from behind by a snow-ball. The action and attitude ofthe grouse were comically suggestive of that very thing. The hawk appearedto be only fooling, for certainly it made no determined strike, but presentlyrose and curved away. An instant later the grouse took wing.Game.?Sandys says of the game qualities of this grouse
:
As an object of the sportsman's pursuit, the sage-grouse is greatly inferior tomost of its relatives. The young, the only ones worth shooting, are great run-ners, and only take wing when compelled to, and once in the air their size isagainst them, although they fly fairly fast. Another objectionable feature istheir ability to carry off shot, which sometimes borders on the marvelous. Alight gun, deadly on other grouse, will hardly serve for these big fellows, theuse of it surely meaning a lot of wounded birds. The coveys usually are small,as the young have many enemies, among which the chief are fierce storms, wet,wolves, foxes, and rapacious birds, while man plays no unimportant part in tnework of destruction.Dwight W. Huntington (1903), however, speaks very highly of itas a game bird, and says of its value as a table bird "that these
308 BULLETIN 162, UNITED STATES NATIONAL MUSEUMbirds, like others, often receive a flavor from their food, and whenthe wild sage is their exclusive diet they have a more or less bittertaste. When, however, the birds are young and have been feeding ongrasshoppers, their flesh is as good as that of the sharp-tails or prai-rie-grouse."As to his method of hunting sage grouse, Huntington writes
:
My shooting of these birds was mostly done from the saddle while on themarch. When we flushed a covey of birds I took a shot at them, and markingthose that flew away to the particular bush where they settled, rode at onceto the spot and sometimes dismounted to shoot at the scattered birds. Uponseveral occasions I went out with a friend especially to shoot them, ridinghere and there (we had no dog) until the horse flushed a covey, and followingthem so long as we could make them take wing. Birds often escaped byhiding in the sage and refusing to fly. The most likely places seemed to bedepressions where the water evidently flowed in wet seasons and little knollsadjacent, but we stumbled upon the birds almost anywhere in the sage, andoften made very good bags. It was next to impossible to miss one, since theshots were always in the open and the marks large. The birds required hardhitting, however, to bring them down, and I would not advise the use ofshot smaller than number 5 or 6. A wounded bird is difficult to recover withouta dog where the sage grows thickly, and I always tried to kill the birds outright.The side shots, or those at quartering birds, are more likely to be fatal thanthose at birds going straight away, since the shot then penetrates the lighterfeathers beneath the wings.Burnett (1905) says:The counties of Albany, Converse, Natrona, and Carbon are the placeswhere grouse are most abundant in Wyoming. A single hunter has beenknown to kill a hundred birds in a day without a dog. The best huntingis found over lands adjacent to springs, down green draws and the bottomsalong streams, and the best time to find coveys is in the morning or eveningwhen the birds are feeding. After feeding they hide either on the feedingground or at some distance from it where the sage is large enough to screenthem from enemies and the rays of the sun.Its large size, the ease with which it can be killed, and the accessi-bility of its haunts combine to make this grouse a popular game birdduring summer and fall when its flesh is most palatable. Conse-quently it is disappearing very fast, notably in California, Oregon,and Washington, where the extension of good roads and the increasein automobiles have made the sagebrush plains more accessible. Tosave this fine bird from extinction, as civilization spreads, the openseason for shooting it must be shortened and the bag limits reduced.Even then, it probably can not be saved except on protected reserva-tions.Fall.?These grouse are usually resident throughout the year wher-ever they are found; but on some of the elevated plateaus, in themore northern portions of their range, the sagebrush, on which theyfeed in winter, becomes buried under the snow; they are thereforeobliged to migrate in search of a food supply.
SAGE HEN" 309In the days of their abundance they used to gather in immensepacks in fall. Bendire (1892) quotes the following from notes sentto him by Dr. George Bird Grinnell
:
In western Wyoming the Sage Grouse packs in September and October. InOctober, 1886, when camped just below a high bluff on the border of Bates Hole,in Wyoming, I saw great numbers of these birds, just after sunrise, flying overmy camp to the little spring which oozed out of the bluff 200 yards away.Looking up from the tent at the edge of the bluff above us, we could see pro-jecting over it the heads of hundreds of the birds, and, as those standing theretook flight, others stepped forward to occupy their places. The number ofGrouse which flew over the camp reminded me of the oldtime flights of Passen-ger Pigeons that I used to see when I was a boy. Before long the narrow valleywhere the water was, was a moving mass of gray. I have no means whateverof estimating the number of birds which I saw, but there must have been thou-sands of them.Winter.?Unless the snow is too deep the sage grouse seek shelterfrom the winter storms and blizzards in the denser clumps of sage-brush on their favorite plains or find protection in the brushy valleysof the streams, in coulees, or in sheltered hollows. Sandys (1904)writes
:
As winter tightens its grip upon the sage lands, the birds of many broodsunite into packs of from fifty to one hundred and odd. The flush of one ofthese large packs is something to be remembered, for great is the tumult ofwings, and piercing the cackling, as the heavy fowl beat the air in frantic effortsto get squared away upon their chosen course. At this season the only wayto get any sport out of them is by using the rifle.DISTRIBUTIONRange.?Western United States and casually in the interior ofsouthwestern Canada.The range of the sage grouse has been greatly restricted throughthe development of the West and through grazing activities, partic-ularly of sheep, which do much to extirpate the birds over wide areas.The full range apparently extended north to (casually) the interiorof southern British Columbia (Osoyoos Lake) ; southern Saskatche-wan (casually Skull Creek, Val Marie, and casually Pinto Creek)
;
North Dakota (Marmarth, Deep Creek, and formerly Fort Bert-hold) ; and formerly northeastern South Dakota (Grand RiverAgency and Fort Sisseton). East to South Dakota (formerly FortSisseton, Indian Creek, formerly Eapid City, and formerly SageCreek) ; northwestern Nebraska (Antelope Creek) ; southeasternWyoming (Marshall, Arlington, and Cheyenne) ; Colorado (Walden,Kremmling, Dillon, Lone Cone, and Dolores) ; and formerly north-ern New Mexico (Tres Piedras). South to formerly New Mexico(Tres Piedras and Tierra Amarilla) ; southern Utah (Grass Valley74564?32 21
310 BULLETIN 162, UNITED STATES NATIONAL MUSEUMand Hamblin) ; Nevada (Belmont and Queen) ; and eastern Califor-nia (Big Pine and the headwaters of the Owens River). West toeastern California (the headwaters of the Owens River, Long Val-ley, Ravendale, Madeline Plains, Eagleville, and Tule Lake) ; Ore-gon (Klamath Falls, Silver Lake, Fort Rock, Silvies River, TurtleCove, and Haines) ; Washington (Rattlesnake Mountains, Yakima,and Ellensburg) ; and (casually) the interior of southern BritishColumbia (Osoyoos Lake).Egg dates.?Washington and Oregon : 16 records, March 11 to May28; 8 records, April 11 to May 6. Idaho, Montana, and Wyoming:25 records, April 25 to June 15 ; 13 records, May 16 to 29. Coloradoand Utah : 11 records, May 10 to June 3 ; 6 records, May 19 to 28.Family PHASIANIDAE, Pheasants, PeacocksPHASIANUS COLCHICUS TORQUATUS GmelinRING-NECKED PHEASANTHABITSContributed by Charles Wendell TownsendAlthough some of the earlier English settlers in North Americacalled the ruffed grouse the pheasant, a name that is still retained inthe southern parts of its range, no true pheasants are native, norwere they successfully introduced into America until 1881, whenJudge O. N. Denny, then American consul general at Shanghai,China, after a previous unsuccessful attempt, sent 30 ring-neckedpheasants to Oregon. Of these 26 survived and were liberated inthe Willamette Valley. Two years later more were sent (Shaw,1908). Although several early attempts at introduction were made,the first successful introduction of pheasants into the East was in1887 by Rutherford Stuyvesant, who brought over a number of birdsfrom England and liberated them on his estate at Allamuchy, N. J.In the nineties, pheasants were brought from England and liberatedin various places in Massachusetts and elsewhere.The bird proved to be remarkably hardy and prolific and spreadrapidly, partly by natural increase and partly by artificial breedingin private and State farms, and by the shipment of eggs and birds tonew sections of the country. The bird thrives in the North, butsouth of Baltimore and Washington, according to Dr. J. C. Phillips(1928), although there have been many attempts at introduction," the stock does not hold out long if thrown on its own resources."W. L. McAtee (1929) says of this bird that itnow has an almost continuous distribution over the Northern States from coastto coast. It has proved hardy as to climatic conditions, wary as to enemies,and without doubt is more numerous than any native game bird in the area
KING-NECKED PHEASANT 311
occupied. The success of the introduction of pheasants in the NorthwesternStates is well known, but how amazingly the birds have thrived in certainother sections is not generally appreciated. In South Dakota, according to theDirector of the State Department of Game and Fish, pheasants increasedsteadily from the first, a fact justifying almost steady lengthening of theopen season and increase in the daily bag limit. The total bag in 1926 wasestimated at a million birds, and in 1927 from one and a half to two millions,a record that has scarcely been approached in all our history by a singlespecies of game bird in a single State.William L. Finley contributes interesting details on the status ofthe bird in Oregon, which, somewhat condensed, are as follows:For more than 20 years the success of the ring-necked pheasant seemedto be complete in Oregon. The numbers of the birds increased and theyspread to all parts of the Willamette Valley and over into other valleys,although thousands of the birds were killed each hunting season. Then therecame a period when the bird seemed to be just holding its own, and for thepast 20 years the population has been on the decrease. Many reasons havebeen given for this decrease, as, for example, too much shooting or an increasein enemies.One reason for the decrease in the number of pheasants in the WillametteValley may be a change in the life of the bird itself. It survives best where itnests, roosts, and lives out in the open fields, where its watchfulness is alwaysa check upon its enemies. In the Willamette Valley during the past 20 yearsthe pheasant has come to be a bird of the woods, often lays its eggs in otherbirds' nests, and is not so good a mother to its chicks as formerly, so that fewersurvive. It was the common thing late in summer or early in fall to flush acovey of 8 to 15 birds, while nowadays it is very rare to see more than 4 to 6young pheasants in a covey.Dr. J. C. Phillips (1928) says:The extraordinary vitality of the first birds set out by the writer at NorthBeverly, Mass., in 1897 and 1898 was a most interesting feature. The broodswere at first large and the species did not appear to meet any natural checksto its spread for a number of years. This initial " vigor," however, seems tohave been lost here as well as in other places where the pheasant has beenplanted for 25 to 30 years.The following information regarding the pheasant from W. H.Hudson (1902) is of interest in this connection:In Britain, where it has been permitted to run free in the woods for the lastsixteen to seventeen centuries, it is still scarcely able to maintain its existencewithout the strictest protection and a great deal of attention on the part of man.It is known that when the birds are left to shift for themselves they soon de-crease in numbers, and eventually die out, except in a few rare cases where theconditions are extremely favorable. How heavy the cost is of keeping pheas-ants in numbers sufficient for the purpose of sport is well known to all thosewho have preserves.The many thousands killed annually may well be the cause of thefailure of these birds to hold their own without artificial aid. Theextent to which this aid was given in Massachusetts in the year 1929
312 BULLETIN 162, UNITED STATES NATIONAL MUSEUMis shown by William C. Adams, director of the Division of Fisher-ies and Game, who writes : " From the four game farms we haveshipped out 13,200 pheasant eggs, 1,554 adult pheasants, and 12,728pheasants at least three months old." Incubators and brooders areused, and the eggs and birds are distributed to individuals and localgame clubs throughout the State for liberation of the birds in thewild.The pheasant is generally believed to be of Asiatic origin and ata very early date Phasianus colchicus was introduced into Europe,although Newton (1893-1896) thinks it not impossible that it mayhave been indigenous to Europe. This species introduced into Eng-land was called the English pheasant. Later the so-called Chinese,or ring-necked, pheasant {Phasianus colchicus torquatus) was intro-duced and hybridized with the pure colchicus, so that for the past 50years nearly all English pheasants, according to F. C. R. Jourdain,have had some trace at least of a white collar. Those first intro-duced into Oregon were P. colchicus torquatus, the Chinese pheasant,or Denny pheasant, as they were sometimes called, after JudgeDenny who introduced them; but those introduced in the easternpart of the United States were the mixed colchicus and torquatusrace then existing in England, and birds and eggs have been widelydistributed over the country. The resulting mixture, however, isgenerally called the ring-necked pheasant, Phasianus colchicustorquatus.Courtship.?The cock ring-necked pheasant is a magnificent bird,and the display of his brilliant plumage and beautiful form to thebest advantage before the demurely plumaged hens may well arousetheir admiration and passion. On English game preserves, whereonly the males are killed, or in many of our own States where onlythe cocks are allowed to be shot, each cock may have a large harem,yet where the hens are not spared polygamy doubtless is still therule. J. G. Millais (1909) notes, however, that in a wild state onerarely meets with more than three hens to a cock. Leffingwell (1928)says : " My opinion is that pheasants are naturally polygamous inthe wild, but that some males may have monogamous tendencies. Inthe spring it is most usual to see one male bird with several females."He thus describes the display
:
The male runs around the female with short steps, usually with the tipof the partly outstretched wing describing an arc on the ground, and stops infront of her. The feathers of the upper back, lower back, rump, and tail areshifted over to the side on which the female is, and the tail partly spread. Theneck is bent and the head kept low. Apparently the air sacs are partly in-flated, for after the pose is held for several seconds, the plumage is allowed tofall back to its natural position as the bird gives out a hissing sound.
RING-NECKED PHEASANT 313Millais (1909) described the courtship display as follows:At this season the usual walk is seen to be more reserved and dignified, andthe whole of the feathers are held out so as to give the bird a puffed appearance
;
the brilliant scarlet patch of skin round the eye is inflated and lowered beyondthe angle of the jaw and the purple ears erected and inclined outwards. Thebone-coloured bill is of a brighter hue and the eye, especially in the Mongoliansubspecies, very brilliant. Thus he proceeds until the moment of show. Thewing nearest the female is then lowered and extended, the scapulars droppeda little, the tail is also spread and turned over towards her, so that she maysee its full beauty. The feathers of the rump are also opened as far as pos-sible, the neck is lowered and curved and the head slightly turned to displaythe extended eye ornaments. If the female walks coquettishly away, or picksabout with apparent indifference, he is not annoyed but walks ahead to stopher and displays the other side of his person.Courtship displays of captive pheasants are easily studied, butunder favorable circumstances these may be seen in the wild. I havethus described my own observations (Townsend, 1920) :In courtship the ear-tufts of the cock are erected and the bare skin aboutthe eyes is prominent and very red. He struts before the hens turning in alldirections to display his gorgeous plumage, or walks, with an exaggerated bob-bing motion. Every now and again he flaps his wings almost inaudibly, crowsand flaps again with a loud clapping sound.The rivalry among the cocks leads to more or less fighting, afterwhich the victor, according to Beebe (1918), crows and flies off.Nesting.?The ring-necked pheasant, as a rule, nests on the groundin the open in fields of grass or grain and in bushy pastures, byhedgerows or roadsides or haystacks, very rarely in the woods andrarely at any great distance from water. The female is so protect-ively colored that one may pass within a few feet of the incubatingbird and not see her, and she trusts so much to this protective colora-tion that she rarely leaves the nest until almost stepped upon.When she does leave she generally skulks quietly away and rarelyflushes. She is much more difficult to see than the clutch of ex-posed eggs. A tuft, of grass or a group of luxuriant weeds helps inthe concealment of the nest, but in a field of grass or grain theeggs lie bare to the sky when the crop is harvested if they are notdestroyed in the process. F. J. Rice writes of such a nesting sit-uation found on August 10
:
The oats had, of course, been harvested at this date and the nest lay inthe stubble, with no concealment beyond some weeds, which grew up 6 or 8inches from the ground. There was but slight pretense of a nest, the nineeggs lying on the earth among the weeds. In an hour [after his dog hadflushed her] the bird was incubating again. She seemed very fearless anddid not leave the nest again until a team and hayrack were driven within 10feet of her.The dog, in hunting about the field, may have happened uponthe bird, for the sitting hen is protected not only by her coloration
314 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbut by her lack of scent. Leffingwell (1928) reports tests on prize-winning dogs conducted by M. C. Ware, secretary of the SouthernIdaho Field Trail Club. Two of these dogs, " in ranging over alarge area, pointed twelve birds, all males. As pheasants are veryabundant here, many females were undoubtedly present but were notfound because of their scentlessness." Newly hatched young, likethe incubating female, appear also to be scentless, for " three newlyhatched pheasants found by Mr. Ware could not be located by thedogs until the birds were seen."Pheasants have been known to lay their eggs in other birds' nests.Leffingwell (1928) reports that A. A. Allen found a nest of a ruffedgrouse containing eight eggs of the owner and four of a pheasant,and that Finley has found pheasant eggs in the nests of domesticfowl, bobwhites, ruffed grouse, and sooty grouse. In England pheas-ants have been observed to use the old nest of another bird in a tree(Van de Weyer, 1919, and Tegetmeier, 1911). The nest itself con-sists generally of a slight natural hollow in the ground, or one madeby the female, lined, sometimes very scantily, with leaves, grasses, orweed stalks.Eggs.?[Author's note: The ring-necked pheasant lays 6 to 14or 15 eggs, usually 10 to 12. They vary in shape from ovate toshort ovate. The shell is smooth with a very slight gloss. Theusual color is a rich, brownish olive, varying from " wood brown "or " avellaneous " to " dark olive-buff " or " olive-buff." Some ofthe darkest eggs are " buffy brown " and very rarely some eggs arepale blue. The measurements of 29 eggs in the author's collectionaverage 41.85 by 33.5 millimeters ; the eggs showing the four extremesmeasure 44.5 by 33.2, 42.6 by 35.1, 39.3 by 31.3, and 40 by 31.1 mil-limeters. Witherby's handbook (1920) gives the average of 35British eggs, a mixture of both races, as 45.93 by 36.04, the maxima49.2 by 37 and 48 by 39, and the minima, 39 by 36.5 and 46.8 by 34.5millimeters.]Young.?The length of incubation is stated to vary between 23and 25 days. Leffingwell (1928) says of 656 birds constituting thefirst hatch at a game club that 81.7 per cent hatched on the twenty-third day, 15.5 per cent hatched on the twenty-fourth day, and 2.7per cent hatched on the twenty-fifth day. In the second hatch 94.5per cent hatched on the twenty-third day, the rest on the twenty-fourth day. As a rule, the female alone incubates, but Tegetmeier(1911) records an instance of a cock pheasant not only building thenest but incubating and hatching the eggs, and he states that theincubation by the male has been observed in several cases of wildbirds.As soon as the down on the chicks has dried after they haveescaped from the eggshell and have fluffed out their feathers, they
RING-NECKED PHEASANT 315show great activity, and the mother bird leads them in search of food.If enemies, including the chief one, man, should disturb the familyat this time, the mother resorts to the " broken-wing " tactics andflops over the ground, endeavoring to lead the enemy away. Atother times the mother springs into the air with a loud whirring ofwings, while the young scatter and hide. Leffingwell (1928) saysthat " normally all the care of the young is undertaken by the femaleuntil the young birds are from six to seven weeks old. After thistime the cock bird occasionally wanders with the flock. I have seencock pheasants with a flock of six or seven week old birds on severaloccasions and as no females were observed it is not unlikely that theywere setting or raising another brood."The mother bird leads her brood about exactly like the domestichen, helps them to find food, and broods them in cold or storm andat night.The gain in weight of the young pheasants is at first rather slow.The weight is doubled soon after the second week, but is increasednearly sevenfold over the initial weight at 3 weeks, and at 5 weeksthe male chick weighs, according to Leffingwell's figure, about 15times his birth weight and is fully feathered. Leffingwell writesme:It is somewhat difficult to define the age at which young pheasants can fly.The birds are able to clear a 1-foot obstruction when they are 4 or 5 days old,but whether or not you can call this flying I do not know. Certainly whenthey are a week old they can fly distances of 4 or 5 feet.I copy the following from my notes of September 1, 1913
:
From my window this morning at 7 I saw a hen pheasant with a number ofhalf-grown young scale down the hill, alight on the edge of the " forest," anddisappear within. A few minutes later I heard a loud, insistent calling, keekee kee, which suggested somewhat a flicker. Then I saw the hen pheasantstanding erect, with head and neck stretched up, in the field, close to the trees.Soon she stopped calling and disappeared in the grass, and three young pheas-ants came sailing down the field to her. She had been drumming up thelaggards.Plumages.?[Author's note: The chick in down is thus describedin Witherby's handbook (1920) :Fore-bead and sides of crown buff to yellow-buff with blackish line or spotsdown sides, centre of crown dark red-brown to blackish-brown ; nape rufous
;
back of neck buff to yellow-buff with short blackish line in centre ; rest of up-per-parts rufous-buff with three wide black lines and wings with black blotchessides of head pale yellow-buff to pale buff with a brownish streak from baseof upper mandible and a black spot on ear-coverts ; under-parts buff white topale buffish-yellow, sometimes with a tawny tinge at base of throat.The same writer describes the juvenal plumage, as follows
:
Crown and back of neck dark brown, feathers with subterminal pale buffmarks giving a spotted appearance; mantle and scapulars brown-black,
316 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMfeathers with margins and central streaks of buff to pale buff, those of upper-mantle and base of back of neck more or less tinged rusty-rufous ; back, rump,and upper tail-coverts browner (less black) and with rather more pale buffmargins and centres and often bars; sides of head pale buff minutely speckledblack ; chin pale buff ; sides and base of throat same but streaked and with con-centric lines of dark brown ; flanks same but with thicker concentric lines orbroken bars of brown to black-brown ; tail feathers small and narrow, buffand rufous closely barred black ; wing feathers much as adult female, but withpale buff barring; wing-coverts as scapulars, but with larger centres, andgreater often with bars of pale buff.The postjuvenal molt begins when the young bird is about halfgrown ; it is a complete molt, except that the two outer primaries oneach wing are retained for a full year. By November or Decemberyoung birds have completed the molt into the first winter plumage,which is practically adult.I can find no evidence of a prenuptial molt in adults, but theyhave a complete postnuptial molt late in summer and early in fall,beginning in July or August and ending in September or October.Old females, whose ovaries have become inactive through old age ordisease, sometimes assume more or less of the male plumage. Ihave seen such a bird in a local collection in Taunton and another inthe collection of W. F. Peacock in Marysville, Calif. This bird, Mr.Peacock told me, had been raised in captivity and had laid some 280eggs. When 6 years old she quit laying, and a post-mortem examina-tion, 2 years later, showed that the ovaries were completelyatrophied. From the time she stopped laying the male plumagegradually developed, until finally she could easily be mistaken fora first winter male, having acquired the head and body plumage andthe long tail but not the comb and wattles of the male.]On the development of the male genital organs, Paul D. Dalke hassent me the following notes:The size and development of the testes in the pheasant have been noted inall the cock birds taken since January 22, 1930. A specimen taken on theabove date had testes that measured 4 by 10 millimeters. The next male wascollected on February 28, 1930. This was a young cock, but the testes had al-ready grown considerably and measured 12 by 18 millimeters. Another youngcock collected on March 5, 1930, had testes that measured 13 by 21 millimeters. Aspecimen of March 28 showed an increase in size of the testes to 15 by 25millimeters. On May 1, 1930, two cock pheasants were collected, one an oldcock and the other of the previous year's crop. The latter had testes measur-ing 16 by 30 millimeters and the former 22 by 35 millimeters.Food.?The food of the ring-necked pheasant consists of insects,weed seeds, wild fruits and berries, and cultivated crops. Occasion-ally small rodents are eaten. The economic status of the bird,whether harmful or beneficial, is dependent on local conditions andseasons of the year, and on the proportion between weed seeds and
RING-NECKED PHEASANT 317injurious insects eaten on the one hand and cultivated crops on theother.Pheasants are at times very destructive to sprouting corn and evento corn in the ear. They also are known to eat tomatoes, beets, peas,beans, and other farm crops, including grain of all sorts. Thestomachs of those shot doing damage in a garden often contain, how-ever, a large number of injurious insect pests as well as many weedseeds. Much of the grain?wheat, oats, rye, barley?eaten by pheas-ants is waste grain that has fallen to the ground during harvesting.In summer, according to an unpublished report kindly sent me byLeffingwell, 20.4 per cent of the contents of 11 pheasant stomachsconsisted of injurious insects, and insects are consumed almost exclu-sively by the broods of young. Stomach examinations made by vari-ous observers show that grasshoppers, crickets, potato beetles,squash bugs, curculio beetles, and larvae of all kinds, including thoseof the gypsy and brown-tail moths and including also the tent cater-pillar, are among the items in the food. With the injurious insects,a certain number of beneficial ones are eaten.McAtee (1912) records that the stomach of one pheasant contained360 larvae of March flies, Bibio, and another 432. " Twenty-threeacorns and 200 pine seeds were taken by the bird who ate the largestamount of mast, and about 800 capsules of chickweed, containingmore than 8,000 seeds were in the stomach of the weed-seedeater." McAtee sums up his report as follows : " What is most evi-dent is that pheasants are gross feeders, their capabilities for goodor for harm are great. If a number of them attack a crop they arelikely to make short work of it, or if they devote themselves to weedseeds or insect pests they do a great deal of good."Lefiingwell (1928) says that "in Minnesota, F. D. Blair, thesuperintendent of game laws, believes that pheasants destroy moremice per bird than do most of the hawks and owls." At the sea-shore, especially when the uplands are covered with snow, pheasantsvisit the salt marshes, where they probably consume small crusta-ceans and mollusks.Clarence Cottam (1929) says: "Except during the coldest winterthose stomachs collected at mid-day contained little food material,while those taken in the morning and evening were full or beingfilled. This suggests that pheasants usually feed during the earlymorning or late afternoon."Paul D. Dalke says in his notes
:
On January 22, 1930, I made a trip to collect pheasants 5 miles south ofAnn Arbor, Mich., where the birds are fairly abundant. I flushed one cockpheasant from a field overgrown with wild sweetclover; the cock had traveledonly 100 feet from the time he left the roost until 2 p. m. While travelingthis 100 feet he had rested three times, as indicated by the hollowed-out forms
318 BULLETIN 162, UNITED STATES NATIONAL MUSEUMin the snow and by the droppings. Although corn was available in an adjacentfield this pheasant had nothing in his crop and only grass in his gizzard.Behavior.?The introduction of the ring-necked pheasant hasadded greatly to the charms of the countryside. To watch a cockpheasant striding along the ground or launching into the air, show-ing his metallic-blue head, his snow-white neck ring, his golden-brown back, and his long magnificent tail is always worth while.The males, as if aware of their conspicuous shape and colors, gener-ally run away or flush at some distance, while the modest-hued hensdepend more on their protective coloration and allow a much closerapproach. Frequenting for the most part cultivated fields andbushy pastures, they are also to be found in swamps and moistthickets and among sand dunes. In the sand their tracks are aseasily seen and recognized as in the snow?three toe marks widelydiverging, with a dot or a line behind and on the inner side made bythe hind toe if the sand or snow is soft. Occasionally the tracksshow that they drag the middle toe, but the birds usually step clear.When the bird is walking, the distance between tracks is generally4 or 5 inches, but a stride of 7 or 8 inches is not uncommon, and Ihave measured several tracks where 18 inches was cleared at eachrunning step. In running the tail is cocked up at an angle of 45?,but when the bird is feeding it is generally held horizontally.Pheasants have been seen swimming across bodies of water (Teget-meier, 1911).When suddenly startled, pheasants flush with a loud metallic whirof the wings, but not so thunderous as that of the ruffed grouse.Occasionally a bird flies off almost silently. They are able to shootup nearly vertically if they are hemmed in by trees or a building,and they make off at a great speed, which has been estimated to reach38 miles an hour (Tegetmeier, 1911). When not much frightenedthey soon set their wings and scale, each primary standing out likefingers on a hand, while the long tail is spread to its utmost andcurved downward for a brake, and their wings are fluttered rapidlyas they approach a landing. In flight the feet are at first drawn upin front but are quickly jerked back and are held extended behindunder the tail. In very short distance flights this backward extensionmay not take place. It is generally believed that a pheasant's flightis limited to somewhat less than a mile, although, when helped by astrong wind, longer distances are accomplished. In a very strongwind a cock pheasant in attempting to fly over my house one Januaryday collided with a chimney and met his instant death.As to the distance to which pheasants wander from their coverts,Leffingwell (1928) states that of 16 banded birds recovered withinthree years " but two exceeded two miles, while the average distance
RING-NECKED PHEASANT 319covered was but one and one-fifth miles. One bird, however, wentsix miles and another three miles." Merriam (1889) records that apair in Oregon traveled in two months 50 miles from the pointwhere they were released.Pheasants, like domestic fowls, are fond of dusting themselves toget rid of lice, and dusting places are common in pastures where thesebirds are found. Pheasants spend the night on the ground and alsoin trees, the latter especially where foxes or other ground enemiesabound. Several times early in the morning I have seen pheasantsroosting in trees, singly, and once as many as five. As the sun rose,their breasts glistened like burnished copper in its rays.Under complete protection, where shooting is not permitted, as inpublic parks, pheasants become very tame, but when persecuted theyquickly develop great wariness, and they seem to be able to distin-guish the harmless farm laborer from the man with a gun. Any un-usual noise, such as blasting, makes pheasants crow, and they areusually sensitive to any shock, whether from an explosion or anearthquake, and respond by crowing. This response to earthquakesor distant explosions is apparently due to the sense of feeling ratherthan of hearing. In Japan pheasants are believed to give warningsof earthquakes. It is found that they respond to earthquake shocksso slight that they are unnoticed by human beings, and the birds mayin this way foretell a more severe earthquake shock that follows.Hartley (1922) states that " during the World War the pheasants inEngland developed into fairly responsible sentinels against Zeppelinattacks. The birds seemed particularly sensitive to far-off explo-sions and a raid generally was heralded by a concerted crowing ofcocks."William Beebe (1918) writes:In the spring the cock pheasant invites his mate or mates to share orappropriate some especially delectable morsel of food. The accompanyingmovement is a picking up and dropping of the food, thus calling it to visualattention, while at the same time a low chuckle or crowing sound is uttered.Voice.?The courtship " song " of the cock pheasant is his crow,which suggests a juvenile bantam rather than the noble pheasant.It is a challenge call by which the cocks announce their territorialholdings. This crow, which consists of a long followed by a shortnote, can be heard from a considerable distance, but when the ob-server is near at hand he hears also a hurried clapping of the wings,which is heard loudest following the crow and not preceding it, asin the case of the domestic cock. If, however, the bird is seen atthis time it will be observed that he flaps his wings two or threetimes almost inaudibly before the crow, and follows the crow with a
320 BULLETIN 162, UNITED STATES NATIONAL MUSEUMrapid succession of five or six flaps, which are audible for somedistance. Leffingwell (1928) states thatthe wings are held rather stiffly and the force of the beat is directed upwardsand inwards, somewhat after the manner of a drumming partridge. The twopreliminary wing beats are given at intervals of about one twenty-fifth of asecond, while those given after the call begin very rapidly but soon diminishin vigor. The force of the latter strokes seem to push the pheasant backwardsagainst its tail, which is partly flattened on the ground and acts as a brace.Crowing is, of course, commonest in the spring of the year, andI have heard one crowing on a pleasant day as early as mid-January.The birds are generally silent in the summer months, but they areoften heard again in October. The young are said to attempt tocrow when 7 or 8 weeks old.Besides the crow, the alarm notes, emitted when the birds arestartled, are most commonly heard. These are loud and hoarse croak-ings, which they emit as they fly away, sometimes in two or threesyllables, written cuck-et or tuck-ee-tuck, sometimes a prolongedand scolding repetition of croaking notes. These croaking notes,suggestive at times of an old domestic hen, may also be given whenthe bird is on the ground. I have heard a cock pheasant in a smallthicket croaking continuously, owing apparently to irritation causedby crows who were scolding him from the trees above. I have alsoheard a querulous queep, queep, queep given by a hen pheasant.I once disturbed a hen pheasant in feeding, and she looked up intime to see a fox creeping toward her from the other side. Thepheasant at once flew off uttering a whistling shriek, and the fox,interrupted in his turn, also departed. Millais (1909) says that "thecock pheasant when he has paired or gathered his wives, makes useof a gentle note or chuckle."The young have a variety of notes. Leffingwell (1928) lists fivedistinct calls in birds up to 7 weeks of age?calls expressive of con-tentment, caution, alarm and fright, and the flock call.Field marks.?The cock pheasant, with his resplendent plumage,white neck ring, and long tail, is easily identified. The hen andyoung might be mistaken for ruffed grouse except for the muchlonger tails.Game.?The cock pheasant is a prize well calculated to delight theheart of the sportsman. Splendid in plumage?a magnificenttrophy?large and heavy, and delicious eating, it tests his skill tothe utmost. In game preserves in England, numerous beaters drivethe pheasants to the quiescent sportsmen and force the birds to flyhigh and at great speed over them. In this country, the sportsmanseeks the bird, going on foot over the fields and shooting the pheas-ant as it flushes and makes off. Dogs are generally used, and al-though the pheasant often lies close to the pointing dogs and allows
RING-NECKED PHEASANT 321himself to be flushed by the sportsman, he may sometimes hide, or,worse still from the point of view of the owner of a well-traineddog, he may run long distances and entice the dog to follow. Onlyby continuous replacement with birds raised on private or Statefarms can the drain on pheasants by hunters be checked, for inthese days of automobiles hunters may visit numerous favorablelocalities many miles apart in the course of one day.A very important asset of the ring-necked pheasant, both fromthe sportsman's and the bird lover's point of view, is that it divertsgunfire from our fast-diminishing grouse and other game birds.As the natural habitat of the ruffed grouse is in woods, whilethat of the pheasant is usually in open fields and pastures, there isno direct conflict between these two birds, and from this pointof view there seems to be no harm resulting to the ruffed grousefrom pheasant introduction. On the other hand, bobwhites, hav-ing much the same habitat as the pheasant, might be displaced by thelatter bird if there were not enough food for both. Dr. George W.Field (1914) does not think there is any evidence that the pheasantinterferes with our native game and says:More conclusive is the testimony from records of the great shootingestates in England, Scotland, and Germany, where for at least two genera-tions accurate records have heen kept of the number of birds produced an-nually on each estate. These records show conclusively that there has beenno diminution in the number of native grouse and quail, and no displacementof these birds by the introduced and naturalized pheasant, which is here pro-duced in larger quantities than on any similar area in the world.In this connection the remarks of W. L. McAtee (1929) are perti-nent:There is little fear, however, that any of the large and highly edible speciesclassed as game birds will continue for any period as pests. Should theyexhibit destructive tendencies their numbers can easily be cut down by theextension of the open season and increase in the bag limits. No bird thatis widely prized for food is ever likely to become destructively abundant inthe United States.There is, however, a subtle danger from the introduced pheasant,that of carrying to our native game blackhead, the heterakis worm,and other diseases and parasites. Prevalence of the heterakis wormin wild pheasants has been shown by Dr. E. E. Tyzzer. McAtee(1929) does not think that this danger should be given much weight" in view of the fact that domestic poultry is constantly being intro-duced, abounds in all parts of the country, and constitutes a sourceand reservoir of most of the diseases to which our game birds aresusceptible. In other words, the disease hazard is scarcely likely tobe notably increased by further introductions of game birds. Thereshould, of course, be proper inspections of imported birds and ex-
322 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
elusion or quarantine when found necessary." Dr. J. C. Phillips(1928), however, says "the interesting point is that the pheasantmay now easily infect territory at a distance from farmyards."Enemies.?Man, of course, is the pheasants' worst enemy in destroy-ing them, but their best friend in conserving the stock that mightotherwise become extinct. All mammals that prey on eggs, and, ifthey are powerful enough, on the sitting or roosting birds, like rats,skunks, weasels, foxes, and coyotes, are more or less destructive topheasants. A few hawks and owls also take toll of pheasants, butthe damage done by these birds is probably insignificant, and theirinfluence may be of value in eliminating the feebler fliers and thediseased birds. Crows, however, undoubtedly eat many pheasanteggs and the far-wandering domestic cat is a very serious menaceto these ground-nesting birds.Winter.?When the ground is covered with a blanket of snow andice, pheasants are obliged to wander far for food. At this time, ifweed seeds are largely covered, they are driven to eat the buds ofbushes and trees and to seek manure and garbage heaps, and even topartake with the poultry on a farm. At such times, if the saltmarshes are still open, they frequent these for small mollusks andcrustaceans. I have found the thinner snow under a tree scratchedaway by pheasants in order to get at the seeds and dormant insectson the ground. Sometimes the long tails of the males are frozento the snow and ice during the night and the birds held captive.DISTRIBUTIONRange.?Introduced and now fairly well established in approx-imately the northern half of the United States and in extremesouthern Canada. North to southern British Columbia (VancouverIsland and Fraser Valley) ; southern Alberta; southern Manitoba;southern Ontario (Kent County and north shore of Lake Erie)
;
central New York and Vermont (Lake Champlain region) ; centralNew Hampshire (Concord, Hanover, and Plymouth) ; and south-western Maine. East to the Atlantic coast. South to Maryland(north of Baltimore); Pennsylvania; Ohio; Kentucky; Missouri;Kansas; Colorado; and California (Inyo, Tulare, and Kern Coun-ties and Santa Clara Valley). West to the Pacific coast.Introductions into the eastern Provinces of Canada and into theStates south of those named above have not been successful.Egg dates.?Washington and Oregon : 5 records, April 13 to June17. California : 2 records, May 3 and June 10. Michigan, April 17.Massachusetts, May 16. Pennsylvania: 2 records, May 12 andJune 4.
merriam's turkey 323Family MELEAGRIDIDAE, TurkeysMELEAGRIS GALLOPAVO MERRIAMI NelsonMERRIAM'S TURKEYHABITSThe wild turkey of the mountain regions of the SouthwesternUnited States and extreme northwestern Mexico was described by Dr.E. W. Nelson (1900) and named in honor of Dr. C. Hart Merriam.He has characterized it as follows : " Distinguished from M. g. feraby the whitish tips to feathers of lower rump, tail-coverts, and tail
;
from M. g. rnexicana by its velvety black rump and the greateramount of rusty rufous succeeding the white tips on tail-coverts andtail, and the distinct black and chestnut barring of middle tailfeathers."Nelson showed in the same paper that the ancestors of our domesticturkeys were neither of the forms that we now call merriami andintermedia but the more southern, strictly Mexican form, M. gallo-pavo gallopavo.That this wild turkey is not nearly so abundant as it was 50 yearsago is shown by the following quotation from Henry W. Henshaw(1874) :The wild turkey is found abundantly from Apache throughout the mountain-ous portion of Southeastern Arizona. In New Mexico it was met with furtherto the north, in the mountains, and I was informed by Colonel Alexander thathe had found them in large numbers in the Raton Mountains, in extremeNorthern New Mexico. It breeds abundantly through the White Mountains,Arizona, and about the middle of August several broods of the young, about two-thirds grown, were met with. Toward the head of the Gila, in New Mexico, thecanons, in November, were found literally swarming with these magnificentbirds ; in many places the ground being completely tracked up where they hadbeen running. As many as eleven were killed by the members of a party duringa day's march.Nesting.?Two brief notes by Major Bendire (1892) are all thathe gives us on the nesting habits of this turkey, which are probablynot very different from those of other wild turkeys. He quotesWilliam Lloyd as saying that " near a river their nests would be madeon small inlets surrounded by reeds ; on the hills in shin-oak clumps."He says that Frank Stephens found a nest on the east slope of theSanta Rita Mountains in Arizona, " in the oak timber, just where thefirst scattering pines commenced, at an altitude of perhaps 500 feet.It was placed close to the trunk of an oak tree on a hillside, nearwhich a good-sized yucca grew, covering, apparently, a part of thenest; the hollow in which the eggs were placed was about 12 inchesacross and 3 inches deep."
324 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMMrs. Florence M. Bailey (1928) says the nest is " on the groundin tall thick weeds or briers, lined with grass, weeds, and leaves."O. W. Howard (1900) found a nest in the Huachuca Mountains inArizona, which he describes as follows
:
The nest was in the bed of the canon at the base of the hill, in a naturaldepression in the soft earth at the side of a rock, and just under a large whiteoak tree. The nest had a lining of leaves and small twigs, with a few feathersfrom the old bird scattered about. The nest was about a mile above the placewhere I had seen the first bird and at about 7,000 feet elevation. Strange tosay. the nest was within a stone*s throw and in plain sight from a well-traveledtrail.Eggs.?The eggs are indistinguishable from those of other wildturkeys. The measurements of 16 eggs average 65.8 by 47.3 milli-meters; the eggs showing the four extremes measure 70.5 by 49,61.7 by 46.7, and 64.5 by 46 millimeters.Food.?Mrs. Bailey (1928) lists the food of this turkey as follows:In winter pinyon nuts, acorns, and juniper berries ; in summer flower buds,grass and other seeds, wild oats, wild strawberries, manzanita berries, rosehaws, fruit of wild mulberry and prickly pear, grasshoppers, crickets, beetles,caterpillars, ants, and other insects. In New Mexico " the crop of a MerriamTurkey killed February 10 on Haut Creek contained 76 juniper berries, 25pinyon nuts, 6 acorns, 30 soft worms an inch long, grass blades and some rock.The crop of a gobbler, weighing about 30 pounds and shot March 25 out of aflock of 50 in the Black Mountains, contained 30 pinyon nuts and 215 juniperberries" (Ligou). The stomach of a specimen collected near the southern endof the range contained fully a half pint of the fruiting panicles of grass(Muhlen'bcrgia) , a few seeds of Bromus, and some grass blades comprising 55per cent; pinyon pine and other pine seeds, 45 per cent. In some localitiesconsiderate ranchmen plant small patches of oats for turkey food (Ligon).Mrs. Bailey quotes Charles Springer as saying
:
At times, and particularly in years when there are few or no nuts, the prin-cipal food of the Merriam Turkey is wild rye, which is plentiful in the canyonsand draws in our mountains and foothills. On the Suree I have often seen wildTurkeys eating the short blades of Kentucky blue grass which grows wild alongthe canyon near the stream and remains green all winter. One of the mostimportant winter foods of the Merriam Turkey is the red kinicklnick berrywhich grows on the high ridges and plateaus in our mountains. When acorns,pinyon, and pine nuts, and other foods may be buried deep under snow, the Tur-keys may find kinickiuick berries on the high ridges and high places from whichthe snow blows off. Mason Chase tells about the wild Turkeys hunting out, orat least finding and appropriating, caches of nuts made by rodents. He saysthis occurred during a time when deep snow covered up all the Turkey's foodexcept the buds of shrubs.Dr. C. Hart Merriam (1890) found it on San Francisco Mountain,Ariz., feeding on wild gooseberries in the balsam belt in August andon pinyon nuts in the cedar belt in September. Major Bendire(1892) says that it also eats the fruits of the giant cactus, " which isalike a favorite article of food with man, bird and beast."
merriam's turkey 325Behavior.?Henshaw (1874) writes:They roost at night in the large cotton-woods by the streams, and soonafter daylight, having visited the stream, they usually betake themselvesto the dry hills, where they feed, in the fall, at least, almost exclusively uponthe seeds of grasses and grasshoppers. I think they return once or twiceduring the day to drink, the dry nature of their food rendering a copioussupply of water necessary. In these wilds, they appear to be wholly unsuspi-cious, and without knowledge of danger from man, and, if not shot at, willallow one to get within a few yards without manifesting any distrust. Theyrarely fly, except when very hard pressed, but, when alarmed, run with suchrapidity as to quickly outstrip the fleetest foot, betaking themselves to thesteep sides of the ravines, which they easily scale, and soon elude pursuit.Apparently, the only dangers they have to fear in these regions are frombirds of prey, which attack the young, but more especially from the panthers.In certain portions of the Gila Canyon the tracks of these animals were verynumerous, and always these sections appeared to have been entirely depopu-lated of Turkeys, an occasional pile of feathers marking the spot where onehad fallen a victim to one of these animals.DISTRIBUTIONRange (entire species).?Southern Ontario; the Eastern, Central,and Southern United States, including the southern Kocky Mountainregion ; and Mexico, except the extreme western and southern parts.The range of the wild turkey has been greatly restricted since theadvent of civilization, so that the species is now extirpated through-out New England and the Great Plains. It is still common locallyin Pennsylvania (largely through introductions) and in some of theSouthern States, as South Carolina, Georgia, Florida, Louisiana,Texas, New Mexico, and Arizona.The complete range of the species extended north to Arizona (BillWilliams Mountain, and San Francisco Mountain) ; formerly Colo-rado (Coventry. Salida, and Buckhorn) ; formerly Nebraska (Valen-tine) ; formerly southeastern South Dakota (Cedar Island, FortRandall, Yankton, and Vermilion) ; formerly Iowa (Grant City,Ames, and Fort Atkinson) ; formerly southern Wisconsin (Newark,Lake Koshkonong, Waukesha, and Racine) ; formerly southern Mich-igan (Grand Rapids, Locke, and Reece) ; formerly southern Ontario(Mitchells Bay, Plover Mills, and Dundas) ; formerly northern NewYork (Niagara County) ; and probably formerly southern Maine(Mount Desert Island). East to probably formerly southern Maine(Mount Desert Island) ; formerly Massachusetts (Ipswich, Montague,and Mount Holyoke) ; formerly Connecticut (Northford) ; formerlyNew Jersey (Sussex County, Raccoon, and Cape May County) ; Vir-ginia (Neabsco Creek and Ashland) ; North Carolina (Walke and theCape Fear River) ; South Carolina (Georgetown, Santee, MountPleasant, and Charleston) ; Georgia (Riceboro, Macintosh, Cumber-74564?32 22
326 BULLETIN 162, UNITED STATES NATIONAL MUSEUMland Island, and St. Marys) ; eastern Florida (Port Orange, OakHill, Malabar, Fort Kissimmee, and Everglade) ; Tamaulipas (Sotola Marina and Forlon) ; and Vera Cruz (Zacuapan, Mirador, andSoledad). South to Vera Cruz (Soledad and Paso del Macho);Hidalgo (Real del Monte) ; and Michoacan (La Salada). West toMichoacan (La Salada) ; Durango (Durango and El Salto) ; Chihua-hua (Colonia Garcia and Cajon Bonito) ; and Arizona (formerlyHuachuca Mountains, formerly Santa Catalina Mountains, SaltRiver Bird Reservation, Sierra Ancha, Baker Butte, Apache MaidMountain, and Bill Williams Mountain).The range as above described is for the entire species, which has,however, been divided into four subspecies. The eastern wild turkey(Meleagris g. silvestris) ranged over the entire eastern part of thecountry north of central Florida and west to eastern Texas, centralKansas, Nebraska, and Oklahoma ; the Florida turkey (M. g. osceola)ranges through southern Florida; the Rio Grande turkey (Af. g.intermedia) is found in central and southern Texas, chiefly betweenthe Brazos and Pecos Rivers, north to the Staked Plains, and innortheastern Mexico; and Merriam's turkey (M. g. merriami) isfound in mountainous regions in New Mexico, Arizona, and Colorado,ranging also into western Texas and northern Mexico.Wild turkeys have been restored to parts of the range from whichthey had been exterminated, notably in Pennsylvania. They alsohave been carried north successfully as far as Minnesota (State gamefarm at Minneapolis)
.
Egg dates.?Michigan: 3 records, February 10 and May 5 and6. Pennsylvania: 5 records, May 5 to June 30. South Carolinaand Georgia: 15 records, March 30 to May 25; 8 records, April 25to May 22. Missouri and Arkansas: 8 records, April 3 to June2. Louisiana, Oklahoma, and Texas (silvest?*is) : 20 records, April9 to July 25; 10 records, May 3 to 16. Arizona, Mexico, and NewMexico {merriami) : 7 records, April 8 to May 8. Texas andMexico (intermedia) : 23 records, March 4 to June 28 ; 12 records,May 1 to June 3. Florida (osceola) : 15 records, March 25 to May22 ; 8 records, April 10 to May 3.MELEAGRIS GALLOPAVO SILVESTRIS VieillotEASTERN TURKEYHABITSWhen the noble red man roamed and hunted unrestrained through-out the virgin forests of eastern North America, this magnificentbird, the wild turkey, another noble native of America, clad in afeathered armor of glistening bronze, also enjoyed the freedom of the
EASTERN TURKEY 327forests from Maine and Ontario, southward and westward. But thecoming of the white man to our shores spelled the beginning of theend for both of these picturesque Americans. The forests disap-peared before the white man's ax, his crude firearms waged warfareon the native game, and the red man was gradually eliminated beforeadvancing civilization. In the days of the Pilgrims and Puritansthe Thanksgiving turkey was easily obtained almost anywhere in thesurrounding forest; the delicious meat of the wild turkey was animportant and an abundant food supply for both Indians and set-tlers; and the feathers of the turkey held a prominent place in thered man's adornment.Thomas Morton (1637), one of the earliest writers, says:Turkies there are, which divers times in great flocks have sallied by ourdoores ; and then a gunne, being commonly in a redinesse, salutes them withsuch a courtesie, as makes them take a turne in the Cooke roome. They daunceby the doore so well.They soon began to disappear, however, for John Josselyn (1672)writes
:
I have also seen threescore broods of young Turkies on the side of a marsh,sunning of themselves in a morning betimes, but this was thirty years since, theEnglish and the Indians having now destroyed the breed, so that 'tis very rareto meet with a wild Turkie in the Woods.Edward H. Forbush (1912) says:In Massachusetts Turkeys were most numerous in the oak and chestnutwoods, for there they found most. food. They were so plentiful in the hills bor-dering the Connecticut valley that in 1711 they were sold in Hartford at oneshilling four pence each, and in 1717 they were sold in Northampton, Mass., atthe same price. From 1730 to 1735 the price of those dressed was in North-ampton about one and one-half penny per pound. After 1766 the price wastwo and one-half pence, and in 1788, three pence. A few years after 1800 itwas four pence to six pence a pound, and about 1820, when the birds had greatlydecreased, the price per pound was from ten to twelve and one-half cents.Wild turkeys made their last stand in Massachusetts in the Holyokerange, where the last one was killed in 1851. According to Dr. D. D.Slade (1888)these birds had the range of a large tract of wild mountainous country, in someparts almost inaccessible and impassable, lying at the base of and comprisingMount Holyoke, and to the Southwest also including Mount Tom and its sur-roundings. I am unable to state the exact period at which this flock becameexterminated but should say it must have been in 1840 or thereabouts.The last turkey in Connecticut was seen in 1813, a few remainedhidden in the Vermont Hills until 1842, and they were said to benumerous along the southern border of Ontario as late as 1856.Albert H. Wright (1914 and 1915) has written a very complete ac-count of the early history of the wild turkey to which the reader is
328 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
referred. Dr. Glover M. Allen (1921) also has given us a very fullhistory of this bird in New England. Both of these exhaustivepapers give far too much information to be included here.Audubon (1840) wrote, as to its status in his time:The unsettled parts of the States of Ohio, Kentucky, Illinois, and Indiana,an immense extent of country to the north-west of these districts, upon theMississippi and Missouri, and the vast regions drained by these rivers fromtheir confluence to Louisiana, including the wooded parts of Arkansas, Ten-nessee, and Alabama, are the most abundantly supplied with this magnificentbird. It is less plentiful in Georgia and the Carolinas, becomes still scarcerin Virginia and Pennsylvania, and is now very rarely seen to the eastward ofthe last-mentioned States. In the course of my rambles through Long Island,the State of New York, and the country around the Lakes, I did not meetwith a single individual, although I was informed that some exist in thoseparts. At the time when I removed to Kentucky, rather more than a fourthof a century ago, Turkeys were so abundant, that the price of one in the marketwas not equal to that of a common barn fowl now. I have seen them offeredfor the sum of three pence each, the birds weighing from ten to twelve pounds.A first-rate Turkey, weighing from twenty-five to thirty pounds avoirdupois,was considered well sold when it brought a quarter of a dollar.In the mountains of central Pennsylvania turkeys have alwaysexisted up to the present time. C. J. Pennock tells me that they"have multiplied greatly within the last 15 years," but that mostof the " stock has been intermixed with domestic birds." This islargely due to the efforts of the game commission in controlling thehunting season and bag limits and by importing birds from otherStates or transferring them from a section where they are plentifulto one where they are scarce. A report of this has been publishedin some detail by Bayard H. Christy and George M. Sutton (1929).M. P. Skinner wrote to me in 1928 :The wild turkey is still found over most of North Carolina wherever thereare undisturbed forests of the kind preferred by the turkey. In the sand hillsthere are still two or three groups living mostly in the swamps and river bot-toms, and totaling perhaps 30 birds in all. They are resident and nonmigratory.In the sand hills, wild turkeys have largely retired to the deep swamps,for they prefer to roost only in trees standing in water; but quite often theyfeed out on the drier upland.James G. Suthard says in his notes, sent to me in 1930
:
This noble game bird formerly bred in Kentucky, at large, but at the presenttime has a very restricted range. It is found in the areas bordering Virginiaand Tennessee, in Taylor, Larue, and Hart Counties in central Kentucky, andalso in the game preserve in Lyon and Trigg Counties. I have some recordsfor Fulton and Hickman Counties. It is possibly found straggling in othercounties, but, because of its retiring habits, I have never seen it, nor do Ihave any authentic records other than those already mentioned. It breeds dur-ing April and May, sometimes late in June.
EASTERN TURKEY 329Wild turkeys are essentially woodland birds. When the EasternStates were largely covered with virgin forests, they ranged widelyover the whole of these districts. As the land became cleared theyoften resorted to clearings, open fields, savannas, or meadows insearch of grasshoppers, other insects, berries, and other foods. Astheir numbers were reduced by persistent hunting, they became veryshy and were forced to retire to the wooded hills and mountains,where in many places they made their last stand. There are manyhills and creeks named for this bird because turkeys were once com-mon there. Turkeys are now found, in the Northern and EasternStates, only in the more remote and heavily wooded mountains, thewildest and least frequented forests, or the most inaccessible swamps,far from the haunts of man. In the Southern States they are muchmore abundant and more widely distributed. M. L. Alexander(1921) says of their haunts in Louisiana, which are typical:The determining factor in the distribution of turkeys is the occurrence ofoaks, wild pecans, beech and other nut-bearing trees. It is chiefly the oaksthat attract them to the flatwoods type of river lands, while the beech, chin-quapin, and certain species of oaks furnish the mast on the slopes of creeks,ravines and small rivers in pine regions. Dogwood, holly, black gum andhuckleberry are among other trees and shrubs, growing chiefly on slopes andridges, that furnish food for turkeys. Such food is not generally available,however, unless there is sufficient undergrowth to protect the birds whilethey are feeding. Late in the winter, after the best of the berries and mastin the bottoms of the hill sections have been picked up, or washed out by therains, the turkeys frequent southerly slopes, with a good cover of brush, scratch-ing in the fallen leaves and other woodland debris for such seeds and insectsas may be concealed there.Courtship.?The courtship display of the turkey gobbler is too wellknown to need any description here. The wild turkey's display issimilar, with the same expansion of body plumage, erection andspreading of the fan-shaped tail, swelling of the naked head orna-ments, and the drooping and rattling of the wing quills, accompaniedby gobbling and strutting.Audubon (1840) mentions a peculiar feature of the gobbler atthis season, the " breast sponge." which fills the upper part of thebreast and crop cavity. This is a thick mass of cellular tissue,which serves as a reservoir of sweet, rich oil and fat, on which thegobbler draws to supply the loss of flesh and energy during themating season.The object of the display and the gobbling notes is, of course, toattract the females. Turkeys are polygamous, the gobbler havingmany mates and serving them all every day during the laying seasonuntil his vigor is exhausted. The females separate from the malesbefore the mating season, and each hen comes to her favorite cockonce each day, for a short time, during the laying season. She
330 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMkeeps the nest concealed from him and shuns him after the eggs arelaid, lest he might break the eggs to prolong his sexual enjoyment.The gobbler often begins to display and gobble before he leaves hisroosting tree. He gobbles, watches, and waits until he sees the henapproaching, or hears her responsive yelp or cluck. He flies downto the ground, struts and gobbles again, and waits for the hen tocome to him. He probably knows how many hens he has in hisharem and keeps on strutting and gobbling until he has served themall. He roosts in the vicinity and repeats the performance every dayuntil the laying season is over or until he becomes emaciated andtakes no further interest in the hens.Audubon (1840), who had far better opportunities for observingthe wild turkey than can ever be had again, writes:I have often been much diverted, while watching two males in fierce conflict,by seeing them move alternately backwards and forwards, as either had obtaineda better hold, their wings drooping, their tails partly raised, their body-feathers ruffled, and their heads covered with blood. If, as they thus struggle,and gasp for breath, one of them should lose his hold, his chance is over, forthe other, still holding fast, hits him violently with spurs and wings, and in afew minutes brings him to the ground. The moment he is dead, the conquerortreads him under foot, but, what is strange, not with hatred, but with all themotions which he employs in caressing the female.When the male has discovered and made up to the female (whether sucha combat has previously taken place or not), if she be more than one year old,she also struts and gobbles, turns round him as he continues strutting, suddenlyopens her wings, throws herself towards him, as if to put a stop to his idledelay, lays herself down, and receives his dilatory caresses. If the cock meeta young hen, he alters his mode of procedure. He struts in a different manner,less pompously and more energetically, moves with rapidity, sometimes risesfrom the ground, taking a short flight around the hen, as is the manner of somePigeons, the Red-breasted Thrush, and many other birds, and on alighting, runswith all his might, at the same time rubbing his tail and wings along theground, for the space of perhaps ten yards. He then draws near the timorousfemale, allays her fears by purring, and when she at length assents, caressesher.Nesting.?Audubon says on this subject, referring to the South-ern States
:
About the middle of April, when the season is dry, the hens begin tolook out for a place in which to deposit their eggs. This place requires to beas much as possible concealed from the eye of the Crow, as that bird oftenwatches the Turkey when going to her nest, and, waiting in the neighbourhooduntil she has left it, removes and eats the eggs. The nest, which consists ofa few withered leaves, is placed on the ground, in a hollow scooped out, bythe side of a log, or in the fallen top of a dry leafy tree, under a thicket ofsumach or briars, or a few feet within the edge of a canebrake, but always ina dry place. The eggs, which are of a dull cream colour, sprinkled with reddots, sometimes amount to twenty, although the more usual number is from tento fifteen. When depositing her eggs, the female always approaches the nestwith extreme caution, scarcely ever taking the same course twice; and when
EASTERN TURKEY 331
about to leave them, covers them carefully with leaves, so that it is very diffi-cult for a person who may have seen the bird to discover the nest. Indeed, fewTurkeys' nests are found.When an enemy passes within sight of a female, while laying or sittingshe never moves, unless she knows that she has been discovered, but croucheslower until he has passed. I have frequently approached within five or sixpaces of a nest, of which I was previously aware, on assuming an air of care-lessness, and whistling or talking to myself, the female remaining undisturbed
;
whereas if I went cautiously towards it, she would never suffer me to approachwithin twenty paces, but would run off, with her tail spread on one side, to adistance of twenty or thirty yards, when assuming a stately gait, she wouldwalk about deliberately, uttering every now and then a cluck. They seldomabandon their nest, when it has been discovered by men ; but, I believe, nevergo near it again when a snake or other animal has sucked any of the eggs. Ifthe eggs have been destroyed or carried off, the female soon yelps again for amale; but, in general, she rears only a single brood each season. Several henssometimes associate together, I believe for their mutual safety, deposit theireggs in the same nest, and rear their broods together. I once found three sit-ting on forty-two eggs. In such cases, the common nest is always watched byone of the females, so that no Crow, Raven, or perhaps even Pole-cat, daresapproach it.Bendire (1892) refers to nests found in Nebraska and Texas inmore open situations. One is described as " a simple affair, on agrassy hillside, in an exposed position, and lined with dead grass."George M. Sutton (1929) describes a nest in Pennsylvania, asfollows
:
On June 6, on a rocky mountainside about twelve miles from Lock Haven,Clinton County, I examined a nest which held seventeen well-incubated eggs.On the day before there had been eighteen eggs in it ; it is thought that a skunkor fox had disturbed the nest, though the female bird evidently had been sittingclosely most of the time. This nest was built among small, angular rocks, and,while not very well hidden from above, it was screened on all sides by thicklaurel, which made photography difficult. The female bird was either very un-suspicious or remarkably brave, for she did not leave her nest while we werenear. Her broad back, with its squamate pattern and dull greenish lights, wasdifficult to discern among the foliage and the intricate interlacing of shadows.When first seen her neck was stretched out at full length in front of her, andher plumage was spread and flattened out noticeably. When she realized shewas being observed she drew her head back and moved it slowly about in asnakelike manner, while she gave forth strange hissing and grunting sounds.When she had become accustomed to us she again stretched her neck out infront of her. Occasionally, when disturbed, she gave a characteristic quit,quit.Eggs.?The normal set for the wild turkey numbers from 8 to 15eggs. The smaller sets are laid by young birds. As many as 18or 20 eggs have been found in a nest, which were probably laid byone bird. Occasionally two, or even three, birds lay in the same nest,taking turns at incubating or guarding the nest; in such cases thenest may contain many more eggs.The eggs are usually ovate in shape, but sometimes they are shortovate, or elongate ovate and quite pointed. The shell is smooth, with
332 BULLETIN" 162, UNITED STATES NATIONAL MUSEUMlittle or no gloss. The ground colors vary from " pale ochraceous-buff " or " pale pinkish buff " to " cartridge buff " or buffy white.They are more or less evenly marked with small spots and fine dotsof " light vinaceous-drab," " pale purple-drab," " clay color," or
" pinkish buff." The measurements of 56 eggs average 62.6 by 44.6millimeters; the eggs showing the four extremes measure 68.5 by 46,64.5 by 48.5, 59 by 45, and 64.7 by 42.4 millimeters.Young.?The period of incubation is 28 days and this duty isperformed by the female alone in seclusion. The male does not evenknow the location of the nest. The following is from Audubon's(1840) matchless account:The mother will not leave her eggs, when near hatching, under any circum-stances, while life remains. She will even allow an enclosure to be made aroundher, and thus suffer imprisonment, rather than abandon them. I once witnessedthe hatching of a brood of Turkeys, which I watched for the purpose of securingthem together with the parent. I concealed myself on the ground within avery few feet, and saw her raise herself half the length of her legs, lookanxiously upon the eggs, cluck with a sound peculiar to the mother on suchoccasions, carefully remove each half-empty shell, and with her bill caress anddry the young birds, that already stood tottering and attempting to make theirway out of the nest. Yes ; I have seen this, and have left mother and young tobetter care than mine could have proved, to the care of their Creator and mine.I have seen them all emerge from the shell, and, in a few moments after,tumble, roll, and push .ach other forward with astonishing and inscrutableinstinct.Before leaving the nest with her young brood, the mother shakes herself in aviolent manner, picks and adjusts the feathers about her belly, and assumesquite a different aspect. She alternately inclines her eyes obliquely upwardsand sideways, stretching out her neck, to discover hawks or other enemies,spreads her wings a little as she walks, and softly clucks to keep her innocentoffspring close to her. They move slowly along and, as the hatching generallytakes place in the afternoon, they frequently return to the nest to spend thefirst night there. After this they remove to some distance, keeping on thehighest undulated grounds, the mother dreading rainy weather, which isextremely dangerous to the young in this tender state, when they are onlycovered by a kind of soft hairy down of surprising delicacy. In very rainy sea-sons, Turkeys are scarce, for if once completely wetted the young seldom re-cover. To prevent the disastrous effects of rainy weather the mother, like askilful physician, plucks the buds of the spice-wood bush and gives them to heryoung.In about a fortnight the young birds, which had previously rested on theground, leave it and fly at night to some very large low branch, where theyplace themselves under the deeply curved wings of their kind and careful parent,dividing themselves for that purpose into two nearly equal parties. After thisthey leave the woods during the day and approach the natural glades or prairiesin search of strawberries and subsequently of dewberries, blackberries, andgrasshoppers, thus obtaining abundant food and enjoying the beneficial influenceof the sun's rays. They roll themselves in deserted ants' nests to clear theirgrowing feathers of the loose scales and prevent ticks and other vermin from
EASTERN" TURKEY 333
attacking them, these insects being unable to bear the odor of the earth inwhich ants have been. The young Turkeys now advance rapidly in growth andin the month of August are able to secure themselves from unexpected attacksof Wolves, Foxes, Lynxes, and even Cougars by rising quickly from the groundby the help of their powerful legs, and reaching with ease the highest branchesof the tallest trees. The young cocks show the tuft on the breast about thistime and begin to gobble and strut, while the young hens pur and leap in themanner which I have already described.C. J. Pennock writes to me that in northern Florida, where theturkeys are somewhat intermediate but rather nearer the northernform, a cold, wet spell late in April or early in May produces con-siderable mortality among the young and that after such an unfavor-able season turkeys are much scarcer for one or more years. Theweather also has much to do with the time of laying. He has seenyoung able to fly as early as May 26 and a brood of very young aslate as July 9. At times he has seen two hens together with theircombined broods of 20 or more young. The young are able to flyup into the trees when about one-third grown. The broods of youngremain with their mothers all through the winter and until the springmating time comes.Plumages.?In the wild-turkey chick the crown is " pinkish cinna-mon " and the back a somewhat lighter shade of the same, fading offto still lighter shades on the breast and flanks ; the crown and upperparts are heavily spotted or blotched with dark, rich browns, " bis-ter " to " Vandyke brown " ; the sides of the head and underparts are
" pale pinkish buff " to " ivory yellow," nearly white on the chin andthroat and almost " straw yellow " on the belly.As with the quail and grouse, the young turkey starts to grow itswings when a small chick ; these are soon followed by the plumage ofthe back, breast, and flanks; the tail comes later, followed finally bythe head and belly. The juvenal feathers of the back are "walnutbrown," edged with "russet," with a broad median "russet" stripe,a whitish tip, and large black areas near the tip; the wing coverts aresimilar, but in duller colors and with less black; the scapulars are
" sayal brown," peppered with black and spotted or barred with blackalong the outer edge and at the tip; the tertials and secondaries are
"hair brown," marked like the scapulars on the outer edge; the pri-maries are "hair brown," mottled and peppered with buffy white;the underparts are " fuscous," with whitish tips and shaft streaks
;
the tail is barred with dusky and " pinkish cinnamon."Before the young bird is fully grown, in September, a postjuvenalmolt takes place; this is a complete molt, except that the two outerprimaries on each wTing are retained for a year. In this first winterplumage the sexes begin to differentiate, the males becoming muchlarger than the females, but the plumages of the two sexes are very
334 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMmuch alike, and they resemble the adult female. Wilson (1832)says:On the approach of the first winter the young males show a rudiment of thebeard or fascicle of hairs on the breast, consisting of a mere tubercle, andattempt to strut and gobble ; the second year the hairy tuft is about three incheslong; in the third the turkey attains its full stature, although it certainly in-creases in size and beauty for several years longer.Audubon's (1840) statement is similar.Wilson (1832) says of the female:Females four years old have their full size and colouring ; they then possessthe pectoral fascicle, four or five inches long (which, according to Mr. Audubon,they exhibit a little in the second year, if not barren), but this fascicle is muchthinner than that of the male. The barren hens do not obtain this distinctionuntil a very advanced age; and, being preferable for the table, the hunterssingle them from the flock and kill them in preference to the others. The femalewild turkey is more frequently furnished with the hairy tuft than the tame one,and this appendage is gained earlier in life. The great number of young henswithout it has no doubt given rise to the incorrect assertion of a few writers thatthe female is always destitute of it.Adults apparently have only one complete postnuptial molt inAugust and September. A fully grown gobbler seldom weighs morethan 20 or 25 pounds, even when in good condition ; there are some,apparently authentic, records of birds weighing between 30 and 40pounds, but such cases must be very rare; reported records of 50pounds are unreliable.Food.?Dr. Sylvester D. Judd (1905a) found that the stomachsand crops of 16 wild turkeys examined by the Biological Surveycontained 15.57 percent of animal matter and 84.43 percent of vegetable matter.The animal food consisted of insects?15.15 percent?and miscellaneous in-vertebrates, such as spiders, snails, and myriapods?0.42 percent. Grasshoppersfurnished 13.92 percent, and beetles, flies, caterpillars, and other insects 1.23 per-cent. The 84.43 percent of the bird's vegetable food was distributed as follows
:
" Browse," 24.80 percent ; fruit, 32.98 percent ; mast, 4.60 percent ; other seeds,20.12 percent ; miscellaneous vegetable matter, 1.93 percent.Judd says that they are very fond of grasshoppers and crickets,and thatduring the Nebraska invasion of Rocky Mountain locusts, Professor Augheyexamined the contents of six wild turkey stomachs and crops collected duringAugust and September. Every bird had eaten locusts, in all amounting to 259.The wild turkey has been known also to feed on the cotton worm (Alabama,argillacea), the leaf hoppers, and the leaf-eating beetles (Chrysomela suturalis).The grasshopper (Arnilia sp.) and the thousand-legs (Julus) form part of theturkey's bill of fare. Tadpoles and small lizards also are included.Of a bird shot in Virginia, he says
:
Ten percent of its food was animal matter and 90 percent vegetable. Theanimal part consisted of 1 harvest spider (Phalangidae), 1 centipede, 1thousand-legs (Julus), 1 ichneumon fly (Ichneumon unifasieulata), 2 yellow-
EASTERN TUEKEY 335jackets (Vespa germanica), 1 grasshopper, and 3 katydids {Cyrtophyllusperspiculatus). The vegetable food was wild black cherries, grapes, berries offlowering dogwood and sour gum, 2 chestnuts, 25 whole acorns (Quercuspalustris and Q. velutina), a few alder catkins, seeds of jewel weed, and 500seeds of tick-trefoil (Meibomia nudiflora). Another turkey, also shot inDecember, had eaten a ground beetle, an ichneumon fly, 2 wheel bugs, 10 yellow-jackets, a meadow grasshopper, 75 red-legged grasshoppers, a few sour-gumberries, some pine seeds (with a few pine needles, probably taken accidentally),several acorns, a quarter of a cupful of wheat, and a little corn.Various other kinds of berries, fruits, and insects are doubtlesseaten when available, as turkeys will eat almost anything they canfind in these lines.Behavio7\?The turkey's ordinary method of locomotion is walkingor running; the long powerful legs enable these birds to travel longdistances and very rapidly on foot. But they are also strong flierswhen hard pressed or when necessity requires it, and can fly for aconsiderable distance or even across wide rivers. What few turkeysI have seen in flight looked to me like huge ruffed grouse, with longtails spread and heavy wings beating rapidly, though the speed ofthese large heavy birds is proportionately much less. Audubon(1840) says:Their usual mode of progression is what is termed walking, during whichthey frequently open each wing partially and successively, replacing themagain by folding them over each other, as if their weight were too great.Then, as if to amuse themselves, they will run a few steps, open both wingsand fan their sides, in the manner of the common fowl, and often take two orthree leaps in the air and shake themselves. During melting snowfalls, theywill travel to an extraordinary distance and are then followed in vain, it beingimpossible for hunters of any description to keep up with them. They havethen a dangling and straggling way of running, which, awkward as it mayseem, enables them to outstrip any other animal. I have often, when on agood horse, been obliged to abandon the attempt to put them up, after followingthem for several hours.While traveling about during fall and winter the sexes gatherinto separate flocks, the females forming the largest flocks; youngmales also flock by themselves and, for the most part, keep awayfrom the old gobblers. When flocks of old and young males happento meet they do not ordinarily quarrel; but they seem to have dif-ferent interests.What few turkeys still survive, in regions where they are muchhunted, have developed a high degree of shrewdness and cunning.An instance of cunning is given by Dr. J. M. Wheaton (1882) asfollows
:
As if aware that their safety depended on their preserving an incognitowhen observed, they effect the unconcern of their tame relatives so long as athreatened danger is passive or unavoidable. I have known them to remainquietly perched upon a fence while a team passed by; and one occasion knewa couple of hunters to be so confused by the actions of a flock of five, which
336 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMdeliberately walked in front of tbem, mounted a fence, and disappeared leisure-ly over a low hill before tbey were able to decide them to be wild. No soonerwere they out of sight, than they took to their legs and then to their wings,soon placing a wide valley between them and their now amazed and mortifiedpursuers.Wild turkeys have a preference for roosting over water, and theywill often go a long way in order to obtain such a roost. The back-water from the overflowing streams when it spreads out widelythrough the standing timber of the river bottoms, affords them greatcomfort; also the cypress ponds to be found in our southern riverdistricts. They evidently fancy that there is greater safety in suchplaces.Voice.?The wild turkey has quite a vocabulary, according to E. A.Mcllhenny (1914), a language with various meanings. If the strut-ting gobbler thinks he has heard the cluck and yelp of a callinghen, cluck, cluck, keow, keow, keow, he drops his broad wings, partlyspreads his tail, and listens; then vut-v-r-r-o-o-o-m-m-i comes thebooming strut, and gil-ooole-obole-ooole. Then let the hen give herlow quavering yelp, keow-keow, keow, and he will yell out in a fierceand prolonged rattle. More calls from the hen, keow, keow, kee,kee, or cluck, keow, ku-ku, one interspersed with loud gobblings, untilthe siren call of the hen, cut-o-r-r-r, cut, cut, keow, keow, keow, indi-cates that she has gone to him and all is quiet. Should any threat-ening danger intrude on this pretty love scene, a warning note isgiven, cluck, put, put, or put, o-r-r-r-r, or perhaps the turkeys walkquietly away saying, quit, quit, in irritated alarm.Enemies.?Although the eggs and young are preyed upon bymany predatory animals and birds, it is only the larger species thatare strong enough to attack an adult turkey. Audubon (1840)writes
:
Of the numerous enemies of the Wild Turkey, the most formidable, exceptingman, are the Lynx, the Snowy Owl, and the Virginian Owl. The Lynx suckstheir eggs and is extremely expert at seizing both young and old, which heeffects in the following manner. When he has discovered a flock of turkeyshe follows them at a distance for some time, until he ascertains the direction inwhich they are proceeding. He then makes a rapid circular movement, getsin advance of the flock, and lays himself down in ambush until the birdscome up, when he springs upon one of them by a single bound and secures it.While once sitting in the woods on the banks of the Wabash, I observed twolarge Turkey-cocks on a log by the river, pluming and picking themselves. Iwatched their movements for awhile, when of a sudden one of them flewacross the river, while I perceived the other struggling under the grasp of aLynx.Game.?It is probably safe to say that the wild turkey is the largestand grandest game bird in the world, certainly in North America. Itis not so well known and not so popular as the quail or ruffed grouse,because comparatively few sportsmen have had an opportunity to
EASTERN TURKEY 337hunt it, on account of its growing scarcity and the remoteness of itshaunts. What few turkeys remain within easy reach of civilizationhave become so highly educated that* it requires considerable ex-perience and skill to outwit them. Their eyes can not easily rec-ognize a stationary object, but they are very quick to detect theslightest movement. Their sense of hearing is very acute, and theyare always on the alert for approaching enemies, especially humanbeings. As a food bird the turkey is unsurpassed both in quantityand quality.The methods employed in hunting turkeys are, or have been, manyand varied. An interesting method of capturing turkeys, in thedays when they were plentiful and unsuspicious, was thus describedby John Hunter in 1824, as quoted by Albert H. Wright (1914) inhis excellent history of this bird
:
The turkey is not valued, though when fat, the Indians frequently take themalive in the following manner. Having prepared from the skin an apt resem-blance of the living bird, they follow the turkey trails or haunts till theydiscover a flock, when they secrete themselves behind a log in such a manneras to elude discovery, partially displaying their decoy, and imitate the gob-bling noise of the cock. This management generally succeeds to draw off firstone and then another from their companions which, from their social andunsuspecting habits, thus successively place themselves literally in the handsof the hunters, who quickly despatch them and await for the arrival of more.This species of hunting, with fishing, is more practiced by the boys than theolder Indians, who seldom, in fact, undertake them unless closely pressed byhunger.A common method of capture, referred to by many writers, wasto trap them in an inclosure, or pen, made of logs. The top wascovered with logs, leaving narrow open spaces between them. Atrench was dug, sloping gradually down, under the log wall and upinto the pen. Corn or other grain was sprinkled along this trenchand plenty of it spread on the inside of the pen to tempt the turkeysto enter. When, after eating all they wanted, they attempted toescape, they constantly looked upward for an opening but seldom, ifever, had sense enough to crawl out the way they had come in. Largenumbers were caught in this way.Audubon (1840) says that as many as 18 turkeys have been caughtin a pen at one time, and as many as 76 within a period of twomonths.One of the most popular methods, which is still widely practiced,is calling the gobbler by imitating the call of the hen during themating season. This requires the utmost skill, experience, practice,and thorough knowledge of the habits and haunts of the birds.Much has been written in various books and numerous articles insporting magazines on how to succeed in this. The instruments usedin calling may be simply the leaf of a tree held between the lips,
338 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthe box or trough call, the splinter and slate, or a new clay pipe;but the commonest and most effective call is made from the wingbone of a hen turkey. The hunter must know how to use theseperfectly, for a false note will drive the turkey away, perhaps neverto return. He must also be able to keep perfectly still for a longtime, with his gun, or rifle, trained on the spot where he expects theturkey to appear, for the slightest visible movement would spoil hischance. He would better be well concealed, but success may be had,even if he is in plain sight, if seated against a stump or tree largeenough to conceal the outline of his body. As to the use of the calls,he had better study the various seductive notes of the hen, theturkey language, or, better still, learn them from an experiencedhunter.Tracking turkeys in the snow on a clear cold winter day is splen-did sport. It has been well described by Edwyn Sandys (1904).In following a flock of turkeys a single track may turn off to oneside ; this means a tired bird, which will soon crouch to rest. If hecarries a shotgun, the hunter should follow this bird, for he willsoon flush it and get a flying shot. But, if carrying a rifle, he shouldfollow the main flock ; sooner or later he will get a long shot at someof them, though it may be a long chase unless the snow is soft anddeep. Should the birds take wing they will fly in a straight line,indicated by the direction of the long steps taken in rising, andthe trail can be taken up again.Coursing turkeys with greyhounds, as practiced in the more openwestern country is exciting sport. It is also vividly described bySandys (1904). The hunter on horseback, accompanied by a goodgreyhound, finds his turkeys feeding out on an open plain and triesto flush one headed for the open. The turkey's first flight is hislongest, hotly pursued by dog, horse, and man. If the bird comesdown and tries to run, he is soon overtaken. His flights and runsgradually grow shorter and shorter, until he becomes exhausted andis caught.Well-trained turkey dogs are useful in chasing winged birds,which a man could never catch. Audubon (1840) says:Good dogs scent the turkeys when in large flocks at a great distance; Imay venture to say half a mile away, if the wind is right. Should the dogbe well trained to the sport, he will set off at full speed on getting the scentand in silence until he sees the birds, when he instantly barks, and, runningamong them, forces the whole flock to take to the trees in different directions.This is of great advantage to the hunter, for, should all the turkeys go oneway, they would soon leave the perches and run again; but when they areseparated by the dog, a person accustomed to the sport finds the birds easilyand shoots them at pleasure.
EASTEBN TURKEY 339Fall.?Turkeys are not migratory, in the strict sense of the word,but they are much given to extensive wanderings, mainly in the falland winter, in search of food, which varies in abundance from oneseason to another. Audubon (1840) writes:About the beginning of October, when scarcely any of tlie seeds and fruitshave yet fallen from the trees, these birds assemble in flocks, and graduallymove towards the rich bottom lands of the Ohio and Mississippi. The males,or, as they are more commonly called, the gobblers, associate in parties from10 to 100, and search for food apart from the females ; while the latter are seeneither advancing singly, each with its brood of young, then about two-thirdsgrown, or in connexion with other families, forming parties often amountingto 70 or 80 individuals, all intent on shunning the old cocks, which, even whenthe young birds have attained this size, will fight with and often destroy themby repeated blows on the head. Old and young, however, all move in the samecourse, and on foot, unless their progress be interrupted by a river, or thehunter's dog force them to take wing. When they come upon a river, theybetake themselves to the highest eminences, and there often remain a wholeday, or sometimes two, as if for the purpose of consultation. During thistime the males are heard gobbling, calling, and making much ado, and are seenstrutting about, as if to raise their courage to a pitch befitting the emergency.Even the females and young assume something of the same pompous demeanour,spread out their tails, and run around each other, purring loudly, and per-forming extravagant leaps. At length, when the weather appears settled, andall around is quiet, the whole party mounts to the tops of the highest trees,whence, at a signal, consisting of a single cluck given by a leader, the flocktakes flight for the opposite shore. The old and fat birds easily get over, evenshould the river be a mile in breadth ; but the young and less robust frequentlyfall into the water, not to be drowned, however, as might be imagined. Theybring their wings close to their body, spread out their tail as a support,stretch forward their neck, and, striking out their legs with great vigour, pro-ceed rapidly towards the shore; on approaching which, should they find it toosteep for landing, they cease their exertions for a few moments, float down thestream until they come to an accessible part, and by a violent effort generallyextricate themselves from the water. It is remarkable, that immediatelyafter thus crossing a large stream, they ramble about for some time, as ifbewildered. In this state, they fall an easy prey to the hunter.Winter.?During winter, when the snow is too deep or soft totravel on the ground, turkeys often remain in the trees for longperiods, subsisting on buds and what fruits, nuts, and berries theycan find above the snow. They are great travelers, however, in lightor on hard snow. When hard pressed for food they sometimes ven-ture into farmyards or grain fields, or along roadsides or railroadtracks where grain has been spilled. At such times they can beeasily baited by scattering corn in such places.
340 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMMELEAGRIS GALLOPAVO OSCEOLA ScottFLORIDA TURKEYHABITSThe Florida wild turkey, which is resident in the southern half ofFlorida, was described by W. E. D. Scott (1890) and named forOsceola, a famous chief of the Seminole Indians. Scott says that itis similar to the northern wild turkey,but perceptibly darker in general tone. Coloring of tail and upper tail-covertssimilar in both forms. The white on the primary and outer secondary quillsrestricted, and the dark color (brownish black) predominating, the white beingpresent only as detached, narrow, broken bars not reaching the shaft of thefeather. The inner secondaries of a generally dirty grayish brown withoutapparent bars, but with brownish vermiculations on the inner web.Referring later to the Caloosahatchie region, he (1892) writes:This is still a very abundant bird in this part of Florida, though said to bediminishing in numbers every year and to be not nearly so plentiful as it wasten or fifteen years ago. During my stay at Fort Myers from November tillMarch, the open season, the birds were constantly offered for sale in the mar-kets, the price being on the average ten cents a pound for dressed birds. A henturkey could generally be bought for from seventy-five cents to one dollar anda gobbler for from one dollar to a dollar and a half. Only a few years backthe regular price paid to the hunters was twenty-five cents each. This I wastold by many reliable people who had lived there a dozen years or more.It would seem that these birds, living as they do at this point in cypressswamps and " bay heads," have a natural protection that will not allow oftheir absolute extermination, but, unless the exceedingly good laws passed bythe last legislature of the State are carefully enforced, the Wild Turkey, stillvery abundant in this region, is doomed to become in a few years as rare asit has already become in the northern part of Florida.Dr. William L. Ralph, in a letter to Major Bendire (1892), states:Fifteen years ago I found the Wild Turkey abundant in most parts of Florida,north of Lake Okeechobee, with perhaps the exception of the Indian Riverregion, but they have gradually decreased in numbers since then, and thoughstill common in places where the country is wild and unsettled, they arerapidly disappearing from those parts, in the vicinity of villages and navigablewaters.One can hardly believe that the Wild Turkeys of to-day are of the samespecies as those of fifteen or twenty years ago. Then they were rather stupidbirds, which it did not require much skill to shoot, but now I do not know of agame bird or mammal more alert or more difficult to approach. Formerly, Ihave often, as they were sitting in trees on the banks of some stream, passedvery near them, both in rowboats and in steamers, without causing them tofly, and I once, with a party of friends, ran a small steamer within 20 yardsof a flock, which did not take wing until several shots had been fired at them.Turkeys are still fairly common in the more remote regions ofFlorida, or where no hunting is allowed, especially around the edgesof the larger cypress swamps, such as the " big cypress " in Collier
FLORIDA TURKEY 341County. There, in 1930, 1 saw a small band of them on the outskirtsof a protected citrus plantation; and one day I saw one cross theTamiami Trail from one tract of pine woods to another; shootingis prohibited for a mile on each side of this road. We often sawtheir tracks around the borders of the pine woods and open savannas,near the cypress swamps ; they feed in such places and roost at nightin the large cypresses.Courtship.?Doctor Ralph says further:These birds are polygamous, and the female takes all the cares and dutiesof incubation upon herself. The gobblers are very pugnacious, and will oftenfight fiercely for the favors of the hens. The love season begins in Floridaabout the middle of February and lasts for about three months, and duringthis period the gobblers frequently utter their call and are then easily decoyedwithin gunshot. Native hunters have informed me that the hens roost bythemselves at this season of the year.Nesting.?On this subject Ralph writes:The nest is a slight depression in the ground, either at the foot of a treeor under a thick bush or saw palmetto. It is lined sparingly with deadleaves and grass, etc., but I could never find out whether this material wasplaced there by the birds or was there originally. I think these birds raisebut one brood a season, though I have found fresh eggs as early as the middleof March and as late as the 1st of May. I have never found more than thirteeneggs in one nest, nor less than eight, unless they were fresh, the usual numberbeing ten. The chicks of this species are very tender, and as they follow theirmothers as soon as hatched I have often wondered how the latter could raiseso many as they do. The natives of Florida say that a hen Turkey will deserther nest if the eggs are handled. Whether this be true or not I do not know,for I never tried to find out but once, and then, though the bird was gone onmy second visit to the nest, I always had a strong suspicion that she was shot,for its whereabouts was known to several persons besides myself.I have a set of 10 eggs in my collection that was taken on March28, 190S, near Everglade; the nest was a hollow in the ground undera saw palmetto, near the Big Cypress. We found a nest, fromwhich the young had hatched, on April 19, 1902, on the border ofJane Green Swamps, Brevard County. It was a mere hollow in thesand, lined with strips of palmetto leaves, under a small cabbagepalmetto; it was well shaded but not particularly well hidden, andcontained the broken shells of nine eggs.Eggs.?The eggs are similar to those of other wild turkeys. Themeasurements of 56 eggs average 61 by 46.3 millimeters; the eggsshowing the four extremes measure 66 by 46.7, 62.5 by 48.8, 56.3by 46.4, and 65.2 by 41 millimeters.Young.?F. M. Phelps (1914) relates the following experience:Late on the afternoon of April 18th, as we were working along an open gladebordering a cypress swamp, the dog began to nose excitedly in the grass. Sud-denly up popped half a dozen little brown cannon-balls, quail I thought, butwhen they alighted in some cypress saplings I saw at once they were young74564?32 23
342 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMTurkeys. The old hen, hard pressed, soon rose from the grass and sailed awayacross the tops of the cypress trees. More youngsters kept popping up untilthere were eleven sitting about in the saplings some twelve or fifteen feet up.Soon one gave a peculiar little quit, and then to my utter astonishment flewstraight away over the tops of the cypress trees after the old hen, and one byone the rest followed. My guide pronounced them to be about two weeks oldand that seemed to me about correct.The ability of very small young to fly is also attested by DonaldJ. Nicholson (1928), who writes:On May 3, my brother Wray and I were going thru the pine and cypresscountry just east of Turner's River about one mile, and just about noon wecame upon a turkey hen with five or six little ones not quite as large as afull-grown Bobwhite. We stood and watched her for a few seconds, and sheran slowly thru the scattered low palmettos with the young scampering along.We conceived the idea that it might be possible to catch one of the youngsters,and began to give chase, but immediately they all rose and flew with strongflight, alighting in the lower limbs of a thirty-foot pine tree. We managedto find two of the young perched on dead branches not far apart, peeringdown upon us ; they did not offer to fly or show any restlessness.Behavior.?Again Nicholson says:One day we were driving along the Tamiami Trail not far from where weencountered the young turkeys, and saw five large gobblers feeding right outin the open, 400 yards from the road. It was a sort of prairie or savanna,among the stands of cypress, and had been burned over ; short grass had grownup. The birds paid not the slightest heed to us and we sat and watched themfor ten or twenty minutes. However, one bird would stand with head and neckerect ; as if on guard while the others fed ; then another would take its place.MELEAGRIS GALLOPAVO INTERMEDIA SennettRIO GRANDE TURKEYHABITSWhen George B. Sennett (1879) first called attention to the char-acters in which the Rio Grande turkey differs from the other racesof wild turkeys, he evidently thought it was an intermediate andshould not be named, for he said, at that time: "All Lower RioGrande specimens, therefore, must be held as gallopavo (the Mexicanform), or a var. intermedia established?an alternative not to bedesired." Later on, however, he (1892) described and named itellioti, in honor of Dr. Daniel G. Elliot. But his earlier name,intermedia, must stand under the law of priority. He says that it
* * * can be distinguished from the other forms by its dark buff edgingson tail and upper and lower tail-coverts, in contrast with the white color onthe same parts of mexicana, and the deep, dark, reddish chestnut of the sameparts in M. gallopavo, the eastern United States bird. The lower back is adeep blue-black and is wanting in those brilliant metallic tints so prevalentin the eastern bird and in the type of mexicana. The primaries of the wing are
BIO GRANDE TURKEY 343black with white bars in contrast with M. gallopavo the primaries of whichare white with black bars.The range or habitat of this race, so far as known at the present time, isrestricted to the lowlands of eastern Mexico and southern Texas. It willprobably not be found south of Vera Cruz, nor is it likely to be met with tothe north beyond the Brazos River of Texas, its range being thus restrictedwithin about ten degrees of latitude. Wherever timber and food are in abun-dance we find this new form common to the coast and lowlands, and we couldnot expect to find it at an altitude exceeding 2,000 feet above sea-level ; whilethe variety mexicana is found only at the higher altitudes from 3,000 to 10,000feet above the sea.The Rio Grande turkey is now known to have been quite widelydistributed in Texas from the central-northern part southward andwestward into northern Mexico, though it has become much scarcerexcept in the wilder portions of the State. Writing of this turkey inKerr County, Tex., Howard Lacey (1911) says:Formerly very common, but getting rather scarce now that the shotgun isbecoming almost as common a piece of furniture as the rifle in the ranchman'shouse. These birds are so foolishly tame when about half grown as they arewild and able to take care of themselves when fully mature; if they were notshot at until fully grown and allowed to roost in peace at night, there is noreason why we should not have them always with us. Armadillos and skunkssometimes roll the eggs out of the nests, and they have plenty of enemiesbesides the boy with the shotgun.Austin P. Smith (1916), referring to the same general region,writes
:
There can be little doubt that, at the present time, Wild Turkeys exist ingreater numbers in Kerr and adjoining counties than in any other part ofTexas. Their abundance may be accounted for, as the result of the encroach-ment of the Cedar and various species of scrubby oaks upon lands formerlyunder cultivation or in pasture; to the decrease in numbers of the Armadillo(Tatu novemcinetum texanum) which of late years have been much hunted forcommercial purposes ; and to the enactment of a law limiting the open seasonand the number that may be killed. During the winter spent in the regionseveral heavy snowfalls occurred. These caused many turkeys to seek openspots in the valleys and along fence rows, often in the vicinity of human habi-tations, and I recall one flock of seven hunting for several hours within a hun-dred feet of the building I lived in.George F. Simmons (1925) says its haunts in the Austin regionare " wild, rough, brushy, country ; dry, big-timbered arroyos run-ning back from watered creeks; hill and valley country; shin oakclumps on hillsides; creek bottoms and lower slopes; wild, less-fre-quented, thinly settled country, particularly in the mountains andnotches in the hills."Nesting.?Simmons describes the nest as a " slight hollow, scrapedout by the bird, lined with grasses and leaves, among low bushes, indense woods along streams, in tangles of briar vines, and in thick
344 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
weed}- places. Very difficult to find, particularly when placed ingrowths of underbrush."Eggs.?The eggs are similar to those of the other wild turkeys.The measurements of 49 eggs average 62.4 by 46.5 millimeters; theeggs showing the four extremes measure 64.8 by 43.2, 64 by 48.6, 57.2by 43.6, and 61.7 by 43.2 millimeters.Food.?Simmons (1925) says of its feeding habits:When pecans are ripe, the birds feed under the pecan trees along the valleys.At other seasons, they wander about wooded slopes in the daytime, feedingamong the cedars and scrub oaks which cover the hillsides and ridges of theland that was once prairie ; at night, they return to the valleys to roost. Feedon nuts, acorns, seeds, grain, berries, plant tops, insects, crickets, and grass-hoppers.Sennett (1879) says that in April "their principal food was thewild tomato, which attains about the size of a cranberry, and whichthey devoured whole, together with insects and larvae."Behavior.?Simmons (1925) writes:Observed singly or in pairs during the breeding season, at other times inflocks of from 12 to 15 or more ; flocks are usually practically all of one sex orthe other. Very wary ; when in danger, it usually sneaks away or runsthrough the underbrush and into thickets, preferring to trust to its stoutlegs rather than taking wing. Usually roost each night in the same locality,birds returning singly and by twos and threes at dusk, until all the birds haveassembled in their favorite places in the tops of larger, taller trees, generallyover water and frequently in partially submerged trees, possibly for protectionagainst prowling coyotes and bob-cats ; big pecan trees along wooded creekvalleys, in washes, and in bottomlands are generally selected. Birds make theirway back to the higher ridges before daylight has half arrived. Males flocktogether during the period in which females are kept busy with eggs or care ofyoung.Vernon Bailey, in Mrs. Bailey's book (1902), gives the followingaccount of the habits of the Rio Grande turkey
:
Over most of the country where the wild turkeys were once plenty theyhave now become scarce or extinct, but in a few places may still be found insomething like their original abundance, living much as their ancestors lived,breeding unmolested, strolling through the woods in flocks, and gathering atnight in goodly numbers in their favorite roosting places. Perhaps the bestof these undevastated regions are on the big stock ranches of southern Texas,where the birds are protected not by loosely formed and unenforced gamelaws, but by the care of owners of large ranches, who would as soon think ofexterminating their herds of cattle as of shooting more than the normal in-crease of game under their control. Here, at least through the breedingseason, the turkeys are not more wary than many of the other large birds,and as we surprised them in the half open mesquite woods along the NuecesRiver, would rarely fly, merely sneaking into the thickets, or at most runningfrom us. The ranchmen say that the turkeys always select trees over waterto roost in when possible, and no doubt they do it for protection in this regionwhere foxes, coyotes, and wildcats abound. On the edge of the flooded bottomsof the Nueces River they roosted in the partially submerged huisache trees.
CHACHALACA 345A loud gobble just at dusk led us to tbeir cover, and crouching low to get thesky for a background we could see the big forms coming in singly or in twosor threes, and hear the strong wing beats as they passed on to alight in thehuisaches out in the water. When the noise of their wings and the rattling ofbranches had subsided, with a few gobbles from different quarters they settleddown for the night. The next morning, as the darkness began to thin anda light streak appeared in the east, a long loud gobble broke the stillness,followed by gobble after gobble from awakening birds in different parts of thebottoms, and before it was half daylight the heavy whish whish of big wingspassed overhead, as the turkeys with strong, rapid flight took their way backto the higher ridges.Family CRACIDAE, Curassows, Guans, ChachalacasORTALIS VETULA VETULA (Wagler)CHACHALACAHABITSThis curious and exceedingly interesting bird, the chachalaca,brings a touch of Central American bird life into extreme southernTexas in the lower valley of the Rio Grande, where so many otherMexican species reach the northern limits of their ranges and wherethe fauna and flora are more nearly Mexican than North American.On May 27, 1923, 1 spent a good long day, from before sunrise untilafter sunset, in the haunts of the chachalaca, wTith R. D. Camp,George F. Simmons, and E. W. Farmer, the last named a chachalacahunter of many years' experience, who knows more about this curiousbird than any man I have ever met. The locality to which heguided us was the famous Resaca de la Palm a, where so many otherobservers have made the acquaintance of the chachalaca, only a fewmiles outside of the city of Brownsville, Tex. This and other resacasin the vicinity are the remains of old river beds of the Rio Grande,which from time to time in the past has overflowed its banks orchanged its course, cutting these winding channels through the wild,open country, chaparral, or forest. Some of these channels were dryor nearly so, but most of them contained more or less water belowtheir gently sloping grassy borders. Above the banks were denseforests of large trees, huisache, ebony, hackberry, and mesquite, witha thick undergrowth of thorny shrubbery, tangles of vines, and anoccasional palmetto or palm tree. In other places almost impene-trable thickets of chaparral lined the banks, with its forbiddingtangle of thorny shrubs of various kinds, numerous cactuses andyuccas. These forests and thickets were teeming with bird life.Along the edges of the watercourses the pretty little Texas king-fishers were seen flying over the water or perched on some deadsnag. In some small trees overgrown with Usnea moss the daintylittle Sennett's warblers were flitting about, reminding me of our
346 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMnorthern parulas. Handsome green jays were sneaking about inthe larger trees, surprisingly inconspicuous in spite of their gaudycolors. Brilliant Derby flycatchers proclaimed their noisy presencein loud, clamorous notes from the tree tops. Sennett's thrashersscolded us in the thickets, and the confiding little Texas sparrowshopped about on the ground near us, scratching among the deadleaves. Many other birds were seen, but the most conspicuous ofall were the doves; the woods and the thickets almost constantlyresounded with the deep-toned notes of the wThite-fronted, the tire-some who cooks for you of the white-winged, and the soft cooing ofthe mourning dove. Such is the home of the chachalaca with someof its neighbors.As we entered the chaparral before sunrise we heard the warningcry of the chachalaca on all sides ; the woods fairly resounded withits cries, some of which sounded like a watchman's rattle, morewooden than metallic in quality; the birds were very shy and seldomseen; occasionally we saw one, perched on some small tree top andgiving its challenge or battle cry; but as soon as it realized that itwas observed, it would sail down into the thicket and keep still.Much of the following information is taken from some very fullnotes obtained from Mr. Farmer. He promised to send me somenotes, but unfortunately he has now gone to the " happy huntinggrounds." My friend Frederic H. Kennard was more successful andhas very kindly placed these notes at my disposal. According to Mr.Farmer's personal knowledge, the chachalaca occurs in Cameron,Hidalgo, Starr, Zapata, Willacy, and Kenedy Counties in Texas; thebirds are never permanently located more than a mile from water;if the water dries up all about, they move; otherwise they stick toone place throughout the year.Courtship.?According to Mr. Farmer, courtship begins aboutMarch 20 in ordinary seasons, with the chachalac challenge calls ofthe male, perched in the tops of the highest trees in the chaparral;other males answer from every direction in competition, each tryingto " outholler " the others. The females can make a similar call, butit is on a higher key and less in volume. The concert begins at aboutsunrise or a little before. The male's call to the female is like thechallenge, but it is less harsh and ends with a soft note. The femalesmay climb up into the tree beneath the male, but in a less conspic-uous place, generally keeping under cover and answering the malein their own way chattering, talking, and scolding. After the malehas " hollered himself out " in the tree, he comes down to the groundand devotes himself to the females, walking about and strutting withhead erect and making a low call hardly to be heard a short distanceaway. If another male appears he is promptly chased off. In Mr.
CHACHALACA 347Farmer's experience with the birds there have always seemed to betwo females to one male in this courtship. The male seems strictlyimpartial. Mr. Farmer has had much experience in rearing youngchachalacas, and says that, so far as he has observed, they alwaysseem to hatch out in the ratio of two females to one male. He alsohas, many a time, watched the courtship performance of the malewith two females, no other male being tolerated in the vicinity.He says that the males fight a great deal at this season. Theyhave no spurs, but fight with bills, feet, and wings, jumping overone another and pecking at one another's backs rather than at theheads, their wing strokes, however, being directed at the head. Atthis season of the year the males frequently appear with most ofthe feathers pecked off their backs.Mr. Kennard writes to me as follows:It is a matter of common report on both sides of the Rio Grande that thechachalacas are used for crossing with game chickens for fighting purposes, theresulting cross being much quicker on its feet than the ordinary game fowl.These reports have, however, never been actually verified by either Mr. Campor Mr. Farmer and are to be doubted.A letter from Mr. Camp confirming this states:I do not agree with any of the statements concerning the crossing of thechachalaca with the domestic fowl. I have traveled hundreds of miles andinvestigated dozens of cases both on this and the other side of the Rio Grande,endeavoring to verify reported hybrids, and at no time have I found a specimenthat I would acknowledge was a cross. Last year I investigated quite exten-sively among the natives in the district 125 miles from here, where the cha-chalacas are very abundant and tame for wild birds. Most all Mexican coloniesin the district had semidomesticated chachalacas running with their barnyardfowl, but none of the natives would acknowledge that he had ever seen a hybrid.Nesting.?The only two nests of the chachalaca that I ever saw werefound near the Kesaca de la Palma referred to. On May 27, 1923,they both held sets of three eggs each, heavily incubated. Mr. Sim-mons found the first one; he had been standing under the tree forsome time, when he heard the bird fly off from the thick foliage overhis head and found the nest Sy2 feet above the ground. I found theother by seeing the bird fly off, and I had to climb up into the verytops of several slender trees to reach the nest, which was about 18feet up and well concealed in the leafy tops. Both nests were verysmall, frail structures, made of sticks and leaves and lined with a fewgreen leaves; they were barely large enough to hold the eggs.According to Mr. Farmer, nest building is started soon aftercourtship has begun. He has seen two birds, probably the male andone female, at work on the nest, while the other female was sittingabout near by, perhaps helping or perhaps only watching. Thenest is a scraggly but strongly built structure of short, stout twigs,so well interlaced as to stand a lot of handling; a particularly well-
348 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMpreserved previous year's nest measured about 24 inches across and10 inches deep. The nest is usually built in an ebony tree, but some-times in a mesquite or other tree, and usually between 5 and 15 feetabove the ground, occasionally as high as 25 feet or as low as 2 feetbut never in a hollow tree. He says that he has seen at least 1,000nests, and that the nest is usually near the edge of the chaparral andnear a resaca, never more than 200 yards from water and alwaysnear a supply of the berries on which the birds feed their young.The nest is generally built out on a limb, but sometimes in a crotchor where the limb of a tree is interlaced with vines.There are two sets of eggs in Col. John E. Thayer's collection,said to have been taken from nests on the ground ; one was " in acane brake, composed of grass, weeds, and other litter," and theother was "on the ground among heavy grass." These were takenfor Thomas H. Jackson, probably by Frank B. Armstrong or hisMexican collectors, who were known to be careless in the make-upof sets and data. Messrs. Farmer and Camp do not mention anyground nests, and George B. Sennett (1878) says:The nest of this species is never found on the ground, but in trees andbushes varying in height from four to ten feet. The structure varies in compo-sition and size according to its location. If it is in a large fork close to thebody of the tree, a few sticks, grasses, and leaves are sufficient, and the struc-ture will not equal in size or strength that of a Mockingbird. This smallsize is by far the most frequent, but I have a nest built upon a fork of twosmall branches, composed entirely of Spanish moss. It is bulky and flat, beinga foot in diameter and four inches deep, with a depression four inches wide andtwo deep.Major Bendire (1892) quotes J. A. Singley as saying:All the nests I found were in mesquite stubs, where the limbs had been cutoff to make brush fences. These limbs are never cut close to the tree and,being close together, form a cavity ; leaves and twigs will fall in this andaccumulate, and the bird occupies it as a nesting site. I did not find a nestthat I could say was built by the bird. When the nest is approached the birdquietly flies off, rarely remaining in sight, and soon calls up its mate.Eggs.?Mr. Farmer says that in his experience the eggs have beeninvariably three in number, and most of the other observers saythree or rarely two. The larger sets in Mr. Jackson's collectionprobably came from Armstrong and may be made-up sets, thoughperhaps sets of four occasionally occur. The eggs of the chachalacaare ovate, short ovate, or elongate ovate. The shell is thick, tough,and roughly granulated. The color is pale creamy white or dullwhite. The measurements of 56 eggs average 58.4 by 40.9 milli-meters; the eggs showing the four extremes measure 65.5 by 47, 53.3by 40.6, and 58.2 by 37.6 millimeters.Young.?Mr. Farmer has hatched a number of eggs under hensand found that 22 days was the longest period of incubation. He
OHACHALACA 349
says that the " sitting bird often leaves the nest to go off and feed,"but he doubts whether the male helps in incubation. " The femalesits fairly close if one does not make too much fuss in approachingand does it quietly and indirectly, pretending to look for somethingelse. When she flies she ' eases ' off the nest quietly and disappearsinto the brush, going off and hiding somewhere where you can notsee her, but where she can see you, whence she ' quarrels and scolds
'
until you leave the vicinity. Often other birds will join in therumpus. If you have a dog along the birds will be much morenoisy, objecting to the dog even more than they do to you."The young are precocial, leaving the nest just as soon as the downis dry. The female carries the young down to the ground clingingto her legs, one at a time, according to Mr. Farmer's observation.This particularly peculiar habit of the young clinging to its moth-er's legs has been verified time and again by both Mr. and Mrs.Farmer when lifting the mother hen off the nest to inspect thebrood on which she was sitting, and sometimes two at a time cling-ing to the mother. Once in a drizzling rain Mr. Farmer heard achachalaca chatter and went to investigate. He found a femalewith young, and, while he watched, saw her carry, one at a time, allthree young up into an ebony tree and leave them perched in aline on a limb about 15 feet above the ground. When about 2 weeksold they can fly perhaps 100 feet, but when they are a week old theycan flutter 8 or 10 feet, and even at this age they are almost im-possible to catch, flitting from bush to bush among the underbrushas they do. Mr. Farmer believes that there is only one brood. Hehas found young birds as late as September or October, and thinksthey are the result of the first nest being disturbed.Plumages.?In the downy young chachalaca the center of thecrown and the occiput are black, tinged with " russet," and there isan isolated black spot on the forehead; the sides of the head andneck are " cinnamon-buff," tinged with " cinnamon " on the neck,and finely mottled with black ; the chin and throat and lower under-pays are white, with a broad band of " cinnamon-buff " across thechest ; the upperparts are mottled with sepia and " cinnamon-buff."Another specimen is similar, but the sides of the crown are " palemouse gray," and the back is tinged with " russet " in the centralblack stripe and with " ochraceous-tawny " on the mantle. In bothchicks, one of which was known to be only 4 days old, the wings arewell started and already reach beyond the tail. The wings and tailin another young bird, about 9 inches long, are so well developed thatit could probably fly.The wings and tail of the juvenal plumage are the first to appear,and they grow so rapidly that the flight stage is reached at an earlyage. These and the upper parts in general are " Saccardo's um-
350 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMber " ; the center of the back and rump are barred with " tawny "
;
the wing coverts are barred with " cinnamon-bufi' "; the remiges aretipped, and mottled on the outer edge, and the rectrices are tippedwith "cinnamon-bull"; the rectrices are decidedly pointed. In the9-inch specimen referred to above, the head, neck, center of breast,and belly are still downy, evidently the last parts to be feathered;the juvenal plumage coming in on the sides of the breast is " Sac-cardo's umber," while that on the flanks and belly is " cinnamon-buff "; these two colors are sharply defined in the juvenal plumageand not intergraded or blended, as in the adult.Apparently the juvenal plumage is worn only a very short time,and a complete molt soon produces a practically adult plumage. Icould find no traces of juvenal feathers in fully grown fall or winterbirds. I have seen evidences of a complete molt in adults in Augustand September. Mr. Farmer says that they molt only once a year,in September and October. He also says that the naked places show-ing on each side of the chin are of a grayish flesh color, alike in maleand female, except during courtship in spring, when the male'spatches become red.Food.?Mr. Farmer says that the food of the chachalaca consistsprincipally of berries, though they do catch bugs, and sometimes inspring when the buds are tender they " bud " hackberry or othertrees. In captivity the tame birds eat bread and crackers or chopped-up meat, and they are especially fond of milk, particularly whenyoung, and will relish any kind of fruits, particularly apples andbananas. They are especially fond of raw, chopped-up rabbit.Behavior.?Mr. Kennard has seen chachalacas fly silently andswiftly over the tops of the chaparral, alight heavily in the top of atree, and hop down from limb to limb without opening the wings.Mr. Farmer once came upon a bunch of 9 or 10 chachalacas follow-ing an opossum, teasing it and attacking it, but paying no attentionto him even after he had killed some of them.Sennett (1878) writes:Several times, when well concealed, I have noticed a pair spring from athicket into a large tree, jump from limb to limb close to the body until theyreached the top, when they would walk out to the end of the branch and begintheir song. They roost in trees, and hunters frequently get them at night.Barely did I see them on the ground. Once, while resting in a mesquite grovewhich looked very much like a peach-orchard on a well-kept lawn, I saw aChachalaca trot out from a neighboring thicket in full view. He seemed lookingfor food on the ground. He discovered me and we eyed each other for amoment, when it turned, ran a short distance, sprang into the lower branches ofa tree, and hopping along from tree to tree disappeared into the thicket aboutfive feet from the ground.Voice.?The remarkable vocal performances of this species are itsmost interesting and striking habits. They are difficult to describe,
CHACHALACA 351but once heard they can never be forgotten. Dr. J. C. Merrill (1878)writes:During the day, unless rainy or cloudy, the birds are rarely seen or heard;but shortly before sunrise and sunset they mount to the topmost branch of adead tree and make the woods ring with their discordant notes. Contrary toalmost every description of their cry I have seen, it consists of three syllables,though occasionally a fourth is added. When one bird begins to cry the near-est bird joins in at the second note, and in this way the fourth syllable is made;but they keep such good time that it is often very difficult to satisfy one's selfthat this is the fact. I can not say certainly whether the female utters thiscry as well as the male, but there is a well-marked anatomical distinction in thesexes in regard to the development of the trachea. In the male this passesdown outside the pectoral muscles, beneath the skin, to within about one inch ofthe end of the sternum ; it then doubles on itself and passes up, still on theright of the keel, to descend within the thorax in the usual manner. Thisduplicature is wanting in the female.Sennett (1879) says:A more intimate acquaintance with this bird enables me to give a betterdescription of its notes than the attempt in my former memoir. The notes areloud and uttered in very rapid succession, and those of the female follow themale's so closely, while so well do they harmonize, although in different keys,that I mistook the first note of one for the last note of the other. It reallyutters but three syllables, thus : Cha-cha-lac, instead of four, cha-cha-lac-ca, asgiven before. It also has a hoarse, grating call or alarm note, uttered in onecontinuous strain and without modulation, something like kak-kak-kak.Dr. T. Gilbert Pearson told me that when he went out to camp onenight with Mr. Farmer in the heart of the chachalaca country, thelatter had told him that there would probably be within earshot oftheir camp at least 500 chachalacas, a statement about which he wasvery skeptical. About sundown the concert began, increasing involume until the din became almost indescribable. Doctor Pearsonwas convinced, and finally suggested to Mr. Farmer that he call it5,000 instead of 500 birds.Game.?The chachalaca has figured largely as a game bird in theBrownsville market. Its flesh is said to make delicious eating. Ithardly comes up to a sportsman's idea of what a game bird shouldbe, though one must have a thorough knowledge of its haunts andhabits to be successful in hunting it. It has been quite extensivelydomesticated on many Mexican ranches, lives contentedly with do-mestic poultry, and becomes very tame and makes a good pet.although often so familiar as to be troublesome.DISTRIBUTIONRange.?Lower Rio Grande Valley in Texas and northeasternMexico.The chachalaca occupies a limited range extending north to south-ern Texas (Rio Grande City, Fort Ringgold, Lomita Ranch, Hidalgo,
352 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMand Brownsville). East to Texas (Brownsville) ; eastern Tamauli-pas (Matamoras, Jimenez, and Aldama) ; and northeastern VeraCruz (Tampico). South to northeastern Vera Cruz (Tampico)
;
and southeastern San Luis Potosi (Valles). West to southeasternSan Luis Potosi (Valles) ; western Tamaulipas (Xicotencatl, Ciu-dad Victoria, and Rio Pilon) ; and southern Texas (Rio GrandeCity).Egg dates.?Texas and Mexico: 73 records, March 21 to August16; 37 records, April 27 to May 27.
Order COLUMBIFORMESFamily COLUMBIDAE, Pigeons, DovesCOLUMBA FASCIATA FASCIATA SayBAND-TAILED PIGEONHABITS
It was a bright, sunny, cold morning, after a frosty night inFebruary, when I first made the acquaintance of this fine bird, theband-tailed pigeon. The sun was shining full of genial warmth onthe tops of the tall sycamores and eucalyptus trees, which grew ina deep arroyo in southern California, but it had not yet penetratedto its shady depths, which still sparkled with white frost. I did notat first recognize the large plump birds, 33 of them I counted, perchedin the tops of two tall sycamores, evidently enjoying their morningsun bath. But with a glass I soon recognized them as pigeons, sawthe white crescent on the neck, and, as they flew, marked them asbandtails, from the bands on the broad square tails. This was oneof the wandering, restless flocks which travel about during thewinter, moving from one place to another as food or fancy leadsthem. Later I saw them flying down the arroyo in a detached flockhigh in the air. And almost daily for some time I saw more orless of them in the same arroyo on the outskirts of Pasadena. Theyremained in the vicinity off and on until the latter part of April.This is one of the birds that was being rapidly killed off, as it wasa favorite game bird in the Pacific Coast States. It was even vergingtoward extinction. But, fortunately, protection came in time to saveit and it has made a wonderful recovery. E. A. Kitchin writes to meof conditions in Washington
:
The Federal protection of these birds in recent years has made wondrouschanges in the Puget Sound country, which always was a natural breedingground. Before this protection the pigeons had become very scarce, so muchso that it was even an event to see one, and these were only seen in the moreisolated parts. The large gulches so numerous on Pnget Sound, covered atthe bottom by thick alder and on the sides by small firs, form the naturalbreeding grounds for these birds. Now, thanks to Government protection, thereis hardly a gulch that does not contain 50 or more pairs of breeding birds.Courtship.?The only note we have on the subject of the courtshipof the bandtail is the following by Harry S. Swarth (1904)
:
353
354 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMDuring the breeding season the male bird is fond of sitting in some elevatedposition, usually the top of a tall dead pine, giving utterance, at frequentintervals, to a loud coo, more like the note of an owl than a pigeon, which canbe heard at a considerable distance; while occasionally he launches himself intothe air with wings and tail stiffly outspread, describes a large circle backto his starting point, uttering meanwhile a peculiar wheezing noise impossibleof description. I had supposed that this noise was made by the outspreadwings, but a male bird which Mr. Howard had in his possession for some timegave utterance to the same sound whenever angered or excited, evidentlyby means of his vocal organs, as we had ample opportunity of observing.Nesting.?Mr. Kitchin, who has had considerable experience withthis pigeon in Washington, contributes the following
:
The flocks arrive the latter part of March or early in April and at once seekthe gulches, where they feed on the seeds of the alder. They apparently havea rather prolonged breeding season, lasting from April through June. Thenesting sites are mainly in the dark fir trees, where their nest of dead fir twigsis placed near the trunk and generally in the lower branches, averaging prob-ably 20 feet from the ground. Occasionally, however, the nest is found inan alder and sometimes on the top of a thick birch overhanging the hillside.When approached the brooding bird has a habit of standing erect in the centerof the nest and by doing so becomes very conspicuous. I have read of thesebirds carrying their egg when disturbed, and although I have flushed many birdsnone showed any inclination to take the egg with it. Knowing where anoccupied nest was, I have approached quietly to perhaps 6 feet before the birdflushed, and at other times I have rushed, in a startling way, taking her by sur-prise, but in neither case was the egg removed.The nest is somewhat loosely made and entirely of dead twigs and, thoughit is not in any way cupshaped it certainly is saucershaped, and the roughnessof the twigs prevents the egg from rolling in the nest. One can tell frombelow whether the nest contains an egg or a squab, as the brooding bird willstand in the center of the nest, astride the egg, while if a squab she will bestanding on the rim.The birds are fond of their old nesting sites and are insistent in using thesite selected. They not only come back to the same tree but will use the samelimb as that used the previous year, even if the first nest has been disturbed.On one occasion the bird selected a hanging bush on the hillside and built hernest near the top. The egg and nest were taken and she at once built anotheron the same site and raised her young. The following year she was again inthe bush, sitting on a slightly incubated egg. This set I took, and by July shehad another nest and egg, which were taken, and I was much surprised, inpassing later in the season, to find a third nest in which she had probably raisedher young to maturity.Grinnell, Bryant, and Storer (1918) give the most comprehensiveaccount available of the nesting habits of this bird in California,from which I quote as follows:Nearly all authentic reports from California agree in stating that the Band-tailed Pigeon nests in trees?almost invariably in black or golden oaks?atheights ranging from eight to thirty feet above the ground. As exceptions,Littlejohn (MS) found a nest in San Mateo County in a Douglas spruce; andin Marin County, J. Mailliard found a nest in a California lilac (Ceanothusthyrsiflorus) overhanging a steep slope. Some early reports from this State
BAND-TAILED PIGEON 355have mentioned ground nests, as have several more recent, but scarcely trust-worthy, accounts from Oregon and Washington ; but there is no late evidenceof the ground nesting habit in California. In a general way the nest resemblesthat of the Mourning Dove, save that it is considerably larger, and sometimesproportionately thicker. It is a crude structure, a mere pile of oak and othertwigs, so loosely arranged that attempts to remove the mass often result in itsfalling to pieces. The average diameter is six or eight inches, while the thick-ness in two recorded instances was one and four inches, respectively. Some-times as few as sixteen or eighteen twigs are all that go to make up the nestand again there may be more than a hundred. The twigs range from asixteenth to a quarter of an inch in diameter and are of various lengths. Theyare laid across one another, with little or no weaving, forming a platform withnumerous interstitial spaces. A slight lining of pine needles was found in onenest. As Gilman well says, it is a marvel how an egg can be kept warm enoughto hatch while resting on such an airy platform in the cool air of a high alti-tude. The nest site, which is almost always on top of a large horizontal limb,seems to be so selected that the incubating bird may flush directly and rapidlyfrom the nest when danger threatens.There is some evidence to indicate that the band-tailed pigeonoccasionally nests on the ground. Major Bendire (1892) quotesO. B. Johnson, referring to Oregon, as saying : " They nest in vari-ous situations, much like the common Dove, Z. carolinensis. I foundone of leaves and moss beside a tree, placed on the ground betweentwo roots; another one upon an old stump that had been split andbroken about 8 feet from the ground; another was in the top of afir (A. grandis), and was built of twigs laid upon the dense flatlimb of the tree, about 180 feet from the ground." This statement,he says, is confirmed by Doctor Cooper, as follows : " In June theylay two white eggs, about the size of those of the house pigeon, onthe ground, near streams or openings, and without constructingany nests." A similar statement is made by Mrs. Irene G. Wheelock(1904).Although the band-tailed pigeon usually nests in widely scatteredpairs, the following account of an Arizona colony, by F. H. Fowler(1903), is interesting:When the breeding season draws near, they betake themselves to shelteredplaces among the lower mountains, and nest in scattered communities, oras I have seen in several cases, a pair will nest apart from the others. Oneof the largest breeding communities I noted was in a little pocket in themountains, about five miles south of Fort Huachuca ; this little place was at thehead of a short canyon, and was indeed an ideal spot for birds, as it waswell wooded and watered. Here a flock of about thirty-five pairs of band-tailsnested in a scattered rookery, probably not averaging a nest to every three orfour acres at the most thickly populated part ; and a great majority of the nestswere even farther apart than this. The nests in this colony were all placedon the forks of low horizontal limbs of live oaks usually not more than twelvefeet up or less than nine, and in no case did I find more than one egg or squabin a nest. The nests were all of that very simple dove-like constructionconsisting of a few sticks placed on a fork of a branch.
356 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMFrancis C. Willard (1916) made the following observations in theHuachuca Mountains, Ariz.
:
There were a few pigeons nesting in the vicinity, and one pair near campwas watched quite closely from the time the nest was begun until the egg waslaid. Nest building was carried on only in the early morning hours, from sun-rise till about S o'clock. Both birds were present, but the female alone seemedto be engaged in the actual construction of the nest, which she went about ina very lackadaisical manner. The pair would sit together on the few sticksalready in place for many minutes; at last the female seemed to rememberthat she was nest building and flew up the mountain side followed by the male.Considerable time was spent on every trip after material, so very few stickswere added each day, and it was not until six days had elapsed that theflimsy platform was completed and the egg laid.Eggs.?Most authorities agree that this pigeon lays, almost invari-ably, only one egg; but there are a number of apparently authenticrecords of two eggs in a nest. The egg is elliptical ovate, generallysomewhat pointed and pure white. The shell is smooth and slightlyglossy. The measurements of 19 eggs average 39.7 by 27.9 milli-meters ; the eggs showing the four extremes measure 43.5 by 30, 39.3by 30.2, and 36.8 by 25.9 millimeters.Yovng.?According to Major Bendire (1892) the period of incuba-tion is " from 18 to 20 days, both sexes assisting. The young growrapidly, and are able to leave the nest when about a month old." Thenesting season is prolonged through 10 months of the year, and theevidence shows that probably several young are raised during the sea-son. Clinton G. Abbott (1927) has published some notes fromAlbert E. Stillman, who had good opportunities for studying thisspecies in San Diego Count}7 . In his notes for September 17, 1922,he writes
:
That day the female left the oak tree in the early morning and returned attwilight ; after quickly feeding the young she left again. Next clay she left atdaybreak and returned at sundown. For more than a week after that I keptthe youngster with me during the day, letting him perch on my finger or hopabout on the cabin floor, returning him to his nest before evening. On themorning of October 2 I found that the young bird had climbed from the nestand was sitting on a branch of the oak tree, where he remained until late in theafternoon. That night he roosted on the high limb of a nearby pine tree. OnOctober 4 he left the neighborhood and I did not see him again.That successive young pigeons are sometimes raised in one nest the sameseason was proved by Bushnell in 1925. He found a nest on March S containingone egg, from which the squab hatched and grew up. Then the pigeon laidan egg in the same nest and started to incubate. The second young birdhatched about the middle of May and lived to leave the nest.Dr. Joseph Grinnell (1928a) watched a pigeon feeding its youngin the Yosemite region on September 29, 1927, of which he says
:
Soon an old bird alighted, coming up the same steep course as the first one, atmid-height of the trees through the forest, and alighted on a branch of the nesttree, on a level with the nest but on nearly the opposite side of the trunk.
BAND-TAILED PIGEON 357After remaining perched quietly for awhile, the old bird then walked along thebranch lengthwise to the trunk, hopped across, fluttering some, to the base ofthe nest branch and walked out on it to the nest.Immediately a commotion began?the young one fluttering its wings spasmod-ically, the old one, not plainly seen because of intervening foliage, evidentlyfeeding it. The process lasted fully three minutes, when the old bird flew di-rectly off from the nest, out into space from a cliff: base, and circling, was seento alight at far distance on a middle branch of a dead tree. We would havetimed the feeding process if we had had any notion of its lasting so long.After feeding, the youngster crouched down motionless and could be seenplainly no more. When being fed, its upraised, fluttering wings showed thequills to be only an inch or so long; it could have been no more than ten daysold.Mrs. Irene G. Wheelock (1904) says that the squab is "fed on athin milky fluid, by regurgitation, for 20 days."'Plumages.?I have never seen a nestling band-tailed pigeon, butMrs. Wheelock (1904) saj^s that its " yellow skin is covered with thesparse, cottony, white down." The juvenal plumage is much like thatof the adult, but it lacks the white collar and the iridescent metalliccolors on the neck; the vinaceous tints are wholly lacking; and thefeathers of the breast and wing coverts are narrowly edged withwhitish, giving a slightly scaly effect. Molting birds are scarce incollections, but apparently young birds have a complete molt duringthe first fall, which produces a practically adult plumage. I havebeen unable to trace the molts of adults, but a complete molt prob-ably occurs, as it does in European pigeons, during summer and fall
;
this may begin as early as May or June and end as late as Octoberor November.Food.?Grinnell, Bryant, and Storer (1918) have published a veryfull account of the food of the band-tailed pigeon, from which I caninclude only a condensed summary. As their food consists mainlyof nuts and berries, which are intermittent crops, the pigeons find itnecessary to wander about considerably, congregating in large num-bers where food is abundant and deserting these same localities dur-ing seasons of scarcity. Acorns seem to be their chief food; prob-ably all the oaks are patronized, but mainly the live oaks, goldenoak, and black oak; the acorn crop lasts through a long season infall and winter. The acorns are swallowed whole and form anattractive food supply in the fall. They resort at times to the apple-like fruits of certain species of manzanita, eating them from the timethey are first formed and green until late in fall, when they are fullyripe. Early in the fall they feed on the fruit of the coffeeberry,elderberry, and chokecherry. In winter they have the toyon, orChristmasberry, and when the nut and fruit crops become exhaustedthey feed on the flower and leaf buds of the same plants, such asmanzanita and oak buds. Early in spring sycamore balls are fre-74564?32 24
358 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMquently eaten; as many as 35 have been counted in the crop of onepigeon. Fruits of dogwood, wild peas, pine seeds, and other seedshave been found in their crops. Considerable cultivated grain iseaten ; this is mainly waste grain, picked up in stubble fields of bar-ley, oats, and corn ; but pigeons have been known to do some damageby pulling up newly sown seed barley ; such records are scarce, how-ever. P. A. Taverner (1926) says that in western Canada, "theyare especially partial to peas and are said to pull up the sproutingseeds. The flocks so engaged are described as being numerousenough to turn the colour of the fields they alight upon from brownto blue. As they are large birds, each one intent on filling a capa-cious crop, their power for damage is not small. In the autumnthey alight on the stooked grain and may take a considerable tollof it."Other observers have noted in their food hazel, pinyon and othernuts, wild grapes, wild cherries, wild mulberries, blueberries, black-berries, raspberries, juniper, cascara, salmon and salal berries, andgrasshoppers and other insects. Their method of feeding on man-zanita berries is thus described by Laurence M. Huey (1913) :Some boys there told me that for the past two weeks a buuch of about onehundred pigeons had been feeding on green manzanita berries in a near-bythicket, and I was much pleased when they offered to take me to the place. Itproved to be about one and one-half miles north of their ranch, due south ofVolcan Mountain, and was the only thicket thereabout having a large crop ofberries. In the morning the birds would begin to arrive a little after sunrise,leaving between eight and nine o'clock ; in the evening they returned about fourand stayed until dark. They seemed always to come from and return to thesame place, at the top of Volcan Mountain among the pine trees.The pigeons seen were apparently always the same bunch, as one bird notedwith a few secondaries missing on the left wing was seen on three out of fouroccasions when the flock was encountered. It was interesting to watch themtrying to alight on the clusters of berries, far too weak to support them, makingmany futile attempts and finally succeeding in reaching the berries only bysettling on a stronger perch and then walking out to the cluster. But howthey did gorge and stuff when they finally got at them.Mr. Willard (1916) writes:A few days later a flock was observed feeding on acorns in a group of largeoak trees (Quercus emoryi). The antics of these birds were more like theacrobatic stunts of parrots than of pigeons. They would walk out on theslender branches till they tipped down, then, hanging by their feet, wouldsecure an acorn and drop off to alight on a branch lower down. In spite oftheir large size, pigeons are surprisingly inconspicuous when thus engaged infeeding among the leaves.M. French Gilman (1903) says of their feeding habits in thegrainfields
:
In March, 1901, great flocks of the pigeons poured into San Gorgonio Passand fed in the barley fields. For about two weeks there were hundreds of thembut they all left as suddenly as they had appeared. Their method of feeding
BAND-TAILED PIGEON 359
was peculiar. Instead of spreading out they kept together, alternately walk-ing and flying. Those behind would fly a few feet ahead of the advance line,alight, and walk along picking up grain until other rear ones would fly aheadand it came their turn again. In this way the flock advanced, some in the airall the time and ground was covered quite rapidly.Behavior.?Except during the nesting season, band-tailed pigeonsare decidedly gregarious, flying about in large, open, or scatteredflocks, formerly in flocks of hundreds, but now more often in flocksof dozens. They are fond of perching for long periods in the topsof tall trees; in the leafless sycamores in winter the flocks are veryconspicuous, but among the thick foliage of live oaks or eucalyptusthey are well hidden. If approached too closely, they will begin toleave, a few birds at a time, with loud flapping of wings, and thereare usually a few laggards that slip away from some unseen spot atthe last moment. Their flight is strong, direct, and very swift, re-minding one of domestic pigeons. According to Grinnell, Bryant,and Storer (1918), "in passage down a mountain side, the flight isinconceivably swift, the wings being held close in to the sides, beat-ing only at long intervals, and the body veering slightly from sideto side in its arrow-like course. This headlong flight produces arushing noise as of escaping steam."Voice.?The cooing notes of the band-tailed pigeon are much likethose of the domestic pigeon. Doctor Grinnell (1905) says that
" their deep monotonous coo'-coo, coo'-coo, coo'-coo, or tuck-oo',tuch-oo' was a frequent sound on Mount Pinos." Mrs. Florence M.Bailey (1902) writes:If you follow the pigeons to their breeding-grounds in some remote canyonyou will be struck by the owl-like hooting that fills the place, and you will locatethe sound here and there along the sides of the canyon at dead treetops, ineach of which a solitary male is sunning himself, at intervals puffing out hisbreast and hooting. The hooting varies considerably. Sometimes it is a calmwhoo'-ivhoo-hoo, tchoo'-hoo-hoo, and others a spirited hoop'-ah-whoo', and againa two syllabled ichoo'-ugh, made up of a short hard hoot and a long coo, as ifthe breath was sharply expelled for the first note and drawn in for the second.The method of uttering the notes is described in detail by JosephH. Wales (1926) as follows:When the male pigeon starts this performance he usually maneuvers aroundfor a firm footing and perhaps opens his bill slightly once or twice. Next hestretches his neck out in a line parallel with the axis of his body, and bendshis head down to a right angle. With his bill open a crack he gives one gaspwhich fills out the skin of his neck until about three times natural size, andat the same time utters a very faint oo which is not usually audible over twentyfeet All of these are preliminary actions, as directly following the first soundcomes the whoo-oo. This hoot is made by a quick expelling of the air fromthe bird's lungs, and is accompanied by a slight downward push which seemsto give abruptness to the first note. The swelled neck skin is not reduced, asthe bill is opened and the lungs are refilled for the following coo. There are
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usually about seven or eight of these hoots in a series, but sometimes as many aseleven. When finished, the male pigeon brings his neck back into its naturalposition and allows the air to escape from under the neck skin. This perform-ance is repeated at irregular intervals through the early morning and the latterpart of the afternoon.Fall.?Illustrating the heavy fall flights that formerly occurred inCalifornia, W. Leon Dawson (1923) writes:In the fall and winter of 1911-12, lured by an unusual crop of acorns, andimpelled, no doubt, by corresponding " crop " failures elsewhere, immensenumbers of Band-tailed Pigeons appeared in the interior valleys of Santa Bar-bara County, centering about the town of Los Olivos. It is probable that prac-tically the entire summer population of California north of the Tehachipe,Oregon, Washington, and British Columbia concentrated at this point. It isnot surprising, therefore, that " millions " of birds should have been reportedin this section, although half a million would probably be much nearer thetruth. What followed on this occasion was a humiliating example of whathuman cupidity, callousness, and ignorance, when unrestrained, will accom-plish toward the destruction of birds. Reports of the birds' abundance spreadrapidly. The " Wild Pigeon " of the East had unexpectedly turned up in theWest. Hunters from the outside flocked to the scene. Every gun was put intocommission. By automobiles and trainloads they came. The country wasaroar with gunfire. The ammunition business jumped in a dozen towns. En-terprising dealers organized shipments to the San Francisco and other mar-kets. W. Lee Chambers, writing for The Condor, reports a Sunday excur-sion of hunters from San Luis Obispo which brought home 1,560 birds. Anotherman, hunting for the San Francisco market, killed 280 pigeons under one oakin one day. The stupid birds, knowing nothing of their offense, flew miserablyfrom one part of the valley to another, but would not, or could not, forsaketheir food. How great the destruction of that winter really was is a matter ofmerest conjecture, but it must have been a very sensible proportion, possiblymore than half the entire species. I passed through this section of the countryon the 1st of the following April and saw only 28 pigeons, but the sides of theroad in many places were so covered with paper waste from cartridge boxes thatI was reminded of a street in Chinatown on the morning after New Year's.Fortunately, this destruction and the agitation which ensued prompted theGovernment to declare a five-year closed season on Band-tailed Pigeons.Game.?Grinnell, Bryant, and Storer (1918) say:The value of the Band-tailed Pigeon as a true game bird is to be concededwithout argument. Its pursuit is of a different type from that offered by anyother game species. An anonymous writer in southern California, who signshimself " Stillhunter," says that the best place for hunting pigeons there isnear a dead tree where the birds are known to alight. For such a situationhe advises using a .22 or 25-20 rifle ; then single birds may be secured withoutfrightening away others in the flock. For sneaking up on birds a " duck gun "is recommended. Ten pigeons are considered a good day's bag. If the fleshhas become " strong " by reason of the birds' acorn diet, soaking in brineflavored with vinegar or lemon will remove the disagreeable taste. After suchtreatment the birds should be broiled, or baked in a pot pie.Enemies.?Band-tailed pigeons apparently have few natural ene-mies, and these have proved of little consequence in reducing their
BAND-TAILED PIGEON 361
numbers. In spite of their slow rate of increase they could holdtheir own against natural' enemies, but they could not long resist theterrible slaughter by man when congregated in their winter quarters.Fortunately, they are now protected against this.Mr. Willard (1916) says that in Arizona " the Prairie Falcon andCooper Hawk take considerable toll from the flocks. These two ter-rors of the air will dash into a tree and grab a pigeon off a branch,rarely making an unsuccessful raid. The Prairie Falcon is the chiefoffender." Mr. Kitchin tells me that, in Washington, " apparentlythe only enemy these birds have during the breeding season is thelocal gray squirrel1, which I know on more than one occasion hastaken possession of the nest, using it as a foundation and adding to itto suit himself, and once I found the egg of the pigeon buried underthe structure that the squirrel had added."Winter.?Grinnell, Bryant, and Storer (1918) say that
?
north of the northern boundary of California the Band-tailed Pigeon is whollymigratory. It seems inevitable that this northern-bred contingent should movesouth into California for the winter season, and there is, therefore, little reasonto doubt the inference that the birds which concentrate in winter in west-central and southern California, represent the entire pigeon population of thePacific coast region. If this be true, it is of course apparent that, as far asthe whole Pacific coast region is concerned, California alone is, in winter,responsible for the existence of the species.DISTRIBUTIONRange.?British Columbia, the Western United States, and CentralAmerica.Breeding range.?The breeding range of the band-tailed pigeonextends north, to southwestern British Columbia (Courtenay andChilliwack). East to British Columbia (Chilliwack) ; Washington(Everett, Seattle, Tacoma, and Kalama) ; Oregon (Beaverton)
;
northeastern California (Lyonsville and Stirling City) ; Colorado(Estes Park and the Wet Mountains) ; New Mexico (Tres Piedras,Pecos Baldy, Sandia Mountains, Capitan Mountains, SacramentoMountains, and Guadalupe Mountains) ; western Texas (Dog Can-yon, Fort Davis, Marfa, and Chisos Mountains) ; and Puebla (LasVegas). South to Puebla (Las Vegas) ; Durango (Otmapa Ranch)and Lower California (Cape San Lucas). West to Lower Cali-fornia (Cape San Lucas, Victoria Mountains, and El Sauz) ; Cali-fornia (Laguna Mountains, Cuyamaca Mountains, Pine Mountain,San Jacinto Mountains, Mount Wilson, Mount Pinos, Lopez Canyon,San Jose, Lagunitas, Gualala, Eureka, and Crescent City) ; Oregon(Lookingglass, Newport, and Astoria) ; Washington (Granville, LaPush, and Neah Bay) ; and southwestern British Columbia (LakeCowichan and Courtenay).
362 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMWinter range.?In winter the species occurs regularly north toCalifornia (East Park and Alta) ; Arizona (Salt River Bird Reser-vation) ; and southern New Mexico (Haut Creek). East to NewMexico (Haut Creek and Silver City) ; and Guatemala (VolcanoToliman). South to Guatemala (Volcano Toliman) ; Chiapas(Pinabete) ; and Lower California (Mount Miraflores). West toLower California (Mount Miraflores, El Sauz, La Laguna, PierceRanch, and Guadalupe Valley) ; and California (El Cajon, LosAngeles, Carpentaria, Fremont Peak, and East Park).The range above described is for the entire species and is occupiedchiefly by the typical race, Golumba f. fasciata. Viosca's pigeon{Columba f. vioscae) is confined to southern Lower California and isapparently nonmigratory.Spring migration.?Early dates of spring arrival are : Colorado,Beulah, May 7, Moraine Hill, May 25, and Gold Hill, June 2; Ore-gon, Mercer, March 5, Corvallis, March 14, Beaverton, April 4,Southerlin, April 8, North Bend, April 10, and Tillamook, April 14;Washington, Clallam Bay, April 9, Menlo, April 12, Vancouver,April 17, and Everett, April 26; and British Columbia, Courtenay,March 31, Sumas, April 4, Burrard Inlet, April 5, Chilliwack, April13, and Hastings, April 26.Fall migration.?Late dates of fall departure are: British Co-lumbia, Courtenay, October 5, and Chilliwack, October 29 ; Washing-ton, Fort Steilacoom, September 25, Clallam Bay, October 15, Argyle,October 20, and Cascades of the Columbia, October 29; Oregon,Forest Grove, October 3, Tillamook, October 10, North Bend, October24, and Newport, October 28; and Colorado, Ouray, September 8,Del Norte, September 20, and Forks of the Rio Grande, September 26.A vertical migration from the higher mountains in Californiaalso is occasionally noted (Escondido, 1920).Casual records.?Band-tailed pigeons are rarely taken outside oftheir normal range. Patch (1922) noted them at Tow Hill, GrahamIsland, of the Queen Charlotte Group, on July 28, 1919, and statesthat there is one record from Bella Coola, British Columbia, indicat-ing that they may at times breed farther north than is now known.One was taken in 1905 near Crescent, Okla., and another on June 2,1912, at Englevale, N. Dak.Egg dates.?Washington and Oregon: 6 records, May 3 to July12. California: 46 records, March 6 to September 24; 23 records,May 10 to July 1. Arizona and New Mexico: 32 records, April 23to October 4 ; 16 records, June 16 to July 14. Southern Lower Cali-fornia: 35 records, January 22 to December 26; 18 records, June 21to July 28.
viosca's pigeon 363COLUMBA FASCIATA VIOSCAE BrewsterVIOSCA'S PIGEONHABITSThis pale race of the band-tailed pigeon was described by WilliamBrewster (1888), based on a study of a series of more than 100specimens collected in southern Lower California and named inhonor of Mr. Viosca, the United States consul at La Paz. He givesit the following characters : " Similar to G. fasciata but with the tailband wanting or only faintly indicated, the general coloring lighterand more uniform, the vinaceous tints, especially on the head, neck,and breast, much fainter and more or less replaced by bluish ash."As to its distribution (1902), he says:This pigeon seems to be strictly confined to the Cape Region, for neither Mr.Bryant nor Mr. Anthony has succeeded in finding it in the central or northernportions of the Peninsula where true fasciata is also apparently wanting.Chester C. Lamb (1926) writes:The Viosca Pigeon is, with one exception, known to occur only in the VictoriaMountains, sometimes known as the Sierra de la Laguna, or in the adjacentfoothills. The exception is Brewster's statement that Mr. Frazar saw largenumbers in San Jose del Cabo in September " passing southward." Duringmy own two years' residence in the Cape district, however, this bird was notseen outside the mountainous district above indicated. I very much doubt thepigeons leaving Lower California at all, as implied by Brewster on Frazar'sreport.I became acquainted with the Viosca Pigeon July 5, 1923, when I made myfirst trip to the Laguna Mountains, and in the next month found them abundant.The following year parts of four rnqnths were spent in their range, and I hadample opportunity to study and observe this isolated race of pigeon. It wascommon throughout the mountains, ranging from an altitude of 1,500 feet tothe tops, some 6,500 feet. At the lower levels the birds are found in thecanyons, where wild grapes and another native fruit grow ; but the type ofcountry they like best, and their real home, is the live-oak region of the highervalleys and canyons. These birds are swift and powerful fliers and it wouldnot take them long to travel for their food, either to the pinyon pines above orto the wild grapes and figs below, whenever they might wish to vary theiracorn diet.Nesting.?The nesting season is very variable or very much pro-longed. Reliable observers have found this pigeon nesting in Janu-ary, February, April, May, June, July, August, September, andDecember. Mr. Lamb (1926) says of its nesting sites:The majority of the numerous nests I examined were in live-oak trees, usu-ally situated on the forks of the larger horizontal limbs, and placed from 10to 20 feet above the ground. Some nests were also found placed among thesmaller branches and near their extremities, but this was exceptional. Avery few nests were found in a small species of white-oak tree that grows onthe hillsides. This oak is peculiar in that in the dry season the leaves turn
364 BULLETIN 162, UNITED STATES NATIONAL MUSEUMbrown and appear dead, but a few days after the first rain, the leaves graduallygrow green again.There are a few pine trees, mostly pinyons, scattered among the oaks insome parts, but only in one instance did I find a pigeon's nest in a pine. Thiswas a well built nest six feet above the ground, against the trunk where a hori-zontal limb grew out. One nest was found on a frond of a leaning fan palmtree. The nest is as a rule carelessly made, of a few coarse twigs, with nonest lining.A nest collected for me by W. W. Brown, in the Sierra de la Laguna,on June 14, 1913, containing one egg, was described as a frail plat-form-like structure of sticks, built near the extremity of a branch, ina pine tree about 40 feet from the ground. An egg in the UnitedStates National Museum, taken by M. A. Frazar, near Pearce's ranch,on July 18, 1887, was presented by Mr. Brewster; it was taken froma nest composed of a few sticks, 18 feet up, on a broken upright branchof a giant cactus.Eggs.?One egg seems to be the almost invariable rule with Viosca'spigeon. If two eggs are ever laid, it must be very rarely, for in morethan 25 nests examined by Mr. Lamb and 8 or 10 by Mr. Brown,only one egg or young was found. The egg is pure white, like thatof the band-tailed pigeon. The measurements of 25 eggs average39.7 by 27.5 millimeters ; the eggs showing the four extremes measure43.2 by 28.5, 41.4 by 29, 36.7 by 26.9, and 38.1 by 26.4 millimeters.Food.?Mr. Lamb (1926) says: "Acorns, wild grapes, pinyon nuts,and a sort of wild fig were, in my experience, their only food in thesummer."Behavior.?Again Lamb writes:At one of my camps in the Victoria Mountains, my work table was placeddirectly under a large live-oak tree which bore an abundant crop of acorns.This was a great attraction to the pigeons as well as to numerous Narrow-fronted Acorn-storing Woodpeckers. It was a marvel to me how such a largebird as a pigeon could alight in this tree, even on its slenderest branches, with-out the least audible flapping of its wings; often I would be unaware of apigeon's presence until it was made known to me by the woodpeckers. Thepigeons and woodpeckers, it appears, are inherent enemies. Let a pigeon alightin this tree, and if a woodpecker is near-by, the latter immediately, withloud cries, sets upon and drives the pigeon away, which departs with a greatflapping of wings. In no case have I seen a pigeon try to defend itself, andone was never seen to take the part of the aggressor. When attacked, a pigeonflies to a near-by tree and often, as soon as the woodpecker's back is turned,so to speak, the pigeon is back again in the oak tree, only to have the samething happen again. It is lucky for the pigeons that woodpeckers are notalways on guard, else they would get but few acorns.Voice. Again he writes:The first bird voices one hears in the early morning in the live-oak regionare those of the Narrow-fronted Woodpeckers, closely followed by the VioscaPigeons, whose mellow tchoo-ichoo (first note short, second long and slightlylower) sounds almost human, jvs if someone were trying to attract attention.
RED-BILLED PIGEON 365From the specimens taken I learned it was only the males that make this sound.At this time the birds perch upon some dead or bare limb, usually at someelevation. They are frequently seen fluttering spirally with short wing-beatsor sailing slowly over some clearing, and then an entirely different note isuttered, at short intervals, hard to describe, but which could be called a sort ofhoarse, guttural croak, sounded for a sustained period.Fall?Brewster (1902) says:At San Jose del Cabo large flocks were observed in September passing south-ward. Mr. Frazar believes that the majority left Lower California that seasonbefore winter set in, although he saw a few on November 15 along the roadbetween San Jose and Miraflores and others at San Jose del Rancho December18-25. None were found on the Sierra de la Laguna between November 27and December 2.COLUMBA PLAVIROSTRIS FLAVIROSTRIS WaglerRED-BILLED PIGEONHABITSThe large, handsome red-billed pigeon, locally known as " bluepigeon," is a Central American species that extends its range intothe United States only in a narrow belt of heavily wooded bottom-lands along the valley of the Rio Grande in Texas and perhaps asfar west as the Graham Mountains in southern Arizona, where MajorBendire (1892) reports the capture of three specimens. I first metthis pigeon in the heavy timber near the Resaca de la Palma, nearBrownsville, Tex. This forest of ebony, huisache, mesquite, andhackberry trees, with its thick undergrowth of thorny shrubs and itstangles of vines, has been more fully described under the chachalaca.This pigeon is a resident in this region for about 10 months eachyear. George B. Sennett (1879) quotes Dr. T. M. Finley as sayingthat at Hidalgo these pigeons were " first noticed on January 24th inflocks ; about the middle of February they were seen in the woods inpairs, and cooing. The last seen of them in 1877 was the latterpart of November. These Pigeons were seen several times consortingwith tame Pigeons in the ebony-trees in the neighborhood of thevillage of Hidalgo." Dr. J. C. Merrill (1878) says: "This largeand handsome pigeon is found in abundance during the summermonths, arriving in flocks of fifteen or twenty about the last week inFebruary. Though not very uncommon about Fort Brown, it ismuch more plentiful a few miles higher up the river, where the densewoods offer it the shade and retirement it seeks."Nesting.?On May 27, 1923, near Resaca de la Palma, we found twonests of the red-billed pigeon in the heavily timbered thickets. Onewas about 10 feet up in a tangle of vines and saplings ; it was a small,frail nest of small twigs, barely strong enough to support the weight
366 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
of the single, fat squab that it held. Another similar nest, in muchthe same kind of a location, held a single fresh egg.Three typical nests are described by Sennett (1878) as follows:The locality was a grove of large trees, with undergrowth, and clumps ofbushes matted with vines. While prying about the thick vines I flushed thebird off its nest, and it alighted in one of the tall trees near by. It took me buta moment or two to examine the nest and shoot the bird. In less than ten min-utes' time I had also its mate. The nest was only eight or nine feet from theground, and set upon the horizontal branches of a sapling in the midst of thevines. It was composed of sticks, lined with fine stems and grasses, had a de-pression of an inch or more, and was about eight inches in outside diameter bytwo and one-half inches deep. It contained one egg, with embryo just formed.Dissection of the bird showed that she would have laid no more.On May 8th, at Lomita Ranche, a few miles above Hidalgo, in the fine grove ofebonies in the rear of the buildings of the ranche, I found two nests. Both werewell up in the trees, one about twenty-five feet and the other about thirty. Thenests were situated close to the body of the trees, on large branches, and werecomposed of sticks and grasses, with an inside depth of about two inches. Onecontained a single egg, far advanced ; in the other also lay a solitary egg, fromwhich a young chick was just emerging. The parents persisted in staying about,notwithstanding we were making a great disturbance, even shooting into thesame trees. Whenever we would go off some distance they would immediatelygo on their nests and seemed loth to leave them at our return. These were theonly ones seen breeding so near habitations. The grove was a common resortfor man and beast, besides being the place where wagons, tools, etc., were keptand repaired.On May 11th, I obtained my fifth and last nest. I found it in the woods at thefork of two roads, a mile or so from the village, down the river. This nestI had discovered a week or so before, complete, but empty. It was situatedabout ten feet from the ground, in one of a thick clump of small trees, at thejunction of several small branches. It was composed of twigs and rootlets,without grasses, and had a depression of one and one-half inches. The bird wasflushed from the nest and shot. Upon examination, the solitary egg showed thatincubation had begun, and dissection of the bird proved that no other eggs weredeveloped for laying.Again (1879) he says:This bird breeds irregularly and lays several times in a season. I foundnests during the whole time of my stay, containing eggs and young in allstages of development, but in no case did a nest contain more than one egg oryoung. The parents are fond and affectionate, and both assist in incubation.A. J. Van Rossem has sent me the following notes on its nestinghabits in Salvador:The breeding season apparently extends throughout the year, for males inbreeding condition were taken in January, April, May, July, September, Novem-ber, and December, and females either laying or about to do so in July, Sep-tember, and November. A nest was found at Lake Olomega on April 11, 1926.It consisted of only a few twigs, barely sufficient to keep the egg from rollingabout, placed on top of two crossing fronds about 6 feet from the ground. Thevery sheltered location was the only thing that prevented the haphazard col-lection of twigs from falling to the ground with the first passing breeze. The
RED-BILLED PIGEON 367
male was on the nest and did not fly off until the hunter was directly beneaththe nest.Eggs.?Apparently the red-billed pigeon usually lays but one egg,though possibly very rarely two. The shape is oval or elliptical oval,the shell is smooth and slightly glossy, and the color is pure white.The measurements of 33 eggs average 38.6 by 27.3 millimeters; theeggs showing the four extremes measure 41.4 by 27.9, 39 by 29.5, 34by 26.5, and 36.8 by 25 millimeters.Plumages.?The squab, which I found in the nest mentionedabove, is described in my notes as nearly naked, the dark, reddish-brown skin being only sparsely covered with short, black pinfeathers.Sennett (1879) says:The young from the egg have the upper parts plumbeous and sparselycovered with dark hair-like feathers. Under parts are pale and naked. Thehalf-grown young have plumage on the body like the adult. Head and flanksdo not become feathered until bird is nearly fledged, and in half-grown youngJust commences to show.Food.?There is a brief statement by Sennett (1879) that "theirfood when " he " saw them was chiefly the hackberry fruit."Col. Andrew J. Grayson (1871) says:This is the largest of our pigeons, and abundant in the Marias, as well as insome localities on the mainland. It is gregarious and frequents large forests,feeding upon various kinds of berries, acorns, etc., etc. It migrates from onepart of the country to another in small flocks. In sonre seasons of the year theflesh of this bird has a bitter, disagreeable taste, caused by some species ofberry or small bitter acorn upon which it subsists.Dr. E. W. Nelson (1899), writing of the birds of the Tres MariasIslands, states
:
On Maria Magdalena they were numerous in some trees near a group ofdeserted houses and in old clearings a short distance back from the shore.They came to these trees to feed upon the ripening fruit, but were rather shy.When one becomes startled and takes wing it makes a loud flapping noisethat alarms its companions, and then all dash swiftly away. They were lessconfiding than most of the birds on the islands, but were not so shy as theirrepresentatives on the mainland. Wild figs and the small fruit of a tree, prob-ably a species of Psidium, or wild guava, were favorite articles of food. Theirloud cooing note is uttered at short intervals and is one of the characteristicsounds in the forests they frequent. They are essentially arboreal in habitsand are rarely seen near the ground.Behavior.?The red-billed pigeon reminds one of a domesticpigeon in its swift, strong, steady, and direct flight and in its similarbut louder cooing notes.Sennett (1878) writes:Like all the Pigeons, it is fond of the water. Any morning will find numbersof all the different species going to and coming from the sand-bars in the river,where they are in the habit of drinking and bathing. The cooing of this bird
368 BULLETIN" 162, UNITED STATES NATIONAL MUSEUM
is clear, short, and rather high-pitched. It is more secluded in its habitsthan any of the others, except the one I have lately found new to our fauna,AecJimoptila albifrons. In point of numbers it is much less numerous than theCarolina and the White-winged Doves ; still, it is quite extensively shot formarket.George N. Lawrence (1874) quotes Colpnel Grayson as follows:This fine species in some localities of Western Mexico is quite abundant,particularly in the region of Mazatlan River. It frequents the larger forests,and feeds upon various kinds of berries, acorns, and the tender buds of sometrees. It is partially gregarious but is often seen solitary and in pairs.Small flocks of from twenty to fifty migrate from one part of the country to an-other in search of its favorite food. I have found it at a considerable height onthe western slope of the Sierras Madres, feeding upon acorns, that are therein abundance in some seasons. The flesh of this pigeon is tough, and some-times bitter to the taste, caused by the bitter acorn, and also by an astringentkind of berry, upon which they may be subsisting at the time. As a gamebird it is inferior to most of our pigeons for the table, but, being a large andhandsome bird it is sure to attract the attention of the gunner. They are noteasily approached, however, being very shy, and without the strictest cautionthe hunter would not be able to fill his bag with this game in a long day'stramp. DISTRIBUTIONRange.?Central America north to the lower Rio Grande Valley;nonmigratory.The range of the red-billed pigeon extends north to southernSonora (Sierra de Alamos) ; northeastern Sinaloa (El Toro) ; north-ern Nuevo Leon (Alamo and Trevino) ; and southern Texas (Car-rizo, Lomita, Hidalgo, and Brownsville). East to Texas (Hidalgoand Brownsville) ; Tamaulipas (Matamoras, Rio Pilon, Rio Cruz,Ciudad Victoria, Aldama, and Altamira) ; Vera Cruz (Tampico,Misantla, Cordoba, and Alvarado) ; Yucatan (Tunkas and ChichenItza) ; and Costa Rica (Guayabo, This, and Dota). South to CostaRica (Dota, Naranjo, and Bolson) ; Salvador (Volcano San Mi-guel) ; Guatemala (Esquintla, San Jose, Retalhuleu, and El Naran-jo) ; Chiapas (Tonala) ; Oaxaca (Topana and Tehuantepec) ; Guer-rero (Acapulco) ; and Jalisco (Las Penas). West to Jalisco (LasPenas) ; Tepic (Cleofas Island and Tres Marias Islands) ; south-western Sinaloa (Escuinapa, Presidio, and Mazatlan) ; and south-ern Sonora (Sierra de Alamos).Casual records.?Bendire (1892) reports three specimens takennear Fort Grant, in the foothills of the Graham Mountains, Ariz.,July 25, 1886.Egg dates.?Texas and Mexico : 79 records, March 1 to August 8 \40 records, April 23 to June 6.
WHITE-CROWNED PIGEON 369COLUMBA LEUCOCEPHALA LinnaeusWHITE-CROWNED PIGEONHABITSThe fine white-crowned pigeon is a permanent resident in theBahamas and West Indies and occurs in the Florida keys as a sum-mer resident only. Audubon (1840) writes:The White-headed Pigeon arrives on the Southern Keys of the Floridas,from the Island of Cuba, about the 20th of April, sometimes not until the1st of May, for the purpose of residing there for a season, and rearing itsyoung. On the 30th of April, I shot several immediately after their arrivalfrom across the Gulf Stream. I saw them as they approached the shore,skimming along the surface of the waters, flying with great rapidity, muchin the manner of the common house species, but not near each other likethe Passenger Pigeon. On nearing the land, they rose to the height of abouta hundred yards, surveyed the country in large circles, then with lessvelocity gradually descended, and alighted in the thickest parts of the man-groves and other low trees. None of them could be easily seen in those darkretreats, and we were obliged to force them out, in order to shoot them, whichwe did at this time on the wing.In creeping among the bushes to obtain a view of them whilst alighted,I observed that the more I advanced, the more they retired from me. Thisthey did by alighting on the ground from the trees, among which they couldnot well make way on wing, although they could get on with much easebelow, running off and hiding at every convenient spot that occurred. Thesemanoeuvres lasted only a few days, after which I could see them perchedon the tops of the trees, giving a preference perhaps to dry branches, butnot a marked one, as some other species are wont to do.Of their haunts there he says:The key on which I first saw this bird, lies about twenty-five miles south ofIndian Key, and is named Bahia-honda Duck Key. The farther south we pro-ceeded the more we saw, until we reached the low. sandy, sterile keys, called theTortugas, on none of which did I see a Pigeon of any kind. On our return fromthe Tortugas to Key West, our vessel anchored close to a small key, in a snugharbour protected from the sea winds by several long and narrow islands wellknown to the navigators of those seas. Captain Day and myself visited thislittle key, which was not much more than an acre in extent, the same after-noon. No sooner had we landed than, to our delight, we saw a great numberof White-headed Pigeons rise, fly round the key several times, and all realightupon it. The Captain posted himself at one end of the key, I at the other, whilethe sailors walked about to raise the birds. In less than two hours we shotthirty-six of them, mostly on the wing. Their attachment to this islet resultedfrom their having nests with eggs on it. Along with them we found Grakles,Red-winged Starlings, Flycatchers, and a few Zenaida Doves.The next morning we thought of calling at this little key on our way, andwere surprised to find that many new comers had arrived there before us.They were, however, very shy, and we procured only seventeen in all. I feltconvinced that this spot was a favourite place of resort to these birds. It beingdetached from all other keys, furnished with rank herbaceous plants, cactuses,and low shrubs, and guarded by a thick hedge of mangroves, no place could be
370 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMbetter adapted for breeding ; and, at each visit we paid it, White-headed Pigeonswere procured.On Jamaica, P. H. Gosse (1847) says:This flue dove is common in almost all situations, but chiefly affects thegroves of pimento, which generally adorn the mountain pens. The sweet aro-matic berries afford him abundant and delicious food during the pimento sea-son; the umbrageous trees afford him a concealment suited to his shy andsuspicious character ; and on them his mate prefers to build her rude platform-nest and rear her tender progeny.Dr. Alexander Wetmore (1916) writes:The white-crowned pigeon was formerly one of the most abundant speciesin Porto Rico, but now is found only in a few localities. Gundlach spoke of itas very common in the seventies, but its numbers have undoubtedly greatlydecreased. The birds occur mainly near the coast, usually in dense swampygrowths, though one was seen near Aibonito ; and the few small areas of forestremaining in the lowlands may account for their diminution in numbers.Around Punta Picua, north of Mameyes, they were found preparing to breedin the swamps, where the growth was so dense that it was hard to get nearthem. They usually came out into the more open portions late in the eveningto feed on the fruit of the icaco (Chrysobalanus sp.), but even then kept wellconcealed in the thick leaves.Courtship.?Audubon (1840) describes its courtship as follows:The White-headed Pigeon exhibits little of the pomposity of the commondomestic species, in its amorous moments. The male, however, struts beforethe female with elegance, and the tones of his voice are quite sufficient to per-suade her of the sincerity of his attachment. During calm and clear mornings,when nature appears in all her purity and brightness, the cooing of thisPigeon may be heard at a considerable distance, mingling in full concord withthe softer tones of the Zenaida Dove. The bird, standing almost erect, fullplumed, and proud of his beauty, emits at first a loud croohoo. as a prelude, andthen proceeds to repeat his coo-coo-coo. These sounds are continued during theperiod of incubation, and are at all times welcome to the ear of the visitor ofthese remarkable islands. When approached suddenly, it emits a hollow, gut-tural sound, precisely resembling that of the Common Pigeon on such occasions.Nesting.?The same gifted author says
:
The nest is placed high or low, according to circumstances; but there arenever two on the same tree. I have found it on the top shoots of a cactus, onlya few feet from the ground, on the upper branches of a mangrove, or quite low,almost touching the water, and hanging over it. In general the nest resemblesthat of the Columba migratoria, but it is more compact, and better lined. Theouter part is composed of small dry twigs, the inner of fibrous roots andgrasses.In the Bahamas, according to Dr. Henry Bryant (1861)
?
It breeds in communities, in some places, as at Grassy Kays, Andros Island,in vast numbers ; here the nests were made on the tops of the prickly pear, whichcover the whole kay ; at the Biminis and Buena Vista Kay, Ragged Island, onthe mangroves ; and at Long Rock, near Exuma, on the stunted bushes. I donot think they ever select a large kay for their breeding place.
WHITE-CEOWNED PIGEON" 371C. J. Maynard (1896) describes an abandoned nesting colony in theBahamas as follows:One of the most remarkable sights that I ever witnessed as regards numbersof birds' nests was on one of the Washerwomen Keys off the South shore ofAndros. These are small, rocky islets, lying on the barrier reef, and are sometwenty-five feet high. On one of these little keys, which did not contain over anacre of land, there were at least ten thousand nests of the White-headed Pigeon.The rocks were mostly covered with a scanty growth of low bushes and with amore luxuriant growth of cacti, and upon both plants and bushes the birds hadplaced their nests, and some were upon elevated portions of rock, while a fewwere placed upon the naked ground. So completely covered was the southernand northern portion of the key that the nests were nowhere over two feet apartand often nearer together than that. Unfortunately, however, all of these nestswere of the previous year, only a single dove being seen. My boatmen informedme that this rookery was occupied by many thousand birds during the past year,and that the spongers were accustomed to visit the place at night and capture thesitting birds. This statement was confirmed by the remains of torches whichwere scattered about the island. Many nests contained eggshells, the contentsof which had been removed by Buzzards, Man-of-war birds or Gulls. The timeof this visit was May 8th, 1884.On the Isle of Pines, W. E. Clyde Todd (1916) says that "thenest is usually built in the top of a royal palm, but along the LosIndios River the birds were found nesting in the mangroves, ratherlow down."Dr. Paul Bartsch writes to me that the nests he found on SanSalvadorwere all placed in mangrove clumps such as are shown in the habitat photo-graph. These clumps stood out in the lakes at some distance from shore andfurnished splendid protection. Furthermore, as a rule, there was a graykingbird's nest in the top branches of these clumps, and the kingbird served asan alarmist. The birds were exceedingly shy, regardless of whether they wereincubating eggs or taking care of young.Eggs.?The white-crowned pigeon ordinarily lays two eggs, butsometimes only one. The eggs are elliptical oval or nearly oval, purewhite, smooth, and quite glossy. The measurements of 35 eggs aver-age 36.8 by 27 millimeters; the eggs showing the four extremesmeasure 40.2 by 28.4, 39.4 by 29.5, 32.3 by 26.2, and 36.8 by 25.2millimeters.Plumages.?I have not seen the downy young of this pigeon, butDoctor Bartsch tells me that the down is of a " light buff color."Audubon (1840) says:The young birds are at first almost black, but have tufts of a soft buff-coloureddown distributed mostly over the head and shoulders. While yet squabs theyhave no appearance of white on the head, and they take about four monthsbefore they acquire their perfect plumage. Smaller size, and a less degree ofbrilliancy, distinguished the female from the male.
372 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMWilson (1832) says:The young are distinguished by duller tints, and the crown is at first nearlyuniform with the rest of their dark plumage ; this part, after a time, changesto grey, then greyish white, and becomes whiter and whiter as the bird growsolder.Gosse (1847) writes of some squabs that he raised in captivity:Both were exceedingly ugly ; long-necked, thin-bodied, the head not wellrounded, the fleshy part of the beak prominent, and its base unfeathered.The whole plumage was blackish ash-colored, each feather slightly tippedwith paler, and the feather of the head terminating in little curled greyfilaments, which added to the uncouth appearance of the birds. In a weekor two I perceived these filaments were gradually disappearing, and aboutthe beginning of October the small feathers began to clothe the base of thebeak ; these feathers were greyish-white, and at the same time the grey huewas beginning to spread up the forehead, I believe, by the dropping of theblack feathers and their immediate replacement by the white ones. Aboutthis time also the general plumage began to assume the blue hue of the adult,in patches ; and on the 12th of October, I first observed the beautiful iridesceutfeathers of the neck, but as yet only on one side. These notes refer to theelder ; the other was about two weeks more backward. On the 16th, I firstheard it coo ; for some time it had now and then uttered a single note, but onthis day it gave the whole Sary-coat-'blue, but short, and in a low tone; andthat only once. By the end of November the white had spread over thewhole crown, as in the adult; but was not yet so pure or so smooth.I have seen several young birds, both males and females, appar-ently in first-winter plumage, taken at various dates from December14 to May 16, that had evidently matured more slowly or were, per-haps, hatched later. They all had dull gray or dirty white crownsand were otherwise like adult females, except that they had no scalymarkings on the neck; the neck and mantle were dull brown, withdarker edgings; the wing coverts and scapulars had narrow lighttips. There was a mixture of new plumbeous, adult plumage in theback, and the wings and tail were either molting or had been re-cently renewed, showing that young birds at least have a completemolt in winter and spring. I have been unable to learn anythingabout the molts of adults.Food.?Gosse (1847) gives a very good account of the feedinghabits of the " baldpate," as he calls this pigeon ; he writes
:
When the pimento is out of season, he seeks other food ; the berries of thesweetwood, the larger ones of the breadnut, and burn-wood, of the bastard cedar,and the fig, and the little ruddy clusters of the fiddle-wood, attract him. He!'< ds early in the morning, and late in the afternoon; large numbers resort to asingle tree (though not strictly gregarious), and when this is observed, thesportsman, by going thither before dawn, and lying in wait, may shoot themone by one, as they arrive. In September and October they are in fine condition,often exceedingly fat and juicy, and of exquisite flavour. In March the clammy-cherry displays its showy scarlet racemes, to which the Bald-pates flock. The
WHITE-CROWNED PIGEON 373Hopping Dick, Woodpecker, and the Guinea-fowl feed also upon it. In April,Sam tells me he has seen as many as thirty, almost covering a tree, feeding onberries which he believes were those of the bully-tree. Late in the year theyresort to the saline morasses, to feed on the seeds of the black-mangrove, whichI have repeatedly found in the craw ; I have even seen one descend to the groundbeneath a mangrove, doubtless in search of the fallen seeds. In general, how-ever, the Bald-pate is an arboreal pigeon, his visits to the earth being very rare.He often feeds at a distance from home ; so that it is a common thing to observe,just before nightfall, straggling parties of two or three, or individuals, rushingalong with arrowy swiftness in a straight line to some distant wood.Doctor Wetmore (1916), on Porto Rico, found that " five stomachsexamined contained vegetable matter only, composed of drupes andfruits of fair size. The icaco and berries of various palms (palmoreal and lluma) are favorites with these birds, while a tree known aspalo bianco {Din/petes sp.) is said to furnish them food in season.No cultivated crop is injured, the bird depending wholly upon wildfruits for its sustenance." Mr. March, as quoted by Baird, Brewer,and Ridgway (1905), states that "they commit serious depredationson the Guinea-corn fields, not only by the quantity they devour, butby breaking down the brittle corn-stalks with the weight of theirbodies."Behavior.?Maynard (1896) writes:The White-headed Pigeons are thoroughly at home among the thick branchesof the trees and shrubbery of the Bahamas, moving about among them as easilyas do the smaller-perching birds, and they make very little noise. When sur-prised by an intruder they will remain perfectly quiet until approached withina few yards, when they will spring rapidly into air, rise to the tips of the wood-land, and dart off with an exceedingly rapid flight; in fact, few, if any birds,can fly any more quickly than do these Pigeons. I have shot several in air, asthey rose from the bushes and darted away, but I never attempted to shoot oneas it passed me at full speed at right angles. When dashing along at this head-long speed they will suddenly alight upon a branch or on the ground, withoutthe beating or fluttering of the wings, which usually attends a similar abruptstoppage in most birds of a similar size and which is so noticeable in ourdomestic pigeon.Dr. Thomas Barbour (1923) says:The White-crowned Pigeon is of irregular appearance in any given locality,its presence depending on the abundance of the fruits upon which it feeds.It is essentially a coastal form, and one which is always gregarious. It roostsin great hordes, usually on some mangrove islet, and bands sally forth eachmorn to feed, returning from their distant foragings at dusk. Then theyrush and swirl into the greater resorts, or palomares, in incredible hosts.Famous roosts are Moraine Cay north of Grand Bahama, where I have shot,and Green Cay, south of New Providence. Gundlach speaks of their seldombeing seen in Cuba except when nesting, which they do at various seasonsof the year. This intermittent appearance is noticed everywhere. They areiii the Florida Keys in summer only, but not every summer in equal numbers
;
in certain of the Bahamas they abound at one season, elsewhere at others.74564?32 25
374 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe fact is, the individual bands are capable of long flights, and move farand wide as food supplies dictate. Great numbers are slaughtered by hunters,who build an ambush near roost or rookery and kill the returning birds asthey fly in just before dark. Unfortunately, this leaves many young birdsto starve.Voice.?In addition to the notes mentioned under courtship, Imight quote what Maynard (1896) says:The notes of this Pigeon are very loud and characteristic, sounding some-thing like wof, wof, wo, co-woo. The first three notes are repeated severaltimes, then the co-tooo is long drawn out; all being In as low a key as thehoot of an owl. The entire cry is cleverly imitated by the Creoles when theywish to decoy the bird within gun shot, but there is a certain tremulousnessin the real notes which cannot be imitated by the human voice.Again Gosse (1847) gives a still different wording of it as " sary-coat-blue, uttered with much energy, the second syllable short andsuddenly elevated, the last a little protracted and descending."Game.?Nearly everyone who has written about the birds of theBahamas or the West Indies has referred to the wThite-crownedpigeon as one of the finest game birds on these islands. Its gamequalities are excellent, as it is rather wild and a very swift flier;and its flesh is delicious on the table. It was formerly much moreabundant than it is now, for it has been shot in enormous numberswhile flying to or from its breeding, feeding, or roosting grounds.As the pigeons were shot most easily on or about their breedinggrounds, many young were left to starve in the nests and many oldersquabs were taken for food or to be raised in captivity. With suchwholesale slaughter the species is rapidly disappearing and is badlyin need of protection. Maynard (1896) relates the following tale:About the first week in July, previous to 1SS4, sportsmen from Nassau hadbeen in the habit of visiting Green Key and shooting the breeding Pigeons asthey flew from their nests to cross to Andros Island, some fifteen miles distant,where they are said to go daily for food and water. Many of the nests of theprevious season which I had examined on Green Key contained broken eggsthat contained the remains of half-formed young, and in some of the nestswere the skeletons of newly hatched young ; the parents of both eggs and younghad doubtlessly been killed as they left the nests. This sight was a mostpiteous appeal to humanity. I was informed by one of my boatmen, who hadaccompanied hunting parties to the key, that so great was the slaughter ofPigeons that many more were killed than were needed, and that he had fre-quently seen hundreds of birds buried in the sand of the beach near wherethey were shot. Upon my return to Nassau I promptly stated the facts as Ihad observed them to the Governor, Sir Henry A. Blake, and, as I have else-where stated in this work, through his ready and sympathetic cooperation alaw was enacted protecting these Pigeons during the breeding season.Winter.?The wanderings of the white-crowned pigeons have beenreferred to in the foregoing quotation from Doctor Barbour. Theseare all winter, or between breeding seasons, wanderings. The birds
WHITE-CROWNED PIGEON 375
are evidently absent from the Florida Keys in winter, and Mr.Todd (1916) says of their exodus from the Isle of Pines:This is a common species everywhere, except in the Cienaga, appearing inflocks late in February, and remaining until the last of September. Althougha few stragglers may be seen through the winter months, the vast majorityof the individuals withdraw at that season from their usual range and accord-ing to native report resort to the " south coast," in great numbers. It is oneof the most numerous birds of the various mountain ridges in the interior of theisland during the breeding-season, which begins in May.This last movement is northward to the larger land area of Cuba,probably to find a better food supply. The birds that breed on theFlorida Keys are probably those that migrate northward to themainland of Florida in winter. Recent information indicates thatthese pigeons migrate to extreme southern Florida occasionally, per-haps regularly, in winter. Gilbert R. Rossignol writes me that hesaw some of these pigeons between Flamingo and Coot Bay fivedifferent times between December 30, 1928, and February 24, 1929.He says : " I recall seeing three at one time and a pair here and therebetween the first and third bridges, but mostly around the secondbridge, where there is considerable open country due to some farm-ing and a burnt district." Frank N. Irving, who was with Mr. Ros-signol on some of these trips, tells me that he saw the white-crownedpigeons in the same region during March, 1928, and January andFebruary, 1929. I wrote to Harold H. Bailey for his experience,and he replied that he has taken several on the mainland at CapeSable and has seen flocks there of 20 or 30 birds, or more, manytimes ; they were feeding in the higher foliage of the Florida hollyand other berry-bearing trees. He has not found a nest on the main-land and thinks they come there to feed only and spend the winternear abundant food. He says that a similar movement takes placein the Bahamas, where the pigeons desert their breeding grounds onthe outlying keys and come to Andros Island to spend the winter.DISTRIBUTIONRange.?Southern Florida, the West Indies, and locally in centralCentral America.The range of the white-crowned pigeon extends north to southernFlorida (probably Dry Tortugas, Key West, Bahia Honda, CapeSable, Coconut Grove, and Indian Key) ; and the Bahama Islands(Abaco Island). East to the Bahama Islands (Abaco, New Provi-dence, Green Cay, and Mariguana Island) ; and the Lesser Antilles(Barbuda Island and Antigua Island) . South to the Lesser Antilles(Antigua and St. Croix Islands) ; Porto Rico (Vieques Island, PuntaPicua, and Mona Island) ; Haiti (San Domingo and Jacmel)
;
376 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMJamaica (Spanishtown) ; Nicaragua (Great Corn Island) ; Honduras(Iiuatan Island) ; British Honduras (Half Moon Cay, Turneff Is-land, and Belize); and Oaxaca (Salina Cruz). West to Oaxaca(Salina Cruz and Tehuantepec) ; Yucatan (Cozumel Island andBuchotz) ; and southern Florida (probably Dry Tortugas).Scott (1889) records a specimen taken at Punta Rassa, Fla., August16, 1886, and Cory (1891) noted them in the winter of 1891 at CaicosIslands of the eastern Bahamas.Although given to considerable wandering, the white-crownedpigeon does not appear to have a regular migration, at least in themain part of its range. Audubon (1840) stated that they arrivedon the Florida Keys from April 20 to May 1. while Maynard (1896)reported their arrival in this region about the 1st of June and theirdeparture late in October. Some appear to winter regularly insouthern Florida.Egg dates.?Bahamas : 48 records, May 21 to December 8 ; 24records, June 18 to 29.COLUMBA SQUAMOSA BonnaterreSCALED PIGEONHABITSThe scaled pigeon is a West Indian species, which is included inour list as an accidental straggler to Key West, Fla., and thereforeextralimital. John W. Atkins (1899) records the incident, as fol-lows :On October 24, 1898, an adult female of this species was shot on the Islandof Key West, and brought to me in the flesh, by a young collector in myemployment, who found it among some Doves in the possession of a dove hunter,who had shot it from a wild fig tree on the outskirts of the town.Dr. Thomas Barbour (1923) says of its haunts in Cuba:In western and central Cuba this beautiful Pigeon is by no means commonat the present time. It is a highland bird but not exclusively confined tomountain ranges. One finds the Torcaza Morada usually perched high on thedead branches of some towering tree, most often on cliffs or steepish slopes.The birds seem sluggish and make short flights, booming their heavy, sonorouscall through the heat of the day. Attempt to approach, and the bird is off,for no Pigeon is more alert. Its flesh is excellent, and the body is heavy beyondother local species. In appearance in the field it is larger and darker than adomestic pigeon, and it has a patch of brilliant metallic feathers on each sideof the neck. It is never terrestrial. Ramsdeu has given an excellent accountof the persecution it suffers in Oriente, where it appears at intervals in greatnumbers. Ramsden also recalls breeding rookeries which Gundlach neverfound. This gregarious habit is beyond a doubt confined to the wild EasternProvince, where the Scaly-naped Pigeon still is more abundant than elsewhere.For never elsewhere have I met with numbers which would allow of killing
SCALED PIGEON 377five-thousand individuals in a couple of weeks in one locality. It still occurs inthe regions mentioned hy Gundlach, in the mountains of Vuelta Abajo and Trini-dad, but in both these highlands it may today be seen regularly ouly in pairs,trios, or small bands, and probably never over a few dozen in a day?and manydays far fewer would be seen. Slaughter for food and sport has already verygreatly reduced this splendid species, and it needs now protection, which prob-ably will not be granted to it, and which, if granted, can not be enforced.In Porto Eico, in 1912, Dr. Alexander Wetmore (1927) "foundthe species common in the hills and mountains, where small tractsof natural forest and extensive coffee fincas offered it shelter, butsaw it seldom on the coastal plain, where there was in the main littleshelter to attract it."Nesting.?As to its nesting habits he says:It is a common belief in Porto Rico that the scaled pigeon is only a migranton the island?a belief promulgated, it may be said, by gunners who desirean open season during the entire year. That this is erroneous was proven onMarch 8, 1912. when without special search I found three nests on El Yunque,while there was no doubt whatever that the dozens of birds flushing on everyhand were breeding. The three nests definitely located were made of sticksloosely piled together and placed about fifteen feet from the ground on horizontallimbs, or on refuse piled on large air plants. Two were empty, while one con-tained a single egg, plain white in color with a slight gloss, which was collected.This egg had had about five days' incubation. It measures 34.8 by 26.7mm. At Maricao, on June 1, a native brought me a young bird about two-thirds grown, and said it was the only one in the nest. Gundlach has saidthat two eggs are laid, but from these instances it would seem that a singleegg in a set is not unusual.A nest found by Harry A. Beatty (1930) on St. Croix " was a veryfrail platform of coarse sticks, situated 25 feet up on the forkedbranch of a mahogany tree. I could see plainly the two glossy whiteeggs through the nest from below."Eggs.?The egg, taken by Doctor Wetmore, referred to above, isoval in shape, slightly elliptical, rather glossy, and pure white. Themeasurements of seven eggs average 36 by 26.7 millimeters ; the eggsshowing the four extremes measure 37.6 by 27.4, 35.5 by 27.5, 31.8 by26.7, and 34.5 by 25.7 millimeters.Plumages.?I have not seen the downy young. Young birds injuvenal plumage, in July, are like the adult, but lack the richlycolored, scaly markings on the hind neck; the head and neck are
" warm blackish brown," shading off to " dark plumbeous " on theback and to " walnut brown " on the upper breast, the latter feathershaving darker edgings ; the wing coverts are edged with " walnutbrown." I have seen adults showing wing molt from September toDecember, but I can learn nothing further from the scantymaterial available.Food.?Doctor Wetmore (1927) says that " it feeds on wild berriesand fruits, with occasional succulent leaves or shoots. The berries
37S BULLETIN 162, UNITED STATES NATIONAL MUSEUM
of various palms, wild figs, the moral (Cordia), and jagua (Genipaamericana), with various wild legumes, are eaten extensively." Hesays elsewhere (1916) :All of the smaller wild fruits in season appear to furnish food, and theseare so abundant that cultivated fields are not molested. The fruits eaten,though sometimes of comparatively large size and with hard stony pits, areswallowed entire. The strong muscular gizzard of the bird has a tremendoustriturating power, however, and the fruits are easily crushed and the meatycenters opened to the processes of digestion.Behavior.?Referring to the habits of the scaled pigeon in PortoRico, Doctor Wetmore writes
:
The dense forests covering the slopes of El Yunque de Luquillo, in the north-eastern part of Porto Rico, harbored great numbers of these birds, whichranged commonly up to 2.500 feet above the sea. In late afternoon and eve-ning, near the Hacienda Catalina, it was a common sight to see them circlingabout high in the air. In spite of their large size, they were difficult to seein tlie trees, even in the thin foliage of the cacao rosetta (Sloanea berteriana)
.
Thus it often happened that bird after bird flew out from amid the limbs, withloudly clapping wings, yet failed to offer a shot, while I peered vainly upwardin search for their hidden companions. When one of the big males chancedto drop in near another, a great flapping of wings ensued until one was forcedto take flight. The ordinary call note was a loud, strongly accented tcho-hoo-hoo, while a burring guttural hoo-o-o-o, given with a throaty rattle, wasalmost startling when heard from directly overhead. Many birds descendedto feed amid the tall trees fringing small streams at the foot of the mountain,and some were encountered in the dense, swampy forests near Punta Picua,beyond Mameyes. Males rest and call at times in the tops of tall, dead trees.Fall.?Doctor Wetmore says thatThe paloma tutca is said to occur in large flocks during fall, and to gather innumbers where wild fruits are ripening, at which time many are killed. Itis common belief that these flocks are entirely migratory, but there can be nodoubt that they come mainly from the forests on El Yunque and elsewherein the interior.Game.?The same writer states
:
The species is the only game bird of importance in the inland region ofPotto Rico and affords excellent sport, as it is wary, strong on the wing, andis found only in the wildest, roughest country. It should be protected fromFebruary 1 to October 15 each year, if not longer, to permit it to breed, asotherwise it cannot maintain its status.Austin H. Clark (1905), referring to the southern Lesser Antilles,writes
:
This is the chief game bird of these islands, and is much hunted. The flightis rapid and powerful, and the birds regularly cross over from one island toanother to feed, returning at night to roost on the smaller keys. Formerlynumbers could be shot any evening about four o'clock from Clifton House, UnionIsland, us they flew from that island over to Prune to spend the night. Theycould be obtained at Hermitage House, Carriacou, in the same manner, as theyflew past, going to one or other of the small keys near by.
PASSENGER PIGEON 379DISTRIBUTIONRange.?The West Indies ; accidental at Key West, Fla. ; non-migratory.The range of the scaled pigeon extends north to Cuba (NuevaGerona and Guantanamo) ; northern Haiti (Massif du Nord, MorneSalnave, and Catare) ; northern Porto Rico (Desecheo Island, Cule-bra Island, and Culebrita Island) ; and the Lesser Antilles (St.Eustatius and Guadaloupe). East to the Lesser Antilles (Guade-loupe, Dominica, Martinique, Santa Lucia, St. Vincent, the Grena-dines, Carriacou, and Grenada). South, to the southern LesserAntilles (Grenada) ; southern Haiti (Selle Mountains and HatteMountains) ; and the Isle of Pines, Cuba (Nueva Gerona). Westto the Isle of Pines (Nueva Gerona).Egg dates.?West Indies : 2 records, March 8 and May 26.ECTOPISTES MIGRATORIUS (Linnaens).PASSENGER PIGEONHABITSContributed by Charles Wendell TownsendThe passenger pigeon, or wild pigeon, as it was often called, isgenerally believed by ornithologists to be extinct. Of the mightyhosts of this splendid bird that swarmed over the country, not asingle individual is thought to remain alive to-day, and yet withinthe memory of men not yet old, the bird was well known, and thepossibility of its extinction was far from their thoughts. Indeed,whenever laws were proposed for conserving the bird, the cry atonce went up that it needed no protection, for its numbers and theextent of country over which it ranged were both so huge that pro-tection seemed unnecessary. Even the tardy protective laws passedby some States were largely disregarded.At last, in 1910, 1911, and 1912, when it was too late, attemptswere made to save the bird, and rewards that totaled more than$1,000 were offered for evidence that it was living and nesting
?
the live bird, not the dead one was sought. But it was all in vain.The passenger pigeon appears to have gone the way of the dodo andthe great auk.James H. Fleming, who has made the most complete and criticalstudies of the recorded specimens of the passenger pigeon, believes(1907) that " for all practical purposes the close of the nineteenthcentury saw the final extinction of the passenger pigeon in the wildstate and there remained only the small flock, numbering in 1903not more than a dozen, that had been bred in captivity by Prof.
380 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMC. O. Whitman of Chicago. These birds, the descendants of a singlepair, had long before that ceased to breed.'' The last of this group,a female in the Cincinnati Zoo, died of old age in September, 1914.The last records of wild birds that are based on specimens aboutwhich there is no doubt appear all to have been taken in 1898?one.an adult male taken at Lake Winnipegosis, Manitoba, on April 14(Fleming, 1903 and 1907) ; an immature male at Owensboro, Ky.,on July 27, 1898 (Fleming, 1907), now in the Smithsonian Insti-tution; and an immature bird taken at Detroit, Mich., on September14, 1898 (Fleming, 1903 and 1907). Fleming (1907) adds in a foot-note : " There is a mature female in the collection of the CarnegieInstitution of Pittsburg, Pa., marked ' Pennsylvania, August 15.1898,' but without further locality." Still another specimen, thefifth for 1898, was a young male shot by Addison P. Wilbur atCanandaigua, N. Y., on September 14, 1898, and is recorded by E. H.Eaton (1910), who states that he saw it killed, and that it "wasunquestionably reared in the spring of 1898, as it was just assumingthe adult plumage."L. E. Wyman (1921). of the Museum of History, Science, and Art,Los Angeles, Calif., records the following : "A mounted specimen ofpassenger pigeon acquired b}r the late F. S. Daggett, in January,1920, and now in the Daggett collection, deposited in this museum,bears the following label : ' Passenger Pigeon, male, No. 315, Coll. ofGeo. S. Hamlin. Shot by a Swede, North Bridgeport, Fairfield Co.,Conn., Aug., 1906.' " This seems conclusive, but under date of Sep-tember 19, 1930, Mr. Fleming writes me, " I suggest that August,1906, is the date the bird was acquired by Hamlin and who thenwrote the label." Mr. Fleming calls attention to the fact that in " TheBirds of Connecticut," by Sage, Bishop, and Bliss (1913), the lastauthentic specimen for the State is recorded for October 1, 1889, and,what is very significant, in notes communicated by Hamlin there isno mention of a specimen taken later than 1892.A specimen in the Cornell University Museum, recorded by S. C.Bishop and A. H. Wright (1917), was shot at Clyde, N. Y., by J. L.Howard, who, when more than 80 years of age in 1915, stated frommemory that it was taken about 6 years previous, that is, in 1909.This date is rendered extremely doubtful from the fact that on thebottom of the mount is the date July 5, 1898, although this may havebeen the record of another bird on the same mount. Mr. Howardgives a circumstantial account of his shooting the bird, and states hehad not seen any passenger pigeons before this for about 1 5 years.It is true that since this time there have been many sight recordsof the bird reported, some of which, at least some of the earlier ones,are doubtless authentic, and at the present day there seems to be a
PASSENGEB PIGEON 381
recrudescence of the belief that the bird still exists. In Science ofFebruary 14, 1930, Prof. Philip Hadley (1930) notes on the au-thority of others?not ornithologists?sight records of these birds innorthern Michigan. Unfortunately, the mourning dove is oftenmistaken for the passenger pigeon, and in the West the band-tailedpigeon has been similarly mistaken. The distinguishing field marksof these birds will be discussed later, but it seems to be a common ideathat the passenger pigeon is easily recognized by its size, which islarger than that of the mourning dove. All ornithologists know,however, in the absence of direct comparison, how deceptive differ-ence in size may be, and they are well aware that not only to anuncritical observer but even to an expert ornithologist, a mourningdove may often look as large as a passenger pigeon. The fact thatthe observer has seen thousands of passenger pigeons in years be-fore, and has handled and plucked them, does not necessarily meanhe is a good judge of the bird. The wish is father to the thought,especially in the unscientific, and it has been proved over and overagain that birds reported as passenger pigeons have turned out to bemourning doves.It may be worth while to enter here the following example amongmany of the manner in which the will to believe in the existenceof the passenger pigeon surmounts all obstacles and all evidence.Dr. C. F. Hodge (1912), who had charge of the offer of $1,000for the location of a nesting pair of passenger pigeons, received aletter from a man in Maine that he had shot a bird " that proved tobe a passenger pigeon," and that he had had it mounted. DoctorHodge sent him descriptions and colored plates of the passengerpigeon and mourning dove, and he underscored in red ink the com-parative lengths of these two birds, and he asked the man if, afterstudy of these, he still thought the bird a passenger pigeon tosend it to him. The specimen arrived and proved to be a mourningdove. In a recent newspaper article, an old-time pigeon trapper isquoted as saying : " No one familiar with this bird could possiblymake a mistake. The mourning dove, although it has the samecoloration, is smaller. I can tell a mourning dove from a pigeonas far as I can see it." With this belief it is easy to see passengerpigeons! This, however, is not conclusive that all the reports areerroneous, and, although the evidence points strongly to the extinc-tion of the passenger pigeon, it is proper to keep an open mind onthe subject and investigate plausible clues.There is a popular idea that the passenger pigeon mysteriouslydisappeared, and that, while still enormously numerous, it suddenlyceased to exist. Its annihilation has been popularly attributed tovarious natural phenomena, and it has even been rumored that the
382 BULLETIN 162, UNITED STATES NATIONAL MUSEUMbird has migrated to South America, but the real cause for itsextinction?man?is not mentioned. The natural phenomena sup-posed to be causative of the extinction are: Epidemics, tornadoes,earty deep snowstorms, forest fires, strong winds, while the birdswere crossing large bodies of water, causing exhaustion and deathby drowning. Some of these were reported while the bird was stillcommon in other localities. Circumstantial accounts were publishedof immense numbers drowned in the Gulf of Mexico, a region be-yond the usual range of the bird. The destruction of the forestsby ax and by fire undoubtedly has been a large detrimental factorin the life history of the pigeons, for the forests supplied theirprincipal food as well as roosting and nesting places. These naturalcauses had acted for countless ages but the passenger pigeon sur-vived, but when the white man arrived on the North American Con-tinent, and especially after the pigeon became a commercial asset,its destruction, unless some curb was put on the slaughter, wasordained.The evidence that man is responsible for the enormous destructionis voluminous and convincing, but why, it may be asked, did not afew escape and continue the race when the numbers became so smallthat the pursuit of the bird was no longer commercially profitable?It is believed by some that the bird still exists in small numbers inremote parts of northern Michigan and in Canada. In answer tothis it may be said that a bird accustomed for ages to living togetherin large numbers and close ranks, whether in feeding, migrating,roosting, or nesting, might find it impossible to continue satisfactorilythese functions with greatly reduced and scattered ranks. It is prob-ably no mere figure of speech to say that under these circumstancessuch a communistic bird would " lose heart," nor is it fanciful tosuppose that sterility might in consequence affect the remnants. Theease and thoroughness with which the squabs were destroyed, even inpreference to the adults, would alone account for the extinction. Justas a forest finally dies of disease, accident, and old age if all theseedlings are destroyed by intensive grazing, so any species of birdis doomed in the same way to extinction if its offspring are an-nihilated before they reach maturity.The probable final stages in the disappearance of the passengerpigeon are well portrayed by W. B. Barrows (1912) as follows:In the opinion of the writer the most probable cause for the disappearanceof the pigeon lies in the fact that, through the clearing of the forests and theincreasing persecution by man, the birds were driven from one place to anotherand gradually compelled to nest farther and farther to the north, and underconditions successively less and less favorable, so that eventually the largerpart of the great flocks consisted of old birds, which, through stress of weatherand persecution, abandoned their nesting places and failed to rear any consider-
PASSENGER PIGEON 383
able number of young. Under such conditions they would naturally becomeweaker, or at least less resistant, each year, and in the attempt to find nestingplaces in the far north they may have been overwhelmed by snow and ice duringone or two of the unusually severe summers that occurred between 1882 and1S90.Dr. Thomas S. Koberts (1919) sums up the whole situation whenhe says of this bird in Minnesota
:
Formerly an abundant summer resident. Rapidly diminished in numbersbetween the years 1878 and 1885, finally disappearing entirely between 1890 and1900. It is now extinct everywhere. All other theories to the contrary, theextermination of this bird was the result of ruthless and wholesale destructionby man.Courtship.?The early writers describe a courtship very much afterthe fashion of the domestic pigeon. Thus Audubon (1840) says:The male assumes a pompous demeanor, and follows the female, whether onthe ground or the branehes, with spread tail and drooping wings, which it rubsagainst the part over which it is moving. The body is elevated, the throatswells, the eyes sparkle. He continues his notes, and now and then rises onthe wing, and flies a few yards to approach the fugitive and timorous female.Like the domestic pigeon and other species, they caress each other by billing,in which action the bill of the one is introduced transversely into that of theother.All that Wilson (1832) has to say on the subject is that " they havethe same cooing notes common to domestic pigeons, but much less oftheir gesticulations."Wallace Craig (1911a) believes that Audubon's description ofthe courtship and voice " came largely by reasoning by analogy fromthe domestic pigeon and from the author's charming but somewhatunscientific imagination," because Craig's careful studies of captivepassenger pigeons in the aviary of Prof. C. O. Whitman showedvarious peculiarities and characteristics, unlike those of the domesticpigeon, characteristics that " all seem connected, directly or indi-rectly, with the extreme gregariousness, the breeding in vastcolonies." Thus he found no bowing or strutting or charging asin other species. The male emits a loud hek and chattering notesand waves his wings in a single sweep, or flaps the wings repeatedly,holding tight to the perch the whole body, head, and tail rising andfalling with each stroke.When close beside the female, the male Ectopistes had a way all his ownof sidling up to her on the perch, pressing hard upon her, sometimes puttinghis neck over her neck, "hugging" her as Professor Whitman expressed it.* * * When the female becomes amorous, instead of edging away fromthe male when he sidles up to her, she reciprocates in the hugging, pressingupon the male in somewhat the same manner that he presses upon her. * * *The act of billing, which occurs in all pigeons before copulation, is in Ectopistesreduced to a mere form. * * * The bills ate quickly clasped, shaken fora fraction of a second, and as quickly separated ; the performance is precisely
384 BULLETIN 162, UNITED STATES NATIONAL MUSEUMlike a brief, quick handshake. It is probable that there is no passing offood from one mouth to the other.The passenger pigeon was essentially an arboreal bird and con-ducted its courtship in the trees. Chief Pokagon (1895), the lastPottawattomie chief, says that about the middle of May, 1850, ata great nesting place in Michigan, " the trees were filled with themsitting in pairs in convenient crotches of the limb, now and thengently flattering their half-spread wings and uttering to their matesthose strange, bell-like waving notes which I had mistaken for theringing of bells in the distance."During the courtship period much fighting occurred among themales in their crowded quarters, but little injury resulted. Craig(1911a) says:The male Ectophtcs was a particularly quarrelsome bird, ever ready tothreaten or strike with his wings (though perhaps not quite so ready withhis beak), and to shout defiance in his loud strident voice. * * * He wasan aggressive, violent threatener, but not a real fighter.Nesting.?As the passenger pigeon approached extinction it nestedin small companies or singly, but until its numbers were greatly re-duced the nestings took place in communities of great size. Theaccounts of these nesting communities by various authors are almostinvariably accompanied with a description of the means used for thecapture and slaughter of the adults and squabs. These will bequoted later under enemies. There are, however, a few descriptionsof the nestings alone. Wilson (1832) says theyare generally in beech woods, and often extend in nearly a straight line acrossthe country for a great way. Not far from Shelbyville in the state of Ken-tucky, about five years ago, there was one of these breeding places, whichstretched through the woods in nearly a north and south direction, was severalmiles in breadth, and was said to be upwards of forty miles in extent ! In thistract almost every tree was furnished with nests wherever the branches couldaccommodate them."More than 100 nests in one tree alone were not uncommon. Mershon(1907) says a game dealer in Detroit had seen a nesting place inWisconsin that extended through the woods for 100 miles.In the spring of 1885 William Brewster and Jonathan Dwightwent to Michigan hoping to see a nesting of the passenger pigeon.In this they were unsuccessful as they found only one nest, butthey collected a great deal of valuable information. Brewster (1889)quotes S. S. Stevens of Cadillac, " a veteran pigeon netter of largeexperience?a man of high reputation for veracity and carefulnessof statement," as follows:The last nesting in Michigan of any importance was in 1881, a few mileswest of Grand Traverse. It was of only moderate size, perhaps eight miles long.The largest nesting he ever visited was in 1876 or 1877. It began near Petosky
PASSENGER PIGEON 385
and extended northeast past Crooked Lake for twenty-eight miles, averagingthree or four miles wide. Nestings usually start in deciduous woods, but duringtheir progress the pigeons do not skip any kind of trees they encounter. ThePetosky nesting extended eight miles through hard-wood timber, then crossed ariver bottom wooded with arbor-vitae, and thence stretched through pinewoods about twenty miles. For the entire distance of twenty-eight miles everytree of any size had more or less nests, and many trees were filled with themNone were lower than fifteen feet above the ground.Mr. Stevens also stated that " so rapidly did the colony extend itsboundaries that it soon passed literally over and around the placewhere he was netting, although when he began, this point was severalmiles from the nearest nest."The nests were simple frail structures, composed of sticks andtwigs crossing one another and supported by forks of the branchesat a height of 10 to 50 feet or more. The nest was often so looselymade that the egg or squab could be seen through it from below.Ruthven Deane (1896) observed the habits of captive birdsbelonging to David Whittaker, of Milwaukee. These birds hadincreased from two pairs procured in 1888 to six males and ninefemales in 1896.When the pigeons show signs of nesting, small twigs are thrown on thebottom of the enclosure, and on the day of our visit, I was so fortunate as towatch the operations of nest building. There were three pairs actively engaged.The females remained on the shelf, and at a given signal which they only utteredfor this purpose, the males would select a twig or straw, and in one instancea feather and fly up to the nest, drop it and return to the ground, while thefemales placed the building material in position and then called for more. Inall of Mr. Whittaker's experience with this flock he has never known of morethan one egg being deposited. * * * The eggs are usually laid from themiddle of February to the middle of September, some females laying as many asseven or eight during the season, though three or four is the average.This fact of one egg to the clutch is also confirmed by Prof. C. O.Whitman. "Wilson (1832) also confirms this, " a circumstance," headds, " in the history of the bird not generally known to naturalists."Audubon and many who have followed and copied him state, how-ever, that two eggs form the clutch, as is the case with the domesticand other pigeons. There are, however, a sufficient number of inde-pendent observations that show that two eggs are often, or, accordingto others, rarely found. It is possible, in the crowded nesting places,that two females may have laid in the same nest. Brewster (1889)questioned Stevens closely on the number of eggs in the nest. " Heassured me that he had frequently found two eggs or two young inthe same nest, but that fully half the nests which he had examinedcontained only one." Mr. Brewster adds : " Mr. Stevens is satisfiedthat pigeons continue laying and hatching during the entire summer.They do not, however, use the same nesting place a second time inone season, the entire colony always moving from twenty to one
386 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMhundred miles after the appearance of each brood of young." Whenfood was plentiful it is believed that three or even four broods ofyoung were raised in a season.Eggs.?[Author's note: The passenger pigeon laid either one ortwo eggs in a set, probably more often only one. Most of theauthentic eggs that I have seen are decidedly elongated, elliptical-ovate in shape, but this may not be the invariable rule. Theshell is smooth and slightly glossy. The color is pure white.The measurements of 32 eggs, apparently authentic, average 38.2 by27 millimeters; the eggs showing the four extremes measure 45.2 by29.7, 33.5 by 26, and 36.2 by 24.9 millimeters.]Young.?The incubation period is 14 days as observed exactly incaptive birds (Deane, 1896). Both birds take part in the incubation,a point that was accurately determined by netters as shown by theircatches. Brewster (1889) quoting Stevens, says:Both birds incubate, the females between two o'clock p. m. and nine or teno'clock the next morning; the males from nine or ten o'clock a. m. to twoo'clock p. m. The males feed twice each day, namely, from daylight to abouteight o'clock a. m. and again late in the afternoon. The females feed onlyduring the forenoon. The change is made with great regularity as to time,all the males being on the nest by ten o'clock a. m. During the morning andevening no females are ever caught by the netters; during the forenoon nomales. The sitting bird does not leave the nest until the bill of its incomingmate nearly touches its tail, the former slipping off as the latter takes itsplace.Kuthven Deane (1896) says:During the first few days, after the young is hatched, to guard against thecold, it is like the egg, concealed under the feathers of the abdomen, the headalways pointing forward. In this attitude, the parents, without changing thesitting position or reclining on the side, feed the squab by arching the head andneck down and administering the food. The young leave the nest in aboutfourteen days.As the nesting of passenger pigeons often began before snowfallshad ceased, the following note by Frank J. Thompson (1881) showingtheir hardihood under such circumstances is of interest:In confinement in Cincinnati early in March, 1S78, two pairs began nesting,the male carrying the material while the female busied herself in placing it.A single egg was soon laid in each nest and incubation commenced. On March16 there was quite a heavy fall of snow and on the next morning I was unableto see the birds on their nests on account of the accumulation of the snowpiled on the platforms around them. Within a couple of days it had dis-appeared and for the next four or five nights a self-registering thermometer,hanging in the aviary, marked from 14? to 19?. In spite of these drawbacksboth of the eggs were hatched and the young ones reared.Brewster (1889) says:The young are forced out of their nests by the old birds. Mr. Stevens hastwice seen this done. One of the pigeons, usually the male, pushes the young
PASSENGEB PIGEON 387
off the nest by force. The latter struggles and squeals precisely like a tamesquab, but is finally crowded out along the branch and after farther feebleresistance flutters down to the ground. Three or four days elapse before itis able to fly well. Upon leaving the nest it is often fatter and heavier thanthe old birds ; but it quickly becomes much thinner and lighter, despite theenormous quantity of food that it consumes.Another point about young birds was brought out by Brewster(1889) from Stevens. He writes:On one occasion an immense flock of young birds became bewildered in afog while crossing Crooked Lake and descending struck the water and per-ished by thousands. The shore for miles was covered a foot or more deepwith them. The old birds rose above the fog and none were killed.As with all the pigeons, the passenger pigeon fed its young atfirst with the so-called " pigeon milk," the secretions from theglandular crop, mixed with the food in the crop, and serving todigest it. It is probable that the young inserted its bill into thatof the mother to obtain this " milk," although some writers statethat the mother inserted her bill into that of the young. In theselarge colonies, communal feeding of squabs that had lost theirmothers has been observed.Plumages.?[Author's note: I have seen but one nestling of thepassenger pigeon ; this is quite completely, but very thinly, coveredwith long, soft, hairlike, " honey-yellow " down. This and two speci-mens showing the development of the juvenal plumage are in theMuseum of Comparative Zoology, in Cambridge, Mass. A smalljuvenal female, 8 inches long, is nearly fledged, but the yellowishdown filaments still adorn the head, neck, and breast ; the crown andupper back are " bister " or " warm sepia," shading off to " natalbrown " on the breast and to " wood brown " on the lesser wingcoverts and scapulars; the feathers of the back, wing coverts, andscapulars are edged with whitish, or pinkish, buff; the greater covertsshade from " fawn color " to " French gray," and are more narrowlyedged ; many of the inner coverts have a large patch of " bister
"
on the outer web ; the inner primaries are tipped and broadly edgedon the outer web with " Mikado brown," the edgings gradually dis-appearing outwardly ; the lower back and rump are " Quaker drab "to " mouse gray " ; the underparts shade off from " wood brown " onthe flanks to whitish on the belly and chin.Another young bird is fully fledged in juvenal plumage; thefeathers of the head, neck, and breast, now fully grown, have nar-row, buffy-white edgings; many of the outer wing coverts, espe-cially the greater, are "French gray"; the tail is shorter than theadult's, the central rectrices are browner and the lateral ones aredarker gray, so that there is less contrast in the tail.
388 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe postjuvenal molt took place during August, September, andOctober, earlier or later according to the date of hatching. It ap-parently involved all the contour plumage, the wing coverts, andthe tail. In the first winter plumage, which was practically adult,the sexes were differentiated. Adults had one complete molt inAugust and September.]Food.?The immense forests of North America, before they weredevastated and wasted by the ax and fire of the white settlers, fur-nished an inexhaustible supply of food to the enormous hosts ofwild pigeons. The principal food was mast?acorns, beechnuts, andchestnuts?and also the fruit or nuts of arry of the forest trees, aswell as wild berries and fruits, such as cherries, raspberries, blue-berries, currants, pokeberries (also called pigeon berries), and straw-berries, grain, seeds of weeds, and grasses. I have the records ofthe stomach contents of three passenger pigeons I shot at Magnolia,Mass., two in September, 1877, and one in July, 1878. All containedsmall pebbles and the hips and seeds of wild roses. S. C. Bishop(1924) reports that in the partially mummified remains of a pas-senger pigeon taken in New York State many }Tears ago, he foundin the crop 25 well-preserved seeds of the sugar maple and close to thebase of each fruit the wing had been sheared off and discarded.Aughey (1878) found that six stomachs of birds, taken in Nebraska,each averaged eight locusts and two other insects and some seeds.Audubon (1840) found three entire acorns in one crop and, in thestomach, fragments of others and three pieces of quartz. Records ofstomach contents of the passenger pigeon are rare. During the nest-ing season insects were largely eaten, especially earthworms, grubs,and grasshoppers. The pigeons were fond of salt and frequentednatural salt licks as well as grounds baited with salt.Pehr Kalm (1911) lists the following as food of the passengerpigeon : Acorns, beechnuts, seeds of red-flowered maple, of Americanelm. mulberries, rye, wheat, buckwheat, but not Indian corn, berries orthe tupelo, as well as of other trees and plants. He also notes thefondness of the pigeons for salt.Wilson (1832) says that besides acorns and beechnuts, " buck-wheat, hemp seed, Indian corn, holly berries, hackberries, huckle-berries, and many others furnish them with abundance of food at allseasons. The acorns of the live oak are also eagerly sought afterby these birds, and rice has been frequently found in individualskilled many hundred miles to the northward of the nearest riceplantation. * * * I have taken from the crop of a single wildpigeon, a good handful of the kernels of beechnuts intermixed withacorns and chestnuts." He calculated that the immense number ofpigeons he observed in a flock in Kentucky would eat 17,424,000
PASSENGER PIGEON 389bushels of mast a day, supposing that each bird consumed a pint.His calculations of the number of birds?2,230,272,000?an incrediblenumber, seems, as will be recorded later, to be a fair one.Ruthven Deane (1896), speaking of observations on captive birds,says:As soon as the young are hatched the parents are fed on earthworms, beetles,grubs, etc., which are placed in a box of earth, from which they greedily feed,afterwards nourishing the young in the usual way, by disgorging the contentsfrom the crop. At times the earth in the enclosure is moistened with waterand a handful of worms thrown in. which soon find their way under thesurface. The pigeons are so fond of these tidbits, they will often pick andscratch holes in their search, largo enough to almost hide themselves.Behavior.?The passenger pigeon was such a spectacular species inits migratory flights, its roostings, and its nestings, in which suchenormous numbers took part, that there are many references to itfrom the times of the earliest pioneers. From this mass of literatureit will be well to enter here some of the important reports, omittingtill later those that deal largely with the slaughter of the bird.Higginson (1630) writing of the region about Salem, Mass., says
:
Upon the eighth of March from after it was faire Daylight until about eightof the clock in the forenoon, there flew over all the towns in our Plantacions soemany flockes of Doves, each flock containing many thousands, and soe manythat they obscured the light, that passeth credit, if but the Truth should bewritten.Wood (1635) in the same region says:These Birds come into the Country to goe to the North parts in the begin-ning of our Spring, at which time (if I may be counted worthy, to be beleevedin a thing that is not so strange as true) I have seen them fly as if theAyerie regiment had been Pigeons, seeing neyther beginning nor ending, length,oi breadth of these Millions of Millions. The shouting of people, the ratlingof Gunnes, and pelting of small shotte could not drive them out of theircourse, but so they continued for foure or five houres together ; yet it mustnot be concluded, that it is thus often ; for it is but at the beginning of theSpring, and at Michaelmas [September 29], when they returne back to theSouthward; yet are there some all the yeare long, which are easily attaynedby such as looke after them.Pehr Kalm (1911) writing of a migration in March, 1740, in Penn-sylvania says:Their number, while in flight, extended 3 or 4 English miles in length andmore than one such mile in breadth, and they flew so closely together thatthe sky and the sun were obscured by them, the daylight becoming sensiblydiminished by their shadow.The big as well as the little trees in the woods, sometimes covering a dis-tance of 7 English miles, became so filled with them that hardly a twig orbranch could be seen which they did not cover; on the thicker branches theyhad piled themselves up one above another's backs, quite about a yard high.74564?32 26
390 BULLETIN 162, UNITED STATES NATIONAL MUSEUMWhen they alighted on the trees their weight was so heavy that not onlybig limbs and branches of the size of a man's thigh were broken straight off,but less firmly rooted trees broke down completely under the load.The ground below the trees where they had spent the night was entirelycovered with their dung, which lay in great heaps.Wilson (1832) thus describes a flock of pigeons that passed overhim as he was on his way to Frankfort, Ky.
:
Coming to an opening by the side of a creek called the Benson, where I hada more uninterrupted view, I was astonished at their appearance. They wereflying with great steadiness and rapidity, at a height beyond gunshot, inseveral strata deep, and so close together, that could a shot have reachedthem, one discharge could not have failed of bringing down several individuals.From right to left as far as the eye could reach, the breadth of this vastprocession extended, seeming everywhere equally crowded * * *. It wasthen half past one. I sat for more than an hour, but instead of diminution ofthis- prodigious procession, it seemed rather to increase both in numbers andrapidity; and, anxious to reach Frankfort before night, I rose and went on.About four o'clock in the afternoon I crossed the Kentucky river, at the town ofFrankfort, at which time the living torrent above my head seemed as numerousand as extensive as ever. Long after this I observed them, in large bodies thatcontinued to pass for six or eight minutes, and these again were followed byother detached bodies, all moving in the same south-east direction, till aftersix in the evening.Wilson calculated that this great mass of birds contained theincredible number of 2,230,272,000 individuals, and his method ofcalculation seems to be a conservative one. He assumed the flock tobe a mile in breadth, although he believed it was much more.Supposing it was moving at the rate of a mile a minute, as it was fourhours in passing, he estimated that its whole length would have been240 miles. He also assumed that each square yard contained threepigeons. As the flock was several strata deep there must have beenmany more than this.Wilson (1832) says:In descending the Ohio by myself, in the month of February, I often restedon my oars to contemplate their aerial manoeuvres. A column, eight or ten milesin length, would appear from Kentucky, high in air, steering across to Indiana.The leaders of this great body would sometimes gradually vary their course,until it formed a large bend of more than a mile in diameter, those behindtracing the exact route of their predecessors. This would continue sometimeslong after both extremities were beyond the reach of sight, so that the whole,with its glittering undulations, marked by a space on the face of the heavensresembling the winding of a vnst and majestic river. * * * . Sometimes ahawk would make a swoop on a particular part of the column, from a great height,when almost as quick as lightning, that part shot downwards out of the commontrack, but soon rising again, continued advancing at the same height as before;this inflection was continued by those behind, who on arriving at this point,dived down almost perpendicularly, to a great depth, and rising followed theexact path of those that went before.
PASSENGER PIGEON 391On another occasion, he says, " while talking with the people withindoors, I was suddenly struck with astonishment at a loud rushingroar, succeeded by instant darkness, which, on the first moment, Itook for a tornado about to overwhelm the house, and everythingaround in destruction. The people, observing my surprise, coollysaid, ' It is only the pigeons.' "Audubon (1840) describes similar great multitudes of passengerpigeons. In the autumn of 1813, while traveling 54 miles on thebanks of the Ohio River between Hardensburgh and Louisville, heobserved great flocks of pigeons flying southwest. He counted theflocks for 21 minutes in the morning and found that 163 had passed.
*' I traveled on," he says, " and still met more the farther I pro-ceeded. The air was literally filled with pigeons ; the light of noon-day was obscured as by an eclipse, the dung fell in spots not unlikemelting flakes of snow, and the continued buzz of wings had a tend-ency to lull my senses to repose." He reached Louisville at sunset.
" The pigeons were still passing in undiminished numbers and con-tinued to do so for three days in succession."Audubon (1840) describes as follows a roosting place on the banksof the Green River in Kentucky
:
It was, as is always the case, in a portion of the forest where the treeswere of great magnitude, and where there was little underwood. I rodethrough it upwards of forty miles, and, crossing it in different parts, found itsaverage breadth to be rather more than three miles * * *. The dung layseveral inches deep, covering the whole estent of the roosting-place. Manytrees two feet in diameter, I observed, were broken off at no great distance fromthe ground, and the branches of many of the largest and tallest had givenway as if the forest had been swept by a tornado. Everything proved to methat the number of birds resorting to this part of the forest must be immensebeyond conception.When the birds were coming to the roost, he continues
?
the noise which they made, though yet distant, reminded me of a hard galeat sea, passing through the rigging of a close-reefed vessel. As the birdsarrived and passed over me, I felt a current of air that surprised me * * *.The pigeons, arriving by thousands, alighted everywhere, one above an-other, until solid masses were formed on the branches all round. Here andthere the perches gave way under the weight with a crash, and, falling to theground, destroyed hundreds of birds beneath, forcing down the dense groupwith which every stick was loaded.A similar account, confirming that of Audubon, is given by Revoil(1869) of his experiences in 1847 of a pigeon roost near Hartford,Ky. As the sun set, the birds began to come.Indeed, the horizon grew dark, and the noise made by the pigeons resembledthat of the terrible Mistral of the Provence, engulfing itself in the gorges of theApennines.When the column of the pigeons passed over my head, I felt a chill, causedall at once by the astonishment and the cold, for this displacement of the air
392 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMproduced a strongly unusual atmospheric draught of air. * * * Thepigeons arrived by millions, precipitating themselves, the ones upon the others,pressed together like the bees in a swarm that escape from the hives in themonth of May. The over-loaded tree-tops of the roosting place broke, and,falling to earth, pulled down at the same time the pigeons and the brancheswhich found themselves below them. The noise was so great that even the nextneighbors could not hear each other if they cried out with all the power oftheir lungs.Sutton (1928) quoting: from an account of a roost by Dr. SamuelP. Bates says : " In the hot summer nights the constant flapping oftheir wings produced by being crowded from their perches, gaveforth a sound not unlike the distant roar of Niagara." John C.French (1919) speaking of a flight from a roost in 1866 in PotterCounty, Pa., says : " Each morning a valley a mile wide between thehills was filled strata above strata, eight courses deep at times, forabout an hour with the multitude of birds flowing westward at therate of a mile a minute, going for food. The roar of wings was likea tornado in the tree tops and the morning was darkened as by aheavy thunder-shower.'1 ' These accounts, coming from different inde-pendent sources, are so similar that we are compelled to believe whatseems to pass belief.Although some of the great flights described above are the migra-tory flights of spring and autumn, as for example those described byHigginson and Wood, many of them are merely flights to and fromthe roosts or nesting grounds to the feeding grounds. The localityof these feeding grounds, as well as the winter residence, dependedon the abundance of mast, which varied from year to year in differ-ent forests. As the mast could not be picked up from the ground ifit were covered with snow, the birds migrated south of the snow line.Thus Mearns (1879) notes that the pigeon was rare in winter in theHudson Valley except in very mild weather when the groundwas bare.Beechnuts were particularly sought by the pigeons. H. J. Jewett(1918) describes as follows the activities of a flock of about 120,feeding on these nuts
:
They lighted in the top of a large beech tree; and, finding the beechnuts hadfallen out of the hulls, dropped in rapid succession from branch to branch tillall had reached the ground. I never have seen more intense activity or seemingsystem in feeding than these birds displayed. They worked in a wing-shapedgroup, moving nervously forward in one direction around the tree, gleaningthe entire nut-eovered space as they went. Those falling to the rear of theflock, where the nuts were picked up. kept flopping across to the front so asto get the advantage of the unpicked ground. A few that wandered apart insearch of scattered nuts kept scurrying about and tilting as they picked themup and then hurried back to the flock as if they feared that the flock would soonbe through feeding and off on the wing. This restless, voracious activity con-tinued till the flock took fright and burst into the air.
PASSENGER PIGEON 393Audubon (1840) thus describes the activities of a larger flock:When alighted, they are seen industriously throwing up the withered leavesin quest of the fallen mast. The rear ranks are continually rising, passingover the main body, and alighting in front, in such rapid succession, that thewhole flock seems still on wing.Herman Behr (1911) speaks of the birds frequenting alder marshesfor food
:
Here they pried under the old leaves, searching for worms or insects, scratch-ing and digging with great energy. Throughout these operations I do notrecall them using their feet once, but always they pried and scratched anddug with their bills.Wallace Craig (1911a) made valuable and interesting studies ofthe expressions of emotion of captive passenger pigeons from whichI have transcribed the following
:
It was eminently a bird of flight ; on the ground it was rather awkward fora pigeon, its legs seeming too short and its massive shoulders too heavy. Thenod of the passenger pigeon was utterly different from that of the mourningdove. The specific manner of nodding seemed an integral part of the bird'sgeneral bearing. The nod consisted of a movement of the head in a circle, back,up, forward, and down, as if the bird were trying to hook its bill over some-thing. Often two or three such nods were given with no pauses between, fol-lowing each other much more rapidly than in the mourning dove, because bodyand tail remained all the while stationary * * *. Ordinary walking paceof the male, 12-13 steps in 5 seconds. In eating, female pecks at rate of12 pecks in 5 seconds on an average, and as head moves through a considerablearc, its motion is very quick.They expressed fear and alarm by beating the wings together inquick succession, making a sound like the rolling beat of a snaredrum.Their flight was direct and made with great velocity. Maynard(1896) says that in twisting and turning they surpassed the Wilson'ssnipe. At times the flocks swept along close to the ground, at othertimes they flew at a great height. Sutton (1928), quoting W. G.Hayes, says : " Usually they flew 10 or 12 feet from the ground. Theyrose in waves to pass over fences and trees, but sometimes they flewfrom 30 to 50 feet in the air without the undulating motion." JamesG. Suthard communicates the following from the notes of Prof. J. J.Glen : " Standing in the open, I watched their flight above in everydirection. They were so close the one to the other that it seemedas if their wings would touch. They were so high above the earththat nothing short of a modern rifle could reach them."Maynard (1896), quoting Edward A. Bowers, describes an im-mense congregation of pigeons at a spring of brackish water inMichigan. "In an incredibly short time the birds begin to come;first in small numbers, then increasing rapidly until, in a few mo-ments, they come in a living avalanche, covering the trees." After
394 BULLETIN" 162, UNITED STATES NATIONAL MUSEUM
a few have cautiously alighted at the mound by the spring, " othersfollow slowly until, at last, a perfect torrent falls upon the spot,covering it so deeply as to endanger the lives of many of them bysuffocation; then the whole enormous body suddenly rises with adeafening roar and alights on the trees. This is repeated until allare satisfied."Voice.?Craig (1911a), as a result of intimate studies of captivebirds, says : " Its voice was loud and strident, the hard notes beingpredominant and the musical notes somewhat degenerated; this beingprobably the result of its living and breeding in colonies so populousthat only the loudest sounds could be heard." He gives at consider'able length and, often with the aid of musical notation, the variouscalls of the bird, which he divides into : " 1. The copulation-note; 2.The keck (a name not used for the note of any other species) ; 3.Scolding, chattering, clucking (these names also peculiar to thespecies) ; 4. The vestigial coo or keeho; 5. The nest-call." The copu-lation note he considers to be essentially the same as that of themourning dove. The keck or kheck is loud and harsh, generallygiven singly, and sometimes it is accompanied with a flapping ofthe wings. The scolding, chattering, and clucking notes have beenwritten kee-kee-kee-kee and tete! tete! and have been said to re-semble the croaking of wood frogs. " In expressing high excite-ment, it becomes loud and high-pitched, and in the excitement offighting especially it becomes very rapid." The vestigial coo or keehowas a mere remnant of the coo of other pigeons. " One sees in this,"says Craig, " probably an adaptation to life in a community sopopulous and hence so noisy that cooing could hardly be heard, andthe pigeon which could best win a female or warn off an interloperwould be the pigeon with the merely loudest voice." The nest callhe describes as " very much blurred?more so than any other noteof this species. A great mixture of high and low tones."Herman Behr (1911) says that feeding birds hailed "newcomerswith a call, peculiar to the occasion. It was a long-drawn and moder-ately loud repetition of one note, which sounded like treet, and thiswould cause the flying birds to alight in nearby trees, giving in theirturn a low call, tret, tret, tret. To me these seemed to be notes ofgreeting, while other sounds were indicative of sex. For instance,the female call note is similar to the treet above, but the male responseis a low oorn, which can not be heard farther than two or three hun-dred feet. My knowledge of these notes is due to the fact that Ilearned to imitate them perfectly, in order to call the birds up withingood shooting distance." Maynard (1896) quotes August Koch asdescribing their note of alarm " sounding something like a laughmade with a child's trumpet."
PASSENGER PIGEON 395Field marks.?The passenger pigeon may easily be confused withthe mourning dove. It is considerably larger, the adult male beingnearly half as long again, but in the female and young, with theirshorter tails, the difference is not so great. As already remarked,and as all ornithologists know, difference in size alone without directcomparison is of little value as a field mark. The best observer,depending on this alone, may easily be mistaken. The passengerpigeon lacks the spot on the neck that is present in adult mourningdoves, but is absent in juveniles. The adult male passenger pigeonhas a much redder breast than the mourning dove. The iris of theadult male passenger pigeon is scarlet, that of the female orange,while the iris of a juvenile bird taken by me on September 4, 1877, isdescribed in my notes as " hazel and gray outside." The iris of themourning dove on the other hand is dark brown. A capital markof the adult male passenger pigeon is its blue rump. To some extentin the female and especially in the young, however, this is obscuredby a brown or olivaceous hue, and is more like the rump of themourning dove. Unlike the mourning dove, the passenger pigeondoes not make a twittering sound with its wings when it rises inflight.It is hardly necessary to consider the band-tailed pigeon, whichmore nearly resembles the domestic pigeon. Its tail is short andsquare. Flocks of this bird in the West have, however, sometimesbeen mistaken for the passenger pigeon.Enemies.?Before the arrival of Europeans in this country preda-tory animals and birds found in the passenger pigeon a large andeasily accessible store of food, and with these the Indian joined inthe feast. Yet, judged from the enormous numbers of pigeons ob-served by the white pioneers, the drain on the pigeons must have beeninsignificant. The white man from the first began destroying thepigeons excessively, and later, with the development of the railroadsand the telegraph and with increasing demands of the markets, thedestruction advanced by leaps and bounds. The pigeon had becomea commercial asset of great value. The Indians, before they werecontaminated by the whites killed no more than they could use them-selves, cooking and eating them, or drying the flesh and trying outthe fat of a moderate number for future use.The following from Kalm (1911) is an interesting reflection onthe difference of treatment of the pigeons by the Indians and so-calledcivilized white man:While these birds are hatching their young, or while the latter are not yetable to fly, the savages or Indians in North America are in the habit of nevershooting or killing them, nor of allowing others to do so, pretending that itwould be a great pity on their young, which would in that case have to starveto death. Some of the Frenchmen told me that they had set out with the inten-
396 BULLETIN 162, UNITED STATES NATIONAL MUSEUMtion of shooting some of them at that season of the year, but that the savageshad at first with kindness endeavored to dissuade them from such a purpose,and later added threats to their entreaties when the latter were of no avail.There are numerous accounts of the slaughter of the pigeons, buta few will suffice. Wilson (1832) described the slaughter in a nest-ing, said to be 40 miles in extent, in Kentucky, as follows
:
As soon as the young were fully grown, and before they left the nests,numerous parties of the inhabitants, from all parts of the adjacent country,came with wagons, axes, beds, cooking utensils, many of them accompaniedby the greater part of their families, and encamped for several days atthis immense nursery. * * * The ground was strewed with broken limbsof trees, eggs, and squab pigeons, which had been precipitated from above,and on which herds of hogs were fattening. Hawks, buzzards, and eagleswere sailing about in great numbers, and seizing the squabs from theirnests at pleasure; while from twenty feet upwards to the tops of the treesthe view through the woods presented a perpetual tumult of crowding andfluttering multitudes of pigeons, their wings roaring like thunder; mingledwith the frequent crash of falling timber ; for now the axe-men were at workcutting down those trees that seemed to be most crowded with nests ; and con-trived to fell them in such a manner that in their descent they might bringdown several others ; by which means the falling of one large tree some-times produced two hundred squabs, little inferior in size to the old ones, andalmost one mass of fat. * * * It was dangerous to walk under the flyingand fluttering millions, from the frequent fall of large branches, broken downby the weight of multitudes above, and which in their descent often destroyednumbers of the birds themselves ; while the clothes of those engaged intraversing the woods were completely covered with the excrements of thepigeons.Wilson also described the destruction of pigeons on migration
:
In the Atlantic states, though they never appear in such unparalleled mul-titudes, they are sometimes very numerous ; and great havoc is then madeamongst them with the gun, the clap-net, and various other implements ofdestruction. As soon as it is ascertained in a town that the pigeons areflying numerously in the neighborhood, the gunners rise en masse; the clap-netsare spread out on suitable situations, commonly on an open height, in an oldbuckwheat field ; four or Ave live pigeons with their eyelids sewed up, arelastened on a moveable stick?a small hut of branches is fitted up for thefowler at the distance of forty or fifty yards; by the pulling of a string, thestick on which the pigeons rest is alternately elevated and depressed, which pro-duces a fluttering of their wings similar to that of birds just alighting; thisbeing perceived by the passing flocks, they descend with great rapidity, andfinding corn, buckwheat, etc., strewed about, begin to feed, and are instantly,by the pulling of a cord, covered with the net. In this manner ten, twenty, andeven thirty dozen have been caught in one sweep. Meantime the air is darkenedwith larger bodies of them moving in various directions; the woods also swarmwith them in search of acorns; and the thundering of musketry is perpetualon all sides from morning to night. Wagon-loads of them are poured intomarket, where they sell from fifty to twenty-five and even twelve cents a dozen.Audubon (1840) describes the killing in a roost in a similar wayand adds iron pots containing burning sulphur, torches of pine knots,and long poles to the instruments of destruction. He says that the
PASSENGER PIGEON 397
next morning " the authors of all this devastation began their entryamongst the dead, the dying, and the mangled. The pigeons werepicked up and piled in heaps until each had as many as they couldpossibly dispose of, when the hogs were let loose to feed on theremainder."John Lewis Childs (1905) says that an old settler in Maine toldhim " that a common way of killing them off was to dig a longtrench in which a quantity of wheat was scattered to attract thebirds. "When they came and settled down to feed, filling the trenchto its utmost capacity, one discharge from some advantageous pointof an old flint-lock musket loaded with a handful of shot wouldoften result in the killing of as many as 75 birds." He adds that
" in those days wild pigeons were hunted for three distinct reasons
?
as sport, as an article of food, and because they were destructive tocrops."Although this early slaughter in many places, largely for indi-vidual use, doubtless reduced the numbers of the passenger pigeonand started it on the road to extinction, the systematic destructionon a large and commercial scale, which began in the forties andreached its highest point late in the sixties and in the seventies ofthe nineteenth century, was responsible for the ultimate result. Thisperfection of extermination was brought about by the increasingdevelopment of the mail and telegraph systems, by which the professional netters, or " pigeoners " as they were called, were informedof and kept in touch with roosting places, in both winter and sum-mer, with flights, and especially with nesting places. A spreadingnetwork of railroads enabled the pigeoner to arrive promptly inthe region of his quarry and to send back to the markets many tonsof the dead bodies of this beautiful bird, as well as thousands ofliving ones to be used in clubs for trap shooting. The decreasingarea of forest suitable for the pigeons tended to concentrate theremainder and render it easier to locate them.Prof. H. B. Roney (1879) estimated that at the time of his writ-ing there were about 5,000 men in the United States who pursuedpigeons year after year as a business. These men had perfectedthe methods of netting, which was generally done at a bed of muckbaited with salt and sulphur, or grain. The net, about 6 feet wideand 20 to 30 feet long, could be sprung by means of ropes and apowerful spring pole from a blind by the operator, and " fliers " (cap-tive birds thrown up into the air) or " stool pigeons " (birds tied toa pole that could be suddenly raised and lowered by a cord, makingthe birds flutter as if alighting) were used to lure down the birdsflying overhead. Professor Roney found that 60 to 90 dozen birds
398 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
a day to the net were a fair average of the numbers caught. Headds
:
Higher figures than these are often reached, as in the case of one trapperwho caught and delivered 2,000 dozen pigeons in ten days, being 200 dozen,or about 2,500 birds per day. A double net has been known to catch as highas 1,332 birds at a single throw, while at natural salt licks, their favoriteresort, 300 and 400 dozen, or about 5,000 birds have been caught in a singleday by one net.At this time, 1879, dead birds were sold in Chicago markets for50 to 60 cents a dozen, 35 to 40 cents at the nestings, while live birdsbrought $1 to $2 a dozen, and Professor Roney estimated that thepigeoner made at these times $10 to $40 a day. As long as moneywas obtained by the sale of the birds, men were ready to slaughterthem. Mershon (1907), in his book on the passenger pigeon, givesa large number of statistics and estimates of the immense slaughter ofthis bird.The last great nesting of this beautiful bird to be desecrated was in1878 at Petoskey, Mich. Here the persecuted birds had gathered inold-time numbers, driven out from roosts and nestings in other places.The nesting was said to have been 28 to 40 miles in length and 3 to 10miles in width. Professor Roney (1879), who visited the nesting atthe time of the slaughter, thus describes it
:
Scarcely a tree could be seen but contained from 5 to 50 nests, according to itssize and branches. Directed by the noise of chopping and of falling trees, wefollowed on, and soon came upon the scene of action. Here was a large forceof Indians and boys at work, slashing down the timber and seizing the youngbirds as they fluttered from the nest. As soon as caught, the heads were jerkedoff from the tender bodies with the hand, and the dead birds tossed into heaps.Others knocked the young fledglings out of the nests with long poles.The Indians were paid a cent apiece for the slain. "Within 100rods of the nests, instead of 2 miles away as the law prescribed, netterswere hard at work, one taking 984 birds in a single day. He sawmen going about close to the nesting, shooting the birds as theyroosted in rows on the branches and passed in clouds overhead.Scores of dead pigeons were left on the ground to decay, and thewoods were full of wounded ones, while many of the squabs, deprivedof their parents, starved to death. It is a terrible picture of crueltyand greed.Roney (1879) adds:For many weeks the railroad shipments averaged fifty barrels of dead birdsper day?thirty to forty dozen old birds and about fifty dozen squabs beingpacked in a barrel. Allowing 500 birds to a barrel, and averaging the entireshipments for the season at 25 barrels per day, we find the railroad shipmentsto have been 12,500 dead birds daily, or 1,500,000 for the summer. Of live birdsthere were shipped 1,116 crates, six dozen per crate, or 80,352 birds. Thesewere railroad shipments only and not including the cargoes by steamer from
PASSENGER PIGEON 399Petoskey, Cheboygan, Cross Village, and other like ports, which were as manymore. Added to this were the daily express shipments in bags and boxes, thewagon loads hauled away by the shotgun brigade, and the myriads of squabsdead on the nest.The profit from these transactions was the lure that led to theannihilation of the passenger pigeon. Barrows (1912) says:Dr. Isaac Voorheis, of Frankfort, Michigan, told the writer personally that in1880 or 1881, when there was a large nesting in Benzie County, he took at onethrow the net 109 dozen and 8 pigeons (1,316 birds), and that six catches ofthe net brought him $650. These birds were kept alive until a schooner loadwas obtained, when they were sent directly to Chicago for trap shooting.A provision dealer at Cheboygan, Mich., is said to have shippedlive pigeons in numbers up to 175,000 a year from 1864 until the end.Kumlien and Hollister (1903) say of Wisconsin:Mr. J. M. Blackford, now residing at Delaware, states the last largecatch of nesters was in 1882. The following spring but 138 dozen were takenin the best pigeon ground in the State, and this was practically the end.Kuthven Deane (1896) wrote in 1895 to N. W. Judy & Co., of St.Louis, Mo., dealers in poultry and the largest receivers of game inthat section, as to their dealings in passenger pigeons and theyreplied : " We have had no wild pigeons for two seasons ; the lastwe received were from Siloam Springs, Ark. We have lost alltrack of them and our netters are lying idle."This firm, Widmann (1907) says, "handled more dead and livepigeons than any other firm in the country, and had their nettersemployed all the year around, tracing the pigeons to Michigan andWisconsin in spring and to the Indian Territory and the south inwinter."This is the last word on the commercial extinction of the passengerpigeon by its greatest of all enemies, so-called civilized man.DISTRIBUTIONRange.?Eastern North America, casual in Bermuda, the WestIndies, and in Mexico.Breeding range.?The breeding range of the passenger pigeon ex-tended north to Montana (" latitude 49?," Great Falls, and theYellowstone River) ; North Dakota (Fort Berthold) ; Manitoba(Waterhen River) ; Minnesota (Lake Itasca and Northern PacificJunction) ; Wisconsin (Stockholm, Shiocton, and West Depere)
;
Michigan (Petoskey, Brant, and Detroit) ; Ontario (Moose Fac-tory) ; Quebec (Fort George and Quebec) ; New Brunswick (GrandFalls); Prince Edward Island; and probably Newfoundland (St.Johns). East to probably Newfoundland (St. Johns) ; Nova Scotia(Halifax and Yarmouth) ; New Hampshire (Conway and Webster)
400 BULLETIN \(i-2, UNITED STATES NATIONAL MUSEUMMassachusetts (Plymouth) ; Connecticut (Portland) ; southeasternNew York (Croton Falls) ; southeastern Pennsylvania (Columbia) ;
-a-ftel-northeastern Virginia (Clarendon). South to northeasternVirginia (Clarendon); northwestern West Virginia (MarionCounty) ; Kentucky (Green County) ; Illinois (Mount Carmel, Hills-boro, and Quincy) ; and eastern Kansas (Neosho Valley). Westto eastern Kansas (Neosho Valley) ; South Dakota (Vermilion andFort Pierre) ; and Montana (Hellgate River, Fort Shaw, and
"latitude 49?").Passenger pigeons also were frequently recorded in summer atpoints well outside the breeding range above outlined. Amongthese records are Mackenzie (Fort Good Hope, by AlexanderMackenzie in 1789; Fort Norman, by Ross; and Fort Simpson, b}7Kennicott) ; Saskatchewan (He a la Crosse, by Hood in 1820; nearthe mouth of the South Saskatchewan River, August 2 to 5, 1858;and Fort Qu' Appelle) ; northern Manitoba (Island Lake, wherein 1903 E. A. Preble received a report of a small flock seen three orfour years previously; York Factory; and Fort Churchill, wheretwo specimens were taken) ; Quebec (Tadousac, one recorded byFleming as taken July 20, 1889; Godbout, one taken June 27, 1859;and Anticosti Island, one record) ; Franklin (one recorded by J. C.Ross as coming aboard their vessel in Baffin Bay at latitude 73%?N.) ; northwestern South Carolina (Caesars Head, two pairs seenby Arthur T. Wayne in the summer of 1882) ; Missouri (New Haven,large flocks observed in 1872 by Dr. A. F. Eimbeck) ; Wyoming (onespecimen taken 40 miles west of Fort Laramie, September 8, 1857) ;and Nevada (West Humboldt Mountains, in September, 1867,according to Ridgway [1873]). Passenger pigeons also have beenrecorded from British Columbia, but the evidence seems rather un-satisfactory {cf. Brooks and Swarth, 1925).Winter range.?The vast numbers of these birds and the extensivearea included in their breeding grounds considered, the winter rangewas greatly restricted. Data bearing upon this phase of the sub-ject, however, are not plentiful. Apparently at this season, thespecies extended north regularly to Arkansas (Rogers) ; southeast-ern Missouri (Attie) ; and northern South Carolina (ChesterCounty). East to South Carolina (Chester County and SineathsStation); and Florida (Amelia Island and Gainesville). Southto Florida (Gainesville) ; Alabama (Greensboro) ; and Louisiana(Mandeville). "West to Louisiana (Mandeville) ; and Arkansas(Judsonia, Fayette, Siloam Springs, and Rogers). In some seasonslarge numbers wintered much farther north as they were recordedas abundant in the winter of 1853-54 at Brookville, Ind. : a flockof about 300 was seen near Harrisburg, Pa., December 25, 1889 ; and
PASSENGER PIGEON 401some wintered near Hartford, Conn., in 1882-83. At this season,they also were recorded at more southern points: Cuba (a femaletaken at Triscornia on Havana Bay, and a male obtained in theHavana market, both specimens being preserved in the MuseoGundlach) ; while four specimens were taken at Jalapa, Vera Cruz.in the winter of 1872-73, and it was reported by Herrera as an occa-sional visitant to the Valley of Mexico (no details). A migrantspecimen was killed in Bermuda on October 24, 1863.Migration.?Despite a rather voluminous literature, actual usefuldata bearing upon the seasonal movements of the passenger pigeonare decidedly scarce. The modern method of recording arrival anddeparture was not developed until the species was so reduced innumbers that its ultimate extinction was already apparent. Forthis reason, many of the dates of later years have no significanceas bearing upon migration, and wherever possible data obtainedprior to 1890 have been used.Spring migration.?Early dates of spring arrival are : NorthCarolina, Raleigh, April 18; Virginia, Highgate, April 20; WestVirginia, French Creek, February 27 and April 10, and Fairview,March 19; District of Columbia (occasionally wintered), Washing-ton, April 3; Pennsylvania (occasionally wintered), Chambersburg,February 13, Erie, March 13, Brockney, March 18, Guths Station,March 20, and Ridgway, March 22 ; New Jersey, Raccoon, March 3,Plainfield, March 12, and Caldwell, March 31; New York (rarelywintered). Locust Grove, March 3, Cornwall-on-Hudson, March 4,Elmira, March 7, Painted Post, March 16, Glasco, March 20, andBuffalo, March 27; Connecticut, Gaylordsville, March 13, and Say-brook, March 21; Massachusetts, Woods Hole, March 20, Ponkapog,March 31, and Amherst, April 7 ; Vermont, Tydeville, April 9
;
New Hampshire, Hollis, April 2; Maine, North Livermore, April28; Illinois (occasionally wintered in southern part), Charleston,February 3, Virden, February 11, Lake Forest, March 4, and Car-thage, March 12; Indiana, Millwood, February 18, Brookville, Feb-ruary 28, Jonesboro, March 1, Brown County, March 7, and Kokomo,March 13; Ohio, Hudson, March 8, West Liberty, March 10, andFayette, March 15; Michigan, Petersburg, March 20, Locke, March22, Battle Creek, March 27, and Plymouth, April 7; Ontario,Kingston, March 21, London, March 24, Port Rowan, March 26,and Toronto, April 13; Iowa, Iowa City, March 13, Burlington,March 15, La Porte, March 16, and Dubuque, March 22 ; Wisconsin,Ripon, March 8, Janesville, March 23, Racine, March 25, Shiocton,March 29, Delavan, April 1, and Green Bay, April 7; Minnesota,Lanesboro, March 22. Bradford, April 1, Minneapolis, April 3, Zum-brota, April 8, and Lake Andrew, April 9; and Manitoba, Aweme,
402 BULLETIN 162, UNITED STATES NATIONAL MUSEUMApril 8, Lake Winnipegosis, April 14, Greenridge, April 17, andOssowo, April 18.Fall migration. ? Late dates of fall departure are: Manitoba,Mount Royal, September 15, and Aweme, September 21 ; Minnesota,Lake Andrews, September 28, Lanesboro, October 5, and Zumbrota,November 15; Wisconsin, Delavan, September 8, and Kelley Brook,September 16; Iowa, Williamstown, September 27, and Keokuk,October 18; Ontario, Ottawa, September 3, and Toronto, October 22;Michigan, Ann Arbor, October 12, and Newberry, October 24;Ohio, Wayne County, September 19 ; Indiana (probably occasionallywintered), Bloomington, September 28; Illinois (occasionally win-tered in southern part), Virden, October 5; Kentucky, Casky, Octo-ber 30; Quebec, Montreal, September 15, and Valley River Rouge,October 7; New Hampshire, Acworth, October 10; Massachusetts(rarely wintered), Worcester, September 25, and Plymouth, October16; Connecticut (rarely wintered), East Hartford, October 19; RhodeIsland, Newport, October 19 ; New York, Ossining, October 1, CrotonFalls, October 16, Locust Valley, November 4, and Orange Lake,November 17; New Jersey, Morristown, September 16, and NewProvidence, November 12; Pennsylvania (occasionally wintered),Linden, October 5, Monroe County, October 25, and Wayne County,November 2; Maryland (occasionally wintered), Laurel, October 11;Virginia, Dunn Loring, October 19; and West Virginia, FrenchCreek, October 20.Egg dates.?Saskatchewan and Manitoba: 6 records, May 21 toJune 23. Minnesota and Wisconsin : 11 records, May 5 to September10 ; 6 records, May 20 to 30.ZENAIDURA MACROURA CAROLINENSIS (Linnaeus)EASTERN MOURNING DOVEHABITSContributed by Winsor Mabeett TylebThe mourning dove must have been one of the first birds thatattracted the attention of the early settlers when this country wasnew and wild. They must have recognized the bird as not farremoved from some of the Old-World species of pigeons, and itsnotes must have recalled to them their old home. The writers of thesetimes speak of the bird familiarly, especially as a game bird thatrelieved the hardships of pioneer life.At the present time, in the Northern States, protected as a songbird, it adds a quiet dignity to our avifauna, while in the SouthernStates it is a common, tame, almost a dooryard bird and a gleanerof fields, except when, during the hunting season, it is shot for food
EASTERN MOURNING DOVE 403and sport. In the West it is an inhabitant both of the plains andthe mountains, ranging commonly to 7,000 feet altitude.And yet, well known and widely distributed as the bird is, it isnot a conspicuous bird of the country at all. It is quiet in voice,neutral in color, and so unobtrusive in deportment that it seemslittle more than a part of the background; a quiet, pastoral bird,reminding us of the man in " The Bab Ballads "?" no characteristictrait had he of any distinctive kind"?or of sweet, lovable, butwholly negative Hero in " Much Ado about Nothing."Spring.?In the parts of the country where the mourning dovespends the winter, one of the early signs of spring is when the winterflocks begin to break up and the doves separate into mated pairs.Just as the mockingbird in the Southern States bursts suddenly intosong and separates winter from spring, so the male mourning dove,who has been silent through the winter, at the first hint of springbegins to coo.As the breeding season approaches, the birds become graduallytamer and, as Wilson (1832) says, they " are often seen in thefarmer's yard before the door, the stable, barn, and other outhouses,in search of food, seeming little inferior in familiarity, at such times,to the domestic pigeon "?a contrast to the wild game bird of theautumn.Courtship.?Very little has been published on the courting actionsof the mourning dove, and apparently no detailed study has beenmade of them. Indeed, many observers who know the bird wellstate that they have seen no courting at all.Barrows (1912), who gives a careful description of a nuptial flight,points out that " although familiar with the mourning clove's habitsin New England, western New York, and elsewhere we have neverseen this peculiar flight except in Michigan." He says
:
An individual leaves its perch on a tree, and, with vigorous and sometimesnoisy flapping (the wings seeming to strike each other above the back), risesobliquely to a height of a hundred feet or more, and then, on widely extendedand motionless wings, glides back earthward in one or more sweeping curves.Usually the wings, during the gliding flight, are carried somewhat below theplane of the body, in the manner of a soaring yellowlegs or sandpiper, andsometimes the bird makes a complete circle or spiral before again flappingits wings, which it does just before alighting. * * * This peculiar evolu-tion is commonly repeated several times at intervals of two or three minutes,and appears to be a display flight for the benefit of its mate, the assumptionbeing that only the male dove soars.Goss (1891) speaks of the courtship thus:During the pairing season the male often circles and sails above his mate,with tail expanded, and upon the ground struts about with nodding head, andfeathers spread in a graceful manner.
404 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMCraig (1911), speaking of the "nest-calling attitude," calls atten-tion to the display of the ornamented tail. He says
:
The male [sits] with his body tilted forward, tail pointing up at a highangle, the head so low that bill and crop may rest on the floor, or if the birdbe in the nest, the head is down in the hollow. Both the voice and the attitudeof the male servo to attract the female, for in all pigeons Hie nest-call isaccompanied by a gentle flipping of the wings, ogling eyes, and a seductiveturning of the head. In addition to these general columbine gestures, Zenai-dura has a special bit of display of his own, for during the first note of thenest-call he spreads his tail just enough to show conspicuously the white markson the outer feathers ; soon as this first note is past, the tail closes and thewhite marks disappear, to flash out again only with the next repetition of thenest-call, before which there is always a considerable interval.Forbush (1927) says that " in courtship the male mourning dovesometimes strikes his feet hard on his perch one after another."James G. Suthard, speaking of the bird in Kentucky, says in hisnotes
:
During the nesting season, the female acts very much like the tame pigeon.The male prances around with his neck feathers all ruffed up, cooing andbilling with the female. I have noticed that he sometimes picks up pieces ofgrass in his courtship antics. The intrusion of another male on one of thesescenes results in a fight whereupon the female usually disappears.Nesting.?The mourning dove uses a very wide choice in selectinga site for its nest. Perhaps the site most nearly typical is not farfrom the trunk on a horizontal branch of an evergreen tree?a pineor cedar?affording a firm foundation for the flimsy nest. The birdfrequently nests on the ground, however, even on a clump of grass,sometimes on the stump of a tree, and there are several recordedinstances where the nest has been found placed on a wooden ledgeattached to an inhabited building. Indeed Gardner (1927) says thatthe birds in Kansas ''preferred the vicinity of buildings to thewooded and secluded canyons of the back country by a ratio of atleast ten to one."The chief requisite, apparently, is a level support that will givestability to the nest, and to acquire this security the dove often makesuse of the experience of another species of bird and builds its owmnest on a nest (for example, that of a robin, brown thrasher, ormockingbird) that has weathered the previous winter.Bendire (1892) cites an extreme instance of this habit. QuotingJ. L. Davidson from Forest and Stream, he says
:
I found a black-billed cuckoo and a mourning dove sitting together on arobin's nest. The cuckoo was the first to leave the nest. On securing thenest I found it contained two eggs of the cuckoo, two of the mourning dove, andone robin's egg. The robin had not quite finished the nest when the cuckoo tookpossession of it and filled it nearly full of rootlets, but the robin got in andlaid one egg.
EASTERN MOURNING DOVE 405As a rule a pair of mourning doves, in contrast to the habit ofthe passenger pigeon, nests well removed from the nests of otherdoves, but Charles E. Stockard (1905) reports in Mississippi aninteresting exception to this rule. " Doves," he says, " often nestedin small colonies. In a clump of about fifteen young pine trees Ionce found nine nests, and in an Osage orange hedge about one-halfmile long twelve nests were located. But most doves nest singly, orwith the nests too far apart to suggest any gregarious nesting habit."Most commonly the nest is made of sticks and is lined with finertwigs. A. D. DuBois, however, in his notes records the use of grass,weed stalks, roots, and a lining of leaves and mentions one nestmade " almost entirely of rootlets and stems lined with finer root-lets (a shallow affair)."The nest, oftenest, perhaps, just a platform of sticks, but firmenough to withstand usage for 30 days, is made apparently entirelyby the female bird. Frank F. Gander in his notes states this to bea fact in the case he describes, and he demonstrates the aid thatthe male bird gives to his mate. He says
:
The bringing of the material was accomplished by the male, who flew tothe ground and searched about until a suitable stick was found. In selectingmaterial the male was very careful and tested the sticks by shaking themvigorously. Perhaps this was as much to test his hold upon the stick as totest the stick itself, as many times sticks were shaken from his beak. Somuch time was consumed in this choosing of a twig that his trips to thenest averaged about one every two minutes. He always approached the nestby the same route, alighting upon a protruding branch, hopping from this toanother, and walking along the latter to the nest. Reaching the nest, heturned the material over to the female, who reached up her beak to receiveit. Sticks were frequently dropped during this exchange. The female placedthe sticks under and about her to construct the nest.Building did not continue uninterruptedly, as the female frequently leftthe nest when the male would pursue her and peck at her until she returned.Work for the day was stopped at about 11 a. m. The nest building was takenup again on the following morning and carried on until about 10 a. m.Continuing, he shows the division of labor during the incubationperiod of 15 days:The male took up his duties at the nest at about 10 in the morning and wasrelieved again at about 3 in the afternoon. The male often left the eggs un-guarded for a few minutes about noon while he flew to a near-by wateringplace to drink.Margaret M. Nice (1922), referring to the building of the nest,corroborates the observation quoted above. She says:Nest building as a rule takes place in the early morning. The male mourn-ing dove gathers the materials and carries them to his mate who arranges them.He takes one piece at a time, and if he happens to drop it. he does not stop butcontinues his journey to the tree and then starts over again,74564?32 27
406 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMMrs. Nice's articles form an exhaustive study of the nesting habitsof the mourning dove and contain many statistical data of the utmostinterest. Readers are referred to these valuable articles for detailedinformation.That the division of labor, with a well-ordered time for relievingeach other on the nest, continues through the incubation period andduring the 15 days that the young spend in the nest, is shown bythe following extracts. Wallace Craig (1911) says:Male and female take regular daily turns in sitting on the eggs or young:the female sits from evening till morning, the male from morning till evening,the exchanges taking place usually about 8.30 a. m. and 4.30 p. m. Thisarrangement is very regular if there is nothing to disturb the birds ; but ifinterloping birds come about, this arouses the anger of the male and he leavesthe nest in order to attack them.Mrs. Nice's (1923) experience corroborates Doctor Craig's observa-tion. She says:The male incubates and broods during the day while the female does thesame during the night, early morning, and late afternoon ; and both parentsregurgitate " pigeon milk " for the young. Two striking differences betweenthe nest behavior of mourning doves and most passerine birds is the almostconstant brooding of the young till near the end of the nest life and the lackof any sanitary care of the nest.And further (1922)
:
As a rule one or the other parent is continuously on the nest from the timethe first egg is laid until the young are fairly well grown.I have approached the nest of a mourning dove and come almostwithin arm's reach of the bird before it flew quietly away, but thereis plenty of evidence that this behavior is not invariable for fre-quently the bird is reported to flop from the nest and resort to theruse of the broken wing.The breeding season is very long; in the Middle States it lastsfrom May to August and rarely to early September. The birdscommonly rear two broods in a season, and Miss A. R. Shermanbelieves that they probably rear three sometimes. In her notes MissSherman says in substance
:
The doves are so numerous and so secretive in their ways that it is not possibleto say whether a pair of birds, which has nested in May or June, breed againlate in June or in July. When a nest is used twice in the same season,however, the assumption is that a pair of birds is using their own nest asecond time.[Author's note : On April 19, 1907, while hunting for hawks' nestsin a grove of tall pines, I rapped on a tree containing a likely lookingold nest and was surprised to see a mourning dove fly from it. Iclimbed to it, more than 40 feet up in a white pine, and found it tobe an old hawk's nest, that had since been used by squirrels, as it
EASTERN MOURNING DOVE 407was full of pine needles and soft rubbish, such as squirrels use.It was quite a large nest, measuring 25 by 15 inches. The doves hadscraped out a hollow in the pine needles and laid one egg. I visitedthe nest a^ ain on April 28, when the dove flew off, as before, and thenest held two eggs. I photographed the nest and collected the eggs.As a pair of great-horned owls and a pair of Cooper's hawks werenesting in these woods, the doves stood a poor chance of raising abrood, or even escaping with their own lives.Eggs.?The mourning dove lays almost always two eggs, but thereare a few records of three, or even four. In shape they vary fromelliptical oval, the commonest shape, to elliptical ovate or ovate. Theshell is smooth with very little gloss. The color is pure white. Themeasurements of 47 eggs average 28.4 by 21.5 millimeters; the eggsshowing the four extremes measure 31 by 22, 29.5 by 23, 26 by 20.5,and 28.5 by 20 millimeters.]Young.?The young of the mourning dove are helpless whenhatched, and during the two weeks they remain in the nest the}rrequire constant care from their parents. They are fed by regurgi-tation during most of their nest life, but solid food, such as insectsand seeds, is gradually substituted, and at the time of leaving thenest it largely replaces the pigeon milk." The contents of the cropof a young bird, examined at the end of its nest life, consisted almostentirely of seeds (principally grass seeds) and less than 2 per cent
"pigeon milk." [See Townsend (1906).]Gabrielson (1922), who studied the nest life from a blind, clearlydescribes the process of regurgitation thus
:
At 7.30 a. in. a squab backed toward the blind and getting from beneath theparent raised its head and mutely begged for food. The adult (presumablythe female) responded immediately by opening her beak and allowing the nest-ling to thrust its beak into one corner of her mouth. She then shut her beakon that of the nestling and after remaining motionless for a short time begana slow pumping motion of the head. The muscles of her throat could be seento twitch violently at intervals, continuing about a minute, when the nestlingwithdrew its beak. The other nestling then inserted its beak and the processwas repeated, 15 seconds elapsing before its beak was removed. With inter-vals varying from 5 to 10 seconds (watch in hand) four such feedings, two toeach nestling, occurred. The nestling not being fed was continually tryingto insert its beak in that of the parent and at the fifth feeding both succeededin accomplishing this at the same time. The nestlings' beaks were inserted fromopposite sides of the parent's mouth and remained in place during the feedingoperation although I could not say whether or not both received food. Whilebeing fed the nestlings frequently jerked the head from side to side and alsofollowed the motion of the parent's beak by raising and lowering themselves bythe use of the legs. They were not more than five days old but had better useof their muscles than the young of passerine birds at from eight to ten days ofage. The entire process described above occupied about six minutes, afterwhich the nestlings crawled back beneath the parent.
408 Bl\LLETIN 10 2, UNITED STATES NATIONAL MUSEUMMiss A. R. Sherman, who has had ample opportunity to study themourning dove and a wide experience as a field observer, gives in hernotes the period of incubation definitely as 15 days.Plumages.? [Author's note: The young squab is fat and unat-tractive, scantily covered with short, white down, through whichthe }rellowish skin shows. The stiff quills of the juvenal plumagesoon appear, giving the young bird an ugly, spiny appearance. Thejuvenal plumage is well developed before the young birds leave thewell-filled nest. In this plumage the upper parts are " buffy brown "to " snuff brown," with faint, whitish edgings on the back and wingcoverts; the scapulars and some of the inner wing coverts have largeblack patches; the underparts are from" pinkish cinnamon " to " lightvinaceous-cinnamon," paler on the belly and grayer on the flanks.A postjuvenal molt of the contour plumage and tail, during fall,produces a first winter plumage, which is like the adult but somewhatduller. Adults have a complete molt in fall.]Food.?Adult mourning doves are essentially seedeaters. Wheatand buckwheat are said to be their favorite grains, but they consumesuch enormous numbers of weed seeds that they prove to be a highlybeneficial species, as the following quotation from Dutcher (1903)shows
:
The examination of the contents of 237 stomachs of the dove shows over 99Tier cent of its food consists wholly of vegetable matter in the shape of seeds,less than 1 per cent being animal food. Wheat, oats, rye, corn, barley, andbuckwheat were found in 150 of the stomachs, and constituted 32 per cent of thetotal food. However, three-fourths of this amount was waste grain pickedup in the fields after the harvesting was over. Of the various grains eaten,wheat is the favorite, and is almost the only one taken when it is in good condi-tion, and most of this was eaten in the months of July and August. Corn,the second in amount, was all old damaged grain taken from the fields afterthe harvest, or from roads or stock yards in summer. The principal and almostconstant diet, however, is the seeds of weeds. These are eaten at all seasonsof the near. In one stomach were found 7,500 seeds of the yellow wood-sorrel(Oxalis strieta), in another, 6,400 seeds of barn grass or fox tail (Chaetocloa)
.
Wayne (1910) records this interesting observation on the dove'sfeeding habits
:
Although this species is supposed to feed upon the ground, this is by no meansalways the case as the birds resort to the pine woods for weeks at a timeto feed upon the seeds of these trees, which they obtain by walking out on thelimbs and extracting them from the cones. The flesh at this time is verystrongly impregnated with a piney flavor.Behavior.?Although mourning doves spend a large part of theyear in flocks, they have a strong tendency in spring to separate intopairs and scatter over the country to nest. Doubtless they owe theirpresent status, perhaps even their existence, to this habit, for, hadthey bred in colonies as the passenger pigeon did, the doves would
EASTERN MOURNING DOVE 409have been subjected at their nests to the wholesale slaughter thatexterminated the pigeon.As we watch a number of doves feeding in a stubble field we soonsee that there is no very strong tie binding together the members ofthe company?no such bond as holds together a flock of sandpipersand suppresses individual action. The doves are spread out over theground, each walking off by itself and feeding more or less alone, likegrazing cattle. When we walk toward them they start into the air,but not all together ; a few, very often only two, fly away ; then, aftera moment, a few more take flight and go off, very likely in anotherdirection. The flock when alarmed, instead of moving off as a unit,breaks up, and the birds retreat individually or in pairs. Thus evenwhen the doves are assembled in numbers there is a tendency tosegregate into pairs?a characteristic of the breeding season.The birds leave the ground very quickly, gaining speed rapidlywith strong, sweeping wing beats and fly with whistling wings,suggesting the whistling flight of the golden-eye.In eastern Massachusetts, where since 1910 the birds have becomewell established, they frequent the dry, sandy, sparsely wooded hill-sides characteristic of this glaciated countiy, and retire to nest in thenear-by pine woods, where they seem much at home, walking easilyamong the branches.Doves often visit gravelly roads and are sometimes seen on the seabeaches. On the dry plains of western Texas (Merriam, 1888) theywere found 3 to 5 miles from the nearest water, and Merriam (1890)describes thus the coming of the doves at dusk to drink
:
Common from the Desert of the Little Colorado to the upper limit of thepine belt. Every evening they assemble at the springs and water holes, com-ing in greatest numbers just at dark, particularly about the borders of theDesert where water is very scarce. On the evening of August 20 we campedfor the night at a small spring about 5 miles west of Grand Falls. At duskhundreds of doves came to drink, and continued coming until it was so darkthat they could not be seen.Voice.?The mourning dove takes its name from its commonnote, a low-toned, moaning coo. This is one of the bird notes that,while fairly loud and perfectly distinct, does not readily attract theattention of one who is not familiar with it. In this respect itresembles the diurnal hooting of the screech owl; both of thesenotes in some strange way are disregarded by the ear until it istrained to detect them. We then recognize them both as familiarsounds of the countryside.A. A. Saunders in his notes describes a typical song thus
:
The sound is well imitated by a low-pitched whistle, but some birds strikenotes lower than I can whistle. The song consists of four notes. The firstis usually twice as long in time as the others, and slurred first upward, and
410 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthen downward. The other three notes are either all on one pitch or slightlyslurred downward. Lengths of songs vary from 3 to 4% seconds. Usually onebird sings a single song and repeats it over and over in just the same manner
;
but I once recorded five different songs from a single bird.Craig (1911) has reported the results of an exhaustive study ofthe bird in confinement. The following are the outstanding featuresof his article. He describes the song, which he terms the " perchcoo " essentially as Saunders does, and adds
:
When delivering his song, the mourning dove does not perform any danceor gesture, as some birds do. He invariably stands still when cooing; evenwhen he coos in the midst of pursuing the female he stops in the chase, standsimmovable until the coo is completed, and then runs on. His attitude is,to be sure, very definite, the neck somewhat arched and the whole body rigidbut the impression it gives one is, not that the bird is striking an attitude, butthat he is simply holding every muscle tense in the effort of a difficultperformance.The female also utters the perch-coo, though less often than the male, andin a thin, weak voice and staccato tones, which, as compared with the male'ssong, form so ludicrous a caricature that on first hearing it I burst outlaughing.To this commonly heard note he adds two others; the nest callof which he says : " This call is much shorter than the song, andmuch fainter, so that the field observer may fail ever to hear it. Itstypical form is of three notes, a low, a high, and a low, thus some-what resembling the first bar of the song, but differing in that thethree notes do not glide into one another, there being a clear breakfrom each note to the next " ; and the copulation note, which " isgiven by both the male and female, immediately after coition; inthe mourning dove it is a faint growling note, repeated two to fourtimes with rests between. So far as I have seen, the mourningdoves, throughout the utterance of these sounds, keep the bill wideopen."Field marks.?The mourning dove in the field appears as a long,slim, gray-brown bird with small, nodding head, whistling wings, andlong, pointed tail. The sparrow hawk resembles the dove very closelyin flight, but it has strong, heavy shoulders, a larger head, andsquarely tipped tail. The little ground dove of the Southern Statesis instantly distinguished from the mourning dove by its stumpytail and the flash of bright color under the wing.Enemies.?Harold C. Bryant (1926) speaks thus of the dove'senemies
:
Apart from man, the dove has other enemies. The duck hawk is swiftenough to overtake the dove, and this bird is probably the dove's most dreadedavian foe. Other predatory species take a toll during the nesting season.Its rapid flight frequently brings the dove in violent contact with telephonewires, and many birds die annually from this cause. Rodent-poisoning opera-tions have in recent years been responsible for the death of many doves; for,
EASTERN- MOURlSriN-G DOVE 411
unlike the quail, the mourning dove and the band-tailed pigeon are both sus-ceptible to strychnine.C. S. Thompson (1901) notes a very long tapeworm wound roundand round in the intestines of an emaciated bird, and Lloyd (1887),writing of the bird in western Texas, speaks of owls as enemies inwinter, " when they frequently change their roosting place, as afriend (Mr. Loomis) suggests, in consequence of being disturbed bythe numerous owls."The cowbird not infrequently selects the dove as a host for itsyoung.Game.?Bryant (1926) shows why, aside from its desirability as atable delicacy, the dove is a popular game bird, affording a rapidlymoving target that demands the utmost skill on the part of thehunter. He says
:
Unless favorably located near a watering place, one bird in three or fourshots makes a good average for all but the most experienced hunter. Thesmall size and great speed make the bird a difficult target. The variety ofshots possible is almost endless. Quartering and side shots are most difficultbecause of the speed of the birds in flight. Then come shots at towering ordescending birds, often dependent on whether they are coming or going. Theeasier straight-away shots are to be expected less often in dove shooting thanin quail shooting.Thus it will be seen that dove hunting gives the best of practice to the loverof wing shooting. No finer test of skill is afforded unless it be in snipeshooting.Fall and winter.?In regions where mourning doves are common,they begin to resume their gregarious habits soon after the breedingseason is over.J. A. Spurrell (1917), speaking of the bird in Iowa, says:From the latter part of July until the doves depart on their fall migration inlate October they select common roosting places, one of which happens to beour orchard. Toward sunset the doves visit some place to drink and then flyto the roosting place from all directions until between five and six hundred areroosting there. They depart again just as it becomes light in the morning,spending the day far away in pastures and grain fields. During the month ofAugust they may be commonly found about salt troughs for cattle, seeming toeat the salt.Stockard (1905) speaks thus of the dove's habit of roosting inMississippi, where they remain during the winter
:
Late in summer they begin roosting in company, and many hundreds comeabout sunset to their chosen place for the night. During this season they areshot in large numbers while flying to the hedge or small wood that has beenselected as a roosting place.Throughout the winter in the Southern States we see the doves, incompanies of a dozen or more, feeding quietly in the stubble and peafields, from which, as we approach, they flush rather wildly and,
412 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
scattering, retreat in twos and threes beyond the surrounding pinetrees.[Author's note: A comprehensive and interesting study of thefall migration of the mourning clove has been published recently byWilliam B. Taber, jr. (1930), to which the reader is referred. F. 0.Lincoln (1930) proves the migratory status of the mourning dovesfrom evidence obtained from returns from banded birds.]DISTRIBUTIONRange.?North America, Central America, and the West Indies.Breeding range.?The mourning dove breeds north to BritishColumbia (Chilliwack and Okanagan Landing) ; Saskatchewan(Qu'Appelle) ; Manitoba (Aweme, Portage la Prairie, and Winni-peg) ; Michigan (Newberry, Mackinac Island, Hillman, and Zion) ;Ontario (Sarnia, Plover Mills, Guelph, Toronto, and Brighton) ;Vermont (Burlington) ; and New Hampshire (Concord and Hamp-ton Falls). East to New Hampshire (Hampton Falls and Sea-brook) ; Massachusetts (Boxford, Boston, North Truro, and WoodsHole) ; New York (Shelter Island and Yaphank) ; New Jersey (RedBank, Spring Lake, and Sea Isle City) ; Delaware (Lincoln) ; Mary-land (Cambridge) ; Virginia (Wallops Island and Spottsville) ;North Carolina (Beaufort and Fort Macon) ; South Carolina (Wav-erly Mills, Mount Pleasant, St. James Island, and Hilton Head) ;Georgia (Savannah, Macintosh, Cumberland, and St. Marys) ; Flor-ida (St. Augustine, Fruitland Park, Longwood, and Fort Mj^ers) ;the Bahamas (Abaco Island, Eleuthera Island, Long Island, andBird Rock) ; and Haiti (Monte Cristi, La Vega, Mahiel, and SanCristobal). South to Haiti (San Cristobal and Port au Prince);Cuba (Trinidad, La Ceiba, and McKinley) ; and central Mexico(Aquas Calientes and Las Penas). West to Mexico (Las Penas andprobably San Bias) ; Lower California (Comondu, San Quintin Bay,and probably Todos Santos Island) ; California (San ClementeIsland, Santa Catalina Island, Santa Cruz Island, San Francisco.Napa, Cahto, and Eureka) ; Oregon (Elkton, Eugene, Corvallis, andPortland) ; Washington (Yakima, Seattle, and Bellingham) ; andBritish Columbia (Chilliwack).Whiter range.?In winter doves are found with regularity northto California (San Geronimo) ; central Arizona (Fort Verde) ;Colorado (Navajo Springs and Pueblo) ; Nebraska (Lincoln) ; Iowa(Wiota, Ames, and Sabula); Illinois (Rantoul) ; Indiana (Cam-den) ; southern Michigan (Manchester) ; Ohio (Oberlin and Cleve-land) ; Pennsylvania (Philadelphia) ; and New Jersey (Morristownand Englewood). East to New Jersey (Englcwood and Newfield)
;
Virginia (Wallops Island and Bowershill) ; Bermuda; North Caro-
EASTERN MOURNING DOVE 413lina (Raleigh and New Bern) ; South Carolina (Summerville, Charles-ton, and Beaufort) ; Georgia (Savannah and Darien) ; Florida(Daytona, Titusville, St. Lucie, and Royal Palm Hammock) ; theBahamas (Abaco Island, Eleuthera Island, Long Island, and BirdRock) ; Haiti (Monte Cristi, La Vega, and San Cristobal) ; andPanama (Calobre). South to Panama (Calobre and Volcano deChiriqui) ; Costa Rica (Azahar de Cartago and San Jose) ; Guate-mala (Duenas and Quezaltenango) ; Oaxaca (Tehuantepec) ; Jalisco(Zapotlan) ; and Lower California (La Laguna). West to LowerCalifornia (La Laguna and Triunfo) ; and California (SantaBarbara, Paicines, Gilroy, San Francisco, and San Geronimo).In addition to this normal winter range, individuals or small num-bers of mourning doves will occasionally spend the winter northalmost to the limits of the breeding range. Among such cases are:British Columbia (Okanagan Landing) ; Idaho (Emmett andGray) ; Minnesota (Lanesboro) ; Wisconsin (Beloit) ; Ontario(Plover Mills) ; New York (Rochester and Rhinebeck) ; and Massa-chusetts (Danvers and Barnstable).The range as described is for the entire species, which has, however,been divided into several races. True macroura is restricted to theGreater Antilles, although occurring in winter along the coast ofCentral America. It has no record in North America. The easternmourning dove {Zenaidura m. carol'men sis) occurs from the Atlanticseaboard west to the eastern edge of the Great Plains. It is foundalso in the Bahamas and on the Gulf and Caribbean coasts of Mexicoand Central America. The western mourning dove (Z. m. margin-ella) occupies the territory from the Great Plains to the Pacificcoast, and south through Mexico to Panama.Spring migration.?Early dates of spring arrival are : New York,Orient, March 1; Canandaigua, March 4; Hamburg, March 6; Ithaca,March 8; Buffalo, March 9; Rochester, March 14; and Rhinebeck,March 23 ; Connecticut, Fairfield, March 10 ; Jewett City, March 12
;
Norwalk, March 14. New Haven, March 18 ; Hadlyme, March 21 ; andSaybrook, March 26; Rhode Island, Providence, March 31. Massa-chusetts, Harvard, March 12; Taunton, March 12; Amherst, March17; Somerset, March 21; and Danvers, March 29; Vermont, Rutland,April 4; Castleton, April 8; Bennington, April 10; and Wells River,April 13; New Hampshire, Charlestown, April 14; Manchester, April16; and Concord, April 20; Maine, Bucksport, March 21; PophamBeach, March 30 ; and Machias, April 6 ; Ontario, Port Dover, March9; Point Pelee, March 17; London, March 17; Brighton, March 19;Harrow, March 20; Preston, March 24; and Guelph, March 25; Wis-consin, Janesville, March 10 ; Beloit, March 16 ; Prairie du Sac, March18; Milwaukee, March 20; North Freedom, March 21; Madison,
414 BULLETIN 162, UNITED STATES NATIONAL MUSEUMMarch 22, and Menomonie, March 23; Minnesota, Redwing, March12 ; Lanesboro, March 15 ; Hutchinson, March 23 ; St. Cloud, March24; Elk River, March 25; and Wilder, March 26; South Dakota,Yankton, March 14; Vermilion, March 23; Sioux Falls, March 25;and Dell Rapids, March 28; North Dakota, Fargo, March 18; Bis-marck, April 3 ; Larimore, April 5 ; Wahpeton, April 6 ; and GrandForks, April 12; Manitoba, Margaret, April 7; Aweme, April 8;Treesbank, April 10; and Pilot Mound, April 21; Saskatchewan,Eastend, April 23; Indian Head, April 29; Muscow, May 4; andWiseton, May 16. Utah, Clinton, April 18, and Salt Lake, April 26;Wyoming, Sheridan, April 13 ; Cheyenne, April 17 ; and YellowstonePark, May 9; Idaho, Deer Flat, April 3; Meridian, April 6; andPocatello, April 22; Montana, Knowlton, April 25; Bozeman, April26 ; Corvallis, April 29 ; and Billings, May 3 ; Alberta, Banff, May 24
;
Nevada, Carson City, April 23 ; Oregon, Albany, March 18 ; Beaver-ton, March 30; Portland, March 30; and Mulino, April 15; Washing-ton, North Yakima, March 21; Camas, April 7; and Pullman, April16; and British Columbia, Okanagan Landing, April 20; BurrardInlet, May 7; and Edgewood, May 15.Fall migration.?Late dates of fall departure are : British Colum-bia, Chilliwack, October 2; Courtenay, October 9; and OkanaganLanding, November 8; Washington, Tacoma, October 1, and Pull-man, October 2; Oregon, Cold Spring Bird Reservation, October14, and Portland, November 19; Nevada, Winnemucca, September9; Alberta, Red Deer River, September 15; Montana, Bozeman,October 2, and Glacier Park, October 30 ; Idaho, Minidoka, Septem-ber 29, and Meridian, November 4; Wyoming, Yellowstone Park,October 2, and Sundance, October 4; Utah, Clinton, September 10;Saskatchewan, Eastend, September 15, and Indian Head, October 1
;
Manitoba, Killarney, October 14; Ninette, October 15; Aweme, Oc-tober 19; and Margaret, October 20; North Dakota, Harrisburg,October 11; Inkster, October 14; Fargo, October 15; and Grafton,November 12; South Dakota, Wall Lake, October 13; Harrison,October 18; and Rapid City, October 24; Minnesota, Hutchinson,October 22; Twin Valley, October 25; Elk River, October 26; andMinneapolis, November 10; Wisconsin, Delavan, October 31; Shioc-ton, November 4 ; Trempealeau, November 17 ; Menomonie, November20; Elkhorn, November 22; and Meridian, November 23; Ontario,Guelph, November 3; Port Dover, November 4; Plover Mills, Novem-ber 14; Harrow, November 21; Windsor, November 27; and PointPelee, December 4; Maine, Gorham, September 26; Mount Desert,October 16; and Machias, November 10; Vermont, Bennington, Sep-tember 1 ; Massachusetts, Harvard, October 12 ; Lunenburg, October22; Amherst, October 28; Braintree, November 7; and Marthas
EASTERN MOURNING DOVE 415Vineyard, November 21; Rhode Island, Block Island, October 22,and South Auburn, November 20; Connecticut, Fairfield, October12; Meriden, October 20; Hartford, October 24; New Haven, Octo-ber 30; and Portland, November 30; and New York, Eochester,October 5 ; Rhinebeck, October 6 ; Geneva, October 12 ; Collins, Octo-ber 19; and Howard, October 24.From the fact that mourning doves winter so far north it mightbe assumed that they do not have an extensive migration. Thereturns from banded birds, however, indicate that the majority ofthese birds move in winter well to the south. For example, onebanded at Wauwatosa, Wis., on June 6, 1929, was shot in DaleCounty, Ala., on January 3, 1930, and another banded at Madison,Wis., on April 12, 1929, was recovered at Jennings, La., on December2, 1929.The region from Texas east to Georgia is the favored winter homeof the species. Eight birds banded at points in Nebraska, Kansas,Minnesota, Missouri, Illinois, Indiana, and Ohio were retaken inTexas; 7 banded in Illinois and Indiana were shot in Louisiana; 10banded in Illinois, Indiana, Michigan, and Ohio were recovered innorthern Florida ; 4 banded in Illinois, Indiana, and Ohio were shotin Alabama; and 14 banded in Illinois, Indiana, Michigan, NewYork, New Jersey, Pennsylvania, and Virginia were recovered inGeorgia. (See Lincoln, 1930.)Casual records.?Casual occurrences of the mourning dove havebeen chiefly north of the regular range, where it appears they maysometimes breed. Two young barely able to fly were noted on theStikine River, Alaska, by Willett, who also records one from Hyda-burg, Prince of Wales Island, on September 1, 1926, and one takenat Sitka, Alaska, on September 14, 1912. Kermode (1913) recordsa specimen from Telegraph Creek, British Columbia. There areseveral records for Quebec, among which are: Godbout, October10, 1881, and June 6, 1882; once at St. Joachim (date ?), and onein the fall of 1887 near Quebec City (Dionne). Newfoundland hasone record on October 16, 1890, but there are a few others on theLabrador coast: Spotted Islands, October 17, 1912: Battle Harbor,October 20, 1912; and Red Bay, September 7, 1891. The specieshas been recorded from New Brunswick at Fredericton, October 14,1899; Hampton, June, 1880; Rothesay, September 30, 1881; andMilkish, October 17, 1881.Egg dates.?Southern New England and New York: 20 records,April 6 to August 8; 10 records, May 20 to June 15. New Jerseyand Pennsylvania : 72 records, April 6 to July 8 ; 36 records, April26 to May 29. Florida : 10 records, March 11 to July 10. Michiganto the Dakotas : 18 records, April 27 to August 9 ; 9 records, May 14
416 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMto June 5. Indiana to Iowa: 58 records, April 4 to September 1;29 records, April 30 to June 7. Texas : 89 records, February 20 toSeptember 24; 45 records, April 21 to May 28. Arizona and NewMexico: 54 records, April 1 to September 2; 27 records, May 1 toJune 6. California: 170 records (every month but October andNovember), January 18 to December 5; 85 records, May 6 to June 19.ZENAIDURA MACROURA MARGINELLA (Woodhouse)WESTERN MOURNING DOVEHABITSThe habits of the western mourning dove are so much like those ofthe eastern race that there is very little to be added to the excellentlife history of the latter contributed by Doctor Tyler.The western race ranges from Manitoba and Oklahoma westward.Being largely an inhabitant of the open plains and more arid regions,it averages slightly paler in coloration, with the upper parts moregrayish, and slightly larger than the eastern race.Courtship.?Frank F. Gander has noted some aggressiveness on thepart of the female, on which he has sent me the following notes
:
On April 7, 1927, I watched a pair of doves mate. The male flew to a barelimb and was closely followed by the female, who pressed close to him andreached up with her beak as if begging for food. She also fluttered her wings alittle and squatted low to the branch. The male seemed inattentive at first, butgradually began to pay more attention, billing with her as if feeding, but therewas none of the straining of regurgitation. After a few moments of this, withthe female continuing to beg and remaining in a squatting posture, the act ofcopulation took place. The wings of the male were slightly raised and used tomaintain his balance. After mating, there was no strutting flight as in thepigeon, but the two birds calmly resumed their respective places on the limband, beyond a slight craning of necks and peering about, showed no signs ofhaving been affected. In a few moments the female again made advancestoward her mate and the billing took place, but the act of mating was notrepeated before the birds flew awayNesting.?Mr. Gander has found nests as high as 40 feet in cotton-wood and eucalyptus trees, and he says in his notes
:
Many years ago I found a set of eggs on a piece of loose bark that hadbecome lodged in the branches of a large tree. The only additions to thissite were two sticks, which were crossed, and the eggs were in the anglethus formed. In southern California, especially, the doves use a nest overand over again, and as some additional material is carried before each setof eggs, it often grows to a rather substantial nest.On April 27, 1927, I saw an adult dove brooding a set of two eggs inthe same nest with a well-grown young one. I have observed this on otheroccasions but have no further data.George Finlay Simmons (1925) says that in Texas the nest isplaced " rarely on leaning corn-stalks, rail fences, tops of rock fences,
ZENAIDA DOVE 417
or ledges in cliffs. Occasionally old nests of the gray-tailed cardinalor the western mockingbird are repaired and used ; one bird with eggswas found occupying an old nest of the Audubon caracara."Eggs.?The 2 eggs, rarely 1 or 3, of the western mourning doveare pure white like those of the eastern bird. The measurements of48 eggs average 28 by 20.9 millimeters; the eggs showing the fourextremes measure 31 by 21.5, 30 by 22, and 25 by 19.5 millimeters*.Behavior.?Simmons (1925) writes:In desert country, from Austin westward, water holes and the cool shadeof the narrowly timbered prairie creeks can be located by watching duringmid-day the direct lines of flight of the swiftly moving doves, which comefrom miles across the prairie, since they frequently nest a long distance fromwater. Hunters make use of this water-seeking habit of the doves by hidingnear a water hole at dawn or dusk and shooting the birds as they drop in towater preparatory to leaving or roosting in the nearby trees where they spendthe night. ZENAIDA ZENAIDA ZENAIDA (Bonaparte)ZENAIDA DOVEHABITSOn April 24, 1903, while crawling on all fours under the thornytangle of tropical shrubs and vines on Indian Key, one of the lowerFlorida Keys, I saw several small doves, with white in the wingsand tail, flitting along ahead of me near the ground or breakingout and flying away over the tops of the bushes. The vegetation wastoo thick to shoot them or even to get a good look at them, but I havealways suspected that they were Zenaida doves, as this is the keywhere Audubon (1840) mentions finding them. Here is what hesays about it
:
The Zenaida Dove is a transient visitor of the Keys of East Florida. Some ofthe fishermen think that it may be met with there at all seasons, but my ob-servations induce me to assert the contrary. It appears in the islands nearIndian Key about the 15th of April, continues to increase in numbers untilthe month of October, and then returns to the West India Islands, whence itoriginally came. They begin to lay their eggs about the first of May. Themales reach the Keys on which they breed before the females, and are heardcooing as they ramble about in search of mates, more than a week before thelatter make their appearance. In autumn, however, when they take their de-parture, males, females, and young set out in small parties together.Dr. Thomas Barbour (1923) says of its haunts in Cuba:This wide-ranging Pigeon is more shy and retiring than the Rabiche, andmore solitary. Nevertheless it is found in varying numbers throughout theisland. Its noisy flight is often startling. It is found rarely in deep forest,though Brooks and I have taken it in the high woods about the Cienaga. It isfar more characteristic of open savanna lands and the shady second-growthmanigua along water-courses in pastures and the outer boundaries of cultivated
418 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUM
fields. It shuns habitations, and is seldom seen in cultivated land; in fact, itfeeds but little on tbe ground. Its flesh is excellent.P. H. Gosse (1847) says that in Jamaica:The open pastures, or the grassy glades of pimento pens, are the favouritehaunts of this pretty Dove, where it walks on the ground singly or in pairs.In such open situations, it can discover, and mark the motions of an intruder,and long before he is within gun-range it is upon the wing. When the rainshave ceased, the increasing drought renders these, as it does many other birds,more familiar; and they may be seen lingering on the borders of streams andponds. Indeed they seem, of all our Doves, to haunt most the vicinity of water
:
particularly those dreary swamps or morasses which are environed by tallwoods of mangrove. In the winter months, when the pastures are burnt upwith drought, we may hear all day long their plaintive cooing, proceedingfrom these sombre groves, though it is not much heard in any other situation.Nesting.?Audubon (1840) writes:Those Keys which have their interior covered with grass and low shrubs,and are girt by a hedge of mangroves, or other trees of inferior height, areselected by them for breeding; and as there are but few of this description,their places of resort are well known and are called Pigeon or Dove Keys.It would be useless to search for them elsewhere. They are by no means soabundant as the White-headed Pigeons, which place their nest on any kind oftree, even on those whose roots are constantly submerged. Groups of such treesoccur of considerable extent, and are called Wet Keys.The Zenaida Dove always places her nest on the ground, sometimes artlesslyat the foot of a low bush, and so exposed that it is easily discovered by any-one searching for it. Sometimes, however, it uses great discrimination, placingit between two or more tufts of grass, the tops of which it manages to bendover so as completely to conceal it. The sand is slightly scooped out, and thenest is composed of slender dried blades of grass, matted in a circular form,and imbedded amid dry leaves and twigs. The fabric is more compact thanthe nest of any other Pigeon with which I am acquainted, it being sufficientlysolid to enable a person to carry the eggs or young in it with security. Theeggs are two, pure white, and translucent. When sitting on them, or whenher young are still small, this bird rarely removes from them, unless anattempt be made to catch her. which she, however, evades with great dexterity.On several occasions of this kind, I have thought that the next moment wouldrender me the possessor of one of these Doves alive. Her beautiful eye wassteadily bent on mine, in which she must have discovered my intention, herbody was gently made to retire sidewise to the farther edge of her nest, asmy hand drew nearer to her, and just as I thought I had hold of her, off sheglided with the quickness of thought, taking to wing at once. She wouldthen alight within a few yards of me and watch my motions with so muchsorrow, that her wings drooped, and her whole frame trembled as if sufferingfrom intense cold. Who could stand such a scene of despair? I left themother to her eggs or offspring.Gosse (1847) describes its nesting habits in Jamaica quite differ-ently, as follows
:
The nest is, as usual, a loose platform of twigs interlaced, with scarcelyany hollow, and no leaves ; it. is often built in an orange, or a pimento, andcontains two eggs of a drab hue. Near the end of March we started a Pea-dove from the centre of a lofty Ebby palm (Elate) in Mount Edgecumbe; it
ZENAIDA DOVE 419immediately alighted on the ground just before my lad, and began to tumbleabout in a grotesque manner, affecting inability to fly. Sam was not to becaught, however ; but calling my attention to the circumstance, we began topeer among the fronds of the tree, where we presently discerned the project-ing ends of the twigs that constituted her nest, the centre of her fears andanxieties. It was inaccessible, however, when discovered.Doctor Barbour (1923) says that " Gundlach found nests fromApril to July," in Cuba, " the usual shabby platform with two eggs,on bunches of epiphytic bromeliads or on some horizontal limb."Stuart T. Danforth (1925) found the Porto Kican form nesting stilldifferently ; he says
:
At the lagoon a few of these doves forsake their usual nesting sites andnest in the cat-tails. I found two such nests in 1924. The first was a crudeplatform of twigs on some bent cat-tail leaves in a dense clump on Las Casitas.It was three feet above the water, which was two feet deep at that place. OnMay 13 it contained two white eggs, but the nest was so frail it fell to piecesbefore the eggs had time to hatch. The second nest was built of dry cat-tailleaves and placed in the cat-tails at a height of four feet from the water, whichwas two inches deep at that time. When I found it on May 27 it containedtwo egg shells from which the young had hatched but the young birds werenowhere to be seen.And all the nests that Dr. Henry Bryant (1861) found in theBahamas " were made in holes in the rocks, and consisted, as isalways the case in this family, of but a few sticks."Eggs.?The eggs of the Zenaida dove are two in number, oval, butmore rounded than doves' eggs usually are, and pure white. Themeasurements of 17 eggs average 29.6 by 22.8 millimeters; the eggsshowing the four extremes measure 34 by 23, 31 by 24.5, and 25.2 by19.8 millimeters.Plumages.?I have not seen enough specimens in immature ortransition plumages to say much on this subject. Young birds injuvenal plumage seem to be similar to adults, but paler or duller,with more whitish on the chin and without the iridescent colors. Anadult, taken July 10, is completing the molt of wings and tail, whichindicates that the complete molt begins early.Food.?Audubon (1840) says of the food of this dove:The flesh is excellent, and they are generally very fat. They feed on grassseeds, the leaves of aromatic plants, and various kinds of berries, not exceptingthose of a tree wThich is extremely poisonous,?so much so, that if the juice ofit touch the skin of a man, it destroys it like aquafortis. Yet these berries donot injure the health of the birds, although they render their flesh bitter andunpalatable for a time. For this reason, the fishermen and wreckers are in thehabit of examining the crops of the Doves previous to cooking them. This,however, only takes place about the time of their departure from the Keys, inthe beginning of October. They add particles of shell or gravel to their food.In Porto Kico, Dr. Alexander Wetmore (1927) found thatthe bulk of the food of this dove consists of seeds, including many wildlegumes, euphorbias, mallows, knotweed and pigweed. Waste grain is also
420 BULLETIN 162, UNITED STATES NATIONAL MUSEUMtaken and various small wild fruits in season. The species is a valuable gamebird and should be carefully protected during the breeding season.Gosse (1817) writes:The Pea-dove subsists on various fruits and seeds : pimento-berries, orange-pips, sop-seeds, castor-oil nuts, physic-nuts, maize, and the smaller seeds ofpasture weeds are some of his resources. His flesh is white and juicy, and whenin good condition is in general estimation.Behavior.?Doctor Wetniore (1927) refers to its habits as follows:The flight is strong and direct and it flushes with a loud clapping of wings.On the ground this species resembles the mourning dove, as it walks quicklyabout with nodding head, and it has a cooing note almost indistinguishablefrom that of the bird mentioned.During the breeding season the males are frequently seen sailing out incircles, with the wings held stiffly, and their cooing notes come from the hillsidesall day long. They are also observed at times walking rapidly about on theground near the females, striking at each other with their wings.Between nine and ten in the morning the Zenaida dove comes in to streams orponds for water, usually in pairs, swiftly flying high in the air. On the gravelbars of the larger rivers they walk about quickly, quenching their thirst andpicking up bits of sand and gravel. Usually they are quite wary, but sometimesprefer to hide and let an intruder pass, rather than fly.The species frequents open country and is thus the only one of the largepigeons in Porto Rico that prospers with the clearing of the land. In somelocalities it is hunted constantly and then is very wild ; elsewhere it is quitetame.Audubon's (1840) account is somewhat different:The flight of this bird resembles that of the little Ground Dove more than anyother. It very seldom flies higher than the tops of the mangroves, or to anyconsiderable distance at a time, after it has made choice of an island to breedon. Indeed, this species may be called a Ground Dove too ; fox-, although italights on trees with ease, and walks well on branches, it spends the greaterportion of its time on the ground, walking and running in search of food withlightness and celerity, carrying its tail higher than even the Ground Dove,and invariably roosting there. The motions of its wings, although firm, producenone of the whistling sound, so distinctly heard in the flight of the CarolinaDove ; nor does the male sail over the female while she is sitting on her eggs,as is the habit of that species. When crossing the sea, or going from one Keyto another, they fly near the surface of the water ; and, when unexpectedlystartled from the ground, they remove to a short distance, and alight amongstthe thickest grasses or in the heart of the low bushes. So gentle are they ingeneral, that I have approached some so near that I could have touched themwith my gun, while they stood intently gazing on me, as if I were an objectnot at all to be dreaded.Voice.?C. J. Maynard (1896) gives an elaborate description ofthe sound-producing apparatus of the Zenaida dove, and thendescribes its notes, as follows:This dove, as might be expected from such a musical apparatus, has a singularnote. The male perches upon a limb of a tree, swells out his throat, and
ZENAIDA DOVE 421
utters his cooing song, which he repeats at rather regular, but protractedintervals.This song begins with two notes, the first uttered with a falling inflection,the second with a rising. The second follows the lirst rather quickly and isnot as prolonged. Both are in a low key. Then follows three other notes,sounding like " Who, who, who," but there is a decided pause between the firsttwo; the last three are given in the same time and in the same key. The notesare all loud, but when softened by distance have a singularly mournful effect.The notes of this bird which I have described above are very loud and ona still morning can be heard for a long distance. The males begin to coow ith the first indication of the dawn, and begin to fly about some time beforesunrise. They also coo at sunset and continue to utter their mournful notesuntil darkness fairly begins.Enemies.?One of the worst enemies of this and other ground-nesting- birds is the mongoose, but fortunately the dove has developedthe tree-nesting habit in certain places, perhaps as a result of pred-atory attacks. Several writers speak of it as an important andpopular game bird and much in need of protection. Mr. Danforth(1925) says:This species is probably preyed upon to some extent by the Mongoose, althoughits nesting habits make it more immune to attack than the Ground Dove. The/jenaida Dove is one of the most-sought-for game birds on the Island, and justlyso, as its meat is of good flavor, and it is so wary that one has to be an expertmarksman to obtain many. But it has a disadvantage in that it frequently flieslong distance after being shot before it falls, making the recovery of the birdsshot very difficult. Often but a small percentage of the birds killed arerecovered, especially when one is hunting in rough country. The few that Ihave shot were lost in this manner.DISTRIBUTIOXRange.?The West Indies, including the Bahama Islands, and thecoast of Yucatan; casual visitor on the Florida Keys. Generallynonmigratory.The range of the Zenaida dove extends north to northern Yucatan(Progreso and Holbox Islands) ; northwestern Cuba (San Cristobaland Habana) ; southern Florida (Indian Key) ; and the BahamaIslands (Great Bahama and Stranger Cay). East to the BahamaIslands (Stranger Cay, Moraine Cay, Abaco Island, Nassau, CatIsland, Watlings Islands, Acklin Island, North Caicos, GrandCaicos, and East Caicos) ; and the Lesser Antilles (Sombrero Is-land, Barbuda Island, Antigua Island, Grande Terre, BarbadosIsland, and Grenada). South to the Lesser Antilles (Grenada andSt. Croix Island) ; southern Porto Rico (Comerio, the CartagenaLagoon, and Mona Island): southern Haiti (San Domingo);Jamaica (Spanishtown, Port Henderson, Cayman Brae, Little Cay-man, and Grand Cayman) ; and the coast of Yucatan (Mujeres74564?32 28
422 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMIsland, La Logartos, and Progreso). West to the coast of Yucatan(Progreso).The Zenaida dove is apparently only a visitor to the FloridaKeys, arriving about the middle of April and remaining until aboutOctober, when they return to the islands of the West Indies.Two birds were reported by Pangburn (1919), as seen at Passa-grille, Fla., February 11, 1918, but as no specimen was obtained,the record is considered doubtful.Egg dates.?Bahamas and West Indies: 10 records, April 6 toDecember 8 ; 5 records, May 13 to June 12.LEPTOTILA FULVIVENTRIS ANGELICA Bangs and PenardWHITE-FRONTED DOVEHABITSThe white-fronted dove was first discovered as an inhabitant ofthe United States by George B. Sennett (1878) in the valley of theRio Grande in southern Texas, of which he writes
:
On April 18th, I obtained my first in a tract of timber a mile below Hidalgo,near the bank of the river. It was shot from the upper branches of the tallesttrees. Scattered about the woods in pairs were Colvmba flavirostris, Red-billed Pigeon, and Melopelia leucoptera, White-winged Dove. On the 19th, an-other was shot in the same locality. Five specimens were secured up to thetime of leaving, and a number of others seen and heard. It is more secludedthan the other Pigeons, and only found among the tallest timber. Seen in thewoods, it resembles M. leucoptera both in size and shape of tail, but can berecognized from it at sight by the absence of the large, white wing-patch.Recently the northern form of this dove, found in northern Mex-ico and southern Texas, has been separated as a distinct subspeciesby Bangs and Penard (1922), given the name angelica, and char-acterized as " similar to Leptotila fulviventris fulviventris Lawrence,but under parts less buffy, the under tail-coverts almost pure white
;
forehead more grayish, less vinaceous ; neck and chest less vinaceous."Spring.?Sennett (1879) writes:Dr. Finley reports the arrival of this Pigeon at the vicinity of Hidalgo andLomita about the middle of February, its departure having taken place inNovember. Although it is less numerous than the Red-billed Pigeon, yet, byits peculiar note, it is easily distinguished from all other species, and can thusbe readily obtained. We heard it daily. It is so much more retiring in itshabits than other Pigeons, that were it not for the peculiarity we mention,it would be met with very seldom. It frequents the dense and heavy growthof timber, and long and frequent were our endeavors to find its nest.Nesting.?On May 27, 1923, we found the white-fronted dovequite common and breeding in the dense forest around the Resacade la Palma, near Brownsville, Tex. These and the mourning andwhite-winged doves were especially abundant and much in evidence
WHITE-FRONTED DOVE 423among the varied bird life of that interesting region, more fullydescribed under the chachalaca. Their characteristic notes were al-most constantly heard, and we found several nests of each of thesespecies. The white-fronted and white-winged doves looked muchalike, but could be easily recognized by the deep-toned notes of theformer and the white wing patches of the latter. The nests and theeggs of these two species were also much alike, but could usually beidentified by seeing the birds leave them. The only set of eggs ofthe white-fronted dove that I collected was taken from a small, frailnest of sticks, 10 feet above the ground, on a horizontal limb of asmall tree, in the dense underbrush of the forest. I attempted tophotograph the bird on a similar nest, not so high up, but somethingdestroyed the eggs before the bird returned.Sennett (1879) had a nest brought to him (with the parent bird)that " was situated in the forks of the bushes, about five feet from theground, was flat and quite large for a Pigeon's nest, and composedof the dead branches, twigs, and bark of pithy weeds." Dr. J. C.Merrill (1879) took a nest that " was about seven feet from theground, supported by the dense interlacing tendrils of a hanging vinegrowing on the edge of a thicket." Major Bendire (1892) says:Mr. William Lloyd writes me that this Pigeon breeds abundantly in theSierra Madre, from southern Chihuahua to Beltran, Jalisco, Mexico, at analtitude of from 1,100 to 2,200 feet. The nests, usually placed in thornyshrubs, Huisache, Acacia farnesianna, 10 to 12 feet from the ground, aresubstantially made of straw. He found eggs as early as May 10, and up toJune 13, when they were much incubated. It frequents deep arroyas mostlyduring the breeding season.According to George N. Lawrence (1874) the white-fronted doveof western Mexico nests on the ground. He quotes Colonel Gray-son as saying that " differing from all our American doves, it de-posits its eggs upon the ground, forming scarcely any nest; theeggs are two, and white; the young soon follow the mother, beforebeing able to fly, like some of the gallinaceous birds."Eggs.?The white-fronted dove lays two eggs, which are easilyrecognized, when first collected, by their color. They are ellipticaloval in shape and somewhat glossy. The color is " cream-buff
"
at first, but it soon fades to " cartridge buff " or dull white. Themeasurements of 49 eggs average 30.6 by 22.9 millimeters; the eggsshowing the four extremes measure 33.7 by 23.5, 29.2 by 24.1, 28 by22.2, and 30.2 by 21.4 millimeters.Plumages.?I have seen no specimens of the downy young. Speci-mens in juvenal plumage, in June, have the crown, mantle, wings, andtail " sepia " ; the tail and wing feathers and scapulars are narrowlyedged with " cinnamon-buff " or " pinkish cinnamon " ; the greaterand lesser wing coverts are more broadly edged with " cinnamon-
424 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
rufous "; the breast is " wood brown." with " pinkish buff " edgings,shading off to grayish or buffy white on the belly.Material in collections is too scanty to illustrate subsequent moltsand plumages. A young bird, taken July 2, is fully grown and hasnearly completed the body molt into a plumage that is practicallyadult, but the flight feathers and wing coverts are still juvenal.Apparently both young and old birds have one complete molt duringa prolonged period in summer and fall.Food.?Little seems to be known about the food of these doves,which probably consists mainly of seeds, fruits, and berries. AustinP. Smith (1910) says: " They feed almost entirely on small herb andgrass seed, rarely partaking of the mesquite or ebony bean."Behavior.?Smith (1910) says:This dove approaches the true pigeons in hulk, but is more eminently ter-restrain than any of the several pigeons I am acquainted with. The White-fronted Dove is a slow-moving bird on the ground and quite unsuspicious ; audas it generally prefers to feed under growth of some sort, proves an easy targetfor the pot-hunter.Lawrence (1874) quotes from Colonel Grayson's notes as follows:This ' " ground dove," as its name indicates, is usually met with upon theground in search of its food, or sometimes resting upon low branches, or oldlogs, and always in the thickest woods, out of which they are seldom seen. Itwalks and runs with great facility upon the ground, whilst its flight is alwayslow amidst the bushes or underbrush as if to conceal itself, and not longcontinued, usually alighting upon the ground beneath a massive canopy o/underbrush, where it continues to walk or run to elude pursuit, or search forits food. When suddenly started from its retreat, the wings whir, accompaniedby a whistling sound, very similar to that of the wood-cock. Its habits aresolitary, never congregating into flocks, and only during the breeding season dowe ever find a pair together.Sennett (1879) considered it less of a ground dove, for he writes:During both seasons that I passed on the Rio Grande, I saw this bird uponthe ground but once, and it was then feeding upon some corn that was scatteredin the roadway, and. so far from its remaining near or on the ground, its habitis to frequent the high branches of tall trees: indeed, on this account almostevery specimen shot was more or less injured in falling, a number being toomuch so to save.Voice.?The notes of the white-fronted dove are characteristic; Irecorded them in my notes as deep-toned. Sennett (1878) says:Its note is somewhat prolonged, ends with a falling inflection, and is exceed-ingly low in pitch. Most of my birds were obtained by following the sound oftheir notes until within range; all were seen sitting quietly in secluded places;all are males, and injured considerably by falling from great heights.Winter.?From November to February this dove is apparentlyabsent from the northern portion of its range.
EASTERN WHITE-WINGED DOVE 425DISTRIBUTIONRange.?Central America, north to the lower Rio Grande Valleyin Texas; nonmigratory except that in the northern part of therange there is probably a slight withdrawal from November toFebruary.The range of the white-fronted dove extends north to Tepic(Santiago) ; northern Nuevo Leon (Rio Salado) ; and southernTexas (Rio Grande City, Lomita, and Sause Ranch). East to Texas(Sause Ranch, Santa Rita Ranch, and Brownsville) ; southernNuevo Leon (Montemorelos) ; Pueblo (Tecali) ; eastern Guatemala(San Geronimo) ; and northwestern Nicaragua (Chinandega).South to northwestern Nicaragua (Chinandega) ; Salvador (LaLibertad) ; western Guatemala (San Jose, Duenas, and Retalhuleu) ;Chiapas (Tonala) ; Oaxaca (Santa Efigenia) ; and Colima (San-tiago). West to Colima (Santiago); and Tepic (San Bias andSantiago).A specimen in the United States National Museum from TresMarias Island, Tepic, can be considered only as a straggler fromthe mainland.Egg dates.?Texas: 42 records, March 30 to July 25; 21 records,May 6 to 27. Mexico : 18 records, April 13 to June 28 ; 9 records,May 20 to June 24.MELOPELIA ASIATICA ASIATICA (Linnaeus)EASTERN WHITE-WINGED DOVEHABITSThe type name of the white-winged dove is now restricted to thebirds found in the West Indies and the eastern part of the rangeof the species. This eastern form is much less numerous than thewestern form and is not nearly so well known. For these reasonsit has seemed best to write a full life history of the western formonly.Baird, Brewer, and Ridgway (1905) say:This species is abundant in Jamaica, where, according to Mr. March, it ismore a lowland than a mountain dove. They are said to be gregarious, usuallykeeping in flocks of from 10 to 20, but in January and in February, in theGuinea-corn season, and at other times when the Cerei are in fruit, they con-gregate in large flocks, often of several hundreds. Their food is principallygrain and seeds, but they are equally fond of the ripe fruit of the differentspecies of Cereus abounding on the savannas and salines during the summer.Inland, the White-Wings, in the same manner as the Baldpate, breed in solitarypairs ; but in the mangrove swamps, and in the islands along the coast, theybreed in company, many in the same trees. The nest is a frail platform of
426 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
sticks, with a slight hollow lined with leaves and bark, and sometimes a fewfeathers.P. H. Gosse (1847) writes:In the early months of the year, when the physic-nut (Jatropha curcas) isripening, and oranges come in, the Whitewing becomes plentiful in openpastures, and the low woods in the neighbourhood of habitations ; the seeds ofthese fruits, and the castor-oil nut, forming the principal part of their food.At this time they are very easily shot, as they walk about on the ground.From the ease with which they are procured, they are a good deal eaten, thoughseldom fat, and rather subject to be bitter.When the rains fall, we see the Whitewings but seldom ; they betake them-selves to the deep woods and impenetrable morasses, when their presence isindicated by their loud stammering coo.Farinaceous and pulpy berries are found in the woods at all seasons, so thatthe Pigeons and other frugivorous birds have not only abundance but variety.Its nest is not very often met with. I am informed that it occasionally buildsin a pimento; Robinson says that it builds also in the orange, and sea-sidegrape, in May, a very slight and narrow platform of rude twigs, and lays twoeggs, of a pale drab hue.We found this dove abundant in Hidalgo and Cameron Countiesin southern Texas, where it evidently was the most numerous bird,next to the omnipresent great-tailed grackle, in the forests andthickets about Brownsville. We found a few nests in the chaparraland in the dense forests around the resacas, which I have alreadydescribed under the chachalaca. The nests were in low trees orbushes and were made of small twigs, grasses, and weeds. George B.Sennett (1878) found one nest made of Spanish-moss. The eggsare like those of the western form, but average a little smaller. Themeasurements of 33 eggs average 29.8 by 22.1 millimeters; the eggsshowing the four extremes measure 33 by 23, 31.5 by 24.5, 26.5 by20.5, and 28 by 19.5 millimeters.DISTRIBUTIONRange.?Southern United States, the West Indies, and CentralAmerica.The range of the white-winged dove extends north to southernCalifornia (Brawley and Palo Verde) ; Arizona (Little Meadows,Big Sandy Creek, Congress Junction, New River, Roosevelt, andGraham Mountains) ; New Mexico (Hidalgo County, Cloverdale,Mesilla Park, and Cliff) ; southern Texas (Del Rio, Uvalde, Castro-ville, San Antonio, Beeville, and probably High Island) ; theBahamas (Great Inagua Island) ; and the Lesser Antilles (St. Bar-tholomew Island). East to the Lesser Antilles (St. BartholomewIsland). South to the Lesser Antilles (St. Bartholomew Island);Haiti (Mount La Laguneat and Gonave Island) ; Jamaica (Spanish-town and Port Henderson) ; Costa Rica (La Palm a) ; Nicaragua
EASTERN WHITE-WINGED DOVE 427(San Juan del Sur) ; Salvador (La Libertad) ; Guatemala (VolcanoAgua and Duenas) ; Oaxaca (Tehuantepec) ; Tepic (Las PenasIsland and San Bias); and Lower California (Cape San Lucas).West to Lower California (Cape San Lucas, San Jose del Cabo,Santa Anita, Triunfo, La Paz, and Comondu) ; western Sonora(Guaymas) ; and southern California (Brawley).Migration.?Although white-winged doves are found in wintermore or less throughout their breeding range, migrating birds havebeen observed to arrive in Arizona as follows: Sabina Canyon,April 6, Otero Creek Canyon, April 12, Oracle, April 15, and Tomb-stone, April 15. Similarly, fall migrants have been observed atPhoenix, October 1, and Tombstone, October 21.The range as above outlined is for the entire species, which has,however, been separated into two subspecies. The eastern white-winged dove (Melopelia a. asiatica) is found from Texas, easternMexico, and Costa Rica, east to the West Indies, and casuallysouthern Florida. The western white-winged dove (M. a. mearnsi)occurs in the Southwestern United States and western Mexico.Casual records.?The occurrence of white-winged doves north oftheir normal range has been noted on numerous occasions. Amongthese records are California, a specimen at Escondido, about Sep-tember 25, 1911, one "heard" at Needles (Stephens, 1903), andone seen at Santa Barbara, November 8, 1922; Washington, onetaken at Puyallup, November 11, 1907; British Columbia, two seenand one taken at Sherringham Point, Vancouver Island, in July,1918 ; Colorado, one shot in the Wet Mountains in September, 1899
;
Texas, the most northwestern record being a specimen at Kerrville,November 25, 1910; Louisiana, one of a pair taken at Venice, aboutNovember 20, 1910, and Grand Isle, May, 1894, and August, 1895;Mississippi, one in Jackson County, on November 13, 1915; Alabama,one taken at Point Clear, about December 23, 1916, one taken atDaphne, about December 2, 1916, and another, also in BaldwinCounty, exact date and locality unknown (A. H. Howell, 1928)Florida, three specimens at Key West on November 14, 18S8, Novem-ber 20, 1895, and November 28, 1895, Kissimmee, November, 1896,and one taken near Orlando in the winter of 1908-9; Georgia, aspecimen at Hoboken, January 6, 1917; and Maine, one taken atLincoln, November 5, 1921.Egg dates.?Texas : 108 records, March 30 to July 14 ; 54 records.May 12 to 29. Arizona : 68 records, April 2 to August 2 ; 34 records,May 18 to June 2. Mexico: 20 records, April 20 to August 5; 10records, May 18 to June 20.
428 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMMELOPELIA ASIATICA MEARNSI RidgwayWESTERN WHITE-WINGED DOVEHABITSThe name mearnsi was applied to the white-winged dove of theSouthwestern United States and Mexico by Robert Ridgway (1915),who described it as " similar to M. a. asiatica but averaging de-cidedly larger, and coloration paler and grayer, the foreneck andchest light drab to hair brown instead of fawn color, the back, etc.,hair brown to deep drab."In the regions where I have met with the white-winged dove, Ifound it to be one of the commonest and decidedly the noisiest ofbirds. Its monotonous cooing and hooting notes were heard almostconstantly in the chaparral and forests, especially early in themorning and toward night.We found it very abundant in certain parts of southern Arizona.We camped for several days on the edge of the mesquite forest, onthe Santa Cruz River, south of Tucson, where we were lulled tosleep at night, or awakened in the morning, with the monotonousnotes of the white-winged doves ringing in our ears. They were mostnoisy at morning and evening, but could be heard at all hours ofthe day and sometimes during the night. These doves were alsocommon in nearly all the canyons, and a few were found in the morefertile valleys of the San Pedro River.Harry S. Swarth (1920) says:Throughout the valleys of southern Arizona the white-winged dove, orSonora pigeon as it is generally known, is an abundant summer visitor.Mosquite-grown bottom lands form the favorite breeding resort, and it isthere or in cultivated fields that the white-wings are to be found in numbers.Anywhere on the desert, however, one is apt to see them, passing overhead,feeding, or resting on the giant cactus or in the shade of the thicker bushes.They also invade the towns to some extent, and may frequently be observedin garden shrubbery or perched on fences or electric wires.Spring.?Although Major Bendire (1892) found it partially resi-dent throughout the year in the vicinity of Tucson, Ariz., and ob-served it during every month of winter, they were not so abundantthen as in summer, many having migrated. Mr. Swarth (1920)says: "The birds, as a rule, arrive in southern Arizona about thethird week of April. Gilman gives the date of arrival at Sacatonas April 20, while I found a bird sitting on eggs near Tucson asearly as April 13."M. French Gilman (1911) writes:Their coming is coincident with the ripening of the berries of the wild jujube,Zizyphus hicioides, upon which they feed greedily as long as the fruit lasts,consuming both ripe and green. They come in such great numbers that the
WESTERN WHITE-WINGED DOVE 429
wheat fields suffer and the loss is considerable. The Indians try to frightenthem away from the fields but do not hunt them. Probably they figure thatammunition would count up more on the debit side than would the wheatdestroyed.Referring to southern California, W. Leon Dawson (1923) says:The White-winged Dove is a tardy migrant, and its numerous arrival in lateApril is quite conspicuous. Flight is conducted at low levels, and occupancyis effected by a progressive invasion rather than by a sudden coup. Thebirds troop across the roads in endless desultory columns, or else rise hastilyfrom a wayside snack ; or, most likely of all, gather upon exposed branchesto mark with curious wooden detachment the passing of the intruder.Courtship.?Dr. Alexander Wetmore (1920) has published a mostinteresting account of his extensive studies of the white-winged dove,from which I shall quote freely ; regarding courtship, he writes
:
In displaying before females males had a curious habit or pose in which theyraised the tail high and tilted the body forward. At the same time the tail wasspread widely and theu closed with a quick flash of the prominent black andwhite markings. In the breeding colonies males at intervals flew out withquick, full strokes of the spread wings, rising until they were thirty or forty feetin the air. The wings were then set stiffly with the tips decurved, while thebirds scaled around above the mesquites in a great circle that often broughtthem to their original perches. The contrasted markings of the wings showedbrilliantly during this flight and the whole was most striking and attractive.In the cooler part of the morning males performed constantly in this mannerover the rookery.Mrs. Florence M. Bailey (1923) describes it as follows:One was seen displaying as he gave his call, as is described by Bend ire.Instead of inflating his chest pouter-pigeon style, as is done by the Band-tails,he puffed out his throat, and, as if about to launch into the air, threw up hiswings as some of the ducks do in courtship display of the speculum, showingthe handsome white wing crescent ; and at the same time curved up therounded fan tail so that its white thumb-mark band showed strikingly?allthis as he gave his loud emotional call
?
Eroo-kroo' -kroo-lcru' . A rather distantanswering call suggested that he was displaying for a prospective mate. Displayactually before a female was witnessed a week later by Mrs. Nicholson whenI was down in the valley. When the call was given without the emotionaldisplay it lapsed almost to monotony, being heard at camp all through theday. Some of the notes were heavily mouthed, while others were muffled.The noise of the flight was volitional. One that I saw, puffed out his chestand started with whacking wings, soaring around, wings and tail spread ; butshortly afterward it or another bird was seen flying by silently.Nesting.?Doctor Wetmore's (1920) account of the nesting habitsfollows
:
On my arrival in June I found them breeding in pairs scattered through thecultivated lands or the open desert, or congregated in large colonies in suitablemesquite monies near the Gila River. One or two pairs were found at inter-vals in cottonwoods beside roads or near ranch houses, but the greatestinterest centered in the large congregations to be found in suitable tracts ofmesquites. These rookeries were often of considerable extent. One located
430 BULLETIN 162, UNITED STATES NATIONAL MUSEUMthree and one-half miles south of Arlington extended over an area a quarter of amile square, while another three miles beyond occupied a grove nearly half amile wide and an equal distance in length. The birds maintained regular flightsacross country and gathered in flocks to feed, so that they were conspicuous fig-ures in the bird life of the region. It was difficult to estimate the number pres-ent, as they were scattered about in dense groves of mesquites, but it is believedthat there were at least two thousand pairs in the largest colony examined.The total number present in the area was large. It appeared that the periodfor breeding among these birds was somewhat irregular. A part of themevidently began to nest soon after their arrival, as a number that were feedingyoung were observed on June 6. Others were nest-building on June 17,so that the entire period of reproduction was somewhat prolonged. In thecolonies nests were scattered about irregularly through the mesquites. Some-times two or three nests were placed in the same tree, or again one pairoccupied a tree alone. There was no crowding and apparently the birds, whilegregarious, were too truculent to permit close proximity of nests. Often twoor three trees, suitable in every way for the primitive needs of these doves,intervened between occupied sites.In most cases the nest, slight in structure, though usually somewhat largerand bulkier than that of the Mourning Dove, was placed in a mesquite, thougha few were observed on the desert in palo verdes. Nests were built on in-clined living limbs where forking of small branches gave a firm, broad support.The site varied from six to twenty feet from the ground, with about eight feetas an average height. In most of those that were examined the structure wascomposed of dead twigs of the mesquite, small in diameter, and from six to teninches long. For the inner layers small twigs were chosen that had been deadfor some time, so that the spines, abundant on mesquite limbs, crumbled at atouch and caused no discomfort to the brooding bird or to the young. Thenest was flat and had merely enough depression to receive the eggs that oftenwere visible through the loosely interlaced twigs at the sides.Gilman (1911), who has had extensive experience with this dove,says:Nests are always, as far as my observation goes, placed in trees or shrubsat varying distances from the ground. The average height was ten feet andextremes ranged from four to twenty-five foot. The only nest as low as fourfeet was built in a mesquite tree and placed on top of an old Thrasher's nest.This may have been a shiftless bird ; but I found several others using old CactusWren's nests as foundation, and one had made use of a deserted Verdin's home.In choice of nesting sites the bird shows a decided preference for mesquite,as about 70 per cent of nests noted were in that plant. About 20 per centwere in willows, and 3 per cent in Cottonwood, Opuntia fulgida or tree cholla,and Prosopis odorata or screw-bean. Baccharis gluten brought up the rear with1 per cent. The dove is usually very wild on the nest, flying off whenever ap-proached as close as twenty-five feet. Rarely is the broken-wing play made,though I have seen a few mild attempts at it, and occasionally one will allow anapproach as close as fifteen feet to the nest before taking flight.In Arizona we found the white-winged doves nesting in mesquiteand hackberry trees in the mesquite forest; the nests were on hori-zontal branches, 10 or 12 feet above the ground, and were madeentirely of grass, weed stems, and straws; a nest found in the SanPedro Valley was 12 feet from the ground in a large willow.
WESTERN WHITE-WINGED DOVE 431Eggs.?The white-winged dove lays two eggs, very seldom threeor only one. Frequently one egg proves to be infertile, resultingin the rearing of only one young. The eggs are elliptical oval or ovalin shape, and the shell is smooth but not glossy. Fresh eggs, evenafter being blown, are often a rich, creamy buff, but this colorvaries greatly, and many eggs are pale creamy white or nearly purewhite; the whitest eggs are probably those that have been incubatedlongest. The color fades very soon, sometimes within a few days,after the eggs are blown. The measurements of 42 eggs average 31.1by 23.3 millimeters; the eggs showing the four extremes measure34 by 24, 27.5 by 21.5, and 33.5 by 21 millimeters.Young.?The period of incubation, according to Major Bendire(1892), is about 18 days, in which "the male relieves the femalesomewhat in these duties, but does not assist to any great extent;he, however, assiduously helps to care for the young." Apparentlyonly one brood is raised in a season in the northern part of itsrange, but farther south in Mexico two or more broods may be raisedduring a longer breeding season.Doctor Wetmore (1920) writes:I was of the opinion that males did not aid in incubation, but this I wasunable to ascertain with certainty. Occasionally I saw both parents perchedon the sides of a nest that contained young, but all birds that were definitelyidentified while engaged in incubating were females. Each mate chose aperch near the nest site, usually from ten to thirty feet away, and remainedthere on guard while the female was sitting, save for the times required tosecure food. Such perches were selected in situations that were well shadedfrom the direct rays of the sun during the heat of the day, and when notoccupied could be readily located by the collection of ordure, often con-siderable in quantity, on the ground beneath.The young birds were fed by regurgitation and at the age of four daysreceived solid food in the form of undigested seeds, in addition to the usualdiet of " pigeon's milk." Fledglings left the nest when between three andfour weeks old, as nearly as I could ascertain. The first young bird able to flywas noted on June 12, and by June 15 birds of this age were fairly common.These young were still dependent upon their parents for food, and thoughable to fly well were undeveloped and small. On first leaving the nest theyperched about in the mesquites, always seeking shade, but in a few dayswere often found on the ground, preferably where the soil was sandy. Therethey walked about in the thin shade of the mesquites, examining bits of sticksand other refuse curiously, often testing such fragments with their bills, orrested quietly, squatting on the earth. In many instances it was found thatthey were heavily infested with small ticks against whose attacks they seemedinexperienced. No ill results from the presence of these parasites were notedand older birds were free from them.Plumages.?He says further:Young White-winged Doves when first hatched were well covered with long,straggling down, that in color was dull white slightly tinged with buff. Thisnatal down was replaced by secondary feather growth so rapidly that it had
432 BULLETIN 162, UNITED STATES NATIONAL MUSEUMdisappeared for the great part at the end of the first week. The featherquills that followed the dowu did not burst until they were quite long as thatfor a time the young were as grotesque as young cuckoos.Young birds in juvenal plumage are much like adults, but theyare grayer on the back and breast and generally paler; there are fainttraces of narrow pale edgings on the mantle; and the throat and sidesof the head are whiter. I have seen adults, from Arizona, under-going a complete molt from August 2 to October 11, and adults, fromJamaica, completing the molt of wings and tail from December 25 toJanuary 19.Food?Doctor Wetmore (1920) gives the following account ofthe feeding habits of the white-winged dove:A purple drupe, one-fourth of an inch in diameter, borne by a spiny shrub(Covdalia spathulata) was a favorite food at this season and the birds alsoate the fruits of the giant cactus as rapidly as they ripened. Various seedswere taken also. Hai-vesting of grain began in this valley about the first ofJune and continued until the end of the month. Fields of wheat or barley thathad been cut recently were attractive to the White-wings as here they found anabundant source of food. The wheat grown in this region shattered (orshelled out) badly during the process of cutting, binding, and shocking, sothat kernels of grain wrere scattered thickly over the fields. Further, therewas much additional waste grain from heads matured or stalks too short tobe bound that fell to the ground when cut. As may be imagined the White-wings sought this food supply eagerly. They were gregarious in feeding asin nesting so that newcomers passing over the grain fields usually decoyedto those already on the ground until many had gathered in one spot. Thegrain stubble was cut high and afforded the feeding bands shelter, as thedoves were short in leg and walked about with the body beat forward. Itwas often the case that not a bird was seen in looking across a field of wheatstubble, though several hundred might be feeding there under shelter of thewheat stalks and the low levees thrown up to direct the flow of the water usedin irrigation while the crop was being grown. White-wings were wary andeasily alarmed while feeding. At times I crawled up under shelter of weedsto watch them at close range. If one of the feeding birds happened to ob-serve some slight motion, the heads of all were up in an instant and all re-mained motionless, while in a minute or so they usually flew hastily in sud-den alarm. Where they were shot at they became even more wary. Afterfeeding, little groups of White-wings often flew up to rest for a time in theshelter of cottonwoods or mesquites.Occasionally, when feeding in fields where wheat had not been shocked adove hopped up on one of the bundles of bound grain and pecked at theheads of wheat, choosing, preferably, those that were short so that they werefirmly held by the twine. Or a flock of half a dozen dropped down on ashock of wheat and fed on the cap sheaves for a few minutes. Usually,however, the birds preferred to feed in the more secure cover of the stubbleand confine their attention to the abundant waste grain as long as this wasavailable. When wheat was not threshed within a short time after it was cutthe-e doves were said to cause serious damage to the grain in the shock.This was particularly true in the case of isolated fields that remained afterthe surrounding crops had been removed. For this reason the White-wingedDoves were in bad repute among many of the ranchers.
WESTERN WHITE-WINGED DOVE 433Various kinds of waste grain, seeds, berries, mesquite beans, andinsects are mentioned by other observers. Dr. Joseph Grinnell(1914) found 33 watermelon seeds and a muskmelon seed in one crop.Behavior.?Doctor Wetmore (1920) writes:Combats among males were frequent, but tbese were bloodless battles, asthe birds merely flapped at one another uttering guttural notes, or whennear at hand struck quickly with one wing. Often one male was at muchtrouble to drive all others from some trees, and once I observed one hustleaway a pair of Mourning Doves that chanced to intrude upon bis domain.White winged Doves start in flight with a loud chipping of wings that isaccompanied by a whistling noise. Wben the birds are well under waytheir passage, while swift and direct, is noiseless. The sound at the startresembles that made by domestic pigeons. The White-wing, like certain tropi-cal doves (for example the White-headed and Scaled Pigeons) in perching incottonwoods or other trees with dense foliage, usually alights among clumpsof leaves on the higher outer branches rather than on dead limbs or in opensituations such as those chosen by Mourning Doves. So well did the birdsconceal themselves that after I had seen half a dozen fly into such a tree,it was not unusual to be unable to pick out a single dove in spite of theirlarge size. In the mesquite they followed the same practice in perching, sothat they were often observed merely as silhouettes through the thin foliage.When perched in trees they remained quiet save when they were calling.Voice.?Wetmore also gives the best impression of the remarkablevocal performances of these doves, as follows
:
In early morning White-winged Doves began to call soon after day break,and when the sun appeared above the horizon were heard cooing in everydirection. At this period of the day many males came out to rest on deadlimbs in openings in the mesquite montes, or flew to more distant perches inmesquites or cottonwoods where they basked in the warm rays of the sun.Others chose perches in the tops of living mesquites where the thin foliagedid not cast an appreciable shade. In mid-forenoon when the heat becameoppressive they retired again to protected stations. Males had two distinctsongs, that were given without apparent choice. One of these efforts may berepresented by the syllables who hoo who hoo-oo'. The first three notes weregruff and abrupt, the last one strongly accented and somewhat prolonged. Theother song, longer and more complicated, may be noted as who, hoo, whoo, hoo,hoo-ah.' hoo-hoo-ah' , who-oo. In this case the song was separated in fiveparts. The first section was short and low, the second louder and almostmerged with the third ; the third and fourth were more musical than theothers and were strongly accented on the last syllable, while the last part waslower and was more or less slurred. At times the doves gave one or the otherof these two songs in repetition for long intervals, or again alternated themrapidly. The longer song was more varied and pleasing to the ear as the otherfrequently was given in a burring, guttural tone that was often unpleasant.In addition to these songs males uttered a low, querulous, muttered note re-sembling queh queh-eh that served as a call to the female, or was given whensquabbling with other males. No females were observed in the act of cooingand I was unable to ascertain their notes. Although males did not coo inunison the effect produced by hundreds of them calling at the same time wasremarkable. Save for one or two birds that might chance to be near at hand,their notes seemed to come from a distance, and were so blended that it was
434 BULLETIN 162, UNITED STATES NATIONAL MUSEUMdifficult to pick out individual songs. In a large colony the volume of soundproduced was so great that it carried readily for a distance of a mile and yetthe tone produced was so soft that it was not deafening when near at hand.On the contrary the whole formed an undertone, continuous, and to my ear notunpleasing, that did not intrude sharply on the senses, of so vague a naturethat faculties perceptive to sound soon became accustomed to it, so that throughconstant repetition it might pass unnoted. Although it filled the air with thesame effect as that produced by the rushing of water, other sounds, the songof a Redwing or a Lucy's Warbler, the cooing of a Mourning Dove or the stamp-ing of a horse, were heard through it clearly even when such noises originatedat some distance. The effect as a whole was most remarkable and, once expe-rienced, lingers long in memory.Major Bendire (1892) says:Their call notes are varied, much more so than those of any other species ofthis family found with us ; they are sonorous, pleasing, and rather musical. Onthis account the natives keep many of them as cage birds, calling them Palomacantador, Singing Dove. They soon become very gentle and reconciled to cap-tivity, feeding readily out of one's hand and allowing themselves to be handledwithout fear.One of their most characteristic call notes bears a close resemblance to thefirst efforts of a young Cockerel when attempting to crow, and this call is fre-quently uttered and in various keys. While thus engaged the performer usuallythrows his wings upward and forward above the head and also spreads his tailslightly. Some other notes may be translated into " cook for you," or "' cookfor two," " cook-kara-coo," besides a variety of calls, one of these a querulousharsh one, resembles somewhat the syllables " chaa-haa."The commonest note, as I recorded it, might be written " who cooksfor you," and I notice that several others have so recorded it. Swarth(1920) very aptly remarks that it is "given with rather insultingemphasis." To my ear it sounds rather like a soft rendering of oneof the common notes of the barred owl. Its monotonous repetitionbecomes rather tiresome, but it is an impressive performance, whichonce heard can never be forgotten. Dawson (1923) says:In uttering this note the bird throws his head well forward and closes hiseyes ecstatically (thereby disclosing a livid blue eyelid), but he does not openhis beak. In defiance of all the masters, he sings through his nose. The effectis charming, it must be admitted, but one can not help wondering what thesound would be if only the bird would " sing out." Chanticleer's effort wouldsurely pale beside it.Fall.?Gilman (1911) writes:As soon as the young are grown both they and the parents congregate inlarge flocks and fly from feeding ground to watering place, thus affording agood chance at wing shooting. One evening in twenty minutes I counted over700 fly past a bridge over a small irrigating canal.The gunner, in these birds, has a good test of his skill, as they fly veryrapidly with seemingly little effort, and the rate of speed is hard to estimate.They will carry off a large load of shot, too, and all things considered are afine game bird.
EASTEEN GROUND DOVE 435Besides the danger from gunner, the Cooper Hawk is a menace, feeding oftenon the fat pigeon. I have seen a Marsh Hawk after a White-wing with a brokenwing, but do not think any but wounded birds are ever attacked by this species.Along in August the big flocks begin to grow less, the birds probably scat-tering out and seeking feeding grounds more distant from the breeding grounds.Toward the first of September they begin to thin out in earnest and by the 15thof the month very few are seen. Individuals may linger a little longer, as in1909 I saw one as late as October 12, and in 1910 the last seen was on September25. A few lingered on a sorghum field up till September 10 of this year, butwere not seen any later.COLUMBIGALLINA PASSERINA PASSERINA (Linnaeus)EASTERN GROUND DOVEHABITSThe gentle little ground dove is one of the most familiar and con-fiding dooryard birds in Florida, where it may be seen walkingbriskly about on its short legs, with a graceful nodding motion of itshead, about the houses, gardens, and more quiet streets in nearlyevery village. It is very tame and will allow a close approach, but, iftoo hard pressed, it will flit away to the nearest cover with a con-spicuous flash of reddish brown in its wings. Besides being verydomestic in its habits and attached to the vicinity of human dwellings,it is fond of sandy, cultivated lands, old weedy fields, cottonfields,pea patches, orange groves, and the borders of woods.Its range extends northward into Georgia and South Carolina,mainly in the coastal counties. Referring to Chatham County, Ga.,W. J. Erichsen (1920) says:A characteristic bird of the Lower Austral zone, this species, while formerlyabundant, is now quite uncommon. Its decrease during the past five yearshas been rapid and the few that now breed are restricted to three or four widelyseparated localities.During the period when it was abundant and generally dispersed in thecounty, I had many opportunities to observe its habits, and while it was to bemet with in equal abundance in country of greatly diverse character, its pre-ferred haunts were sparsely timbered woodland containing low and denseundergrowth.This species is non-migratory, passing its entire life in or very near the localityat which it was hatched. So attached to certain localities does it become thateven if the undergrowth is cleared and the land cultivated the bird remains,nesting on the ground among the vegetables.Courtship.?The courtship of the ground dove is a very simpleaffair, much like that of the domestic pigeon. The male struts be-fore the female, puffing out his feathers, bowing his head, and mak-ing a soft cooing sound. After they are mated the pair may often
436 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMbe seen sitting on a branch with their bodies touching in a mostaffectionate attitude. Donald J. Nicholson writes to me:Early in the spring the mating is begun and it is a common sight to seetwo or three males chasing a female on the ground. The female runs alongand the male closely follows, taking short flights to catch her, when shearises and alights just ahead. When she has made her choice, they fly tosome elevated spot in a tree and the mating is consummated. I have neverseen them copulating on the ground. They utter a soft peculiar note whenchasing mates. They have a habit of flitting their wings, when running along.Nesting.?Erichsen (1920) writes:In its choice of nesting sites, it exhibits a very wide range. It mostfrequently selects a low bush, either thinly or densely foliaged. Other sit-uations in which I have found nests include the top of a low stump ; highup on a horizontal limb of a large pine ; and, frequently, upon the ground.An instance of its nesting on the ocean beach came under my observationMay 13, 1915, on Ossabaw Island. In this case there was no attempt at nestbuilding, the eggs being deposited in a slight depression in the sand ; andwhen breeding on the ground in woodland or cultivated fields, little or nomaterial is assembled. In fact, nest building occupies little of the time andattention of this species, as when placed in trees or bushes the nest is simplya slight affair of a few twigs loosely interlaid. Further evidence of thisbird's disinclination to build a nest for the reception of its eggs is found inthe fact that I once found a set in a deserted nest of the cardinal (Cardinaliscardinalis cardinalis) .So gentle and confiding are these birds that it is often possible to touchthem while on the nest, especially if incubation is advanced. Upon droppingoff the nest they always simulate lameness, dragging themselves over theground with drooping wings in an effort to draw the intruder away. I amof the opinion that they remain mated for life, since they are observed through-out the year most frequently in pairs.Mr. Nicholson, who has " examined hundreds of nests," says in hisnotes
:
Nests are built on the ground as frequently as in vines, bushes, or trees,or along the tops of fences. One foot to 10 or 12 feet above the ground isthe usual height.The nests are delicate-looking structures, made usually of fine rootlets orgrasses, and seldom any sticks are used, saddled on a limb, or among dead vines.The diameter measures from 2% to 3 inches across, by 1 inch to 2 inchesthick, with scarcely any depression for the eggs, the eggs always showing abovethe rim of the nest.A nest that I found on Murrays Key, Bay of Florida, on April 3,1908, consisted of merely a few straws in a slight hollow in theground, under and between two tussocks of grass, which were archedover it; it was located in an open space in the brush, with smallshrubs and weeds about it. Maynard (1896) "always found thenests in orange groves; the neat domiciles are placed on the lowerlimbs of trees." Audubon (1840) says that the nest "is large forthe size of the bird and compact. Its exterior is composed of dry
EASTERN GROUND DOVE 437twigs, its interior of grasses disposed in a circular form ;" he found anest " placed on the top of a cactus not more than two feet high."Dr. T. Gilbert Pearson (1920) writes:There is no bird in the United States that to my knowledge breeds over solong a period of the year as does the ground dove. In my experience withthese birds in Florida, I have found their nests occupying varying situationsduring different seasons of the year. Thus on February 28 and March 3, Ihave found nests located on the tops of partially decayed stumps of pinetrees, only about 2 feet from the ground. Later in the season I have seennumerous nests placed on the ground, usually in fields of weeds or in standinggrain. Fields of oats seem to be especially favored with their presence duringmidsummer. Late in July, August and on to the latter part of September,I have found their nests on horizontal limbs of large orange trees, on the levelfronds of palms, and on the cross-bars or rails, as commonly used for supportsof the widespreading scuppernong grape-vines.Most observers have noted that when a ground dove's nest isapproached, the brooding bird quickly leaves the nest and fluttersalong the ground, attempting to lure the intruder away by feigninglameness. But Doctor Pearson (1920) writes:Occasionally an individual is found that declines to expose her treasureswithout an argument. As the inquiring hand comes close to the nest, shedoes not strike with her bill, nor even indulge in loud scolding, but withruffled feathers raises her wings in a threatening attitude, as if she would crushthe offending fingers if they came too close. Surely a puny, hopeless bit ofresistance ; nevertheless it shows that a stout heart throbs within the featheredbreast of the little mother.Mr. Nicholson has proved to his satisfaction that the same nestis used for a second or even third brood in a single season, by appar-ently the same pair of birds.Eggs.?The ground dove lays almost invariably two eggs, veryrarely three and apparently never less than two. The eggs areusually elliptical oval in shape, sometimes oval, and rarely ovate.They show little or no gloss and are pure white in color. The meas-urements of 34 eggs average 21.9 by 16.2 millimeters; the eggs show-ing the four extremes measure 24.4 by 16.6, 22.8 by 17.2, 20.3 by 15.2,and 21.7 by 15.1 millimeters.Young.?Incubation is said to last from 12 to 14 days, both parentsassisting. The breeding season is so prolonged, from February toNovember, that probably three or four broods, certainly two or three,are raised in a season. The young remain in the nest until they areready to fly. Nicholson says, in his notes, that " when disturbed theyoung fly from the nests with a strong flight which is marvelous forthe first attempt. After the young have hatched, the nest is veryuntidy; the droppings piled high on the outer edge of every nest,sometimes to half an inch or more in depth."74564?32 29
438 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMNicholson noted that the young are brooded by one or the otherparent, or shaded from the hot sun, until they are well grown andfeathered. When somewhat advanced in age they frequently comeout from under the brooding parent to sun themselves and stretchtheir legs and wings.Feeding begins within two hours after hatching; Nicholson saw ayoung bird break the shell and watched it until it was fed. Theyoung are fed entirely on regurgitated food, which is given by bothparents. Nicholson says it is a. common act for a ground dove tofeed both young simultaneously and that the young are more often fedthis way than singly. He has succeeded in photographing the act.Plumages.?The nestling ground dove is scantily clothed with long,stringy, hairlike down of a dull gray color. In the juvenal plumagethe young bird is much like the adult, but browner with duller mark-ings above and fewer or none below; the upper parts vary in colorfrom "snuff brown" to "cinnamon-brown," brightest on the wingand tail coverts, with conspicuous black spots in the wing coverts;the underparts vary from " buffy brown " to " wood brown." I havebeen unable to trace the subsequent molts.Food.?Doctor Pearson (1920) says:The ground dove's food consists largely of small seeds which it gathers in thegarden, on the lawn, by the roadside, in the field, and other places whereweeds or grasses are found. Naturally many insects are also picked up in theirtravels, particularly in the spring and summer. Small wild berries are alsoconsumed. So far as known they never adversely affect the interests of man-kind, even in the slightest degree, and wherever found they are protected bystatute and by the still stronger law of public sentiment.Behavior.?When disturbed the ground dove rises on whistlingwings; its flight is low and direct, but not protracted to any greatdistance; it generally amounts to only a short dash into the nearestcover. It is very much attached to certain restricted localities, inwhich it may be regularly found, and to which it soon returns afterbeing disturbed. It is well named, for it is decidedly terrestrial inits habits, spending most of its time on the ground, where it walksquickly, with a pretty nodding motion of its head and with anelevated tail. It is, however, often seen perched on a fence, thebranch of a tree, or the roof of a building.Nicholson tells me that the incubating or brooding male assumesa fighting attitude when the nest is approached, with wings raisedhigh above his back and uttering an angry, nasal, rasping note. Oneallowed himself to be lifted from the nest, to which he clung, makingangry notes and striking with repeated heavy downward strokesof the wings, but never striking with the bill.Voice.?The soft, cooing notes of the ground dove are the char-acteristic sounds that one hears in its Florida haunts ; their mourn-
EASTERN GROUND DOVE 439ful character has given it the local name of " mourning dove " inmany places. Nicholson says that
'
; their cooing is done entirely froman elevated position ; a house top, fence, telegraph wire, dead or livingtrees. I do not know if the female coos or not. For hours at a timeit is kept up, but with rest periods, of course. Four or five malescan be heard at one time within a 400-yard space."Enemies.?The gentle little ground dove is too small to be of anyaccount as a game bird, and, because of its sociable and confidingnature, there is a strong sentiment in favor of its protection every-where. It has little, therefore, to fear from man ; but it has plentyof natural enemies, such as cats, foxes, skunks, opossums, hawks,and snakes. It seems well able to take care of itself, however, andits numerous broods, though small, are enough to keep up its numbers.DISTRIBUTIONRange.?Southern United States and Central America.The range of the American races of the ground dove extends north,to southern California (Brawley and opposite Ehrenberg, Ariz.) ;Arizona (Big Sandy Creek, Fort Verde, Beaver Creek, H BarRanch, and Pima) ; southern New Mexico (Mesilla Park) ; Texas(Pecos, Devils River, and Seguin) ; Louisiana (New Orleans) ; Ala-bama (Autaugaville and Montgomery) ; and North Carolina (David-son County). East to North Carolina (Davidson County) ; SouthCarolina (Waverly Mills, Sullivans Island, and Frogmore) ; Georgia(Savannah, Blackbeard Island, and Macintosh) ; and Florida (Wa-tertown, San Mateo, De Land, Canaveral, Micco, Eden, Lake Worth,Palm Beach, Fort Lauderdale, and Miami). South to Florida(Miami, Royal Palm Hammock, Vaca Key, Big Pine Key, KeyWest, Boca Grande, and Marquesas Keys) ; Guatemala (Duenas andSanta Maria) ; Guerrero (Chilpancingo) ; Jalisco (Chapala and Za-potlan) ; Tepic (San Bias and Tres Marias Islands) ; and LowerCalifornia (Cape San Lucas). West to Lower California (Cape SanLucas, San Jose del Cabo, La Paz, Espiritu Santo Island, and SanFelipe) ; southwestern Arizona (Yuma) ; and southern California(Winterhaven and Brawley).The range as above outlined is for only the two American races,the eastern ground dove {ColvmibigalUna passerina passerina) , and theMexican ground dove (C. p. pallescens) , the first of which is foundin the South Atlantic and Gulf States west to Louisiana, whilepallescens is found in Texas, New Mexico, Arizona, southern Cali-fornia, and south to Guatemala. This species has been separated intomany other geographic races, extending from Bermuda, throughthe West Indies, to northern South America (Colombia and Vene-zuela).
440 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe species is generally nonmigratory, although there is someevidence of a movement in the Southwestern United States thatapparently involves only a part of the ground-dove population.Casual records.?There are several records of occurrence north ofthe normal range. Among these are : California, one found dead atSalinas in June, 1913, and one obtained at San Francisco in May,1870; Arkansas, a pair seen regularly for three years (last 1927), atRogers, according to a report from D. E. Merrill to the BiologicalSurvey ; Iowa, one reported as " seen " at Des Moines, on June 10,1922, by Clifford H. Pangburn; northern Alabama, one shot atLeighton, on May 4, 1889; northern Georgia, one at Rising Fawn,May 9, 1885 ; western North Carolina, one seen in Buncombe County,May 29, 1891; Virginia, one at Lynchburg on November 4, 1900;Maryland, one near the mouth of Broad Creek, October 14, 1888;District of Columbia, one at Washington, September 1, 1844; Penn-sylvania, one shot in Lancaster County in 1844; New Jersey, onetaken near Camden in the autumn of 1858 ; and New York, one takenfrom a flock of seven, near New York City, in October, 1862.Egg dates.?Florida: 73 records, February 27 to October 22; 37records, April 16 to June 2. South Carolina and Georgia: 20records, February 22 to October 19 ; 10 records, May 17 to June 10.Texas : 58 records, March 30 to October 1 ; 29 records, May 3 to June28. Arizona : 25 records, May 17 to October 8 ; 13 records, June 2 toAugust 11. Mexico: 34 records, March 7 to October 18; 17 records,May 3 to August 5.COLUMBIGALLINA PASSERINA PALLESCENS (Baird)MEXICAN GROUND DOVEHABITSA slightly paler form of the ground clove is found along oursouthwestern borders and in Mexico. We found it common orabundant in suitable places in southern Texas and in southern Ari-zona, where its haunts and habits seemed to be like those of theeastern bird. In Arizona it was breeding commonly in the valley ofthe San Pedro River and I recorded it as abundant in the mesquiteforest south of Tucson. It is a common dooryard bird in the small,quiet villages, very tame and confiding, as it runs about in thegardens or along the streets and is equally familiar about the ranches,barnyards, and cultivated fields. It is also common in the well-watered woodlands in the river bottoms and in the willows along theirrigation ditches.Grifling Bancroft (1930) says of its haunts in Lower California:The presence of water seems to be the determining factor in the distributionof this little dove. It is common wherever there are irrigation ditches or pools
MEXICAN GROUND DOVE 441
or available water in any form. As a consequence its occurrence is locallyconcentrated.The birds breed near open water. They carry their demand for its proximityso far that, assuming my notes to represent a fair average, four-fifths of theirnests are within fifty feet of a place to drink. I was surprised when this factbegan to develop and I found myself looking about for water whenever I flushedone of the doves from eggs or young. Seldom, indeed, was it not close at hand.There is a marked contrast here with the birds of southern Sonora. There theyabound on the open mesas and breed freely twenty to fifty miles or more fromwater.Courtship.?W. Leon Dawson (1923) writes:Business-like always, the ground dove is not less diligent in courtship. Thecall note oo woo vk, oo woo uk, sounds a little hard and unromantic in com-parison with that of the larger doves. The sound is very penetrating, but it isso low-pitched that some people fail to observe it. The singer is discreet, andthe sound usually ceases upon the appearance of the ever-despicable human.Yet at close quarters with his lady love, the workaday swain knows how to betender. At such times he trails after his enamorata with trembling wings andcries kool kooul. The daily visit to the drinking pool is the recognized occasionfor amours.Nesting.?A typical nest of the Mexican ground dove, which wrefound and photographed in the San Pedro Valle}^, Ariz., on May 17,1922, is shown on Plate 93. It was found while exploring a narrowstrip of small willows along the banks of an irrigation ditch. It waswell made, for a dove's nest, of coarse dry grasses and placed in themain crotch of a small willow, 6 feet above the ground. The twoeggs in it were fresh.Herbert Brown, as quoted by Major Bendire (1892), says:They lay two eggs, and nest in trees or bushes. A nest found June 11, 1887,was constructed of a few dead twigs and grass placed on a limb of a willownear the ground ; the female was on the nest. One found June 19 was also in awillow tree 20 feet from the ground and out on a limb 15 feet from the bodyof the tree, and made of a few dried stalks of alfalfa. It contained two eggsand the female was on the nest. A third nest, found June 26, containing twoeggs, was made of long stems of dry grass and placed about ten feet from theground. Whether this was a first laying I can not say. The nests are almostflat. I do not think I ever saw a cavity more than half an inch deep.Major Bendire (1892) says further:All of the nests seen by me were placed on bushes or on trees, from 3 to 21feet off the ground, not a single one was found on the ground.The first one found, on May 30, was placed in a syringa bush, about 3 feet fromthe ground. The little platform of small twigs and grass stems was veryslight, about 4% inches in diameter, and almost perfectly flat. The eggs werefresh.Other nests, subsequently noticed, were placed in various trees and bushes,mostly in mesquite thickets, a few in willows, and two in walnut trees. A nestfound July 28 was placed in a tree of this kind, about 20 feet from the ground.The tree was leaning, and some young sprouts had grown out from the maintrunk, among which the nest was placed. The eggs were fresh, probably a
442 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
second laying. All the nests examined by me were found in the creek bottomsor else close by, generally in clumps of mesquite bushes.Griffing Bancroft (1930) says of its nesting sites in Lower Cali-fornia :The Mexican ground dove is an unobtrusive little fellow, blending his colora-tion into his background on every occasion, and carrying his reticence into hischoice of nesting sites. He certainly does like concealment for his home, farmore so than the birds of the opposite mainland. Trestled grapes are plentifulin this part of Lower California. Commonly a beam of palm wood three or fourinches wide is supported on uprights at a height of five feet and a vine is trainedto grow over this structure. The favorite nesting site of the dove is on the flatsurface of the beam. The bird is snuggled in among the leaves, ideally protectedand hidden.Another popular haunt is in the palm jungles. At a height of from four to tenfeet and on the stems of the vertical leaves of the date palms numbers ofthese birds build. They seek the shadows that come from heavy vegetationor crossed leaves. Most of the fan palms have been trimmed, their leavesbeing cut largely for roofing material. The stubs left are generally about afoot long, smooth and well cupped. Here, hidden from below and concealedfrom the sides, many of these doves raise their young. The preferred heightis twelve to fifteen feet above the ground.This dove is tame, flushes at close range, and plays cripple most artistically.The laying season begins the middle of April. The nests are the most sub-stantial of any of the local Columbidae and often attain a thickness of aninch or more. They are built of comparatively long and fine materials, palmfibre and grass stalks being the favorites. They are well matted and thestrands are twisted spirally to form a flat disc to which is added somewhatfiner material in the center. Fifty-four eggs collected from Santa Agueda toSan Joaquin average 21.9X16.3 millimeters.John C. Fortiner (1920) reports three interesting records of winternesting of this dove in Imperial County, Calif. On December 21,1919, he found a dove brooding a single squab in its nest in aeucalyptus tree; the nest was well built and was placed on somelodged bark, well hidden from view from the ground. A secondnest was found on January 22, 1920, containing one young bird;
"this nest was also in a eucalyptus tree, about 18 feet from theground, and was a rebuilt mourning dove's nest. This second nestwas watched, and on February 14 was seen to have a sitting grounddove on it. The two eggs it contained were collected the next dayand found to have been incubated already several days."Again (1921) he writes:The Mexican ground dove appears to be partial to old nests, using its own orthat of a mourning dove generally; but I have seen a pair trying a Sonorared-winged blackbird's nest ; and during 1921 a pair has used an old AbertTowhee's nest for three broods, beginning to sit January 30, on the first eggs,and June 21, on the third set.M. French Gilman (1911) says:The nests are fairly well made for doves and are composed mostly of rootletsand small twigs. One nest rather more pretentious than usual was made of
BAHAMA GROUND DOVE 443
rootlets, grass stems and blades, leaf stems with veins attached, small twigs,horse hair, and a few feathers. It was compact and fairly well made, with adecided cup in the center measuring nearly an inch deep, and two inches acrossfrom rim to rim. One was an old nest re-vamped, and another was merely asuperstructure over an old Abert Towhee's nest. The very late date before men-tioned was probably the second brood, as the nest was an old one re-lined,possibly a last year's nest, but more likely an earlier nest of the same year.Eggs.?The Mexican ground dove lays two eggs, rarely only one.They are just like eggs of the eastern bird. The measurements of56 eggs average 21.5 by 16.5 millimeters; the eggs showing the fourextremes measure 23.5 by 16.5, 23 by 17.5, and 20 by 15 millimeters.Young.?Probably two, and perhaps three, broods are raised in aseason. Both sexes share in the duties of incubation, which is saidto last 14 days. Fortiner (1920) gives the following interestingaccount of the behavior of the young
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The nesting birds were not disturbed, and two weeks later the two old dovesand the young were discovered feeding on the ground. They soon flew to atree, where the young bird was fed by regurgitation, but by one of the parentsonly. No time was available for observation until the following Sunday, whenthe three doves were again seen feeding, and later all three flew to an umbrellatree, where the young dove was fed by both parent doves. The young dove, afterbeing fed once, hopped onto the old bird's back, then down to the limb on whichthe old dove was perched ; then, when not being fed, it extended its wing outover the parent dove and gently tapped the back of its parent until it was fedagain. It then flew to where the other parent dove was perched, where it wentthrough the same actions. Whether this is typical of the behavior of youngground doves I am unable to say.Food.?The food of this dove consists of seeds, waste grain, andvarious berries.Voice.?The call notes are described under courtship. WilliamLloyd, according to Major Bendire (1892), gives them as pas-cual,pas-cual, pas-cual. George F. Simmons (1925) describes the voice,which is seldom heard, as " intense cooing ; mellow, soft, crooning,floating coos; a single long drawn-out ventriloquistic, misleadingwoo, uttered at short intervals. Begins its moaning aboutmid-afternoon."COLUMBIGALLINA PASSERINA BAHAMENSIS (Maynard)BAHAMA GROUND DOVEHABITSThe form of the ground dove found on the Bermuda and BahamaIslands was described by Charles J. Maynard (1896) as "similar inform and general coloration to the ground dove, but somewhatsmaller and paler; the color on the lower parts and on the wingsabove being much less ruddy and the top of the head is more ashy,
444 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMthis color often extending well down onto the neck above. The billis wholly black, not red at base."Its habits apparently do not differ from those of the commonground dove, for he writes
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This is an exceedingly common and familiar bird throughout all of theBahama Islands which I have visited, being equally abundant in the groundsabout the houses, even in the city of Nassau, in the open spaces in the scrubremote from settlements, as well as on the most desolate and unfrequentedkeys, provided they are sufficiently wooded to afford the birds shelter. In thecity of Nassau, and in other towns and settlements, they are very tame, feed-ing about the houses either in pairs or in small flocks of from half a dozen toa dozen individuals.The Bahama ground dove breeds everywhere about the more open portionsof the scrub. The nest, as far as I have observed, is always placed on treesor bushes, the latter being most often chosen as a nesting site.Eggs.?These are also similar to those of the mainland birds. Themeasurements of the only three eggs available are 24 by 17.5, 20.7 by15.7, and 21.3 by 15.7 millimeters.SCARDAFELLA INCA INCA (Lesson)INCA DOVEHABITSContributed by Charles Wendell TownsendThe charming little Inca dove, sometimes called the scaly dove,or the long-tailed dove, after characteristic features, is a bird ofTropical and Lower Sonoran Zones and occurs in the United Statesonly in Arizona, New Mexico, and Texas. Formerly confined insouthern Texas to the region between San Antonio and the RioGrande, these doves, according to A. E. Schutze (1904), on accountof long droughts, have " moved north and eastward to a countrywhere they found food and water in abundance.'' According toG. F. Simmons (1925), the first record of this dove for Austin, Tex.,was in 1889, while, by 1909, they had become common nesters inthat region. Wherever found, it is resident, although in Texas,according to the same author, " a few birds move southward incolder, winter weather." Appearing to delight in human com-panionship, the Inca dove is rarely found at a distance from townsor the neighborhood of houses.Courtship.?Frank Stephens (1885) says of the courtship: "Isaw a little group on the ground, the males strutting around thefemales, carrying their tails nearly vertical and cooing." Althoughthe Inca dove may be heard cooing in every month in the year, thecooing is most in evidence during the courtship season. At this
INCA DOVE 445time, also, the dove is very pugnacious, and the rivalry is intenseamong them.M. F. Gilman (1911) says:The Inca dove could never have inspired the term " dove of peace," asthey are pugnacious to a fault and fight like little fiend:-;. Two of them willface each other with one wing on guard, held straight above the body, thenclose in and mix it, buffeting with wings till the sound of the blows is audibleat a distance of fifty yards. The bill is also used with bloody results aboutthe head. I have been told that one will sometimes kill the other, but neversaw such an extreme case. When arranging for a fight the combatants uttera sort of growl, if it may be so described; a very guttural, ? anger-expressingsound.Bryant (1891), quoting A. J. Grayson, says, "They exhibit themost ardent attachment for their mates and may often be seen caress-ing each other in a loving manner."Nesting.?The Inca dove delights to nest near houses or barnseven in villages and towns. Indeed, nests are rarely found exceptin the vicinity of mankind, and this familiarity is shown by thefact, according to M. F. Gilman (1911), that " the birds are generallyquite tame on the nest, rarely flying off till the intruder comes closerthan arm's length," and, he adds, " they are so accustomed to humanpresence that the broken-wing subterfuge is rarely resorted to."A curious instance of this familiarity with man and his works isgiven by George F. Simmons (1925), who reported that "a nestwas on a trolley wire at a switch in the eastern part of Austin,where about every seven minutes street-cars raised it from six totwelve inches above its normal position."The usual location of the nest is on a horizontal fork or flattenedlimb of a tree or in a bush, and it is generally within 10 or 12 feetof the ground, varying in height from 4 to 25 feet. Shade treesplanted about dwellings are commonly used. Umbrella trees, cot-tonwoods, elms, sycamores, fruit trees, mesquites, live oaks, acacias,thorn bushes, prickly-ash, and even Opuntia cactuses are all used forthis purpose, and, according to A. J. Grayson, quoted by W. E.Bryant (1891), "not infrequently they construct a nest under thesheds of the houses, if a suitable beam is found."A. J. Van Rossem, in his notes from Salvador, says
:
Inca doves breed the year round, nor does there appear to be any notableincrease or decline of this activity correlated with season. The numberof broods raised per year is not known to us, but because of the activity ofthe species as a whole it is not difficult to conjecture four or five. There is nocessation of nesting because of the fall molt. Males and females alike appearto have no dormant period whatsoever. This statement is based upon speci-mens taken every month in the year besides others inadvertently shot but notpreserved, and observations of numerous nests. Eggs were seen in July,August, September, October, November, February, and April. Nests of theusual slight dove construction were seen in orange trees, balanced on palm
446 BULLETIN" 162, UNITED STATES NATIONAL MUSEUMfronds, in mimosa thickets, and even in hanging fern baskets around thecorridor of an occupied ranch house. Some attempt at concealment wasusually noticeable, but this was frequently offset by the unsecretive mannerin which the parents left or approached the nest. Two eggs were the invariablerule.According to Bendire (1892), who quotes Herbert Brown, "thenests are as a rule much better constructed than those of the Mexicanground dove. The cavity is about half an inch deep, and the mate-rials used, fine dead twigs, are much more compactly put togetherthan in the nests of the latter." Simmons (1925), describes the nestconstruction as follows:Small, rather compact, firmly matted, almost flat platform or shallow saucerof weed stems, tiny twigs, dried grass, rootlets, a few straws, grass seedstems, bits of Indian tobacco weed, and sometimes bits of Bermuda grass,Spanish and bull moss, mesquite leaves, and a few feathers from the birds;occasionally nests contain string, horse-hair, or strips of cedar bark. Com-monly unlined ; rarely lined with grass stems, Spanish moss or a few smallInca dove feathers.Simmons has frequently found Inca doves using the nests of theirown species, of the western mourning dove, and of the westernmockingbird, after being slightly repaired and relined. Gilman(1911) found two nests each of which was built on top of an oldnest of a cactus wren.The dimensions of the nests, according to Simmons (1925), varyfrom 1.8 to 3.4 by 3.6 inches, with a height of 1.15 inches and aninside depth of 0.5 inch. He speaks of one nest " about the sizeof a silver dollar." Bendire (1892) describes a nest he found in athick mesquite bush as " a slight platform of twigs and grasses about5 inches in diameter."Eggs.?[Author's note: The Inca dove lays almost invariablytwo eggs. These are elliptical oval, smooth with very little gloss,and pure white. The measurements of 34 eggs average 22.3 by 16.8millimeters; the eggs showing the four extremes measure 24.3 by16.8, 22.9 by 18, 20 by 16, and 21.8 by 15.5 millimeters.]Young.?The duration of incubation is not known. It is prob-ably not far from two weeks, the same as the incubation period ofthe Mexican ground dove. Two broods are generally raised in aseason, sometimes three, and occasionally four. E. W. Quillin andR. Holleman (1918) record a case of a pair rearing four broods inone season in the same yard. Gilman (1911) says: "The past sea-son I noted four cases where two broods were raised in the samenest, and two cases where a last year's nest was relined and used."F. C. Willard observed one pair that laid five sets of eggs; he col-lected three sets and allowed the pair to raise two broods. Othersma}T have been raised later, as he was away after June 1.
INCA DOVE 447Plumages.?The following observations on plumage changes underartificial conditions are of great interest and possible significance,and are therefore recorded here. William Beebe (1907) subjectedwith other birds some Inca doves to a warm, superhumid atmos-phere and found that the plumage, with each succeeding molt,became darker and developed iridescence. He says:When the concentration of melanin has reached a certain stage, a changein color occurs, from dull dark brown or black to a brilliant iridescent bronzeor green. This iridescence reaches its highest development on the wing covertsand inner secondaries, where, in many genera of tropical and subtropical doves,iridescence most often occurs.In other words, by subjecting the Inca dove, which belongs in agenus of tropical origin, to the humidity of the Tropics, it revertedback in the lifetime of the individual to an ancestral type. Thisis certainly a most surprising result and the experiment should berepeated. Mr. Beebe's skins of Inca doves showing these remark-able changes are on exhibition in the park of the New YorkZoological Society.[Author's note: I have never seen the nestling of this dove.A small young bird, about two-thirds grown, in juvenal plumageis much like the adult; it is more heavily barred with black ordusky on the breast and flanks, with more buffy color on the belly
;
the feathers of the back, scapulars, and wing coverts have a heavierterminal black bar and a subterminal band of " cinnamon-buff."A. J. Van Kossem tells me that the postjuvenal molt occurs soonafter the young bird attains its full size; and that the completepostnuptial molt of adults comes at any time between July andOctober, inclusive, with individual variation.]Food.?"Weeds abound in back yards and near dwelling housesand barns, and in eating the seeds of these plants Inca doves givegood service. They also eat wheat and other small waste grains thathave fallen, but they are apparently unable to manage whole Indiancorn. Their familiarity permits them to mingle with poultry andpartake of their food.Water is, of course, an essential part of their diet, and referencehas alreacty been made to the extension of their range in times ofdrought in order that they may obtain water. M. F. Gilman (1911)says : " They are rather dainty in their drinking, rarely using thechickens' drinking vessel but perching on the hydrant and catch-ing the drops of water as they leak from the pipe. To do thisthey nearly have to stand on their heads, but that does not botherthem at all."Behavior.?In summer these birds may be seen singly or in pairs,or rarely in family groups, but in fall and winter they gather insmall flocks, which sometimes number as many as 50 individuals.
448 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMI shall always remember my first sight of this charming littledove. I was sitting alone in a small park in Tucson, Ariz., at Christ-mastime when 15 of these birds appeared and walked about withina few feet of me, picking at seeds on the ground. When they stoopedover for this purpose, their breasts nearly touched the earth, becausetheir legs were so short. As they walked their long tails generallysloped gently downward so as barely to skim the ground, but attimes the tails were cocked up nearly vertically. When disturbed bya passer-by, they flew up rapidly into the tree overhead, their wingsmaking a twittering sound. In the tree they sat in pairs or threes,affectionately snuggled together like love birds, their heads sunk intheir breasts, their tails pointing straight down. They went tosleep at once. This was at 4 p. m. ; at 4.20 they awoke and droppedto the ground to feed again.G. F. Breninger (1897) writes:The strange way in which Inca doves go to roost at night has recently cometo my notice. Nearly a month ago, when the air at night was still chilly, Isaw seven of these little doves perched on a limb side by side. This in itselfwas not strange, but directly upon the backs of the first row sat three moredoves. At another time I saw five in the lower row and two on top. Anexamination of the ground beneath showed it to be a resort to which thesebirds gathered to spend the cold nights of the winter months.F. C. Willard writes of watching an Inca dove in a grape arbor ina walled-in garden in Tombstone, Ariz., when four others alighted onthe same lattice bar
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The newcomer at the far end immediately began to assert herself by sidlingup to her next door neighbor and striking it with her wing. She soon forcedhim to fly and in like manner went down the line forcing each of them to seekanother perch. Having thus made room for herself, she crouched as if contentto call it a day and take a nap.They are not all love birds
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The following account of the dove's behavior is by M. F. Gilman(1911) :The vivacious little Inca dove is the cream of the dove family, and is in thepublic eye or ear most of the time. Whether sitting on a barbed wire fence oron a clothes line, with long tail hanging down perfectly plumb, or marchingaround in a combative manner with tail erect at right angles to the body, orrushing around busily and hurriedly, not to say greedily, feeding with thechickens in the back yard, it shows a decided individuality and arouses interestand affection.The faint twittering sound of the wings sometimes heard in flighthas already been mentioned, but as a rule the flight is noiseless. It isa quick and jerky flight.Voice.?Simmons (1925) describes the voice thus:Monotonous, tiresome, extremely mournful, rather short two-syllabled, hardlittle coo, quite different from the soft, soothing manner of the western mourn-
INCA DOVE 449ing dove, a slowly uttered coc-coo or co-o-o-h coo-o-o, the first slightly shorter,high-pitched, coarser, and with o as in go; the second lower, with a typical oosound, as in moon.Myron H. and Jane Bishop Swenk (1928) give the coos in musicalnotation, and they describe them as follows
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The call of the Inca dove is a monotonous, unvaried, rather plaintive coo-oo-coo or whoo-oo-ivhoo, rapidly repeated over and over. There is a blowingquality to it. We heard the call all through the winter, but it became louderand more insistent as the nesting season approached in March and April. It isvery different from the soft, drawled coo-oo-coo, coo-coo, coo of the mourningdove.During the very hot months of July and August the monotonousrepetition from morning to night of the Inca's coo is much dislikedby those with overwrought nerves.I have quoted under Courtship M. F. Gilmairs (1911) descriptionof the " growl " of the fighting birds, and he adds that " in animatedtalk, gossip perhaps, they excitedly utter sounds like cut-cut-ca-doo-ca-doo. In all quite a vocabulary is at their command."Field marks.?The chief field marks of the Inca dove are the longtail with its white edges and' the scaled appearance of the feathersover much of the body, which is due to the darker outline of theiredges. By these two marks the Inca dove may be distinguishedat once from the Mexican ground dove that occurs in the sameregion. Half of the Inca's length is in the tail. The chestnut-brownof the wing coverts, which is concealed or nearly concealed whenthe wings are closed, is prominent in flight.DISTRIBUTIONRange.?Southwestern United States and Central America; non-migratory.The range of the Inca dove extends north to Arizona (Wickenburg,Eice, and Safford) ; New Mexico (Silver City) ; and southern Texas(Kerrville, Austin, and Columbus). East to Texas (Columbus andSanta Maria) ; Tamaulipas (San Fernando de Presas, Ciudad Vic-toria, and Tampico) ; northern Guatemala (Lake Peten) ; Honduras(San Pedro); and Nicaragua (Chinandega). South to Nicaragua(Chinandega) ; Salvador (La Libertad) ; western Guatemala(Duenas) ; Oaxaca (Tehuantepec) ; Jalisco (Guadalajara) ; andsouthern Sinaloa (Escuinapa, Presidio, and Mazatlan). West toSinaloa (Mazatlan) ; Durango (Rio Sestin) ; Sonora (Opodepe) ;and Arizona (Tubac, Tucson, Sacaton, Phoenix, and Wickenburg)
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Casual records.?Although repeatedly listed as a bird of LowerCalifornia, there are apparently only two records. Dr. Witmer Stone(1905) reported seeing " a very few in the upper Hardy River
450 BULLETIN 162, UNITED STATES NATIONAL MUSEUM
region " of the Colorado River delta, in February and March, 1905,and there is a specimen in the British Museum taken at La Paz.Egg dates.?Arizona: 37 records, February 28 to October 21; 19records, April 15 to May 25. Mexico : 42 records, March 11 to October14; 21 records, March 23 to April 21. Texas: 12 records, April 10to August 10 ; 6 records. April 19 to May 28.OREOPELEIA CHRYSIA (Bonaparte)KEY WEST QUAIL-DOVEHABITSAudubon (1840) gives a graphic account of his discovery of thebeautiful Key West quail-dove in the dense, tangled, thorny thicketsof Key West. He named it the Key West pigeon, but used thespecific name montana, which is now applied to the ruddy quail-dove.It was originally discovered in Jamaica and now is rarely seen onKey West, as one taken there in 1889 by J. W. Atkins was the onlyone found by that keen observer in three years of careful field work.It was probably only a straggler, or a rare summer visitor, on KeyWest, retiring to Cuba in winter. Dr. Thomas Barbour (1923)says of its haunts in Cuba
:
The Torito is found in dry upland woods as well as in the low country, andI have flushed a good many in the low but thick forest of the limestone hills,or sierras, of Pinar del Rio. I also shot one once on the Sierra de Casas of theIsle of Pines, in low, scrubby second-growth {manigua) , hardly to be called aforest. In the cayos within the Zapata Swamp it was far less common thanthe ruddy quail dove ; nevertheless we often shot a few for food as well asto skin.Nowhere abundant, indeed a rather rare bird throughout its considerablerange, the Key West quail dove is one of the species which sooner or laterwill completely disappear.Nesting.?Audubon (1840) says:The nest of the Key West pigeon is formed of light dry twigs, and muchresembles in shape that of the Carolina dove. Sometimes you find it situatedon the ground, when less preparation is used. Some nests are placed on thelarge branches of trees quite low, while others are fixed on slender twigs.There is a set of two fresh eggs in Col. John E. Thayer's collec-tion, taken by A. H. Verrill on Inagua Island, Bahamas, April 21,1905 ; the nest was on the ground, composed of loose leaves. I havea similar set, taken by Mr. Verrill, at this locality on the previousday; the nest was on the ground, made of leaves and trash. MajorBendire (1892) gives Gundlach's description of the nest, as follows:The nest, consisting of a slight platform of sticks, is usually placed on thetop crown of certain parasitic creepers, found in the more open but shadyprimitive forests. The eggs are two in number, of a pale ochre yellow color,
KEY WEST QUAIL-DOVE 451
and measure 31.5 by 24 millimeters. I found nests between tbe months ofFebruary and July.Dr. Juan Vilaro wrote Major Bendire that " the nest is placedin high trees as a rule, usually Cui-ujeyes. It commences breedingin February and lays until July. The eggs are two in number, ochra-ceous white in color, and measure 31 by 24 millimeters."Eggs.?Audubon (1840) was evidently mistaken in reporting theeggs of this dove as white, to which no one else agrees. Bendire(1892) evidently never saw the eggs, which he quotes as " ochra-ceous white." Eggs in my collection, and others that I have seen, arecream color, or a pale shade of " cream-buff " ; and this color isfairly permanent in cabinet specimens. The measurements of eighteggs average 30 by 22.7 millimeters; the eggs showing the fourextremes measure 32.8 by 22.6, 31.3 by 23.6, 27.7 by 22.4, and 31.4by 22.3 millimeters.Plumages.?C. J. Maynard (1896) describes the nestling as " darkashy brown, becoming considerably lighter below; feet pink; billyellow, red at base; and iris red in all stages." Audubon (1840)says : " The young, when fully feathered, are of a dark gray colorabove, lighter below, the bill and legs of a deep leaden hue. I aminclined to believe that they attain their full beauty of plumage thefollowing spring."Eidgway (1916) describes the young asvery different in coloration from adults. Above rufous-cinnamon or pecanbrown, the scapulars, interscapulars, and wing-coverts narrowly tipped orterminally margined with cinnamon-buff, the pileum and hindneck duller (morebrownish) with indistinct, very narrow lighter tips to the feathers, the fore-head light grayish brown ; a dull white malar-subocular stripe, as in adults
;
foreneck and chest grayish brown or drab, the feathers margined with dullcinnamon ; rest of under parts mostly pale grayish buffy.Food.?Audubon (1840) says: "Their food consists of berries andseeds of different plants, and when the sea-grape is ripe, they feedgreedily upon it." Doctor Vilaro reports " fruits, seeds, and smallsnails " among the food.Behavior.?Audubon (1840) writes:The flight of this bird is low, swift, and protracted. I saw several after-wards when they were crossing from Cuba to Key West, the only place inwhich I found them. It flies in loose flocks of from five or six to a dozen, withflappings having an interval apparently of six feet, so very low over the sea,that one might imagine it on the eve of falling into the water every moment.It is fond of going out from the thickets early in the morning, for the purposeof cleansing itself in the shelly sand that surrounds the island ; but the instantit perceives clanger it flies off to the woods, throws itself into the thickest partof them, alights on the ground, and runs off with rapidity until it thinks itselfsecure. The jetting motions of its tail are much like those of the Carolinadove, and it moves its neck to and fro, forward and backward, as pigeons arewont to do.
452 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMThe cooing of this species is not so soft or prolonged as that of the commondove, or of the Zenaida dove, and yet nut so emphatical as that of any truepigeon with which I am acquainted. It may be imitated by pronouncing thefollowing syllables: lYhoe-whoe-oh-olb-oh. When suddenly approached by m:iu,it emits a guttural gasping-like sound, somewhat in the manner of the commontame pigeon on such an occasion. They alight on the lower branches of shrubbytrees, and delight in the neighbourhood of shady ponds, but always inhabit, bypreference, the darkest solitudes.Doctor Barbour (1923) says:This ground dove has habits much like those of the Perdiz and is oftencaught for food by the same means. Its flesh is excellent, although less es-teemed than the Perdiz. It is known as " Torito," the little bull, from itshabit of bobbing, or " Barbequejo," from the moustache-like markings. This,like the following species, is also called " Boyero," or Ox Driver, for its note, anoft repeated and prolonged monosyllablic coo, somewhat resembles the noiseconstantly made by men urging their oxen to strain to a heavy load.The Geotrygons, as I still like to call them, walk slowly about on the groundwith the head usually pulled in and not extended, and not bobbing except whendisturbed or frightened. Then they bob vigorously, as does the Perdiz all thetime, and this species is, I think, the shyest and most prone to take flight ofany of the group.L. J. K. Brace (1877), referring to the habits of the Key Westquail dove in the Bahamas, writes:This beautiful bird is frequently met with in the coppices underneath thetrees of which it delights to feed, preferring for this purpose those parts whichare rather open beneath, and less choked up with undergrowths, its habit beingto feed almost exclusively on the ground, on berries and seeds, more particularlyon the berries of the " poison wood," on the fruit of which, amongst others, thePatagioenas leucocephala feeds also. On being flushed, it scarcely, if ever, fliesto any distance, generally alighting after a short curved flight. Its note ispeculiarly mournful, being an expiring groan, which is rather startling to hearif the cause of it is not known. DISTRIBUTIONRange.?Key West, Fla., and the northern West Indies. Non-migratory.The range of the Key West quail dove is greatly restricted. Itextends north to southern Florida (Key West only) and the BahamaIslands (New Providence). East to the Bahama Islands (NewProvidence) ; San Domingo (Aguacate and Samana Bay) ; and pos-sibly Porto Rico (Mona Island). South to possibly Porto Rico(Mona Island) ; and Cuba (Isle of Pines). West to Cuba (Isle ofPines) and southern Florida (Key West).Audubon reported that this species left Key West about the middleof October, but other observers have since recorded their presenceas late as the middle of November.Egg dates.?Bahamas and Cuba : 9 records, April 5 to May 2.
RUDDY QUAIL-DOVE 453OREOPELEIA MONTANA (Linnaeus)RUDDY QUAIL-DOVEHABITSOn December 8, 1888, a boy at Key West, Fla., shot a red dove,which was sold with some mourning doves and plucked. That keenornithologist, J. W. Atkins, secured the remains, the head and somewing and tail feathers, and sent them to W. E. D. Scott (1889) whopublished the first Florida record of this West Indian species. Asecond record for the same locality was published by Ned Hollister(1925), when he reported the receipt at the National ZoologicalPark of a living specimen of this dove, caught by Ross E. Sawyer,in his backyard in Key West, about May, 1923.Dr. Thomas Barbour (1923) says of the haunts of the ruddyquail-dove in Cuba:With habits essentially like those of the preceding, this forest beauty is muchmore abundant and more confiding. By standing watching some little sunlitglade, or lying flat on one's belly on the damp forest floor, patience wasgenerally rewarded by a shot at the ruddy quail doves, provided one chose asuitable haunt in which to lie in wait. None of the quail doves occur in allsituations where, from the character of the terrain, one might expect them.They were really abundant, however, in the low woods between Zarabanda andSan Francisco de Morales and the Zapata Swamp, and equally so in a very finestretch of damp woods which I have visited but once, far to the south ofBolondron.Dr. Alexander Wetmore (1927), referring to Porto Rico, writes:The ruddy quail-dove is an inhabitant of dense growths of jungle and findscover to its liking mainly in the hills and mountains above the coastal plain.It is probable that its distribution is governed somewhat by the abundanceof the mongoose near the coast, since from its terrestrial habits the dove issubject to depredation by this mammal.At times the ruddy quail-dove is seen in coffee plantations, where these arenot kept too clean of brush, but it is usually found in areas of dense secondgrowth on the slopes of hills. As such cover becomes restricted in area, thesedoves grow steadily less abundant. To observe them it is necessary to walknoiselessly along footpaths, crouching low to obtain what vision may be hadof the ground beneath the dense brush.Nesting.?Writing of the tropical wild life of British Guiana,William Beebe (1917) gives a good description of the nesting habitsof this dove, as follows
:
Though one of the tropical jungle residents, the red mountain dove wasseldom seen, for it merged so completely with its surroundings that one passedit by, time after time, without ever knowing that such a bird existed. If itwere discovered, careful watch had to be kept or it would seemingly disappearwhere it sat. The nest was equally difficult to find and usually could only bediscovered by frightening the bird from the eggs. If it thought there were a74564?32 30
454 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
chance to escape undetected, the parent would quietly slip from the nest to theground, run a few steps and noiselessly flutter to a protecting branch withoutthe hunter being aware of its presence.The nests were built away from the ground, the distance varying from a footto five feet. The bird usually selected the head of an old rotted stump or thefork of a low outhanging branch, or possibly the horizontal surface of an oldgnarled liana that ran close to the ground. The nest itself was a concave plat-form of twigs lined with leaves on which rested the two dark, cream-coloredeggs. The nest in the accompanying illustration was lined in the same way,but some of the leaves were green and freshly picked so that the whole struc-ture had an effect of not existing at all in the green mass of foliage that grewaround it. The habit of mingling green leaves with brown was doubly significantfrom the fact that other nests found on stumps and lianas, where there was nosurrounding green, were lined only with dead brown leaves which made themjust as hard to see in their individual locality. The coloring of the eggs was noaid, for they nearly matched the leaves on which they lay.P. H. Gosse (1847) found two nests in Jamaica, of which he writes:One day in June I went down with a young friend into a wooded valley atContent to look at a partridge nest. As we crept cautiously toward thespot, the male bird flew from it. I was surprised at its rudeness; it wasnothing but a half dozen decayed leaves laid on one another, and on two orthree dry twigs, but from the sitting of the birds it had acquired a slighthollowness, about as much as a skimmer. It was placed on the top (slightlysunk among the leaves) of a small bush not more than 3 feet high, whoseglossy foliage and small white blossoms reminded me of a myrtle. Therewere two young recently hatched, callow and peculiarly helpless, their eyesclosed, their bills large and misshapen ; they bore little resemblance to birds.On another occasion I saw a male shot while sitting; the nest was thenplaced on a slender bush, about 5 feet from the ground. There were but twoeggs, of a very pale buff color; sometimes, however, they are considerablydarker.A set of two eggs in my collection was taken by A. H. Verrill onDominica, May 2, 1906 ; the eggs were laid on leaves on the ground.Eggs.?Two eggs form the usual set. They are oval, smooth withvery slight gloss, and vary in color from pale " cream-buff " to
" salmon-buff," which fade only slightly. The measurements of 25eggs average 28.5 by 21.4 millimeters; the eggs showing the fourextremes measure 32.6 by 22.2, 30.2 by 23.4, and 24 by 19.5 millimeters.Plumages.?I have never seen a nestling of this quail dove. Asmall but fully feathered young bird, in juvenal plumage in July,has the crown, back, scapulars, and wing coverts " brownish olive,"with a greenish luster; the wing coverts and scapulars are tippedwith " tawny," most broadly on the median and greater coverts,which are also edged with it; the remiges are edged with "rufous-cinnamon," and the rectrices and upper tail coverts are tipped withit ; the breast is " hair brown," with " tawny " tips, the belly buffywhite and the chin whitish with buffy tips.
RUDDY QUALL-DOVE 455Apparently a postjuvenal molt takes place in fall, which is prob-ably complete, for first winter birds of both sexes are much likethe adult female. I have not seen enough material to trace the molts.Food,.?Gosse (1847) writes:It is often seen beneath a pimento picking up the fallen berries ; the physicnut, also, and other oily seeds afford it sustenance. I once observed a pairof these doves eating the large seeds of a mango that had been crushed.With seeds, I have occasionally found small slugs, a species of Vaginulus,common in damp places, in its gizzard.In the Short Cut of Paradise, where the sweet wood abounds, the partridgeis also numerous ; in March and April-, when these berries are ripe, theirstomachs are filled with them. Here, at the same season, their cooing re-sounds, which is simply a very sad moan usually uttered on the ground, buton one occasion we heard it from the limb of a cotton tree at Cave, onwhich the bird was sitting with its head drawn in ; it was shot in the veryact.Doctor Wetmore (1927) says:During the orange season these doves feed mainly on the seeds of the wildsweet oranges, secured from fruit that has fallen to the ground and has partlydecayed, enabling the birds to peck open the skins and reach the seeds at thecenter. They do not touch this fruit except when on the ground, and can notopen oranges except when the skin is soft through decay, so that no injury inorange groves may be charged to them. Near Manati they were eating thefruits of the manchineel.Behavior.?Regarding the habits of the ruddy quail dove, DoctorWetmore writes
:
If the doves feel that they are liable to observation, they rest motionless,and at such times it is almost impossible to detect them. If approached tooclosely, they rise and dart into the dense growth. At other times they walkrapidly to one side, with quickly nodding heads, and it is then that they maybe momentarily visible. Their flight begins with a loud fluttering of feathers,but after a few feet they set their wings and sail away on noiseless pinions.Occasionally they were seen on low limbs in the trees, perhaps six to tenfeet from the ground, but this was unusual.Voice.?The same writer says that during the nesting season themales " give utterance to a low, resonant note of such character thatit seems always to come from a distance, though the singer may benear at hand; this resolves itself into a deep coo-oo-oo, with apeculiar undertone as of the humming of wind across the end of agun-barrel?a striking sound and one whose source is difficult tolocate." DISTRIBUTIONRange.?Northern South America, eastern Central America, andthe West Indies, accidental in Key West, Fla., and in western Mex-ico; nonmigratory.The range of the ruddy quail dove extends north to Hidalgo(Potrero) ; British Honduras (Orange Walk) ; Cuba (San Cristobal
456 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMand Trinidad) ; San Domingo (Puerto Plata) ; and Porto Rico(Mona Island, Aguadilla, Manati, and Vieques Island). East toPorto Rico (Vieques Island) ; the Lesser Antilles (GuadeloupeIsland, Dominica Island, Grenada Island, and probably Trinidad) ;British Guiana (Bartica) ; and eastern Brazil (Para, Capim River,Mirador, Cantagallo, Murungaba, Ypanema, and Iguape). Southto Brazil (Iguape and Mattogrosso) ; northwestern Bolivia (Apolo) ;and Peru (Tarma). West to Peru (Tarma, Ucayali River, Yuri-maguas, Chamicuros, and Nauta) ; Ecuador (Sarayacu and Pa-ramba) ; northern Colombia (Bonda) ; Panama (Chiriqui) ; CostaRica (Boruca, Terraba, Orosi, Angostura, and San Carlos) ; Nica-ragua (Chinandega) ; southern Vera Cruz (Esperanza) ; and Hidalgo(Chiquihuite Mountain and Potrero).Casual records.?Miller (1905), reporting on a collection of birdsmade in southern Sinaloa, Mexico, cites the capture of a specimenat Arroyo de Limones, on April 21, 1904, and indicates that others(" stragglers ") were seen by the collector. There are two recordsfor Key West, Fla., a specimen shot by a hunter on December 8,1888, and another captured alive in May, 1923. This bird was sentco the National Zoological Park, where it lived until March 5, 1926.Egg dates.?West Indies : 15 records, March 24 to June 13 ; 8records, April 26 to May 11.STARNOENAS CYANOCEPHALA (Linnaeus)BLUE-HEADED QUAIL-DOVEHABITSThe beautiful blue-headed quail-dove, one of the handsomest onour list, is a Cuban species, which occurs, as a rare straggler only,on some of the Florida Keys. No one seems to have recorded iton the keys since Audubon's (1840) experience with it, of which hesays:A few of these birds migrate each spring from the Island of Cuba to theKeys of Florida, but are rarely seen, on account of the deep tangled woods inwhich they live. Early in May, 1832, while on a shooting excursion with thecommander of the United States Revenue Cutter Marion, I saw a pair of themon the western side of Key West. They were near the water, picking gravel,but on our approaching them they ran back into the thickets, which were onlya few yards distant. Several fishermen and wreckers informed us that theywere more abundant on the " Mule Keys " ; but although a large party andmyself searched these islands for a whole day, not one did we discover there.I saw a pair which I was told had been caught when young on the latterKeys, but I could not obtain any other information respecting them, than thatthey were fed on cracked corn and rice, which answered the purpose well.
BLUE-HEADED QUAIL-DOVE 457Major Bendire (1892) quotes from Dr. Jean Gundlach as follows:It is not uncommon in the extensive forest, especially in such in which theground is rocky, but is scarcely ever found in cultivated fields or open prairiecountry. It moves slowly, with the neck contracted and tail erected, whilesearching for food among the dead leaves on the ground. This consists ofseeds of various kinds, berries, and occasionally small snails. After feeding,it usually flies into a tree and perches on a leafless horizontal limb, or on oneof the numerous parasitic vines, to rest. In tbe early mornings, should itsplumage, perchance, have become wet while traveling through the dew-ladenshrubbery, it selects a sunny spot to dry itself. From time to time this doveutters her call note, consisting of two hollow-sounding notes, hu-up, the firstsyllable long drawn out, the second short and uttered very quickly. Besidesthis note a low muttering is occasionally heard. Their call notes are deceptive,appearing near when distant, and distant when close by. Its flight is noisywhen starting, similar to that of the European partridge, from which it receivesits misleading name " Perdiz."Dr. Thomas Barbour (1923) writes:The blue-headed quail dove, on account of its brilliant blue crown, can notbe confused with any other species. Formerly it was a common denizen of allthe lowland forests of the island, where the soil was not too dry. To-day itis greatly reduced in numbers, both because it is so extensively trapped forfood and because the forests are being constantly cut away. There are twoordinary methods of trapping ground doves in general use among the countrypeople in Cuba. One involves the use of a casilla, a cage made of boughsor twigs, tied one upon the other, but at different intervals so that the completestructure is pyramidal and about two feet square and a foot high. This is putout in the open woods and baited with tripa de quhra, the mushy inner pulp ofthe wild calabash, which is full of seeds. The casilla is tilted, and a " figure-four " drops the contrivance when it is touched by the bird fussing aboutinside. Sometimes a small dish of water serves for bait. Another methodis to erect a net on hoops of creeper, and put bait beneath, where a decoy eitheralive or stuffed is often put out conspicuously. The hunter, in hiding, imitatesthe hup-up of the bird by means of a small hollow gourd.The bird is called " Perdiz " because of its firm white flesh and the noise itmakes when flushed. In common with the other ground doves, it prefers torun away from an annoyance rather than take flight. I have collected a goodmany by lying prone on the forest floor and simply watching for the birds towalk about. Much of the lowland forest in Cuba is flooded during the rainyseason, often for several feet, and this eliminates the very low undergrowtn,so that one may often see long distances with the eyes near the level of theground. Standing up, it is impossible to see off at all, so thick are the vinesand creepers. In 1915 I found Perdizes very common in the low woods, aboutfive miles inland from Jucaro and Palo Alto. I shot a good many, and theguajiros had dozens caged to sell to the planters about Ciego de Avila, who eatthem. This forest today is largely gone. About the cayos of the Cienagawhere I got the other ground pigeons in numbers, the blue-headed doves werevery rare, although I shot a few specimens. In Oriente the bird is still commonwhere it has not been trapped too hard, and here it occurs in the highlandforest where also suitable open woods are sometimes to be found.Nesting.?Major Bendire (1892) states that the blue-headed quail-dove " nests in April and May ; the nest is a simple affair, consisting
458 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUM
of a few twigs. It is usually placed in the tops of parasitic vines,Tillandsiay There are two sets of eggs in Col. John E. Thayer'scollection, taken by Oscar Tollin in Cuba on April 18, 1906 ; one nestwas 2 feet above the ground and was made of small sticks ; the othernest was in a small bunch of grass.Eggs.?The eggs of this dove are very scarce in collections ; I knowof only seven that appear to be authentic. Two eggs in my collection,laid by a bird in captivity in Florida, are between oval and ellipticaloval, smooth but not glossy, and pure white, quite unlike otherquail-doves' eggs; they measure 31.8 by 24.8 and 32.7 by 24.7 milli-meters. The three eggs, one set of two and one set of one, in theThayer collection are also white and are considerably larger, measur-ing 37.5 by 30.2, 37.9 by 30.2, and 37.9 by 28.9 millimeters. Two simi-lar eggs in P. B. Philipp's collection measure 38.5 by 27.7 and 37.7by 28.9 millimeters. These seven eggs average 36.3 by 28, and theextremes are indicated above.Plumages.?We know very little about the plumage changes of thisdove. The nestling is apparently unknown. In the Museum ofVertebrate Zoology, at Berkeley, Calif., I examined two young birds,about one-third grown, which had been hatched in an aviary. Theyare fully feathered in juvenal plumage, except that the chin andthroat are naked. They are strikingly like adults in color patternand colors. The colors above are duller; the crown is duller blue;the white stripe below the eye is present; there is a black patch onthe lower throat, bordered with white spots ; the feathers of the backand wing coverts are narrowly edged with rufous.DISTRIBUTIONRange.?Cuba, including probably the Isle of Pines; accidentalat Key West, Fla.Very little is known concerning the range of the blue-headedquail-dove. It is apparently restricted chiefly to Cuba (Guama,Trinidad, and Habana). Bangs and Zappey (1905) state that whileit has not been seen on the Isle of Pines by a naturalist, the nativesreport its presence. Cory (1892) states that it is claimed to havebeen introduced in Jamaica but was exterminated by the mongoose.Its only claim for inclusion in the North American list is the state-ment of Audubon that he saw two at Key West, Fla., in May, 1832,and that he also saw a pair in captivity alleged to have been cap-tured on the "Mule Keys."Egg dates.?Cuba ; 4 records, April 14 to June 12.
LITERATURE CITEDAbbott, Clinton Gilbert.1927. Notes on the nesting of the band-tailed pigeon. The Condor, vol. 29,no. 2, pp. 121-123, illus., Mar-Apr.Adams, A. Leith.1873. Field and forest rambles, with notes and observations on the naturalhistory of eastern Canada, 333 pp., illus. London.Alexander, Mark Leigh.1921. Wild life resources of Louisiana, their nature, value, and protection,164 pp., illus. New Orleans.Allen, Glover Morrill.1921. The wild turkey in New Eugland. Bull. Essex County Orn. ClubMassachusetts, vol. 3, no. 1, pp. 5-1S, Dec.Allen, Joel Asaph.18S6. The masked bob-white (Colinus ridgwayi) of Arizona, and its allies,Bull. Amer. Mus. Nat. Hist., vol. 1, no. 7, pp. 273-290.18S9. Note on the first plumage of Colinus ridgwayi. The Auk, vol. 6, no. 2,p. 189, Apr.American Ornithologists' Union.1910. Check-list of North American birds, ed. 3 (revised), 430 pp.1931. Check-list of North American birds, ed. 4, 526 pp.Anderson, Rudolph Martin.1907. The birds of Iowa. Proc. Davenport Acad. Sci., vol. 11, pp. 125-^17,1 map, Mar.Anthony, Alfred Webster.1889. New birds from Lower California, Mexico. Proc. California Acad.Sci., ser. 2, vol. 2, pp. 73-82, Oct. 11.1899. Hybrid grouse. The Auk, vol. 16, no. 2, pp. 180, 181, Apr.1903. Migration of Richardson's grouse. The Auk, vol. 20, no. 1, pp. 24-27,Jan.Atkins, John W.1899. Columba corensis at Key West, Florida. The Auk, vol. 16, no. 3, p.272, July.Audubon, John James.1840-1S44. The birds of America, 7 vols. New York and Philadelphia.Aughey, Samuel.1878. Notes on the nature of the food of birds of Nebraska. 1st Ann. Rep.U. S. Ent. Comm. for 1877, Appendix 2, pp. 13-62.Bailey, Alfred Marshall.1926. A report on the birds of northwestern Alaska and regions adjacentto Bering Strait, part 9. The Condor, vol. 28, no. 3, pp. 121-126,illus., May-June.1927. Notes on the birds of southeastern Alaska. The Auk, vol. 44, nos. 1-3,pp. 1-23, 184-205, 351-367, illus., Jan.-July.Bailey, Florence Merriam.1902. Handbook of birds of the Western United States, 511 pp., 36 pis.,601 figs. Boston. 459
460 BULLETIN" 16 2, UNITED STATES NATIONAL MUSEUMBailey, Florence Merriam.1918. Wild animals of Glacier National Park. The birds, pp. 103-199, pis.22-37, figs. 19-94. Washington.1923. Birds recorded from the Santa Rita Mountains in southern Arizona.Pacific Coast Avifauna no. 15, 60 pp., illus.1928. Birds of New Mexico, SOT pp., 79 pis., 135 figs., 60 maps. Santa Fe.Baird. Spencer Fullerton ; Brewer, Thomas Mayo ; and Ridgway, Robert.1905. The land birds of North America, 64 pis., 593 figs. Boston.Bancroft, Griffing.1930. The breeding birds of central Lower California. The Condor, vol. 32,no. 1, pp. 20-49, illus., Jan.-Feb.Bangs. Outram.1912. A new subspecies of the ruffed grouse. The Auk, vol. 29, no. 3. pp. 378,379, July.Bangs, Outram, and Penard, Thomas Ei>wari>.1922. The northern form of Lcptotila fulviventris Lawrence. Proc. NewEngland Zool. Club, vol. 8, pp. 29, 30, May 8.Bangs, Outram, and Zappey, W. R.1905. Birds of the Isle of Pines. Amer. Nat., vol. 39, no. 460, pp. 179-215,illus., Apr.Barbour, Thomas.1923. The birds of Cuba. Mem. Nuttall Orn. Club, no. 6, pp. 1-141, 4 pis.,June.Barlow, Chester.1899. Another chapter on the nesting of Dendroica Occidentalls, and otherSierra notes. Bull. Cooper Orn. Club, vol. 1, no. 4, pp. 59, 60, illus.,July-Aug.Barlow, Chester (with supplementary notes by W. W. Price).1901. A list of the land birds of the Placerville-Lake Tahoe Stage Road.The Condor, vol. 3, no. 6, pp. 151-184, illus., Nov.-Dec.Barrows, Walter Bradford.1912. Michigan bird life, 822 pp., 70 pis., 152 figs. Lansing.Beatty, Harry A.1930. Birds of St. Croix. Journ. Dept. Agr. Porto Rico, vol. 14. no. 3,pp. 135-150, 1 map, July.Beebe, William.1907. Geographical variation in birds with especial reference to the effectsof humidity. Zoologica, vol. 1, no. 1, pp. 1-41, illus., Sept. 25.1918-1922. A monograph of the pheasants, 4 quarto vols., pis. (part col.),maps. London.Beebe, William ; Hartley, G. Inness ; and Howes, Paul G.1917. Tropical wild life in British Guiana, vol. 1, 504 pp., 143 figs. NewYork.Behr, Herman.1911. Recollections of the passenger pigeon. Cassinia, no. 15, pp. 24-27.Belding, Lyman.1892. Food of the grouse and mountain quail of central California. Zoe,vol. 3, no. 3, pp. 232-234, Oct.1903. The fall migration of Oreortyx pictus plumiferas. The Condor, vol. 5,no. 1. p. 18, Jan.-Feb.Bkndire, Ch arlks Emil.1892. Life histories of North American birds. U. S. Nat. Mus. Spec. Bull.No. 1, vol. 1, 446 pp., 12 pis.
LITERATURE CITED 461Bendire. Charles Emil.1894. Tympanuchus americanus attwateri Bendire. Attwater's or Southernprairie hen. The Auk, vol. 11, no. 2, pp. 130-132, Apr.Bent, Arthur Cleveland.1912. A new subspecies of ptarmigan from the Aleutian Islands. Smith-sonian Misc. Coll., vol. 56, no. 30, 2 pp., Jan. 6.1912a. Notes on birds observed during a brief visit to the Aleutian Islandsand Bering Sea in 1911. Smithsonian Misc. Coll., vol. 56, no. 32,29 pp., Feb. 12,Bishop, Louis Bennett.1900. Descriptions of three new birds from Alaska. The Auk, vol. 17, no. 4,pp. 113-120, Apr.Bishop, Sherman Chauncy.1924. A note on the food of the passenger pigeon. The Auk, vol. 41, no. 1,p. 154. Jan.Bishop, Sherman Chauncy, and Wright. Albert Hazen.1917. Note on the passenger pigeon. The Auk, vol. 34, no. 2, pp. 208, 209,Apr.Bishop, Watson L.1890. Canada grouse in captivity. Forest and Stream, vol. 34, no. 19,p. 367, May 29.Bond, Frank.1900. A nuptial performance of the sage cock. The Auk, vol. 17, no. 4, pp.325-327, illus., Oct.Bowles, John Hooper.1901. Mice as enemies of ground-nesting birds. The Condor, vol. 3, no. 2,p. 47, Mar.-Apr.Brace, L. J. K.1S77. Notes on a few birds observed at New Providence, Bahamas, not in-cluded in Dr. Bryant's list of 1859. Proc. Boston Soc. Nat. Hist.,vol. 19, pp. 240, 241, May 16.Bradbury, William Chase.1915. Notes on the nesting of the white-tailed ptarmigan in Colorado. TheCondor, vol. 17, no. 6, pp. 214-222, illus., Nov.-Dec.Breninger, George Frank.1S97. A roosting method of the Inca dove. The Osprey, vol. 1, no. 8, p. Ill,Apr.Brewster, William.1874. The drumming of the ruffed grouse. American Sportsman, vol. 4.no. 1, p. 7, Apr. 4.1S82. A remarkable specimen of the pinnated grouse (Cupidonia cupido).Bull. Nuttall Orn. Club, vol. 7, no. 1, p. 59, Jan.1S85. The heath hen of Massachusetts. The Auk, vol. 2, no. 1, pp. 80-84.Jan.1885a. Additional notes on some birds collected in Arizona and the adjoin-ing province of Sonora, Mexico, by Mr. F. Stephens in 1S84 ; with adescription of a new species of Ortyx. The Auk, vol. 2, no. 2, pp.196-200, Apr.1887. Further notes on the masked bob-white (Colinus ridgicayi). TheAuk, vol. 4, no. 2, pp. 159, 160, Apr.1888. Descriptions of supposed new birds from Lower California, Sonora,and Chihuahua, Mexico, and the Bahamas. The Auk, vol. 5, no. 1,pp. 82-95, Jan.
462 BULLETIN 162, UNITED STATES NATIONAL MUSEUMBrewster, Wiixiam.1889. The present status of the wild pigeon (Ectopistes migrator ius) asa hird of the United States, with some notes on its habits. TheAuk, vol. 6, no. 2, pp. 285-291, Apr.1890. The heath hen. Notes on the heath hen (Tympanuchus cupido) ofMassachusetts. Forest and Stream, vol. 35, no. 10, p. 188, Sept. 25.1895. A remarkable plumage of the prairie hen (Tympanuchus americanus).The Auk, vol. 12, no. 2, pp. 99, 100, pi. 2, Apr.1902. Birds of the Cape region of Lower California. Bull. Mus. Comp.Zool., vol. 41, no. 1, pp. 1-241, 1 map, Sept.1925. The birds of the Lake Umbagog region of Maine. Bull. Mus. Comp.Zool., vol. 66, pt. 2, 402 pp.Brooks, Allan Cyril.1907. A hybrid grouse, Richardson's and sharp-tail. The Auk, vol. 24, no. 2,pp. 167-169, pi. 4, Apr.1912. Some British Columbia records. The Auk, vol. 29, no. 2, pp. 252, 253,Apr.1926. The display of Richardson's grouse, with some notes on the speciesand subspecies of the genus Dendragupuis. The Auk, vol. 43, no. 3,pp. 281-287, illus., July.1927. Notes on Swarth's report on a collection of birds and mammals fromthe Atlin region. The Condor, vol. 29, no. 2, pp. 112-114, Mar-Apr.1929. On Dendragapus obscurus obscurus. The Auk, vol. 46, no. 1, pp. 111-113, Jan.1930. The specialized leathers of the sage hen. The Condor, vol. 32, no. 4,pp. 205-207, July-Aug.Brooks, Allan Cyril, and Swarth, Harry Schelwaldt.1925. A distributional list of the birds of British Columbia. Pacific CoastAvifauna No. 17, 158 pp., 2 col. pis., 38 figs.Brooks, Winthrop Sprague.1915. Notes on birds from east Siberia and Arctic Alaska. Bull. Mus.Comp. Zool., vol. 59, no. 5, pp. 361-^13, Sept.Brown, Herbert.1885. Arizona quail notes. Forest and Stream, vol. 25, no. 23, p. 445,Dec. 31.1900. The conditions governing bird life in Arizona. The Auk, vol. 17, no. 1,pp. 31-34, Jan.1904. Masked bob-white (Colinus ridgicayi). The Auk, vol. 21, no. 2, pp.209-213, Apr.Bryant, Harold Child.1926. Life history and habits of the western mourning dove. CaliforniaFish and Game, vol. 12, no. 4, pp. 175-180, illus., Oct.Bryant, Henry.1861. A list of birds seen at the Bahamas, from Jan. 20 to May 14, 1S59,with descriptions of new or little known species. Proc. BostonSoc. Nat. Hist., vol 7, pp. 102-134.Bryant, Walter E.1891. Andrew Jackson Grayson. Zoe, vol. 2, no. 1, pp. 34-68, Apr.Bryant, Walter Pierce.1889. A catalogue of the birds of Lower California, Mexico. Proc. Calif.Acad. Sci., 2d ser., vol. 2, pp. 237-320.Burnett, Leonard Elmer.1905. The sage grouse, Centrocercus urophasianus. The Condor, vol. 7,no. 4, pp. 102-105, illus., July.
LITERATURE CITED 463Butler, Amos William.1898. The birds of Indiana. 22d Ann. Rep. Indiana Dept. Geol. and Nat.Res. for 1897, pp. 515-1187, illus.Cameron, Ewen Somerled.1907. The birds of Custer and Dawson Counties, Montana. The Auk, vol. 24,nos. 3, 4, pp. 241-270, 389-^06, illus., July, Oct.Capen, Elwin A.1886. Oology of New England, 116 pp., 25 col. pis. Boston.Cassin, John.1856. Illustrations of the birds of California, Texas, Oregon, British andRussian America, 29S pp., 50 col. pis. Philadelphia.Chapman, Frank Michler.1902. List of birds collected in Alaska by the Andrew J. Stone Expedi-tion of 1901. Bull. Amer. Mus Nat. Hist., vol. 16, pp. 231-247.1904. A new grouse from California. Bull. Amer. Mus. Nat. Hist., vol.20, art. 11, pp. 159-162.1908. Camps and cruises of an ornithologist, 432 pp., 250 figs., New York.Childs, John Lewis.1905. Personal recollections of the passenger pigeon. The Warbler, ser. 2,vol. 1, no. 3, pp. 71-73, July 20.Christy, Bayard Henderson, and Sutton, George Miksch.1929. The turkey in Pennsylvania. The Cardinal, vol. 2, no. 5, pp. 109-116,illus., Jan.Clark, Austin Hobart.1905. Birds of the southern Lesser Antilles. Proc. Boston Soc. Nat. Hist.,vol. 32, no. 7, pp. 203-312, Oct.1907. Eighteen new species and one new genus of birds from eastern Asiaand the Aleutian Islands. Proc. U. S. Nat. Mus., vol. 32,pp. 467-475.1910. The birds collected and observed during the cruise of the UnitedStates Fisheries steamer Albatross in the North Pacific Ocean,and in the Bering, Okhotsk, Japan, and Eastern Seas, from Aprilto December, 1906. Proc. U. S. Nat Mus., vol. 38, pp. 25-74, 3 figs.,Apr. 30.Cleaves, Howard Henderson.1920. A partridge Don Quixote. Bird-Lore, vol. 22, no. 6, pp. 329-334,illus., Nov -Dec.Colvtn, Walter.1914. The lesser prairie hen. Outing, vol. 63, pp. 608-614, illus., Feb.1927. In the realm of the prairie hen. Outdoor Life, Jan., pp. 17-19, illus.Cooke, Wells Woodbridge.1888. Report on bird migration in the Mississippi Valley in the years 1884and 18S5. U. S. Dept. Agr. Div. Econ. Orn. Bull. 2, 313 pp., 1 map.1897. The birds of Colorado. State Agr. College Bull. 37, 143 pp., Mar.Cory, Charles Barney.1891. Lists of birds collected by C. L. Winch in the Caicos Islands andInagua, Bahamas, during January and February, and in Abaco,in March, 1891. The Auk, vol. 8. no. 3, pp. 296-298, July.1S92. Catalogue of West Indian birds, 163 pp., 1 map, Boston.Cottam, Clarence.1929. The status of the ring-necked pheasant in Utah. The Condor, vol. 31,no. 3, pp. 117-123, May-June.
464 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMCoues, Elliott.1874. Birds of the Northwest. U. S. Geol. Surv. Terr., Misc. Publ. no. 3,791 pp.Cbaig, Wallace.1911. The expression of emotion in the pigeons. II, The mourning dove(Zenaidura macroura Linn.). The Auk, vol. 28, no. 4, pp. 398-407, Oct.1911a. The expressions of emotion in the pigeons. Ill, The passengerpigeon (Ectopistes migratorius Linn.). The Auk, vol. 28. no. 4, pp.40S-427, Oct.Criddle, Nobman.1930. Some natural factors governing the fluctuations of grouse in Munitoba. Can. Fiekl-Nat., vol. 44, no. 4, pp. 77-80, Apr.Danfobth, Stuart Taylob.1925. An ecological study of Cartagena Lagoon, Porto Rico, with specialreference to the birds. Journ. Dept. Agr. Porto Rico, vol. 10,no. 1, 136 pp., figs. 2-45, 1 map.Dawson, William Leon.1896. Notes on the birds of Okanogan Co., Washington. Wilson Bull., no.10, pp. 1-4, Sept. 30.1923. The birds of California, 3 vols., part col. pis. San Diego, Los Angeles,San Francisco.Dawson, William Leon, and Bowles, John Hoopeb.1909. The birds of Washington, 2 vols., illus. Seattle.Deane, Ruthven.1896. Some notes on the passenger pigeon (Ectopistes migratorius) inconfinement. The Auk, vol. 13, no. 3, pp. 234-237, July.Devany, J. (L.1921. The spruce drummer. Can. Field-Nat., vol. 35, no. 1, pp. 16, 17, Jan.Dickey, Donald Rydeb, and Van Rossem, Adkiaan Joseph.1923. Description of a new grouse from southern California. The Condor,vol. 25, no. 5. pp. 168, 169, Sept.-Oct.Dixon, Joseph Scattebgood.1927. Contribution to the life history of the Alaska willow ptarmigan.The Condor, vol. 29. no. 5, pp. 213-223, illus., Sept.-Oct.1930. Jays or California quail? Nature Mag., vol. 15, no. 1, pp. 42, 43, illus.,Jan.Dutcheb, William.1903. The mourning dove. Nat. Assoc. Audubon Soc. Educ. Leaflet 2, 3 pp.,1 pi., Mar. 2. New York.Dwight, Jonathan, jb.1900. The moult of the North Amerii an Tetraonidae (quails, partridges, andgrouse). The Auk, vol. 17, nos. 1, 2. pp. 34-51, 143-166, illus., Jan.,Apr.Eaton, Elon Howabd.1910. Birds of New York. New York State Mus. Mem. 12, pt. 1, Waterbirds and game birds, 501 pp., 42 pis.Edson, John Melton.1925. The Hooters of Skyline Ridge. The Condor, vol. 27, no. 6, pp. 226-229,Nov.-Dec.Edwabds, George.1755. A letter to Mr. Peter Collinson, F. R. S., concerning the pheasant ofPennsylvania, and the Otis Minor. Philos. Trans. Roy. Soc. Lon-don, vol. 48, pt. 2, for 1754, pp. 499-503, 2 pis.
LITERATURE CITED 465
Elliot, Daniel Gibaud.1896. Descriptions of an apparently new species and subspecies of ptarmiganfrom the Aleutian Islands. The Auk, vol. 13, no. 1, pp. 24-29, pi. 3,Jan.1897. The gallinaceous game birds of North America. 220 pp., 46 pis.,New York.Erichsen, Walter Jefferson.1920. Observations on the habits of some breeding birds of ChathamCounty, Georgia. Wilson Bull. no. 113, vol. 32, no. 4, pp. 133-139,Dec.Field, George Wilton ; Graham, George H. ; and Adams, William C.1914. Special report of the Board of Commissioners on Fisheries and Game,under Chapter 70 of the Resolves of 1913, relative to the habitsof those birds commonly known as pheasants. Commonwealthof Massachusetts, House Rep. no. 2049, pp. 1-14, Jan.Finley, William Lovell.1896. The Oregon ruffed grouse. Oregon Nat., vol. 3, no. 7, pp. 102, 103,illus., July.Fisher, Albert Kenrick.1893. Report on the ornithology of the Death Valley Expedition of 1891,comprising notes on the birds observed in southern California,southern Nevada, and parts of Arizona and Utah. North Amer.Fauna no. 7, pp. 7-158, May.Fleming, James Henry.1903. Recent records of the wild pigeon. The Auk, vol. 20, no. 1, p. 66, Jan.1907. The disappearance of the passenger pigeon. Ottawa Field Nat.,vol. 20, no. 12, pp. 236, 237, Mar.1930. Ontario bird notes. The Auk, vol. 47, no. 1, pp. 64-71, Jan.Forbush, Edward Howe.1912. A history of the game birds, wild-fowl and shore birds of Massachus-etts and adjacent States, 622 pp., illus., Boston.1927. Birds of Massachusetts and other New England States, vol. 2, 461pp., 27 col. pis. Boston.Fortiner, John C.1920. Winter nesting of the ground dove. The Condor, vol. 22, no. 4, pp.154, 155, July-Aug.1921. The doves of Imperial County, California. The Condor, vol. 23, no. 5,p. 168, Sept.-Oct.Fowler, Frederick Hall.1903. Stray notes from southern Arizona. The Condor, vol. 5, no. 3, pp.68-71, May-June.French, John C.1919. The passenger pigeon in Pennsylvania, its remarkable history, habits,and extinction, with interesting side lights on the folks and forestlore of the Alleghanian region of the old Keystone State, 257 pp.,illus. Altoona.Fuertes, Loins Agassiz.1903. With the Mearns quail in southwestern Texas. The Condor, vol. 5, no.5, pp. 113-116, illus., Sept.-Oct.Gabrielson, Ira Noel.1922. Short notes on the life histories of various species of birds. WilsonBull., vol. 34, no. 4. pp. 193-210, illus., Dec.
466 BULLETIN 162, UNITED STATES NATIONAL MUSEUMGardner, Leon Lloyd.1927. Habits of blue jays and doves in central Kansas. The Auk, vol. 44,p. 104, Jan.Giddings, H. J.1897. A quail's egg in a towhee's nest. The Osprey, vol. 2, no. 2, p. 26, Oct.Gilman, Marshall French.1903. More about the band-tailed pigeon (Columba fasciata). The Condor,vol. 5, no. 5, pp. 134, 135, Sept.-Oct.1911. Doves on the Pima Reservation. The Condor, vol. 13, no. 2, pp. 51-56,Mar.-Apr.1915. A forty-acre bird census at Sacaton, Arizona. The Condor, vol. 17, no.2, pp. 86-90, Mar.-Apr.Goss, Nathaniel Stickney.1891. History of the birds of Kansas, 693 pp., 35 pis. Topeka.Gosse, Philip Henry.1S47. The birds of Jamaica, 447 pp. London.Grayson, Andrew Jackson.1871. On the physical geography and natural history of the Tres Mariasand of Socorro, off the western coast of Mexico. Proc. Boston Soc.Nat. Hist., vol. 14, pp. 261-302, June 7.Green, Charles deBlois.1928. The fluttering habit of the Richardson's grouse (Dendragapus o&-scurus richardsoni). The Murrelet, vol. 9, no. 3, p. 67, Sept.Grinnell, Joseph.1900. Birds of the Kotzebue Sound region. Pacific Coast Avifauna, no. 1,80 pp., 1 map.1904. Midwinter birds at Palm Springs, California. The Condor, vol. 6, no.2, pp. 40-45, Mar.-Apr.1905. Summer birds of Mount Pinos, California. The Auk, vol. 22, no. 4,pp. 378-391, Oct.1906. The Catalina Island quail. The Auk, vol. 23, no. 3, pp. 262-265, July.190S. Catalina quail. The Condor, vol. 10, no. 2, p. 94, Mar.-Apr.1910. Birds of the 1908 Alexander Alaska Expedition, with a note on theavifaunal relationships of the Prince William Sound district. Univ.California Publ. Zool., vol. 5, no. 12, pp. 361-428, pis. 33, 34, 9 figs.,Mar. 5.1914. An account of the mammals and birds of the lower Colorado Valley.Univ. California Publ. Zool., vol. 12, pp. 51-294.1916. A new ruffed grouse from the Yukon Valley. The Condor, vol. 18,no. 4, pp. 166, 167, July-Aug.1926. Another new race of quail from Lower California. The Condor, vol.28, no. 3, pp. 128, 129, May-June.1928. A distributional summation of the ornithology of Lower California.Univ. California Publ. Zool., vol. 32, pp. 1-300.1928a. September nesting of the band-tailed pigeon. The Condor, vol. 30,no. 1, pp. 126, 127, Jan.-Feb.Grinnell, Joseph; Bryant, Harold Child; and Storer, Tracy Irwin.1918. The game birds of California, 642 pp., 16 col. pis., 94 figs. Berkeley.Grinnell, Joseph ; Dixon, Joseph ; and Linsdale, Jeax M.1930. Vertebrate natural history of a section of northern California throughthe Lassen Peak region. Univ. California Publ. Zool., vol. 35, 594pp., 181 figs. (1 col. map), Oct.
LITERATURE CITED 467Grinnell, Joseph ; Stephens, Frank ; Dixon, Joseph ; and Heller, Edmund.1909. Birds and mammals of the 1907 Alexander Expedition to south-eastern Alaska. Univ. California Publ. Zool., vol. 5, no. 2, pp. 171-264, pis. 25, 26, figs. 1-4, Feb. 18.Grinnell, Joseph ; and Storer, Tracy Irwin.1924. Animal life in the Yosemite, 752 pp., 60 pis., 65 figs., 2 eol. maps.Berkeley.Griscom, Ludlow.1926. Notes on the summer birds of the west coast of Newfoundland.The Ibis, ser. 12, vol. 2, no. 4, pp. 656-6S4, pis. 11-13, Oct.Gross, Alfred Otto.1928. The heath hen. Mem. Boston Soc. Nat. Hist., vol. 6, no. 4, pp. 491-588, 12 pis., May.Gurney, John Henry.1884. Hybrid between Pedioecetcs phasi(melius and Cupidonia cunido.The Auk, vol. 1, no. 4, pp. 391, 392, Oct.Hadley, Philip.1930. The passenger pigeon. Science, new ser., vol. 71, no. 1833, p. 187,Feb. 14.Hadzor, Rae T.1923. A grouse refuge. Bird-Lore, vol. 25, no. 5, p. 318, Sept.-Oct.Hagerup, Andreas Thomsen.1891. The birds of Greenland, 62 pp. Boston.Hall, Henry Marion.1929. An Indian summer day with ring-necks. Forest and Stream, vol. 99,no. 10, pp. 735, 756, 757, illus. Oct.Hantzsch, Bernhard.1929. Contribution to the knowledge of the avifauna of north-easternLabrador. Can. Field-Nat., vol. 43, no. 1, pp. 11-18, Jan. (Trans-lated by M. B. A. and R. M. Anderson.)Hartley, George Inness.1922. The importance of bird life, 316 pp., illus. New York.Henshaw, Henry Wetherbee.1874. Report upon the ornithological collections made in portions of Ne-vada, Utah, California, Colorado, New Mexico, and Arizona, dur-ing the years 1871, 1872, 1873. and 1874. Wheeler's Rep. Geogr.and Geol. Expl. and Surv. West 100th Meridian, vol. 5, Zoology,chap. 3, pp. 133-507, 15 pis.Higginson, Francis.1630. New England's plantations, 11 unpaged leaves, 7th and 8th leaves.London.Hill, Grace A.1922. With the willow ptarmigan. The Condor, vol. 24, no. 4, pp. 105-108,illus., July-Aug.Hodge, Clifton Fremont.1905. The drumming grouse. Country Calendar, vol. 1, no. 7, pp. 640-644,illus.1911. The passenger pigeon investigation. The Auk, vol. 28, no. 1, pp. 49-53,Jan.1912. A last word on the passenger pigeon. The Auk, vol. 29, no. 2, pp.169-175, Apr.Holland, Harold May.1917. The valley quail occupying nests of the road-runner. The Condor,vol. 19, no. 1, pp. 23, 24, illus., Jan.-Feb.
468 BULLETIN 162, UNITED STATES NATIONAL MUSEUMHollistee, Ned.1925. Another record of the ruddy quail-dove at Key West. The Auk, vol.42, no. 1, p. 130, Jan.Holzner, Frank Xavier.1596. Habits of the valley partridge. The Auk, vol. 13, no. 1, p. 81, Jan.Howard, Ozra William.1900. Nesting of the Mexican wild turkey in the Huachuca Mts., Arizona.The Condor, vol. 2, no. 3, pp. 55-57, illus., May-June.Howe, Carlton Durant.1904. A tame ruffed grouse. Bird-Lore, vol. 6, no. 3, pp. 81-85, illus
,
May-June.Howell, Alfred Brazier.1917. Condition of game birds in east-central California. The Condor, vol.19, no. 6, pp. 186, 187, Nov.-Dec.Hudson, William Henry.1902. British birds, with their structure and classification, 363 pp., 130pis. (8 col.).Huey, Laurence Markham.1913. With the band-tailed pigeon in San Diego County. The Condor, vol.15, no. 4, pp. 151-153, July-Aug.1927. Where do birds spend the night? Wilson Bull., vol. 39, no. 4, pp.215-217, Dec.Huntington, Dwight W.1597. The sage grouse. The Osprey, vol. 2, no. 2, pp. 17, 18, illus., Oct.1903. Our feathered game, 396 pp., illus. in col. New York.Jewett, Hibbard, J.1918. Memories of the passenger pigeon. Bird-Lore, vol. 20, no. 5, p.351, Sept.-Oct.Johnson, R. A.1929. Summer notes on the sooty grouse of Mount Rainier. The Auk, vol.46, no. 3, pp. 291-293, pis. 12, 13. July.Jones, Lynds.1903. A bob-white covey. Wilson Bull., no. 45, vol. 10, no. 4, pp. 105, 106,Dec.Josselyn, John.1672. New-England's rarities discovered, 114 pp., 1 pi., 12 figs. London.Judd, Sylvester Dwight.1905. The bobwhite and other quails of the United States in their economicrelations. U. S. Dept. Agr. Biol. Surv. Bull. 21, 66 pp., 2 cols, pis.,10 figs.1905a. The grouse and wild turkeys of the United States, and their eco-nomic value. U. S. Dept. Agr. Biol. Surv. Bull. 24, 55 pp., 2pis. (1 col.).Kalm, Pehr.1911. A description of the wild pigeons which visit the southern Englishcolonies in North America, during certain years, in incrediblemultitudes. The Auk, vol. 28, no. 1, pp. 53-66 (Translated by S. M.Gronberger from Kongl. Vetenskaps-Akademiens Handlingar forar 1759, vol. 20. Stockholm.)Kellogg, Louise.1916. Report upon mammals and birds found in portions of Trinity, Sis-kiyou and Shasta Counties, California, with description of a newDipodomys. Univ. California Publ. Zool., vol. 12, no. 13, pp. 335-398, pis. 15-18, Jan. 27.
LITERATURE CITED 469Kentwood.1S96. Martha's Vineyard heath hen. Forest and Stream, vol. 47, no. 18,pp. 343, 344, Oct. 31.Kermode, Francis.1914. Report of the Provincial Museum of Natural History for the year1913, pp. 1-32, illus.Keyes, Charles Rollin.1905. Some bird notes from the central Sierras. The Condor, vol. 7, no. 1,pp. 13-17, illus., Jan.Knight, Ora Willis.1908. The birds of Maine, 693 pp., illus., 1 map. Bangor.Kobbe, William Hoffman.1900. The birds of Cape Disappointment, Washington. The Auk, vol. 17, no.4, pp. 349-35S, Oct.Kumlien, Ludwig, and Hollister, Ned.1903. The birds of Wisconsin. Bull. Wisconsin Nat. Hist. Soc, new ser.,vol. 3, nos. 1-3, Jan-July.Lacey, Howard.1911. The birds of Kerrville, Texas, and vicinity. The Auk. vol. 28, no. 2,pp. 200-219, 1 map, Apr.LaHontan, Baron de.1703. New voyages to North America, 2 vols. London.Laing, Hamilton Mack.1913. Out with the birds, 249 pp., illus. New York.1925. Birds collected and observed during the cruise of the Thicpval in theNorth Pacific, 1924. Victoria Mem. Mus. Bull. 40.Lamb, Chester Converse.1926. The Viosca pigeon. The Condor, vol. 28, no. 6, pp. 262, 263, Nov.-Dec.Lawrence, George Neweold.1853. Descriptions of new species of birds of the genera Ortyx Stephens,Sterna Linn., and Icteria Vieillot. Ann. Lyceum Nat. Hist., vol. 6,pp. 1-4.1874. The birds of western and northwestern Mexico, based upon collec-tions made by Col. A. J. Grayson, Capt. J. Xantus and Ferd. Bis-choff, now in the museum of the Smithsonian Institution, at Wash-ington, D. C. Mem. Boston Soc. Nat. Hist., vol. 2, pp. 265-319,Apr. 25.18S9. Remarks upon abnormal coloring of plumage observed in severalspecies of birds. The Auk, vol. 6, no. 1, pp. 46-50, Jan.Leffingwell, Dana Jackson.1928. The ringed-neck pheasant, its history and habits. Occ. Pap. CharlesR. Conner Mus. no. 1, pp. 1-35, illus., Apr.Lewis. Elisha Jabrett.1885. The American sportsman, ed. 3, 553 pp.. illus. Philadelphia.Lewis, Evan.1904. The nesting habits of the white-tailed ptarmigan in Colorado. BirdLore, vol. 6, no. 4, pp. 117-121, illus., July-Aug.Lewis, Harrison Flint.192S. Notes on the birds of the Labrador Peninsula in 1928. Can. Field-Nat., vol. 42, pp. 191-194.74564?32 31
470 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMLincoln, Frederick Charles.1918. A strange case of hybridism. Wilson Bull. no. 102, vol. 30, no. 1,pp. 1, 2, illus. Mar.1930. Migratory status of mourning doves is proved by banding. U. S.Dept. Agr. Yearbook for 1930, pp. 386-388, map.Lloyd, William.18S7. Birds of Tom Green and Concho Counties, Texas. The Auk, vol. 4,no. 3, pp. 181-193, July.MacFarlane, Roderick Ross.1905. List of birds and eggs observed and collected in the North-West Ter-ritories of Canada, between 1880 and 1894, in Through the Mac-kenzie Basin, by Charles Mair, pp. 287-^90, illus. London.MACGILLIVRAY, WILLIAM.1837-1852. A history of British birds, 5 vols., illus. London.MacMillan, Donald Baxter.1918. Four years in the white north, 426 pp., illus. New York.Macoun, John, and Macoun, James Melville.1909. Catalogue of Canadian birds, ed. 2, 761 pp. Can. Dept. Mines. Geol.Surv. Branch.Manniche, Arner Ludvig Valdemar.1910. The terrestrial mammals and birds of northeast Greenland. Biolog-ical observations. Meddelelser om Gr0nland, vol. 45, no. 1, pp.1-199, pis. 1-7 (part col.), figs. 1-20.Maynard, Charles Johnson.1896. The birds of eastern North America, rev. ed., 721 pp., 40 col. pis., 122figs. Newtonville, Mass.McAtee, Waldo Lee.1929. Game birds suitable for naturalizing in the United States. U. S.Dept. Agr. Circ. 96, pp. 1-23, figs. 1-14, Nov.McAtee, Waldo Lee, and Beal, Foster Ellenborough Lascelles.1912. Seme common game, aquatic, and rapacious birds in relation to manU. S. Dept. Agr. Farmers' Bull. 497, pp. 1-30, 14 fi?s., May 6.McGregor, Richard Crittenden.1906. Birds observed in the Krenitzin Islands, Alaska. The Condor, vol. 8.no. 5, pp. 114-122, 1 map, Sept.McIlhenny, Edward Avery.1914. The wild turkey and its hunting, 245 pp., illus. Garden City.Mearns, Edgar Alexander.1879. Notes on some of the less hardy winter residents in the HudsonRiver Valley. Bull. Nuttall Orn. Club, vol. 4, no. 1, pp. 33-37, Jan.1914. Diagnosis of a new subspecies of Gambel's quail from Colorado.Proc. Biol. Soc Washington, vol. 27, p. 113, July 10.Merriam, Clinton Hart.1885. Change of color in the wing-feathers of the willow grouse. The Auk.vol. 2, no. 2, pp. 201-203, Apr.1888. What birds indicate the proximity to water, and at what distance?Auk, vol. 5, no. 1, p. 119, Jan.1889. Introduced pheasants. Rep. Comm. Agr. for 1S88, pp. 484-^88.1890. Results of a biological survey of the San Francisco Mountainregion and desert of the Little Colorado, Arizona. Annotated list oi'birds of the San Francisco Mountain plateau and the desert ofthe Little Colorado River, Arizona. North Amer. Fauna no. 3, pt. 4,pp. 87-101. Sept. 11.
LITERATURE CITED 471Merrill, James Cushing.1S78. Notes on the ornithology of southern Texas, being a list of birdsobserved in the vicinity of Fort Brown, Texas, from February, 1876,to June, 1878. Proc. U. S. Nat. Mus., vol. 1, pp. 118-173.Mershon, William Butts.1907. The passenger pigeon, 225 pp., col. pis. New York.Mlllais, John Gullle.1909. The natural history of British game birds. 142 pp., 8 pis. London.Miller, Waldron DeWitt.1905. List of birds collected in southern Sinaloa, Mexico, by J. H. Batty,during 1903-1. Bull. Amer. Mus. Nat. Hist., vol. 21, art. 22, pp.339-369.Minot, Henry Davis.1877. The land birds and game birds of New England. 456 pp., 1 pi. Salem.Morss, Charles B.1923. The signal: An episode of grouseland. Field and Stream, vol. 28,no. 6, pp. 689-691, 757-762, illus., Oct.Morton, Thomas.1637. New English Canaan, ed. 2, 188 pp. London.Moselet, Edwin Lincoln.1928. Bob-white and scarcity of potato beetles. Wilson Bull., vol. 40, no. 3,pp. 149-151, Sept.Murphy, Robert Cushman.1917. Natural history observations from the Mexican portion of the Colo-rado Desert. Abstract of Proc. Linnaean Soc. New York, nos. 28, 29,pp. 1-114, pis. 1-6, Dec. 11.Neff, Johnson A.1923. Some birds of the Ozark region. Wilson Bull., vol. 35, no. 4, pp.202-215, map, Dec.Nelson, Edward William.1887. Report upon the natural history collections made in Alaska, betweenthe years 1877 and 1881. Arctic ser., Signal Service, U. S. Army, pt.1, Birds, pp. 19-226, 12 col. pis.1899. Natural history of the Tres Marias Islands. Birds of the Tres MariasIslands. North Amer. Fauna no. 14, pp. 21-62, Apr. 29.1900. Description of a new subspecies of Meleagris gallopavo and proposedchanges in the nomenclature of certain North American birds. TheAuk, vol. 17, no. 2, pp. 120-126, Apr.Newton, Alfred.1893-1896. A dictionary of birds, 1088 pp., illus. London.Nice, Margaret Morse.1910. Food of the bobwhite. Journ. Econ. Ent, vol. 3, no. 3, pp. 295-313,June.1922-1923. A study of the nesting of mourning doves. The Auk, vol. 39, no.4, pp. 457-474, pi. 18, Oct. ; vol. 40, no. 1, pp. 37-58, Jan.Nicholson, Donald John.1928. Actions of baby Florida wild turkeys. Florida Nat., vol. 2, no. 1,p. 32, Oct.Noble, George Kingsley.1919. Notes on the avifauna of Newfoundland. Bull. Mus. Comp. Zool.,vol. 62, no. 14, pp. 543-568, Mar.Nuttall, Thomas.1832. A manual of the ornithology of the United States and of Canada.Land Birds, 683 pp., illus. Cambridge, Mass.
472 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMObebholser, Harry Church.1923. Notes on the forms of the genus Orcortyr Baird. The Auk, vol. 40, no.1, pp. 80-84, Jan.Osgood, Wilfred Hudson.1901. New subspecies of North American birds. The Auk, vol. 18, no. 2, pp.179-185, Apr.1904. A biological reconnaissance of the base of the Alaska Peninsula.North Amer. Fauna no. 24, 86 pp., 7 pis., Nov. 23.Pangburn, Clifford Hayes.1919. A three months' list of the birds of Pinellas County, Florida. TheAuk, vol. 36, pp. 393-405.Patch, Clyde A.1922. A biological reconnaissance on Graham Island of the Queen Char-lotte Group. Can. Field-Nat., vol. 36, pp. 101-105, 133-136.Payne, E. B.1897. Quail's eggs in a nieadowlark's nest. The Osprey, vol. 2, no. 4, p. 65,Dee.Pearson, Thomas Gilbert.1920. The ground dove. Bird-Lore, vol. 22, no. 2, pp. 126-129, illus.,Mar.-Apr.Pembebton, John Roy.1928. The nesting of Howard's grouse. The Condor, vol. 30, no. 6, pp. 347,348, illus., Nov.-Dec.Peters, James Lee.1923. A new quail from Lower California. Proc. New England Zool. Club,vol. 8, pp. 79, 80, May 16.Phelps, Frank Mills.1914. The resident bird life of the Big Cypress Swamp region. WilsonBull., no. 87, vol. 26, no. 2. pp. 86-101, illus., June.Phillips, John Charles.1928. Wild birds introduced or transplanted in North America. U. S.Dept. Agr. Techn. Bull. 61, 64 pp.Pokagon, Chief Simon.1895. The wild pigeon of North America. Chautauquan, vol. 22, no. 2,pp. 202-206, Nov.Pbeble, Edward Alexander.1908. A biological investigation of the Athabaska-Mackenzie region. NorthAmer. Fauna no. 27, 574 pp., 25 pis., 16 figs., map, Oct. 26.Quillin, Roy W., and Holleman, Ridley.1918. The breeding birds of Bexar County, Texas. The Condor, vol. 20, no.1, pp. 37-44, Jan.-Feb.Revoil, Benedict Henry.1869. Chasses dans l'Amerique du Nord. (See Bird-Lore, vol. 30, no. 5,pp. 317-320, Sept-Oct, 1928.)Rich, Walter Hebiiert.1907. Feathered game of the Northeast, 432 pp., illus. New York.1909. The " Hungarian Partridge "?the gray partridge. Journ. Maine Orn.Soc, vol. 11, no. 2, pp. 33-38, illus. June.RlDGWAY, ROBEBT.1912. Color standards and color nomenclature, 43 pp., 53 col. pis. Wash-ington, D. C.1915. Descriptions of some new forms of American cuckoos, parrots, andpigeons. Proc. Biol. Soc. Washington, vol. 28, pp. 105-108,May 27.
LITERATURE CITED 473Ridgway, Robert.1916. The birds of North and Middle America, U. S. Nat. Mus. Bull. 50,pt. 7, 543 pp., 24 pis.Riley, Joseph Harvey.1911. Descriptions of three new birds from Canada. Proc. Biol. Soc. Wash-ington, vol. 24, pp. 233-235, Nov. 28.1912. Birds collected or observed on the expedition of the Alpine Club ofCanada to Jasper Park, Yellowhead Pass, and Mount Robson region.Can. Alpine Journ., special number, pp. 47-75, pi. 1, 1912.Roberts, Thomas Sadler.1919. A review of the ornithology of Minnesota. Research publications ofthe Univ. of Minnesota, vol. 8, no. 2, 100 pp., illus., May.Roney, H. B.1879. A description of the Michigan pigeon nesting of 1878 and the workof protection undertaken by the East Saginaw and Bay City gameprotection clubs. Chicago Field, vol. 10, no. 22, pp. 345-347, Jan. 11.Rowan, William.1926. Comments on two hybrid grouse and on the occurrence of Tym-panuchus americanus americanus in the Province of Alberta. TheAuk, vol. 43, no. 3, pp. 333-336, pis. 17, 18, July.Sage, John Hall; Bishop, Louis Bennett; and Bliss, Walter Parks.1913. The birds of Connecticut. Connecticut State Geol. and Nat. Hist.Surv., Bull. 20, Pub. Doc. no. 47, pp. 1-370.Samuels, Edward Augustus.1883. Our northern and eastern birds, 600 pp., illus., part col. pis. NewYork.Sandys, Edwyn, and Van Dyke, T. S.1904. Upland game birds, 429 pp., illus. London.Sawyer, Edmund Joseph.1923. The ruffed grouse, with special reference to its drumming. Roose-velt Wild Life Bull., vol. 1, no. 3, pp. 355-384, illus.Schutze, Adolf Emil.1904. The Inca dove in central Texas. The Condor, vol. 6, no. 6, p. 172.Nov.-Dec.Sclater, William Lutley.1912. A history of the birds of Colorado, 576 pp., 17 pis., map. London.Scott, William Earl Dodge.1889. Records of rare birds at Key West, Florida, and vicinity, with a noteon the capture of a dove (Oeotrygon montana) new to NorthAmerica. The Auk, vol. 6, no. 2, pp. 160, 161, Apr.1890. Description of a new subspecies of wild turkey. The Auk, vol. 7, no. 4,pp. 376, 377, Oct.1892. Notes on the birds of the Caloosahatchie region of Florida. The Auk,vol. 9, no. 3, pp. 209-218, July.Sennett, George Burritt.1878. Notes on the ornithology of the lower Rio Grande of Texas fromobservations made during the season of 1877. Bull. U. S. Geol.and Geogr. Surv. Terr., vol. 4, art. 1, pp. 1-66.1879. Further notes on the ornithology of the lower Rio Grande of Texas.Bull. U. S. Geol. and Geogr. Surv. Terr., vol. 5, no. 3, pp. 371^40.1892. Description of a new turkey. The Auk, vol. 9, no. 2, pp. 167-169, Apr.Seton, Ernest E. Thompson.18S5. Notes on Manitoban birds. The Auk, vol. 2, no. 3, pp. 267-271, July.
474 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMShaw, William Thomas.190S. The China or Denny pheasant in Oregon, with notes on the nativegrouse of the Pacific Northwest, 24 pp., 15 pis. Philadelphia.Simmons, George Finlay.1915. On the nesting of certain birds in Texas. The Auk, vol. 32, no. 3, pp.317-331, pis. 21, 22, July.1925. Birds of the Austin region, 387 pp., illus. Austin.Simpson, Gene M.1914. Pheasant farming. Oregon Fish and Game Comm. Bull. 2, 47 pp.,illus., 1 col. pi.Skinner, Milton Philo.1927. Richardson's grouse in the Yellowstone Park. Wilson Bull., vol. 39,no. 4, pp. 208-214, Dec.Slade, Daniel Dennison.1888. The wild turkey in Massachusetts. The Auk, vol. 5, no. 2, pp. 204, 205,Apr.Smith, Austin Paul.1910. Miscellaneous bird notes from the lower Rio Grande. The Condor,vol. 12, no. 3, pp. 93-103, May-June.1916. Additions to the avifauna of Kerr Co., Texas. The Auk, vol. 33, no. 2,pp. 187-193, Apr.Smith, Everett.1883. Birds of Maine. Forest and Stream, vol. 20, pp. 25-61.Snyder, John Otterbein.1900. Notes on a few species of Idaho and Washington birds. The Auk, vol.17, no. 3, pp. 242-245, July.Spiker, Charles J.1929. The Hungarian partridge in northwest Iowa. Wilson Bull., vol. 41.no. 1, pp. 24-29, illus., 1 map, Mar.Spurrell, John A.1917. Annotated list of the water birds, game birds, and birds of prey ofSac County, Iowa. Wilson Bull., no. 100, vol. 29, no. 3, pp. 141-160,Sept.Stejneger, Leonhard.18S4. A new subspecies of willow grouse from Newfoundland. The Auk, vol.1. no. 4, p. 369, Oct.Stephens, Frank.1885. Notes on an ornithological trip in Arizona and Sonora. The Auk,vol. 2, no. 3, pp. 225-231, July.1903. Bird notes from eastern California and western Arizona. The Con-dor, vol. 5, no. 3, pp. 75-78, May-June.Stockard, Charles Rupert.1905. Nesting habits of birds in Mississippi. Auk, vol. 22, no. 2, pp. 146-158, Apr.Stoddard, Herbert Lee.1922. Notes on birds from southern Wisconsin. Wilson Bull., vol. 34, no2, pp. 67-79, June.1925. Progress on cooperative quail investigation : 1924.1926. Report on cooperative quail investigation: 1925-26, with preliminaryrecommendations for the development of quail preserves, 62 pp.,5 pis.1931. The bobwbite quail, its habits, preservation, and increase. 559 pp.,69 pis. (part col.), 32 figs. New York.
LITERATURE CITED 475Stone, Witmer.1905. On a collection of birds and mammals from the Colorado Delta,Lower California. Proc. Acad. Nat. Sci. Philadelphia, vol. 57, pp.676-690, Sept.Sutton, Geoege Miksch.1928. The birds of Pymatuning Swamp and Conneaut Lake, CrawfordCounty, Pennsylvania. Ann. Carnegie Mus., vol. 18, no. 1, pp.19-239, pis. 2-11 (in col.), Mar.1929. Photographing wild turkey nests in Pennsylvania. The Auk, vol. 46,no. 3, pp. 326-328, pi. 18, July.Swainson, William, and Richardson, John.1831. Fauna Boreali-Americana, vol. 2, Birds, 523 pp.. col. pis. and figs.London.Swarth, Harry Schelwaldt.1904. Birds of the Huachuca Mountains, Arizona. Pacific Coast Avifaunano. 4, 70 pp.1909. Distribution and molt of the Mearns quail. The Condor, vol. 11, no. 2.pp. 39-43, illus., Mar.-Apr.1912. Report on a collection of birds and mammals from Vancouver Island.Univ. California Publ. Zool., vol. 10, no. 1. pp. 1-124, pis. 1-4,Feb. 13.1920. Birds of the Papago Saguaro National Monument and the neighboringregion, Arizona, 63 pp., 8 pis. Washington, D. C.1921. The Sitkan race of the dusky grouse. The Condor, vol. 23, no. 2, pp.59, 60, Mar.-Apr.1922. Birds and mammals of the Stikine River region of northern BritishColumbia and southeastern Alaska. Univ. California Publ. Zool. rvol. 24, no. 2, pp. 125-314.1924. Birds and mammals of the Skeena River region of northern BritishColumbia. Univ. California Publ. Zool., vol. 24, no. 3, pp. 315-394,illus.1926. Report on a collection of birds and mammals from the Atlin region,northern British Columbia. Univ. California Publ. Zool., vol. 30^no. 4, pp. 51-162, pis. 4-8 (1 in col.), 11 figs., Sept. 24.Swenk, Myron Harmon, and Swenk, Jane Bishop.1928. Some impressions of the commoner winter birds of southern Arizona.Wilson Bull., vol. 40, no. 1. pp. 17-29, Mar.Tarer, William Brewster, jr.1930. The fall migration of mourning doves. Wilson Bull., vol. 42, pp. 17-28.Taverner. Percy Algernon.1914. A new subspecies of Dendragapus (Dendragapus obseums flemingi)from southern Yukon Territory. The Auk. vol. 31, no. 3, pp. 385-388,July.1926. Birds of western Canada. Can. Dept. Mines, "Victoria Mem. Mus.Bull. 41, 360 pp., 84 pis.1929. A study of the Canadian races of rock ptarmigan (Lagopus rupestris)
.
Ann. Rep. Nat. Mus. Canada for 1928. pp. 28-3S.Taylor, Walter Penn.1920. A new ptarmigan from Mount Rainier. The Condor, vol. 22, no. 4, pp.146-152, illus., July-Aug.Taylor, Walter Penn, and Shaw, William Thomas.1927. Mammals and birds of Mount Rainier National Park. U. S. Dept.Int., 249 pp., 109 figs.
476 BULLETIN 162, UNITED STATES NATIONAL MUSEUMTegetmkieb, William Bernhard.1911. Pheasants, their natural history and practical management, ed. 5,276 pp., illus. (part col. plates). London.Thompson, Charles S.1901. Further tape-worm observations. The Condor, vol. 3, no. 1, p. 15,Jan.-Feb.Thompson, Ebnest Evan (Seton, Ernest Thompson-).1S90. The birds of Manitoba. Proc. U. S. Nat. Mus., vol. 13, pp. 457-643,pi. 38.Thompson, Frank J.1881. Breeding of the wild pigeon in confinement. Bull. Nuttall Orn. Club,vol. 6, no. 2, p. 122, Apr.Todd, "Walteu Edmond Clyde.1916. The birds of the Isle of Pines. Ann. Carnegie Mus., vol. 10, no. 12,art. 11, pp. 146-296, pis. 22-27, Jan. 31.Town send, Charles Wendell.1906. Note on the crop contents of a nestling mourning dove. The Auk, vol.23, no. 3, pp. 336, 337, July.1912. The case of a crow and a ruffed grouse. The Auk, vol. 29, no. 4, p.542, Oct.1920. Supplement to the birds of Essex County, Massachusetts. Mem. Nut-tall Orn. Club, no. 5, 196 pp., 1 pi., 1 map.Turner, Lucien McShan.1886. Contributions to the natural history of Alaska. Signal Seiwice, U. S.Army, No. 2, pt. 5, Birds, pp. 115-196, 10 col. pis.Tyler, John Gbipper.1913. Some birds of the Fresno district, California. Pacific Coast Avifaunano. 9, 114 pp.Van DeWeyer, B.1919. Pheasant breeding in sparrow hawk's nest. British Birds, vol. 13, no.3. p. 87, Aug. 1.Van Dyke, Theodore Strong.1892. The valley quail of California. Outing, vol. 19, pp. 485-488, Mar.Van Rossem, Adriaan Joseph.1925. Flight feathers as age indicators in Dendragapus. The Ibis, ser. 12.vol. 1, no. 2, pp. 417-422, figs. 10-12, Apr.Vreeland, Frederick K.1918. The drumming of the ruffed grouse. Forest and Stream, vol. 88, no. 4.pp. 199-201, 244, illus., Apr.Wales, Joseph Howe.1926. The coo of the band-tailed pigeon. The Condor, vol. 28, no. 1, p. 42,illus., Jan.-Feb.Warner, Edward Pearson.1911. The history of a ruffed grouse's nest. Bird-Lore, vol. 13, no. 2, p. 99.Mar-Apr.Wayne, Arthur Trezevant.1910. Birds of South Carolina. Contr. Charleston Mus. no. 1, 254 pp.,1 map.Wetmore, Alexander.1916. Birds of Porto Rico. U. S. Dept. Agr. Bull. 326, 140 pp., 10 pis.,Mar. 24.1920. Observations on the habits of the white-winged dove. The Condor,vol. 22, no. 4, pp. 140-146, July-Aug.
LITERATURE CITED 477Wetmoee, Alexandeb.1927. The birds of Porto Rico and the Virgin Islands. Scientific Surv. ofPorto Rico and the Virgin Islands, vol. 9, pt. 3, pp. 245-406. NewYork Acad. Sci.Wheaton. John Maynabd.1882. Report on the birds of Ohio. Rep. Geol. Surv. Ohio. vol. 4, pt. 1,>logy, pp. 187-628.Wheelock, Ibene Geosvenob.1904. Birds of California, 578 pp., 10 pis., 78 figs. Chicago.Wicks, Morie Laxgley, jk.1897. Partnership nesting of valley partridge and long-tailed chat. The Auk,vol. 14, no. 4. p. 404, Oct.Widmann, Otto.1907. A preliminary catalog of the birds of Missouri, 288 pp. St. Louis.WlLLABD, FBANCIS COTTLE.1910. Nesting of the band-tailed pigeon in southern Arizona. The Condor,vol. IS. no. 3, pp. 110-112 illus., May-June.WlLLETT, GeOBGE.1914. Birds of Sitka and vicinity, southeastern Alaska. The Condor, vol. 16.no. 2, pp. 71-91, illus., Mar.-Apr.Williams, John J.1 903. On the use of sentinels by valley quail. The Condor, vol. 5, no. 6, pp.146-148, illus., Nov.-Dec.Wh^son Alexandeb.1832. American ornithology, vol. 1, 408 pp., illus. London.Withebby. Habry Foebes, and others.1919-1920. A practical handbook of British birds, vol. 1, 532 pp.. 17 pis.(part col.). London.Wood, William.1635. New England prospect, Chap. 8, pp. 22-27. London.Weight, Albebt Hazen.1914-1915. Early records of the wild turkey. The Auk, vol. 31, nos. 3, 4, pp.334-358, and 463-473, July, Oct. ; vol. 32, nos. 1, 2, pp. 61-81, and207-224, Jan., Apr.Wyman, Lutheb Eveeet.1921. A very late record of the passenger pigeon (Ectnpistes migratorius).The Auk, vol. 38, no. 2, p. 274, Apr.Yabbell, Wiltjam.1871-1885. A history of British birds, ed. 4, 4 vols., illus., revised andenlarged by Alfred Newton and Howard Saunders. London.

U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 1
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 2
Nesting Site and Nest of BobwhiteBerkley, Mass., June 28, 1915; both photographs by the author. Referred to on page 13.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 3
Male Bobwhite on NestOn the grounds of the Portland Golf Club, Oregon, 1915. Presented by Mr. and Mrs. William L.Finley.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 4
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 5
Nest of Florida BobwhiteOrlando, Fla., April 25, 1925. Presented by Jack II. Connery.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 6
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 7
Nest of Mountain QuailJennings Lodge, Oreg., May 7, lfe'OS. Presented by Mr. and Mrs. William L. Finley.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 8
Downy Young Mountain Quail HidincSoda's ill*-. Oreg. Presented by Leslie L. Haskin.
Immature Mountain QuailJennings Lodge, I ?reg. Presented by Mr. and Mrs. William L. Finley.
U. S. NATIONAL. MUSEUM BULLETIN 162 PLATE 9
Downy Young Scaled QuailPresented by Miss Angeline M. Keen
Nest of Arizona scaled QuailCochise County, Ariz., May 25, 1922. Photograph by the author. Referred to on page 53.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 10
Valley Quail FeedingIn yard of J. Eugene Law, Altadena, Calif., March 24. l!)29. Photograph by the author. Referredto on page 62.
Female Valley Quail<>n nest in unusuallj open situation, near Claremont, Calif. Presented bj Wright M, Pierce.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 11
Nest of Valley QuailNear Claremont, Calif., May 26, 1916. Presented by Wrighf M. Pierce.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 12
Two Views of valley Quail and Her NestVentura County, Calif. Presented by Donald R. Dickey.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 13
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 14
Nesting Site and Nest of Gambel's QuailPima County, Ariz. Presented by Frank C. Willard.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 15
Haunts and Nesting site of mearnss QuailHuachuca Mountains, Ariz. Presented by Frank ('. Willard.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 16
Hungarian PartridgeCorvallis, Oreg. Presented by Glaus J. Murie.
Nest of Hungarian partridgeWaukesha County, Wis.. July, 1927. Presented by Dr. Alvin R. Calm.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 17
v * . $&female dusky GrouseLittle Gros Ventre River, Wyo., July 7, 1904. Presented by E. R. Warren.
Nest of Dusky Grouse under SagebrushGunnison County, Colo., June 21, 1900. Presented by E. R. Warren.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE If
Two Views of Courtship Display of Richardson's GrouseFrom drawings loaned by Maj. Allan Brooks.
U. S. NATIONAL MUSEUM BULLETIN 16?. PLATE 19
Nesting Site of Sooty GrouseNear Dayton, Oreg., May 15, 1905. Presented by William L. Finley and II. T. Bohlman.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 20
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Nest of Sooty GrouseMulin<>. Oreg., April 15, 1914. Presented by Alexander Walker.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 21
Female Sooty GrousePresented by Mr. and Mrs. William L. Finley.
Nest of Sooty GrouseSodaville, Oreg., April 22, 1919. Presented by Leslie L. Haskin.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 22
Female Canada Spruce Grouse on NestNew Brunswick, Canada, Presented by P. B. Philipp.
Female Canada Spruce Grouse on NestPresented by L. W. Brownell.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 23
hudsonian Spruce Grouse and its NestBelvedere, Alberta, June 13, 1923. Both photographs presented bj J, Fletcher Street.
U. S NATIONAL MUSEUM BULLETIN 162 PLATE 24
Male Franklins GrousePresented by L. W. Brownell.
Female Franklins Grouse
( Uacier National Park, August, llL's. Presented by William L, Finley.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 25
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U. S. NATIONAL. MUSEUM BULLETIN 162 PLATE 27
DRUMMING OF THE RUFFED GROUSE?CONTINUEDPresented by Dr Arthur A. Allen. Frames 100 to 103 show the beginning of the terminal roll, wherethe wings are blurred. Note the end of the roll, in frame 120, followed by a forward and downwardstroke, in frame 122, to help the bird regain its balance, as it pitches forward and the tail springsupward in frames 122 and 123. Referred to on page 1 15.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 21
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 29
Female Ruffed Grouse on NestOakland County, Mich. Presented by Walter E. Hastings.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 30
Ruffed-grouse Nest in pine SlashBerkley, Mass.. May 21, 1903.
RUFFED-GROUSE NEST IN OAK WOODSRehoboth, Mass., May 18, 1921. Both photographs by the author. Referred to on page 148.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 31
Ruffed-grouse Chick 2 Days Old
Ruffed-grouse Chick 5 Days OldBoth photographs presented by Dr. Arthur A. Allen.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 32
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 33
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 34
Ruffed-grouse Chicks Hiding
i Irand Traverse County. Mich.. June 12, L927. Presented by Walter E. Hastings.
IMMATURE RUFFED GROUSE 130 DAYS OLD. IN FIRST WINTERPlumagePresented by Dr. Arthur A. Allen.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 35
Ruffed Grouse beginning to StrutPresented by W. J. Breckenridge and reproduced by courtesy of the Museum of Natural History,University of Minnesota.
Ruffed Grouse in Full DisplayPresented by Dr. Arthur A. Allen.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 36
U. S. NATIONAL MUSEUM BULLETIN 162 FLATE 37
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Partially Concealed Nest of Willow ptarmiganSt. Michael, Alaska, June 8, 1914.
?A?! ' *Open Nest of Willow ptarmiganYukon Delta, Alaska, June 1 1, l'.'i t. Both photographed bj !'. Sej mour Bersey for the author.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 39
Willow-ptarmigan ChickYukon Delta, Alaska, Juno 23, 1914.
Female Willow ptarmigan on NestYukon Delta, Alaska, June 16, 1914. Both photographed by F. Seymour Hersey for the author.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 40
Female willow ptarmigan Brooding YoungPresented by Joseph Dixon, by courtesy of the Museum of Vertebrate Zoology, University of California.
MALE WILLOW PTARMIGAN IN LATE-SUMMER PLUMAGEMount McKinley National Park, Alaska, September, 1926. Presented by Mr. and Mrs William L.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 41
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 42
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IMMATURE ROCK PTARMIGANSavage River, Alaska, September 6, 1922
Young Rock ptarmigan hidingIgiak Bay, Alaska, July 11, 1924. Both photographs by Olaus J. Murie; presented by the UnitedStates Biological Survey.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 44
Two Views of Rock Ptarmigan, Changing into Winter PlumageMount MeKinley National Park, Alaska, September, 1926. Presented by Mr. and Mrs. WilliamFinley. Note the concealing coloration in the upper figure.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 45
Nesting Site and Nest of Chamberlain's PtarmiganAdak fsland, Alaska, June 27, 1911. Photographs by the author. Referred to on pat'.' 222.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 46
Nesting Site and nest of sanfords PtarmiganTanaga Island, Alaska, June 2.r>, 1911. Photographs by Hollo II. Heck. Referred to on page 22
U. S NATIONAL MUSEUM BULLETIN 162 PLATE 47
Female Southern White-tailed Ptarmigan on NestIdaho Springs, Colo. Photograph taken by Evan Lewis and presented by E. K. Warren. Noteconcealing coloration.
Nest of Southern White-tailed ptarmiganPresented by the Colorado Museum of Natural History.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 48
Southern White-tailed Ptarmigan ChickGunnison County, Colo., July 8, 1902. Presented by E. R. Warren.
Larger Chick of Washington White-tailed PtarmiganMount Rainier, W ash . August 2, 1929. Presented by Dr. Oayle B. Pickwell.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 49
Southern White-tailed ptarmigan in Winter PlumageMarch 13, 1902.
Southern White-tailed ptarmigan Beginning to acquire NuptialPlumageMay 30, 1900. Both photographs taken by E. R. Warren in Gunnison County, Colo., and presentedby him.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 50
Female Southern White-tailed Ptarmigan Brooding ChicksGunnison County, Colo., July 11, 1927.
White-tailed PtarmiganGlacier National Park, Mont.. June 18, 1927. Both photographs presented by E. K. Warren.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 51
Washington White-tailed PtarmiganMounl Rainier, Wash., August, 1919.
Nest of Washington White-tailed ptarmiganMount Rainier, Wash., July 11, 1919. Both photographs presented by Mr. and Mrs. William L. Finlej .
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 52
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 53
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 54
Prairie Chicken Approaching Her NestJune 7, 1929.
Prairie Chicken on Her NestJune ?"', 1929. Botb photographs presented by Pr. A. 0. <in>ss. by courtesy of the Wisconsin Con-servation Commission.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 55
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Two Views of Male Prairie Chickens FightingPhotographs taken by Walter W. Bennett in northwestern Nebraska and presented by him.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 55
Typical Grass Nest of Prairie ChickenBismarck, X. Dak., May 24, 1925. Presented by Russell Keid.
Cjtfr&i It%^Brush-land Nest of Prairie ChickenKittson County, Minn , in timber on the outskirts of a village. Presented by the Rev. P. B.Pe ibody.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 57
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 58
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 59
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Heath Hen in Initial Stage of BoomingApril 12, 1923.
Heath Hen at Climax of Booming PerformanceMarch 29, 1927. Both photographs taken by Dr. A. 0. Gross, Marthas Vineyard, Mass., and pre-sented by him. Referred to on page 271.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 60
Nesting Site and Nest of the Heath Hen
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Female Heath hen on NestBoth photographs taken by Dr. George W. Field, Marthas Vineyard, Mass.. and presented by theMassachusetts Division of Fisheries and Game.
U. S. NATIONAL MUSEUM BULLETIN 1C2 PLATE 61
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Male Lesser Prairie ChickenOctober. 1925.
Nest of Lesser Prairie Chicken
-Tune 2, 1920. Referred to on page 281. Both photographs taken by Walter < '<>1\ in in Seward County,Kans., and presented by him.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 62
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Habitat of Prairie Sharp-tailed GrouseBabcock region, Wisconsin, July 29, 1930. Presented by Dr. A. O. Gross, by courtesy of the Wis-consin Conservation Commission,
Grass Nest of Prairie Sharp-tailed GrouseNear Lake Winnipegosis, Manitoba, June 2, 1913. Photographed by the author.
Prairie Sharp-tailed Grouse on Her Nest
Brush Nest of Prairie Sharp-tailed GrouseBoth photographs taken by the author near Crane Lake. Saskatchewan, June ?i, 1905. Referred to(H) page 293.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 64
Two Views of Prairie Sharp-tailed Grouse in WinterPhotographs by H. H. Pittman.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 65
Three Views of Prairie Sharp-tailed Grouse in Fighting attitudesduring CourtshipPresented by \V. J. Rreekenridge, by courtesy of the Museum of Natural History, University ofMinnesota.
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Front and Side Views of Courtship Display of Sage HenNear Lower Klamath Lake, Calif., May 12, 1917. Presented by William L. Finley.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 63
NEST OF SAGE HENNear Spencer, Idaho. Presented by Henry T. Rust.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 69
Sage-hen Chick
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Young Sage hen HidingBoth photographs presented by the Rev. P. B. Peabody.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 70
Nest and nesting Site of Ring-necked PheasantKavnham, .Mass., May 16, L920. Photographed by the author.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 71
Nest of Ring-necked PheasantNear Dayton, Oreg., May 15, 1905. Presented by William L. Finley and B. T, Bohlman,
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 72
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RING-NECKED PHEASANT CHICKDayton, Ureg.. May 15, 1905. Presented by William L. Finley and H. T. Bohhnan.
Male Ring-necked PheasantCorvallis, Oreg. Presented by 0. J. .Murie.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 74
Nest of Wild TurkeyXear Falls Church, Va., May 10. 1903. Presented by Dr. Paul Bartsch.
Wild Turkey on Her NestClearfield County, Pa., May 5, 1928. Presented bj l>r. George M. Sutton.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 75
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 77
Nest of Merriam's TurkeyAt fool of a live-oak on a hillside, Santa Rita Mountains, Ariz., June 15, l y M.Presented by Frank Stephens.
Nest and Eggs of Merriams TurkeyHuaehuca Mountains, Ariz., July 1, 1900. Photograph taken by 0. \V. Howard.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 78
Flock of Band-tailed Pigeons in Characteristic Resting PosePresented by Mrs. \V. I.eon Dawson.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 79
BAND-TAILED PIGEONSCorvallis, Oreg. Presented by 0. J. Murie.
BAND-TAILED PIGEON PORTRAITPresented by Joseph H. Wales.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 80
Nest and Egg of band-tailed pigeonIn A. M. [ngersoll's collection. Photograph presented by him.
Nest of Band-tailed PigeonHuachuca Mountains, Ariz. Presented by Frank C. Willard.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 81
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 83
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U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 84
Young Mourning Doves in Their Nest
nest and Eggs of Mourning DoveBoth photographs taken near Buffalo, X. v., Maj 8, 1927, and presented by S. A. Crimes.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 85
Nest of Mourning Dove in Old hawk's NestMiddleboro, Mass., April 28, 1907. Referred to on page 406.
Nest of Mourning Dove on the GroundNear Stump Lake, N. Dak., June 5, 1901. Both photographs by the author.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 86
Mourning Dove on Her Nest on Low stumpRehoboth, Mass., May 29, 1920. Photograph by the author.
Nest of Western Mourning Dove Under Fallen TreeNear Brownsville, Oreg. Presented by Mrs. Lillian <!. Haskin.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 87
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Immature Western White-winged Doves
Young Western White-winged Doves in NestBoth photographs taken near Tucson, Ariz., and presented by Mr. and Mrs. William L. I-inlcv.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 90
Kra* nest of Western White-winged DoveNear Tucson, Ariz. Presented by Mr. and Mrs. "William L. Finley.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 91
MALE GROUND DOVE AT FEEDING STATION
'V??rFemale Ground Dove Entering Nest on the Ground underPalmettoBoth photographs taken at St. Cloud, Fla., and presented by Dr. Arthur A. Allen.
U. S. NATIONAL MUSEUM BULLETIN 162 PLATE 92
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Nest of Mexican Ground DoveFairbank, Ariz., May 17, 1922. Photograph by the author. Referred to onpage 441.
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Nest of Mexican Ground DoveXear Tucson, Ariz. Presented by -Mr. and Mrs. William L. Finley.



INDEX
Abbott, Clinton G., x.on band-tailed pigeon, 356.achrustera, Lophortyx californica, 61,72.Adams, A. L., on Hudsonian sprucegrouse, 126, 127.Adams, W. C, x.on ring-necked pheasant, 312.alascensis, Lagopus, 190, 200.Alaska ptarmigan, 200.Alaska spruce grouse, 129.albus, Lagopus lagopus, 178, 190.Alexander, Miss Annie M., 195.Alexander, M. L., on eastern turkey,329.Alexander's ptarmigan, 194.alexandrae, Lagopus lagopus, 191, 194.Allen, A. A., x.on ruffed grouse, 142.Allen, Ena A., 29.Allen, G. M., on eastern turkey, 328.Allen, J. A., on masked bobwhite, 37, 38.alleni, Lagopus lagopus, 190, 191.Allen's ptarmigan, 191saltipetens, Lagopus leucurus, 233, 234.americanus, Tvmpanuchus cupido, 242,263.Anderson, R. M., on greater prairiechicken, 260.on Reinhardt's ptarmigan, 213.angelica, Leptotila fulviventris, 422.Anthony, A. W., on Richardson'sgrouse, 97, 99, 101.on San Pedro quail, 51.on sooty grouse, 109.on southern white-tailed ptarmi-gan, 235.Arizona scaled quail, 51.asiatica asiatica, Melopelia, 425, 427.mearnsi, Melopelia, 427, 428.asiatica, Melopelia asiatica, 425, 427.atkhensis, Lagopus rupestris, 212, 218.Atkins, J. W., 450, 453.on scaled pigeon, 376.atratus, Canachites canadensis, 129,135.Attwater, H. P., 263.attwateri, Tympanuchus cupido, 263.Attwater's prairie chicken, 263.Audubon, J. J., on blue-headed quail-dove, 456.on eastern bobwhite, 25.on eastern ground dove, 436.on eastern turkey, 328, 329, 330,332, 334, 335, 336, 337, 338, 339.on heath hen, 277, 279.on Hudsonian spruce grouse, 127.
Audubon, J. J., on Key West quail-dove, 450, 451.on passenger pigeon, 383, 385, 388,391, 393, 396.on ruffed grouse, 143, 146, 158.on white-crowned pigeon, 369, 370,371, 376.on Zenaida dove, 417, 418, 419, 420.Aughey, Samuel, on greater prairiechicken, 257.on passenger pigeon, 388.Badger, M. C., x.Bahama ground dove, 443.bahamensis, Columbigallina passerina,443.Bailey, A. M., on Alexander's ptar-migan, 195, 196.on Dixon's ptarmigan, 224, 225.on Sitka grouse, 120.on willow ptarmigan, 188.Bailey, Mrs. Florence M., on Arizonascaled quail, 52, 54, 55, 56.on band-tailed pigeon, 359.on Columbian sharp-tailed grouse,289.on dusky grouse, 91, 92, 93.on Franklin's grouse, 138.on Gambel's quail, 74.on gray ruffed grouse, 171.on Mearns's quail, 88, 90.on Merriam's turkey, 324.on Richardson's grouse, 100, 101.on southern white-tailed ptarmigan,235, 236, 238.on Texas bobwhite, 35.on western white-winged dove, 429.Bailey, H. H., on white-crownedpigeon, 375.Bailev, Vernon, on Rio Grande turkey,344.Baird, Brewer, and Ridgway, on eas-tern white-winged dove, 425.on Gambel's quail, 80.on northern sharp-tailed grouse,286.on Oregon ruffed grouse, 174.on white-crowned pigeon, 373.Baker, J. H., x.Bancroft, Griffing, on masked bob-white, 39.on Mexican ground dove, 440, 442.on San Lucas quail, 72, 73.on San Quintin quail, 72.Band-tailed pigeon, 353.Bangs, Outram, on Nova Scotia ruffedgrouse, 177. 479
480 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMBangs and Penard, on white-fronteddove, 422.Banks, J. W., on Canada ruffed grouse,167.Barbour, Thomas, on blue-headedquail-dove, 457.on Key West quail-dove, 450, 452.on ruddy quail-dove, 453.on scaled pigeon, 376.on white-crowned pigeon, 373.on Zenaida dove, 417, 419.Barlow, Chester, on plumed quail, 45.Barlow and Price, on plumed quail, 50.Barnes, C. T., on valley quail, 62.Barrows, W. B., on eastern mourningdove, 403.on passenger pigeon, 382, 399.Bartlett, Moses, on rock ptarmigan,209.Bartram, John, on ruffed grouse, 142,143.Bartsch, Paul, x.on white-crowned pigeon, 371.Bates, S. P., on passenger pigeon, 392.Beatty, H. A., on scaled pigeon, 377.Beebe, William, on Inca dove, 447.on ring-necked pheasant, 313, 319.on ruddy quail-dove, 453.Behr, Herman, on passenger pigeon,393, 394.Belding, Lyman, on plumed quail, 50, 51.Bendire, C. E., on Arizona scaled quail,53, 55, 56.on Attwater's prairie chicken, 263.on band-tailed pigeon, 355, 356.on blue-headed quail-dove, 457.on Canada ruffed grouse, 167, 168.on Canada spruce grouse, 132.on chachalaca, 348.on chestnut-bellied scaled quail, 59.on Columbian sharp-tailed grouse,288, 289, 290.on dusky grouse, 91, 92.on eastern bobwhite, 14, 28.on eastern mourning dove, 404.on eastern turkey, 331.on Florida bobwhite, 33.on Florida turkey, 340.on Franklin's grouse, 136, 137.on Gambel's quail, 75, 77, 78, 81.on gray ruffed grouse, 172.on heath hen, 273.on Hudsonian spruce grouse, 122,123.on Inca dove, 446.on Key West quail-dove, 450, 451.on lesser prairie chicken, 282.on masked bobwhite, 38, 39.on Mearns's quail, 85, S9, 90.on Merriam's turkey, 323, 324.on Mexican ground dove, 441, 443.on mountain quail, 41.on northern sharp-tailed grouse,285, 286.on Oregon ruffed grouse, 175.on plumed quail, 44, 46.on Richardson's grouse, 100.
Bendire, C. E., on rock ptarmigan, 203.on ruffed grouse, 148.on sage hen, 303, 304, 305, 306,309.on sooty grouse, 104, 107.on southern white-tailed ptarmi-gan, 235, 239, 240.on Texas bobwhite, 35, 36.on valley quail, 64, 70.on western white-winged dove,428, 434.on white-fronted dove, 423.Benson, H. C, on masked bobwhite, 39.Bent, A. C, 122.on Hudsonian spruce grouse, 124,127.Berner, Glenn, x.on greater prairie chicken, 257,260.Biological Survey, Bureau of, x, 12.Bishop, L. B., on Alaska spruce grouse,129.Bishop, S. C, on passenger pigeon,388.Bishop, S. C, and Wright, A. H., onpassenger pigeon, 380.Bishop, W. L., on Canada sprucegrouse, 132, 133."Blue grouse," 103.Blue-headed quail-dove, 456.Bobwhite, eastern, 9.Florida, 32.masked, 36.Texas, 34.Bonaparte, C. L., 300.Bonasa umbellus sabini, 166, 174.umbellus thayeri, 166, 177.umbellus togata, 166.umbellus umbelloides, 166, 171.umbellus umbellus, 141, 166.umbellus yukonensis, 166, 177.Bond, Frank, on sage hen, 300.Booth, E. J., x.Bowers, E. O., on passenger pigeon,393.Bowles, J. H., on California quail, 60.on mountain quail, 41.on Oregon ruffed grouse, 176.on sooty grouse, 106, 110.Brace, L. J. K., on Key West quail-dove, 452.Bradbury, W. C, on southern white-tailed ptarmigan, 236.Brandt, H. W, on Alaska sprucegrouse, 129.on Kellogg's ptarmigan, 228, 229.on northern sharp-tailed grouse,285.on willow ptarmigan, 179, 181, 188.Breninger, G. F., on Inca dove, 448.Brewer, T. M., on eastern bobwhite, 15.(See also under Baird.)Brewster, William, on Canada ruffedgrouse, 167, 169, 170.on Canada spruce grouse, 131, 132,133, 134, 135.on greater prairie chicken, 256.
INDEX 481Brewster, William, on heath hen, 264,266, 273.on masked bobwhite, 37.on passenger pigeon, 384, 385, 386,387.on ruffed grouse, 143, 149, 154.on San Lucas quail, 72.on Viosca's pigeon, 363, 364.Broley, C. L., on prairie sharp-tailedgrouse, 297.Brooks, Allan, xi.on blue grouse, 103.on Fleming's grouse, 102.on Richardson's grouse, 98.on sage hen, 302.Brooks, W. S., on Kellogg's ptarmigan,230.Brown, D. E., on Oregon ruffed grouse,175.on sage hen, 303.Brown, Herbert, on Gambel's quail, 81,82.on Inca dove, 446.on masked bobwhite, 36, 37, 39.on Mexican ground dove, 441.Bryant, H. C, on eastern mourningdove, 410, 411..(See also under Grinnell.)Bryant, Henry, on white-crowned pig-eon, 370.on Zenaida dove, 419.Bryant, W. E., on Inca dove, 445.Bryens, O. M., x.on greater prairie chicken, 244.Burnett, L. E., on sage hen, 302, 308.Butler, A. W., on eastern bobwhite, 25,26.Cahoou, J. C, on masked bobwhite, 39.California quail, 59.californica achrustera, Lophortvx, 61,72.californica, Lophortyx, 59, 61, 62.catalinensis, Lophortyx, 61, 70.plumbea, Lophortyx, 61, 71.vallicola, Lophortyx, 61, 62.californica, Lophortyx californica, 59,61, 62.CaUipepla squamata castanogastris, 58.squamata pallida, 51, 58.squamata squamata, 58.Cameron, E. S., on prairie sharp-tailedgrouse, 292.on sage hen, 302.Camp, R. D., 345.campestris, Pedioecetes phasianellus,287, 291.canace, Canachites canadensis, 128,129, 131.Canachites canadensis atratus, 129, 135.canadensis canace, 128, 129, 131.canadensis canadensis, 120, 128.canadensis osgoodi, 129.franklini, 136.Canada ruffed grouse, 166.Canada spruce grouse, 131.
canadensis atratus, Canachites, 129,135.canace, Canachites, 128, 129, 131.canadensis, Canachites, 120, 128.osgoodi, Canachites, 129.canadensis, Canachites canadensis, 120.128.Cantwell, G. G., on willow ptarmigan,180.carolinensis, Zenaidura macroura, 402,413.Carpenter, W. L., on Mearns's quail,88.on sage hen, 303.Carriger, H. W., on plumed quail, 45.Cassin, John, on Gambel's quail, 73,80, 81.castanogastris, Callipepla squamata,58.Catalina quail, 70.catalinensis, Lophortyx californica, 61,70.Centrocercus urophasianus, 300.Chachalaca, 345.chamberlaini, Lagopus rupestris, 212,221.Chamberlain's ptarmigan, 221.Chapman, F. M., on Alaska sprucegrouse, 130.on greater prairie chicken, 244.on Kenai white-tailed ptarmigan,234.on northern white-tailed ptarmi-gan, 232.on Sierra grouse, 114.Chestnut-bellied scaled quail, 58.Chicken, Attwater's prairie, 263.greater prairie, 242.lesser prairie, 2S0.Childs, J. L., on passenger pigeon, 397.Christy, B. H., and Sutton, G. M., oneastern turkey, 328.chrysia, Oreopeleia, 450.Clark, A. H., on Alaska ptarmigan, 201.on Chamberlain's ptarmigan, 221.on Evermann's ptarmigan, 231.on Nelson's ptarmigan, 216.on scaled pigeon, 378.Clark, John N., 14.Cleaves, H. H., on ruffed grouse, 156.Coffin, R. L., x.colchicus torquatus, Phasianus, 310.Colinus ridgwayi, 36.virginianus cubanensis, 32.virginianus floridanus, 32.virginianus texanus, 31, 32, 34.virginianus virginianus, 9, 31, 32.Columba fasciata fasciata, 353, 362.fasciata vioscae, 362, 363.flavirostris flavirostris, 365.leucocephala, 369.squamosa, 376.Columbian sharp-tailed grouse, 288.columbianus, Pedioecetes phasianellus,287, 288.Columbidae, 353.Columbiformes, 353.
482 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMColumbigallina passerina bahamensis,443.passerina pallescens, 439, 440.passerina passerina, 435, 439.Colvin, Walter, on lesser prairiechicken, 280, 281, 282, 283, 284.Cornpton, L. V., x.confinis, Oreortyx picta, 43, 51.Conover, H. B., on willow ptarmigan,178, 182.Cooke, W. W., on dusky grouse, 95.on greater prairie chicken, 244, 260.Cooperative Quail Investigation, 12.Cory, C. B., on white-crowned pigeon,376.Cottam, Clarence, on ring-neckedpheasant, 317.Coues, Elliott, on Columbian sharp-tailed grouse, 290.on dusky grouse, 91, 94.on Gambel's quail, 74, 78, 82.on sage hen, 306, 307.on southern white-tailed ptarmi-gan, 235, 238.Cracidae, 345.Craig, Wallace, on eastern mourningdove, 404, 406, 410.on passenger pigeon, 383, 384, 393,394.Cram, Eloise B., 29, 163.Criddle, Norman, on prairie sharp-tailed grouse, 298.Crone, J. V., x.cubanensis, Colinus virginianus, 32.cupido americanus, Tvmpanuchus, 242,263.attwateri, Tympanuchus, 263.cupido, Tympanuchus, 263, 264.cyanocephala, Starnoenas, 456.Cyrtonyx montezumae mearnsi, 84.Dalke, P. D., x.on ring-necked pheasant, 316, 317.Danforth, S. T., on Zenaida dove, 419,420.Davidson, J. L., on eastern mourningdove, 404.Dawson, W. L., on band-tailed pigeon,360.on Franklin's grouse, 138.on Mexican ground dove, 441.on Oregon ruffed grouse, 175, 176.on plumed quail, 44, 47, 49.on Sierra grouse, 115.on sooty grouse, 104, 105.on valley quail, 62, 63.on western white-winged dove, 429,434.Day, William, on heath hen, 277.Deane, Ruthven, on passenger pigeon,385, 386, 389, 399.Dendragapus, 103.fuliginosus fuliginosus, 103, 113,114.fuliginosus howardi, 114, 117.fuliginosus sierrae, 114.fuliginosus sitkensis, 113, 114, 119.obscurus flemingi, 96, 102.
Dendragapus obscurus obscurus, 91,95, 96.obscurus richardsoni, 95, 96.Denny, O. N., 310, 312.Devany, J. L., on Hudsonian sprucegrouse, 121.Dickey, D R., and van Rossem, A. J.,on Mount Pinos grouse, 118, 119.Dixon, Joseph, on Alexander's ptarmi-gan, 195.on Dixon's ptarmigan, 223, 224.on valley quail, 68.on willow ptarmigan, 180, 182,185, 186.(See also under Grinnell.)dixoni, Lagopus rupestris, 202, 213, 223.Dixon's ptarmigan, 223.Doolittle, A. E., on greater prairiechicken, 259.Dove, Bahama ground, 443.blue-headed quail, 456.eastern ground, 435.eastern mourning, 402.eastern white-winged, 425.Inca, 444.Key West quail, 450.Mexican ground, 440.ruddy quail, 453.western mourning, 416.western white-winged, 428.white-fronted, 422.Zenaida, 417.Doves, 353.DuBois, A. D., on eastern mourningdove, 405.Dusky grouse, 91.Dutcher, William, on eastern mourningdove, 408.Dwight, Jonathan, 384.on eastern bobwhite, 16.on rock ptarmigan, 207.on Texas bobwhite, 35.on willow ptarmigan, 183, 184.East, Ben, x.Eastern bobwhite, 9.Eastern ground dove, 435.Eastern mourning dove, 402.Eastern turkey, 326.Eastern white-winged dove, 425.Eaton, E. H., on passenger pigeon, 380.Ectopistes migratorius, 379.Edson, J. M., on sooty grouse, 111.Edwards, George, on ruffed grouse, 142.Elliot, D. G., 342.on Evermann's ptarmigan, 231,232.on Hudsonian spruce grouse, 126,127.on prairie sharp-tailed grouse, 291.on sage hen, 306.on southern white-tailed ptarmi-gan, 239.on Townsend's ptarmigan, 220.Erichsen, W. J., on eastern grounddove, 435, 436.European partridge, 1.
INDEX 483
everrnanni, Lagopus rupestris, 212, 230.Evermann'6 ptarmigan, 230.Falrchild, William, on greater prairiechicken, 259.Farley, F. L., x.on prairie sharp-tailed grouse, 291.Farmer, E. W., 345, 346.on chachalaca, 347, 348, 349, 350.fasciata, Columba fasciata, 353, 362.fasciata vioscae, Columba, 362, 363.Field, G. W., on greater prairie chicken,249.on heath hen, 273.on ring-necked pheasant, 321.Figgins, J. D., on Alaska sprucegrouse, 130.on Kenai white-tailed ptarmigan,234.Finlev, T. M., on red-billed pigeon,365.Finlev, W. L., on European partridge,
' 3.on Oregon ruffed grouse, 174, 175.on ring-necked pheasant, 311.on sage hen, 301, 306.Fisher, A. K., on valley quail, 68.flavirostris, Columba flavirostris, 365.Fleming, J. H., 102.on passenger pigeon, 379, 380.flemingi, Dendragapus obscurus, 96,102.Fleming's grouse, 102.Florida bobwhite, 32.Florida turkey, 340.floridanus, Colinus virginianus, 32.Forbush, E. H., on Canada sprucegrouse, 131, 134.on eastern bobwhite, 10, 22, 28, 30.on eastern mourning dove, 404.on eastern turkey, 327.on Hudsonian spruce grouse, 127.on ruffed grouse, 148, 150, 152,153, 154, 157, 158, 159.Fortiner, J. C, on Mexican grounddove, 442, 443.Fowler, F. H., on band-tailed pigeon,355.franklini, Canachites, 136.Franklin's grouse, 136.French, J. C., on passenger pigeon, 392.Fuertes, L. A., on Mearns's quail, 89.fuliginosus, Dendragapus fuliginosus,103, 113, 114.fuliginosus howardi, Dendragapus, 114,117.sierrae, Dendragapus, 114.sitkensis, Dendragapus, 113, 114,119.fulviventris angelica, Leptotila, 422.Gabrielson, I. N., on eastern mourningdove, 407.Gale, Denis, on dusky grouse, 91, 92.on southern white-tailed ptar-migan, 239.Galliformes, 1.
gallopavo intermedia, Meleagris, 326,342.merriami, Meleagris, 323, 326.osceola, Meleagris, 326, 340.silvestris, Meleagris, 326.Gambel, William, 73.gambeli, Lophortyx gambeli, 73.Lophortyx sanus, 83, 84.Gambel' s quail, 73.Gander, F. F., on eastern mourningdove, 405.on western mourning dove, 416.Gardner, L. L., on eastern mourningdove, 404.Giddings, H. J., on eastern bobwhite, 14.Gillespie, J. A., x.Gilman, M. F., on band-tailed pigeon,358.on Gambel 's quail, 77.on Inca dove, 444, 445, 446, 447448.on Mexican ground dove, 442.on western white-winged dove,428, 430, 434.Glen, J. J., on passenger pigeon, 393.Goss, N. S., on eastern mourning dove,403.on prairie sharp-tailed grouse, 297.Gosse, P. H., on eastern white-wingeddove, 426.on ruddy quail-dove, 454, 455.on white-crowned pigeon, 370,372, 374.on Zenaida dove, 418, 420.Gray ruffed grouse, 171.Grayson, A. J., on Inca dove, 445.on red-billed pigeon, 367, 368.on white-fronted dove, 423.graysoni, Ortyx (Colinus), 37.Greater prairie chicken, 242.Green, C. de B., on Richardson'sgrouse, 99.Grinnell, G. B., on sage hen, 306, 309.on southern white-tailed ptarmi-gan, 240.Grinnell, Joseph, 223, 224.on Alexander's ptarmigan, 195,197.on band-tailed pigeon, 356, 359.on Catalina quail, 70, 71.on Gambel's quail, 81.on Hudsonian spruce grouse, 126.on Kellogg 's ptarmigan, 227, 229.on San Quintin quail, 71, 72.on Valdez spruce grouse, 135, 136.on willow ptarmigan, 185.on Yukon ruffed grouse, 177, 178.Grinnell, Bryant, and Storer, on band-tailed pigeon, 354, 357, 359, 360,361.on Columbian sharp-tailed grouse,288, 289.on Gambel's quail, 80.on plumed quail, 43, 45, 49, 50.on sage hen, 304.on valley quail, 63, 66, 67, 68.
484 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMGrinnell, Dixon, and Linsdale, onSierra grouse, 117.Grinnell and Storer, on plumed quail,44.on Sierra grouse, 115, 117.Griscom, Ludlow, on rock ptarmigan,209.Gross, A. O., x, 242, 264.on prairie sharp-tailed grouse, 293,294, 295.on ruffed grouse, 161, 162.Ground dove, Bahama, 443.eastern, 435.Mexican, 440.Grouse, 91.Alaska spruce, 129."blue," 103.Canada ruffed, 166.Canada spruce, 131.Columbian sharp-tailed, 288.dusky, 91.Fleming's, 102.Franklin's, 136.gray ruffed, 171.Hudsonian spruce, 120.Mount Pinos, 117.northern sharp-tailed, 285.Nova Scotia ruffed, 177.Oregon ruffed, 174.prairie sharp-tailed, 291.Richardson's, 96.ruffed, 141.Sierra, 114.Sitka, 119.sooty, 103.Valdez spruce, 135.Yukon ruffed, 177.Gundlach, Jean, on blue-headed quail-dove, 457.Gurney, J. H., on greater prairiechicken, 256.Hadley, Philip, on passenger pigeon,381.Hadzor, R. T., on ruffed grouse, 156.Hagerup, A. T., on Reinhardt's ptarmi-gan, 215.Hall, F., on greater prairie chicken,262.Handley, C. O., on eastern bobwhite,18, 19.Hantzsch, Bernhard, on rock ptarmi-gan, 205, 210, 211.Hardy, Manly, on Canada ruffedgrouse, 168.on Hudsonian spruce grouse, 122.Haskin, L. L., on plumed quail, 49.on sooty grouse, 105, 113.Hayes, W. G., on passenger pigeon,393.Heath hen, 264.Henderson, A. D., on Hudsonian sprucegrouse, 122.Henshaw, H. W., on Mearns's quai,85, 87, 88.on Merriam's turkey, 323, 324.on plumed quail, 48.on valley quail, 69. '
Herman, F. J., x.Hersey, F. Seymour, 106.on Allen's ptarmigan, 192, 193,194.on rock ptarmigan, 204, 209.on willow ptarmigan, 180.Higginson, Francis, on passenger pi-geon^ 389.Hill, Grace A., on willow ptarmigan,185.Hodge, C. F., on passenger pigeon, 381.on ruffed grouse, 143, 144.Holden, C. H., jr., on sage hen, 307.Holland, H. M., on valley quail, 63.Holleman, Ridley. (See under Quillin.)Hollister, Ned, on ruddy quail-dove,453.(See also under Kumlien.)Holzner, F. X., on valley quail, 64.Horefall, R. B., x.on eastern bobwhite, 22.Howard, O. W., 118.on Merriam's turkey, 324.howardi, Dendragapus fuliginosus, 114,117.Howe, C. D., on ruffed grouse, 155.Howell, A. B., on Sierra grouse, 115.Hoyle, C. E., on heath hen, 276, 277.Hudson, W. H., on ring-necked pheas-ant, 311.Hudsonian spruce grouse, 120.Huev, L. M., on band-tailed pigeon,358.on valley quail, 67.Hungarian partridge. (See Europeanpartridge.)Hunter, John, on eastern turkey, 337.Huntington, D. W., on sage hen, 300,307, 308.on valley quail, 69.Inca dove, 444.inca, Scardafella inca, 444.intermedia, Meleagris gallopavo, 326,342.Irving, F. N., x.on white-crowned pigeon, 375.Jewett, H. J., on passenger pigeon, 392.Johnson, O. B., on band-tailed pigeon,355.Johnson, R. A., on sootv grouse, 104,111.Jones, Lynds, on eastern bobwhite, 21.Jones, MyrnaF.,29.Josselyn, John, on eastern turkey, 327.Jourdain, F. C. R., 3.on European partridge, 4, 6.on ring-necked pheasant, 312.Judd, S. D., on Arizona scaled quail, 55.on eastern bobwhite, 18, 19, 24.on eastern turkey, 334.on Gambel's quail, 79.on greater prairie chicken, 257,258.on plumed quail, 48.on prairie sharp-tailed grouse, 295.on ruffed grouse, 152, 153, 154.
INDEX 485Judd, S. D., on sage hen, 305.on sooty grouse, 109.on southern white-tailed ptarmi-gan, 238.Kalm, Pehr, on passenger pigeon, 388,389, 395.Keen, Miss A. M., x.Kellogg, Louise, on mountain quail, 42.kelloggae, Lagopus rupestris, 202, 213,227.Kellogg's ptarmigan, 227.Kenai white-tailed ptarmigan, 234.Keniston, Allan, on heath hen, 276.Kennard, F. H., on chachalaca, 347,350.Kentwood, on heath hen, 266.Key West quail-dove, 450.Keyes, Charles R., on plumed quail, 45.Kitchin, E. A., x.on band-tailed pigeon, 353, 354,361.Knight, O. W., on Canada ruffedgrouse, 169.Knobbe, W. H., on Oregon ruffedgrouse, 175.Koch, August, on passenger pigeon, 394.Kumlien, Ludwig, and Hollister, Ned,on passenger pigeon, 399.Lacey, Howard, on Rio Grande turkey,343.lagopus alascensis, Lagopus, i90, 200.albus, Lagopus, 178, 190.alexandrae, Lagopus, 191, 194.alleni, Lagopus, 190, 191.ungavus, Lagopus, 190, 197.Lagopus lagopus alascensis, 190, 200.lagopus albus, 178, 190.lagopus alexandrae, 191, 194.lagopus alleni, 190, 191.lagopus ungavus, 190, 197.leucurus altipetens, 233, 234.leucurus leucurus, 232, 233.leucurus peninsularis, 233, 234.leucurus rainierensis, 233, 240.mutus, 202.rupestris atkhensis, 212, 218.rupestris chamberlaini, 212, 221.rupestris dixoni, 202, 213, 223.rupestris evermanni, 212, 230.rupestris kelloggae, 202, 213, 227.rupestris nelsoni, 212, 215.rupestris reinhardi, 212, 213.rupestris rupestris, 202, 212.rupestris sanfordi, 212, 225.rupestris townsendi, 212, 220.rupestris welchi, 203.LaHontan, Baron de, on ruffed grouse,143.Laing, H. M., on Chamberlain'sptarmigan, 222.on Evermann's ptarmigan, 231, 232.on prairie sharp-tailed grouse, 292,297, 299.on Turner's ptarmigan, 220.Lamb, C. C, on Viosca's pigeon, 363,364.
Law, J. E. on valley quail, 62.Lawrence, A. G., on European par-tridge, 2.Lawrence, G. N., on greater prairiechicken, 255.on red-billed pigeon, 368.on Texas bobwhite, 34.on white-fronted dove, 423, 424'.Leffingwell, D. J., on European par-tridge, 2.on ring-necked pheasant, 312, 314,315, 317, 318, 320.Leptotila fulviventris angelica, 422.Lesser prairie chicken, 280.leucocephala, Columba, 369.leucurus altipetens, Lagopus, 233, 234.leucurus, Lagopus, 232, 233.peninsularis, Lagopus, 233, 234.rainierensis, Lagopus, 233, 240.leucurus, Lagopus leucurus, 232, 233.Lewis, E. J., on heath hen, 278.Lewis, Evan, on southern white-tailed ptarmigan, 236.Lewis, H. F., on Ungava ptarmigan.198, 200.Lewis and Clark, on sage hen, 300.Lincoln, F. C, x.on greater prairie chicken, 256, 260.Lingley, James, on Hudsonian sprucegrouse, 122.Linsdale, J. M. (See under Grinnell.)Littlejohn, Chase, on Alexander's ptar-migan, 197.Lloyd, William, on Arizona scaled quail,56.on eastern mourning dove, 411.on Mearns's quail, 85.on Merriam's turkey, 323.on Mexican ground dove, 443:on Texas bobwhite, 36.Lophortyx californica achrustera, 61, 72.californica californica, 59, 61, 62.californica catalinensis, 61, 70.californica plumbea, 61, 71.californica vallicola, 61, 62.gambeli gambeli, 73.gambeli sanus, 83, 84iLord, J. K., on Oregon ruffed grouse,174.Lowell, A. C, on valley quail; 70.MacFarlane, R. R., on Kellogg'sptarmigan, 228.on northern sharp-tailed grouse-285.on rock ptarmigan, 206.on willow ptarmigan, 180.Macgillivray, William, on Europeanpartridge, 6.MacMillan, D. B., on Reinhardt'sptarmigan, 213* 215.on rock ptarmigan, 206.Macoun, John, on northern sharp-tailedgrouse, 286.macroura carolinensis, Zenaidura, 402,413.marginella, Zenaidura, 413, 416.74564?32- -32
486 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMMailliard, Joseph, on Gambel's quail,81.Manniche, A. L. V., on Reinhardt'sptarmigan, 213, 214.on rock ptarmigan, 208.marginella, Zenaidura macroura, 413,416.Masked bohwhite, 36.Mavnard, C. J., on Bahama grounddove, 443, 444.on eastern ground dove, 436.on Key West quail-dove, 451.on passenger pigeon, 393, 394.on white-crowned pigeon, 371,373, 374, 376.on Zenaida dove, 420.McAfee, W. L., on ring-necked pheas-ant, 310, 317, 321.McCabe, Mrs. Eleanor B., 136.McCabe, T. T., x.on Franklin's grouse, 136, 141.on gray ruffed grouse, 172.McCall, G. A., 73.McClintock, Norman, x.McGregor, R. C., on Nelson's ptarmi-gan, 217.Mcllhenny, E. A., x.on eastern turkey, 336.Mearns, E. A., on Olathe quail, 84.on passenger pigeon, 392.on ruffed grouse, 151.mearnsi, Cvrtonvx montezumae, 84.Melopelia asiatica, 427, 428.Mearns's quail, 84.Meleagrididae, 323.Meleagris gallopavo intermedia, 326,342.gallopavo merriami, 323, 326.gallopavo osceola, 326, 340.gallopavo silvestris, 326.Melopelia asiatica asiatica, 425, 427.asiatica mearnsi, 427, 428.Merriam, C. H., on Allen's ptarmigan,191, 193.on eastern mourning dove, 409.on Merriam's turkey, 324.on ring-necked pheasant, 319.merriami, Meleagris gallopavo, 323, 326.Merriam's turkey, 323.Merrill, J. C., on chachalaca, 351.on red-billed pigeon, 365.on white-fronted dove, 423.Mershon, W. B., on passenger pigeon,384.Mexican ground dove, 440.migratorius, Ectopistes, 379.Millais, J. G., on ring-necked pheas-ant, 312, 313, 320.Minot, H. D., on eastern bobwhite, 26.Monson, G. W., x.on greater prairie chicken, 261.montana, Oreopeleia, 453.montezumae mearnsi, Cyrtonyx, 84.Morse, G. W., x.Morss, C. B., on ruffed grouse, 159.Morton, Thomas, on eastern turkey,327.on heath hen, 266.
Moseley, E. L., on eastern bobwhite, 19.Mount Pinos grouse, 117.Mountain quail, 40.Mourning dove, eastern, 402.western, 416.Murie, O. J., on rock ptarmigan, 209,210, 211.on Ungava ptarmigan, 198, 199.Murphy, R. C., on Gambel's quail, 80.Murraj\ J. J., x.mutus, Lagopus, 202.Nelson, E. W., on Arizona scaled quail,55.on Kellogg's ptarmigan, 230.on Mearns's quail, 90.on Merriam's turkey, 323.on red-billed pigeon, 367.on willow ptarmigan, 187, 189.on Yukon ruffed grouse, 178.nelsoni, Lagopus rupestris, 212, 215.Nelson's ptarmigan, 215.Newberry, Doctor, on sage hen, 306.Newton, Alfred, on ring-necked pheas-ant, 312.Nice, Mrs. M. M., on eastern bobwhite,19.on eastern mourning dove, 405, 406.Nicholson, D. J., on eastern grounddove, 436, 437, 438.on Florida bobwhite, 33.on Florida turkey, 342.Noble, G. K., on rock ptarmigan, 203,209.Norris-Elye, L. T. S., x.Northern sharp-tailed grouse, 285.Northern white-tailed ptarmigan, 232.Nova Scotia ruffed grouse, 177.Nuttall, Thomas, on heath hen, 266.Oberholser, H. C., on mountain quail,40, 41.obscurus, Dendragapus obscurus, 91,95, 96.obscurus flemingi, Dendragapus, 96,102.obscurus, Dendragapus, 91, 95, 96.richardsoni, Dendragapus, 95, 96.Ogilvie, F. M., on European partridge, 3.Olathe quail, 84.Oregon ruffed grouse, 174.Oreopeleia chrysia, 450.montana, 453.Oreortyx picta confinis, 43, 51.picta palmeri, 40, 41, 42, 43.picta picta, 40, 42, 43.picta plumifera, 40, 43, 51.Ortalis vetula vetula, 345.Ortyx (Colinus) graysoni, 37.picta, 40.osceola, Meleagris gallopavo, 326, 340.Osceola, Seminole chief, 340.Osgood, W. H., on Alaska spruce grouse,129, 131.on Nelson's ptarmigan, 215, 217.on southern white-tailed ptarmi-gan, 234.osgoodi, Canachites canadensis, 129.
INDEX 487pallescens, Columbigallina passerina,439, 440.pallida, Callipepla squamata, 51, 58.pallidicinctus, Tympanuchus, 280.Palmer, T. S., 4i.palmeri, Oreortyx picta, 40, 41, 42, 43.Pangburn, C. H., on Zenaida dove, 422.Partridge, European (Hungarian), 1.Passenger pigeon, 379.passerina bahamensis, Columbigallina,443.pallescens, Columbigallina, 439,440.passerina, Columbigallina, 435, 439.Payne, E. B., on eastern bobwhite, 14.Peabody, P. B., x.Peacock, W. F., on ring-necked pheas-ant, 316.Pearson, T. G., on chachalaca, 351.on eastern ground dove, 437, 438.Pedioecetes phasianellus campestris,287, 291.phasianellus columbianus, 287, 288.phasianellus phasianellus, 285, 287.Pemberton, J. R., on Mount Pinosgrouse, 118.Penard, T. E. (See under Bangs.)peninsularis, Lagopus leucurus, 233,234.Pennock, C. J., on eastern turkey, 328,331.Perdicidae, 1.Perdix perdix perdix, 1.Perrett, W. W., x.Peters, J. L., on San Lucas quail, 72.phasianellus campestris, Pedioecetes,287, 291.phasianellus columbianus, Pedioecetes,287, 288.phasianellus, Pedioecetes, 285, 287.phasianellus, Pedioecetes phasianellus,285, 287.Phasianidae, 310.Phasianus colchicus torquatus, 310.Pheasant, ring-necked, 310.Pheasants, 310.Phelps, F. M., on Florida turkey, 341.Phillips, J. C, on European partridge,1,2.on ring-necked pheasant, 310, 311,322.Pickwell, G. B., x.picta confinis, Oreortyx, 43, 51.palmeri, Oreortyx, 40, 41, 42, 43.picta, Oreortyx, 40, 42, 43.plumifera, Oreortyx, 40, 43, 51.picta, Oreortyx picta, 40, 42, 43.Ortyx, 40.Pigeon, band-tailed, 353.passenger, 379.red-billed, 365.scaled, 376.Viosca's, 363.white-crowned, 369.Pigeons, 353.plumbea, Lophortyx californica, 61, 71.Plumed quail, 43.
plumifera. Oreortyx picta, 40, 43, 51.Pokagon, Chief Simon, on passengerpigeon, 384.Poling, Otho C, 38.on Mearns's quail, 85.Prairie chicken, Attwater's, 263.greater, 242.lesser, 280.Prairie sharp-tailed grouse, 291.Preble, E. A., on Hudsonian sprucegrouse, 125.on willow ptarmigan, 185.Price, W. W. (.See under Barlow.)Proud, William, on valley quail, 64.Ptarmigan, Alaska, 200.Alexander's, 194.Allen's, 191.Chamberlain's, 221.Dixon's, 223.Evermann's, 230.Kellogg's, 227.Kenai white-tailed, 234.Nelson's, 215.northern white-tailed, 232.Rainier white-tailed, 240.Reinhardt's, 213.rock, 202.Sanford's, 225.southern white-tailed, 234_Townsend's, 220.Turner's, 218.Ungava, 197.willow, 178.Quail, Arizona scaled, 51.California, 89.Catalina, 70.chesfcnut-bellied scaled, 58..Gambel's 73.Mearns's, 84.mountain, 40.Olathe, 84.plumed, 43.San Lucas, 72.San Pedro, 51.San Quintin, 71.valley, 62.Quail-dove, blue-headed, 456.Kev West, 450.ruddy, 453.Quails, 1.Quillin, R. W., and Holleman, Ridley,on Inca dove, 446.Rabbi tts, Gower, x.Rainier white-tailed ptarmigan, 240.rainierensis, Lagopus leucurus, 233,.240.Ralph, W. L., on Florida turkey, 340,341.Rapp, F. W., on eastern bobwhite, 14.Rathbun, S. F., 2, 3, S, 105, 106.Red-billed pigeon, 365.reinhardi, Lagopus rupestris, 212, 213.Reinhardt's ptarmigan, 213.Revoil, B. H., on passenger pigeon, 391.Rice, F. J., on ring-necked pheasant,313.
488 BULLETIN 16 2, UNITED STATES NATIONAL MUSEUMRich, W. H., on Canada spruce grouse,135.on European partridge, 8.richardsoni, Dendragapus obscurus,95, 96.Richardson's grouse, 96.Richmond, C. W., xi.Ridgway, Robert, 37.on Key West quail-dove, 451.on western white-winged dove,428.(See also under Baird.)ridgwayi, Colinus, 36.Riley, J. H., on Alaska ptarmigan, 201.on Franklin's grouse, 139, 140.on gray ruffed grouse, 172.on Ungava ptarmigan, 197.Ring-necked pheasant, 310.Rio Grande turkey, 342.Roberts, T. S., on passenger pigeon,383.Rock ptarmigan, 202.Roney, H. B., on passenger pigeon,397, 398.Rosen, W. M., x.Rossignol, G. R., on white-crownedpigeon, 375.Rowan, William, on greater prairiechicken, 256.Ruddy quail-dove, 453.Ruffed grouse, 141.jupestris atkhensis, Lagopus, 212, 218.chamberlaini, Lagopus, 212, 220.dixoni, Lagopus, 202, 213, 223.evermanni, Lagopus, 212, 230.kelloggae, Lagopus, 202, 213, 227.nelsoni, Lagopus, 212, 215.reinhardi, Lagopus, 212, 213.rupestris, Lagopus, 202, 212.sanfordi, Lagopus, 212, 225.townsendi, Lagopus, 212, 220.welchi, Lagopus, 203.rupestris, Lagopus rupestris, 202, 212.
sabini, Bonasa umbellus, 166, 174.Sage hen, 300.Samuels, E. A., on ruffed grouse, 148.Sandys, Edwyn, on Canada sprucegrouse, 134.on dusky grouse, 194.on eastern bobwhite, 15, 21, 23,24, 30.on eastern turkey, 338.on Oregon ruffed grouse, 176.on plumed quail, 50.on prairie sharp-tailed grouse,291, 298.on ruffed grouse, 150.on sage hen, 307, 309.on southern white-tailed ptar-migan, 239, 240.on willow ptarmigan, 179, 187,188Sanford, L. C, 225.sanfordi, Lagopus rupestris, 212, 225.Sanford's ptarmigan, 225.San Lucas quail, 72.San Pedro quail, 51.
San Quintin quail, 71.sanus, Lophortyx gambeli, 83, 84.Sargent, Luman, 132.on Canada ruffed grouse, 170.Saunders, A. A., on eastern bobwhite,11, 22, 23.on eastern mourning dove, 409.on Franklin's grouse, 140, 141.on prairie sharp-tailed grouse, 297.on Richardson's grouse, 100, 101.Savary, W. B., x.Sawyer, E. J., on ruffed grouse, 145,150, 160.Scaled pigeon, 376.Scardafella inca inca, 444.Schmidt, E. W., x.Schutze, A. E., on Inca dove, 444.Sclater, W. L., on duskv grouse, 92.Scott, W. E. D., on Florida turkey, 340.on ruddy quail dove, 453.on white-crowned pigeon, 376.Sennett, G. B., on chachalaca, 348,350, 351.on chestnut-bellied scaled quail,58, 59.on red-billed pigeon, 365, 366, 367.on Rio Grande turkey, 342, 344.on white-fronted dove, 422, 423,424.Seton, E. T., on Canada ruffed grouse,168, 170.on gray ruffed grouse, 173.on prairie sharp-tailed grouse, 292,294, 297, 299.Sherman, Miss Althea R., on easternbobwhite, 21.on eastern mourning dove, 406,407.Sherwood, W. E., x.Shoemaker, F. H., x.Sierra grouse, 114.sierrae, Dendragapus fuliginosus, 114.silvestris, Meleagris gallopavo, 326.Sim, Robert J., 21.Simmons, G. F., 345.on Arizona scaled quail, 52, 53, 56.on Attwater's prairie chicken, 263,264.on eastern bobwhite, 13.on Inca dove, 444, 445, 446, 448.on Mexican ground dove, 443.on Rio Grande turkey, 343, 344.on Texas bobwhite, 34, 35.on western mourning dove, 416,417.Singley, J. A., on chachalaca, 348.Sitka grouse, 119.sitkensis, Dendragapus fuliginosus, 113,114, 119.Skinner, M. P., on eastern bobwhite,11.on eastern turkey, 328.on grav ruffed grouse, 171, 172,173.'on Richardson's grouse, 96, 99,100, 101.Slade, D. D., on eastern turkey, 327.
INDEX 489Smith, A. P., on Rio Grande turkey,343.on white-fronted dove, 423, 424.Smith, Everett, on Hudsonian sprucegrouse, 121.Smith, L. W., on ruffed grouse, 148.Snyder, J. O., on Franklin's grouse,140.Sooty grouse, 103.Southern white-tailed ptarmigan, 234.Spiker, Charles J., on European par-tridge, 3, 7, 8.Springer, Charles, on Merriam's turkey,324.Sprunt, Alexander, jr., on greaterprairie chicken, 245.Spurrell, J. A., on eastern mourningdove, 411.on greater prairie chicken, 256,260.squamata, Callipepla squamata, 58.squamata castanogastris, Callipepla,58.pallida, Callipepla, 51, 58.squamata, Callipepla, 58.squamosa, Columba, 376.Squires, W. A., x.Starnoenas cyanocephala, 456.Stejneger, Leonhard, on Allen's ptarmi-gan, 191.Stephens, Frank, 37.on Inca dove, 444.Stevens, O. A., on greater prairiechicken, 257, 260.Stevens, S. S., on passenger pigeon,384, 385.Stevenson, D. H. 226.Stockard, C. R.. on eastern bobwhite,13.on eastern mourning dove, 405,411.Stoddard, H. L., on eastern bobwhite,12, 13, 14, 15, 17, 18, 20, 23, 24,25, 28, 29.on greater prairie chicken, 245.Stone, Witmer, on Inca dove, 449.Storer, T. I. (see under Grinnell).Street, J. Fletcher, on Hudsonianspruce grouse, 126, 127.Sugden, J. W., x.Suthard, J. G., x.on eastern mourning dove, 404.on eastern turkey, 328.on passenger pigeon, 393.Sutton, G. M., on eastern turkey, 331.on passenger pigeon, 392.on ruffed grouse, 148.(See also under Christy.)Swainson and Richardson, on northernsharp-tailed grouse, 285, 286.Swarth, H. S., on Alaska ptarmigan,200. 201.on Alexander's ptarmigan, 195.on Allen's ptarmigan, 191.on band-tailed pigeon, 353.on blue grouse, 103.on gray ruffed grouse, 171, 172,173. "
Swarth, H.S., on Kellogg's ptai-migan,227.on Mearns's quail, 89.on Oregon ruffed grouse, 175.on rock ptarmigan, 202.on Sitka grouse, 120.on sooty grouse, 109, 112.on western white-winged dove,428.Swenk, M. H. and Jane B.on Inca dove, 449.Taverner, P. A., on band-tailed pigeon,358.on Fleming's grouse, 102.on Kellogg's ptarmigan, 227.on rock ptarmigan, 202.Taylor, W. P., on Rainier white-tailedptarmigan, 240, 241.Tegetmeier, W. B., on ring-neckedpheasant, 314, 318.TerriU, L. M., x.Tetraonidae, 91.texanus, Colinus virginianus, 31, 32,34.Texas bobwhite, 34.Thayer, J. E., 177.on masked bobwhite, 38.thayeri, Bonasa umbellus, 166, 177.Thompson, C. S., on eastern mourningdove, 411.F. J., on passenger pigeon, 386.Tindall, C. W., x.Todd, G. W., on Mearns's quail, 86.Todd, W. E. C, on white-crownedpigeon, 371, 375.togata, Bonasa umbellus, 166.Tomkins, I. R., x.torquatus, Phasianus colchicus, 310.Townsend, C. W., x, 120, 310, 379, 444jon ring-necked pheasant, 313, 315.on ruffed grouse, 158.townsendi, Lagopus rupestris. 212,220.Townsend's ptarmigan, 220.Trippe, T. M., on dusky grouse, 91, 94.on southern white-tailed ptarmi-gan, 238.Turkey, eastern, 326.Florida, 340.Merriam's, 323.Rio Grande, 342.Turkeys, 323.Turner, L. M., on Hudsonian 6prucegrouse, 122, 123, 125, 126, 127.on Nelson's ptarmigan, 216, 217.on rock ptarmigan, 203, 205, 206,208, 210.on Turner's ptarmigan, 219.on Ungava ptarmigan, 197, 198,199, 200.on willow ptarmigan, 184, 188.Turner's ptarmigan, 218.Tuttle, Miss F. May, x.Twitchell, A. H., on Alaska sprucegrouse, 130.Tyler, J. G., on valley quail, 63.Tyler, W. M., x, 402.
490 BULLETIN 162, UNITED STATES NATIONAL MUSEUMTympanuchus cupido americanus, 242,263.cupido attwateri, 263.cupido cupido, 263, 264.pallidiciuctus, 280.Tyzzer, E. E., 321.
umbelloides, Bonasa umbellus, 166, 171.umbellus, Bonasa umbelloidee, 166,171.Bonasa umbellus, 141, 166.umbellus sabini, Bonasa, 166, 174.thayeri, Bonasa, 166, 177.togata, Bonasa, 166.umbellus, Bonasa, 141, 166.yukonensis, Bonasa, 166, 177.Ungava ptarmigan, 197.ungavus, Lagopus lagopus, 190, 197.urophasianus, Centrocercus, 300.Valdez spruce grouse, 135.Valley quail, 62.vallicola, Lophortyx californica, 61, 62.Van Dyke, T. S., on valley quail, 68.van Rossem, A. J., on Inca dove, 445,447.on red-billed pigeon, 366.on sooty grouse, 108.(See also under Dickey.)vetula, Ortalis vetula, 345.Vilaro, Juan, on Kev West quail dove,451.vioscae, Columba fasciata, 362, 363.virginianus, Colinus virginianus, 9,31, 32.virginianus cubanensis, Colinus, 32.floridanus, Colinus, 32.texanus, Colinus, 31, 32, 34.virginianus, Colinus, 9, 31, 32.Vreeland, F. K., on ruffed grouse, 144.Wales, J. H., x.on band-tailed pigeon, 359.Warburton, Stanton, jr., on willowptarmigan, 180, 188.Ware, M. C, 314.Warner, E. P., on ruffed grouse, 148.Wayne, A. T., on eastern mourningdove, 408.welchi, Lagopus rupestris, 203.Western mourning dove, 416.Western white-winged dove, 428.Wetmore, Alexander, 222.on ruddy quail-dove, 453, 455.on scaled pigeon, 377, 378.on western white-winged dove,429, 431, 432, 433.on white-crowned pigeon, 370, 373.on Zenaida dove, 419, 420.Wheaton, J. M., on eastern turkey, 335.Wheelock, Mrs. Irene G., on band-tailed pigeon, 355, 357.
Wheelock, Mrs. Irene G., on plumedquail, 46.on sage hen, 304.on Sierra grouse, 116, 117.on valley quail, 64.Whitaker, J. R., on Allen's ptarmigan,192, 193, 194.on rock ptarmigan, 203, 209.White-crowned pigeon, 369.White-fronted dove, 422.White-winged dove, eastern, 425.western, 428.Whitman, C. O., on passenger pigeon,385.Wicks, M. L., on valley quail, 63.Widmann, Otto, on passenger pigeon,399.Willard, Frank C, xi.on Arizona scaled quail, 52, 53, 56.on band-tailed pigeon, 356, 358,361.on Gambel's quail, 74, 76, 77.on Inca dove, 446, 448.on Mearns's quail, 86, 89.Willett, George, on Dixon's ptarmigan,223.Williams, J. J., on valley quail, 66.Williams, R. S., on Richardson'sgrouse, 100.on sage hen, 305.Willow ptarmigan, 178.Wilson, Alexander, on eastern mourn-ing dove, 403.on eastern turkey, 334.on passenger pigeon, 383, 384, 385,388, 390, 391, 396.on white-crowned pigeon, 372.Witherby, H. F., on European par-tridge, 5.on ring-necked pheasant, 315.Wood, N. A., x.Wood, William, on heath hen, 265.on passenger pigeon, 389.Worden, John, on greater prairiechicken, 261, 262.Wright, A. H., on eastern turkey, 327,337.(See also Bishop, S. C.)Wyman, G. H., on sage hen, 305.Wyman, L. E., on passenger pigeon,380.Wythe, Miss M. W., x.Yarrell, William, on European par-tridge, 5, 6.Yukon ruffed grouse, 177.yukonensis, Bonasa umbellus, 166, 177.Zenaida dove, 417.Zenaida zenaida zenaida, 417.Zenaidura macroura carolineusis, 402,413.macroura marginella, 413, 416.c





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