//SMITHSONIAN INSTITUTIONUNITED STATES NATIONAL MUSEUMBulletin 144
THE AMERICAN BATS OF THE GENERAMYOTIS AND PIZONYX
BYGERRIT S. MILLER, Jr.Curator, Dhisioti of Mammals, United States National MuseumANDGLOVER M. ALLENCurator of Mammals, Museum of Comparative ZoologyCambridge, Massachusetts
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UNITED STATESGOVERNMENT PRINTING OFFICEWASHINGTON1928

SMITHSONIAN INSTITUTIONUNITED STATES NATIONAL MUSEUMBulletin 144
THE AMERICAN BATS OF THE GENERAMYOTIS AND PIZONYX
BYGERRIT S. MILLER, Jr.Curator, Division of Mammals, United States National MuseumANDGLOVER M. ALLENCurator of Mammals, Museum of Comparative ZoologyCambridge, Massachusetts
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UNITED STATESGOVERNMENT PRINTING OFFICEWASHINGTON1928
ISSUED MAY 2 5 1928
ADVERTISEMENTThe scientific publications of the National Museum include twoseries, known, respectively, as Proceedings and Bulletin.The Proceedings, begun in 1878, is intended primarily as a mediumfor the publication of original papers, based on the collections ofthe National Museum, that set forth newly acquired facts in biology,anthropology, and geology, with descriptions of new forms and revi-sions of limited groups. Copies of each paper, in pamphlet form,are distributed as published to libraries and scientific organizationsand to specialists and others interested in the different subjects.The dates at which these separate papers are published are recordedin the table of contents of each of the volumes.The Bulletin, the first of which was issued in 1875, consists ofa series of separate publications comprising monographs of largezoological groups and other general systematic treatises (occasion-ally in several volumes), faunal works, reports of expeditions,catalogues of type-specimens, special collections, and other materialof similar nature. The majority of the volumes are octavo in size,but a quarto size has been adopted in a few instances in whichlarge plates were regarded as indispensable. In the Bulletin seriesappear volumes under the heading ContHhutions from the UnitedStates National Herbarium, in octavo form, published by theNational Museum since 1902, which contain papers relating to thebotanical collections of the Museum.The present work forms No. 144 of the Bulletin series.Alexander "VVetmore,Assistant Secretary, Smithsonian Institution.Washington, D. C, Afril 18, 1928.
PREFATORY NOTEThe revision of the American bats of the Genus Myotis wasoriginally begun by the senior author alone. A preliminary reviewwas made by him on the basis of the available material in the UnitedStates National Museum (including the collections of the Bureau ofBiological Survey, United States Department of Agriculture), theField Museum of Natural History, and the British Museum (NaturalHistory). On account of the pressure of other duties, however, hewas unable to bring the study to completion. The notes and mate-rial were then placed in the hands of the junior author who took upthe revision ab initio and prepared a complete draft of the generalaccount of the species. Later, by conference and correspondence,the more difficult aspects of the subject were jointly considered andall divergencies of opinion were thus reduced to a basis of mutualaccord. The monograph in its present form sets forth the finalresults of all this work. m

CONTENTS
PageIntroduction 1Material and acknowledgments 2Abbreviations and measurements 3Differential characters of the American species of Myotis 4List of the forty-six recognized forms of American Myotis, with theirtype localities 10Geographical relations of the American species of Myotis 11Nomenclature of the American species of Myotis 15The Genera Myotis and Pizonyx and their American species 30Genus Myotis 30Genus Pizonyx 33Key to the American species and subspecies of Myotis and Pizonyx__ 34Species of Myotis occurring in Temperate North America 38Myotis lucifugus 38Myotis lucifugus lucifugus 48Myotis lucifugus alascensis 47Myotis lucifugus carissima 50Myotis lucifugus phasma 53Myotis lucifugus fortidens 54Myotis yumanensis 61Myotis yumanensis yumanensis 65Myotis yumanensis sociabilis 68Myotis yumanensis saturatus 70Myotis yumanensis lutosus 72Myotis austroriparius 76Myotis grisescens 80Myotis velifer 86Myotis velifer velifer 89Myotis velifer Incautus 92Myotis velifer peninsularis 93Myotis occultus ^^-^.?^_ji^_' 97Myotis keenii 101Myotis keenii keenii 104Myotis keenii septentrionalis 105Myotis evotis 111Myotis evotis evotis 114Myotis evotis chrysonotus 116Myotis milleri 118Myotis thysanodes 122Myotis thysanodes thysanodes 126Myotis thysanodes aztecus 128Myotis sodalis 130Myotis volans 135Myotis volans volans '. 139Myotis volans longicrus 140Myotis volans interior 142Myotis volans amotus 145
VI CONTENTSThe Genera Myotis and Pizonyx and tlieir American species?Continued.Si^ecies of Myotis occurring in Temperate North America?Continued. PageMyotis califoruicus 148Myotis californicus californicus 151Myotis californicus cauriuus 155Myotis californicus pallidus 157Myotis californicus mexicanus 159Myotis subulatus 164Myotis subulatus subulatus 168Myotis subulatus melanorhinus 169Myotis subulatus leibii 171Species of Myotis occurring in Tropical and South America 175Myotis nigricans 175Myotis nigricans nigricans 177Myotis nigricans extremus 181Myotis nigricans nesopolus 182Myotis nigricans dominicensis 183Myotis chiloensis 189Myotis chiloensis chiloensis 190Myotis chiloensis dinellii 191Myotis chiloensis atacamensis 192Myotis chiloensis oxyotus 193Myotis chiloensis alter 194Myotis ruber 197Myotis albescens 200Myotis simus 205[Myotis pilosus] 208Species of Pizonyx 209Pizonyx vivesi 209Index 215ILLUSTKATIONSPlate (facing p. 7)[All figures natural size]FiGUBE 1. Pizonyx vivesi, Guaymas, Sonora, Mexico.2. Large and small cheekteeth in two South American species ofMyotis {a, M. chiloensis, b, M. nigricans).3. Large and small cheekteeth in two North American species ofMyotis (a, M. velifcr, b, M. lucifugus)
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4. Maximum size of skull in American Myotis (M. velifer incautus).5. Minimum size of skull in American Myotis {M. nigricans domini'censis, type)
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6. Robust general form and well developed sagittal crest {M. velifervelifer).7. Slender form and slightly developed sagittal crest {M. evotis).8. Rostrum large relatively to brain case ; region between caninesbroader than interorbital constriction (M. occultus; No. 137098 typeof species, No. 125787 type of M. "baileyi").9. Rostrum small relatively to brain case; region between canines nar-rower than interorbital consti'iction {M. albescens).
CONTENTS VnFigure 10. Brain case high {M. volans; from left to right the specimens are:type of species, type of M. " capitaneus " and type of M. volanslongicrus)
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11. Brain case low (M. subuhttus Say, not Harrison Allen and recentauthors).12. Myotis lucifugus for comparison with M. keerm, fig. 13.13. Myotis keenii {=M. suMlattis H. Allen and recent authors, not Say)for comparison with M. lucifugus, fig. 12. (No. 22922 type ofspecies, No. 38633 lectotype of M. keenii septentrionalis)
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14. Myotis californicus for comparison with M. subulatus (Say, not H.Allen and recent authors).15. Myotis subulatus (Say) for comparison with M. californicus, fig. 14and M. lucifugus, fig. 12 (No. 86444 type of " Vespertilid oiliola-brum," No. 18684 type of M. subulatus melanorhinus, No. 150275type of M. " winnemana " )
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Text figxirb (p. S)[All figures greatly enlarged, not to scale]FiGUBE 1. Upper molars of Myotis and Pizonyx: a^ m^ of Myotis lucifugus;b, m' of Myotis myotis; c, m" of Myotis lucifugus; d, tn' of Pizonyxvivesi; e, in' of Myotis thysanodes.MAPSMap 1. (p. 38). Distribution of Myotis lucifugus.1. M. lucifugus lucifugus.2. M. lucifugus alascensis.3. M. lucifugus carissvttia.4. M. lucifugus phasma.5. M. lucifugus fortidens.2. (p. 62). Distribution of Myotis yumanensia.1. M. yumanensis yumanensis.2. M. yumanensis sociabilis.3. M. yumanensis saturatus.4. M. yumanensis lutosus.3. (p. 81). Distribution of Myotis grisescens.4. (p. 87). Distribution of Myotis velifer.1. M. velifer velifer.2. M. velifer incautus.3. M. velifer peninsularis.5. (p. 98). Distribution of Myotis occultus.6. (p. 102). Distribution of Myotis keenii.1. M. keenii keenii.2. M. keenii septentrionalis.7. (p. 112). Distribution of Myotis evoiis.1. M. evotis evotis.2. M. evotis chrysonotus.8. (p. 122). Distribution of Myotis thysanodes.1. M. thysanodes thysanodes.2. M. thysanodes aztecus.9. (p. 131). Distribution of Myotis sodaMs.
VIII CONTENTSMap 10. (p. 136). Distribution of Myotis volans.1. M. volans volans.2. M. volans longicrus.3. M. volant interior.4. M. volans oifnotus.11. (p. 149). Distribution of Myotis caUformeus.1. M. californicus californicus.2. M. californicus caurinus.3. M. californicus pallidus.4. M. californicus niexicanus.12. (p. 165). Distribution of Myotis suiulatus.1. M. suhulatus suiulatus.2. M. subulatus melanorhinus.3. M. subulatus leibid.13. (p. 210). Distribution of Pizonyx vivesi.
THE AMERICAN BATS OF THE GENERA MYOTIS ANDPIZONYX
By Gerrit S. Miller, Jr.Curator, Division of Mammals, United States National MuseumandGlover M. AllenCurator of Mammals, Museum of Comparative Zoology, Cambridge,Massachusetts
INTRODUCTIONThe bats of the genus Myotis, though small and inconspicuousmammals, present many features of unusual interest from the pointof view of systematic zoology. At nearly every point in its exces-sively wide range the genus is represented by several species oftenpuzzlingly alike in superficial appearance though readily distinguish-able from each other when the true differential characters are oncerecognized. As these animals everywhere subsist, so far as known,on small, soft-bodied insects, and their habits appear to be subject tono conspicuous changes from species to species or from district todistrict, we are forced to regard the process of specific differentia-tion in the group as a whole as primarily dependent on some otherfactor than any influence which might be exercised directly by theenvironment. This is equally true of the species in most genera ofbats; but in no other genus do we find the process worked out in somuch variety of detail over a territory so nearly world-wide inextent.In striking contrast to Myotis the nearly related genus Pizonyx isnot at present known to have any geographical " range " in the truesense, as the few existing specimens have all been collected on islets inthe Gulf of California and on the neighboring mainland of Sonora.Its peculiarities may prove to have an adaptational significance asthe animal has become specialized in a manner which somewhat paral-lels the occasionally fish-eating Noctilio. The food habits are, how-ever, not yet known. 1
2 BULLETIN 144, UNITED STATES NATIONAL MUSEUMMATERIAL AND ACKNOWLEDGMENTSWhen the North American Vespertilionidae were reviewed byMiller, 30 years ago/ the available collections, though much betterthan any previously studied, were still far from suiRcient to elucidatemany of the questions as to distribution and relationship of themembers of the genus Myotk, 1,322 specimens of which were thenenumerated. These were referred to 16 forms. The material onwhich the present study of this genus is based is much more abundantand well prepared than has ever before been brought together.From North America we have been able to list no less than 4,504specimens, representing 34 forms. So far as this region is concernedwe probably have now a fairly adequate knowledge of practicallyall the species of Myotis, though much work remains to be done onthe subject of local races and the exact limits of ranges. Withregard to South America the subject is in a less satisfactory state;but it is sufficiently far advanced to make us believe that, for theneotropical species, we have at least a good general picture whichlater may be filled in with more detail. We have examined 970specimens, referable, in our present opinion, to 12 forms. Of the5,474 specimens of Myotis which have passed through our hands wemay say that they represent practically all of the American materialavailable in the larger museums of this continent and a very impor-tant series of South American specimens lent by the British Museum.Acknowledgments are gratefully made to the following personsfor the loan of specimens belonging to them or in their charge : Mr.H. E, Anthony, of the American Museum of Natural History, NewYork; Prof. C. D. Bunker, Kansas University Museum of NaturalHistory, Lawrence, Kans. ; Prof. Manton Copeland, Bowdoin College,Brunswick, Me.; Dr. Lee R. Dice, University of Michigan, AnnArbor, Mich. ; Mr. Donald R. Dickey, 514 Lester Avenue, Pasadena,Calif.; Dr. Joseph Grinnell of the Museum of Vertebrate Zoology,University of California, Berkeley, Calif.; Mr. Samuel Henshaw,director of the Museum of Comparative Zoology, Cambridge, Mass.
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Miss Mary E. McLellan, California Academy of Sciences, San Fran-cisco, Calif. ; Dr. Wilfred H. Osgood, of the Field Museum of NaturalHistory, Chicago, 111.; Dr. Witmer Stone and Mr. Wharton Huber,of the Academy of Natural Sciences of Philadelphia, Pa. ; and Mr.Oldfield Thomas, of the British Museum (Natural History), London,England. The collection of the Academy of Natural Sciences ofPhiladelphia contains the types of several of Harrison Allen'sspecies; that of the American Museum includes a number of typesof both North American and South American forms; while that of
'North Amer. Fauna, No. 13, Oct. 16, 1897.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 3the University of California is especially rich in series of west coastspecimens. Finally, particular mention should be made of the use-fulness of the South American material belonging to the BritishMuseum.For the preparation of the maps we are indebted to the courtesyof the Biological Survey, United States Department of Agriculture.ABBREVIATIONS AND MEASUREMENTSThe following abbreviations are used, in the lists of specimensexamined and the tables of measurements, to indicate the severalinstitutions of whose collections the material forms a part
:
A. M. N. H.?American Museum of Natural History, New York City.A. N. S. P.?Academy of Natural Seieuces of Philadelphia, Pa.B. M.?British Museum (Natural History), London, England.C. A.?California Academy of Sciences, San Francisco, Calif.F. M.?Field Museum of Natural History, Chicago, 111.K. U.?Kansas University Museum of Natural History, Lawrence, Kans.M. C. Z.?Museum of Comparative Zoology, Cambridge, Mass.U. C.?Museum of Vertebrate Zoology, University of California, Berkeley,Calif.U. M.?University of Michigan, Ann Arbor, Mich.U. S. N. M.?United States National Museum, Washington, D. C.When no collection is specifically indicated the specimens alludedto are in the National Museum.Cranial measurements have all been made by the authors. Thetables of external measurements have been prepared by Mr. H. H.Shamel of the National Museum. It is to be understood that, unlessthere is a specific statement to the contrary, every individual includedin the tables of external measurements has the joints of the fingersin adult condition and that every skull has the basal suture closed.Wear of teeth as an index to age is indicated as follows in the tablesof cranial measurements: 0=no wear visible; l^wear just visible butnot obvious; 2= wear obvious but not obscuring the structural char-acters of the upper molars; 3= wear so far progressed as to obscurethe structural characters of the upper molars. In measuring skullsthe incisors have been excluded from both " greatest length " and
" mandible." The " occipital depth " does not include the sagittalcrest or the auditory bullae. The teeth rows were measured fromfront surface of canine to hind surface of posterior molar. Lengthand breadth of individual teeth were taken with dissecting micro-scope and eyepiece micrometer. Quotation marks indicate that cer-tain external measurements are given as recorded by the collectorsof the specimens. The absence of measurements of the ear showsthat the external measurements of a specimen were taken from thedried skin.
4 BULLETIN 144, UNITED STATES NATIONAL MUSEUMDIFFERENTIAL CHARACTERS OF THE AMERICAN SPECIES OF MYOTISAs the American members of the genus Myotis show no excessivedevelopments of structure, the differential characters by which thespecies may be recognized are not always readily understood. Themore important of these characters will therefore be passed brieflyin review.Ear.?The ear is usually about long enough to reach the end ofthe muzzle when laid forward. In Myotis volans it is slightlyshortened, but in such species as M. keenii {= M. subulatus H. Allenand recent authors, not of Say), M. evotis^ M. thysanodes^ and M.chiloensis, it is elongated suflEiciently to extend rather noticeably be-yond the tip of the muzzle. The tragus is of characteristic slenderlancet shape, varying little in its proportionate height, which isnearly half that of the ear. While the length of the ear relativelyto that of the head is often a very important specific character, wehave found the actual form of less value in distinguishing betweennearly related members of the genus.Wing and membranes.?Usually the third to fifth metacarpals andthird to fifth fingers are slightly graduated; but in a few speciesthese metacarpals tend to be nearly equal, as are also the fourth andfifth digits. Otherwise we have not found that characters derivedfrom the bony elements of the wing are of much service in determin-ing species. This remark, of course, is not intended to allude to theactual lengths of forearms and of digits, as these measurements arevery significant.The point of attachment of the wing membrane to the lower limb,a feature of importance in the Old World members of the genus, isa character which does not contribute much to the determination ofAmerican species of Myotis. In all of these except the North Amer-ican M. grisescens and the South American M. mnus (also M.pilostvs, if this is actually an American bat; see pp. 25, 208) themembrane is inserted on the side of the foot near the base of thephalanges ; in M. grisescens and M. simus the point of insertion is atthe ankle. While the type of insertion thus remains surprisinglyconstant in the American species of. Myotis., a very peculiar conditionhas been developed in Pizonyx (see p. 33).Foot.?The length of the foot relatively to that of the tibia variesmuch both individually and from species to species. When a fewspecimens of one kind are measured the variations in ratio of footlength to tibia length will be found to group themselves about a meanwhich is an important specific character. In the American speciesthe means of this ratio range from about 40 to about 60.
AMEEICAN BATS OF THE GENERA MYOTIS AND PIZONYX 5Among the Old World members of the genus there are species inwhich the foot is more enlarged than in any of those known fromAmerica (ratio of foot to tibia rising as high as 64 and T6), the wingattachment is at a point a little above the ankle, and the fur israther dense and woolly in appearance. The subgeneric name Leu-conoe has been applied to these bats; and Thomas formally raisedthe group to generic rank in 1915.^ The occurrence of various inter-mediate conditions, however, seems to us to make generic recognitionundesirable. Though Thomas has referred Myotis Jucifugus cai^-sima to the Leucono'e group, the nearest approach to this type of batin America is probably Myotis grisescens of the southeastern UnitedStates, the American species with the highest ratio of foot to tibia,and one of the two in which the wing membrane comes off from thetarsus instead of from the side of the foot. A large-footed animalresembling Myotis inacrotarsus of the Philippine Islands has beendescribed under the name Vespertilio pilosus from a supposed SouthAmerican locality. No second specimen of this bat has been taken,and the species must for the present be regarded as doubtfullyAmerican. *The presence or absence of a keel on the outer edge of the calcar isa character of importance in the identification of species. This struc-ture is a widening of the long spur that helps to spread the marginof the interfemoral membrane. It is convex in outline, beginning torise at a distance from the heel equal to about the width of the foot.Fur and color.?The body is well furred, and the hairy coveringusually extends thinly on the under side of the wing to a line joiningthe knee and the distal half or third of the humerus. In Myotisvolans it reaches the elbow. In some species there is a fringe ofshort hairs on the free edge of the interfemoral membrane; thisfringe is slightly developed in M. evotis^ very conspicuously in M.thysmiodes. Usually the fur is soft and silky, but sometimes, as inthe South American M. simus and the North American M. austro-riparius, it assumes a decidedly velvety or woolly texture. The tipsof the longer hairs may be burnished and glistening (as in Myotislucifugus) or dull (as in M. califomicus) . When the hairs of theback are smoothly parted two well-contrasted color zones are ordi-narily exposed, a darker slaty area involving the bases of all thehairs and a lighter distal area formed by the tips of the shorter hairsand determining the general hue of the upper parts. Occasionally thecontrast is not very noticeable, and rarely (as in M. grisescens or informs which have undergone unusual general darkening) it may beessentially absent. Ordinarily the tips of the longer dorsal hairs donot form an obvious third color band ; hence the absence of a tricolor
?Journ. Bombay Nat. Hist. Soc, vol. 23, p. 607.
6 BULLETIN 144, UNITED STATES NATIONAL MUSEUMpattern is often a convenient character by which to distinguish aMyotis from a Pipistrellus in parts of America where the two generaoccur together. One North American species of Myotis, however{M. sodalis), has definitely tricolored dorsal fur, but the pattern isnot so noticeable as it is in the species of Pipistrellus with which theanimal is associated. In the pallid desert race of Myotis califomicusan obvious tricolor pattern is also met with.The actual color is never bright. It ranges in different speciesfrom a very dark, almost blackish brown {niffricans, cMloen-sis)through dull brown to a decided reddish brown {t^her, califomicus),or, in desert races, to a very pale buff. The underparts are usuallypaler than the back, sometimes rather noticeably so (albescens),but there is never any striking color pattern. The ears and mem-branes vary from dull brownish to whitish or to almost black, some-times (albescens, nigricans) nearly concolor with the back, some-times (evotis, subulatus Say, not of H. Allen) strongly contrastedwith it. Immature individuals are normally darker and duller thanadults. No sexual differences in color have been detected, but twocolor phases independent of sex?a darker and a lighter, or dullerand brighter?may not infrequently be observed. For example, inM. lucifugus an olive and a bronzy phase are present. In theSouth American M. ruber there is likewise the dark brown and therusty phase ; and rusty individuals of M. nigricans occasionally occur.In M. evotis as well as in M. suhulatus (Say) the black pigmenta-tion of the ears and membranes is a character usually retained evenunder desert conditions that bleach the color of the fur, while in thearid-country forms of M. lucifugus the ears are less intensely pig-mented and the interfemoral membrane becomes whitish, with oftena contrasting white border along the fr'ee edge of the wing.Color differentiation among the species of Myotis is not verygreat. In the six species known to occur in the eastern United Statesthe general brownish color is so nearly alike that it is often difficultto identify individual specimens by this character alone, althougha comparison of enough skins shows that each animal has its dis-tinctive hue. This general similarity has led to great confusion inthe identification of specimens.Skull.?In both form and size the skull presents variations whichare useful in distinguishing species. Plate 1 has been prepared withthe special object of giving a clear idea of these features. The greatdifference in size which separates Pizonyx from the largest knownAmerican Myotis is at once evident on comparison of Figure 1(Pizonyx) with Figure 4 (Myotis velifer incautus). Figures 4 and5 show the extremes of known variation in size of skull among theAmerican species of Myotis (M. velifer incautus and M. nigricansdominicensis) . Further inspection of the plate shows that the

U, S. NATIONAL MUSEUM BULLETIN 144 PLATE I
10hh
IZ.
15
l^
// 1 1 f #/^
W9 w^ ^P* /5 ^* w^ w^'
Skulls of American Bats of the Genera Myotis and Pizonyx. Natural SizeFor explanation of plate see page VI
AMEEICAN BATS OF THE GEKEEA MYOTIS AND PIZONYX 7general outline of the skull may be narrower {evofis, fig. 7, keenii^fig. 13) or broader {velifer^ fig. 6, occuUus, fig. 8) ; that the brain casemay be high {volans, fig. 10) or low {suhulatus, fig. 11) ; that thearea of the rostrum as compared with that of the brain case may belarger {occultus^ fig. 8) or smaller {albescens, fig. 9) ; that the breadthof the rostrum across the roots of the canines may be more {occultus)or less {albescens) than the interorbital constriction; and that thesagittal crest may be conspicuous {velifer, figs. 4, 6) or essentiallyabsent {Tucifugus^ fig. 12). Other important characters which canbe less successfully illustrated by photographs are : The greater or lessbreadth of the palate, including the tooth rows (a measurement besttaken at the point of contact between the outer borders of rnr andni^) as compared with the length of the upper tooth row (canine tothird molar, both inclusive), and the greater or less crown area ofthe upper molariform teeth as compared with the area of the bonypalate between the tooth rows {chilo'ensis and nigricans^ fig. 2;velifer and lucifugus, fig. 3).Teeth.?The teeth vary in their general size relatively to the sizeof the skull; but a character of this kind is impossible to describewith any degree of accuracy. The difference in this respect betweensmaller toothed animals like Myotis ludfugus and M. nigricans., andlarger toothed animals like M. velifer and M. chUoensis is, however,appreciable on direct comparison (pi. 1, figs. 2 and 3).The incisors and canines, both upper and lower, and the lowermolars vary slightly in form; but we have not found that theirpeculiarities give much aid in the discrimination of species. NoAmerican Myotis has yet been discovered in which the second tri-angle of nu is so much reduced as it is in the type of the genus, thepalearctic M. myotis.The development and relative position of the two minute premolarsboth above and below is variable individually as well as in differentspecies. Miller has published figures of the upper small premolars offour specimens of Myotis thysanodes,, a species which appears to beespecially subject to variation in this respect.^ Usually, there iseither a slight crowding, or else the posterior of the small teeth maybe drawn inward from the tooth row, particularly in the case of theupper jaw, a condition well developed in the peculiar South AmericanMyotis simus. The most progressive form, as regards the reductionof the tooth-formula, is Myotis occultus. In this animal there isa decided tendency toward the loss of the posterior of the smallpremolars in the upper jaw, while in the lower jaw the correspondingtooth is usually crowded inward away from the line. Occasionalindividuals of the other species or races may also lack one of theseminute teeth; in most cases it is the second and smaller of the two
?North Amer. Fauna, No. 13, p. 82, Oct. 16, 1897.
8 BULLETIN 144, UNITED STATES NATIONAL MUSEUMwhich disappears. An anomaly which has been twice observed (inM. thysanodes No. 52228, U. S. N. M., and M. lucifugus carissimaNo. 38029, U. S. N. M.) consists in the coalescence of the two smallteeth on one side of the upper jaw; and in one instance (see p. 53)we have found a third small tooth welded to the anterior margin ofthe large premolar.The upper molars present some important structural peculiaritieswhich should be carefully understood.In the first and second of these teeth the most complicated typeof cusp development is the one which is found fully displayed inMyotis lucifugus (fig. Ic). In this type the hypocone is a largeand obvious element of the crown plan, its base well indicated as aswelling distinct from thatwhich forms the base ofthe protocone, its pointsometimes but not alwaysstanding out from the highlongitudinal loph whichconnects the summit of theprotocone with that of theless elevated hypocone. Anoticeable metaloph extendsfrom the base of the meta-cone across the deep valleywhich lies between the meta-cone and hypocone. Thisridge passes up the innerside of the hypocone to thelongitudinal loph (as in the figure) or in some instances to theactual summit of the hypocone itself. On the anterior marginof the crown a conspicuous triangular protoconule lies in the spacebetween the protocone and paracone. This small cusp is connectedwith the base of the paracone by a short but very obvious ridgewhich may be spoken of as the paraloph. The opposite extreme,so far as the American members of the genus is concerned, is seenin Myotis thysanodes (fig. le). Here the inner side of the toothis somewhat narrowed, chiefly at the expense of the hypocone;the metaloph, protoconule, and paraloph have completely disap-peared, leaving a deep smooth-floored valley between the hypoconeand metacone and another between the protocone and paracone. Inthe type of the genus, the palearctic Myotis myotis^ this reductionprocess is carried so far that the hypocone is normally smaller than inM. thysanodes, and in some individuals it practically does not existas an element distinguishable from the base of the protocone. Apeculiar condition is sometimes seen in the genus Pizonyx (fig. Id),Here the hypocone and metaloph are essentially as in Myotis
Fig. 1. MOLAK TKETH OF MYOTIS AND PiZONYX : a,ms of Myotis lucifugus ; &, m^ of Myotismyotis ; c, m 2 of myotis lucifugus ; d, m =OF Pizonyx vivesi ; e, m= of Myotis thy-sanodes. All gkkatly bnlabged, not to scale
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 9lucifugus, but the protoconule is absent and the paraloph mayextend from the base of the paracone well up toward the summit ofthe protocone.In the third upper molar the process of reduction seems to beconcentrated on the posterior segment of the tooth. The anteriorportion in Myotis lucifugus (fig. la) remains essentially as in thefirst and second teeth. The metastyle is, however, completely sup-pressed, and the loph extending to it from the metacone is reducedto a mere trace running downward and directly backward along theposterior surface of the metacone. The metaloph has also disap-peared; but the hypocone is still visible as a minute remnant. Ex-treme reduction (fig. 16) takes the form of still greater antero-posterior shortening of the crown, together with the complete andfinal elimination of the metaconule and its loph as well as the hypo-cone. In this stage, however, the protoconule usually persists, evenwhen it has completely disappeared from ni^ and ?;r. This condi-tion is best shown by Myotis w,yotis, the species figured, but it isapproached in the American M. thysanodes and M. evotis.The crowns of the upper molars bear no cingulum on the outerside, but the lingual and hinder borders are margined by a cingulumwhich, beginning at about the level of the protoconule on the anteriorside, extends around the lingual border and out along the posteriormargin to the metastyle. This cingulum is subject to much varia-tion in development. Usually its continuity is nearly or quite brokenat the antero-lingual portion of the base of the protocone. Less oftenthere is a break at the postero-lingual bulge of the base of the hypo-cone. In rare instances the cingulum may be reduced, along theentire lingual border of the crown, to a series of irregularly devel-oped enamel nodules. Still more rarely it may bear an incipientcusp in the region of the hypocone. As is the case with the secondarycusps the cingulum shows a more reduced condition in the typespecies of the genus, Myotis myotis, than it does in any of the knownAmerican forms.Were the extreme types of cusp development isolated they mightwell be considered as furnishing characters of generic or subgenericimportance, but so many intermediate conditions occur that it seemsimpossible to attribute any such weight to these structures. Ap-parently the more complicated type is the one which is to be re-garded as representing the condition primitive for the genus Myotis.It occurs in all the American species except M. evotis and M. thy-sanodes. Among the Old World forms we have found it in speci-mens determined, with varying degrees of authenticity, as adversus,hocagei, capaocinii, ca7%7nonensis, dasycneme, daubentonii., hilde-gardcB, 'macrotarsus, and irvwrcMs. A more simplified structure re-58518?28 2
10 BULLETIN 144, UNITED STATES NATIONAL MUSEUMsembling in different degrees that which is seen in the Americanevotls and thysanodes occurs in at least six Old World species;hechsteinii^ emarginatus^ mystacinus^ nattereri^ 7nifopicttis, and #n-coJor. The extreme type of reduction we have seen in the myotisgroup only {blythii, cMnensis, 7nyotis, oxygriathus). This group,apparently, is not connected by intermediate stages of structure withany other members of the genus. It might, therefore, be separatedas a subgenus or genus Myotis in the restricted sense. The nameLeucono'e Boie would then be available for the great mass of speciesnow called Myotis. At present, however, we are not convinced ofthe desirability of following this course, as the Old World membersof the genus are still insufficiently known.LIST OF THE FORTY-SIX RECOGNIZED FORMS OF AMERICAN MYOTIS,WITH THEIR TYPE LOCALITIES
A. NORTH AMERICANMYOTIS LUCIFUGUS (p. 38) :M. lucifugus lueifKffus (p. 43). Probably Georgia.M. lucifugus alascenMs (p. 47). Sitka, Alaska.M. lucifugus carissima (p. 50). Yellowstone Park. Wyo.M. lucifugus phasma (p. 53). Snake Rivex-, Colo.M. lucifugus fortidens (p. 54). Teapa, Tabasco, Mexico.MYOTIS YUMANBNSIS (p. 61) :M. ifumanensis yumancnsis (p. 65). Fort Yuma, Calif.M. yumanensis sociaMUs (p. 68). Hamilton, Wash.M. yumanensis saturatus (p. 70). Old Fort Tejon, Calif.M. yumanensis lutosus (p. 72). Patzcuaro, Michoacan, Mexico.MYOTIS AUSTRORIPARIUS (p. 76). Tarpon Springs, Fla.MYOTIS GRISESCENS (p. 80). Nickajack Cave, Tenn.MYOTIS VELIFER (p. 86) :M. velifer velifer (p. 89). Guadalajara, Jalisco, Mexico.M. velifer incautus (p. 92). San Antonio, Tex.M. velifer peninsulari^ (p. 93). San Jos6 del Cabo, Lower California,Mexico.MYOTIS OCCULTUS (p. 97). Near Needles, Calif.MYOTIS KEENII (p. 101) :M. keenii keenU (p. 104). Massett, Queen Charlotte Islands, British Colum-bia, Canada.M. keenii septentrionalis (p. 105). Halifax, Nova Scotia.MYOTIS EVOTIS (p. Ill) :M. evotis evotis (p. 114). Near Colville, Wash.M. evotis chrysonottis (p. 116). Kinney Ranch, Sweetwater County, Wyo.MYOTIS MILLERI (p. 118). La Grulla, Lower California, Mexico.MYOTIS THYSANODES (p. 122) :M. thysanodes thysanodes (p. 126). Old Fort Tejon, Calif,M. thysanodes aztecus (p. 128). San Antonio, Oaxnca, Mexico.MYOTIS SODALIS (p. 130). Wyandotte Cave, Ind.
AMERICAN BATS OF THE GENEEA MYOTIS AND PIZONYX HMYOTIS VOLANS (p. 135) :M. volans volans (p. 139). Cape St. Lucas, Lower California, Mexico.M. volans longicrus (p. 140). Puget Sound, Wash.M. volans interior (p. 142). Twining, N. Mex.M. volans amotus (p. 145). Cofre de Perote, Vera Cruz, Mexico.MYOTIS CALIFORNICUS (p. 148) :M. californicus californicus (p. 151). California.M. californicus caurinus (p. 155). Massett, Queen Charlotte Ids., BritishColumbia, Canada.M. californicus pallidus (p. 157). Vallecito, San Diego County, Calif.M. califomious mexicanus (p. 159). " Les terres chaudes de la province deMexico."MYOTIS SUBULATUS (p. 164) :M. suhulatus subulatus (p. 168). La Junta, Colo.M. sxibulatus melanorliinus (p. 169). San Francisco Mountain, Ariz.M. subulatus leibii (p. 171). Erie County, Ohio.
B. SOUTH AMEKICANMYOTIS NIGRICANS (p. 175) :M. nigricans nigricans (p. 177). Fazenda de Aga, near Rio Iritiba, prov-ince of Espirito Santo, eastern Brazil.M. nigricans extremus (p. 181). Huehuetan, Chiapas, Mexico.M. nigricans nesopolus (p. 182). Near Willemstad, Curacao, Dutch WestIndies.M. nigricans dominiccnsis (p. 183). Dominica, British West Indies.MYOTIS CHILOENSIS (p. 189) :M. chiloensis ohilocn^is (p. 190). Islets on the eastern side of ChiloeIsland, southern Chile.M. chiloensis dinellii (p. 191). Tucuman, Catamarca Province, Argentina.M. chiloensis atacamensis (p. 192). Atacama, Chile.M. chiloensis oxyotus (p. 193). Mount Chimborazo, Ecuador, at 9,000-10,000 feet latitude.M. chiloensis alter (p. 194). Palmeiras, Parana, Brazil.MYOTIS RUBER (p. 197). Paraguay.MYOTIS ALBESCENS (p. 200). Paraguay.MYOTIS SIMUS (p. 205). Sarayacu, on the Ucayali River, Loreto, easternPeru.[MYOTIS PILOSUS (p. 208). ?Montevideo, Uruguay.]GEOGRAPHICAL RELATIONS OF THE AMERICAN SPECIES OF MYOTISThat the genus Myotis is most abundantly represented in the tem-perate parts of America rather than the tropics, is indicated by thefact that of the 19 species here recognized as definitely pertaining tothe New World, 14 {austroHpanits, californicus, evotis, grisescens,keenii, lucifugus, niilleH^ occultus^ sodaZis^ subulatits, thysanodes^velifer, volans, yumanensis) are exclusively North American, whileonly 5 {albescens, chiloensis, nigineans, ruber, svnius) are confined tothe neotropical region. It must be remembered, however, that ourpresent knowledge of the South American members of the genus isstill far from complete.
12 BULLETIN 144, UNITED STATES NATIONAL MUSEUMThe most widespread American species is Myotis lucifugiis, whoserange extends to tree limit in the north and to southern Mexico inthe south. It becomes modified into five geographic races under thechanging conditions of this geographical area, but its structuralcharacters remain very constant. The fact, therefore, is rather re-markable that a genus whose members may be capable of pushingtheir range so far should be absent from the West Indies except inthe southern Lesser Antilles (Grenada, Dominica), where the tropi-cal Myotis nigricans has gained a foothold, having undoubtedly comein from northern South America. A representative of the same speciesoccurs on the island of Curacao, Such a distribution, however, is inline with the pronounced northern affinities of the genus; it alsoserves, perhaps, to emphasize the long isolation of the Antilles fromthe continent. We have too little accurate knowledge of the dis-tribution of Myotis in tropical America to say whether or not theknown species are mainly found at the higher levels and in coolforest, but such may not improbably be the case.In the northern continent a general correspondence may be tracedbetween some of the American species and those of the Eastern Hemi-sphere. There appears to be little doubt that the long-eared Ameri-can Myotis keenii, M. evotis, and M. thysanodes are not distantlyrelated to the Old World M. nattereri (with its Japanese race horrv-hinus) and M. emarginatus. The Palearctic Myotis daubentonii hascharacters allying it to M. lucifugus, while a Chinese species, M.frater, seems to be the Old World counterpart of M. volans. Simi-larly, the Chinese M. tnoupinensis may be regarded as the OldWorld representative of M. californicus^ which it closely resemblesin its small size, bodily proportions, delicate feet, keeled calcar, andlong fur. The range of the American species volant and californiousnow reaches as far north as the coast of southern Alaska, and that ofevotis and yumanensis nearly, if not quite, as far. Presumably, thenorthward extent of their representatives in the Old World will befound to be limited by corresponding isotherms on the Asiatic side,and this fact, if it could eventually be substantiated, would be in ameasure indicative of the higher temperature that must once haveprevailed to the northward at the time when the ranges of these nowslightly differentiated species were continuous from Asia to Americaby way of a northern land connection providing favorable climaticconditions. For it is not unreasonable to suppose that by some suchimmigration these American species were derived from Asia andthat by subsequent evolution specific differences have arisen. On theother hand, there is no obvious explanation of the fact that the verylarge species, such as the European and Asiatic Myotis nvyotis andM. chinensis have no representatives in the New World; but in thisconnection it should be recalled that these bats are not known to
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 13
occur in either the British Islands or Japan, a fact which may indi-cate that they are more sedentary in habit than some of their smallerrelatives.That the American Myotis stock came originally from the north-ern part of the Old World is made to appear probable by the follow-ing circumstances : The greater variety and higher degree of differ-entiation among the Palearctic members of the genus, an indicationof greater age for this portion of the group; the occurrence of thegreatest number of American species in western North America, theregion nearest the supposed Asiatic source of supply; and finallythe diminution in number of species as we go away from this westernregion, either toward the east or toward the south.Three North American species of Myotis have geographicalranges which extend across the continent from ocean to ocean. Theyare M. lucifugus with a north-south distribution from tree limit tosouthern Mexico, M. keenU ( = the subulatus of H. Allen and recentauthors) with a range from the southeastern United States to BritishColumbia and southern Alaska, and M. suhidatics (of Say) prac-tically confined to the United States. All of the others are restrictedeither to the East or to the West.North America east of the Mississippi is inhabited by three speciespeculiar to the region, all of them rather imperfectly known as todetails of distribution, though structurally they are well differenti-ated from each other and from the three wide ranging types whichoccur with them. One of these, 31. grisescens, appears to be peculiarto the limestone cave region of the central and southern United States,another, also apparently a cave bat, M. sodalis, has been found in theSouthern States and in Vermont, while the third, M. austroripariiis,Las been observed in only two regions, one in Indiana and the otheron the west coast of Florida.In contrast to the eastern United States, western North America,the portion of the New World nearest the supposed Asiatic center ofdispersal, has no less than eight species not yet known to occur else-where. One of these (milleri) has thus far been collected at asingle locality only, in Lower California; the others have wide andwell-defined ranges in the region west of the plains. Four of these{califomicus, evotis, volcms, and yunianensis) extend from the north-west coast region to southern Mexico, two {thysanodes and velifer)do not range so far to the north, while the remaining species, occultus,still imperfectly known, appears to be restricted to the southwesternUnited States and the adjoining parts of Mexico.Of the five species definitely known to inhabit South America,three, chiloensis, nigricans, and i^her, are generally distributed andnot strongly differentiated from each other. Their nearest relativesto the north appear to be M. lucifugus and M. yvjmanensis. Prob-
14 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
ably as wide ranging is the more highly specialized M. aR>escen8^The fifth species, M. shnus^ one of the most strikingly characterizedAmerican members of the genus, is known from two localities about250 miles apart on the lowlands east of the Andes.A feature which is noticeable among the American members ofthis group is the usually inverse ratio between distribution and dif-ferentiation. The four nearly related and slightly specialized speciesMyotis lucifugus^ M. yumanensis, M. chiloen^sis, and M. nigHccmscover practically the entire American area inhabited by these bats;at the opposite extreme of structural modification we find Pizonyx,an animal whose area of dispersal is restricted, apparently, to thecoast and islands of the Gulf of California. The range of Myotisvelifer includes most of Mexico and the adjoining part of the UnitedStates ; that of the more highly specialized M. ocauZtus is, accordingto present knowledge at least, decidedly less extensive. The seem-ingly narrow ranges of such aberrant species as Myotis grisescensand M. simvus are in noticeable contrast with the large areas occupiedby M. keerm and M. subulatus. On the other hand the range ofthe highly specialized M. volans is practically coincident with thatof the rather primitive M. yumanensis.Where species spread over areas of strikingly different climaticconditions, they usually show a corresponding color variation. Thisseems to be chiefly a response to the degree of atmospheric saturation?or dryness. Thus along the humid coast of the Northwest, fromsouthern Alaska to California, no less than five different Myotis arerepresented by darkened (" saturate ") local forms. In the semiaridregions farther eastward, the color becomes less intense over a widearea of country from the interior of California northeastward, whileunder the desert conditions of the Southwest and parts of the interiorof the United States, the same species become extremely pale. Stillfarther south, in southern Mexico, they again darken in color ; whilein the eastern United States, they may be of a different color still,neither saturate nor pallid.In the following table are listed the geographical races of six NorthAmerican species of Myotis grouped according to the general climatic-areas inhabited.
Species
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 15The fact that the wide-ranging species break up so readily intosubspecies corresponding to areas of different climatic conditionsperhaps indicates that the bats of this genus are very little migratoryand that such migration as there may be probably consists in nothingmore than a local withdrawal to certain not-distant caves for hiber-nation, or in some cases in an invasion of near-by alpine heightsduring the warm months at the close of the breeding period.NOMENCLATURE OF THE AMERICAN SPECIES OF MYOTISNo less than 102 trivial names have been given to American batsof the genus Myotis. Only 39 of these are here regarded as the validdesignations of recognizable forms. We are proposing 7 new namesfor species and races not hitherto discriminated. Tiie total numberis thus raised to 109 names for 46 forms, an average of nearly2.4 names to a form. This average, among the genera of Americanmammals, which have been recently revised, is high.* It may betaken as some index to the difficulty and confusion which haveattended the study of these bats.The following are the names which have been applied to speciesand subspecies of American Myotis:Aenobarbus (VespertUio) Temminck, Monogr. de Mamm., vol. 2, p.247, pi. 69, fig. 4, 1840. The specimen on which this name was basedis presumably still in the Leiden Museum; it came from an unknownlocality in South America. Though the tooth formula is not given,Temminck's figure of the head shows the rjharacteristic long narrowtragus of Myotis, while the proportions of tail to total length andthe description of the color leave little doubt that the name is asynonym of M. albescens.Affinis (VespertUio) H. Allen, Monogr. Bats North Amer., Smith-sonian Misc. Coll., No. 165, p. 53, June, 1864. Examination of thetype specimen (No. 5342, U. S. Nat. Mus.) shows it to be referableto the eastern race of Myotis lucifugus.Agilis {VespertUio) H. Allen, Proc. Acad. Nat. Sci. Philadelphia,1866, p. 282. This name, based on specimens from Mirador, VeraCruz, was applied to the Mexican representative of M. califomicus.It is antedated by VespertUio meaeicamis Saussure, 1860.Alascensis {Myotis livcifugus) Miller, North Amer. Fauna, No. 13,p. 63, October 16, 1897. The only name applied to the dark sub-species of M. lucifugus occuring on the humid northwest coast.Albescens {VespertUio) E, Geoffroy, Ann. Mus. d'Hist. Nat. Paris,vol. 8, p. 204, 1806. This name, based on the " Chauve-souris
* In some of the other groups it is as follows : Ochotona, 1.04 ; Thomomys, 1.1 ; Lepori-dw, 1.3; Oryzomys, 1.3; Spilogale, 1.3; Peromyscus, 1.5; Vespertilionine bats other thanMyotis, 1.7 ; Talpidm, 1.8.
16 BULLETIN 144, UNITED STATES NATIONAL MUSEUMdouzieme " of Azara's Essais sur 1' Histoire Naturelle du Paraguay,1801, was the first to be applied to the species which now bears thename Myotis albescens.Albicinctus {Myotis) G. M. Allen, Journ. Mamm., vol. 1, p. 2,November 28, 1919. This name proves to be a synonym of Myotislucifugus cmissimn\ the pale subspecies of M. lucifugus from thesemiarid parts of the western United States.Altifrons [Myotis) Hollister, Smithsonian Misc. Coll., vol. 56, No.26, p. 3, December 5, 1911. The type specimen is from Henry House,Alberta. It is not distinguishable from other examples of the darkM. volans longici^us.Altipetens {Myotis yunumensis) H. W. Grinnell, Univ. CaliforniaPubl. Zool., vol. 17, p. 9, August 23, 1916. At first described as arace of Myotis yuniaiiensis, the animal was later correctly recognizedby its describer as a pale form of M. lucifugus (Univ. CaliforniaPubl. Zool., vol. 17, p. 263, January 31, 1918). The name is antedatedby Myotis carissima Thomas, 1904.Amotus {Myotis longicrus) Miller, Proc. Biol. Soc. Washington,vol. 27, p. 212, October 31, 1914. The tenable name for the Mexicansubspecies of Myotis volans ; type from Cofre de Perote, Vera Cruz.Arsinoe {Vespertilio) Temminck, Monogr. de Mamm., vol. 2, p. 247,1840. The specimen which formed the basis for this name came fromSurinam, and was said by Dobson (1878) to be in the LeidenMuseum. The six cheek teeth in each jaw indicate that it was aMyotis, while the description as a whole applies to the Vespertilioalbescens of GeofFroy, 1805,Atacamensis {Vespertilio) Lataste, Actes Soc. Sci. Chile, Santiago,vol. 1, p. 79, 1892. This is the tenable name for the small gray-haired race of Myotis chiloensis occurring in the arid western coastregion of South America. Lataste briefly pointed out some of themore important characters. He attributes the name to Philippi, whohad so labeled the specimens in the Santiago Museum. Philippi'sdescription and plate did not appear until 1896, however, hence theauthorship of the name must date from Lataste on the basis of hisshort diagnosis.Austroriparius {Vespertilio lucifugus) Rhoads, Proc. Acad. Nat. Sci.Philadelphia, p. 157, May 22, 1897. Misled by the immaturity of thetype specimen, Miller (1897) placed this name in the synonymy ofMyotis lucifugus lucifugus. Additional material shows that theanimal is a distinct species, known at present from the vicinity ofTarpon Springs, Fla. (the type locality), and from Mitchell,Indiana. A specimen in the British Museum, collected by Drum-mond, is labeled " North America." It may have come from theinterior of Canada.
AMEEICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 17Baileyi (Myotis) HoUister, Proc. Biol. Soc. Washington, vol. 22,p. 44, March 10, 1909. The type specimen (from Ruidoso, N. Mex.),is a large individual of M. occultus HoUister in the olive phase ofpelage.Bayleyi {Myotis) Lydekker, Zool. Eecord, 1909, Mammalia, p. 59,1913. This is a misprint for haileyi HoUister.Bondas {Myotis) J. A. Allen, BuU. Amer. Mus. Nat. Hist., vol. 33,p. 384, July 9, 1914, This name was given to the form of M. nigri-cans from Bonda, Santa Marta, Colombia. The animal seems, how-ever, to be identical with M. nigricans nigricans.Brasiliensis {Vespertilio) Spix, Sim. et Vespert. Brasil. Sp. Nov.,p. 63, pi. 36, fig. 8, 1823. The figure and description indicate thatthis bat is identical with Myotis nigricans, also from Brazil. Thisis the first name applied to the species, but it is preoccupied by V.brasiliensis Desmarest, 1822, for a different animal, an Eptesicus. Itwas replaced by Vespertilio spixii Fischer in 1829, but Wied hadmeanwhile (1826) described the species as V. nigricans.Brevirostris {Vespertilio) Maximilian zu Wied, Verzeich. beobacht.Saugeth. N. Amer., Arch. f. Naturgesch., 1861, vol. 1, p. 195. Thereis almost nothing in the description of this bat that is diagnostic,except the yellowish tint of the belly. This and the general account,as well as the locality (Freiburg, Pa.) leave little doubt that Wied'sanimal was Myotis lucifugus. The type was said to be lost.Californicus {Vespertilio) Audubon and Bachman, Journ. Acad.Nat. Sci. Philadelphia, ser. 1, vol. 8, p. 285, 1842. Although nolocality beyond " California " is given, the original descriptionapplies to the small reddish-brown Myotis of the southern part ofthat State. The small feet are specially mentioned and the " lightyellowish-brown" color above, slightly darker beneath. While thismight refer as well to a race of Myotis subulatus (Say, not recentauthors). Miller (1897) has definitely assigned the name to thesmaller species.Capitaneus {Myotis) Nelson and Goldman, Proc. Biol. Soc. Wash-ington, vol. 22, p. 28, March 10, 1909. This name was given to thetypical race of M. volans before the latter name had been correctlyallocated through a reexamination of the type (from Cape St. Lucas,Lower California). Since the type of Myotis capitaneus is alsofrom Lower California (near Comondu) the name becomes a syno-nym of M. volans.Capucinus {Vespertilio) Philippi, Arch. f. Naturgesch., 1866,vol. 1, p. 114. Trouessart (Catal. Mamm. viv. foss., suppl., p. 94,1904) includes this as a species of Myotis, but Lataste (Actes Soc.Sci. Chile, Santiago, vol. 1 (1891), p. 90, 1892), who examinedPhilippi's type in Santiago, had shown that it is a Histiotus identicalwith H. Tnagellamicus.
18 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCarissima {Myotis) Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol.13, p. 383, May, 1904. This is the first name applied to the palesubspecies of M. lucifugus inhabiting the semi-arid portions of thewestern United States (type from Yellowstone Park, Wyo.). Theword is intended as an equivalent to the surname of the collector ofthe type. Darling. (Italian cosa carissima.)Carolii (VespertiUo) Temminck, Monogr. de Mamm., vol. 2, p. 237,1840. The description of this bat was based on specimens fromPhiladelphia and New York. It unquestionably applies to Myotislucifugus LeConte, 1831.Caucensis {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol.33, p. 386, July 9, 1914. The description is based on a large specimenof M. nig7'icans from Cauca, Colombia.Caurinus {Myotis californicubs) Miller, North Amer. Fauna, No. 13,p. 72, October 16, 1897. This is the first name applied to the darkrace of M. cdtifoimicus inhabiting the humid northwest coast dis-trict (type from Massett, Queen Charlotte Islands, British Columbia).Chiloensis {VespertiUo) Waterhouse, Zool. Voyage H. M, S. Beagle,pt. 2, Mamm., p. 5, pi. 3, 1838. This is the first name given to arather widely ranging South American species related to Myoticnigricans. The type was from Chiloe Island.Chiriquensis {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist.,vol. 20, p. 77, February 29, 1904. As first pointed out by Goldman(Smithsonian Misc. Coll., vol. 69, No. 5, p. 213, April 24, 1920), theshortness of the forearm, which was supposed to distinguish thisspecies from M. nigricans, is an accidental character due to mutila-tion in both wings of the type skin. The specimen (from Chiriqui,Panama) is identical with the Central American bat which we regardas M. nigricans nigricans.Chrysonotus {VespertiUo) J. A. Allen, Bull. Amer. Mus. Nat. Hist.,vol. 8, p. 240, November 21, 1896. This name is available in a sub-specific sense for the pale race of Myotis evotis which occupies thegreater part of the animal's range. The type was from SweetwaterCounty, Wyo.Ciliolabrum {VespertiUo) Merriam, Proc. Biol. Soc. Washington,vol. 4, p. 1, December 7, 1886. Though long considered a form ofMyotis caUfomicus, the bat to which this name was applied provesto be the typical race of a distinct species, the animal first describedas VespertiUo suhvJatus by Say in 1823 but not clearly recognizeduntil 1918, when its true characters were pointed out by Mrs. Grinnellunder the name oHnomus (Univ. Calif. Publ. Zool., vol. 17, p. 290,January 31, 1918).Cincinnus {VespertiUo) H. Allen, Bull. U. S. Nat. Mus., No. 43(1893), p. 97, footnote, March 14, 1895. A misspeUing of co^winnus,
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 19here used inadvertently no doubt, but appearing in the index as well;as in the text.Cinnamomeus (Vespertilio) Wagner, Schreber's Saugthiere, Suppl.,vol. 5, p. 755, 1855. Wagner believed that Rengger was correct inconsidering Azara's " Chauve-souris cannelle " a Noctilio. Since
-^zara's description was the basis of Vespertilio rube?' of E. Geoffroy,Wagner proposed cijinamoTiieus as a substitute for the name imber asused by D'Orbigny and Gervais for a red Myotis from Corrientes,Argentina. Azara's animal, however, is now known to have beenthis same red Myotis, so that the two names, 'ruber and cinnmnwnev/S,were actually based on one bat.Concinnus {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadel-phia, 18G6, p. 280. This name, applied to specimens from San Sal-vador, proves to have been based on the large form of Myotisnigncans inhabiting Central America and not, as orginally supposedby Miller (Proc. Biol. Soc. Washington, vol. 13, p. 154, June 13,1900), on the Mexican race.Crassus {Vespertilio) F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat.Paris, vol. 1, p. 18, 1832. This name is practically impossible toallocate with certainty. It was based on a specimen sent from NewYork, and is probably a synonym of Myotis lumfiigas.Dinellii {Myotis) Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 10,p. 493, December, 1902. This is the only name applied to the bright-colored race of Myotis chilohisis occurring in the drier parts ofnorthwestern Argentina.Domesticus {Vespertilio) Green, in Doughty's Cabinet of Nat. Hist.,vol. 2, p. 290, 1832. The description as well as the house-hauntinghabit of this bat leave little doubt that it is the same animal aaIjeConte's V. lucifugus described the previous year.Dominicensis {Myotis) Miller, Proc. Biol. Soc. Washington, vol. 15,p. 243, December 16, 1902. This is the form of M. nigricans foundin the island of Dominica, Lesser Antilles.Durangse {Myotis californicus) J. A. Allen, Bull. Amer. Mus. Nat.Hist., vol. 19, p. 612, November 12, 1903. An examination of thetype and other specimens from Durango on which the descriptionwas based,, shows that all are typical M. ymianensis ywmxinensis.Durangose {Myotis califomious) Trouessart, Cat. Mamm. viv. foss.,suppl., p. 93 1904. A modification of durangcB.Esmeraldse {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol.33, p. 385, July 9, 1914, We are unable to distinguish the form ofMyotis nigincans on which this name was based from the typical:subspecies.Evotis {VespertiUo) H. Allen, Monogr. Bats North Amer., Smith-sonian Misc. Coll., No. 165, p. 48, June, 1864. This is the first namegiven to the long-eared Myotis of the western United States.
20 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExiguus {Vespertilio) H, Allen, Proc. Acad. Nat. Sci. Philadelphia^1866, p. 281. The description of this bat leaves little doubt that it isidentical with M. nigricans (described as Vespertilio ooncinnus onthe previous page). The type specimen was from Aspinwall, nowColon, Panama.Exilis {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadelphia^1866, p. 283. This is a synonym of Myotis californicus. The typecame from Cape St. Lucas, Lower California.Ferrugineus iyespertilio) Temminck, Monogr. de Mamm., vol. 2, p.239, 1840. Trouessart in his Catal. Mamm. viv. foss., suppl., 1904,,p. 90, includes this as a valid species of Myotis^ following Dobson.The original description, based on specimens in alcohol from Suri-nam, states that there were-* teeth posterior to the canine, whichindicates probably an Eptesicus. Moreover, the name VespertUioferinigimeus is preoccupied by the V. fernjigineus of C. L. Brehm,1827, which is a synonym apparently of some European species ofNyctalus. It therefore requires no further consideration here.Gayi {VespertUio) Lataste, Actes Soc. Sci. Chile, Santiago, vol. 1(1891), p. 81, 1892. The specimen figured and described by Gervaisin Gay's Historia de Chile, ZooL, vol. 1, p. 42, 1847, as VespertUiochiloensis was slightly unusual in having the posterior minute pre-molar of the upper jaw crowded into the angle behind the frontcorner of the large third premolar. For this reason Lataste believedit to be a distinct species and named it VespertUio gayi. Gervaismentions that it is often taken in the houses at Valdivia, Chile.The name is a synonym of Myotis chiloensis chiloensis.Griisescens {Myotis) Howell, Proc. Biol, Soc. Washington, vol. 32,p. 46, March 10, 1909. This is the name of a large gray Myotis ofthe southeastern United States, distinguished among Americanspecies by the fact that the wing arises from the tarsus rather thanfrom the base of the toes.Gryphus {VespertUio) F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat.Paris, vol. 1, p. 15, 1832. Although Cuvier's description offers verylittle that is diagnostic, it is likely that this name is a synonymof Myotis lucifugus. That the animal was a member of the genusMyotis is rendered probable by the lancet-shaped tragus and the twominute premolars in each jaAv (in addition of course to the largerthird premolar) ; the size and color apply fairly well also. Thespecimen came from near New York.Henshawii {VespertUio nitidus) H. Allen, Bull. U. S. Nat. Mus.,No. 43 (1893), p. 103, March 14, 1894. This name was proposed ina tentative way for two specimens collected by H. W. Henshaw nearWingate, N. Mex., in 1872 (entered together under the number12450 (U. S. Nat. Mus.). These, though referred to "F. nitidus cilio-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 21)Idbruni''^ {^^Myotis subulatus subulatus), were recognized as insome way different, and the name hensJmwii was proposed in casefurther study confirmed this conclusion. It seems not unlikely thatthey were examples of the southwestern race of Myotis subulatus^for which Merriam's name nielanor^hinus, 1890, has priority. Thecotypes can not now be found in the National Museum collection,though they were indicated as present by Miller in his Lists of 1911and 1923, probably on insufficient evidence.Hypothrix (Vespertilio) D'Orbigny and Gervais, Voy. dans I'Amer-Merid., vol. 4, pt. 2, Mammiferes, pp. 14 (footnote), 16, 1847. Thespecies on which this name was founded is said by its authors to becommon in the Mojos country of eastern Bolivia. It is apparentlyidentical with Myotis nigricans nigricans.Hypothryx {Vespertilio) D'Orbigny and Gervais, Voy. dans. FAmer.Merid., vol. 4, pt. 2, Mammiferes, p. 16, 1847. Apparently a mis-print for hypothrix^i the spelling that occurs on p. 14 (footnote).Incautus {Vespertilio) J. A. Allen, Bull. Amer. Mus. Nat. Hist.,vol. 8, p. 239, November 21, 1896. This is the name of the slightlydifferentiated pale northern race of Myotis velifer. The type lo-cality is San Antonio, Tex.Interior {Myotis longicrus) Miller, Proc. Biol. Soc. Washington,vol. 27, p. 211, October 31, 1914. This now stands as M. volans inte-rior, for the pale race of the long-legged bat of the arid West. Thetype locality is Twining, Taos County, N. Mex.Isidori {Vespertilio) D'Orbigny and Gervais, Voy. dans I'Amer.Merid., vol. 4, pt. 2, Mammiferes, p. 16, 1847. The description of thisbat, based on a specimen from Corrientes, Argentina, leaves no doubtthat it is the same as M. albescens Geoffroy, 1805.Jaliscensis {Myotis califamicus) Menegaux, Bull. Mus. d'Hist,Nat. Paris, vol. 7, p. 321, 1901. The cotypes of this species, whichcame from near Lake Zacoalco, Jalisco, Mexico, are in the ParisMuseum, where Miller examined them in 1904 and found them tobe examples of Myotis velifer velifer.Keaysi {Myotis ruber) J. A. Allen, Bull. Amer. Mus. Nat. Hist.,vol. 33, p. 383, July 9, 1914. This is a name based on the red phaseof Myotis nigricans nigricatns.Keenii {Vespertilio subulatus) Merriam, Amer. Nat., vol. 29, p. 860,September, 1895. This is the first name unquestionably based on themoderately long-eared species which has been generally but incor-rectly known as Myotis subulatus, that is, the subulatus of HarrisonAllen, 1864, but not of Say, 1823. The type locality, Massett, QueenCharlotte Islands, British Columbia, lies within the area occupied bythe dark northwest coast form of the species.Kinnamon {Vespertilio) Gervais, Exped. Amer. du Sud du Castel-nau, Zool., Mammiferes, p. 84, pi. 15, fig. 1, 1855. The description of
22 BULLETIN 144, UNITED STATES NATIONAL MUSEUMthe teeth and color, as well as the measurements (forearm 40 mm.)leave no doubt that this animal is identical with M. tniher (Geoffrey),.1805. The type was from Capella Nova, Brazil.lanceolatus (Vespertilio) Maximilian zu Wied, Reise in das InnereNord-America, vol. 1, p. 364, footnote, 1839. The name wasbased on two specimens of a small bat from Bethlehem, Pa. Itwas originally proposed as a substitute for Vespertilio subulahiSyshould the bat which Maximilian so designated prove to be differ-ent from Say's animal. Maximilian's subulatus, however, appearsto be the same as M. lucifugiis LeConte, 1836.Leibii {Vespei^tilio) Audubon and Bachman, Journ. Acad. Nat.Sci. Philadelphia, ser. 1, vol. 8, p. 284, 1842. These authors carefullydescribed a small bat from Erie County, then Michigan, now Ohio,pointing out the obvious characters of small size, small foot, long tail,,and contrasting black ears and wings that distinguish the small black-faced bat of the eastern United States. The name has been considereda synonym of Myotis lucifucjus^ though the original authors showedtheir familiarity with that animal by redescribing it under the nameF. virgimmnus in the same paper. In recent years the discovery ofadditional specimens of leihii proves that it is a valid form, the east-ern race of Myotis subulatus Say, 1823 (not the subulatus of HarrisonAllen, 1864, and recent writers). The name, therefore, in the formMyotis subulatus leibii, replaces Myotis winnemaiia Nelson, 1913.Leucogaster {Vespertilio) Maximilian zu Wied, Beitr. z. Naturg.Brasil, vol. 2, p. 271, 1826. Wied's description as well as his excellentcolored figure in the Abbildungen zur Naturg. Brasil, pt. 13, 1829,leave no doubt that this animal is the same as the V. albescens of E.Geoffroy, 1805. His specimen was from the Moucouri River, Brazil.Longicrus {Vespertilio) True, Science, vol. 8, p. 588, December 24,1886. This is the valid name for the dark subspecies of Myotis volansoccurring in the humid northwest coast district. The type came fromPuget Sound, Washington.Lucifugus {Vespertilio) LeConte, McMurtrie's Cuvier, AnimalKingdom, vol. 1, p. 431, 1831. Though the original description con-tains little that is useful for allocating this name, the later and moredetailed account (Proc. Acad. Nat. Sci. Philadelphia, 1855, p. 436)shows that LeConte had in mind the common small brown Myotisof eastern North America, to which the name is currently applied(see Miller, North Amer. Fauna, No. 13, p. 29, October 16, 1897).The type locality is assumed to be Georgia, at or near LeConte'shome in the vicinity of Riceboro, Liberty County.Macropus {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadel-phia, 1866, p. 288. This name is a synonym of Myotis yimianensisof the same author, published two years previously. The type was
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 23
collected at Fort Mohave ("Majaor" in original description; seeLyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull.U. S. Nat. Mus., No. 62, p. 271, January 28, 1909) on the ColoradoRiver, western Arizona. The name is preoccupied by V. viacroj)tisGould, 1854, for an Australian bat.Maripensis {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol.33, p. 385, April 20, 1914. Though it has been regarded as a fullspecies, this bat is apparently nothing else than true Myotis nigricans.Some of the specimens are less dark than usual, but we are unable todefine the form. The type locality is Maripa, Rio Caura, Venezuela.Melanorhinus (Vespertilio) Merriam, North Amer. Fauna, No. 3,p. 46, September 11, 1890. This appears to be the valid name for theform of Myotis subulatus Say (not of H. Allen and subsequentwriters) of the southwestern United States and Mexican border.The type came from San Francisco Mountain, Arizona, and has beengenerally regarded as a specimen of M. colifornieus.Mexicanus {Vespertilio) Saussure, Rev. et Mag. de Zool., ser. 2,vol. 12, p. 282, 1860. This is the first name applied to the largerichly colored Mexican race of Myotis californicus. The type wassaid to have come from " les terres chaudes de la province de Mexico."Micronyx {Myotis) Nelson and Goldman, Proc. Biol. Soc. Wash-ington, vol. 22, p. 28, March 10, 1909. A name based on the LowerCalifornian form of Myotis evotis, apparently not distinguishablefrom M. evotis chrysonotiis.Milleri {Myotis) Elliot, Publ. Field Columbian Mus., zool. ser.,vol. 3, p. 172, April, 1903. This is the only name applied to apeculiar local form, related to Myotis evotis but apparently a distinctspecies, known only from a few specimens taken in the San PedroMartir Mountains, Lower California.Mundus {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadelphia,1866, p. 280. An examination of the specimen on which the descrip-tion of this species was based, shows it to be an immature example ofMyotis albescens Geoffroy. It came from Maracaibo, Venezuela.Nesopolus {Myotis) Miller, Proc. Biol. Soc. Washington, vol. 13,p. 123, April 6, 1900. This is the only name given to the form ofMyotis nigricans inhabiting the island of Curagao.Nesopotus {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol.33, p. 384, April 20, 1914. A misprint for nesopolus.Nigricans {Vespertilio) Maximilian zu Wied, Beitr. z. Naturg.Brasil, vol. 2, p. 266, 1826. This is the first valid name applied tothe common small blackish Myotis of tropical South America. Thetype was from the Rio Iritiba, Espirito Santo, Brazil.Nitens {Vespertilio) Wagner, Schreber's Saugthiere, suppl., vol. 5,p. 810, pi. 51, fig. 4, 1855. Referred by Trouessart to Myotis., and
24 BULLETIN 144, UNITED STATES NATIONAL MUSEUMpossibly a synonym of M. nigricans; "Wagner's figure of the tragus,howBA^er, does not show the form typical of this genus.Nitidus {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadelphia,1862, p. 247. From the original description as well as from themore detailed later account with figures (Bull. U. S. Nat. Mus., No.43 (1893), p. 94, pi. 12, March 14, 1894), there is no doubt that thisname applies to the bat which had already been described as Ves-pertilio califormicus by Audubon and Bachman. This fact DoctorAllen himself partly recognized; but he regarded a new name asdesirable on account of the supposed impossibility of identifyingVespertilio californicus with full certainty. His specimens werefrom Monterey, Calif., and Fort Steilacoom, Wash., therefore in-cluding representatives of two geographical races. Miller (1897),the first reviser, selected Monterey as the type locality andgave a new name (caurinus) to the dark northwestern form. Forreasons which appear to us to be insufficient Lyon and Osgood (Bull.U. S. Nat. Mus., No. 62, p. 272, January 28, 1909) reversed the de-cision of Miller by regarding one of the specimens from FortSteilacoom as the type. (On this subject see also H. W. Grinnell,Univ. California Publ. Zool., vol. 17, p. 283, January 31, 1918.)Nubilus (Vespertilio) Wagner, Schreber's Siiugthiere, suppl., vol.5, p. 752, 1855. This was described by Wagner from Natterer's speci-mens, the same apparently that served Temminck for his descriptionof Vespertilio albescefis (not of Geoffroy). They were collected insouthern Brazil and with their f cheekteeth, slender tragus, andwings from the base of the toes were evidently representatives of aspecies of Myotis. Wagner describes the coloring thus : " Die Haaresind am Grunde schwarzbraun, auf der Oberseite allmahlig in's russigKastanienbraun iibergehend, auf den Bauch mit gelbbraunlichenSpitzen." The forearm was 1 inch 4 lines long or about 37 mm.From this description it seems clear that Wagner's bat was, like Tem-minck's albescens^ the M. ruber of Geoffroy, 1805.Obscurus {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadel-phia, 1866, p. 281. As first shown by Goldman (Proc. Biol. SocWashington, vol. 27, p. 102, May 11, 1914), the type of this species,from Lower California, proves to be the same as Vespertilio ymnan-ensis H. Allen, 1864.Occultus {Myotis) Hollister, Proc. Biol. Soc. Washington, vol. 22,p. 43, 1909. This name is applicable to a peculiar and imperfectlyknown species inhabiting the southwestern United States and adjoin-ing parts of Mexico.Oregonensis {Vespertilio) H. Allen, Monogr. Bats North America,Smithsonian Misc. Coll., No. 165, p. 61, June, 1864. This was origi-nally a manuscript name written by LeConte on the label of a speci-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 25men of Myotis californieus bearing no more definite indication oflocality than the words " United States." In placing it formally inprint, H. Allen mentions three other specimens (one from Cape St.Lucas, Lower California, and two from Fort Yuma, Calif.), forwhich, if they should prove to be distinct from his V. nitidus {=M.californieus) , " this name will be reserved." "While the first of thesespecimens is doubtless best considered as true Myotis californian^^the two others are M. californieus follidus^ so that the form as origi-nally understood was a composite. In view of these facts it seemsbest to follow Miller, 1897, as first reviser, in relegating the nameoregonensis to the synonymy of M. californieus californieus.Orinomus {Myotis) Elliot, Publ. Field Columbian Mus., zool. ser.,vol. 3, p. 228, June, 1903. This bat, described from La Grulla, SanPedro Martir Mountains, Lower California, is the same as M. suh-ulatus melanorhinus (Merriam), 1890, from San Francisco Moun-tain, Arizona,Oxyotus (Vespertilio) Peters, Monatsber. k. Akad. Wiss. Berlin, p.19, 1866. This is the first name based on the long-eared form ofMyotis chiloensis occurring in the Andes. The type came fromChimborazo, Ecuador.Pallidus {Myotis californieus) Stephens, Proc. Biol. Soc. Washing-ton, vol. 13, p. 153, June 13, 1900. This is the pale race of Myotiscalifornieus inhabiting the more arid parts of the Southwest. Thetype came from Vallecito, San Diego County, Calif.Parvulus {Vespertilio) Temminck, Monogr. de Mamm., vol. 2, p.246, 1840. Dobson states that he directly compared Temminck's typeof Vespertilio parvulus in the Leiden Museum with an alcoholicspecimen of Myotis nigHcans and considered them as representativesof one species. The type was collected by Natterer in Brazil.Peninsularis {Myotis) Miller, Ann. and Mag, Nat, Hist., ser, 7,vol, 2, p. 124, 1898. This is the only name which has been applied tothe small form of Myotis velifer inhabiting Lower California,Pernox {Myotis) Hollister, Smithsonian Misc, Coll,, vol, 56, No, 26,p. 4, December 5, 1911. The specimen on which this name was basedproves to be a large, richly colored individual of M. lucifugusalascensis. It was collected at Henry House, Alberta, Canada.Pilosus {Vespertilio) Peters, Monatsber. k. Akad. Wiss. Berlin, 1869,p. 408. The origin of the wings from the tibia and from the middleof the back close to the spine, as well as the very large size of thefoot, are characters so different from those of any other Americanmember of the genus, and so like those of the Philippine Myotisinacrotarsus., that until the occurrence in South America of an ani-mal possessing these peculiarities can be established by the discoveryof a second specimen, it is difficult to avoid the conclusion that the58518?28 3
26 BULLETIN- 144, UNITED STATES NATIONAL MUSEUMtype probably came from the Far East rather than, as supposed,from Montevideo, Uruguay.Polythrix {Vespertilio) I. Geoffroy, Ann. des Sci. Nat., ser. 1, vol. 3,p. 443, 1824. There is little in the original description that wouldcertainly identify this bat, which in size is said to be slightly largerthan the common pipistrelle of Europe, its color " brun-marrontres fonce " above ; below, the same with a slight touch of grayish
;
forearm 38 mm. The original specimens were sent from southernBrazil with others from Rio Grande do Sul and Minas Geraes.Gervais examined the types in 1855 and referred them to Myotis(as a subgenus) since they appeared to have teeth back of thecanines. He describes their pelage as " brun, legerement nuance deroussatre." The color and size indicate that they are specimens o*Myotis ruber, to the synonymy of which the name poJythiiic maytherefore be referred.Punensis {Myotis) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol.33, p. 383, July 9, 1914. Four specimens of Myotis, two from PunaIsland, Ecuador, and two from Daule on the neighboring mainlandwere regarded by J. A. Allen as distinct from M. nigricans. Theirsomewhat pale color appears to us to be within the limits of indi-vidual variation in true nigricans.ftuercinus {Myotis californicus) H. W. Grinnell, Univ. CaliforniaPubl. ZooL, vol. 12, p. 317, December 4, 1914. Mrs. Grinnell en-deavored to restrict the name M. californicus to the form of thesmall-footed brown bat occurring in "that portion of Californianorth of about latitude 36 degrees and west of the desert divides,"and applied the new term quercinus to the form of southern andsouthwestern California. Unfortunately, however, Audubon andBachman's original characterization of the color of californicus as
"light yellowish brown" hardly applies to those specimens northand west of 36? so well as to those south of it, and Miller in 1897had already described the darker northwestern animal as caurinus.It seems best therefore to regard quercinus as a synonym of cali-fornicus. The type locality is Seven Oaks, San Bernardino County,Calif.Ruber {Vespertilio) E. Geoffroy, Ann. Mus. d'Hist. Nat. Paris,vol. 8, p. 204, 1806. This is the first name applied to the large russetor reddish-brown Myotis of South America, the " chauve-sourisonzieme " of Azara. It is assumed that Azara's description, on whichthe name is based, was taken from Paraguayan specimens.Salarii {Vespertilio) F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat. Paris,vol. 1, p. 15. 1832. The original description of this bat, based on aspecimen sent by Milbert from the vicinity of New York, is notsufficient for a clear determination. Apparently, however, it appliesto Myotis lucifugus.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 27Saturatus {Myotis yumanensis) Miller, North Amer. Fauna, No. 13,p. 68, October 16, 1897. This is the only name based on the darksubspecies of yumanensis inhabiting the humid area of the northwestcoast of the United States and British Columbia. The type localityis Hamilton, Skagit County, Wash.Septentrionalis (VespertiNo gryphus var.) Trouessart, Catal. Mamm.viv. fOSS., p. 131, 1897. Trouessart established this name by latinizingH. Allen's "northern form of VespertiUo gryphus'''' (Monogr. BatsNorth Amer. (1893), p. 80, March 14, 1894). The animal to which itwas applied is the wide-ranging eastern long-eared bat for whichthe name Myotis subulatus has been current during the past 30years. Say's VespertiUo subulatus^ 1823, proves to have been notthis species but the one which has received the names leibii, cilio-labruni^ melanorMnus, orinoTrms, and winnemana. As Trouessart'sname is the only one which was certainly based on the easternlong-eared Myotis it must now be applied to that animal, but in thetrinomial form Myotis keenii septentiionalis.Simus {Myotis) Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol. 7,p. 541, June, 1901. This is the first and only name based on a verydistinct and peculiar South American species readily distinguishablefrom all other known members of the genus.Sociabilis {Myotis yumanensis) H. W. Grinnell, Univ. CaliforniaPubl. ZooL, vol. 12, p. 318, December 4, 1914. This is the dull brown-ish race of the large-footed M. yumanensis, inhabiting the semi-arid region of the western United States, from Montana to the east-ern part of the Pacific Coast States. The type locality is Fort Tejon,Calif.Spixii {VespertiUo) Fischer, Synopsis Mamm., p. Ill, 1829. Thisname was proposed as a substitute for VespertiUo brasiliensis ofSpix, 1823 (not of Desmarest, 1822). It is with little doubt asynonym of Wied's V. nigy^cans published three years previously.Subulatus {VespertiUo) Say, Long's Exped. to the Rocky Moun-tains, vol. 2, p. 65, footnote, 1823. The specific name subulatus hasbeen misapplied since 1864 to the common long-eared Myotis of east-ern North America by practically all writers who have recognizedthe animal as a distinct species. Recently accumulated evidenceshows, however, that it must now be restored to a different and stillimperfectly known bat ; that is, to the animal which, since its originaldiscovery by Say, has received in its several subspecific forms, thenames leibii (1842), ciUolabrum (1886), melanorhinus (1890), ori-nomus (1903), and winnemana, (1913). The reason for this changeis twofold: First, the type locality of VespertiUo subulatus, on theArkansas River near the present town of La Junta, Otero County,southeastern Colorado, is outside of the known range of the easternlong-eared bat, and second, the original description, when read with
28 BULLETIN 144, UNITED STATES NATIONAL MUSEUMspecimens of the eastern animal and of those species actually knownto occur in Colorado at hand for comparison, is immediately seen toapply much less to the former than to a definite one of the latter,namely, the bat on which Merriam based the name ciliolahrum in1886.Tenuidorsalis {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Phila-delphia, p. 283, 1866. An examination of the type (from Cape St.Lucas, Lower California) shows that the specimen on which this namewas based, is identical with Myotis californicus.Thomasi {Myotis) Cabrera, Bol. Soc. Espan. Hist, Nat., Madrid,vol. 1, p. 370, 1901. The structural characters ascribed to this batconvince us that the name applies to the subspecies of M. chilo'ensisoccurring in the Andean region from northern Ecuador to Boliviawhich Peters had previously called Vespertilio oxyotus.Thysanodes {Myotis) Miller, North Amer. Fauna, No. 13, p. 80,October 16, 1897. This is the tenable name for the large Myotis withlong ears and well-fringed interfemoral membrane occurring in thewestern United States and Mexico. The type locality is Old FortTejon, Calif.Velifer {Vespertilio) J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 3,p. 177, December 10, 1890. This is the name of the large drab-grayMyotis with large foot, common in the southwestern United Statesand through Mexico to Guatemala. The type locality is Guadala-jara, Jalisco, Mexico.Virginianus {Vespertilio) Audubon and Bachman, Journ. Acad.Nat. Sci. Philadelphia, ser. 1, vol. 1, p. 93, 1841. This is undoubtedlya synonym of Myotis luci-fugus. The original description andmeasurements apply fairly well to that species and the characteristicdark shoulder spot is especially mentioned. The specimen came fromthe " mountains of Virginia," where M. lucifugus is common.Volans {Vespertilio) H. Allen, Proc. Acad. Nat. Sci. Philadelphia,p. 282, 1866. The correct disposition of this name remained for nearlyfifty years unsettled. It was originally applied to a bat from CapeSt. Lucas, Lower California, but the description was quite inade-quate, and its author subsequently considered it a synonym of hisVespertilio nitldus. Miller later placed both names in the synonymyof Myotis califomicus; but the rediscovery of the type and its ex-amination by Goldman in 1914 proved volans to be the first nameapplicable to the short-eared, long-legged Myotis of western NorthAmerica, then currently known as M. longicrus. The name Myotisvol-ans is therefore tenable in a specific sense, while longicrus remainsavailable for the dark race of the humid northwest coast.Winnemana {Myotis) Nelson, Proc. Biol. Soc. Washington, vol. 26,p. 183, August 8, 1913. This name was given to the small-footed,
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 29black-faced bat, with keeled calcar, inhabiting the eastern UnitedStates (type from Plummer Island, Md.). It is, however, asynonym of Vespertilio leibii based by Audubon and Bachman ona specimen from Ohio many years before, a name whose allocationhas long been misunderstood on account of the apparent rarity ofthe animal to which it was applied.Yumanensis (VespertUio) H. Allen, Smithsonian Misc. Coll., No.165, p. 58, June, 1864. This name, based on specimens from FortYuma, Calif., is the first one applied to the small, large-footedMyotis of the western United States and Mexico.
THE GENERA MYOTIS AND PIZONYX AND THEIR AMERICAN SPECIESGenus MYOTIS KaupMyotis Kaup, Skizzirte Entw.-Gesch. u. Natiirl. Syst. d. Europ. Thierw., vol. 1,p. 106, 1829.?Miller, Ann. and Mag. Nat. Hist., ser. 6, vol. 20, p. 382,October, 1897; North Amer. Fauna, No. 13, p. 55, October 16, 1897.?Tbouessabt, Catal. Mamm. viv. foss.. p. 1283, 1899.
?
Miller, Key LandMamm. Northeastern North Amer., Bull. New York State Mus., vol. 8, No.38 (October, 1900), p. 148, November 21, 1900.?Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 400, March,1901 ; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ. 57,zool. ser., vol. 2, p. 517, June, 1901.
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Miller and Rehn, Proc. Boston Soc.Nat. Hist., vol. 30, p. 255, December 27, 1901.?Elliot, Land and SeaMamm. Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 570,1904.
?
Tbouessabt, Catal. Mamm. viv. foss., suppl., p. 89, 1904.
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Elliot,Check List Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser.,vol. 6, p. 473, 1905.?Miller, Fam. and Gen. Bats, Bull. U. S. Nat. Mus.,No. 57, p. 200, June 29, 1907; List North Amer. Land Mamm. 1911, Bull.U. S. Nat. Mus., No. 79, p. 54, December 31, 1912 ; List North Amer. RecentMamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 68, April 29, 1924.Nystactes Kaup, Skizzirte Entw.-Gesch. u. Natiirl. Syst. d. Europ. Thierw., vol.1, p. 108, 1829 (bechsteinii). Not Niystactes Gloger, 1827.Leuconoe Boie, Isis, p. 256, 1830 (Die Wasserfledermause, a group defined byBoie in Isis, 1825, p. 1206, as including Vespertilio daubcntonii, V. nattereri,V. dasycnetne, and V. mystacinus).?Dobson, Monogr. Asiat. Chiropt., p.126 (subgenus of Vespertilio? Myotis including dasycneme and daubentoniiof Bole's original species. No type designated) ; Catal. Chiropt. Brit,Mus., p. 289, 1878 (subgenus of Vespcrtilio= Myotis)
.
?Thomas, Ann. andMag. Nat. Hist., ser. 7, vol. 13, p. 382, May, 1904 (subgenus of Myotis; M.daubentonii selected as type) ; Journ. Bombay Nat. Hist. Soc, vol. 23,p. 607, 1915 (genus).Vespertilio Keysebling and Blasius, Wiegmann's Archiv fiir Naturgesch., 1839.vol. 1, p. 306 (not of Linnaeus, 17.58).?H. Allen, Monogr. Bats NorthAmer., Smithsonian Misc. Coll., No. 165, p. 46, June, 1864.
?
Dobson, Catal.Chiropt. Brit. Mus., p. 284, 1878.?H. Allen, Monogr. Bats North Amer.,Bull. U. S. Nat. Mus., No. 43 (1893), p. 70, March 14, 1894.Selysius Bonaparte, Iconografla della Fauna Italica, Introd., p. 3, 1841 ( Vesper-tilio mystacinus)
.
Oapaccinius Bonaparte, Iconografla della Fauna Italica, Indice Distrib.. p. 1.1841 (Vespertilio capacoinii).Trilatitus Gray, Ann. and Mag. Nat. Hist., p. 258, December, 1842 (Vespertiliohasseltii, V. macellus=a^versus, and V. blepotis=Minioptcrus sp.).Tralatitus Gervais, Diet. Univ. d'Hist. Nat., vol. 13, p. 213, 1849 (Modificationof Trilatitus Gray, 1842).Br'achyotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, new ser..vol. 2, p. 131, 1856 (Vespertilio mystacinus, V. daubentonii, and V. dasyc-neme). Not Brachyotua Gould, 1827.30
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 31Isotus KoLENATi, Allgcm. deutsch. Naturhist. Zeitung, Dresden, new ser., vol. 2,p. 131, 1856 {Vespertilio nattereri and V. emarginatus)
.
Tralatitius Gray, Ann. and Mag. Nat. Hist., ser. 3, vol. 17, p. 90, February, 1866.(Modification of Trilatittis Gray, 1842.)Pternopterus Petees, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 706,1867 (subgenus of Vespertilio=Mi/otis, type, V. loMpes Peters=VespertUiomuricola Hodgson),Exochitrics Fitzingeb, Sitzungsber. kais. Akad. Wissenesch., Wien, math.-natur-wissensch. Classe, vol. 62, p. 75, 1870 (Vespertilio macrodactyl us, V. hors-fieldii^=V. adversus, and V. macrrjtarsus.)Aeorestes Fitzinger, Sitzungsber. kais. Akad. Wissensch., "Wien, math.-natur-wissensch. Classe, vol. 62, p. 427, 1870 (Vespertilio villosissimus, V. al-bescens, and V. nigricans).Comastes Fitzinger, Sitzungsber. kais, Akad. Wissensch., Wien, math.-natur-wissensch. Classe, vol. 62, p. 565, 1870 (Vespertilio capaccinii, V. mega-podius, V. dasycneme, and V. limnophilus)
.
Euvcspcrtilio Acloque, Fauiie de France, Mammiferes, p. 38, 1899 (Vespertilioemarginatus, V. mystacinus, V. murinus?myotis, V. nattereri, and V. bech-steinii).Rickettia Bianchi, Ann. Mus. Zool. Acad. Sci., St. Petersburg, vol. 21, p.Ixxviii, 1916 (Vespertilio ricketti).Dichromyotis Bianchi, Ann. Mus. Zool. Acad. Sci., St. Petersburg, vol. 21, p.Ixxviii, 1916 (Vespertilio formosus).Paramyotis Bianchi, Ann. Mus. Zool. Acad. Sci., St. Petersburg, vol. 21, p.Ixxix, 1916 (Vespertilio bechsteinii)
.
Genotype.?Vespertilio tnyotis Borkhausen.Characters.?Vespertilionine bats with the maximum known2?2 1?1 3?3number of teeth present, the dental formula : i ^-g> ^ f?r> V"^ 5-01O O i"~l o?
o
3?3w*K3o = 38; sternum, skull, teeth, and external characters primi-tive for the group, none of them presenting any features of unusualspecialization.Distnhution.?Entire mainland of the Eastern and WesternHemispheres to the limits of tree growth ; on the larger islands therange extends to Japan, the British Islands, Newfoundland, Mada-gascar, the Philippines, Samoa, and Australia, and, from SouthAmerica, to the southern Lesser Antilles.Rema/rks.?The bats of the genus Myotis constitute the most ex-tensively distributed of any comparable group of land mammals.Its range is approached in area by that of only two other genera,the nearly related Eptesieus and Piyistrellus. A seeming anomaly indistribution is the fact that the range is not yet known to include theGreater Antilles. There is no unlikelihood, however, that furtherresearch may show that some inconspicuous member of the genusactually inhabits one or more of these islands or that the remainsof an extinct representative may be found as part of the imperfectlyknown fauna whose traces have been left in the caves. Coincident
32 BULLETIN 144, UNITED STATES NATIONAL MUSEUMwith this almost universal distribution is a notably generalized typeof structure. The family Vespertilionidse, it is true, stands highamong the bats in the development of those general characters whichdistinguish the Chiroptera from nonvolant mammals; but withinthe family the genus Myotis is conspicuous for the retention of themost primitive dental formula?lacking only one upper incisor, oneupper premolar, and one lower premolar of the complete eutherianarmature?and for the absence of the specialized features of cranialstructure and individual tooth form which characterize many of thebat genera with limited range. More than 100 forms of Myotishave been described. Thus the genus is one of the largest as well asthe most widely distributed of the entire order. For the most partthe species do not differ conspicuously from each other. They areseldom noticeable in color and usually small in size.It is probable that a further study of these bats will show the de-sirability of arranging them in several genera and subgenera.Thomas (1915) has already used Leuconoe as a genus. At about thesame time (1916), Bianchi, dealing with the Siberian members of thegenus, applied the name Capaccinius, in a generic sense, to a groupcontaining the species capaccinii, dasycneme, longipes, taiwanensis,pequinius^ and ricketti (the last of which he separated subgenericallyas Richettia). In the genus Myotis proper he recognized, in addi-tion to the typical group, the subgenera Dichromyotis^ Ptemopterus^Parainyotis^ Isotiis, and Seh/sius. We have shown (p. 9) that theMyotis vyyotis group has rather important dental characters whichmay prove to separate it definitely from the rest of the genus ; but, inour opinion, no final results can be attained without a detailed studyof the species inhabiting all parts of the world.A satisfactory arrangement of the descriptions which follow wouldrequire that the least specialized members of the genus be first dealtwith, then those showing successively wider departure from a general-ized condition. Such an ideal method is impracticable, however, onaccount of the varying degi'ee of development shown by the structuralcharacters in the different species. Thus in Myotis lucifugus^ thoughthe ears are not unusually long, the feet are relatively large, while inM. ccMfomicus the ears are elongated and the feet small; again inM. keenii { = subulatus of recent authors) the feet are nearly aslarge as in M. lucifugus but the ears are conspicuously lengthened.In Myotis thysanodes^ the species with the most aberrant condition ofthe molar cusps, the general structure shows no very high specializa-tion. In other words, the different parts of the body vary independ-ently, so that there is seldom an obvious correlation between theirdegrees of development from one species to another. We have con-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 33
sequently not been able to find any recognizably different generaltotals of specialization for the members of the group as comparedwith one another, or any main line of modification along which theevolution of the American species has progressed. Since a lineararrangement which would represent a natural sequence is thus clearlyout of the question, for the present at least, the order of enumerationwhich we have chosen is in the main an arbitrary one. It begins withthe longest-known and most wide-ranging American species, Myotisludfugus^ as a fairly generalized representative of the genus. Theother large-footed species follow, then those with small foot andkeeled calcar. Finally we have found it convenient to give separatetreatment to the tropical and South American members of the genus.There is practically no overlapping of the ranges of these and of theforms which occur in temperate North America; furthermore theyare much less completely known.Genus PIZONYX MillerPizomjx Miller, Proc. Biol. Soc. Washington, vol. 19, p. 85, June 4, 1906;Fam. and Gen. Bats, Bull. U. S. Nat. Mus., No. 57, p. 202, June 29, 1907;List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 59,December 31, 1912.
?
Elliot, Check-List Mamm. North Amer., suppl., p. 157,1917.?MiLLEB, List Recent Mamm. North Amer. 1923, Bull. U. S. Nat. Mus.,No. 128, p. 73, April 29, 1924.Gciivotype.?Myotis vivesi Menegaux.Chctracters.?Like Myotis but foot so enlarged that, with the claws,it about equals the length of the tibia ; toes and claws so greatly com-pressed that the width of claw is only about one-eighth the height ofthe claw at base; wing membrane abruptly narrowed in region ofknee ; a large glandular mass in wing membrane near middle of fore-arm; teeth showing a general tendency to heightening of cusps; sec-ond premolar, both above and below, larger than first.Distribution.?Islands and coast of the Gulf of California, Mexico.Remarks.?The genus Plzonyx, though nearly related to Myotis,is well differentiated by the remarkable development of the foot, andby the other characters which have just been mentioned. In size andgeneral appearance, apart from the enlarged feet, the single knownspecies of Pizonyx bears a resemblance to Myotis myotis of thepalearctic region. This resmblance is, however, purely superficial.That Pizonyx can not have been derived from some member of theMyotis myotis group is shown by its retaining the normally developed7)}?., whereas a reduction of this tooth is one of the main character-istics of M. m^yotis and its allies (see p. 10).
34 BULLETIN" 144, UNITED STATES NATIONAL MUSEUMKEY TO THE AMERICAN SPECIES AND SUBSPECIES OF MYOTIS AND PIZONYXFoot greatly enlarged, its length about equal to that of tibia or skull, thedigits and claws conspicuously compressed ; wing membrane abruptly nar-rowed at level of knee ; a glandular mass in membrane between forearm andfifth digit; second premolar, both above and below, larger than first.Pizonyx (p. 33).Size large, exceeding that of any known American species of Myotis;forearm 60 mm. or more; greatest length of skull about 22 mm. (Gulfof Californiu) P. vivesi (p. 209).Foot not greatly enlarged, its length decidedly less than that of tibia orskull; the digits and claws not conspicuously compressed; wing membranenot narrowed at level of knee ; no glandular mass in membrane between fore-arm and fifth digit ; second premolar, both above and below, smaller thanfirst [size in the known American species medium or small ; forearm notknown to attain a length of 60 mm.] Myotis (p. 30).Species Occukking in Tropical and South AmericaWing from tibia and sides of back -close to the median line ("Uruguay" butprobably not American) M. pilosus (p. 208).Wing from foot and sides of body.Breadth of rostrum across canines equal to or greater than interorbitalconstriction; wing attached at ankle M. simus (p. 205).Breadth of rostrum across canines less than interorbital constriction ; wingattached at base of outer toe.Maxillary cheek teeth unusually small relatively to area of palate; skullwith rostrum shortened and brain case enlarged and rounded.M. albescens (p. 200).Maxillary cheek teeth not unusually small relatively to area of palate;skull with normal rostrum and brain case.A well developed sagittal crest always present in adults ; color usuallyreddish M. ruber (p. 197).A well developed sagittal crest rarely present in adults ; color seldomreddish.Skull small, its greatest length usually ranging from 13 to 14.5; teethsmall, the crown of m' usually less than 1.25 by 1.70 mm. ; earusually not extending beyond nostril when laid forward.M. nigricans (p. 175).Greatest length of skull usually more than 14 mm. ; foot usually lessthan half as long as tibia M. n. nigricans (p. 177).Greatest length of skull never as much as 14 mm. ; foot usually morethan half as long as tibia.Skull extremely small ; forehead rising with unusual abruptness.M. n. dominicensis (p. 183).Skull not extremely small ; forehead rising normally.Color probably averaging darker (.southern Mexico).M. n. extremus (p. 181).Color probably averaging lighter (Island of Curagao).M. n. nesopolus (p, 182).Skull large, its greatest length usually ranging from 14 to 16 mm.
;
teeth large, the crown of w' usually more than 1.30 by 1.70 mm.ear always extending beyond nostril when laid forward.M. chiloensls (p. 189).
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 35General color grayish M. c. atacamensis (p. 192).General color not grayish.Size large, greatest length of skull usually more than 15 mm.M. c. alter (p. 194).Size medium, greatest length of skull usually less than 15 mm.Color distinctly yellowish M. c. dinellii (p. 191).Color dark brown.Burnished tips of hairs on back rather conspicuous.M. c. oxyotus (p. 193).Burnished tips of hairs on back obsolete.M. c. chiloensis (p. 190).Species Occubeing in Temperate North AmericaUnder side of wing furred to level of elbow ; skull with rostrum shortened andocciput unusually elevated (western North America) M. volans (p. 135).Size small, forearm about 35 mm., length of skull less than 13 mm. (LowerCalifornia) M. v. volans (p. 139).Size larger, forearm 37-40 mm. ; length of skull more than 13 mm.Color buffy or ochraceous (interior of western United States).M. V. interior (p. 142).Color dark brown.Upper parts dark reddish brown (northwest coast region).M. V. longicrus (p. 140).Upper parts ochraceous tawny (southern Mexico) M. v. amotus (p. 145).Under side of wing not furred to level of elbow ; skull with normal rostrumand occiput.Foot small, the ratio of its length to that of tibia usually ranging from about40 to 46.Hairs of back with long shiny tips; third metacarpal not so long as fore-arm; skull larger, with flattened braincase and gradually rising pro-file (United States) M. subulatus (p. 164).Belly warm buff, back ochraceous tawny (eastern United States).M. s. leibii (p. 171).Belly whitish.Upper parts very pale, light buff to warm buff (Interior United States).M. s. subulatus (p. 168).Upper parts less pale, bright golden above (southwestern UnitedStates) -M. s. melanorhinus (p. 169).Hairs of back dull-tipped ; third metacarpal usually as long as forearm
;
skull smaller, with rounded braincase and abruptly rising profile (Mex-ico and western United States) M. californicus (p. 148).Size larger, forearm usually 33.6 to 36 mm. (southern Mexico).M. c. mexicanus (p. 159).Size smaller, forearm usually 30 to 34 m.Fur of back distinctly tricolor, the hairs with a dark base succeeded bya pale ring and a pale reddish tip (deserts of western UnitedStates) __?M. c. pallidus (p. 157).Fur of back not distinctly tricolor.Upper parts ochraceous tawny (western United States).M. c. californicus (p. 151).Upper parts nearly mahogany red (northwest coast region).M. c. caurinus (p. 155).
36 BULLETIN 144, UNITED STATES NATIONAL MUSEUMFoot normal or large, the ratio of its length to that of tibia usually rangingfrom about 48 to 60.Wing membrane attached to tarsus ; fur of back without obviously darkenedbasal area; ratio of foot to tibia usually about 60 (Indiana and Illinoisto Alabama and Georgia) M. grisescens (p. 80).Wing membrane attached to side of foot; fur of back with obviously dark-ened basal area ; ratio of foot to tibia usually less than 57.Fur of back with an obvious tricolor pattern ; calcar usually with a smallbut evident keel (eastern United States) M. sodalis (p. 130).Fur of back without an obvious tricolor pattern ; calcar normally withno trace of keel.Ear when laid forward extending noticeably beyond tip of muzzle.Free border of uropatagium with inconspicuous, scattered, stiff hairs(central and northern North America) M. keenii (p. 101).Color darkened ; a saturate northwest coast form.M. k. keenii (p. 104).Color normal ; a normal eastern and central form.M. k. septentrionalis (p. 105).Free border of uropatagium usually with a noticeable fringe of stlflEhairs.Size larger, forearm usually 41 to 46 mm. ; ear not exceptionallyenlarged; fringe conspicuous (Mexico and western UnitedStates) M. thysanodes (p. 122).Under parts whitish, upper parts "warm buff" (western UnitedStates and northern Mexico) Iff. t. thysanodes (p. 126).Under parts huffy, upper parts darker, " tawny olive " (southernMexico) M. t. aztecus (p. 128).Size smaller, forearm usually 33 to 40 mm.; ear exceptionally en-larged; fringe not conspicuous (western United States andnorthern Mexico).Skull with noticeably flattened brain case; forearm 33 to 36 (SanPedro Martir Mountains, Lower California).M. milleri (p. 118).Skull with normal brain case ; forearm usually 37 to 40.M. evotis (p. 111).Upper parts darker, near clay color (northwest coast region).M, e. evotis (p. 114).Upper parts lighter, flaxen or cinnamon buff (range of thespecies except northwest coast region).M. e. chrysonotus (p. 116).Bar when laid forward not extending noticeably beyond tip of muzzle.Cheek teeth robust, the breadth of the maxillary molars, as com-pared with that of the intervening palate, greater than usualin American members of the genus.Brain case flattened; pelage glossy (southwestern United States).M. occultus (p. 97).Brain case highly arched; pelage dull M. velifer (p. 86).Color darker (Mexico and western United States).M. V. velifer (p. 89).
AMEEICAN BATS OF THE GENERA MYOTIS AND PIZONYX 37Color paler.Size maximum for the species ; greatest length of skull 15.8 to17.6 mm. (Kansas, through Texas and New Mexico, toDurango) M. v. incautus (p. 92).Size minimum for the species; greatest length of skull 14.2 to15.6 mm. (Lower California M. v. peninsularis (p. 93).Cheek teeth normal, the breadth of the maxillary molars as comparedwith the intervening palate not greater than usual in Americanmembers of the genus.Fur above dense, wooly ; a low but evident sagittal crest alwayspresent in adults (Florida; Indiana; Saskatchewan?).M. austroriparius (p. 76).Fur above normal, silky ; a sagittal crest rarely present.Fox-earm ranging from 32 to 37 mm. ; greatest length of skullranging from 13.2 to 14.2 mm.; hairs of back without con-spicuous burnished tips (western North America).M. yamanensis (p, 61).Color paler.Above pale buffy, below almost white (deserts of GreatBasin) M. y. yumanensis (p. 65).Above tawny olive, below dull whitish, tinged with buffy(western United States) M. y. sociabilis (p. 68).Color darker.Membranes and ears blackish brown, fur dull sepia above(northwest coast) M. y. saturatus (p. 70).Membranes and ears dull brownish, fur dark cinnamonbrown above (southern Mexico) M. y. lutosus (p. 72).Forearm ranging from 36 to 40 mm. ; greatest length of skullranging from 14.3 to 15.3 mm. ; hairs of back with conspicu-ous burnished tips (North America in general).M. lucifugus (p. 38).Small premolars unusually crowded (Mexico and westernTexas) M. 1. fortidens (p. 54).Small premolars not unusually crowded.Color above ranging from blackish to very dark brown(northwest coast) M. 1. alascensis (p. 47).Color not ranging from blackish to very dark brown.Belly with a distinct bufiCy wash ; membranes not edgedwith white (region east and north of Rocky Moun-tains) M, 1. lucifugus (p. 43).Belly whitish ; membranes edged with white.General color above cinnamon buff, ears black, edgesof membranes often whitish (semiarid westernUnited States) M. 1. carissima (p. 50).General color above pale pinkish buff, ears pale brown-ish (Great Basin) M. 1. phasma (p. 53).
38 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
I. SPECIES OF MYOTIS OCCUERING IN TEMPERATE NORTH AMERICAMYOTIS LUCIFUGUS (LeConte)(Synonymy under subspecies)Distribution.?From the limit of tree growth in Labrador, Can-ada, and Alaska, southward to Tabasco, Mexico.
Map 1.
?
Distribution of Mtotis lucifdgus ; 1, M. lucifdgus lucifugcs; 2, M.LUCIFUGDS ALASCENSIS ; 3, M. LUCIFUGUS CABISSIMA ; 4, M. LUCIFUGUS PHASMA :5, M. LUCIFDGUS FORTIDHNSDiagivosis.?Size medium among the American species; forearmabout 38 (36 to 40) mm.; total length usually about 80 to 86 mm.,tail about 36 mm., averaging 45 per cent of the total length (averageratio of tail to head and body in 10 specimens from Nova Scotia,77.8; in 10 from Montana, 76.2; in 10 from the northwest coast,75.8) ; greatest length of skull ranging from 14.0 to 15.8; maxillarytooth row ranging from 5.0 to 5.8 mm.; lower tooth row (excluding
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 39incisors) usually more than 5.6 (5.4 to 6.2) ; ear reaching nostrilwhen laid forward, its height from meatus usually 13 to 15 mm.;wing from the side of foot, which is large, exceeding one-half thetibia (ratio of foot to tibia ranging from about 53 to about 55.7) ;calcar without keel ; hairs of back with long glossy tips which impartto the pelage a conspicuous metallic sheen. Skull with graduallyrising forehead and broad brain case which is usually not providedwith a distinct sagittal crest. Upper molars with full complementof secondary cusps and ridges, but with a narrow, inconspicuousand sometimes incomplete cingulum on inner border of crown.Ears.?The ear is of moderate length, when laid forward reach-ing to the nostril ; its anterior edge is convex, becoming nearlystraight in the upper third to the bluntly rounded tip; below thetip the .external margin is slightly concave or nearly straight inthe upper half, then convex in the lower half which forms a pro-jecting, but not abrupt, shoulder. The tragus is about half thetotal height of the ear (8:15 mm.). At its outer base is a smallrounded lobe, marked off by a shallow notch. Above this point andopposite the inner base is the widest region. The inner margin ofthe tragus is nearly straight, the outer very slightly convex, andfaintly crenulate to the narrowly rounded tip.
'Wing and Tnemhrames.?The wing membrane arises from the sideof the foot near the distal end of the metatarsal. The third to fifthmetacarpals are graduated, the third longest, the fourth slightlyshorter, the fifth still shorter. The basal phalanges and total lengthof the digits are likewise graduated in the same order. Taking thethird finger as 100, the fourth and fifth fingers are, respectively, as86 and 80 (60:52:48 mm.). When folded, the third metacarpalusually falls short of the elbow by about 2.5 mm. The extreme tipof the tail is free. Uropatagium with minute hairs along the freeborder. These hairs are few and widely scattered, very inconspicuous.Foot.?The foot is rather large, the ratio of its length to thatof the tibia ranging from about 53 to nearly 56. Calcar long (about17 mm.), exceeding in length the free border of the interfemoralmembrane, terminating in a minute lobule ; its free edge quite with-out indication of a keel. Outer toe a little shorter than the others.A slight web between bases of toes; terminal joints with a few verysmall stiff hairs.Fur and color.?The pelage is rather full, but not fleecy, long andfine above, the longer hairs attaining a length of about 9 to 10 mm.On the ventral surface of the wing membrane the fur extends thinlyfrom the knee to the distal three-fourths of the humerus. The inter-femoral membrane is thinly haired about to a line joining the knees.Characteristic are the long glossy tips of the dorsal hairs which
40 BULLETIN 144, UNITED STATES NATIONAL MUSEUMmay completely conceal the dark bases and give a uniformly shiningappearance to the coloration of the upper side. In a well-madeskin, this glossy sheen to the fur is usually sufficient in itself toseparate the species from others with which it occurs. A darkshoulder spot is usually present.Shall.?The skull, viewed from above (pi. 1, p. 7, fig. 12), has therostrum relatively short (distance from anterior margin of alveolusof inner incisor to narrowest part of interorbital constriction decidedlyless than that from narrowest part of constriction to lambda) andnarrow (maxillary breadth at m^ usually more than 1 mm. less thanbreadth of brain case) ; anterior breadth across roots of caninesobviously less than interorbital constriction, and the forehead, asseen in profile, has a slight gradual upward slope. The brain caseis somewhat flattened and subcircular in dorsal view, its area notice-ably greater than that of rostrum. Even in adults the sagittal crestis seldom developed (see footnote, p. 133), though it is sometimesindicated by a low median ridge, which, posteriorly at least, isnarrow and sharply defined. The lambdoid crests usually do notquite meet at the vertex.Teeth.?The teeth are of normal size relatively to the rostum andpalate (pi. 1, p. 7, fig. 36). Length of maxillary tooth row lessthan greatest palatal width across molars. Upper molars with fullmaximum development of secondary cusps and ridges (fig. la and 1<?,p. 8) , the protoconule always present and well developed, the hypoconewith base well marked off from that of the protocone and with apexsometimes distinct from the inner loph; cingulum variable in itsdevelopment, usually obvious on lingual margin of crown, but rarelyif ever continuous around the antero-lingual face of the protocone;exceptionally the cingulum may be almost or quite absent. The twosmall upper premolars are usually somewhat drawn inward fromthe tooth row, but the tips of both teeth are visible from the outerside.Remarks.?As a species Myotis lucifugus has a broader area ofdispersal than any other member of its genus in the Western Hemi-sphere. Over an astonishingly wide range it holds its structuralcharacters very closely, with remarkably little variation in size andproportions. In color, however, it shows the usual response to dif-ferences in climate, tending to become dark in areas of high humidityand pallid under conditions of dryness. It is usually associated withat least moderate tree growth and particularly delights to feed alongsmall water courses in the forest. It is therefore less common in ourarid Southwest, if not altogether absent from areas of actual desertconditions; in such places it is outnumbered by the smaller, duller-colored M. 'i/umanensis. In many localities, however, the two speciesoccur together along the borders of thinly wooded country, and
AMEEICAN BATS OF THE GENEEA MYOTIS AND PIZONYX 41Allen collected local forms of both at the same spot by an alpinestream at the limit of trees, 11,000 feet, on Mount Whitney, Cali-fornia. At this locality the animals were probably summer invadersfrom lower altitudes, M. yumanensis perhaps from the desert at theeast base of the Sierra, whence additional examples were later ob-tained. Myotis lucifugus is a social species, breeding in colonies,sometimes in caves, frequently in dark recesses of buildings. Itwinters in caves in the temperate parts of its range.Throughout eastern North America this is by far the commonestspecies of Myotis^ but in western North America it is less abundant,especially at lower elevations and in desert country.In the Eastern Hemisphere Myotis lucifugus is represented byMyotis dauhentonii. The only external characters by which trueMyotis dauhentonii and the larger footed forms of M. lucifugus canwith certainty be distinguished appear to be that in Europeanspecimens {a) the toes are elongated and the claws distinctly en-larged, (6) the insertion of the wing membrane is rather on the basalthan the terminal half of the metatarsus, and (c) the fur of theupperparts is not conspicuously glossy and the color of the underpartsis more contrastedly paler than that of the back. The skulls of thetwo species are alike in size and in all essential characters; but intypical M. dauhentonii the dorsal profile from the middle of thebrain case backward tends to be evenly convex, while in M. lucifugusthere is usually some indication of a concavity at front of occiput,the extreme posterior region tending to rise a little above level ofmiddle of brain case. We have not been able to find any tangibledifferences between the teeth of the two animals (compare teeth ofM. dauhentonii, figured by Miller,^ and those of M. lucifugus, figuredby H. Allen ).^ The resemblance betAveen the two species was notedby H. Allen in 1894,^ but we have not seen it mentioned by any otherrecent author.The occurrence of Myotis lucifugus in Kamchatka was recordedby Hahn in 1905? on the basis of a specimen in alcohol (No. \}\l%U. S. Nat. Mus.) collected at Petropavlovsk by Mr. F. Wliymper andbrought to Washington by Dr. W. H. Dall. (It was entered in thecatalogues Jan. 10, 1873). Another individual, not mentioned byHahn, was taken by Grebnitzki at the same i^lace in July, 1889(No. 155523 U. S. Nat. Mus.). Both specimens are in poor conditionand neither of them has a perfect skull. After careful examinationwe are unable to distinguish them, on the basis of the characters
" Cat. Mammals Western Europe, Brit. Mus., p. 186, Nov. 23, 1912.?Monogr. Bats North Amer. (1893), pi. 11, Mar. 14, 1894.* Monogr. Bats North Amer., p. 79.Proc. Biol. Soe. Washington, vol. 18, p. 254, Dec. 9, 1905.58518?2S 4
42 BULLETIN 144, UNITED STATES NATIONAL MUSEUMwhich they show, from M. lucifugus. There is a strong probability,however, that they represent the animal which Ognev has recentlydescribed from Vladivostok as Myotis daubentonii usswie7isis,^ andin the absence of more complete evidence we prefer not to regardtheir presence in Kamchatka as undoubted proof that an Americanbat has extended its range into the Old World.Nomenclature.?Without doubt the earliest tenable name for thisspecies is lucifugus LeConte (1831) as pointed out by Miller (1897).The original description, though all too general and brief, appliedwell enough to this bat, and in a more amplified account, LeConte(1855) made it appear reasonably probable that the species was thecommon short-eared Myotis of the eastern United States, though hesupposed that his specimens represented two animals, one of whichhe wrongly took to be the same as Say's Vespertilio suhulatus. TheBritish Museum now possesses a skin of the species (No. 7.1.1.534,formerly No. 4741, U. S. Nat. Mus.), received by Tomes manyyears ago from the Smithsonian Institution. There is no question asto the identity of this specimen, and the original record in the cata-logue of the United States National Museum shows that it camedirectly from LeConte. It was entered as Vespertilio lucifugv^s, butunfortunately without indication of locality. Several writers subse-quently characterized supposed new species which can be no otherthan this bat. Among these forms are three described by F. Cuvierin 1832 {Vespertilio gryphus, V. salarii^ and V. crassus), all from
" New York ; " the first and second sent to Paris by Milbert, the lastby Lesueur. The essential parts of the descriptions are much alike,and seem in all probability to refer to slight variations in the samespecies, allowing for a discrepancy in giving &t^assus only two pre-molars. Temminck's Vespertilio carolii^ from Philadelphia and NewYork, is unquestionably a Myotis as shown by the number of teeth;and the rest of the description fits M. lucifugus perfectly. Green'sF. domestieus and Audubon and Bachman's V. virginianus appearalso to be the same. An immature specimen with supposedly peculiarcharacters served as the basis for H. Allen's Vespertilio affinis, but anexamination of the type specimen proves that the name is a synonymof lucifugus. LeConte's incorrect identification (1855) of an indi-vidual variation in the short-eared species lucifugus with the Ves-pertilio suhulatus of Say was the cause of much confusion amongsubsequent authors (H. Allen, 1864, and Dobson, 1878, excepted).Thus the blanket name " Vespertilio suhulatus " came to be usedrather indiscriminately for the two common small brown bats of theeastern United States until the specific distinctness of Myotis luci-fugus and M. " suhulatus " ( = 3/. keenii septentnonalis of the pres-ent paper) was finally made clear by Miller in his review of 1897.
? Journ. Mamm., vol. 8, p. 146, May 12, 1927.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 43MYOTIS LUCIFUGUS LUCIFUGUS (LeConte)Vespertilio lucifugus LeConte, McMurtrie's Cuvier, Animal Kingdom, vol. 1,appendix, p. 431, 1831 ; Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 436,1855.?H. Allen, Mouogr. Bats North Amer., Smithsonian Misc. Coll., No.165, p. 55, fig. 51-52, June, 1864.?True, Proc. U. S. Nat. Mus., vol. 7, p. 603,1885.?Miller, Proc. Boston Soc. Nat. Hist., vol. 28, p. 39, April, 1897.Vespertilio gryphus F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat. Paris, vol. 1, p.15, 1832 (New York).-Temminck, Monogr. de Mamm., vol. 2, p. 259, 1840.?H. Allen, Monogr. Bats North Amer., Bull. U. S. Nat. Mus., No. 43 (1893),p. 75, March 14, 1894 (part).? Vespertilio salarii F. Cuvier, Nouv. Ann. Mus. di'Hist. Nat. Paris, vol. 1,p, 16, 1832 (New York).?Temminck, Monogr. de Mamm., vol. 2, p. 259,1840.? Vespertilio crassus F. Cuvier, Nouv. Ann. Mus. d'Hist. Nat. Paris, vol. 1,p. 18, 1832 (New York).
?
Temminck, Monogr. de Mamm., vol. 2, p. 261,1840. JVespertilio domesticus Green, Doughty's Cabinet of Nat. Hist., vol. 2, p. 290,1832 (western Pennsylvania).? Vespertilio lanceolatus Maximilian zu Wied, Reise in das Inuere Nord-Amer.,vol. 1, p. 364, footnote, 1839 (Bethlehem, Pa.).Vespertilio carolii Temminck, Monogr. de Mamm., vol. 2, p. 237, 1840 (Phila-delphia and New York).
?
Dobson, Catal. Chiroptera Brit. Mus., p. 325, 1878.
?True, Proc. U. S. Nat. Mus., vol 7, p. 603, 1885.Vespertilio virginianus Audubon and Bachman, Jouru. Acad. Nat. Sci. Phila-delphia, ser. 1, vol. 1, p. 93, 1841 (mountains of Virginia).Vespertilio subulatus LeConte, Proc. Acad. Nat. Sci. Philadelphia, 1855, p. 435(not of Say, 1832).Vespertilio hrevirostris Maximilian zu Wied, Verzeich. beobacht. Siiugeth. N.Amer., p. 19, 1860; Arch. f. Naturgesch., 1861, vol. 1, p. 195 (Freiburg, Pa.).Vespertilio affinis H. Allen, Monogr. Bats North Amer., Smithsonian Misc.Coll, No 165, p. 53, fig. 48-50, June, 1864 (Fort Smith, Ark.).?Lyon andOsgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No.62, p. 270, January 28, 1909.Vespertilio gryphus lucifugus H. Allen, Monogr. Bats North Amer.,- Bull. U. S.Nat. Mus., No. 43 (1893), p. 78, March 14, 1894.Vespertilio albescens afflnis H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 93, March 14, 1894.?Trouessart, Catal. Mamm.viv. foss., p. 132, 1897.Vespertilio gryphus var. lucifugus Trouessart, Catal. Mamm. viv. foss., p. 131,1897.MyoHs lucifugus Miller, North Amer. Fauna, No. 13, p. 59, October 16, 1897.
?
Trouessart, Catal. Mamm. viv. foss., p. 1284, 1899.?Miller, Prelim.List New York Mamm., Bull. New York State Mus., vol. 6, No. 29 (October,1899), p. 365, November 18, 1899; Key Land Mamm. Northeastern NorthAmer., Bull. New York State Mus., vol. 8, No. 38 (October, 1900), p. 149,November 21, 1900.
?
Elliot, Synops. Mamm. North Amer., Field Columb.Mus., publ. 45, zool. ser., vol. 2, p. 402, March, 1901; List Land and SeaMamm. North Amer., Field Columb. IVIus., publ. 57, zool. ser., vol. 2, p. 517,June, 1901.
?
Miller and Rehn, Proe. Boston Soc. Na4;. Hist., vol. 30. p.255, December 27, 1901.?Elliot, Land and Sea Mamm. Middle Amer.,Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 580, 1904.
?
Trouessart,Catal. Mamm. viv. foss., suppl., p. 92, 1904.?Elliot, Check List Mamm.North Amer^ Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 478, 1905;
44 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCatal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115, zool.ser., vol. 8, p. 504, 1907 (part).
?
Hahn, Mamm. of Indiana, 33d Ann. Rep.Dept. Geol. and Nat. Resources of Indiana, 1908, p. 621, 1909.
?
Cory, Mamm.of Illinois and Wisconsin, Field Mus. Nat. Hist., publ. 153, zool. ser., vol,11, p. 455, 1912 (part).?G. M. Allen, Bull. Mus. Comp. Zool., vol. 52,p. 44, June, 1908.llyotis lucifugus lucifugus Miller, Key Laud Mamm. Northeastern NorthAmer., Bull. New York State Mus., vol. 8, No. 38 (October, 1900), p. 149,November 21, 1900.
?
Setton, Life Hist. Northern Anim., p. 1148, 1909.
?
Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79,p. 55, December 31, 1912; List North Amer. Recent Mamm. 1923, BulLU. S. Nat. Mus., No. 128, p. 68, April 29, 1924.?Bailet, North Amer.Fauna, No. 49 [December, 1926], p. 213, January 8, 1927 (North Dakota).Type locality.?Georgia; probably the LeConte plantation, nearRiceboro, Liberty County.Type specimen.?None designated. The British Museum has amade-over skin received with the Tomes collection, that was origi-nally No. 4741 of the United States National Museum. It is labeled
:
VespertUio " lucifugus^'' United States, Maj. LeConte. This is atypical Myotis lucifugus lucifugus as now understood, and may wellbe one of the specimens on which LeConte based his original accountof the animal.Distribution.?The entire forested portion of North America northof the southern border of the United States except in the RockyMountain region and on the Pacific .coast of California, Oregon,Washington, British Columbia, and southern Alaska. (See map 1,p. 38.)Diagnosis.?Color above ranging from yellowish brown (bronzy)to olive brown; a dark area at the shoulder; belly gray with a richbuffy suffusion; membranes not pale-edged; ratio of foot to tibiausually ranging from about 53 to 54.Description.?Two extremes or phases of color are found in theeast. In the one the fur above is a glossy yellowish brown almostbronze, in the other it is browner (U. S. N. M., No. 159411 fromYoung Harris, Georgia) nearly " ochraceous-tawny " (of Ridgway,1912) varying individually in the depth of the bronzy tint to an olivebrown (near "Dresden brown"). At the shoulder there is a smallcontrasting area of darker, nearly blackish brown, forming an ill-defined spot. Bases of the hairs above and below blackish or darkplumbeous. The hairs of the lower surface are tipped with a richbuff, nearly the chamois of Ridgway, those at the extreme posteriorpart of the body without dark bases. Immature individuals aresooty gray above, the glossy tips of the hairs inconspicuous. Earsand membranes blackish brown.Measui'evients.?For measurements see tables, pages 56 and 59.
AMEEICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 45Specwiens exatrdned.?Total number 744, from the followinglocalities
:
ALASKA: Anchorage, 1 head (XI.S.N.M.) ; Bristol Bay, 1 ale. (U.S.N.M.) ;Iliamna Lake, 1 ale. (U.S.N.M.) ; Kodiak Island, 3 skins, 5 ale. (U.S.N.M.).ALBERTA: Canadian National Park, 3 skins (A.M.N.H.).ARKANSAS: no exact locality, 1 ale. (M.C.Z.) ; Fort Smith, 1 ale., typeof alfinis (U.S.N.M.).ATHABASCA: Athabasca River, 1 skin (U.S.N.M.).CANADA: Jame.s Bay, 1 ale. (U.S.N.M.).?CONNECTICUT : Greenwich, 1 ale. (U.S.N.M.); New Haven, 1 nlc.(U.S.N.M.).DISTRICT OF COLUMBIA: Washington, 7 skins, 20 ale, 1 skeleton,(U.S.N.M.).
-GEORGIA: Young Harris, 1 skin, 1 ale. (U.S.N.M.); no exact locality,1 ale. ( M.C.Z. ).ILLINOIS: Evanston, 1 ale. (M.C.Z.); Golconda, 3 skins (F.M.) ; OgleCo., 5 skins (M.C.Z.) ; Warsaw, 50 ale. (U.S.N.M.) ; West Northfield,1 skin (U.S.N.M.).INDIANA: Bloomington, 2 skins (U.S.N.M.); Coon Cave. 3 skins(U.S.N.M.); Mitchell, 11 skins (U.S.N.M.); Indianapolis, 3 ale.(A.N.S.P.) ; Wheatland, 3 ale. (U.S.N.M.).IOWA: Grinnell, 1 skin (M.C.Z.) ; McGregor, 5 ale. (U.S.N.M.).XABRADOR: Makkovik, 1 skin (M.C.Z).MACKENZIE: Salt River, 1 skin (U.S.N.M.).MAINE: Cupsuptic River, 1 skeleton (U.S.N.M.); Eastport, 1 ale,(M.C.Z.) ; Lincoln, 3 ale. (A.N.S.P.) ; Norway, 3 ale. (M.C.Z) ; Upton,1 ale. (M.C.Z.); Waterville, 3 ale. (M.C.Z.).^MANITOBA: Hartney, 1 skin (U.C.) ; 1 skin (U.S.N.M.) ; Lake Winnipeg,1 ale. (B.M.).JtfARYLAND: Plummer Island, 1 ale. (U.S.N.M.); Seneca River, 1 ale,(U.S.N.M.).MASSACHUSETTS : Cambridge, 1 skin (M.C.Z.) ; Cotuit, 2 skins (M.C.Z.) ;Harvard, 1 skin (M.C.Z.) ; Lanesboro, 2 ale. (M.C.Z.) ; Littleton, 1 skin(M.C.Z.) ; Wareham. 1 skin (M.C.Z.) ; Woods Hole, 1 ale. (U.S.N.M.).
-MICHIGAN: Charity Island, 10 skins (U.M.) ; Chippewa Co., 4 skins(U.M.) ; Douglas Lake, (Cheboygan Co.), 1 skin (U.M.) ; Isle Royal,Washington Harbor, 10 skins (U.M.) ; Lake Gogebic, Ontonagon Co.,2 skins, 2 ale. (U.M.) ; Leland, Leelanau Co., 1 skin (U.M.) ; LindsleyLake, Gogebic Co., 5 skins, 2 ale. (U.M.) ; Little Girl's Pt. (GogebicCo.). 1 ale. (U.M.) ; Gogebic County (no exact locality), 2 skins (M.C.Z.) ;Michigamme, 16 skins (U.S.N.M.) ; New Baltimore, 8 ale. (U.M.) ; SouthLyon (Oakland Co.), 1 ale. (U.M.).
.MINNESOTA: Albert Lea, 1 skeleton (U.S.N.M.); Devil's Lake (RamseyCo.), 1 ale. (U.M.) ; Elk River, 9 skins (U.S.N.M.), 2 skins (U.M.) ;Fort Snelling, 4 .skins (U.S.N.M.) ; Moorhead, 1 skin (U.S.N.M.) ; ^IslandLake, Lyon Co., 1 ale. (B.M.).JMISSOURI: Columbia, 5 ale. (U.S.N.M.) ; Montgomery City, 1 skin, 2 ale.(U.S.N.M.) ; no exact locality, 82 ale, (M.C.Z.).JVEBRASKA: Fort Pierre, 1 skin (B.M.), 1 skull (U.S.N.M.).NEW BRUNSWICK: Restigouche, 1 skin (M.C.Z.); Trousers Lake, 1skin (A.M.N.H.).
46 BULLETIN 144, UNITED STATES NATIONAL MUSEUMNEW HAMPSHIRE: Antrim, 1 skin (M.CZ.) ; Lacoiiia, 1 ale. (U.S.N.M.) ;Shelburne, 2 ale. (M.C.Z.).NEW JERSEY: Penngrove, 3 ale. (U.S.N.M.); Delaware Water Gap, 1skin (A.N.S.P.).NEW YORK: Big Moose Lake, 1 ale. (U.S.N.M.) ; Catskill Mountains, 2ale. (U.S.N.M.) ; Elmsford, 5 skins, 3 ale. (A.M.N.H.), 1 skin (F.M.) ;Fourth Lake, 1 ale. (U.S.N.M.) ; Howe's Cave, Schoharie County, 51 ale.(U.S.N.M.) ; Keene Valley, 1 skin (U.S.N.M.) ; Lake George, 2 skins, 2.ale. (U.S.N.M.); Locust Grove, 7 ale. (U.S.N.M.) ; Lyon's Falls, 4 ale.(U.S.N.M.) ; Oneida Lake, 4 skins (A.N.S.P.) ; Peterboro, 1 ale.(U.S.N.M.) ; Salem Center, 3 skins (A.M.N.H.) ; Stamford, 1 skia(A.M.N.H.) ; St. Regis Lake, 1 skull (U.S.N.M.) ; Tupper Lake, 1 skin,2 ale. (M.CZ.) ; Tuxedo, 3 ale. (A.M.N.H.) ; West Point, 2 ale. (U.S.N.M.) ;Westport, 1 skeleton (U.S.N.M.) ; no exact locality, 2 skins (B.M.).NEWFOUNDLAND: Bay St. George. 4 skins (M.CZ.) ; Nicholsville, 4rskins (M.CZ.).NORTH CAROLINA: Cherokee, 1 ale. (U.S.N.M.) ; Roan Mountain, 1 skin(U.S.N.M.) ; Weaverville, 2 skins (A.M.N.H.) ; Bertie County, 1 skin(A.N.S.P.).NORTH DAKOTA: Cannon Ball, 4 skins (U.S.N.M.) ; Fort Totten, 2 skins-(U.S.N.M.) ; Goodall, 1 skin (U.S.N.M.) ; Hankinson, 1 skin (U.S.N.M.) ;Stump Lake, 1 skin (U.S.N.M.), approaching carissima; Towner, 28 skins.(U.S.N.M.), approaching carissima.NOVA SCOTIA: Halifax, 41 ale. (U.S.N.M.); no exact locality, 1 ale.(B.M.).OHIO: Circleville, 1 skin (U.S.N.M.).ONTARIO: Linwood, 1 skin (A.N.S.P.); Macdiarmid, Lake Nipigon, 2skins (Roy. Ont. Mus. Zool.) ; Muskoka, 1 skin (M.CZ.) ; Ottawa, 2 ale.(U.S.N.M.) ; Waterloo County, 11 skins (F.M.).PENNSYLVANIA: Big Tink Pond, 1 skin (A.M.N.H.); Kring's Station,Cambria County, 1 skin (A.N.S.P.) ; Lake Ganoga, Sullivan County, 1skin (A.N.S.P.) ; Round Island, Clinton County, 2 ale. (A.N.S.P.).PRINCE EDWARD ISLAND: 1 skin, 1 ale. (U.S.N.M.) ; Mount Herbert,1 ale. (U.S.N.M.).QUEBEC: Godbout, 7 ale. (U.S.N.M.) ; Lac aux Sables, 1 skin (A.N.S.P.) ;Rupert House, James Bay, 1 ale. (U.S.N.M.).SOUTH CAROLINA: Beaufort, 2 ale. (U.S.N.M.).SOUTH DAKOTA: Cedar Island. 1 ale. (U.S.N.M.).TEXAS: Fort Hancock, 1 skin (U.S.N.M.).VERMONT: Brandon, 5 skins (U.S.N.M.); Proctor, 2 skins (U.S.N.M.);Weathersfield, 1 skin (M.CZ.).VIRGINIA: Aylett, 4 ale. (U.S.N.M.); Falls Church, 4 skins, 67 ale,(U.S.N.M.), 2 skins (U.C.) ; Four Mile Run, 1 skin (U.S.N.M.) ; Middle-town. 2 skins, 2 ale. (U.S.N.M.); Riverton, 1 skin (U.S.N.M.); SpringHill, 2 skins (U.S.N.M.).WteST VIRGINIA: Franklin, 1 skin, 1 ale. (U.S.N.M.); White SulphurSprings, 4 skins (U.S.N.M.), 5 skins (M.CZ.), 6 skins (F.M.).WISCONSIN: Beaver Dam, 2 skins (F.M.) ; Cassville, 6 skins, 8 .ale.(U.S.N.M.) ; Delavan, 2 skins (U.S.N.M.), 3 skins (M.CZ.), 1 skiu(U.C.) ; Devil's Lake, 1 skin, 5 ale. (U.S.N.M.) ; De Sota, 1 ale.(U.S.N.M.); Herbster, 1 skin (U.SIN.M.) ; Lake St. Germain, 1 skin(U.S.N.M.) ; Long Lake, 2 skins (U.S.N.M.) ; Madison, 6 ale. (U.S.N.M.) ;Milwaukee, 1 skin (U.S.N.M.) ; Potosi, 1 skin (U.S.N.M.) ; Racine, 1 skin(U.S.N.M.).
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 47WYOMING: Fremont Peak, 3 skins (U.S.N.M.), approaching carissima;Sand Creek, Crook County, 2 skins (U.S.N.M.).YUKON: Yukon River, Caribou Crossing, 1 skin (U.S.N.M.) : Yukon River,50 miles below Fort Selkirk, 1 skin (U.S.N.M.).Remarhs.?Among the smaller brownish bats of eastern NorthAmerica Myotls lucifugus lucifugus may usually be at once recog-nized, when in fresh adult pelage, by the long glossy tips to the hairsof the back taken in connection with the presence of the dark shoulderspot and the faintly yellowish belly. Variations in the intensityof the coppery or bronzy tint are found among specimens from anylocality in various parts of the eastern United States; but west ofthe Mississippi, or roughly, west of the ninetieth meridian, the bronzyextreme is almost wholly lacking, the more brassy or " ochraceous-tawny " prevails, and the less intensely colored individuals usuallyare dull whitish below lacking the buffy tone.The typical subspecies has the widest range of any North Ameri-can form of Myotis and it probably penetrates farther north thanany other American bat. Here the limits of its distribution seemto correspond roughly with those of the coniferous forest, fromsoutheastern Labrador westward to James Bay, and thence northwest-ward in the interior along the edge of the Barren Grounds nearly tothe Arctic Ocean (50 miles from the mouth of the CoppermineRiver), finally reaching the Yukon Basin and the Pacific coast ofthe Alaskan Peninsula (Kodiak Island). It intergrades with thedarker race alascensis on the coast of southern Alaska and in BritishColumbia. The western border of its range in Canada probably runseast of the Eocky Mountains to Saskatchewan and Manitoba, thenceacross the United States through the Dakotas to central Colorado.In the dry interior of the northern United States, as in Colorado,Montana, and Wyoming, it gives place gradually to the paler sub-species carissima. Throughout much of eastern North AmericaMyotis lucifugus lucifugus seems to be the commonest member ofthe genus. MYOTIS LUCIFUGUS ALASCENSIS MillerVespertilio gnjphus lucifugus H. Allen, Monogr. Bats North Amer., BuU. U. S.Nat. Mus., No. 43 (1893), p. 78, March 14, 1894 (part, specimens fromWashington).
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Trouessakt, Catal. Mamm. viv. foss., p. 131, 1897 (part).Myotis lucifugus alascensis MiLLEai, North Amer. Fauna, No. 13, p. 63, Octo-ber 16, 1897. Elliot, Synops. Mamm. North Amer., Field Columb. Mus.,publ. 45, zool. ser., vol. 2, p. 402, March, 1901.
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Milleu and Rehn, Proc.Boston Soc. Nat. Hist., vol. 30, p. 256, December 27, 1901.?Trouessabt,Catal. Mamm. viv. foss., suppl., p. 92, 1904.
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Elliot, Check List Mamm. NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 479, 1905.
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Lyonand Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus.,No. 62, p. 270, January 28, 1909.?Seton, Life Hist. Northern Anim., p. 1148,1909.?Miller, List North Amer. Land Mamm. 1911, Bull. U, S. Nat. Mus.,No. 79, p. 55, December 31, 1912.?H. W. Gbinnell, Univ. California Publ.
48 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
Zool., vol. 17, p. 267, January 31, 1918.?J. Geinnexl, Univ. California Publ.Zool., vol. 21, p. 314, January 27, 1923.
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Miller, List North Amer. RecentMamm. 1923, Bull. U. S. Nat. Mus.. No. 128, p. 69, April 29, 1924.Myotis yumanensis saturatus Osgood, North Amer. Fauna, No. 21, p. 36, Sep-tember 26, 1901 (not of Miller, 1897; see H. W. Gbinnell, Univ. Calif.Publ. Zool., vol. 17, p. 432, April 25, 1918).Myotis pernox Hollister, Smithsonian Misc. Coll., vol. 56, No. 26, p. 4, December5, 1911 (Henry House, Alberta, Canada).
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Miller, List North Amer. LandMamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 56, December 31, 1912.?Eluot,Check List Mamm. North Amer., suppl., p. 155, 1917.
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Miller, List NorthAmer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 69, April29, 1924.Type locality.?Sitka, Alaska.Type specimen.?Adult female, in alcohol. No. 77416, UnitedStates National Museum (Biological Survey collection), collected atSitka, Alaska, August 5, 1895, by Clark P. Streator. Originalnumber 4754.Distribution.?Moist coastal region of western North America,from the archipelago of southern Alaska south through BritishColumbia and thence southeastward following the saturate area intonortheastern Washington (Blue Mountains) and extreme westernMontana, and again coastwise across western Washington and Oregonto the northwest coastal strip of California (Humboldt County).To the north and west of this area intergradation takes place withMyotis lucifugus lucifugus which occurs in nearly typical form onKodiak Island and in the interior of Alaska. In the dry lowlandsof eastern Washington and Oregon, that is, to the east of the coastranges that serve to withdraw much of the atmospheric moisture
-coming from the Pacific Ocean, the transition is fairly rapid to themuch j^aler M. lucifugus carissivia. A similar transition to carissimais seen from the Mount Shasta region of northern California, east-ivard and southward to the edge of the Great Basin.Diagnosis.?A dark, " saturate " race ; size as in typical Myotislucifugus.Description.?Similar to typical Myotis lucifugus but the generalcolor and especially the under-fur is so darkened that the wholeappearance is sooty with inconspicuous dark bronzy gloss on the back.The fur of the upperparts is everywhere blackish at the base becom-ing dark brown subterminally with a mahogany cast, nearly "lightseal brown" of Ridgway (1912). The extreme tips of the longhairs are glossy, reflecting the light in such a way as to give theimpression, especially when the hair is disarranged, of bronzy streaks.The shoulder spot is without these glossy tips but is not otherwisedifferent from the rest of the body in color. Below, the tips of thehairs are darker buff than in the typical form, with a distinct brown-ish cast especially on throat and chest; in some Alaskan specimens
AMERICAiSr BATS OF THE GENERA MYOTIS AND PIZONYX 49
almost uniform sooty brown. Individuals in imperfect pelage, inwhich the long glossy-tipped hairs are not wholly grown out, are avery dark sooty color, in contrast to the grayer tint of specimens ofM. lucifugus luoifugus in corresponding condition. The ears, nose,and membranes are very dark, almost black.Measurements.?For cranial measurements see table, page 60.Specimens examined.?Total number 95, from the followinglocalities
:
ALASKA: Admiralty Island, Mole Harbor, 1 skin (U.C.) ; BaranoffIsland. Red Bluff Bay, 10 skins (U.C.) ; Boca de Quadra, 1 .skin (U.C.) ;Ketchikan, 1 skin, 2 ale. (U.S.N.M.) ; Loring, 1 skin, 3 ale. (U.S.N.M.) ;Revillagigedo Island, 2 skins (U.C.) ; Sitka, 7 ale. including the type(U.S.N.M.).ALBERTA: Henry House, 2 skins (U.S.N.M.), not typical (type andparatype of pe'rnox).BRITISH COLUMBIA: Field, 1 skin (U.S.N.M.); Okanagan, 1 skin(M.C.Z.) ; Queen Charlotte Islands., 4 ale. (U.S.N.M.), Skidegate, 4skins, 1 ale. (U.S.N.M.) ; Sicamous, 1 ale. (A.N.S.P.) ; Vernon, 1 skin(M.C.Z.)
.
CALIFORNIA: Castle Lake, Siskiyou County, 1 skin (U.C), not typical;Eureka, Humboldt County, 1 skin (U.C).IDAHO: Felton Mills, 1 skin (Chas. R. Conner Museum).MONTANA: Columbia, 1 ale. (U.S.N.M.); Corvallis, 4 skins (U.C),.5 skins (U.S.N.M.) and 1 skin not typical (U.S.N.M.).OREGON: Bend, 3 skins (U.S.N.M.), not typical; Blue River, 1 skin(U.S.N.M.), not typical; Cornucopia, 1 skin (U.S.N.M.); East PineCreek, 1 skin (U.S.N.M.), not typical; Lapine, 1 skin (U.S.N.M.);McKenzie Bridge, 2 skins (U.S.N.M.) ; Mohler, Tillamook County, 1skin (U.C).WASHINGTON: Boulder Lake, 1 skin (F.M.) ; Butte Creek, Blue Moun-tains, Columbia County, 1 skin (U.C.) ; Carson, Skamania County, 4skins (U.S.N.M.) ; Chilliwack River, Whatcom County, 3 ekins(U.S.N.M.) ; Godman Springs, 5 skins (S. H. Lyman) ; Hannegan Pass,Whatcom County, 1 skin (U.S.N.M.) ; Husum, Klickitat County, 1 skin(U.S.N.M.) ; Lyle, Klickitat County, 1 skin (U.S.N.M.) ; Mount Ranier,Pierce County, 5 skins (U.S.N.M.) ; Soleduc River, Olympic Mountains,1 skin (U.S.N.M.), 1 skin (F.M.) ; Trout Lake, Klickitat County, 1 skin(U.S.N.M.) ; Wenatchee Lake, Chelan County, 2 skins (U.S.N.M.) ; What-com Pass, Whatcom County, 1 skin (U.S.N.M.).RemMrhs.?There is apparently no difference in size betweenMyOtis Iwdfugus al-ascensis and the tjrpical race; the length of ear^noted in the original description as slightly greater, appears onfurther study to be the same in both forms. Occasional individualsrepresenting the extreme of size do, however, occur. Thus, twospecimens from Henry House, Alberta, are distinctly larger than theaverage; in color one is intermediate between typical lucifugv^s andalascensis, while the other can not be distinguished from richlycolored examples of the eastern subspecies. The darker specimen
50 BULLETIISr 144, UNITED STATES NATIONAL MUSEUM
served as the type of Myotis pemox Hollister. Until there is evi-dence that they represent a form with a definite range, it now seemsmost satisfactory to regard these specimens as large individuals ofthe present race, showing intergradation toward tnie Myotis luci-fugus. Two specimens taken in Bitter Root Valley, western Mon-tana (Corvallis), are similarly dark, while a third (No. 160181,U.S.N.M.), also from Corvallis, is a typical carissima in color. Theirsize is normal. Seven skins from Oregon (Bend, 3; Blue River, 1;Lapine, 1; McKenzie Bridge, 2) present some of the same peculiari-ties. Five of them are indistinguishable from true Myotis lucifugus^two (one from Lapine and one from McKenzie Bridge) are darker.None of them shows an increase in size. One skin from Donner, Calif.(No. 100377, U.S.N.M.), is like eastern lucifugus^ while two othershave the color characteristic of M. I. carissima. Whether the westernspecimens indistinguishable in color from true Myotis lucifugusshould be regarded as indicating that the range of the typical sub-species actually extends southward along the mountains between theareas occupied by the races carissima and alascensis^ or whether theyare best explained as intermediates between the two western racesis a question which can not now be answered. The second alterna-tive, however, appears to be the more probable.MYOTIS LUCIFUGUS CARISSIMA ThomasMyotis yumanensis saturatus Merkiam, North Amer. Fauna, No. 16, p. 89,October 28, 1899 (not of Miller, 1897).
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Stephens, California Mammals, p.267, 1906 (part).?J. Grinnell, Proc. California Acad. Sci., ser. 4, vol. 3,p. 277, August 28, 1913 (part).Ml/Otis yumanensis Rehn, Proc. Acad. Nat. Sci. Philadelphia, 1904, p. 590 (notof H. Allen, 1864).Myotis (Leuconoc) carissima Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol.13, p. 383. May, 1904.Myotis carissima Miller, List North Amer. Land Mamm. 1911. Bull. U. S. Nat.Mus., No. 79, p. 56. December 31, 1912.
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Elliot, Check-list Mamm. NorthAmer., suppl., p. 154, 1917.
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Miller, List North Amer. Recent Mamm. 1923,Bull. U. S. Nat. Mus., No. 128, p. 70, April 29, 1924.Myotis yumanensis altipetens H. W. Grinnell, Univ. California Publ. Zool.,vol. 17, p. 9, August 23, 1916 (Merced Lake, Yosemite Park, California).Myotis lucifugus canssi>tna Gary, North Amer. Fauna, No. 42, p. 43. 1917(Colorado).
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Bailey, North Amer. Fauna, No. 49 [December^ 1926], p.215, January 8, 1927 (North Dakota).Myotis lucifugus altipetens H. W. Grinne3.l, Univ. California Publ. Zool.. vol.17, p. 263, January 31, 1918.?J. Grinnell, Univ. California Publ. Zool.,vol. 21, p. 214, January 27, 1923.
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Miller, List North Amer. Recent Mamm.1923, Bull. U. S. Nat. Mus., No. 128, p. 70, April 29, 1924.?Grinnell andStorer, Anim. Life in the Yosemite, p. 55, 1924.Myotis albicinctus G. M. Allen, Jour. Mamm., vol. 1, p. 2, November 28, 1919(Mt. Whitney, California).
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Miller, List North Amer. Recent Mamm.1923, Bull. U. S. Nat. Mus., No. 128, p. 68, April 29. 1924.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 51Type locality,?Yellowstone Park, Wyo.Type specimen.?Adult female, in alcohol. No. 4.4.25.1, BritishMuseum (Natural History), collected at Yellowstone Lake, Yellow-stone National Park, Wyo., September, 1903, by J. ffolliott Darling.Dlstrihution.?Semi-arid portions of the western United Statesfrom southern and eastern Montana and parts of Wyoming west toeastern Oregon and the Sierras of California.Intergradation with typical Myotis lucifugus takes place in theDakotas, and with M. lucifugus alascensis in the Sierras of northernCalifornia and in the eastern half of Oregon and Washington. Inthe mountains of western Montana, two specimens taken at Corvallisare dark enough to be referable to M. I. alascensis^ yet a third is socontrastingly pale and with white-edged membranes as to be typicalof M. I. carissima. This individual had perhaps come from a lowerlevel or had wandered in from the south. Not infrequently specimensare found in eastern Oregon or northern California that might, with?qual propriety, be referred to either race, it becoming, therefore, amatter of individual judgment which name shall be given to them.Yet the transition is rather abrupt in most places. Montana speci-mens tend to have a faint trace of the bronze tint characterizing theeastern M. lucifugus lucifugus while those from the northern SierraNevada of California have a golden brown cast. As pointed out byMrs. Grinnell (1918) this bat is chiefly an inhabitant of the forestedareas at the higher levels in California, and has been taken at 11,000feet on Mount Whitney at the upper limit of trees. San BernardinoMountains, Calif., where several specimens were taken at about 7,000feet, represent the southern limit of its range as now known.Diagnosis.?Color paler than in Myotis lucifugus lucifugus., thegeneral hue of the upper parts golden instead of bronzy or olive,the belly cartridge buff rather than light chamois; membranes tend-ing to be pale-edged; ears dark; foot larger than in the typical race,the ratio of its length to that of tibia about 55.7.Description.?Entire dorsal surface a pale " cinnamon-buff," thehair of the back with long glossy tips which give a more or lessgolden sheen. Shoulder spot ochraceous tawny ; lower surface a palebuff, very nearly the cartridge buff of Ridgeway. The hairs bothabove and below have dark blackish-plumbeous bases except at theposterior border of the venter where they are white throughout.Ears, feet, and proximal portion of wing membranes black contrast-ing with the pallid pelage; or in some specimens becoming browner.The edges of the interfemoral membrane and parts of the posterior
?edge of the wing membrane may be whitish, sometimes forming adistinct border.Measurements.?The dimensions of the type are given b}' Thomasas: forearm 28 mm.; head and body 45; tail 36; ear 13 by 8; third
52 BULLETIN 144, UNITED STATES NATIONAL MUSEUMmetacarpal 33; first phalanx 11.5; second phalanx 10.5; third phalanx:7.2; tibia 16; foot with claws 11; calcar 16.For other measurements, see tables, pages 58 and 60.Specwiens examined.?Total number 225, from the following-localities :CALIFORNIA: Bear Lake, San Bernardino Mountains, 1 skin (U. C.) ;Bluff Lake, San Bernardino Mountains, 4 skins (U. C.) ; Castle Lake^Siskiyou County, 1 skin (U. C.) ; Donner, 3 skins (U.S.N.M.), not typical;:Feather River, north fork, 3 ale. (U.S.N.M.) ; Gilmore Lake, 1 skin?(U. C.) ; Ice Caves, 6 miles west of Tule Lake, 2 skins (U.S.N.M.), nottypical; Independence Lake, 2 skins (U. C.) ; Keeler, Inyo County, 7 ale.(U.S.N.M.) ; Lincoln Creek, Sierra County, 7,000 ft., 1 skin (U.S.N.M.) ;Lone Pine, Inyo County, 1 skin (U.S.N.M.) ; Mammoth, Mono County,.1 skin (M. C. Z.) ; Mount Shasta, 4 skins (U.S.N.M.), 3 skins (U. C.),.approaching alascensis; Mount Tallac, 2 skins (U. C), 1 skia(A. N. S. P.), 1 skin (M. C. Z.) ; Mount Whitney, 1 skin (M. C. Z.),.type of albicinctus ; Prattville, 1 skin (U.S.N.M.) ; Santa Ana River,San Bernardino Mountains. 1 skin (U. C.) ; Susanville, 1 skin (U.S.N.M.) ;Warren Peak, Warner Mountains, 3 skins (U. C.) ; Yosemite Park, Sskins (U. C), including type of altipetens.COLORADO: Conejos River. 1 skin (U.S.N.M.); Meeker, 2 skins(U.S.N.M.); Steamboat Springs, 1 skin (U.S.N.M.).IDAHO: Birch Creek, 1 ale. (U.S.N.M.).MONTANA: Big Belt Mountains, near Fort Logan, 2 skins (U.S.N.M.);Big Timber, 7 skins (U.S.N.M.) ; Corvallis, 1 skin (U. C), not typical;Crow Agency, 4 skins (U.S.N.M.) ; Cut Bank, 25 miles N. W. of (TetonCounty), 37 ale. (U.S.N.M.) ; Glasgow, 45 skins (U.S.N.M.), not typicaLNEVADA: Little High Rock Canyon, Washoe County, 2 skins (U. C.) ;Pyramid Lake, 1 ale. (U.S.N.M.) ; Ruby Valley, 1 skin (U.S.N.M.).NEW MEXICO: Sierra Grande, Union County, 1 skull only (U.S.N.M.)(determination not positive).OREGON: Baker County, 1 ale. (U.S.N.M.) ; Klamath Falls, 9 skins (U. C),a little darker than typical; Malheur Lake. 1 skin (U.S.N.M.) ; Narrows,1 skin (M. C. Z.) ; Paulina Lake, 1 skin (U.S.N.M.) ; Riverside, 1 skin(U.S.N.M.) ; Sheaville, 1 skin (U.S.N.M.) ; Steen Mountains, 3 skins(U.S.N.M.), nearing lucifugus lucifugus; Voltage, 3 skins (U.S.N.M.);Watson. 1 skin (U.S.N.M.).UTAH: Bear River, mouth of, 2 skins (young), 18 ale. (U.S.N.M.) ; Bounti-ful, 1 ale. (U.S.N.M.) ; Provo City, 1 ale. (U.S.N.M.).WASHINGTON: Stehekin, Okanogan County, 5 skins (U.S.N.M.).WYOMING: Fremont Peak, 1 ale. (U.S.N.M.); Geyser Basin, 1 ale.(U.S.N.M.) ; Sand Creek, 1 skin (U.S.N.M.) ; Yellow.stone Park, Mam-mouth Hot Springs, 17 skins, 2 ale. (U.S.N.M.).Remarks.?The United States National Museum contains an excel-lent series of Myotis lucifugus cainssivia collected at the type locality,,where it is evidently common. Three of these, an alcoholic (No.153635) and two skins with skulls (Nos. 208562 and 209858), weresent to Mr. Oldfield Thomas, who has very kindly compared themwith the type in the British Museum, and pronounces them undoubt-edly the same. Since the type is in alcohol, its general dark appear-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 53
:ance gives little clue to the natural color. Thomas says : " Colorabove and below (in spirit) uniform smoky blackish, the tips of thehairs indistinctly buffy or pale brown." This account applies suffi-ciently well to the appearance of a dull colored individual seen inalcohol, but it gives no idea of the color characteristic of the race.For this reason, Mrs. H. W. Grinnell was misled into supposing thatthe pale California bat was different from Myotis carismna. Shetherefore renamed it altipetens on the basis of a specimen fromYosemite Park, California. Specimens frequently occur with a well-marked though narrow white edge along the posterior margin of thewing membrane, and the terminal part of the interfemoral membranemay also be more or less whitish. It was an extreme example of thisstyle that served for the type of M. alMciiwfus G. M. Allen, while thetype of altipetens H. W. Grinnell is similarly marked. Both areuncjuestionably to be referred to the present subspecies.The specimen No. 32029, from Geyser Basin, Wyoming, alreadymentioned (p. 8) on account of its abnormal teeth {p7n^ right absent,p^n - and />m ^ left coalesced) is further peculiar in the general slen-derness of its skull. It appears, however, to be referable to the pres-ent form. MTOTIS LUCIFUGUS PHASMA. new subspeciesMyotis yumanensis Caby, North Amer. Fauna, No. 33, p. 207, 1911 (not of H,Allen, 1864).?Wabben, Mammals of Colorado, p. 273, 1912 (part).Type.?Adult female, skin and skull No. 148159 United StatesNational Museum (Biological Survey collection), collected at SnakeRiver, south of Sunny Peak, Routt County, Colorado, August 28,1906, by Merritt Cary. Original number, 792.Distribution.?^Arid portions of the Great Basin, Limits of rangenot known.Presumably this race will be found to occupy all of the more aridparts of the Great Basin, where no doubt it will prove to be of localoccurrence, depending on proper conditions. The Colorado localitiesprobably represent the extreme northeastern limits of its range inthe desert valleys of western Colorado (Upper Sonoran Zone). Thethree secured by Cary were captured " in deserted ranch buildingsafter nightfall, where they were not at all common, being greatly out-numbered by M. evotis.''^ Its presence in southern Utah can not bedoubted, but no specimens are at hand. Two immature bats from theBear River in the northern part of the state, though still in darkpelage, seem better referred to carissinia. A single specimen fromInyo County, California, though in poor pelage, is, however, appar-ently phasma.) thus carrying the range across to the Californian des-erts. The type-specimen of M. alhicinctus G. M. Allen, from Mount
54 BULLETIN 144, UNITED STATES NATIONAL MUSEUMWhitney, California, though very pale in general coloration, is refer-able to ca7issivm.Diagnosis.?In general like Myotis lucifugus carissima but color offur even more pallid and ears light brown instead of blackish.Descrij)tion.?Very pallid, the general effect above a uniform palebuff with a pronounced pinkish cast and a faint golden gloss ; belowbuffy white. In detail: the fur of the body is everywhere darkplumbeous at the base, that of the back with a narrow ring of
" ochraceous-buff " and a " light-buff " glossy tip. The ochraceous-buff ring showing through the paler glossy tips produces the pinkishtint. The edges of the ears and their bases exteriorly are clothedwith short hair, pale " ochraceous-buff " throughout. Short stiffhairs about the upper lip between the eye and the muzzle are duskybrown. The usual darker shoulder-spot is hardly apparent, but ismarked by an area of pale cinnamon. Ears and membranes palebrownish (not blackish as in M. lucifugus carissitna) , the edge ofthe interfemoral membrane, especially near the tip of the tail, andthe posterior rim of the wing membranes whitish.MeaMiretnents.?For measurements see tables, pages 58 and 61.Specimens exatiuTied.?In all 4, from the following localities:CALIFORNIA: Argus Mountains, Inyo County, 1 skin (F.M.).COLORADO: Lily, Routt County, 1 skin (U.S.N.M.) ; Snake River, RouttCounty, 2 skins (U.S.N.M.).Remarks.?The desert race phasma is the opposite extreme fromthe saturate Myotis lucifugus alascensis of the moist Pacific coast.Between the two is interposed the form carissima which, althoughintermediate, is so constant in color over a wide area as to constitutea subspecies with a definite geographic range. So closely doesMyotis lucifugus phnsma parallel typical M. yumanensis in color thatGary (1911) very naturally considered his Colorado specimens asbelonging to that species. They are readily separated however, bythe longer glossy-tipped fur above, the longer forearm and wing-bones, and by the larger skull with its less elevated brain case,characters diagnostic of Myotis lucifugus.MYOTIS LUCIFUGUS FORTIDENS, new subspeciesType.?^Adult female (in alcohol) No. 88.8.8.18, British Museum(Natural History). Collected at Teapa, Tabasco, Mexico, by H. H.Smith, January 5, 1888. Presented by Messrs. Salvin and Godman.Distribution.?From southern Mexico north to extreme westernTexas.At present only two specimens are known which can be referred tothis race. The exact limits of the range are therefore problematical.Diagnosis.?Size and color about as in Myotis lucifugus lucifugus;skull with sagittal crest probably more frequently developed than in
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 55the other races (present in both of the known specimens) ; molari-form teeth, both above and below, enlarged, so that the distancebetween the canine and pm^ is reduced, and a noticeable crowdingof the two small premolars takes place.Color.?The type, after long immersion in alcohol shows suchobvious indications of fading that its color can not be regarded asnormal. It resembles that of similarly faded old specimens! ofMyotis lucifugus lucifugus. The second specimen, a skin from FortHancock, Tex., is not distinguishable from the less glossy individualsof the northeastern race.Skull.?The skull does not differ from that of Myotis lucifuguslucifugus except that the presence of the sagittal crest in the twoknown specimens is an indication that this ridge is more usually welldeveloped than in the northern races.Teeth.?The teeth resemble those of the typical race in form, butthe size of the molars and large premolar is obviously increased. Inthe type specimen the crown of m ^ measures 1.40 by 1.85 mm. ; inthe Fort Hancock specimen it is slightly larger, 1.45 by 1.85 mm.The corresponding measurements in Myotis lucifugus lucifugus andM. I. carissima are usually 1.20 to 1.35 by 1.50 to l.TO. While theincrease in size of the individual cheek teeth does not produce alengthening of the entire tooth row as measured from front of canineto back of last molar (see tables of measurements), it increases thedistance from front of pm * to back of m ^ as compared with the otherraces of M. lucifugus., with the result that the space occupied by thetwo small premolars between the canine and the large premolar isshortened. In this reduced space only one small tooth is visiblewhen the maxillary series is viewed from the side. This crowdingis so great in the type specimen that the posterior small tooth hasbeen eliminated.Measu/retiients.?For measurements see tables, pages 58 and 61.Specimens examined.?Total number 2, from the following locali-ties:TEXAS: Fort Hancock, El Paso County, 1 &kin (U.S.N.M.).TABASCO: Teapa, 1 ale. (B.M.).Remarks.?Although the material is unsatisfactory, the two speci-mens indicate rather clearly the existence of a Mexican race ofMyotis lucifugus. This form will probably be found to occur alongthe border of the United States south of the region occupied by thelight-colored, small-toothed subspecies ca7^sirna and phasma.The Fort Hancock specimen agrees perfectly with the type in thelarge size of the molariform teeth as compared with those of theother races of M. lucifugus. Its upper premolars are abnormal in away that we have never seen in any other bat. The large tooth is
56 BULLETIN 144, UNITED STATES NATIONAL MUSEUMcompletely separated from the first molar by a noticeable space. Itsmain axis is tilted forward so that the line of the posterior cuttingedge would, if continued forward, strike the shaft of the canine nearthe tip. The two small premolars are present, but the second isunusually minute and is crowded beneath the inner anterior borderof the large premolar. To the anterior border of the large premolaris soldered a small tooth essentially like a normal small premolar andnot in the least resembling a milk tooth, its tip level with the tip ofthe main cusp of the large premolar and functioning with it as shownby the worn areas on the two summits. These abnormal conditionsare fully developed and symmetrical on the two sides.External measurements of Myotis ludfugus
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 57External measurements of Myotis lucifugus?Continued
Locality
Myotis lucifuguslucifugus?Con.
Number
Illinois:
58 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis luelfugxis?Continued
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 59Cranial measurements of Myotis ludfugus
Locality
Myotis lucifaguslucifugusYukon;Below Fort Sel-kirkCaribou CrossingAlaska:Kodiak IslapdDoDoDoDoOntario:Waterloo County.DoDoManitoba: Hartney..Quebec: James Bay.Nova Scotia:HalifaxDo..DoDoDoNew York:Locust GroveCatsklU Moun-tains ,Howe's CaveDoMinnesota:Elk RiverDoDoDo...Do ,Indiana:Mitchell ,DoDoDo..Do..DoDoDoDoDoBloomingtonDoWest Virginia:White SulphurSprings. ,Do....DoDo....Do.... ,Do..DoDoDoVirginia:MiddletonDoRlvertonGeorgia: YoungHarrisNorth Dakota:TownerDoDo...DoDoDo.-DoDo..Do...
Number
99364 U.S.N.M.9936391563015659156462752627537309 F.M.8600860132969 U.C.38595 U.S.N.M.15450515450915451215451616451918735783000187308187333187387187389187391187392236137153629153622153628153620153621153624153625153626153627153628153633153632
8391783918839205612 F.M.5613 F.M.5614561556165617243086 U.S.N.M.24308787447159411 U.S.N.M.207725 U.S.N.M.207740207739207736207726207729207730207733207718
60 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis lucifugus?Continued
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 61Cranial measurements of Myotis hicifugus?Continued
Locality
Myotis Incifngnscarissima?Con.Washington:StehekinDo-DoDoOregon
:
Paulina LakeRiversideSteen Mountains.DoCalifornia:Qilmore LakeYosemite Park...DoWarner Moun-tainsDo..DonnerDo...Ice CavesDo...PrattvilleLincoln CreekNear Susanville..Myotis IncifugnsphasmaColorado:Snake RiverDo..Lily...California: ArgusMountains.Myotis lucifugusfortidensTabasco: TeapaTexas: Fort Hancock.
Number
22989622989723014123014220490321390921609421609512036 U. C.23036 U. C.3 23034 U. C.11351 U. C.11352 U. C.100377 U. S. N. M.10050397731977329809410037898093
148158< 14815914816713295 F.M.
8. 8. 8. 18 B. M.36121
14.014.414.314.514.814.414.414.314.614.815.014.614.914.214.514.714.314.614.414.2
15.014.914.614.5
15.014.8
^J3
13.113.013.613.713.513.613.513.313.913.814.113.914.013.213.613.913.513.513.813.5
O ?
13.814.0
9.09.09.0
14. 1 8.
9
14. 2 9.13.913.7
9.69.4
O 1.1ut toas p
3.83.84.04.04.03.93.93.84.04.14.04.03.94.04.03.73.73.83.83.8
3.63.83.63.7
3.83.8
7.37.27.27.27.67.27.17.27.57.57.47.17.47.47.37.27.57.27.27.2
7.17.17.07.4
7.47.0
5.25.05.25.25.25.05.14.95.25.15.65.14.95.05.05.05.05.15.3
5.25.0
10.210.010.410.510.810.10.0
10.610.511.010.710.610.610.310.510.610.410.810.3
11.011.110.810.3
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5.05.05.45.35.15.25.15.15.35.45.55.35.35.25.25.25.25.25.35.2
5.7S.3
5.45.5
5.55.65.85.85.45.65.65.75.66.05.95.86.05.65.55.55.75.55.8
5.6.05.8
5.86.2
3^
5.55.35.75.75.65.55.55.65.85.65.95.65.65.65.76.75.85.65.85.6
6.36.05.95.5
5.86.0
' Type of M. yumanensis altipetens H. W. Orinnell. Type.MYOTIS YUMANENSIS (H. Allen)(Synonymy under subspecies)
Disf7'i'bution.?Western North America from southern BritishColumbia to the southern part of the Mexican plateau (at least asfar as the State of Michoacan) ; eastward over the arid region of theUnited States from western Texas northward, including probablythe whole of the Great Basin, to western Montana.Diagnosis.?In general like Myotis lucifugus but smaller and withrelatively somewhat longer tail (average ratio of tail to head andbody in 8 specimens from Yuma, Ariz., 81.1; in 10 from Old FortI'ejon, Calif., 81.2; in 10 from Montana, 87.2; and in 10 from SanLuis Potosi, 83.4) ; forearm about 34 mm. (32 to 37 mm.) ; greatestlength of skull ranging from 13.0 to 14 2; maxillary tooth row rang-
62 BULLETIN 144, UNITED STATES NATIONAL MUSEUMing from 4.6 to 5.2; lower tooth row (excluding incisors) usually less ,than 5.5 mm. (5.0 to 5.6 ram.). Foot slightly less enlarged thanin M. Itwifuffus, its ratio to tibia usually ranging from about 50 to
Map 2.
?
Distribution of Myotis yumanensis : 1, M. yumanensis ydmanhnsis ;2, M. yumanensis sociabilis ; 3, M. yumanensis satubatds ; 4, M. yumanensisLUTOSUSabout 54. Skull with abruptly rising forehead. Fur of back dullas compared with that of M. lueifugics, the longer hairs lacking con-spicuous burnished tips.
AMEKICAN BATS OF THE GENERA MYOTIS AND PIZONYX 63Ears.?The ears are of moderate length; when laid forward theyreach to the nostril (alcoholic specimens). When the ear is helderect, the lower third of its external border forms a strongly juttingshoulder, above which the margin is very slightly convex, then veryslightly concave to the bluntly rounded tip. The result is that theupper two-thirds of the ear conch appears to be narrowed and witha nearly straight posterior margin. In Myotis ludfugus the basalshoulder is much less pronounced and the conch itself is wider sothat the outer edge is more nearly straight from base to tip, the upperpart of the ear is not so narrowed, and its general appearance is lessscimitar-shaped.The tragus differs from that of Myotis ludfugus in the size andform of the basal lobe; this lobe is larger in proportion and nearlysemicircular in outline; the tragus is about one-half the height ofthe ear from meatus (7:13.5 mm.). The anterior edge is nearlystraight, the posterior slightly convex above the base, then taperingin the upper third to the bluntly rounded tip. The upper third ofthe posterior edge is often slightly crenulate.Wing and Tneyiibranes.?The Aving membrane arises from the tarsusjust proximal to the bases of the toes. The metacarpals are distinctlygraduated, the third longest, the fourth slightly shorter, the fifthagain shorter (about 2 mm. less than the third). When folded, thethird metacarpal falls from 1 to 2 mm. short of the elbow (as com-pared with 2-5 mm. in M. luoifugus) . Taking the third fmger as 100,the fourth is 85, the fifth 79 (56:48:44.5 mm.). The first phalanxof the fourth finger is about 1 mm. shorter than the second (8:9 mm.) ;but in the fifth finger the opposite relation obtains, for the first is avery little longer than the second (7.5 : 7) , or they are practically equal(7.5:7.5). The minute terminal vertebra of the tail is the only onefree from the membrane. Under a lens may be seen numerous shortstiff hairs springing in several rows from close along the free edgeof the interfemoral membrane ; they do not, however, form a fringe.Foot.?The foot is proportionally stout and long, the ratio of itslength to that of tibia ranging from about 50 to about 54. The inner-most toe is slightly but evidently shorter than the others, the tip ofits claw reaching only a little beyond the base of the claw of digit 2.The long calcar extends three-fifths of the distance from the heel tothe tail, and ends in a minute lobule. It is quite without projectingkeel.Fur.?The pelage is full and of medium length, the hairs about5 mm. long on the lower back, bicolored ; dark bases are evident every-where except on the lower abdomen. The fur extends out on to theinterfemoral membrane slightly beyond a line joining the knees,above and below. Underneath, the wings are thinly haired nearly
64 BULLETIN 144, UNITED STATES NATIONAL MUSEUMthree-fourths the distance to the elbow. The paler tips of the dorsalhairs are shorter and less glossy than those of Myotis lucifugus^ withthe result that M. yvmumensis usually appears very dull colored incontrast to M. lucifugus with its deeper, more glossy fur. It is ordi-narily possible to separate out the two species in a mixed series byrecourse to this character alone.Skull.?The skull, as compared with that of Myotis lucifugus, isnot only smaller in size (total length 13,2 to 14.2 mm. instead of 14.0to 15.8 mm.) but different in proportions. When the skull is viewedin lateral profile the alveolar line appears to rise from behind for-ward at a more abrupt angle, the forehead seems more pronouncedand higher, so that the rostrum is separated from the brain case by amore obvious concavity, and the occiput is relatively more elevated.The entire brain case as seen from above has a more distinctly globu-lar aspect. The temporal ridges, even in adults, are barely discern-ible, but sometimes they unite to form a low crest. This crest is sorarely developed that we have found it in only 9 of 70 adult skulls.Even when obvious the crest is noticeably flat-topped and muchbroader than high, while that of M. lucifugus, posteriorly at least,is of the usual narrowly ridgelike form if developed at all. Theposterior outline of the occiput when viewed from above is distinctlymore abrupt in its curvature than that of M. lucifugus. All thesedifferences are, in such small objects, very minute and at first some-times difficult to appreciate, but they are, nevertheless, striking whencarefully studied.Teeth.?The teeth resemble in general those of Myotis lucifugus.Maxillary tooth rows rather narrow in proportion to the width ofpalate between them, in this character agreeing perfectly with thoseof M. lucifugus. Upper molars with the full complement of second-ary ridges and a well-developed protoconule. The cingulum at itsmaximum development resembles that of M. lucifugus lucifugus, butmany individuals occur in which it is greatly reduced, frequently tosuch a degree as to be nearly or quite absent from the entire lingualborder of the crowns. The small premolar teeth are remarkably uni-form in their relations. In the upper jaw p^ and p^ are both drawninward a little from the tooth row so that the anterior small tooth isoverlapped by the cingulum of the canine and the posterior is over-lapped to a slightly less degree by the cingulum of p^. In cross-sec-tion p^ is only a little less than p^ but in profile it is much lower. Thecusps of both are broadly triangular and blunt in side view, that ofthe smaller tooth {p^) reaching a level not exceeding half the heightof the anterior tooth. In Myotis lucifugus they are usually moreslender and lancet-shaped.Remarks.?This species is structurally much like Myotis lucifugusbut its size is uniformly less and the foot is, on the average, less
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 65
enlarged. The pelage is shorter and not so glossy (on account of theshorter glistening tips of the long hairs) and the skull is less flat-tened and with a slightly more upturned muzzle. The ears arethinner and narrower and usually less heavily pigmented. All thesedifferences are, of course, best appreciable on direct comparison.Very little is recorded concerning the natural history of Myotisyvmmnensis. It is probably less of a forest haunter than Myotislucifuffus, preferring more open country with scattered tree growth,though representatives of the two species occur together on thePacific coast, as well as in Montana and elsewhere. Mrs. Grinnell(1918) records a large breeding colony that frequented the old ruinedbuildings of Fort Tejon, Kern County, Calif. The adult males ap-parently leave the colony when the young are born, and duringsummer they may wander up to a considerable altitude in the openwoods of the dry mountain country. In the San Bernardino Moun-tains, Calif., Mrs. Grinnell notes the capture of a male at 8,500 feetelevation, and another was taken at 11,000 feet on the east slope ofMount Whitney, Calif., along a small watercourse at the limit of treegrowth (G. M. Allen, 1919). A. B. Howell (1920) recounts the dis-cover}^ of a large colony that frequented an abandoned mine tunnelin California. A single young is the rule, and the breeding season isearly, at least in Texas, where on May 26, at Del Rio, Gaut found acolony with young clinging to the old ones (Bailey, 1905).Of the geographical races into which Myotis yimumeTisis is dividedthe most widespread is a dull-brownish form occupying a consider-able area of the interior from at least western Montana south throughcentral California. In the humid coastal area from the central partof California into Washington and southern British Columbia thiscolor becomes considerably darker, and again, in the desert countryof southeastern California, Arizona, and parts of Texas it changesto an extreme of pallor. In the southern part of the Mexican high-lands, again, there is a dark-brown race, the limits of whose distribu-tion are yet undetermined. Throughout this range, however, thereis very little variation in size, a character which, together with itsdull, short pelage and narrow-tipped ears, will usually serve todistinguish Myotis yumanensis from Myotis lucifugus where the twospecies occur together.MYOTIS YUMANENSIS YUMANENSIS (H. Allen)Vespertilio yumanensis H. Allen, Monogr. Bats North Amer., SmithsonianMisc. Coll., No. 165, p. 58, fig. 54-56, June, 1864.?Dobson, Catal. ChiropteraBrit. Mus., pp. 328, 329, 1878.?H. Allen, Monogr. Bats North Amer., BuU.U. S. Nat. Mus., No. 43, (1893), p. 72, March 14, 1894.Vespertilio obscurus H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 281(Lower California).
?
Milleb, North Amer. Fauna, No. 13, p. 69, October 16,
66 BULLETIN 144, UNITED STATES NATIONAL MUSEUM1897 (as synonym of M. californicus)
.
?Lyon and Osgood, List Type-Sp.Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 272, January 28,1909.?Goldman, Proc. Biol. Soc. Washington, vol. 27, p. 102, May 11, 1914.Vespertilio macropus H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 288,not of Gould, 1854 (Fort Mohave, Colorado River, Arizona).
?
Lyon andOsgood, List Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat Mus., No. 62,p. 271, January 28, 1909.Vespertilio nitidus (pedomorphic variety) H. Allen, Monogr. Bats North Amer.,Bull. U. S. Nat. Mus., No. 43 (1893), pp. 72, 73, March 14, 1894.Vespertilio albescens H. Allen, Monogr. Bats North Amer., Bull. U. S. Nat. Mus.,No. 43 (1893), p. 87, March 14, 1894 (part, not of Geofifroy, 1805).?Tboues-SAET, Catal. Mamm. viv. foss., p. 132, 1897 (part).Vespertilio nitidus macropus H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 100, March 14, 1894.?Troxjessaet, Catal.Mamm. viv. foss., p. 130, 1897.Mi/Otis yumanensis Millee, North Amer. Fauna, No. 13, p. 66, October 16, 1897(part).
?
Elliot, Synops. Mamm. North Amer., Field Columb. Mus., publ.45, zool. ser., vol. 2, p. 403, March, 1901 (part) ; List Land and Sea Mamm.North Amer., Field Columb. Mus., publ. 57, zool. ser., vol. 2, p. 517, June,1901 (part).?MiLLEK and Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p,256, December 27, 1901 (part).
?
Trouessart, Catal. Mamm. viv. foss.,suppl., p. 92, 1904.
?
Elliot, Field Columb. Mus., publ. 91, zool. ser., vol. 3,p. 318, March, 1904 (part) ; Land and Sea Mamm. Middle Amer., FieldColumb. Mus., publ. 95, zool. ser., vol. 4, pt. 2, p. 576, 1904 (part) ; CheekList Mamm. North Amer., Field Columb. Mus.. publ. 105, zool. ser., vol. 6, p.475, 1905 (part).?Stephens, California Mammals, p. 267, 1906 (part).?Elliot, Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115,zool. ser., vol. 8, p. 501, 1907 (part).
?
Lyon and Osgood, List Type-Sp.Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 291, January 28,1909.
?
Warren, Mammals of Colorado, p. 273, 1910.
?
Bailey, North Amer.Fauna, No. 35, pp. 19, 33, September 5, 1913.?A. B. Howell, Journ. Mamm.,vol. 1, p. 173, August 24, 1920.?J. Grinnell, Univ. California Publ. Zool.,vol. 21, p. 314, January 27, 1923.Myotis yumanensis yumanensis Miller, List North Amer. Land Mamm. 1911,Bull. U. S. Nat. Mus., No. 79, p. 56, December 31. 1912 (part).?J. Grin-nell, Proc. California Acad. Sci., ser. 4, vol. 3, p. 276, August 28, 1913.?H.W. Grinnell, Univ. California Publ. Zool., vol. 17, p. 273, January 31,1918.?Miller, List North Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus.,No. 128, p. 69, April 29, 1924.Myotis californicus durangce J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 19,p. 612, November 12, 1903 (San Gabriel, Rio Sestin, Durango, Mexico) ;Bull. Amer. Mus. Nat. Hist., vol. 20, p. 210, May 28, 1904.?Elliot, Land andSea Mamm. Middle Amer., Field Columb. Mus.. publ. 95, zool. ser., vol.4, pt. 2, p. 579, 1904 ; Check List Mamm. North Amer., Field Columb. Mus.,publ. 105, zool. ser., vol. 6, p. 478, 1905.
?
Mieler, List North Amer. LandMamm. 1911. Bull. U. S. Nat. Mus., No. 79, p. 56, December 31, 1912;List North Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 71,April 29, 1924.Myotis californicus difrangoa; Trouessart, Catal. Mam. viv. foss., suppl., p. 93,1904 (variant).Myotis yumanensis sociaMlis G. M. Allen, Journ. Mamm., vol. 1, p. 1, Novem-ber 28, 1919 (not of H. W. Grinnell).Myotis yumaensis yumaensis Strecker, Check-List Mamm. Texas, The BaylorBulletin, Baylor University, Waco, Texas, vol. 29, No. 3, p. 9, August. 1926.
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 67Type locality.?Old Fort Yuma, Imperial County, Calif., on rightbank of Colorado River, opposite the present town of Yuma, Ariz.Type specimen.?In his original description, H. Allen (1864) men-tions by number, four specimens (5387, 6019-21, U.S.N.M.) in alco-hol, all from Fort Yuma, Imperial County, Calif., collected by Maj.Gen. George H. Thomas. Lyon and Osgood (p. 291) were unable totrace these specimens 20 years ago, and we have fared no better.Distribution.?Interior desert country of the Great Basin, fromthe eastern edge of the Sierra Nevada of California eastward intowestern Texas, and south into Lower California and in Mexico tothe State of Durango.Northward range yet to be determined; but the pale desert raceevidently merges into the darker M. yuTnanensis sociabilis somewherein Utah and Nevada. Skins are lacking to determine the exactstatus of the race inhabiting southern Lower California. The sup-posed occurrence of Myotis yuiiianensis yunianensis in Colorado(Gary, 1911) turns out to rest on a misidentification of M. lucifugusphasma.., though it is not unlikely that the smaller animal will eventu-ally be found in the western part of that State.Diagnosis.?Palest subspecies of Myotis yvmunnen^is; butfy above,nearly white beneath.Description.?Color above very pale buffy, nearly pinkish buff orpale, dull, cream buff (RidgAvay, 1912), the hairs dark, near fuscous,for the basal half. Below, pale cartridge buff, the basal half of thehairs fuscous except in the posterior region where they are whitishthroughout. The membranes are pale brownish, translucent, usuallyedged with white along the interfemoral border and at the tip ofthe tail as well as on the edges above the ankles. Specimens inimmature pelage are less pallid, nearly cinnamon buff above.Measuretnents.?For measurements see tables, pages 73, 75.SpeciTnens exa/mined.?Total number 354, from the followinglocalities
:
ARIZONA: Ehrenberg, 13 ale. (U.S.N.M.) ; Fort Mohave. 1 skin (U.S.N.M.),type of macropus; Fort Verde, .5 skins (A.M.N.H.) ; Gila Bend, 3 skins(U.S.N.M.) ; Huachuca Mountains, Cochise County, 1 skull (F.M.) ;Jacob's Pool, 1 skin (U.S.N.M.) ; Rice (San Carlos Indian Reservation),2 skins, 2 ale. (U.S.N.M.), nearly typical; Safford, 1 ale. (U.S.N.M.);Tucson, 1 skin (U.S.N.M.) ; Yuma, 9 ale. (U.S.N.M.).CALIFORNIA: Argus Mountains, Inyo County, 1 skin (F.M.) ; CarrellCreek, Inyo County, 1 skin (U.C.) ; Fort Yuma, Imperial County, 4skins, 2 ale. (U.S.N.M.) ; Lone Pine, Inyo County, 1 skin (M.C.Z.), 2ale. (U.S.N.M.) ; Mount Whitney, Tulare County, 11,000 ft., 1 skin(M.C.Z.),1 ale. (U.S.N.M.) ; Owens Lake, Inyo County, 4 ale. (U.S.N.M.) ;Potholes, Imperial County, 1 skin (U.C), 1 ale, 2 skins (M.C.Z.).
68 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCOLORADO: Colorado River, Horse Shoe Bend, 1 ale. (U.S.N.M.).DURANGO: Arroyo de Buoy, 1 skin (A.M.N.H.) ; Rio Sestin, 6 skins,including type of durangoe (A.M.N.H.) ; San Gabriel, 5 ale. (A.M.N.H. >.LOWER CALIFORNIA: [?Cape St. Lucas], 2 ale. (U.S.N.M.), includingtype and paratype of oJ)scurus; Rancho San Antonio (west base of SanPedro Martir Mountains), 8 skins, 10 ale. (U.S.N.M.) ; Rio Pescadero,1 skin (U.S.N.M.).NEW MEXICO: Clayton, 1 skin not typical (U.S.N.M.) ; Rinconada, 1 ale.(U.S.N.M.).NEVADA: Colorado River, 1 ale. (U.S.N.M.) ; Pyramid Lake, 24 skins, 198ale. (U.S.N.M.).SONORA: Colonia Lerdo, 6 skins, 18 ale. (U.S.N.M.) ; no exact locality, 1ale. (U.S.N.M.).TEXAS: Del Rio, Valverde County, 9 skins (U.S.N.M.); Marathon, 1skin (U.S.N.M.).f MYOTIS YUMANENSIS SOCIABILIS H. W. GrinnellMyotis yumanensis Miller, North Amer. Fauna, No. 13, p. 66, October 16, 1897(part).
?
Trouessakt, Catal. Mamm. viv. foss., p. 1283, 1899 (part).
?
Elliot, Synops. Mamm. North Amer., Field Columb. Mus., publ. 45, zool. ser.,vol. 2, p. 403, March, 1901 (part) ; List Land and Sea Mamm, North Amer.,Field Columb. Mus., publ. 57, zool. ser., vol. 2, p. 517, June 1901 (part).Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p. 256, December27, 1901 (part).
?
Elliot, Field Columb. Mus., publ. 91, zool. ser., vol. 3,p. 318, March, 1904 (part) ; Land and Sea Mamm. Middle Amer., FieldColumb. Mus., publ. 95, zool. ser., vol. 4, p. 576, 1904 (part).
?
Trouessabt,Catal. Mamm. viv. foss., suppl., p. 92, 1904 (part).
?
Elliot, Check ListMamm. North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 475,1905 (part).
?
Stephens, California Mammals, p. 267, 1906 (part).
?
Elliot,Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115, zool. ser.,vol. 8, p. 501, 1907 (part).
?
Miller, List North Amer. Land Mamm. 1911,Bull, U. S. Nat. Mus., No. 79, p. 56, December 31, 1912 (part).?J, GRiNNEax,Proc. California Acad. Sci., ser. 4, vol. 3, p. 276, August 28, 1913 (part),?J. Geinnell and Swahth, Univ. California Publ. Zool., vol. 10, p. 380,October 31, 1913.Myotis lucifugus longicrus J. Grinnell, Univ. California Publ, Zool,, vol. 5,p. 158, December 31, 1908 (not of True).Myotis yumanetisis sociabilis H. W. Grinnell, Univ. California Publ. Zool., vol.12, p. 318, December 14, 1914; Univ. California Publ, Zool., vol. 17, p. 276,January 31, 1918,?A, B. Howexl, Journ, Mamm., vol. 1, p. 173, December4, 1920.?J. Grinnell, Univ. California Publ. Zool., vol. 21, p. 314, January27, 1923.?Miller, List North Amer. Recent Mamm, 1923, Bull. U. S. Nat.Mus., No, 128, p, 69, April 29, 1924,Myotis yumanensis subsp,, Dice, Journ, Mamm., vol. 1, p. 11, November 28,1919.Type locality.?Old Fort Tejon, Tehachapi Mountains, KernCounty, Calif.Type specimen.?Adult female, skin and skull. No, 5158, Museumof Vertebrate Zoology, University of California, collected at OldFort Tejon, Tehachapi Mountains, Kern County, Calif., July 23,1904, by Joseph Grinnell.
AMERICAISr BATS OF THE GENERA MYOTIS AND PIZONYX 69Distribution.?From southeastern British Columbia and westernMontana to the eastern base of the Cascade Mountains in centralWashington and Oregon, thence southward in California (exceptingthe humid coastal strip which extends as far south as San LuisObispo County) to the coast of the southern part of the state.Diagnosis.?Color intermediate between that of the pallid Myotisyumanensis yumanensis of the deserts and the dark M. yummiensissaturatibs of the humid northwest coast.Color.?General color above very near tawny-olive of Ridgway(1912). Below dull whitish with a tinge of buffy. Except about theanal region the hairs are everywhere dark at their bases, nearlyfuscous. Ears and membranes translucent, pale brownish in color.Specimens in immature pelage are darker above, nearly sepia. Theterminal part of the interfemoral membrane is often whitish, and oc-casionally the extreme edge of the wing membranes as well.Measurements.?For measurements see tables, pages 73 and 75.Specimens examined^?Total number 248, from the followinglocalities
:
BRITISH COLUMBIA: Kamloops, 1 skin, nearly typical (A.N.S.P.) ; MountLehman, 1 skin (A.M.N.H.) ; Sicamous, 3 ale. (A.N.S.P.).CALIFORNIA: Alturus Lake, Modoc County, 1 ale. (U.S.N.M.) ; Beswick,Siskiyou County, 2 skins (U.S.N.M.), nearly typical; Capistrano, OrangeCounty, 1 skin (U.S.N.M.) ; Cassel, Shasta County, 1 ale. (U.S.N.M.) ;Chalk Peak, Monterey County, 1 skin (U. C.) ; Chico, Sacramento River,Butte County, 1 skin (U.S.N.M.) ; Eagle Lake, Lassen County, 2 skins, 2ale. (U.S.N.M.) ; Fort Tejon, Kern County, 48 skins including type (U.C), 1 skin (U.S.N.M.), 4 skins (A.M.N.H.), 1 skin (F. M.), 1 skin(A.N.S.P.), 11 ale. (U.S.N.M.) ; Fresno, Fresno County, 4 ale. (U.S.N.M.) ;near Fruto, Glenn County, 1 skin (U. C.) ; Nevada City, Nevada County,2 ale. (U.S.N.M.) ; near Oroville, Butte County, 1 skin (U. C.) ; SanBernardino Mountains, San Bernardino County, 1 ale. (B. M.) ; San LuisRey, San Diego County, 8 ale. (U.S.N.M.) ; Tulare, Tulare County. 30 ale.(U.S.N.M.) ; no exact locality, 4 ale. (U.S.N.M.).IDAHO: Payette Valley, 1 skin (U. C).MONTANA: Belton, 1 skull (U.S.N.M.); Corvallis, 36 skins, 48 ale., 1skull (U.S.N.M.) ; Flathead Lake, Flathead County, 7 ale. (U.S.N.M.) ;Powderville, 1 skull (U.S.N.M.).OREGON: Klamath Falls, 8 skins (U. C), 1 ale. (U.S.N.M.) ; Lone Rock, 2ale. (U.S.N.M.).WASHINGTON: Chelan, 3 ale. (U.S.N.M.) ; Stehekin, 3 skins (U.S.N.M.) ;Fort Walla Walla, 1 ale. (U.S.N.M.).WYOMING: Fremont Peak, 1 skin (U.S.N.M.)."NORTHWESTERN AMERICA": 1 skin (B. M.).Remarks.?Myotis yvmumensis sociahilis is intermediate in colorbetween the typical form of Myotis yumamensis and the dark M.yimmnensis saturatus of the northwest coast, but the color charactersare fairly uniform over a wide area. In geographic variation MyotisyuTTianensis parallels other members of the genus ; for example, Myotis
70 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
volans and M. lucifugus, in which there is a saturate dark form con-fined to the humid northwest coast, a pallid race characteristic of thehot arid interior and an intermediate form covering a wide area be-tween. The present race is usually distinguishable at a glance fromMyotis yumamensis saturatus by its brownish ears and membranesand lighter brown fur, and from the typical yummiensis by its dullbrown instead of pale buffy color. In a series from western Montana(Corvallis) the forearm is unusually long, 20 specimens averaging35.5, with extreme as high as 36.7. In M. htcifugtts from the sameplace, the forearm is 39 mm. Specimens from Shasta and SiskiyouCounties, Calif., are so perfectly intermediate between socidbilis andsaturatus that they might be equally well placed under either form.The more exact definition of the eastward range and the determina-tion of the area of intergradation with the typical subspecies are yetto be worked out in detail.MYOTIS YUMANENSIS SATURATUS MillerMyotis yumanensis saturatus Miller, North Amer. Fauna, No. 13, p. 68, October16, 1897.
?
Trouessart, Catal. Mamm. viv. foss., p. 1283, 1899.
?
Elliot, FieldColumb. Mus., publ. 32, zool. ser., vol. 1, p. 276, March. 1899 ; Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 403, March,1901 ; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ.57, zool. ser., vol. 2, p. 517, June, 1901.
?
Miller and Rehn, Proc. BostonSoc. Nat. Hist., vol. 30, p. 256, December 27, 1901.?Trouessart, Catal.Mamm. viv. foss., suppl., p. 93, 1904.
?
Elliot, Check List Mamm. NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 475, 1905.
?
Stephens, California Mammals, p. 267, 1906.
?
Elliot, Catal. Mamm. FieldColumb. Mus., Field Columb. Mus., publ. 115, zool. ser., vol. 8, p. 502, 1907.Lyon and Osgood, List Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat.Mus., No. 62, p. 273, January 28, 1909. Miller, List North Amer. LandMamm. 1911, Bull. U. S. Nat. Mus., No. 19, p. 56, December 31, 1912.?J.Grinnell, Proc. California Acad. Sci., ser. 4, vol. 3, p. 277, August 28, 1913(part).?H. W. Grinnell, Univ. California Publ. Zool., vol. 17, p. 278, Jan-uary 31, 1918.?G. M. Allen, Journ. Mamm., vol. 1, p. 1, November29, 1919.?Miller, List North Amer. Recent Mamm. 1923, Bull. U. S.Nat. Mus., No. 128, p. 70, April 29, 1924.Type locality.?Hamilton, Skagit County, Wash.Type specimen.?Adult male, skin and skull No. ^|fg| UnitedStates National Museum (Biological Survey collection), collected atHamilton, Skagit County, Wash., September 13, 1889, by T. S.Palmer. Original number 392.Distribution.?Humid northwest coast from southern BritishColumbia south to south-central California (San Luis Obispo County)and to a varying distance inland west of the higher mountains.Diagnosis.?Like Myotis ywnanensis yumanensis and M. y. soci-ahilis but color much darker.
AMERICAlSr BATS OF THE GENERA MYOTIS AND PIZONYX 71Color.?Above, the tips of the hairs are usually a dull sepia, thebuses slate-black; below, the tips of the hairs are buffy in the mid-ventral region, washed on the throat and sides with sepia, their basesslate-black except for a small area at the base of the tail where theyare buffy throughout. The ears and membranes are blackish brown,very dark. In some lights the paler tips of the hairs on the back areglossy, with a brassy sheen, but not to the same extent as in the raceof Myotis lucifugus occurring with it. Frequently the whole lowersurface inclines to a dull buffy tint. The general effect is a small,large-footed, dull-haired bat of uniformly dark color. Immaturespecimens are even darker, a dull blackish brown above, washedbelow with smoky.Measurements.?For measurements see tables, pages 74 and 75.Specimens examined.?Total number 64, from the following lo-calities :BRITISH COLUMBIA: Chilliwack, 1 skin (F. M.) ; Kamloops, 1 skin(U.S.N.M.) ; Okanagan, 1 skin (B. M.), nearly typical; Port Moody,1 skin (U.S.N.M.) ; Shuswap. 1 skin (U.S.N.M.) ; Westminster, 2 skins(A. M. N. H.).CALIFORNIA: Baird, Shasta County, 1 skin (U.S.N.M.), nearly typical;Cuddeback, Humboldt County, 1 skin (U. C.) ; Lake Leonard, MendocinoCounty, 1 skin (U. C.) ; Menlo Park, San Mateo County, 1 ale.(U.S.N.M.) ; Nicasio, Marin County, 2 ale. (U.S.N.M.) ; San CarpojoCreek, 7 miles north of Piedras Blancas, San Luis Obispo County, 1 skin(U. C).OREGON: Blaine, Tillamook County, 3 skins (U. C), 2 skins (A. Walker) ;Columbia River, 1 skin (A.N.S.P.) ; Crooked River, Crook County,1 ale. (U.S.N.M.) ; McKenzie Bridge, Lane County, 3 skins (U.S.N.M.) ;Mohler, Tillamook County, 1 skin (U. C.) ; Roseburg, Douglas County,1 ale. (U.S.N.M.) ; Twelve Mile Creek, 1 ale. (U.S.N.M.) ; Vida, LaneCounty, 1 skin (U.S.N.M.) ; Wedderburn. Curry County, 1 skin (U.S.N.M.) ;Wilson River, Tillamook County, 1 skin (U.S.N.M.) ; no exact locality, 1skin (A.N.S.P.).WASHINGTON : Goldendale, 2 ale. (U.S.N.M.); Hamilton, 1 skin(U.S.N.M.) ; Husum, 1 skin (U.S.N.M.) ; Lake Cushman, 1 skin(U.S.N.M.), 1 skin (M. C. Z.), 8 skins, 5 ale. (U. M.) ; Lake Quinault,2 skins (U.S.N.M.) ; Lake Washington, 1 skin (U.S.N.M.) ; Mabton,1 ale. (U.S.N.M.) ; Neah Bay, 1 ale. (U.S.N.M.) ; Nesqually Flats, 5 ale.(A.N.S.P.) ; North Yakima, 1 ale. (U.S.N.M.) ; Roy, 2 skins (U.S.N.M.),not typical; Stevenson. 1 skin (U.S.N.M.).Remarks.?The form of Myotis yumanensis occurring on theNorthwest Coast closely parallels the race {alascensis) of Myotislucifugus which is found in the same region, so that, in collectionsof skins from northwestern localities, the two are often confused.Usually saturatv^ is recognizable by its smaller size, narrower ear,and duller coat, without reference to the skull.
72 BULLETIN 144, UNITED STATES NATIONAL MUSEUMMYOTIS YUMANENSIS LUTOSUS, new snbspecieaMyotis yumanensis Millee, North Amer. Fauna, No. 13, p. 66, October 16, 1897(part).Type.?A^AM\t female, skin and skull, No. 50783, United StatesNational Museum (Biological Survey collection), from Patzcuaro,Michoacan, Mexico. Collected July 17, 1892, by E. W. Nelson.Distribution.?Southern portion of the Mexican highlands.Diagnosis.?Color darker and richer than in Myotis yumanensissociahHis; underparts distinctly washed with brownish.Color.?The color differs conspic'uously from that of the typicalform of Myotis yumanensis., being a dark cinnamon brown above,nearly Front's brown (Ridgway, 1912), the bases of the hairs inadults but very little darker. Below, the extreme bases of the hairs,except in the anal region, are blackish brown, paling into fuscous intheir upper part and tipped with dull gray, the entire ventral surface,especially at the sides, having a faint brownish wash. Ears andmembranes dull brownish, the tip of the tail paler.Measureiivents.?For measurements see tables, pages 74 and 76.Specimens examined.?Total number 33, from the following lo-calities :JALISCO: 2 ale. (B.M.).MICHOACAN: El Molino, 1 ale. (U.S.N.M.) ; Patzcuaro, 2 skins, 7 ale.(U.S.N.M.).SAN LUIS POTOSI: Ahualulco, 9 ale. (U.S.N.M.); La Parada (Hda.),3 ale. (U.S.N.M.) ; Jesus Maria, 7 ale. (U.S.N.M.).ZACATECAS: San Juan Capistrano, 1 skin, 1 ale. (U.S.N.M.).Remarks.?In its dull brownish (muddy) color this Mexican batresembles Myotis yumunen^is sociabilis, but the brown is slightlyricher, and in the adults there is very little contrast between the tipsand the bases of the hairs above. The darker belly with its distinctbrownish wash is also apparently characteristic. An immature speci-men from Zacatecas, Mexico, is slightly paler and the hairs of theupper parts are rather distinctly bicolor. Between the ranges ofthese two dark forms, typical yum/inensis with its pale colorationintervenes. Thus far very few specimens of the Mexican race havebeen available. Hence the exact definition of its range must awaitfurther coUectins:.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 73External measurements of Myotis yumanensis
Locality
Myotis yumanensisyumanensisNevada:Pyramid LakeDoDoDoDoDoDoDoDoDoCalifornia:Fort YumaDoLone PineDoOwens LakeDoDoArizona:Ehrenburg--DoDoDoDo...SaffordYumaDoDoDoDoDoDoDoNew Mexico:RinconadaLower California:Rancho San Antonio..DoDoDoSonora:Colonia LerdoDoDo.Do.Do.Do.Do.Do.Do.Do.Myotis yumanensissociabilisMontana:CorvallisDoDoDoDoDo...DoDo...DoDoDoDoFlathead LakeDoWashington:ChelanDo.58518?28 6
Number
245289245291245292245299245309245310245323245325245338245345615436154428971289722895228955289571319661319681319711319721319752064155307699681996839968599687
133723147616147620147621147622137021137022137025137028137029137031137032137034137036137038
20370020370120370520370920371020371120371220371420371520371620368620372873170731753031130312
44.041.540.544.041.540.040.040.042.540.0 34.042.541.544.043.04444.542.546.544.043.044.539.042.044.546.545.545.044.043.041.043.041.5
37.033.'53427.034.033.036.035.0
35.035.536.530.033.032.034.533.033.536.037.034.032.536.835.435.637.236.536.033.236.034.0
16.015.016.516.515.014.015.015.516.015.015.016.017.015.01616.0
16.515.015.516.014.014.015.015.514.516.015.015.015.015.515.041.0 37.5 14.540.0 34.5 15.541.5 35.5 15.040.5 36.0 15.04442.54343.04143.043.040.543.041.0
42.040.046.044.044.240.244.845.043.042.041.444.042.042.0
35.537.035.037.532.034.535.033.536.035.0
37.035.537.038.037.037.838.037.832.036.033.834.634.036.2
15.015.515.516.016.015.015.515.515.515.0
16.515.516.017.216.016.416.016.015.015.416.016.015.817.29 43.0 32.2 15.4V 42.6 34.81 14.6
8.0 35.57. 6 34. 59.0 36.08.0 34.07.58.58.0 3436.036.58.5 338.5 33.58.58.08.08.58.08.58.08.08.58.08.0 33.58.0 35.58.08.08.58.58.08.58.08.59.08.58.08.58.58.57.5
35.035.034.535.033.535.033.035.035.035.0
34.034.535.533.035.036.036.034.535.535.034.035.035.535.534.0
8.5i 37.08.5; 36.59.0; 37.09.0: 38.08.4! 36.68.41 36.29.2I 37.09.2i 37.08.21 36.48.2! 37.09.4 35.48.08.48.0 32.435.037.07.4 36.08.4 36.2
8.0 36.67.0 34.08. 37. ., .7.0! 36.0 6.07. 0; 34.8.0 34.07.5 35.08.0 36.07. 5| 36.8.5; 35.5
Bs; s
33.532.034.033.532.031.532.533.033.0;33.5
31.530.532.032.031.030.531.031.532.032.06. 32. 5, 30. 56.0 31.5; 30.034.0, 32.030.5 28.032. 5i 31.032.5; 3133.5 31.0
7.06.56.05.55.56.56.56.56.05.55.56.07.07.07.06.05.55.57.0
32.031.532.033.032.532.531.032.530.533.032.531.532.033.06.5 31.56. 32. 55.5 33.06.5 31.06.5 32.57.06.56.56.56.06.06.56..07.06.0
7.07.57.57.07.06.06.67.07.07.07.06.86.46.4
3133.032.531.532.532.532.532.5
33.034.033.536.033.832.833.433.832.834.031.232.433.033.6
31.030.031.030.530.530.529.030.529.531.530.530.030.031.030.031.031.528.031.033.0 32.533.0 32.030.031.029.030.031.030.030.030.5
32.032.032.533.631.631.032.631.830.032.431.031.231.032.0
a
74 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotia yum^nensis?Continued
Locality Number 5
Eh A 03 a)Myotis jramanensis socia-bills?ContinuedWashington?Continued.ChelanFort Walla WallaOregon:KlamathLone RockDoCalifornia:Tulare.DoDoDoDo...DoDoDoDoDoOld FortTejon.DoDoDoDoDoDoDoDoMyotis yamanensissaturatnsCalifornia:Menlo ParkNicasioDoOregon:RoseburgTwelve Mile Creek ,Crooked River ,Washington:Qoldendale ,Do...HamiltonMabton ,Neah BayYakima ,Myotis yumanensls IntosusZacatecas: Hda. San JuanCapistranoSan Luis Potosi:AhualulcoDoDoDoDoDoDoDoDoDoTaJiscoDoMichoacan:PatzcuaroDoDoDoDo....
30313150131873957929979300306993070430705307063070730709307143071630721307262982029826298302984829850298522985329855I 5158 U.C.
1425446045860469
793059262892631J 24306926249142292623
923845232452325523265232752328523295233062331523325225572. 10. 7.
3
B. M.72.10.7.4B. M.52176522335226252263'5^783
43.842.644.240.042.843.642.044.044.844.444.043.843.041.441.243.042.438.044.041.439.446.243.038.0
43.043.843.441.442.840.040.242.4'47"43.239.043.4
37.841.039.445.042.042.044.046.841.437.843.043.642.0
44.842.843.441.2'49"
34.035.833.833.031.236.033.634.433.034.234.035.436.435.034.633.835.828.033.034.035.039,36.036.0
32.635.434.434.036.232.233.235.0"30"33.431.636.0
28.033.436.435.037.436.037.035.035.035.437.632.234.0
34.033.633.233.0"36"
15.016.016.014.215.215.014.015.215.015.015.415.014.616.015.014.815.216.015.015.414.415.416.016.0
15.215.414.615.216.2
15.016.015.014.815.415.0
13.015.014.814.414.415.014.814.014.614.014.413.415.0
14.815.415.014.015.0
7.8.07.26.28.4
8.28.08.47.68.2
8.07.08.48.48.07.49.08.28.48.09.0
8.48.47.48.47.29.07.88.28.68.07.0
8.27.07.48.07.88.68.07.08.07.0
8.28.08.08.0
36.836.035.233.234.035.633.435.634.434.634.434.634.435.034.036.036.033.035.034.834.035.035.033.7
34.434.434.034.435.835.034.434.033.035.833.036.0
32.034.034.634.234.436.035.034.235.434.634.632.435.2
34.435.034.833.833.4
6.46.06.06.07.06.46.06.26.06.46.26.07.06.86.06.06.06.26.06.26.46.06.2
34.6 32.033.0 31.032.0 30.431.6 30.831.8 30.031.68: 30.230.02 31.030.24 30.031.230.230.429.831.832.030.031.032.031.031.029.6
6.86.25.85.46.26.05.86.05.05.46.66.8
6.26.06.26.46.07.06.26.46.06.06.47.0
32.031.831.033.232.633.431.231.432.030.32.0
28.631.831.632.032.032.832.432.032.831.832.630.032.0
6.2 32.86.0 32.06. 31. 46.8 30.86.2 31.2
13.013.213.012.011.613.012.412.413.413.213.213.412.013.013.012.212.012.012.412.213.014.012.4
30.231.029.830.631.031.430.030.0
14.012.412.413.013.213.012.413.031.029.231.0
27.029.629.830.030.231.431.030.231.629.831.029.031.0
31.230.830.229.830.0
12.612.214.4
12.012.012.013.614.013.012.413.013.412.614.614.013.8
13.212.413.413.2
' Type; from H. W. Grinnell, 1918. Type.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 75Cranial measurements of Myotis yumanensis
Locality
76 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis yumanensis?Continued
Locality Number
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 77
orbital constriction, narrower brain case, and in the constant presence(in adults) of a low but perfectly formed sagittal crest.ExteiTiol characters.?The external characters as shown by a seriesof 12 skins from Florida and four from Indiana do not differ ap-preciably from those of Myotis lucifugus hicifugus otherwise than inthe short, dense, woolly quality of the fur, and the dull colorationof the dorsal surface. Hairs at middle of back, in fresh pelage,usually about 6 to 7 mm. in length instead of 9 to 10 mm. In someindividuals there appears to be tendency for the wing membrane tobe inserted at or near the ankle, but this may be due to a peculiarityin the make-up of the skins.Fur amd color.?The fur of this species is distinctive by reason ofits thick, woolly appearance, resulting from the fine and rather uni-form texture without a conspicuous over fur. This quality of thefur, together with the lack of long burnished tips to the hairs of theback as well as the absence of strong contrast in color between thetips and the bases of the hairs, serves to distinguish the animal readilyfrom M. lucifugus which it otherwise somewhat resembles. Theactual elements of the color are essentially the same as in Myotis luci-fugus lucifugus, but the shortness and inconspicuousness of the bur-nished hair tips on the upperparts give a noticeable dull and luster-less effect usually ranging in tint from sudan brown to Saccardo'sumber (Ridgway, 1912). On parting the hairs the plumbeous basalarea of the fur is seen to be less extensive and less dark than in M.lucifugus. In some individuals the back has a peculiar drabby castwhich we have never seen in M. lucifugus. Underparts dull buff,much as in M. I. lucifugus but the palest individual paler than usualin the related animal.Skull.?The skull is more slender throughout than that of Myotislucifugus. While this character is most obvious in the interorbitalregion (interorbital breadth in 16 skulls varying from 3.6 to 4.0 asagainst 4.0 to 4.4 in 60 skulls of true M. lucifugus) it can also beappreciated on direct comparison of the breadth of the rostrum overbases of canines, as well as of the lacrymal breadth and the breadth ofbrain case. A low but perfectly definite and sharp-edged sagittalcrest is present in all of the adult specimens examined, this characteralone being sufficient to distinguish the series from any similar seriesof Myotis lucifugus. The crest resembles that present in the other-wise very different species, M. velifer and M. grisesceriM?. When thebrain case is viewed from behind, side by side with specimens of M.lucifugus from the eastern United States, the narrower, relativelyhigher form in Myotis au^troripaHus is at once evident. In thisview the summit of the arch in M. lucifugus is nearly always obvi-
78 BULLETIN" 144, UNITED STATES NATIONAL MUSEUM
ously flattened, while in M. austroHparius it is distinctly convex.When the skull is seen from below its general narrowness is evidentbut there appear to be no structural peculiarities to distinguish itfrom the skull of M. ludfugus lucifugus.Teeth.?The teeth are about the same size as those of Myotis lucifugvslucifuffti^, therefore smaller than those of M. grisescens and M, velifer.In structure they show no very obvious peculiarities. The uppermolars have the full complement of secondary cusps and ridges, andthe cingulum on the inner margin of their crowns is almost invariablyas much developed as in the maximum condition seen in Myotis lud-fugus (2 exceptions among 16 specimens), a character usually sharedby M. grisescens and less frequently by M. velifer. The two uppersmall premolars are very slightly crowded, the anterior with thetransverse axis of its crown perceptibly greater than the longitudinalaxis. The anterior tooth is about twice the height of the posterior.The lower canine is relatively short and stout, hardly exceeding thethird premolar. The two small anterior lower premolars are fullyin the tooth row, the posterior tooth only slightly exceeded in heightand cross section by the anterior.Measurements.?Foi- measurements see tables, page 80.Specvmens examined.?Twenty-three from the following localities
:
?CANADA: 1 skin (B.M.).FLORIDA : Tarpon Springs, Pinellas County, 2 skins including type, 4 ale.(A.N.S.P.) ; Bird Key, Tampa Bay, Pinellas County, 12 skins (Copeland).INDIANA: Mitchell, 4 skins (U.S.N.M.).Remarks.?Myotis austrariparius bears a general resemblance toseveral of the brownish species occurring in eastern North America,but may be distinguished by its large and strong hind foot withoutkeel on the calcar, and by its thick, almost wooly fur of a dull cinna-mon or drab tint. In the drab pelage it somewhat resembles theslightly larger Myotis grisescens, but in this species there is no obvi-ous basal darkening of the hairs of the back, and the origin of thewing membrane is from the ankle instead of from the metatarsalsnear the base of toes. From M. lueifugus, with which it is doubtlessoften associated, it is distinguished by the lack of noticeable burn-ished tips to the hairs, and by the uniform presence of a distinctsagittal crest on the skull. From Myotis velifer, a species with simi-larly dull unburnished fur, it differs in smaller size, narrow, weakrostrum, and the normal (not enlarged) cheek teeth. From Myotiskeenii septentrionalis it is distinguished by its larger feet, less elon-gate ears, and crested skull.Although the species was described thirty years ago, its true char-acters are now for the first time recognized. The first specimens re-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 79corded (Rhoads, 1894) were collected by Mr. W. S. Dickinson in1892 at Tarpon Springs, Florida, and were identified by HarrisonAllen as Vespertih'o gryphus {^Myotis lucifugus). Three yearslater Rhoads, noticing that its dense, lusterless, dull-colored fur wasunlike that of northern Myotis lucifugus^ separated the Florida ani-mal as a subspecies, under the name austroriparius.He then submitted his material to Miller who saw that the twoskins, one of them the type, pertained to immature individuals andthat the specimens in alcohol did not differ appreciably, except intheir short fur and dull color, from true M. lucifugus. Believingthat the pecularities of the series were all due to immaturity andthe effects of alcohol Miller placed Khoads' name in the synonymyof lucifugus^ where it has remained until now. The animal was nextcollected by Hahn in Indiana, but was confused by him with Myotis
'''' suhulatu^'''' {=M. keenii septentnondlis) . This error was detectedby Allen while examining Hahn's material, now in the United StatesNational Museum, during the preparation of this monograph. Thecharacters, when once seen, were so obvious that we unhesitatinglydrew up a description of Hahn's bat as a new species. It remainedthus until the manuscript was nearly ready to go to press, when,fortunately, Mr. Copeland sent us his series of twelve skins of adultsfrom Bird Key, Tampa Bay, south of Tarpon Springs. Study ofthis material soon showed two things: that the austroriparius ofRhoads was specifically distinct from Myotis 1/acifugus, and thatthe animal was identical with the bat subsequently taken in Indianaby Hahn.When he placed the name austroriparius in the synonymy ofMyotis lucifugus Miller called attention to the likelihood that Le-Conte's animal might prove to be the same as the one described byRhoads, should the latter ever be shown to possess distinctive char-acters.^*^ This possibility has not yet been disposed of, as we stillhave no knowledge as to the forms of Myotis actually occurring insouthern Georgia. It may be said, however, that LeConte has writtennothing in either of his descriptions of Vespertilio lucifugus whichpoints toward the present animal, and also that the only knownspecimen that can be reasonably supposed to have been determinedby LeConte (see p. 44) is a typical example of Myotis lucifugushicifugus as now understood.
^? North Amer. Fauna, No. 13, p. 21, October 16, 1897.
80 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotia austroriparius
Locality
AMERICAN" BATS OP THE GENERA MYOTIS AND PIZONYX 81
fig. (head), 1912.?Miller, List North Amer. Land Mamm. 1911, Bull. U. S.Nat. Mus., No. 79, p. 54, December 31, 1912.?A. H. Howell, North Amer.Fauna, No. 45, p. 23, October 28, 1921.
?
Elliot, Check List Mammals NorthAmer., suppl., p. 154, 1917.
?
Millek, List North Amer. Recent Mamm. 1923,Bull. U. S. Nat. Mus., No. 128, p. 67, April 29, 1924.Type locality.?Nickajack Cave, Marion County, Tenn. ?Type.?Adult male, skin and skull. No. 157517 United StatesNational Museum (Biological Survey Collection), collected in Nicka-jack Cave, near Shellmound, Marion County, Tenn., August 31, 1903,by A. H. Howell.
Map 3.
?
Disteibution of Myotis ohiskscensDistribution.?Limestone area from extreme southern Indiana andIllinois south to Tennessee, Georgia, and central Alabama, westwardto southwestern Missouri and northern Arkansas.The range is still imperfectly known, but, as indicated by specimensnow at hand, it apparently does not meet that of Myotis veliferincautus at any point, nor does it extend east of the Alleghenies.The occurrence of this animal in Kentucky can hardly be doubted,though no specimens are available. This distribution is probablyconditioned in part by the presence of* large limestone caves availablefor permanent habitation, and the occurrence of the caves is in turndependent on the geological structure of the country. For unlikemost other cave-haunting Myotis, of eastern North America at least,
82 BULLETIIT 144, UNITED STATES NATIONAL MUSEUMthis species frequents these caverns regularly in summer, as well asduring the hibernation period, so that the grisescens population ismore concentrated than in the case of those species (as M. lucifugus,M. keenii septentHonalis) that largely desert the caves in sunmier.]y[owell (1909, 1921) has given a brief account of these Tennesseecaves and their bat population.Diagnosis.?Size decidedly greater than in Myotis lucifugus^ butgeneral proportions not strikingly different (ratio of tail to head andbody in 10 topotypes, 76.4) ; forearm usually more than 40 mm.
;
greatest length of skull ranging from 15.5 to 16.5 mm., maxillary toothrow ranging from 5.8 to 6.2 mm., lower tooth row more than 6 mm.(6.2 to 6.6 mm.) ; skull with an obvious sagittal crest in adults; cheekteeth proportioned to the palate as in M. lucifugus., not enlarged as inM. velifer; first and second upper molars with protoconule unusuallywell developed; a broad, conspicuous cingulum on inner marginof crown. Differs from all other known North American species inthe insertion of the wing membrane at the tarsus instead of the sideof the foot and in the absence of dark bases to the hairs of the back.Ears.?The ears are about as in Myotis velifer: when laid for"\vardreaching the nostril or slightly beyond it. Tragus as in M. velifer^with bluntly pointed tip.Wing and mnemhranes.?^Wing membranes from the base of thetarsus instead of from the base of the toes as in all known NorthAmerican species, a character which, though obvious in alcoholicspecimens, is often obscure in skins if the membrane is stretchedlaterally in drying. Metacarpals graduated, the third longest, thefourth and fifth successively shorter. Taking the third finger as100, the fourth is 86, the fifth 80. When the wing is folded the thirdmetacarpal falls 4 mm. short of the elbow.Foot.?Except for their larger size the foot and calcar resemblethose of Myotis lucifugus^ though the foot is longer proportionallyto the tibia. In 10 topotypes the ratio of foot to tibia averages 60.1
;
in 10 specimens from Illinois it averages 58.8.Fur and color.?The fur is more velvety than in Myotis lucifugus,and the hairs of the dorsal surface are strikingly characterized bybeing of essentially the same tint throughout instead of conspicuouslydarker at the base. There are two color phases: dusky and russet.The former is represented by the series of 20 skins from the typelocality, Nickajack Cave. These average darker than the other avail-able speciijiens, and are a uniform smol^ (about "chsetura drab")above to the roots of the hairs ; below paler, the hairs " dark mousegray " at their bases with dull whitish tips, those of the chin lighter,and those between the thighs whitish throughout.Specimens from Indian Cave, Tenn., and from Rosiclare, lU., areof a slightly warmer tint. In a series of 12 skins from Rogersville,
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZON"YX 83Ala. (U.S.N.M.), both phases are represented by adults: The duskyas above described ; and the russet, a nearly uniform " cinnamon-brown " above to the roots of the hairs, the lower surfaces with thewhitish tips replaced by a pale buff, contrasting with the darkerbases of the hairs.Skull.?The skull differs from that of Myotis lucifugus in its largersize. In shape and proportions the crania of the two species are prac-tically alike, but the evident sagittal and lambdoid crests make theskull of Myotis grisescens readily distinguishable.Teeth.?The two small upper premolars stand nearly in the toothrow so that both are clearly visible in side view, while from below,a line connecting the posterior angle of the canine and the anteriorcorner of the large p* passes through the outer part of both the otherpremolars. In tii^ and m^ the protoconule is unusually well de-veloped, but in actual structure the small cusps and their accompany-ing ridges do not differ from those which occur in Myotis lucifugits.Cingulum on inner side of crown in m^ and rnr well developed andconspicuous, but apparently never extending around the antero-lingual base of the protocone. At the postero-lingual base of thehypocone the cingulum frequently becomes so thickened as to re-semble a low cusplike tubercle applied to the base of the hypocone.This tubercle may in rare instances (No. 202022, U.S.N.M.) becomeso large that it bears at its summit a distinctly worn area produced inm- hy the protoconid of m.^ and in m^ by the protoconid of mg. Tlieteeth of the lower jaw sliow no special characteristics.Measurements.?For measurements see tables, pages 84 and 85.Specmiens examiined.?Total number 258, from the followinglocalities
:
ALABAMA: Anniston, Calhoun County, 2 ale. (M. O. Z.) ; Fort Deposit,Marshall County, 10 skins, 8 ale. (U.S.N.M.) ; Rogersville, LauderdaleCounty, 12 skins, 49 ale. (U.S.N.M.).ARKANSAS: [? Osase River], 1 ale. (M. C. Z.).ILLINOIS: Rosiclare, Hardin County, 12 skins, 33 ale. (F. M.).MISSOURI: Columbus, 1 ale. (U.S.N.M.): Marble Cave, Stone County,12 skins, 3 ale. (U.S.N.M.).TENNESSEE: Indian Cave, Grainger County, 13 skins (Univ. Wisconsin),15 ale. (U.S.N.M.), 1 skin (M. C. Z.) ; New Market, Grainger County.1 skin (M. C Z.) ; Nicka.1aek Cave, Marion County, 21 skins, 62 ale.(U.S.N.M.), 1 ale. (A. N. S. P.) ; Shell Mound, 1 ale. (U.S.N.M.).Habits.?An interesting point in connection with the fact thatcolonies of Myotis grisescens frequent the same caves in both summerand winter is the segregation of the sexes at certain times of year.The breeding females seem to congregate apart from the males. Theyoung are probably born in early July, and when these are wellgrown a breeding colony naturally contains the adult females as wellas immature bats of both sexes. But in August, the colonies seem
84 BULLETIN" 144, UNITED STATES NATIONAL MUSEUMto contain adult males as well as adult females. Thus of 21 batsfrom Nickajack Cave, Tenn., August 31, 15 were adult males, 6 adultfemales; and of 9 from Rosiclare, 111., August 14, 4 were adult males,5 adult females. Presumably pairing takes place in the fall, afterwhich the sexes probably segregate until the following August. Atall events, 17 specimens taken April 14 at Indian Cave, Tenn^were all females. Again, a series of 18 from Fort Deposit, Ala.,June 18-19, proved to be all males, while of 12 adults taken a littlelater, July 9, at Rogersville, Ala., all but 1 were females.Remarks.?Superficially^ Myotis gi'isescens resembles M. velifer inits large size and dull fur. It is, however, a very distinct animal.Apparently is has no near relatives, the insertion of the wing mem-brane at the ankle joint and the peculiar color pattern of the hairson the back distinguishing it sharply from all known Americanspecies.Abnormalities in the number or position of the two minute upperpremolars are interesting as indicating the evolutionary tendencyin connection with the reduction of the tooth row. Thus in 16067(Field Museum) from Rosiclare, 111., these two teeth are so placedthat instead of standing in the tooth row, the smaller {j)^) on eachside is drawn inward while the anterior tooth practically fills theentire space between the canine and p* so that the smaller tooth ishardly visible in side view.Specimens of Myotic grisescens were collected in late June 1892,at Marble Cave, Mo., by Vernon Bailey, and were referred by Miller(1897) to Myotis velifer. Of three individuals taken, "one wascaught in the cave 150 feet below the surface of the earth ; the otherswere shot as they came out of the mouth of the cave in the evening "(Bailey, 1905). The true characters of the animal were detected byHowell, in the series of specimens which he secured in NickajackCave on August 31, 1908.External measurements of Myotis grisescens
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 85External measurements of Myotis grisescens?Continued
Locality Number
.a
86 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis grisescens-?Continued
Locality
Tennessee?Continued
.
Indian Cave, GraingerCountyDoDoDoDoDoDoAlabama:Fort DepositDoDoDoDoDoDoDoDoDoRogersviUe ,DoDoDoDoDoDoDoDo
Number
894895896
900201698201699201700201701201702201703201704201705201706201707202020202021202022202024202025202026202029202030202031
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 87Ears.?The ears are moderately long, reaching, when laid forward,to the nostril or slightly beyond it; tragus slender, about one-halfthe height of the ear, its anterior edge nearly straight, broadest atbase, narrowing in the terminal half to a bluntly rounded tip, andprovided with small lobule at its base posteriorly.
Map 4.
?
Distribution of Myotis velifer : 1, M. velifer velifer ; 2, M. vehfeeiNCAUXus ; 3, M. velifer peninsdlakisWing (md membranes.?^Wing membrane arising from base of toes.Third and fourth metacarpals usually subequal, the fifth decidedlyshorter; when folded the third metacarpal reaches to within 2.5to 3 mm. of the elbow. Taking the third finger as 100, the fourthis 86, the fifth 80 (72 : 62 : 57.5 mm.) . Tail involved in the membraneexcept for the minute terminal and half the penultimate vertebrae
88 BULLETIN 144, UNITED STATES NATIONAL MUSEUMwhich are free. Border of uropataguiin sprinkled with widely spacedstiff hairs, scarcely visible without the aid of a lens.Foot.?The foot is strong, usually a little more than half the lengthof the tibia, the ratio of its length to that of tibia averaging 51.2in 10 specimens from Arizona, 53.4 in a like number from Michoacan,and 54.1 in 9 from Lower California. Calcar well developed, andterminating distally in a minute lobule, the skin along its free edgethickened on the basal half but without forming a distinct keel.Fur and color.?The pelage is of moderate length, neither close noryet full and fluffy, the longer hairs on the back averaging about 6 mm.in length, their tips not burnished. The general coloration is uni-formly dull sepia or drab above, somewhat paler below, the bases ofthe hairs everywhere dark except at the sides of the belly underneath,where the hairs are whitish throughout.Skvll.?The skull is characterized by its broad rostrum, the areaof which, when skull is viewed from above, is not conspicuously lessthan that of the brain case. (PI. 1, p. 7, fig. 6.) Distance from ante-rior border of alveolus of inner incisor to narrowest region of inter-orbital constriction about equal to that from constriction to lambda.Breadth of rostrum across roots of canines greater than interorbitalconstriction. Maxillary breadth at m^ (average in 20 specimens, 6.9)nearly equal to breadth of brain case (average in 20 specimens, 7.2).Brain case higher and narrower than that of Myotis lucifugiis. Inrear view the lambdoid crests are sharply defined. Seen from abovethey form a sharp transverse ridge posteriorly which meets the well-marked sagittal crest at the occiput and cuts off a raised triangulararea. In side view the rostrum appears nearly flat anteriorly ; behindthis region the forehead rises more abruptly than in Myotis lucifugus.Teeth.?In their general structure the teeth resemble those ofMyotis lucifugus. The secondary cusps and ridges (see fig. 1, p. 8)are all fully developed but the cingulum is usually less distinct thanin the smaller animal. The hypocone tends to be lower relatively tothe protocone than is generally the case in M. lucifugus, a peculiaritywhich is best appreciated when the teeth of the two animals are com-pared from their lingual aspect. The greatest width of the palatemeasured across the outer borders of the upper molars slightly ejc-ceeds the maxillary tooth row (front of canine to back of vi^) andis practically equal to or is at most minutely less than the mandibulartooth row (exclusive of incisors). The maxillary rows of cheek teethas a whole are obviously more robust (broader in proportion to theintervening area of palate) than is usual in other large Americanmembers of the genus. This character can not be described withsufficient definiteness to insure the identification of a single specimentaken by itself, but it ig at once obvious on comparison of a few*skulls of Myotis velifer with similar series of M. thysanodes or M.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 89grisescens or with a large individual of M. lueifugus as in Plate 1(p. 7), Figures Za and 3&. First and second upper premolars slightlydrawn in from the tooth row so that the second is partly hidden inexterior view behind the front corner of the third premolar.HahitS'?Myotic velifer is a colonial species, typically a cave-dweller, so that its local distribution and abundance may frequentlybe conditioned in part by the presence of caves suitable for habita-tion. Ward (1891, 1904) found great numbers in a state of semi-hibernation in the tunnel-like caverns formed in lava, on the vol-cano of Perote, in Jalapa, Vera Cruz, Mexico, during the month ofFebruary. The animals are quick to take advantage of the artificialshelter offered by old buildings and are therefore considered as
"house bats " in parts of Texas and Arizona. Bailey (1905) has de-scribed how they came from a considerable distance to drink at anartificial pool in the dry country at Carlsbad, Tex., flying appar-ently from the limestone hills several miles away, " straight for thewater pool without a crook or turn." Mine tunnels are also used asroosting places.Remarks.?In its structural details Myotis velifer suggests a largeMyotis lucifugus, with the cheek teeth broadened but not sensiblyaltered in structure. The form of the skull is less flattened than inthe smaller animal, the rostrum is broader, the interorbital constric-tion relatively narrower, and the brain case fuller and more globose,with sharply defined sagittal crest (compare figs. 3a and 3&, pi. 1,p. 7). Externally it resembles Myotis grisescens^ which was con-fused with it by Miller in 1897, but it is readily distinguishable bythe normal insertion of the wing membrane on the foot, and by thedarkened bases of the hairs on the back.MYOTIS VELIFER VELIFER (J. A. Allen)Vespertilio albescens Tbue, Proc. U. S. Nat. Mus., vol. 7, p. 603, 1885 (part, notof Geoffroy).Vespertilio velifer J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 3, p. 177, De-cember 10, 1890.?H. L. Waed, Amer. Nat., vol. 25, p. 744, August, 1891.Vespertilio albescens velifer H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 92, March 14, 1894.?Trouessaet, Catal.Mamm. viv. foss., p. 132, 1897.Myotis velifer Millek, North Amer. Fauna, No. 13, p. 56, October 16, 1897.
?
Trouessart, Catal. Mamm. viv. foss., p. 1285, 1899.
?
Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 401, March,1901; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ.57, zool. ser., vol. 2, p. 517, June, 1901.
?
Miller and Rehn, Proc. Boston Soc.Nat. Hist., vol. 30, p. 255, December 27, 1901.
?
Elliot, Land and Sea Mamm.Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 573, 1904.?Trouessart, Catal. Mamm. viv. foss., suppl., p. 92, 1904.?Ward, Trans.Wisconsin Acad. Sci., vol. 14, p. 647, 1904. Bailey, North Amer. Fauna,58518?28 7
90 BULLETIN 144, UNITED STATES NATIONAL MUSEUMNo. 25, p. 208, October 24, 1905.?Elliot, Check List Mamm. North Amer.,Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 473, 1905; Catal. Mamm.Field Columb. Mus., Field Columb. Mus., publ. 115, zool. ser., vol. 8, p. 500,1907.
?
Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus.,No. 79, p. 93, December 31, 1912; List North Amer. Recent Mamm. 1923,Bull. U. S. Nat. Mus., No. 128, p. 93, April 29, 1924.Myotis californicus jaliscensis Menegaux, Bull. Mus. d'Hist. Nat. Paris, vol.7, p. 321, 1901 (near Lake Zacoalco, Jalisco, Mexico, type in Paris Mu-seum).
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 31. p. 121,August 27, 1903.
?
Elliot, Land and Sea Mamm. Middle Amer., FieldColumb. Mus., publ. 95, zool. ser., vol. 4, p. 579, 1904.
?
Trouessart, Catal.Mamm. viv. foss., suppl., p. 93, 1904.
?
Elliot, Check List Mamm. NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 477, 1905.Type locality.?Santa Cruz del Valle, Guadalajara, Jalisco,Mexico.Type specimen.?Adult male, skin and skull. No. 2696 Amer. Mus.Nat. Hist., collected at Santa Cruz del Valle, Guadalajara, Jalisco,Mexico, September 7, 1889, by Dr. A. C. Buller.Distribution.?From the highlands of Guatemala northward towestern Arizona and southern California.Diagnosis.?Size slightly below the maximum for the species;greatest length of skull 15.6 to 17.0 mm. ; color dark.Descrnptimi.?Topotypes are a dull " sepia " (Ridgway, 1912)above, the hairs with their basal three-fourths about " blackishbrown " ; below much paler, the tips of the hairs pale cream-buff,their dark " plumbeous-black " bases showing through slightly exceptat the borders of the thighs, where the hairs are pale cream buffthroughout. Specimens from La Palma, Michoacan, Mexico, averagea very little browner than two examined from Guadalajara. One inworn pelage from Lake Atitlan, Guatemala, is not appreciably differ-ent from topotypes. Northward from central Mexico to Arizonathere seems to be very little change in tint. A skin from Tamaulipas,Mexico, and others from Tucson and the Mexican border (monument77) of Arizona are scarcely distinguishable, and seem best referredto true velifer^ while a series from Roosevelt County, Ariz., con-tains specimens some few of which are nearly as pallid as inincautus. while one or two are about as dark as typical Mexican ex-amples, so that the series is taken to show that intergradation betweenthe typical form and the eastern M. velifer incautus occurs in thisregion.3Ieasuremenis.?For measurements see tables, pages 94 and 96.Ward (1904) presents a detailed study of the variations in pro-portions among a series of 167 bats of this species from Las Vegas,Vera Cruz. The forearm measurement averaged about 42 mm.Specimens examined.?Total numbei- 372, from the followinglocalities
:
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 91ARIZONA: Big Sandy Creek, 3 skins, IG ale. (U.S.N.M.) ; Ehrenburg, 1skin, 1 ale. (U.S.N.M.) ; Gila Bend, Marieopa County, 1 skin (U.S.N.M.) :Huachuca Mountains, 1 skin (F. M.), 2 ale. (M. C. Z.) ; Montezuma Well,2 skins (U.S.N.M.) ; Nantan Plateau, 1 skin (U.S.N.M.) ; Roosevelt, 10skins, 39 ale. (U.S.N.M.) ; Santa Rita Mountains, 1 skin (F. M.) ; Tucson,1 skin, 1 ale. (U.S.N.M.) ; Tucson, 20 miles southwest, 1 skin (U. C.) ; noexact locality, 1 ale. (U.S.N.M.).CALIFORNIA: Needles, 3 skins (U. C).DURANGO: Huasamota, 1 ale. (U.S.N.M.); Rio Sestin, 2 skins (A. M.N. H.) ; San Gabriel, 11 ale. (A. M. N. H.).?" ECUADOR": 2 ale. (B. M.).GUANAJUATO: Guanajuato, 25 ale. (U.S.N.M.).GUATEMALA: Lake Atitlan, Panajaehel, 1 skin (F. M.) ; Ciudad Vieja, 5ale. (B. M.), 1 ale. (U.S.M.C.) ; Sierra Sta. Elena, 1 ale. (F. M.).HIDALGO: Tulancingo, 1 skin, 3 ale. (U.S.N.M.).JALISCO: Atenquique, 12 skins (A. M. N. H.) ; Guadalajara, 2 skins(U.S.N.M.), 2 ale. (B. M.), 3 skins including type (A. M. N. M.) ; LakeZacoalco, 4 ale, (Paris), cotypes of jaliscensis, examined by Miller in1904; Las Canoas, 5 skins (A. M. N. H.) ; San Marcos, 1 ale. (A. M.N. H.) ; Tonila, 1 skin (A. M. N. H.) ; Zavala, 7 ale. (A. M. N, H.).LOWER CALIFORNIA: San Bernardino Ranch. 4 skins, 1 ale. (U.S.N.M.).MEXICO STATE: Ixtlapalapa, 2 ale. (U.S.N.M.) ; Lerma, 1 ale. (U.S.N.M.) ;Mexico, 1 skin (U.S.N.M.) ; Valley of Mexico, 2 ale. (M. C. Z.).MICHOACAN: Acambaro, 1 ale. (U.S.N.M.) ; El Molino, 1 ale. (B. M.) ;Lake Chapala, La Palma, 17 skins, 72 ale. (F. M.), 6 ale. (U.S.N.M.) ;Negrete, 2 ale. (U.S.N.M.); Patzcuaro, 3 skins, 8 ale, (U.S.N.M.), 5skins, 3 ale. (F. M.).MORELOS: Morelos, 10 ale. (U.S.N.M.).NEW MEXICO: Fort Wingate, 1 ale. (U.S.N.M.).OAXACA: Isthmus of Tehuantepec, 1 ale. (B. M.) ; Tehuantepee, 1 ale.(B. M.).PUEBLA: Esperanza, 2 ale. (F. M.).QUERETARO: Jalpan, 2 ale. (U.S.N.M.).SAN LUIS POTOSI: Ahualuleo, 1 ale. (U.S.N.M.); Rio Verde, 1 ale,(U.S.N.M.).SONORA: San Bernardino Ranch, 4 skins, 1 ale. (U.S.N.M.).VERA CRUZ: Las Vegas, 1 ale. (U.S.N.M.) ; Orizaba, 2 skins (U.S.N.M.).5 skulls, 28 ale. (F, M.), 4 ale. (A. N. S. P.) ; Xuchil, 2 ale. (F. M.).ZACATECAS: Hacienda San Juan Capistrano, 2 skins (U.S.N.M.).Remarks.?Typical Myotis velifer is widely distributed on theMexican table-land. The most southern record seems to be that ofa specimen from Lake Atitlan, Guatemala, altitude 5,150 feet. Skinsfrom central and northern Mexico are fairly uniform in tint, showinglittle or no approach to the pallid form incoMtus until the Texasborder is reached. A single skin from near sea-level at Soto LaMarina, Tamaulipas, though not quite so dark as the average of truevelifer is yet not so pallid as typical incautti^. Others from northernSonora (San Bernardino Ranch at junction of the river of that namewith the Cajon Bonito Creek) on the Mexican boundary are nearlytypical, as are also three others from Big Sandy Creek, in western
92 BULLETIN" 144, UNITED STATES NATIONAL MUSEUMArizona, a skin from Tucson, Ariz., and one each from the HuachucaMountains and the Santa Rita Mountains of the southeastern part ofthat State. It is in eastern Arizona and western Texas that inter-gradation takes place with the pale form, incautus. In this regionthe range of variation in tint among individuals from one localityis sometimes so great that it would be possible without violence toassign the extremes of a series to one or other of the two forms. Thusof two excellent skins from near the junction of the Pecos and RioGrande Rivers, Tex., one is as dark as the average true velifer, whilethe other is as pale as typical incautus. Again, a series of 10 skinsfrom Roosevelt, Ariz., is about intermediate; for while the darkestindividuals are of nearly the same dark tone as typical velifer, themajority are paler, though none is quite as pallid as incautus.Since the two forms hardly differ except in tint, it is often not pos-sible to assign alcoholic specimens with certainty to either, but indoubtful cases we have been guided by the indications of the seriesof skins.There are two alcoholic specimens in the collection of the BritishMuseum labelled " Ecuador," that were received many years agofrom the Fraser collection. These appear to be perfectly typicalMyOtis "velifer^ but since there is no exact locality given and no otherspecimens are known from any part of South America, it seemslikely that there has been some mistake, and that the locality " Ecua-dor " was supplied by inference, since Louis Eraser's name is so com-monly associated with the zoology of that Republic.MYOTIS VELIFER INCAUTUS (J. A. Allen)Vespertilio sp., J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 8, p. 71, April 22,1896.Vespertilio incautus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 8, p. 239,November 21, 1896.
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol.30, p. 307, December 27, 1901.Vlespertilio] albescens? H. Allen, in Miller, North Amer. Fauna, No. 13, p. 59,October 16, 1897 (Texas specimens).Myotis velifer Miller, North Amer. Fauna, No. 13, p. 56, October 16, 1897 '(part).
?
Strecker, Check-List Mamm. Texas, The Baylor Bulletin, BaylorUniversity, "Waco, Texas, vol. 29, No. 3, p. 9. Au^ist, 1926.Myotis incautus Miller, Proc. Biol. Soc. Washington, vol. 15, p. 155, June 20,1902.?J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 19, p. 611, November12, 1903.?Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 31, p. 121,August 27, 1903.
?
Trouessart, Catal. Mamra. viv. foss., suppl., p. 92, 1904.
?
Bailey, North Amer. Fauna, No. 25, p. 209, October 24, 1905.
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Elliot, CheckList Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6,p. 473, 1905.?HovPELL, Proc. Biol. Soc. Washington, vol. 22, p. 46, March 10,1909.?Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus.,No. 79, p. 54, December 31, 1912; List North Amer. Recent Mamm. 1923,Bull. U. S. Nat. Mus., No. 128, p. 67, April 29, 1924.?Steecker, Cheek-ListMamm. Texas, The Baylor Bulletin, Baylor University, Waco, Texas, vol.29, No. 3, p. 9, August, 1926.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 93Type locality.?San Antonio, Bexar County, Tex.Type specimen.?Adult male, skin and skull, No. -H-H-f Amer. Mus.Nat. Hist., collected at San Antonio, Tex., October 10, 1896, by H. P.Attwater.Distribution.?Open arid country from Texas and New Mexiconortheastward to Kansas; south in Mexico as far as Durango. (Seemap 4, p. 87.)Diagnosis.?Size maximum for the species and for the Americanmembers of the genus (pi. 1, p. 7, fig. 4) ; greatest length of skull15.8 to 17.6 mm.; color pallid.Description.?Compared with that of true Myotis velifer the furof M. V. incautits is usually of a much paler tint in mature animals.A paratype is nearly "tawny olive," with the bases of the hairs notquite so dark as in M. velifer velifer; below, the pale tips of the hairsare a trifle clearer, essentially the cartridge buff of Ridgway (1912).Measurements.?For measurements see tables, pages 95 and 96.Specimens examined.?Total number 80, from the following locali-ties :KANSAS: Medicine Lodge, 1 skin (U.S.N.M.) ; Sun City, 11 skins, 7 ale.(U.S.N.M.).NEW MEXICO: Carlsbad, 14 skins (U.S.N.M.).OKLAHOMA: Cache Creek, Comanche County, 2 skins (U.S.N.M.) ; FortReno, 3 ale. (U.S.N.M.).TAMAULIPAS: Soto la Marina, 1 skin (U.S.N.M.).TEXAS: East Painted Cave, 1 ale. (U.S.N.M.) ; Japonica, 2 skins, 5 ale.(U. S. N. M.) ; Lampasas, 1 ale. (U.S.N.M.) ; Langtry, 2 skins (U.S.N.M.) ;New Braunfels, 1 ale. (U.S.N.M.) ; Pecos River mouth, 4 ale. (U.S.N.M.) ;San Antonio, 1 skin (U.S.N.M.), 4 skins, including type (A. M. N. H.) ;Sonora, 10 ale. (U.S.N.M.) ; Toyahvale (Davis Mountains), 7 ale.(U. M.) ; Uvalde, 1 ale. (U.S.N.M.).UTAH: Thistle Valley, 2 ale, young (U.S.N.M.).MYOTIS VELIFER PENINSULARIS MillerMyotis peninsularis Miller, Ann. and Mag. Nat. Hist., ser. 7, vol. 2, p. 124,August, 1898.
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Trotjessart, Catal. Mamm. viv. foss., p. 1285, 1899.
?
Milleband Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p. 255, December 27, 1901.
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Elliot, Land and Sea Mamm. Middle Amer., Field Columb. Mus., publ. 95,zool. ser., vol. 4, p. 573, 1904.
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Tbouessart, Catal. Mamm. viv. foss., suppl.,p. 92, 1904.
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Elliot, Check List Mamm. North Amer., Field Columb. Mus.,publ. 105, zool. ser., vol. 6, p. 475, 1905.
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Miller, List North Amer. LandMamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 55, December 31, 1912 ; ListNorth Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 67,April 29, 1924.Type locality.?San Jose del Cabo, Lower California, Mexico.Type specimen.?Adult female, skin and skull, No. 98.3.1.59 BritishMuseum (Natural History), collected at San Jose del Cabo, LowerCalifornia, Mexico, August 12, 1896, by Loye Miller. Dane Coolidgecollection, No. 718.
94 BULLETIN 144, UNITED STATES NATIONAL MUSEUMDistribution.?Southern end of the Lower California peninsula.(See map 4, p. 87.)Diagnosis.?Size minimum for the species; forearm usually lessthan 40 mm.; gieatest length of skull 14.2 to 15.6 mm.; color pallid.Color.?Two color phases are exhibited in the original series : Oneduller, the other more intense; the duller specimens are not dis-tinguishable except by size from dull specimens of the large Myotisvelifer incautus, but the brighter-colored individuals are more red-dish above, approaching " clay color."Skidl and teeth.?Except for their smaller size, the skull and teethare counterparts of those of typical Myotis velifer.Masureinents.?For measurements see tables, pages 95 and 97.
^Specinnens examined.?Total number 25, from the followinglocalities
:
LOWER CALIFORNIA: La Paz, 1 ale. (U.S.N.M.) ; Miraflores, 5 skins(A. M. N. H.) ; San Jos6 del Cabo, 5 skins (B. M.), 5 skins (U.S.N.M.) ;Santa Anita, 3 ale. (U.S.N.M.) ; Santiago, 6 skins (C. A.).Remarks.?It was supposed at first that " the extremely short tailof the new species," Myotis peninsularis, was "its most stronglymarked character," as seemed to be indicated by the collector's fleshmeasurements. But this apparent peculiarity was doubtless due tosome peculiar way of measuring, for in alcoholic specimens of theanimal no such feature appears (ratio of tail to head and body in 4specimens in alcohol 82.2, as compared with 81.9 in 10 from VeraCruz and 79.9 in 10 from Arizona).One skin in the original series (93553 U.S.N.M.) is albinistic witha considerable sprinkling of white hairs on the nape and especiallyon the lower back and the sides, where a noticeable patch of whiteis seen just behind each shoulder.External measurements of Myotis velifer
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 95External measurements of Myotis velifer?Continued
Locality
96 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotic velifer
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 97Cranial measurements of Myotis velifer?Continued
Locality
98 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
List. Mamm. North Amer., siippl., p. 154, 1917.
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Miller, List North Amer.Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 67, April 29, 1924.Myotis hayleyi Lydekker, ZooI. Record, 1909, Mammalia, p. 59, 1910 (misprint).Type locality.?^West side of the Colorado River, 10 miles aboveNeedles, San Bernardino County, Calif.Type specimen.?Adult male, skin and skull. No. 137098 UnitedStates National Museum (Biological Survey collection), collected onthe west side of the Colorado River, 10 miles above Needles, Calif.,May 14, 1905, by N. Hollister. Original No. 2237.
Map 5.
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Distribution of Myotis occcltdsDistrihution.?From central and soutliAvestern New Mexico, across ^the southern half of Arizona southwestward on the Mexican highlands as far at least as southern Sinaloa.Diagnosis.?Size and general external characters about as in Myotisvelifer peninisularis (average ratio of tail to head and body in 7specimens, 81.8), but color strongly ochraceous and hairs of back withobviously burnished tips which impart to the fur a gloss like thatpresent in M. lucifugus; skull with rostrum even more enlarged relatively to area of brain case than in M. velifer (compare figs. 8 and 6of pi. 1, p. 7) ; brain case lower and more flattened than in M.hjucifvigus lucifug^is, not higher and narrower as in M. iMlifer; cheek
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 99teeth enlarged as in M. velifer^ but small premolars showing an obvi-ous tendency toward crowding and elimination.Ears, merribraTies, and feet.?The general features of external formdo not differ appreciably from those of Myotis velifer. Averageratio of foot to tibia in 10 specimens, 51.9.Fur and color.?The fur is more full and soft than that of Myotisvelifer., its quality essentially as in M. hicifug'us. Tips of hairs onupper surface of body distinctly burnished.Color above bright ochraceous tawny. Below ochraceous buff,sometimes with a grayish tinge ; bases of the hairs both above andbelow blackish-slate. Ears and membranes brownish. This phase isbrightest in the series of six skins from near Yuma, in the collectionof the Museum of Vertebrate Zoology, University of California. Aduller, olive phase essentially like the ordinary coloring of typicalMyotis iMcifugus also occurs in specimens from parts of Arizona andNew Mexico. Immature individuals may be nearly wood-brownabove, and grayish below. The ears and membranes of some indi-viduals are blackish.Skull.?The skull is well differentiated from that of any otherAmerican species of Myotis by the combination of low, flattenedbrain case and enlarged rostrum. When the skull is viewed fromabove (pi. 1, p. 7, fig. 8) the area of the rostrum as compared withthat of the brain case is greater than in any other American memberof the genus yet discovered. This character is obvious when theskull is compared with that of Myotis velifer (pi. 1, p. 7, fig. 6) ; itis very conspicuous on comparison with M. lucifugus (pi. 1, p. 7, fig.12) and still more so on comparison with M. volans., one race ofwhich occurs in the same region as M. occmltus and bears a super-ficial external resemblance to it. Breadth of rostrum across caninesdecidedly greater than interorbital constriction. Distance from an-terior border of alveolus of inner incisor to narrowest region ofinterorbital constriction frequently exceeding that from narrowestpart of constriction to lambda. Sagittal crest always present inadults but rarely as high as in average specimens of Myotis velifer.Teeth.?The larger cheek teeth of Myotis occultus present the samefeature of unusual robustness which characterizes those of M. veliferand distinguishes these two species from the other American mem-bers of the genus. Secondary cusps and ridges as in Myotis velifer.The small premolars are peculiar in the tendency which they showtoward crowding and elimination. The second premolar in theupper jaw {p^), rarely found to be missing in other AmericanMyotis., is more often absent than not (seventeen times in 24 speci-mens) ; and if it is present, it is usually pushed out of the tooth rowinto the angle between the first and fourth premolars, where it is notvisible in side view. In the lower jaw the second premolar is nor-
100 BULLETIN 144, UNITED STATES NATIONAL MUSEUMmally present (lacking on both sides in 2 out of 24 skulls), but itis crowded inward from the tooth row, or sometimes stands quite inthe row though closely pressed between the two other premolars.Measurements.?For measurements see tables, pages 100 and 101.Specimens examined.?Total number 27, from the following locali-ties:ARIZONA: Fort Verde, 1 skin (A.M.N.H.) ; Fort Whipple, 1 skin (U.S.N.M.); Lakeside, 3 ale. (U.S.N.M.) ; Mojave Desert, 1 skin (U.S.N.M) ;Montezuma Well, 1 skin (U.S.N.M.) ; San Francisco Mountain, 1 ale.(U.S.N.M.).CALIFORNIA: Imperial County, ten miles above Needles, 1 skin, 1 ale.(U.S.N.M.), type of occultus; 4 miles south of Potholes, 1 skin (U.C.) ;5 miles northeast of Yuma, 5 skins (U.C).NEW MEXICO : Luna, 3 skins (U.S.N.M.) ; Ruidosa, 1 skin, type of J)aileyi,2 ale. (U.S.N.M.) ; Zuni Mts., Bear Ridge, 4 skins (U.S.N.M.).SINALOA: Escuinapa, 1 skin (A.M.N.H.).Remm^hs.?The combination of large rostrum, flattened brain caseand burnished fur differentiates Myotis occultus very sharply fromother American species. Unusually robust cheek teeth are also foundin M. velifer but in this animal the brain case is high and the rostrumis less enlarged. Elimination of the second upper unicuspid toothis here more frequent than in any other known Americaii species.Externally Myotis occultus differs from M. velifer in the glossy fur,ochraceous color of the back, and conspicuously buff underparts.The series of specimens now available shows that the characterswhich Hollister regarded as distinguishing Myotis ha/Ueyi from M.occultus lie within the range of individual variation.External measurements of Myotis occultus
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYtJf 101Cranial measurements of Mi/otis occultus
Locality Number
102 BULLETIN 144, UNITED STATES NATIONAL MUSEUM I
straight, its outer edge with a small rounded basal lobe, succeeded bya shallow notch, above which it is nearly straight to the acute tip.Wing and rtiemhraiies.?^Wing membranes from the base of thetoes. Metacarpals 3, 4, and 5 practically equal in length (in an ab-normal individual nic 4 of the right side is 2 mm. shorter than 3and 5). When folded, the third metacarpal falls 2 mm. short ofthe elbow. Fourth and fifth fingers of nearly equal length; takingthe third finger as 100, the fourth is 84, and the fifth is 82 (61:52:51 mm.). The edge of the uropatagium is essentially bare,though with the aid of a lens, a few short hairs may sometimes be
Map 6.
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Distribution of Myotis kebnii : 1, M. keenii kebnh ; 2, M. kebniiSEPTKNTRIONALIS
seen at irregular intervals, along its border, occurring singly, andnever forming a fringe. Terminal caudal vertebra and most of thepenultimate vertebra free from membrane.Foot.?T\iQ foot is usually a little more than half the length of thetibia; in 6 specimens from the Pacific coast the ratio of foot totibia averages 51.2; in 10 from the northeastern part of the animal'srange it averages 50.4; and in a series containing the same numberfrom Illinois, Indiana, Wisconsin, Tennessee, and Missouri it aver-ages 50.5; in exceptional individuals it falls as low as 45.2. Calcarlong, slightly thickened at the base, terminating in a small lobulesituated about one-half the distance from the foot to the tip of tail.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 103Fur and colm\?Pelage fine and full, the longest hairs on the backabout 9 mm. in length. The quality and distribution of the fur areessentially as in Myotis ludfugus^ but the longer hairs tend to beless burnished; consequently the back is usually less glossy, a char-acter not always obvious in single specimens but quite apparent whenseries of skins of the two animals are compared. The general colorapproximates that of two of the races of Myotis lucifugus?thetypical form in the east, and M. Jucifugtis ala-scensis on the northwestcoast.Skull.?The skull is narrower in proportion to its length than thatof Myotis litcifug'us.i so that the distance from the last molar to thetip of the hamular process exceeds the distance between the lastmolars instead of equalling it as it does in M. lucifugus, while thelength of the maxillary tooth row (front of canine to back of lastmolar) very slightly exceeds the greatest palatal width including theupper molars whereas in M. lucifugus the reverse is true. In dorsalview the less robust rostrum is usually an evident feature of theskull as compared with that of M. lucifugus, and the width of theinterorbital constriction less evidently exceeds that of the rostrumacross roots of canines (compare pi. 1, p. 7, figs. 13 and 12). Fromthe skull of Myotis evotis that of M. keemi is distinguishable by itssmaller size and relatively smaller auditory bullae.Teeth.?As compared with the teeth of Myotis lucifugus, those ofM. keenii may in most instances be distinguished by rather obviouspeculiarities in the structure of the upper molars. The protoconuleand metaloph are less developed, the metaloph usually confined tothe bottom of the valley between the bases of the hypocone and meta-cone. The main cusps on the lingual side of the crown tend to beless robust, a feature which is especially noticeable in the protocone;this cusp, when compared with the protocone of Myotis lucifuguslucifugus, having a peculiar shrunken appearance, difficult to de-scribe, but appreciable after comparison of a few specimens of thetwo animals. The cingulum, on the other hand, is better developed,rather frequently extending uninterruptedly around the entire baseof the protocone, a condition which occurs in 13 out of 44 skullsselected at random, while it is not once found in 30 skulls of M. luci-fugus lucifugus. The structure of the teeth in Myotis keenii isessentially similar to that in M. evotis, but the size of the molars isobviously less than in the longer-eared animal, the crown of jn?usually about 1.25 to 1.35 by 1.60 to 1.75 mm. instead of 1.45 to 1.50by 1.95 to 2.00 mm., a difference appreciable to the unaided eye, evenwithout direct comparison of specimens.Remarks.?This bat may readily be distinguished from Myotislucifugus, with which it is everywhere associated, by the approxi-mately equal instead of obviously graded metacarpals, nearly equal
104 BULLETIN 144, UNITED STATES NATIONAL MUSEUMfourth and fifth fingers, the longer ears and tragus, less glossy fur,more buffy color, the size and proportions of the skull, and the morereduced condition of the smaller molar cusps. From Myotis evotis,together with which it occurs in the western part of its range, it isdistinguished by slightly smaller size, less elongated ears, and smallerteeth. MYOTIS KEENII KEENII (Merriam)VespertiUo subulatus keenii Merriam, Amer. Nat., vol. 29, p. 860, September,1895.Myotis lucifugus alascensis Miller, North Amer. Fauna, No. 13, p. 63, October16, 1897 (part, specimen from Wrangell, Alaska).Myotis subulatus keenii Milleh?, North Amer. Fauna, No. 13, p. 77, October 16,1897.
?
Trouessart, Catal. Mamm. viv. foss., p. 12S4, 1899.
?
Elliot, Synops.Mamm. North Amer., Field Colum. Mus., publ. 45, zool, ser., vol. 2, p. 405,March, 1901 ; List Land and Sea Mamm. North Amer., Field Columb.Mus., publ. 57, zool. ser., vol. 2, p. 518, June, 1901.
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Osgood, North Amer.Fauna, No. 21, p. 37, September 26, 1901.
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Miller and Rehn, Proc. BostonSoc. Nat. Hist., vol. 30, p. 258, December 27, 1901.?Trouessart, Catal.Mamm. viv. foss., suppl., p. 94, 1904.
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Elliot, Check List Mamm. NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 478, 1905.
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Lyonand Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus.,No. 62, p. 271, January 28, 1909.
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Miller, List North Amer. Land Mamm.1911, Bull. U. S. Nat. Mus., No. 79, p. 58, December 31, 1912; List NorthAmer. Recent Mamm. 1923, Bull, U. S. Nat Mus., No. 128, p. 72, April 29,1924.Type locality.?Massett, Graham Island, Queen Charlotte Islands,British Columbia, Canada.Type spechnen.?Young adult female (in alcohol), No. 72922 U. S.Nat. Mus. (Biological Survey Collection), collected at Massett,Graham Island, Queen Charlotte Islands, British Columbia, Canada,1894, by J. H. Keen.Distribution.?Humid northwest coast region from northern Brit-ish Columbia and southeastern Alaska to northwestern Washington.(See map 6, p. 102.)Characters.?A saturate, northwest-coast form ; ears conspicuouslyblack.Color.?Upperparts a rich glossy brown, nearly the Dresdenbrown or buckthorn brown of Ridgway, the fur very dark slatybrown beneath the surface; underparts distinctly more buffy thanback; ears black; membranes very dark brown or black.Measurments.?For measurements see tables, pages 109 and 110.SpeciTTiens examined.?Total number 7, from the following localities
:
ALASKA: Wrangell, 1 skin (U.S.N.M.).BRITISH COLUMBIA: Massett, Queen Charlotte Islands, 3 ale.(U.S.N.M.) ; Telegraph Creek, 1 ale. (U.S.N.M.).WASHINGTON: Lake Cushman (Mason County), 1 skin (U. M.) ; SolDue Hot Springs (Clallam County), 1 skin (U.S.N.M.).
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 105Remarks.?In color the typical race of Myotis keenii appears to benot essentially different from M. evotis evotis and M. lucifugusalascensis. The longer ears serve to distinguish it readily from thecoast race of M. liicifugus, and the smaller upper molars (crownof m^ 1.30 by 1.75 mm. in type, 1.35 by 1.60, and 1.30 by 1.T5 mm.,respectively, in two topotypes) are diagnostic as compared withthe local form of M. evotis (crown of m^ usually about 1.45 to 1.50by 1.95 to 2.00 mm.).MYOTIS KEENII SEPTENTRIONALIS (Trouessart)
Vespertilio subulatus H. Allen, Monogr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 51, figs. 45-46, June, 1864 (not of Say, 1823, or LeConte,1855).?DOBSON, Catal. Chiroptera Brit. Mus., p. 324, 1878.?True, Proc.U. S. Nat. Mus., vol. 7 (1884), p. 603, 1885.?Milled, Proc. Boston Soc. Nat.Hist., vol. 28, p. 39, April, 1897.Vespertilio gryphus lucifugus Evebmann and Butleb. Proc. Indiana Acad. Sci.,1893, p. 134, 1894.Northern form of Vespertilio gryphus H. Allen, Monogr. Bats North Amer.,Bull. U. S. Nat. Mus., No. 43 (1893), p. 80, March 14, 1894.Vespertilio gryphus var. septentrionalis Trottessabt, Catal. Mamm. viv. foss.,p. 131, 1897.Vespertilio gryphus septentrionalis Milleb and Rehn, Proc. Boston Soc. Nat.Hist., vol. 30, p. 308, December 27, 1901.Myotis subulatus Miller, North Amer. Fauna, No. 13, p. 75, figs. 13a, 15&,October 16, 1897 (not of Say, 1823).
?
Troxtessabt, Catal. Mamm. viv. foss.,p. 1284, 1899.?MnxEE, Prelim. List New York Mamm., Bull. New YorkState Mus., vol. 6, No. 29 (October, 1869), p. 366, November 18, 1899; KeyLand Mamm. Northeastern North Amer., Bull. New York State Mus., vol. 8,No. 38 (October, 1900), p. 149, November 21, 1900.
?
Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45. zool. ser., vol. 2, p. 40.5, March1901; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ.57, zool. ser., vol. 2, p. 518, June, 1901.
?
Miller and Rehn, Proc. BostonSoc. Nat. Hist., vol. 30, p. 257, December 27, 1901.
?
Elliot, Land and SeaMamm. Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p.580, 1904.
?
Trouessabt, Catal. Mamm. viv. foss., suppl., p. 93, 1904.
?
Elijot,Check List Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser.,vol. 6, p. 478, 1905 ; Catal. Mamm. Field Columb. Mus., Field Columb. Mus.,publ. 115, zool. ser., vol. 8, p. 504, 1907.?G. M. Allen, Bull. Mus. Comp.Zool., vol. 52, p. 45, July, 1908.?Hahn, The Mamm. of Indiana, .33d Ann.Rep. Dept. Geol. and Nat. Resources of Indiana, 1908, p. 623, 1909.?Cory,Mamm. Illinois and Wisconsin, Field Mus. Nat. Hist., publ. 153, zool. ser.,vol. 11, p. 460, 1912.?BAILEY, Proc. Biol. Soc. Washington, vol. 36, p. 137,March 28, 1923.Myotis subulatus subulatus Miller, List North Amer. Land Mamm. 1911, Bull.U. S. Nat. Mus., No. 79, p. 58, December 31, 1912 ; List North Amer. RecentMamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 72, April 29, 1924.?Bailey,North Amer. Fauna, No. 49 [December, 1926], p. 216, January 8, 1927(North Dakota).Type loccdity.?Halifax, Nova Scotia.58518?28 8
106 BULLETIN 144, UNITED STATES NATIONAL MUSEUMType specinien.?Harrison Allen ^^ listed eight individuals of his
"Var. (5) Northern form of VespertUio gryphvs'''' which served asthe basis for Trouessart's name septentrionalis. As no less than fiveof these were from Nova Scotia one of the specimens taken by DoctorGilpin at Halifax (No. xgWs^ U. S. Nat. Mus.) may appropriately bechosen as the lectotype.Distribution.?Eastern North America from Newfoundland andQuebec south to Tennessee and South Carolina; west to NorthDakota, Missouri, and Arkansas. (See map 6, p. 102.)Gharacters.?Color approximately like that of Myotis lucifugushicifugus, not darkened as in the typical northwest-coast rate; earsbrown.Color.?The color is not essentially different from that of M. luci-fugus lucifugus in the bronzy phase, but the brown hair tips areneither so long nor so glossy. Consequently series of skins of thetwo species from the same region have an obviously unlike appear-ance due in part to the slight difference in color and in part to theless complete covering over of the slaty under fur in M. keeniiseptentrionalis.Measurements.?For measurements see tables, pages 109 and 110.Specimens examined.?Total number 169, from the followinglocalities
:
ARKANSAS: 2 ale. (M. C. Z.) ; Delight, 1 skin (U.S.N.M.).CONNECTICUT: Mount Carmel, 1 ale. (M. C. Z.) ; New Haven, 1 ale.(U.S.N.M.).DISTRICT OP COLUMBIA: Washington, 2 skins, 1 ale. (U.S.N.M.).GEORGIA: Young Harris, 1 skin (U.S.N.M.).ILLINOIS: Chieago, 1 ale. (U.S.N.M.) ; Wabash County, 1 ale. (U.S.N.M.).INDIANA: Mitchell, 4 skins (U.S.N.M.) ; Terre Haute, 1 ale. (U.S.N.M.) ;Wheatland, 1 ale. (U.S.N.M.).KENTUCKY: Burchette's Cave, 2 ale. (M. C. Z.) ; Eubanks, 2 ale.(U.S.N.M.).LAKE SUPERIOR: 3 ale. (M. C. Z.).MAINE: Eastport, 2 ale. (U.S.N.M.), 2 ale. (M. C. Z.) ; Norway, 1 ale.(M. C. Z.) ; St. George, 1 skin (M. C. Z.).MARYLAND: Cabin John Bridge, 1 skin (U.S.N.M.); Muirkirk, 1 ale.(U.S.N.M.) ; Plummer Island, 5 skins (U.S.N.M.).MASSACHUSETTS: Brookline, 1 ale. (M. C. Z.) ; Feltonville, 5 ale.(M. C. Z.) ; Harvard, 1 skin (M. C. Z.) ; Shirley, 1 skin (M. C. Z.) ;Springfield, 1 ale. (M. C. Z.) ; Ware, 1 ale. (M. C. Z.) ; Wilmington,1 skin (U.S.N.M.).MICHIGAN: Boynes Falls (Charlevoix County), 1 ale. (U. M.) ; Chip-pewa County, 2 ale. (U. M.) ; Detroit, 1 ale. (A. N. S. P.) ; DouglasLake (Cheboygan County), 1 skin (U. M.) ; Isle Royal, 3 skins (U. M.) ;Macinac Island, 3 skins (B. M.) ; Porcupine Mountains (OntonagonCounty), 3 skins (U. M.) ; South Lyon (Oakland County), 1 ale. (U. M.).
"Monogr. Bats North Amer. (1893), p. 81, Mar. 14, 1894.
AMEBICAlSr BATS OF THE GENERA MYOTIS AND PIZONYX 107MINNESOTA: Elk River, 2 skins (U.S.N.M.).MISSOURI: Marble Cave, 5 ale. (U.S.N.M.) ; Upper Missouri River, 1 skin,1 ale. (U.S.N.M.).NEW BRUNSWICK: Grand Manan, 3 ale. (M. C. Z.) ; no exact locality,1 skill (B. M.).NEWFOUNDLAND: Levels Hills, 2 skins (M. C. Z.) ; Romain's Brook,1 skin (M. C. Z.) ; Spruce Brook, 1 skin (M. C. Z.).NEW HAMPSHIRE: Bethlehem, 2 ale. (M. C. Z.).NEW YORK: Berkshire, 2 ale. (M. C. Z.) ; Fallsburg, 1 ale. (M. C, Z.) ;Hammondville, 12 ale. (U.S.N.M.) ; Hemlock Lake, 1 ale. (U.S.N.M.) ;Highland Falls, 1 skin (A. M. N. H.) ; Lake George, 4 ale. (U.S.N.M.) ;Long Island, 1 ale. (A. M. N. H.) ; Peterboro, 1 skin (U.S.N.M.) ; Stam-ford, 1 skin (A. M. N. H.).NORTH CAROLINA: Cherokee, 1 ale. (U.S.N.M.).NORTH DAKOTA: Fort Buford, 1 skin (U.S.N.M.); Fort Union, 1 ale.(U.S.N.M.).NOVA SCOTIA: Annapolis, 1 ale. (U.S.N.M.) ; Halifax, 2 ale. (U.S.N.M.).OHIO: Port Clinton, 1 ale, (U.S.N.M.).ONTARIO: Casselman, 2 ale. (U.S.N.M.); Lake of Bays, 2 skins(U.S.N.M.) ; Maiden, 1 skin (B. M.) ; Miehipicoten, 1 skin (U.S.N.M.) ;Ottawa, 1 ale. (U.S.N.M.) ; St. Catherine's, 1 ale. (U.S.N.M.).PENNSYLVANIA: Bradford, 1 ale. (U.S.N.M.) ; Center County, 1 skin(U.S.N.M.) ; Chester County, 1 ale. (A. N. S. P.) ; Markleton, SomersetCounty, 6 skins (U.S.N.M.) ; 1 ale. (B. M.) ; Round Island. ClintonCounty, 1 ale. (A. N. S. P.) ; Somerset County, 3 ale. (U.S.N.M.) ;Walnut Hill, Montgomery County, 2 ale. (A. N. S. P.).QUEBEC: Anticosti Island, 1 ale. (M. C. Z.), 1 ale. (U.S.N.M.) ; Godbout,5 ale. (U.S.N.M.) ; Lake Edward, 1 ale. (M. C. Z.).SOUTH DAKOTA: Elk Mountain, 1 ale. (U.S.N.M.).TENNESSEE: Bellamy's Cave, 3 ale. (U.S.N.M.); Hickman County, 2skins (A. M. N. H.).UNITED STATES: (no exact locality) 1 skin (U.S.N.M.).VERMONT: North Hartland, 1 skin (A. M. N. H.) ; Proctor, 2 skins(U.S.N.M.) ; Weathersfleld, 2 skins (M. C. Z.).VIRGINIA: Alexandria, 1 ale. (U.S.N.M.); Dismal Swamp, 1 ale.(U.S.N.M.); Endless Cavern, 1 skin (U.S.N.M.); Luray, 1 skin(U.S.N.M.) ; Orkney Springs, 1 skin, 1 ale. (U.S.N.M.).WEST VIRGINIA: Aurora, 2 ale. (U.S.N.M.); Braxton County, 1 skin(B. M.).WISCONSIN: Bayfield, 1 ale. (U.S.N.M.); Sumpter, 1 skin (F. M.) ;Gotham, 1 ale. (U.S.N.M.).Remarks.?Harrison Allen applied Say's name VespertiUo suhu-latus to the eastern long-eared Myotis in 1864. He was followedwithout question by Dobson in 1878. When writing the secondedition of his Monograph (1894) Allen, no longer regarding theanimal as specifically distinct from Myotis lucifugus., united it andM. lucifugus as "varieties" of a bat which he called Vespertiliogryphus. These he designated respectively as "Northern form ofVespertiUo gryphus " and " Vespertilio gryphus lucifugus. ''"' In orderto enter the supposed races in his Catalogus (1897) Trouessart Latin-
108 BULLETIN 144, UNITED STATES NATIONAL MUSEUMized the " Northern form " as " Vespertilio gryphus septentrionalis^''wrongly attributing to Allen the new name which he thus established.Later in the same year, Miller, again recognizing the animal as adistinct species, restored the specific name subulatus^ but called atten-tion to the fact that, while there was no doubt as to the identity ofthe bat which Harrison Allen had in hand in 1864, the contrary wastrue with regard to the one originally named by Say in 1823. Afterquoting Say's description in full Miller says (1897, p. 36)
:
While there is nothing in this account that refers unquestionably to thelonger eared of the two species of Myotis inhabiting the eastern United States,the name has passed current for this animal so long that, after careful con-sideration of all the evidence, I am unwilling to substitute for it Trouessart'sname septentrionalis, the only one unequivocally based on the species. Say'sVespertilio subulatus came from the Arkansas River, near the present townof La Junta, Colorado. The bats of this region are not well known, but atpresent Myotis evotis, M. californicus ciliolabrum, and M. lucifugus longicrusare the only members of the genus Myotis which may confidently be expectedto occur there. From the known range of Myotis subulatus to the north andwest, however, its regular occurrence in Colorado is' by no means impossible.* * * I am merely continuing the usage of the past thirty-four years, not,however, without grave misgivings that the reasons for so doing are in realityunsound.Miller supposed that the eastern long-eared Myotis might occur insoutheastern Colorado. It is still possible that such is the case; butthirty years of collecting since his paper was published have failedto prove that the animal's range extends nearer to the type localityof Vespertilio subulatus than Arkansas, Missouri, and North Dakota.Furthermore it can easily be shown that Say's description containsstatements which are not applicable to this animal but which dis-tinctly indicate a species whose range is known to include easternColorado, namely the small bat described as Vespertilio ciliolabrum.by Merriam in 1886. The eastern long-eared Myotis is a definitelybrown animal, somewhat lighter and more yellowish below than onthe back ; Say's bat was " dull cinereous " above and " yellowish-white " below. These expressions do not apply to the subulatus ofHarrison Allen and subsequent authors, but they might readily havebeen based on specimens of " ciliolabnmi " and notably on an individ-ual from Colorado Springs which Warren recorded in 1910.^^ Bya curious accident Warren misidentified his specimen as an exampleof the eastern long-eared Myotis. Consequently he called it Myotissubulatus, thinking that he had the subulatus of Harrison Allen,Dobson, and Miller. Actually he had Say's original subulatus, theanimal to which we now restore this long-misapplied name. Troues-
" Mnminals of Colorado, p. 275.
AMEEICAN BATS OF THE GENEKA MYOTIS AND PIZONYX 109
sart's name septentrionalis therefore becomes available for the easternbat which is now generally known as Myotis subulatus, but in theform Myotis keenii septentrionalis, Merriam having described thenorthwest-coast race of the species as VespertUio suhulatus kenmitwo years before Trouessart Latinized Harrison Allen's vernacularterm. External measurements of Myotis keemi
110 BULLETrN" 144, UNITED STATES NATIONAL MUSEUMCranial measurement of Myotis keenii
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX HIMYOTIS EVOTIS (H. Allen)(Synonymy under subspecies)Distribution.?From Vera Cruz and Lower California north tosouthern British Columbia, thence eastward in the arid portions ofthe United States to Colorado and North Dakota.Diagnosis.?Like Myotis keenii but slightly larger (forearmusually 37 to 41 mm.; greatest length of skull 15.0 to 16.4 mm.) andwith longer tail (average ratio of tail to head and body in 4 speci-mens from the northwest coast 90.3, in 10 from Colorado 96.9) andmuch larger ear (height from meatus usually more than 20 mm.).Auditory bullae slightly enlarged; a small and rather inconspicuoussagittal crest nearly always present in adults; crown area of uppercheek teeth distinctly greater than in M. keenii (crown of m ^ usuallyabout 1.45 to 1.50 by 1.95 to 2.00 mm.). A thin and inconspicuousfringe of minute hairs along the free border of the interfemoral mem-brane.Ears.?The ears are long, when laid forward extending 5-7 mm,beyond the tip of the nose (in alcoholic specimens). Tragus corre-spondingly long, about 47 per cent of the height of ear (9:19 mm.),slender, tapering; its inner border is nearly straight, bendingslightly outward at the tip. A small rounded lobe is at its outer base,succeeded by a distinct notch, above which is the broadest point.The outer margin is very slightly convex basally then very faintlyconcave toward the top. The intense black color of the ears usuallypersists in dried specimens.Wing and msTribranes.?Wing membrane from the base of the toes.Metacarpals 3, 4, and 5 usually about equal in length ; sometimes thethird is a very little longer than the others. When folded, the thirdmetacarpal reaches to within 2 mm. of the elbow. The fourth andfifth fingers are of nearly equal length; taking the third as 100, thefourth is 83, the fifth 82 (61:51:50 mm.). Terminal vertebra ofthe tail free. The free border of the uropatagium is thinly fringedabove and below with minute hairs; these are usually not obviouswithout a lens, yet they project beyond the edge and grow in littleclusters of from 2-4. This fringe is much less developed than inMyotis tJiysanodes, yet it is often perfectly definite; and even whenno true fringe is formed the presence of numerous scattered hairsalong the free border of the membrane is usually characteristic ofM. evotis as compared with M. keenii.Foot.?The foot is proportioned to the tibia nearly as in Myotiskeenii; that is, it is usually somewhat less than half as long as thetibia, but is not infrequently more than half as long. In 10 specimensfrom the northwest-coast reeion the ratio of foot to tibia averages
112 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
Map 7.
?
Distribution of Myotis evotis : 1, M. evotis kvotis ; 2, M. bvotisCHKYSONOTUS
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 11349.5 ; in 10 from Colorado it averages 48.3. Calcar long, ending in aminute lobule, sometimes with a rudimentary keel.Fur and color.?The pelage is full and soft, the individual hairsabout 10 mm. long on middle of back. General color a light brown,varying in the subspecies. Membranes and ears usually contrastedbrownish black.Skull.?The skull (pi. 1, p. 7, fig. 7) is essentially identical withthat of Myotis keenii except that it is slightly larger, the auditorybullae are a little more inflated, and the sagittal crest is more con-stantly developed (though rarely very conspicuous). Upper profilecurving very gradually from rostrum to summit of skull. Adultsshow a low but evident sagittal crest, most obvious where it crossesthe depression just in advance of the occiput. Lambdoid ridges lowbut evident, meeting at the occiput when fully developed. Viewedfrom above the brain case is oval, bulging posteriorly beyond thelambdoid ridges. The length of the maxillary tooth row, as in Myotiskeenii, equals or slightly exceeds the palatal width including themolars.Teeth.?In general the teeth resemble those of Myotis keenii., butthe upper molars show a stronger tendency to reduction of the second-ary cusps and ridges; their crown area is obviously greater, thecrown of m? usually about 1.45 to 1.50 by 1.95 to 2.00 mm. instead of1.25 to 1.35 by 1.60 to 1.75 mm. The protoconule is always low andsometimes practically absent; the metaloph is rudimentary, ofteneliminated completely so as to leave a perfectly smooth valley be-tween the bases of the protocone and metacone. Protocone and hypo-cone more robust than in M. keenii, rarely if ever shrinking awayfrom the margin of the crown sufficiently to permit the cingulum topass uninterruptedly around the base of the protocone. The smallupper premolars are seldom crowded but stand either directly in thetooth row or very slightly drawn in.Remarks.?^Among the American members of the genus, Myotisevotis is readily distinguishable by its moderate size combined withits greatly enlarged ears. Usually the ears, in well-prepared skins,are black in sharp contrast with the rather pale general color of thefur, a feature which when present, renders them doubly conspicuous.While obviously related to Myotis keenii this animal shows ahigher degree of specialization in the more enlarged ears and audi-tory bullae, and in the more pronounced tendency to reduction of thesecondary cusps and ridges of the first and second upper molars.The series of skins available for study is not very extensive, asMyotis evotis appears to be nowhere a common bat. So far as itgoes the material indicates that the species is divisible into tworather ill-defined geographical races, a darker typical form confined
114 BULLETIN 144, UNITED STATES NATIONAL MUSEUMto the humid northwest-coast region and a lighter form occupyingthe rest of the animal's range. That all of the lighter individualsrepresent a single geographical race appears to be highly improbable,but the material now at hand is not sufficient to form the basis forany satisfactory subdividing.Hahits.?The long-eared bat occurs over a wide area chiefly westof the Rocky Mountains in the United States, and apparently morein the thinly forested country than in wooded sections. Judgingfrom specimens in collections, it is unusual to find them except asscattered individuals. Gary (1911), however, obtained a series ofadult and immature examples, in August, 1906, from a deserted ranchhouse near Sunny Peak, Routt County, Colo. At other places in thesame State, he secured additional specimens about ranch buildingsbut failed to find them about ledges or cliffs. Where buildings arenot available this species probably finds shelter in crevices or per-haps in trees. At all events it seems not to be reported from caves.In Colorado, Young (1908) records it from an altitude of 7,000 feet.MYOTIS EVOTIS EVOTIS (H. Allen)Vespertilio evotis H. Allen, Monogr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 48, fiji. 42-43, June. 1864.?Dobson, Catal. Chiroptera Brit.Mus., p. 324, 1878.?True, Proc. U. S. Nat. Mus., vol. 7 (1884), p. 602,1885.?Lyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull.U. S. Nat. Mus., No. 62, p. 290, January 28, 1909.Vespertilio albescens evotis H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 89, March 14, 1894.?Tbouessabt, Catal. Mamm.viv. foss., p. 132, 1897.Vespertilio nitidus evotis H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), pi. 12 and explanation, March 14, 1894 (cor-rected in erratum slip).Myotis evotis Miller, North Amer. Fauna, No, 13, p. 77, October 16, 1897.
?
Tbouessabt, Catal. Mamm. viv. foss., p. 1284, 1899.
?
Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 406, March,1901 ; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ.57, zool. ser., vol. 2, p. 518, June, 1901.
?
Miller and Rehn, Proc. BostonSoc. Nat. Hist., vol. 30, p. 258, December 27, 1901.?Elliot, Land and SeaMamm. Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 574,1904.
?
Tbouessabt, Catal. Mamm. viv. foss., suppl., p. 94, 1904.
?
Elliot,Check List Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser.,vol. 6, p. 474, 1905.
?
Lyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat.Mus., Bull. U. S. Nat. Mus., No. 62, p. 290, January 28, 1909.?Miller, ListNorth Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 59,December 31, 1912 ; List North Amer. Recent Mamm. 1923, Bull. U. S. Nat.Mus., No. 128, p. 73, April 29, 1924.Type locality.?Puget Sound.^^Type specimen.?The original description was based on 13 speci-mens : 3 from " Upper Missouri " (the identity of which is uncertain)
;
"See H. Allen, Monogr. Bats North Amer., Bull. U. S. Nat. Mus., No. 43 (1893), pp.90-91. Mar. 14. 1894.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 115three from " Puget Sound " ; 2 from east of Colville, Wash. ; 1 fromMonterey, Calif. ; 2 from Cape St. Lucas, Lower California, Mexico
;
1 from New Mexico; and 1 without locality. All are therefore co-types, althou<;h some at least probably represent another species.In a subsequent paper, H. Allen (1894) described a specimen fromEaston, Wash., as representing what he considered " typical V. evotisof the monoorraph" (1864). Easton lies about 55 miles inland fromPuget Sound. We are informed by Mr. Vernon Bailey that thefaunal aspect of the locality is such that the occurrence of the coastform or of an intermediate between the coast form and chrysonotusis to be anticipated. The specimens described by H. Allen can notnow be determined as to subspecies.Miller (1897), overlooking this definite statement of H. Allen,selected Monterey, Calif., as the type locality, whereby the specimen(which can not now be found in the National Museum) from thatplace became, according to Lyon and Osgood (1909), the lectotype.However, H. Allen's earlier (1894) action can only be interpreted asfixing the type locality in western Washington. He originally hadthree specimens from this region, all taken at " Puget Sound." Oneof these is still in the National Museum (No. 5391). A specimenlabeled " Puget Sound " now in the collection of the Academy ofNatural Sciences of Philadelphia (No. 1808) was received manyyears ago from the Smithsonian Institution and is probably anotherof H. Allen's cotypes.Distribution.?Humid coast region from southern British Co-lumbia and western Washington southward to northwestern Cali-fornia. (See map 7, p. 112.)Intergradation with the subspecies chu^sonotus characterized by itsclearer, golden color takes place in northern California. A skin fromSisson, Siskiyou County, Calif. (No. 95454 U.S.N.M.), is practicallyindistinguishable in color from those taken in British Columbia eastof the coast range. These are somewhat paler than those from themore saturate area near the seaboard and are more or less inter-mediate between the typical race and M. evotis chrysonotus.Diagnosis.?General hue of upper parts dull, nearly the clay colorof Ridgway (1912), without golden cast.Color.?Color above, a uniform yellowish buff with a distinctolivaceous cast, nearly " clay color " (Ridgway, 1912), the tips of thehairs faintly glossy. A blackish-brown spot, usually rather illdefined, at the shoulder. Lower surfaces pale buffy. The bases ofthe hairs both above and below are plumbeous black. Ears blackish,membranes blackish brown.In skins from Washington and British Columbia the dull yellowtips of the long hairs of the back seem shorter than in the southern
116 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
race, so that the dark bases show through. The general effect is aslightly olivaceous and distinctly duller tint as compared with thebright golden brown of Californian specimens.Immature specimens have the pelage thinner, the dorsal surfacepaler buff, and the ventral surface clearer whitish.Measurements.?For measurements see tables, pages 120 and 121.Specimens exaimn?d.?Total number 42, from the following lo-calities :ALBEETA: Jasper, 1 skin (A. M. N. H.), not typical.BRITISH COLUMBIA: Cranbrook, 3 skins (C. B. Garrett) ; Rock Creek,1 skin (B. M.) ; Shuswap, 2 skins (U.S.N.M.) ; Vernon, 1 skin (A.N.S.P.) ;Victoria, 1 ale. (A.N.S.P.).CALIFORNIA: Beswick, Siskiyou County, 1 skin (U.S.N.M.); MountSanhedrin, Mendocino County, 2 skins (A.N.S.P.) ; Sisson, SiskiyouCounty, 1 skin (U.S.N.M.) ; South YoUa BoUy Mountain, 1 skin(U.S.N.M.) ; no exact locality, 1 skin (B. M.).OREGON: Blaine, Tillamook County, 1 skin (Alex. Walker coll.) ; BuckCreek, 1 ale. (U.S.N.M.) ; Fremont, 1 skin (U.S.N.M.) ; McKenzie Bridge,Lane County, 1 skin (U.S.N.M.) ; Tillamook, Tillamook County, 1 skin(Alex. Walker coll.) ; Wallowa Lake, 1 ale. (U.S.N.M.).WASHINGTON: Bartholomew, 4 skins (S. H. Lyman); Easton, 2 ale.(U.S.N.M.) ; Martin, 1 ale. (A.N.S.P.) ; Godman Springs, 9 skins (S. H.Lyman) ; Puget Sound, 2 ale. (B. M.), 1 ale, topotype (U.S.N.M.), 1 ale,topolype? (A.N.S.P.) ; South Touchet, 1 skin (S. H. Lyman.)MYOTIS EVOTIS CHRYSONOTUS (J. A. Allen)Vespertilio evotis H. Aixen, Monogr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 48, fig. 42-43, June, 1864 (part).
?
Cooper, Proc. CaliforniaAcad. Sei., vol. 4, p. 5, 1868.?J. A. Aixen, Bull. Amer. Mus. Nat. Hist., vol.5, p. 202, August 18, 1893.
?
Mesibiam, North Amer. Fauna, No. 3, p. 46,September 11, 1890.
?
Bryant, Zoe, vol. 1, p. 358, February, 1891.Myotis evotis Miller, North Amer. Fauna, No. 13, p. 77, October 16, 1897(part).?Meebiam, North Amer. Fauna, No. 16, p. 88, October 28, 1899.?Elliot, Field Columb. Mus., publ. 91, zool. ser., vol. 3, p. 320, March,1904.?Stone, Proc. Acad. Nat. Sci. Philadelphia, (July, 1904), p. 579,October 17, 1904.
?
Stephens, California Mammals, p. 267, 1906.
?
Elliot,Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115, zool.ser., vol. 8, p. 501, 1907.
?
Lyon and Osgood, List Type-Sp. Mamm. U. S. Nat.Mus., Bull. U. S. Nat. Mus., No. 62, p. 290, January 28, 1909.?Gary, NorthAmer. Fauna, No. 33, p. 207, August 17, 1911 (part).
?
Warren, Mammalsof Colorado, p. 276, 1910.?J. Grinnell, Proc. California Acad. Sci., ser. 4,vol. 3, p. 278, August 28, 1913.?Bailey, North Amer. Fauna, No. 35, p. 33,September 5, 1913.?Gary, North Amer. Fauna, No. 42, p. 26, October 3,1917.?H. W. Grinnell, Univ. California Publ. Zool., vol. 17, p. 293, Janu-ary 31, 1918.
?
Bailey, North Amer. Fauna, No. 49 [December, 1926], p.216, January 8, 1927 (North Dakota).Vespertilio chrysonotus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 8, p. 240,November 21, 1896. Miller, North Amer. Fauna, No. 13, p. 77, October16, 1897 (in synonymy of M. evotis).?Miller and Rehn, Proc. Boston Soc.Nat. Hist., vol. 30, p. 308. December 27, 1901.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 117Myotis mlcronyx Nelson and Goldman, Proc. Biol. Soc. Washington, vol. 22,p. 28, March 10, 1909 (Comondu, Lower California, Mexico).
?
Milled, ListNorth Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 59, De-cember 31, 1912.
?
Elliot, Check-List Mamm. North Amer., suppl., p. 156,1917.?Miller, List North Amer. Recent Mamm. 1923, Bull. U. S. Nat, Mus?No. 128, p. 73, April 29, 1924.Type locality.?Kinney Ranch, Sweetwater County, Wyo.Type specimen.?Adult female, skin only, No. 11645, AmericanMuseum of Natural History, collected at Kinney Ranch, Sweet-water County, Wyo., July 21, 1895, by W. W. Granger.Distributio7i.?Range of the species except the humid northwest-coast region. (See map 7, p. 112.)Diagnosis.?Color not darkened as in the typical race.Color.?Similar to Myotis evotis evotis, but general color averag-ing paler ; the tips of the hairs above tending to be golden or flaxen(pale buffy) those of the lower surfaces often nearly wliite, lackingthe buffy wash of typical evotis from the northern Pacific coast.Measurements.?^For measurements see tables, pages 120 and 121.Specimens examined.?Total number 80, from the following locali-ties:ARIZONA: Chiricahua Mountains, 1 skin (A. M. N. H.) ; HuachucaMountains, 1 skin (F. M.) ; Paradise, Cochise County, 2 skins (U.S.N.M.) ;San Francisco Mountains, 1 ale, 2 skins (U.S.N.M.) ; Santa CatalinaMills, 1 skin (A. M. N. H.) ; Springerville, 1 skin (U.S.N.M.) ; Tucson,1 skin (U.S.N.M.) ; White Mountains, Cooley's, 1 skin (A. M. N. H.) ;Williams, 1 ale. (U.S.N.M.).CALIFORNIA: Belmont, 1 skin (A. N. S. P.) ; Dulzura, 2 ale. (A. N. S. P.),1 skin, 2 ale. (A. M. N. H.) ; Inyo Mountains, 1 ale. (U. S. N. M.) ; MountTallac, 1 skin (M. C. Z.) ; Mount Whitney, 1 skin (F. M.) ; OwensLake, 1 ale. (U.S.N.M.) ; San Bernardino Mountains, 1 skin (U.S.N.M.) ;San Joaquin River, 1 ale. (U.S.N.M.) ; San Rafael, 1 skin (B. M.) ; TwinOaks, 1 ale. (U.S.N.M.).CHIHUAHUA: San Luis Mountains, 1 skull (U.S.N.M.).COLORADO : Ashbaugh's Ranch, Montezuma County, 1 skin, 1 ale.(U.S.N.M.) ; Coventry, 5 ale. (U.S.N.M.) ; Dolores, 1 ale. (U.S.N.M.) :Loveland, 1 ale. (U.S.N.M.) ; Snake River, 6 skins, 3 ale., 1 skull(U.S.N.M.).IDAHO: Albion, 2 skins (U.S.N.M.) ; Tamarack, 1 skin (U.S.N.M.).JALISCO: Los Masos, 1 skin (A. M. N. H.).LOWER CALIFORNIA: Comondu, 1 skin, type of micronyx (U.S.N.M.).MONTANA: Big Belt Mountains, 4 miles south of Fort Logan, 1 skin(U.S.N.M.) ; Billings, 1 ale. (U.S.N.M.) ; Buffalo, 2 skins (U.S.N.M.) ;Highwood Mountains, 1 skin, approaching true evotis (U.S.N.M.) ; HotSprings, 1 ale. (U.S.N.M.).NEVADA: Cottonwood Range, 1 ale. (U.S.N.M.) ; Little Owyhee River. 1skull (U.S.N.M.) ; Pahranagat Valley, 1 ale. (U.S.N.M.).NEW MEXICO: Fort Wingate, 2 ale. (U.S.N.M.) ; Mimbres Mountains, 1skin (U.S.N.M.) ; Sapello Canyon, San Miguel County, 10,000 feet, 1ale. (A.N.S.P.) ; Vermejo River, 1 skin (U.S.N.M.) ; Fort Wingate, 2ale. (U.S.N.M.), 1 ale. (B. M.).
118 BULLETIN 144, UNITED STATES NATIONAL MUSEUMNORTH DAKOTA: Grinuell, 1 skin (U.S.N.M.).OREGON: Camp Harney, 1 ale. (U.S.N.M.) ; Cedar Mountains, MalheurCounty, 1 skin (U.S.N.M.) ; Cornucopia, Wallowa County, 1 skin(U.S.N.M.) ; Disaster Peak, Malheur County, 1 skin (U.S.N.M.) ; Sisters,Cross County, 1 skin (U.S.N.M.) ; Twelve Mile Creek, Crook County, 1 ale.(U.S.N.M.) ; Wallowa Lake, 1 ale. (U.S.N.M.).SOUTH DAKOTA: Corral Draw, 1 ale. (A.M.N.H.).VERA CRUZ: Perote, 7800 feet, 1 ale. (U.S.N.M.).WYOMING: Buffalo Lake, 1 ale. (U.S.N.M.) ; Kinney Ranch. 1 skin, type(A. M. N. H.).Remarks.?As noted by Miller (1897, p. 80), the type specimen ofMyotis evotis chrysonotus has the tail injured so that its shortness incomparison with that of typical Myotis evotis can not be regarded asa valid character. Indeed the similarity of the type to Californianexamples was pointed out by the original describer, who remarkedthat the animal would probably prove to be only a subspecies ofMyotis evotis, a suggestion which is well borne out by the largerseries now^ available. The lighter color as compared with the typicalform seems to be the only diagnostic feature of the race. At present,however, it m'ust be recognized that the status of the subspecies ofMyotis evotis is far from satisfactory. Specimens from localities sofar apart as Montana, Lower California, and Vera Cruz are notlikely to represent one race; but the series of skins now availablehas not enabled us to come to any final conclusions, though it seemsnot improbable that the existence of a central and southern coast form(TTiicronyx), an interior form (true ckrysonotus) , and a southernMexican form will eventually be demonstrated.In parts of western Colorado this bat appears to be less uncommonthan usual. Cary (1911) observes that he has not found it aboutcliffs or ledges, the favorite resort of Pijnstrellus hespeims, but lisu-ally in the vicinity of ranch buildings. Bats of this kind frequentlyfly into houses in pursuit of insects attracted by a light. Cary founda small colony, apparently of adult females and nearly full-grownyoung, inhabiting a deserted ranch house at Sunny Peak, RouttCounty, Colo., in August, 1906. Other than this there appear to beno recorded accolmts of breeding colonies.MTOTIS MILLERI ElliotMyotis milleri Eluot, Field Columb. Mus., publ. 74, zool. ser.. vol. 3, p. 172,April, 1903 ; Land and Sea Mamm. Middle Amer., Field Columb. Mus.,publ. 95, zool. ser., vol. 4, p. 575, 1904 ; Check List Mamm. North Amer.,Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 474. 1905; Catal.Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115, zool. ser.,vol. 8, p. 501, 1907.?Miller, List North Amer. Land Mamm. 1911, Bull.U. S. Nat. Mus., No, 79, p. 59, December 31, 1912; List North Amer.Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 73, April 29, 1924.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 119Type locality.?La Grulla, San Pedro Martir Mountains, LowerCalifornia, Mexico.Type specimen.?Adult male (skin and skull), No. 10840, FieldMuseum of Natural History, collected at La Grulla, San Pedro Mar-tir Mountains, Lower California, September 15, 1902, by EdmundHeller.Distribution.?San Pedro Martir Mountains, Lower California,Mexico.Diagnosis.?Externally resembling Myotis evotis but smaller, fore-arm 34 to 37 mm. instead of 36 to 40 mm., third finger 54 to 58 mm.instead of 62 to 67 mm., and tibia 16.0 to 17 mm. instead of 17 to20 mm. Skull smaller than that of Myotis evotis, the brain case lesselevated, noticeably flat-topped, the sagittal crest absent in each ofthe six specimens examined (nearly always present in adults ofM. evotis). Teeth small, the crown area of the upper molars equalto that of the upper molars of M. keenii (crown area of m^ in sixspecimens, 1.25 to 1.35 by 1.60 to 1.75).Color.?The color does not differ appreciably from that of Myotisevotis cKrysonotus.Measurements.?For measurements see tables, pages 120 and 121.Spedtnens examiTied.?Seven, all from the San Pedro MartirMountains, Lower California, Mexico (2 skins, 1 skull, F.M., in-cluding type; 4 ale, A. M. N. H.).Remarks.?The characters of Myotis lyiilleri appear to be wellmarked and to indicate the existence, in the San Pedro Martir Moun-tains, of a distinct species related to Myotis evotis and M. keenii.The large conspicuously contrasted black ears and the distinctlyfringed interfemoral membrane resemble M. evotis. The generaldimensions and small teeth are essentially as in M. keenii. The flat-tened brain case gives the skull a form unlike that of either of therelated species and somewhat intermediate between the forms charac-teristic of Myotis keenii and M. suhulatus. The specimens of Myotismilleri differ conspicuously from the type of Myotis niicranyx Nelsonand Goldman (collected at Comondu) which has the high brain caseand large teeth (m^ 1.45 by 1.95 mm.) characteristic of Myotis evotis.
120 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis evotis and M. mMleri
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 121Cranial measurements of Mi/Otis' evotis and M. milleri
Locality
Myotis evotis evotisBritish Columbia:Eock CreekShuswapCranbrookDoWashington:Blue MountainsDoDoDoDoDoDoDoDoDoDoDoDoDoOregon:FremontCornucopiaSistersMcKenzie BridgeCamp HarneyDisaster PeakCedar MountainsCalifornia: SissonMyotis evotis chrysonotusIdaho:AlbionDoMontana:Big Belt MountainsBuffaloDo.Highwood Mountains..North Dakota: QrinnellColorado:MoguilSnake RiverDoDoDo >.....Do -DoDoLovelandCalifornia:Mount TallacSouth Yolla Bolly Moun-tainsSan BernardinoLower California: Com-monduArizona:Paradise ,DoSpringerville ,WilliamsChihuahua: San Luis Moun-tainsVera Cruz: PeroteMyotis milleriLower California:San Pedro MartirMountainsDoDoDoDoDo
Number Sex
9.10.27.2, B-M.6713851, Garrett54, GarrettC. 15, LymanC. 16C. 22C. 45C. 52C. 10C. 26C. 47A. 1B.26B.35B. 39B. 42C. 5420490820921820439620439514450020798820798995454
170099170100233473229526229527170027209037149200148166147660148161148162148163148164148165386809405 M.C.Z.138030 U.S.N.M.187430*
> 146044124615124616224921120156304488541
2 10846 F.M.108476425 A. M. N. H.642364246426
16.016.216.115.615.415.415.215.415.415.415.215.415.015.415.215.215.615.015.616.215.916.015.515.615.5
16.416.216.016.016.2
15.615.716.016.115.715.816.116.016.1
15.415.8
16.0
15.916.0
15.014.815.015.015.215.0
as.
15.015.015.014.914.214.414.414.814.214.614.414.614.214.814.414.214.614.615.015.215.015.014.414.214.2
15.415.515.214 615.215.014 614 615.015.014 614 415.015.015.415.014 414 915.015.015.814 815.4
14 213.813.814 614 2
9.2
9.2
9.0
3.8
5.05.25.05.05.04 85.05.00| 4 82j 5.45.00, 5.041 5.22| 5.07.2 5.07.2, 5.07.4i 5.07.6i 5.27.2! 5.07.2 5.07.2 5.2
3.63.83.83.63.8334 7.03.8 7.2
11.211.611.611.211.011.011.011.010.810.611.01011.010.810.611.010.811.211.011.211.211.4ii."o10.811.2
11.010.8U.611.511.811.211.47.2 5.0 11.67.2 5.0 11.07.0, 5.01 11.07.0! 5.o! 11.27.21 5.2! 11.07.2| 5.0; 11.27.0| 5.0! 11.05.0 11.05.0 11.43.8 7.5
3.6 7.0
5.0 11.511.2
11.211.6n.212.011.811.211.8
4 6 10.84.6 10.04 6?10'4 6 11-04 6' 10. 61 5. 8
6.0 6.26.0 6.86.0 6.4
7.1
' Type of Myotis micronyx Nelson and Goldman.58518?28 9 Type.
122 BUUjETIN 144, UNITED STATES NATIONAL MUSEUMMYOTIS THTSANODES MUler(SjTionymy under subspecies)Distribution.?From southeastern Washington, central California,Arizona, and New Mexico southward to the highlands of Oaxaca,southern Mexico: one record from Missouri.
Map 8. -Distribution of Myotis thysanodes : 1, M. thysanodes thysanodes ;2, M. thysanodes aztecusDiagnosis.?General appearance much as in Myotis evotis but sizegreater, about equalling that of M. velifer (forearm 40.6 to 46.0 mm.
;
greatest length of skull 16.2 to 17.2 rmn.) ; tail relatively shorter(average ratio of tail to head and body in 10 specimens from Call-
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 123fornia and Arizona 80.9, in 10 from New Mexico 75.2, and in 10from San Luis Potosi 76.5) ; ear less enlarged (usually about 16to 18 mm. in length) ; foot relatively longer (ratio of its length tothat of tibia usually more than 50) ; and fringe of hairs along freeborder of interfemoral membrane well developed and conspicuous.Maxillary tooth row about 6.5 mm. in length, exceeded by thegreatest width of palate including molars.Ears,?The ears are elongated, when laid forward reaching from3 to 5 mm. beyond the nose. Tragus long and slender, about 60 percent of the height of the ear, its anterior border very slightly convex.The posterior border has a distinct squarish lobule at its base, suc-ceeded by a right-angled notch, above which the outline is at firstconvex with a slightly crenulate edge, then faintly concave belowthe tip.Wing and jnembranes.?The wing membrane arises from the sideof the foot near the base of the toes. The third and fourth meta-carpals are usually nearly equal, while the fifth is a trifle less, butthere is a slight tendency for the fifth to be of equal length withthe fourth. When folded, the third metacarpal falls about 1.6-2mm. short of the elbow and is thus proportionately longer than inMyotis evotis in which the discrepancy is about the same but theforearm is shorter. Taking the third finger as 100, the fourth is 82,and the fifth 79 ; proportions which are essentially like those found inM. evotis. The minute terminal joint of the tail alone is free fromthe membrane. Free edge of the uropatagium bordered both aboveand below by a conspicuous fringe of short stiff hairs which occurin small tufts or clumps, about 15 hairs to each tuft. This fringeis usually quite evident to the eye without the help of a lens.Foot.?The foot is proportionally larger than in Myotis evotis, itslength usually more than half that of the tibia. In 9 topotypes theratio of foot to tibia averages 53.7 ; in 10 specimens from New Mexicoit averages 55.2; in 10 from San Luis Potosi, 53.3; and in 6 fromOaxaca, 56.9. Calcar heavy, without distinct keel, but the skin alongits free margin is thickened and compressed to an evident edge. Itterminates distally in a minute lobule.Fur and color.?The pelage is full and rather long, the hairs about9 mm. in length at the middle of the back. The hair is dark at thebase with slightly buff tips above, and whitish tips below, that atthe sides of the abdomen without dark bases.Skull.?The skull resembles that of Myotis evotis but is immedi-ately distinguishable by its greater size, better developed sagittalcrest, less inflated auditory bullae, and less narrowed rostrum (ratioof greatest breadth across molars to condylobasal length about 44instead of about 40; greatest breadth across molars usually muchgreater than length of maxillary tooth row) ; mandible obviously
124 BULLETIN 144, UNITED STATES NATIONAL MUSEUMlarger and more robust. As compared with the skull of Myotis veli-fer, with which it agrees in general size, that of M. thysaTwdes ismore slender and delicate, the brain case narrower and more oval inoutline. The sagittal crest is about as well developed as in M. veliferbut the occiput is inflated posteriorly so that the lambdoid crests donot meet at the vertex to cut off a raised triangular area. As seenfrom above, therefore, the hind margin of the skull is not truncatebut is bowed backward ; while in rear view the vertex is not peakedbut is evenly arched. In palatal aspect the distance between the lastmolar and the audital bulla is obviously less, both relatively andabsolutely than in Myotis velifer; the palate ends in a well-defined aprojection medially.Teeth.?The molars are normal in the relation of their width tothat of the palate, not enlarged as in Myotis velifer and M. occultus^though distinctly larger than in M. evotis. In the first and secondtooth there is no metaloph and the protoconule and paraloph areusually absent (fig. 1 e, p. 8), though either or both may be slightlydeveloped, particularly in m^. This extreme simplification of struc-ture is not known in other American species of Myotis^ though it isoccasionally approached in M. evotis. The third molar, as in all theother known American members of the genus, retains a distinctprotoconule. Cingulum of m} and 7n? usually wide and distinct,better developed than in Myotis velifer, though not actually passingaround the antero-lingual base of the protocone. At the postero-lingual base of the hypocone the cingulum occasionally becomesthickened and tubercle-like, a condition especially well shown in thetype.The two small upper premolars are usually both visible in sideview, though drawn inward slightly from the tooth row; there issomewhat more variation in their exact position than in relatedspecies. In one specimen (52186 U.S.N.M.) both the minute upperpremolars are missing on the left side while on the right side thelarger only is present, almost completely filling the space betweencanine and p*, while in another (62228 U.S.N.M.) the two smallteeth are united."Remarks.?Externally Myotis thysaxnodes may be distinguishedfrom M. evotis by its greater size and less enlarged ears. Its furis not so full nor so golden in color, a difference fairly obvious ondirect comparison. The ears are usually brownish instead of black.The free edge of the tail membrane is so thickly edged with shortcoarse hairs that they form a fringe which is clearly obvious tothe naked eye, whereas in Myotis evotis the fringe is less denseand not distinctly visible without a lens. The presence of this
" Figured by Miller, North Amer. Fauna, No. 13, p. 82, Oct. 16, 1897.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 125
conspicuous fringe distinguishes Myotis thysanodes from all otherknown American members of the genus. In the reduced conditionof the secondary cusps and ridges of m} and m?' Myotis thysa/nodesshows the culmination of a series of changes whose earlier stagesare seen in M. keemi and whose development may be traced in M.evotis.The possibility that Myotis thysanodes is the American representa-tive of the palearctic M. nattereri has been suggested by Thomas.^'The range of the Old "World species is, however, across temperateEurasia from western Europe to Japan (subspecies toiribvrms)whereas that of Myotis thysanodes in America is more southern,almost subtropical. An equally near relationship is perhaps to befound between Myotis nattereri and M. evotis. The northern limitsof these two species in the Old and New Worlds respectively reachabout the same latitude, so that the ancestor of M. evotis may haveeasily invaded North America from Asia during a Pliocene connec-tion of the two continents. Whatever may have been their exacthistory it seems evident that the three animals represent a commonparent stock.Two subspecies are recognized: The typical form ranging fromWashington and California to northern and central Mexico, and adarker-colored race occurring in southern Mexico and here describedas new.Habits.?^Beyond the account supplied by Dr. T. S. Palmer (Miller,1897) of the colony at Old Fort Tejon, Calif., nothing seems tobe recorded as to the habits of this bat. Doctor Palmer found thisspecies and M. ywnanensis in numbers in the dark attic of an oldadobe house, July 3. Young in various immature stages as well asadults of both species were secured, among which Myotis thysanodesappears to have been in the majority. Dr. Joseph Grinnell visited thelocality 13 years later but failed to find this bat. On September 28,1903, James H. Gaut, collecting for the United States BiologicalSurvey, obtained several in the ruins of Gran Quivera, Mesa Jumanes,N. Mex. A note on the label of specimens collected by Salle inOaxaca in 1857, states that they too were found in an old ruin. Thelate Dr. E. A. Mearns captured two adult females on June 28 inthe San Luis Mountains (on the Mexican boundary) that containedeach a large fetus nearly ready for birth. During the breedingseason the females appear to remain in colonies by themselves apartfrom the males until late summer at least. Such a breeding colonywas discovered by Mr. Vernon Bailey on August 9, 1913, at Clover-dale Hills, N. Mex., and 83 specimens were preserved, most of themin alcohol. An examination of these, evidently taken at random,
??Proc. Zool. Soc. London, 1905, p. 388.
126 BULLETIN" 144, UNITED STATES NATIONAL MUSEUMshows that 53 are adult females and the rest young animals, malesand females in about equal proportions, most of them nearly fullgrown but with the metacarpal epiphyses still showing immaturity.No adult males had as yet joined the colony. Under natural condi-tions this is doubtless a cave-haunting species. Myotis thysamodesseems to be less common or at least more sporadic than Myotis velifevjfor though the two animals occur over much the same general area,collections contain far fewer examples of this species than of thelatter. MYOTIS THYSANODES THYSANODES MillerVespertilw subulatus H. Allen, Monosrr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 51, June, 1864 (part; specimen supposed to have beencollected at St. Louis, Missouri, listed on page 53).Vespertilio albescens velifer H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 93, March 14, 1894 (part; specimen fromDulzura, California).Vespertilio albescens evotis H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 90, March 14, 1894 (part; specimen No. 29827,U. S. N. M., Fort Tejon, California).Myotis thysanodes Miller, North Amer. Fauna, No. 13, p. 80, figs. 16-17 (teeth),October 16, 1897.
?
Trouessabt, Catal. Mamm. viv. foss., p. 1285, 1899.
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p. 258, December27, 1901.
?
Elliot, Land and Sea Mamm. Middle Amer., Field Columb. Mus.,publ. 95, zool. ser., vol. 4, p. 572, 1904.
?
Trouessabt, Catal. Mamm. viv.foss., suppl., p. 94, 1904.
?
Elliot, Check List Mauim. North Amer., FieldColumb. Mus., publ. 105, zool. ser., vol. 6, p. 479, 1905.
?
Stephens, CaliforniaMammals, p. 268, 1906.
?
Elliot, Catal. Mamm. Field Columb. Mus., FieldColumb. Mus., publ. 115, zool. ser., vol. 8, p. 505, 1907.
?
Lyon and Osgood,Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 273,January 28, 1909.
?
Miller, List North Amer. Land Mamm. 1911, BulLU. S. Nat. Mus., No. 79, p. 59, December 31. 1912.?J. Grinnell, Proc.California Acad. Sci., ser. 4, vol. 3, p. 278, August 28, 1913.?H. W. Grinnell,Univ. California Publ. Zool., vol. 17, p. 297, January 31, 1918.?J. Grinnell,Univ. California Publ. Zool., vol. 21, p. 314, January 27, 1923.?Miller, ListNorth Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 73, April29, 1924.?GRINNE3LL and Stober, Anim. Life in the Yosemite, p. 57, 1924.Myotis evotis thysanodes Elliot, Synops. Mamm. North Amer., Field Columb.Mus., publ. 45, zool. ser., vol. 2, p. 406, March, 1901 ; List Land and SeaMamm. North Amer., Field Columb. Mus., publ. 57, zool. ser., vol. 2, p. 518,June 1901.T^pe locality.?Old Fort Tejon, Tehachapi Mountains, KernCounty, Calif.Type speciinen.?Adult female (in alcohol). No. 29827, UnitedStates National Museum (Biological Survey collection), collectedat Old Fort Tejon, Tehachapi Mountains, Kern County, Calif., July i5, 1891, by T. S. Palmer. Original number, 235.Distribution.?From southeastern Washington, central California,Arizona, and New Mexico southward into northern and central
;
Mexico; exact limits of range not known. (See map 8, p. 122.)
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 127Diagnosis.?General color a pale yellowish brown.Color.?Upper parts a uniform "warm buff" (Kidgway, 1912),the tips of the hairs slightly shining ; basally the hairs are " fuscousblack" and a small area of this color is sometimes present at theshoulder. Below, the general tone is dull whitish due to the fuscous-black bases of the hairs showing through the buffy white tips. Alonga narrow strip at the sides of the abdomen the hairs are whitish to theroots.Measurements.?For measurements see tables, pages 128 to 130.Spedmens examined.?Total number 193, from the followinglocalities
:
ARIZONA: Beale's Spring, 1 ale. (U.S.N.M.) ; Flagstaff, 1 ale.(A. M. N. H.) ; Springerville, 1 skull (U.S.N.M.).CALIFORNIA: Old Fort Tejon, 14 ale., including type (U.S.N.M.);Walker Pass, 1 ale. (U.S.N.M.).CHIHUAHUA: San Luis Mountains, 2 skins (U.S.N.M.).JALISCO: La Laguua, 1 skin (U.S.N.M.), not typical; Los Masos, 7 skins(A. M. N. H.), not typical.MICHOACAN: Patzeuaro, 2 skins, not typical. 2 ale. (U.S.N.M.).MISSOURI: St. Louis, 1 ale. (U.S.N.M.).NEW MEXICO: Carlsbad, 1 ale. (U.S.N.M.) ; Cloverdale, Grant County, 18skins, 65 ale. (U.S.N.M.) ; Copperton, Valencia County, 1 skin (U.S.N.M.) ;Espanola, 1 ale. (U.S.N.M.) ; Gallup, McKinley County, 1 skin (U.S.N.M.) ;Mesa Jumanes, 3 skins (U.S.N.M.) ; "Mountains of New Mexico," 1 skin(U.S.N.M.) ; Sacramento Mountains, Otero County, 1 ale. (A. N. S. P.).SAN LUIS POTOSI: Hacienda La Parada, 6 skins, 56 ale. (U.S.N.M.);1 ale. (B. M.).WASHINGTON: Anatone, Asotin County, 1 skin (U.S.N.M.).ZACATECAS: Hacienda San Juan Capistrano, 1 skin, 3 ale. (U.S.N.M.).Reinarks.?It seems characteristic of Myotis thysanodes thysanodesto be found somewhat sporadically. In California the animal hasbeen recorded from only five localities, Dulzura, San Diego County(Miller, 1897), Fyffe and Limekiln, El Dorado County (H. W.Grinnell, 1918), Old Fort Tejon, Kern County, and Walker Pass,Kern County. Yet its range may be expected to include most of theaustral zone area of the State. A single individual has been takenin Asotin County, southeastern Washington, but the species has ap-parently not yet been detected in Oregon or Nevada. An alcoholicspecimen in the United States National Museum (No. 3W/5 ); re-ceived many years ago (catalogued in 1861), is labeled: " St. Louis,Missouri, Dr. Engleman." If there is no mistake about this record,St. Louis marks the extreme known eastward limit of the animal'srange.In northern and central Mexico the color becomes graduallydarker, so that specimens from San Luis Potosi and Zacatecas arepractically intermediate between the typical form and the southernrace. Five specimens from San Luis Potosi, Mexico, are not typical.
128 BULLETIN 144, UNITED STATES NATIONAL MUSEUMOne is as pale above as the average of New Mexican skins, one isnearly as dark as those from Oaxaca, and the others are interme-diate, but all are perhaps best referred to the subspecies thyscmodes.In the series of 20 skins from Cloverdale Hills, southwestern NewMexico, one or two are more intensely colored than the average.MYOTIS THYSANODES AZTECUS, new snbspedesType.?Skin and imperfect skull, No. 58.6.2.3 British Museum(Natural History), collected at San Antonio, Oaxaca, Mexico.Tomes collection, 1860.Distribution.?Southern Mexico (Oaxaca). (See map 8, p. 122.)Diagnosis.?Similar to the typical form, but darker in color.Color.?The color is a uniform " tawny olive " above, the tips ofthe hairs somewhat shining, their bases dark, nearly fuscous black;a dark fuscous spot at the shoulder. Below, the paler tips to thehairs are tinged with buflfy, producing a duller effect than the whitishtips of typical Myotis thysanodes.Skull.?The skulls of the specimens from Oaxaca are all damagedposteriorly and have the zygomatic arches broken. They are never-theless obviously shorter of rostrum than in the typical subspecies.Measurements.?For measurements, see tables, pages 128 to 130.Specimens exaymined.?Total number 6, from the followinglocalities
:
OAXACA: Hacienda de Cinco Seiiores, 5 skins (B. M.) ; San Antonio, 1skin (B. M.).External measurements of Myotis thysanodes
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 129External measurements of Myotis thysanodes?Continued
Locality
130 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial tneoisurenientg of Myotis thysanodes?Continued I
Locality
Myotis thysanodes thysan-odes?ContinuedSan Luis Potosi:Hda. la ParadaDoDoDoDoJalisco: La LagunaMichoaean:PatzcuaroDo ,Myotis thysanodes aztecnsOaxaca:San AntonioHacienda de Cinco SefioresDoDoDo
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 131though not always keeled calcar, and by the pinkish-gray color andloose texture of the fur. Forearm 36 to 40.6 mm. (usually 38 to 39mm.). Skull with a narrower brain case and more pronounced sag-ittal crest than in Myotis lucifugus.Ears.?The ear is about as in Myotis lucifugus., of medium size, itslower anterior border convex, then nearly straight in its upper half,with a broadly rounded summit, below which the outer margin is
Map 9.
?
Distribution of Myotis sodalisvery slightly concave, then passes gradually into the basal shoulderwithout forming a notch or abrupt transition. Laid forward theear reaches to the tip of the nostril. Tragus rather short and blunt,curving slightly forward, its total height a little less than half thetdtal height of the ear from outer corner to tip (about as 6 : 14 mm.).The posterior edge is shghtly crenulat*.^Ying and nnemhranes.?Wing from the side of the foot, beginningjust below the base of the outer toe at the head of the metatarsal.The metacarpals are regularly graduated, the third longest, thefourth and fifth successively a trifle shorter. The lengths of thefingers are similarly graduated. Taking the third finger as 100, thefourth and fifth are respectively as 73 and 67 (69.5:50.5:46.5 mm.),
132 BULLETIISr 144, UNITED STATES NATIONAL MUSEUMthat is, the fourth and fifth fingers are relatively short. Whenfolded, the third metacarpal falls short of the elbow by about 4.5 mm.On the under side of the wing the fur extends thickly as far as aline joining the knee and the middle of the humerus, and on theupper side of the interfemoral membrane as far as a line joining theextended knees. The minute terminal vertebra of the tail and aboutone-half of the penultimate vertebra project free.Foot.?The foot is smaller and more delicately formed than inMyofis lueifugus^ its length a little more than one-half that of thetibia. In 10 specimens from Proctor, Vt., the ratio of foot to tibiaaverages 50.6; in 10 from Center County, Pa., 49.3; in 10 from Mam-moth Cave, Ky., 51.7. The calcar is long, about 16.5 mm., andnearly equals the free border of the interfemoral membrane. Itusually has a low keel, but this is occasionally obsolete or absent.The calcar ends in a minute projecting lobule.Fur and color.?The texture of the fur is extremely fine and fluffy
;
the hairs have a tendency, due perhaps to a slight crinkling, to standout from each other a little, as in Pipistrellus suhflavus. In com-parison with the bronzy burnished-tipped fur of Myotis lucifugusthe pelage of this species is dull grayish chestnut, though on closeinspection the longer tips of the hairs are seen to be slightly polished.Series of skins of the two animals are very different in appearance.The color is distinctive. On the upper surface tTie basal two-thirds of the hair is fuscous-black, then comes a narrow grayishband succeeded by a cinnamon-brown tip, so that there is a dis- ^tinctly tricolor effect, while the grayish band showing through thecinnamon-brown tips gives a peculiar hoary appearance at a short jjdistance. Below, the fur is slaty basally, the hairs with grayish- *white tips, washed more or less heavily with cinnamon brown, par-ticularly at the flanks, instead of slightly yellowish as in M. luci-fugus. The general effect is a pinkish white below and a dullchestnut gray above. The membranes and ears are blackish brown.One unusually bright skin from Proctor, Vt., is dark cinnamonabove, nearly " sayal brown " of Ridgway, slightly washed with thesame on the sides of the chest.JSkull.?The skull in general resembles that of Myotis lucifuguslucifugus with which it agrees in length (usually 14.4 to 15 mm.).In form it differs in certain details that become evident on closecomparison. The most striking of these is the smaller, narrowerbrain case (6.6 to 7.2 mm. instead of 7.1 to 7.6 mm.), which insteadof being high and with a broadly flattened top is narrower, lower,(4.6 to 5.1 mm. instead of 4.9 to 5.5 mm.) and more arched trans-versely than in M. I. lucifugus. A slight but perfectly definite sagit-tal crest is normally present in adults, whereas in M. I. lucifugusthe brain case is so broad-topped that the temporal muscles rarely
AMEKICAN BATS OF THE GENEEA MYOTIS AND PIZONYX 133
meet to form a sharp ridge. ^? Correlated with this difference isthe more tapering front end of the brain case, a slightly narrowerinterorbital constriction (3.5 to 3.9 mm. instead of 4.0 to 4.4 mm.)with more graceful lines; and a slightly larger rostrum as com-pared with the brain case, features which are obvious to the eye,and consistently borne out by actual measurements. The lengthof the maxillary tooth row is essentially the same as in Myotialucifugus (usually 5.2 to 5.6 mm.), but the maxillary breadth at m^is distinctly less (usually 5.3 to 5.8 mm. instead of 5.5 to 6.1). ? Thesecranial differences between the two species become obvious on com-parison of the tables of measurements pages 134 and 135.Teeth.?The teeth are so similar to those of Myotis liwifuguslucifugus that we have been unable to detect any positive charactersby which they may be recognized. Apparently the two small upperpremolars tend to be more nearly equal in area when seen in crownview, but this feature is not constant. The cingulum and the secon-dary cusps and ridges are developed as in M. lucifugus. In a speci-men from Arkansas (No. 5831 F. M.) the second upper premolar isabsent on both sides.Measurements.?For measurements see tables, pages 134 and 135.Specimiens exmnined.?Total number 443, from the followinglocalities
:
ALABAMA: Anniston, 1 ale. (M. C. Z.).ARKANSAS: Greenway, 2 skins (F. M.).ILLINOIS: Rosiclare, 6 skins, 4 ale. (F. M.) ; no exaet locality, 1 ale.(U.S.N.M.).INDIANA: Bicknell, 1 ale. (U.S.N.M.) ; Wheatland, 3 ale. (U.S.N.M.) ;Wyandotte Cave, 3 ale. (U.S.N.M.), 14 skins, 3 ale. (F. M.), 10 skins.including type, (M. C. Z.).KENTUCKY: Mammoth Cave, 269 ale. (U.S.N.M.), 6 ale. (A. N. S. P.),1 ale. (B. M.). 20 ale. (M. C. Z.).MICHIGAN: Grosse Isle, 1 skin (U.S.N.M.),NORTH CAROLINA: Roan Mountain, 1 ale. (U.S.N.M.).PENNSYLVANIA: Center County, (cave), 2 skins, 25 ale. (U.S.N.M.).TENNESSEE: Hickman Co., 1 skin (A. M. N. H.).VERMONT: Brandon, 2 skins, 9 mummie.s (U.S.N.M.) ; Proctor, 21 skins.37 ale. (U.S.N.M.).Remarks.?That Myotis sodalis has been so long overlooked is dueno doubt to the general resemblance which the animal bears toMyotis lucifugus^ with which species the specimens of it in museumshave generally been confused; when once its characters are recog-nized, however, there is no doubt as to its identity. Not only is thetricolor fur, with the characteristic pinkish-gray tinge distinctive,but the slightly smaller size of the foot and the normal presenceof an obvious keel on the edge of the calcar are further diagnostic.
" In 20 skulls of Myotis sodalis from Mammotli Cave the sagittal crest is present eighteentimes, while in 20 skulls of M. lucifugus lucifugus it is present only three times and thenmuch less well developed than in the average of the other species.
134 BULLETIN 144, UNITED STATES NATIONAL MUSEUMIt is true that in distorted or badly preserved specimens the keel maynot always be evident, and the color of the fur is not to be made outin bleached alcoholic material; but the smaller foot is usually suffi-cient to separate Myotis soctalis from M. lucifugus^ and the skull,in its smaller size, more slender form and slight median crest, islikewise sufficiently diagnostic.It is a curious fact that although this bat occurs over much ofthe eastern United States, nearly all the known individuals weretaken during hibernation. A series was secured in a cave at Proctor,Vt., in early April, where a few Myotis lucifugus were found atthe same time. A great number annually winter in the WyandotteCave, Ind. Two or three lots of bats from this station include thisspecies only. Where these bats go with the coming of spring and inwhat way their habits differ from those of Myotis lucifugus it isimpossible to say. Yet it seems probable that some difference inhabits exists, for Myotis sodalis has not been captured as M. lucifugusso often is, either in small colonies in buildings or as single individualsshot while flying about the edges of ponds and along streams.External measurements of Myotis sodalis
Locality
Vermont:ProctorDo -Do...Do-DoDoDoDoDoDoPennsylvania:Center CountyDoDo.DoDoDoDoDoDoDoIndiana:BicknellWheatlandDo.DoWyandotte CaveDo.Do'DoKentucky:Mammoth CaveDoDoDoDoDo..DoDo-._Do...Do
?Type.
Number
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 135Cranial measurements of Myotis sodalis
Locality
136 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
Map 10.?Distribution of Mtotis volans : 1, M. volans volans ; 2, M. volansLONGICEUS ; 3, M. VOLANS INTERIOR ; 4, M. VOLANS AMOTUS
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 137its greatest length 12.2 to 15.0 mm. 5 maxillary tooth row rangingfrom 4.5 to 6 mm. ; lower tooth row (excluding incisors) 4.9 to 6 mm.Ears low and bluntly rounded, barely reaching the nostril when laidforward. Foot decidedly less than one-half the length of tibia, theratio of its length to that of tibia about 41 ; calcar distinctly keeled.Fur on the under side of wing membrane extending out to the levelof the elbow.Ears.?The ear is short, barely reaching the nostril when laidforward, its anterior margin strongly convex, its upper half beveledabruptly backward to the broadly rounded tip. Tragus short, abouthalf the height of ear from meatus (5 : 10 mm.) ; at its base a smallrounded lobe, above which the anterior and posterior margins con-tinue nearly parallel half way to the tip whence the posterior out-line is sharply beveled forward to the tip.
'Wing and Tnemhranes.?The wing membranes arise from the baseof the toes. Third metacarpal longest; second, fourth, and fifthabout equal, or the two latter graduated so that 4 is shorter than 3,and 5 is shorter than 4. When folded the third metacarpal falls 1to 1.5 mm. short of the elbow. Taking the third finger as 100, thefourth is 83, the fifth 77 (66 : 55 : 51 mm. ; specimen from MenloPark, Calif.). Tail involved in membrane except at its extreme tip,where the terminal vertebra alone projects. Free border of uropa-tagium without fringing hairs.Foot.?The foot is delicately formed, its length decidedly less thanhalf the length of the tibia (ratio of foot to tibia in 2 specimensfrom Lower California 40.8; in 10 specimens from Nicasio, Calif,,41.7; in 10 from localities within the area inhabited by the interiorrace, 40.7). Calcaneum about as long as the free edge of the uropa-tagium, ending in a distinct but minute lobule. At about the lengthof the tarsus from the ankle, a low elongate keel arises.Fur.?Pelage full and long, about 7 mm. on lower back. Above,the hair extends out on to the interfemoral membrane over an areaapproximately the length of the femur. Below, the wing membraneis usually well furred as far out as a line joining the elbow and theknee. Membranes blackish brown.Skull.?The skull is of characteristic form, small and delicate withthe rostrum shortened and the profile of the brain case elevatedabruptly, giving a '' pug-nosed " effect (pi. 1, p. 7, fig. 10). Occiputhigh and slightly inflated. Temporal ridges slightly developed butusually traceable with a lens. They unite to form a low sagittal crestanterior to the occiput. Posterior to this point of union they bowoutward, to continue back with a rather characteristic outline, cut-ting off a somewhat triangular area whose sides are convex instead of58518?28 10
138 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
straight or concave. The lambdoid crests are transverse, sometimesmeeting at the occiput to form a sliort, backwardly directed angle.Teeth.?Molars proportioned to palate about as in Myotis liM)i-fugus. The secondary cusps and ridges show an obvious tendencyto reduction; protoconule often less well developed than in Myotislucifugus, sometimes reduced to a mere thickening of the anteriormargin of the crown; metaloph usually present but rarely if everlong enough to reach the crest of the ridge connecting protocone withhypocone; cingulum as in M. lucifugus. The two small upper pre-molars are nearly equal in cross-section; the posterior, however, isslightly the smaller and about two-thirds the height of the anteriortooth. They are usually both drawn slightly in from the tooth row,the second commonly more than the first, though in some individualsthey are nearly in the row, and again the second may be so crowdedinward as hardly to be visible from the exterior. The premolars ofthe lower jaw stand completely in the tooth row without crowding, thefirst with a cross-section about one-third greater than the second,which stands about two-thirds as high. As in Myotis lucifugus thewidth of palate including the upper molars slightly exceeds thelength of the maxillary tooth row (front of canine to back of lastmolar), and about equals the length of the mandibular tooth row(excluding incisors).Remarks.?This species, though superficially resembling Myotislucifugus.^ is very distinct from other American bats. Its shortrounded ears, small foot, well-developed calcaral keel and obviousextension of the fur on the ventral side of the membranes to a linejoining the elbow and knee, are characters which, combined, distin-guish it at once externally. The skull is further diagnostic with itsshortened rostrum and large, high brain case, with the convex outlinesof the temporal ridges meeting at the occiput.Little is known concerning the habits of Myotis volans. It seemsto frequent open forest, ranging in summer to an altitude of at least11,000 feet in the Sierra Nevada. Apparently it is not social to anygreat degree and in general it seems to avoid caves. Dr. JosephGrinnell notes a specimen with a single fetus taken at Santa RosaPeak, Calif. ; a female from Anatone, Wash., June 23, also containeda single young.The specimen on which Harrison Allen, in 1866, based his Vesper-tilio volans was for many years lost sight of. The name was for atime relegated to the synonymy of Myotis califorrdcus (Miller, 1897).It was not until 1914 that Goldman, on reexamining the type, foundit to be a representative of the species whose characters had been firstclearly indicated by True, in 1886, under the specific name longicrus.Both names, however, remain in use, as they were applied to differentsubspecies. Miller, in 1897, regarded this species as a form of Myotis
AMEKICAISr BATS OF THE GENEKA MYOTIS AND PIZONYX 139luGifugtis, but in 1914 he showed that it is distinct. Four subspeciesare noAv recognized.MYOTIS VOLANS VOLANS (H. Allen)Vespertilio volans H. Allen, Froc. Acad. Nat. Sci. Philadelphia, 186G, p. 282.
?
DoBsoN, Catal. Chiroptera Brit. Mus., p. 329, 1878.?H. Allen, Monogr. BatsNorth Amer., Bull. U. S. Nat. Mus., No. 43 (1893), p. 94, March 14, 1894 (assynonym of "Vespertilio nitidus "=^ califomicus).?MiLLsat, North Amer.Fanua, No. 13, p. 69, October 16, 1897 (as synonym of Myotls californicus)
.
?
?
Lyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat.Mus., No. 62, p. 273, January 28, 1909.Myotis capitaneus Nelson and Goldman, Proc. Biol. Soc. Washington, vol. 22,p. 28, Marchi 10, 1909 (San Jorge, near Comondu, Lower California).MiLLEB, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79,p. 56, December 31, 1912.
?
Goldman, Proc. Biol. Soc. "Washington, vol. 27,p. 102, May 11, 1914.Myotis volans Goldman, Proc. Biol. Soc. Washington, vol. 27, p. 102, May 11,1914.?MiLLEB, List North Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus.,No. 128, p. 71, April 29, 1924.Type locality.?Cape St. Lucas, Lower California, Mexico.Type specimen.?Adult female in alcohol, No. 5398, United StatesNational Museum, collected at Cape St. Lucas, Lower California,Mexico, by John Xantus. According to Lyon and Osgood, in 1909,the alcoholic portion of the type could not then be found ; it appears,however, to be in the Academy of Natural Sciences at Philadelphia.Distribution.?So far as known the typical form is confined toLower California. (See map 10, p. 136.)Diagnosis.?Size smallest of the races, forearm about 35 mm., tibia17. Skull delicately formed, distinctly smaller than in the othersubspecies, its greatest length less than 13 mm.Description.?Color above a clear reddish buff?about " clay color "of Ridgway with a tinge of cinnamon?slightly paler on the head andneck; below dull whitish washed with buffy; the bases of the hairsboth above and below plumbeous black, except ventrally along theborders of the membranes where the dark bases are lacking. Tips ofthe hairs above only slightly glossy ; membranes dark blackish brown.There is no dark spot at the shoulder.In color the only available skin (type of Myotis capitaneus) differsvery little if at all from the type of the race interior, which, however,is the brightest of the series representing that subspecies.Measurements.?^For measurements see tables, pages 146 and 147.Specimens examined.?Total number 6, from the followinglocalities
:
LOWER CALIFORNIA: Cape St. Lucas, 1 skull, type (U.S.N.M.), 1 ale.without skull, type (A.N.S.P.) ; San Jorge, 1 skin, type of capitaneus(U.S.N.M.) ; Santa Anita, 4 ale, all but one immature (U.S.N.M.).
140 BULLETIN 144, UNITED STATES NATIONAL MUSEUMMYOTIS VOLANS LONGICRUS (True)Vespertilio longicrus True, Science, vol. 8, p. 588, 1886.Vespertilio lucifugus C. H. Townsend, Proc. U. S. Nat. Mus., vol. 10, p. 182,1887 (see H. W. Grinnell, 1918).Vespertilio nitidus longicrus H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 103, March 14, 1894.?Tbouessabt, Catal.Mamm. viv. foss., p. 130, 1897.Myotis lucifugus longicrus Miller, North Amer. Fauna, No. 13, p. 64, October16, 1897.
?
Tbouessart, Catal. Mamm. viv. foss., p. 1284, 1899.
?
Mebriam,North Amer. Fauna, No. 16, p. 89, October 28, 1899.
?
Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 402, March,1901.
?
Elliot, List Land and Sea Mamm. North Amer., Field Columb. Mus.,publ. 57, zool. ser., vol. 2. p. 517, June, 1901.
?
Miller and Rehn', Proc. Bos-ton Soc. Nat. Hist., vol. 30, p. 256, December 27, 1901.?Elliot, Land andSea Mamm. Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4,p. 581, 1904.
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Trouessart, Catal. Mamm. viv. foss., suppl., p. 92, 1904.
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Stone, Proc. Acad. Nat. Sci. Philadelphia (July, 1904), p. 579, October 17,1904. Elliot, Check List Mamm. North Amer., Field Columb. Mus., publ.105, zool. ser., vol. 6, p. 479, 1905 (part).
?
Stephens, California Mammals,p. 265, 1906 (part).?Seh-on, Life Hist. Northern Anim., p. 1148, 1909(part).
?
Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat.Mus., No. 79, p. 55. December 31, 1912.?J. Grinnell, Proc. CaliforniaAcad. Sci., ser. 4, vol. 3, p. 276, August 28, 1913 (part).?Bailey, WildAnim. Glacier Nat. Park (Dept. of the Interior; Nat. Park Service), p.100, 1918.Myotis longicrus Lyon and Osgood, List Type-sp. Mamm. U. S. Nat. Mus., Bull.U. S. Nat. Mus., No. 62, p. 271, January 28, 1909 (combination wronglyattributed to True).
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Dice, Journ. Mamm., vol. 1, p. 11, November 28, 1919.Myotis longicrus longicrus Millee, Proc. Biol. Soc. Washington, vol. 27, p.211, October 31, 1914.?H. W. Grinnell, Univ. California Publ. Zool., vol.17, p. 267, January 31, 1918; Univ. California Publ. Zool., vol. 17, p. 431,April 25, 1918.?J. Grinne3.l, Univ. California Publ. Zool., vol. 21, p.314. January 27. 1923.
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Miller, List North Amer. Recent Mamm. 1923,Bull. U. S. Nat. Mus., No. 128, p. 68, April 29, 1924.?Grinnell and Storer,Anim. Life in the Yosemite, p. 56, 1924.Myotis altifrons Hollister, Smithsonian Misc. Coll., vol. 56, No. 26, p. 3. De-cember 5, 1911 (Henry House, Alberta, Canada).
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Miller, List NorthAmer. Land Mamm. 1911. Bull. U. S. Nat. Mus., No. 79, p. 56, December 31,1912. Hollister, Canadian Alpine Journ., special number (1912), p. 7,February 17, 1913.
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Miller, List North Amer. Recent Mamm. 1923, Bull.U. S. Nat. Mus., No. 128, p. 69, April 29, 1924.Type locality.?^Vicinity of Puget Sound, Wash.Type specmien.?Siibadult female in alcohol (skull removed). No.
?5MM5 United States National Museum, from "the vicinity of PugetSound," Washington. David Starr Jordan, collector. Thoughnot designated by number in the original description, the singlespecimen which served as the basis for the name is clearly identifiablein the United States National Museum (see Lyon and Osgood, 1909).Distribution.?Humid coast region from Admiralty Island, Alaska,south to Monterey County, Calif, (see map 10, p. 136). 1
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 141This is the dark form of the humid coast region of the Northwest.The most northerly locality known for it is Mole Harbor, AdmiraltyIsland, in southwestern Alaska, whence the University of Californiahas a specimen taken June 9, 1907. Thence southward it occurs inBritish Columbia as well as in western Alberta, where at HenryHouse, the type specimen of Myotis altifrons was captured. Thislast, however, proves to be inseparable from M. v. longicrus. South-ward again, typically dark examples are found from the CascadeRange to the coast in the western half of Washington and Oregon,while in California it is present coastwise in Humboldt, Mendocino,Sonoma, and Marin Counties as far south at least as San Mateo andMonterey Counties. Inland from this area there is a lessening ofthe saturate appearance, and adult specimens show a distinctly lighteror ochraceous-tawny cast. This is true of skins from eastern Oregon,and the Sierra Nevada of California from Mount Shasta southwardto Mount Whitney. These are more or less intermediate between M.V. longicrus and M. v. interior^ and occasional specimens from thecentral portions of the Sierra may be referred with almost equalpropriety to either form, as has been shown by Mrs. Grinnell (1918,for Mono County). On the whole, however, the Sierra Nevadaseries seem best placed with the brighter race, interior. Immatureindividuals are, of course, much darker than adults and lack almostentirely the ruddy golden tint of adults of M. v. inferior, so that theirgeneral appearance is nearly as in worn adults of M. v. longicrus.Diagnosis.?Darkest of the subspecies, general color dark reddishbrown above, smoky below ; size greater than in typical Myotis volans(forearm 37 to 40; tibia 18 to 19.6; greatest length of skull 13.7 to14.6 mm.).Color.?In fresh unworn pelage the tips of the longer hairs are avery dark reddish brown above?near cinnamon brown of Ridg-waj?recalling the color of Myotis lucifugus Imcifugus; below, smokybrown, paler posteriorly; the bases of the hairs blackish; ears andmembranes blackish.Most of the skins available were collected in late summer and seemto have lost much of the long reddish tips of the hairs through wear,and in general are a dark smoky brown ( a little darker than Prout'sbrown), paler below.A specimen from Admiralty Island, Alaska, is blackish brown withvery little of the cinnamon tipping on the back, but is closely matchedby some of the skins from Washington.Measuretnents.?For measurements, see tables, pages 146 and 147.Specimens exa/mined.?Total number 127, from the followinglocalities
:
ALASKA: Admiralty Island, Mole Harbor, 1 skin (U.C).ALBERTA: Henry House, 1 skin, type of altifrons (U.S.N.M.).
142 BULLETIN 144, UNITED STATES NATIONAL MUSEUMBRITISH COLUMBIA: Cranbrook, 1 skin, not typical (Garret) ; VancouverIsland, EiTington, 1 skin (U.G.).CALIFORNIA: Cazadero, Sonoma County, 1 skin, not typical (U.C.) ; ChalkPeak, Monterey County, 1 skin (U.C.) ; Guerneville, Sonoma County, 2skins (U.C.) ; Hurleton, Butte County, 1 skin, nearly typical (U.S.N.M.) ;Menlo Park, San Mateo County, 2 ale. (U.S.N.M.) ; Mount Sanhedrin,Mendocino County, 1 skin (A. N. S. P.) ; Mount Veeder, Sonoma County,1 skin, nearly typical (U.S.N.M.) ; Nicasio, Marin County, 2 skins, 71 ale.(U.S.N.M.), 1 ale. (B.M.) ; Pacheco Pass, Santa Clara County, 1 skin(U.S.N.M.) ; Pesoadero Creek, San Mateo County, 1 skin (U.C.) ; PointReyes, Marin County, 1 ale. (U.S.N.M.) ; Sherwood, Mendocino County,1 skin (U.C).OREGON: Baker County, 2 ale. (U.S.N.M.) ; Cascade Mountains, easternbase, 1 skin (U.S.N.M.) ; McKenzie Bridge, 3 skins (U.S.N.M.).WASHINGTON: Bartholomew (Blue Mountains), 10 skins (S. H. Lyman) ;Beaver Creek, Whatcom County, 1 skin (U.S.N.M.) ; Bumping Lake,Yakima County, 1 skin, 1 ale. (U.S.N.M.) ; Carson, 2 skins (U.S.N.M.) ;Chilliwack River, Whatcom County, 1 skin (U.S.N.M.) ; Entiat, 1 ale.(U.S.N.M.) ; Godman Springs (Blue Moimtains), 6 skins (S. H. Lyman) :Kirkland, 1 skin (U.S.N.M.) ; Lake Cushman (Mason County), 1 ale.(U.M.) ; Mount Angeles, Clallam County, 1 skin (U.S.N.M.) ; MountRainier, 1 skin (U.S.N.M.) ; Oroville, 1 ale. (U.S.N.M.) ; Port Townsend,1 ale. (U.S.N.M.) ; Puget Sound, 1 ale, type (U.S.N.M.).MYOTIS VOLANS INTERIOR MillerMyotis lucifugus longicrus Miller, North Amer. Fauna, No. 13, p. 64, October16, 1897 (part).?Tbouessabt, Catal. Mamm. viv. foss., p. 1284, 1899 (part).?Elliot, Synops. Mamm. North Amer., Field Columb. Mus., publ. 45, zool.ser., vol. 2, p. 402, March, 1901 (part) ; List Land and Sea Mamm. NorthAmer., Field Columb. Mus., publ. 57, zool. ser., vol. 2, p. 517, June, 1901(part).
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p. 256,December 27, 1901 (part).
?
Elliot, Land and Sea Mamm. Middle Amer.,Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 581, 1904 (part).?Trouessart, Catal. Mamm. viv. foss., suppl., p. 92, 1904 (part).
?
Elliot,Check List Mamm. North Amer., Field Columb. Mus.. publ. 105, zool. ser.,vol. 6, p. 479, 1905 (part). Stbs'Hens, California Mammals, p. 265, 1906(part).?J. Grinnell, Univ. California Publ. Zool., vol. 5, p. 158, December31, 1908.?Young, Proc. Acad. Nat. Sci. Philadelphia, p. 408, October 14,1908.?Seton, Life-Hist. Northern Anim., p. 1148, 1909 (part).?Wareen,Mammals of Colorado, p. 273, 1910.
?
Cary, North Amer. Fauna, No. 33,p. 206, August 17, 1911.
?
Miller, List North Amer. Land Mamm. 1911, Bull.U. S. Nat. Mus., No. 79, p. 55, December 31, 1912 (part).
?
Grinnell andSwarth, Univ. California Publ. Zool., vol. 10, p. 380, October 31, 1913.?J.Grinnell, Proc. California Acad. Sci., ser. 4, vol. 3, p. 276, August 28,1913 (part).Myotis longicrus interior Miller, Proc. Biol. Soc. Washington, vol. 27, p. 211,October 31, 1914.?H. W. Grinnell, Univ. California Publ. Zool., vol. 17,p. 271, January 31, 1918.?G. M. Allen, Journ. Mamm., vol. 1, pi 4, Novem-ber 28, 1919.?Miller, List North Amer. Recent Mamm. 1923, Bull. U. S.Nat. Mus., No. 128, p. 69, April 29, 1924.Type locaZity.?Twining, Taos County, New Mexico.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 143Tyfe specimen.?^Adult male (skin and skull), No. 133426, UnitedStates National Museum (Biological Survey collection), collected atTwining, Taos County, New Mexico, July 23, 1904, by Vernon Bailey.Original number 8182.Distribution.?More arid parts of the species' range from easternWashington and Oregon to Wyoming, south to Colorado, New Mex-ico, northern Chihuahua and southern California. (See map 10,p. 136.)Diagnosis.?Size essentially as in Myotis volans longicrus; colorlighter and more yellowish.Color.?Nearly " ochraceous-buff " to " ochraceous-tawny " above,pale buffy below ; the bases of the hairs everywhere blackish brown.Ears and membranes blackish.Measurements.?For measurements see tables, pages 146 and 147.Specimens examined.?Total number 123, from the followinglocalities :ARIZONA: CMricahua Mountains, 1 skn (U.S.N.M.), 1 skin (F. M.) ;Cooley's, White Mountains, 1 skin (A. M. N. H.) : Fort Wtiipple, 1 skin(U.S.N.M.) ; Keam Canyon, 1 skin (U.S.N.M.) ; Little Spring, nearFlagstaff, 2 skins, 1 ale. (U.S.N.M.) ; San Francisco Mountain, 1 ale.(U.S.N.M.); Santa Rita Mountains. 1 skin (U.S.N.M.).CALIFORNIA: Benton. Mono County. 1 skin (U. C.) ; Cabazon, RiversideCounty, 1 skin (U. C.) ; Cuyamaca Mountains, San Diego County, 1 skin(U. C.) ; Dudley, Mariposa County. 4 skins (U. C.) ; Dulzura. San DiegoCounty, 1 ale. (A. N. S. P.) ; Dutcli Flat, Placer County. 1 skin (U. C.) ;Fort Tejon. Kern County. 6 skins (F. M.) ; Fyffe, Eldorado County. 2skins (U. C.) ; Hot Springs Pass, Mono County, 1 skin (M. C. Z.) ;Independence. Inyo County. 1 skin (U. C.) ; Inyo Mountains, 2 skins(F. M.) ; Kearsage Pass, Inyo County, 1 skin (U. C.) ; Little Onion Val-ley, Inyo County, 3 skins (U. C.) ; Los Angeles, 1 ale. (U.S.N.M.) ;Monache Meadows. Tulare County, 1 skin (U. C) ; Mount Lassen, 1 ale.(U.S.N.M.) ; Mount Shasta. 1 skin (U.S.N.M.) ; Mount Whitney, 2 skins(M. C. Z.), 5 skins (F. M.) ; Nevada City, Nevada County, 1 ale.(U.S.N.M.) ; Owens Lake, 1 ale. (U.S.N.M.) ; Pasadena, 2 skins (U. C.) ;Panamint Mountains, Inyo County, 4 skins (U. C.) ; Pine Creek, LassenCounty, 2 skins (U.S.N.M.) ; San Bernardino Mountains, 3 skins (U. C), 1skin (A. N. S. P.) ; San Emigdio, Tulare County, 1 ale. (U.S.N.M.) ; SanJacinto Mountains, 1 ale. (U.S.N.M.) ; Santa Rosa Peak, San BernardinoCounty, 3 skins (U. C.) ; Walker Pass, Eldorado County, 1 ale.(U.S.N.M.) ; White Mountains, Inyo County, 5 skins (U. C).CHIHUAHUA: Colonia Garcia, 1 skin, approaching amotus (U.S.N.M.);San Luis Mountains, 1 skin (U.S.N.M.).COLORADO: Coventry, 1 skin (U.S.N.M.); Grand Junction, 1 ale.(U.S.N.M.).IDAHO: Inkom, 1 skin (U.S.N.M.); Malad. 1 skin (U.SN.M.) ; Mission,1 ale. (U.S.N.M.) ; Snake River desert, 3 skins (U.S.N.M.) ; Warren, 1skin (U.S.N.M.).MONTANA: Buffalo, 1 skin (U.S.N.M.) ; Emigrant Gulch, 1 skin, not typi-cal (U.S.N.M.) ; Florence, 1 skin (U.S.N.M.).
144 BULLETIN 144, UNITED STATES NATIONAL MUSEUMNEVADA: Cottonwood Range, 1 skin (U.S.N.M.) ; Humboldt County, 1skin (U. C.) ; Panaca, 1 ale. (U.S.N.M.).NEW MEXICO: Raton Range, 1 skin (U.S.N.M.) ; Cantonment Bergwyn,1 ale. (U.S.N.M.) ; Costilla River, 2 skins (U.S.N.M.) ; Santa ClaraCanyon, 1 skin (U.S.N.M.) ; Santa Fe, 1 ale. (U.S.N.M.) ; Twining, 5skins, ineluding type (U.S.N.M.) ; Willis, 2 skins (U.S.N.M.).OREGON: Fremont, 5 skins, 1 skull, approaehing longicrus (U.S.N.M.);Paulina Lake, 4 skins (U.S.N.M.) ; Silver Lake, 1 skin (U.S.N.M.) ;Ironside, Malheur County, 1 skin (A. M. N. H.).WASHINGTON: Anatone, 4 skins (U.S.N.M.).WYOMING: Afton, 1 skin (U.S.N.M.); Lake Fork, 1 ale. (U.S.N.M.);Laramie. 1 skin (U.S.N.M.) ; Otto, 2 skins (A. M. N. H.) ; RattlesnakeMountains, 1 skin (U.S.N.M.).Remarks.?The type specimen of Myotis volans interior is a brightochraceous buffy identical in color with the type of Myotis ''ca'pi-taneus " representing the small M. volans volans of Lower California.Others from New Mexico, however, are much redder, ochraceoustawny. Palest of all are skins of adults from the Inyo Mountainsof southeastern California and a single skin from 11,000 feet on theeast side of Mount Whitney at the southern end of the Sierra Ne-vadas. These are probably the extreme of this pallid dry-countryrace, the Mount Whitney specimen perhaps a summer immigrantfrom the deserts below. Specimens from Pasadena, PasadenaCounty, southward to southern California seem best regarded asmfeWor, as well as the majority of those from the Sierra Nevada,particularly the more pallid individuals from its southern part. Theseries from the Sierra Nevada is clearly intermediate between theinterior race and M. volo/ns longicrus of the coast, but most of themhave a distinctly tawny hue, instead of the saturate appearance ofthe latter, and it is for this reason that we have included them underthe form intemor. A small series from Old Fort Tejon, Kern County,Calif., is darker than most specimens of interior^ yet not so darkas M. V. longicrus. In the drier parts of eastern Washington simi-lar dull yellowish-brown individuals are found. Three from Anatonein the extreme southeast corner of the State are clearly interiorrather than the saturate smoky form of the coast, while of two fromWhatcom County, one is as dark as M. v. longicrus and the othershows an approach to interior in its decidedly redder cast. EasternOregon affords a similar series, which, on account of their distinctlypaler and reddish to ochraceous tint are placed Avith the latter form,though clearly intermediates.In immature specimens the bright tips of the long hairs are few andshorter than in adults, so that the bases of the hairs everywhere showthrough and darken the general appearance. The pallid colorationof Colorado specimens was noticed by Cary (1911). No. 160595(U.S.N.M.), from Wyoming, lacks the minute p^ on the left-hand
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 145
side. The other small premolar nearly fills the space between canineand p*. MYOTIS VOLANS AMOTUS MUlerMyotis lucifugus longicrus Miller, North Amer. Fanua, No. 13, p. 65, October16, 1897 (part; specimen from Vera Cruz).Mi/Otis longicrus amotus Milleb, Proc. Biol. Soc. Washington, vol. 27, p. 212,October 31, 1914 ; List North Amer. Recent Mamm. 1923, Bull. U. S. Nat.Mus., No. 128, p. 69, April 29, 1924.Type locality.?Cofre de Perote, Vera Cruz, Mexico. Altitude12,500 feet.Type specimen.?Adult female, skin and skull, No. 54437, UnitedStates National Museum, from Cofre de Perote, Vera Cruz, Mexico.Collected May 27, 1893, by E. W. Nelson.Distrihufion.?Southern Mexico, including the States of Vera Cruzand Jalisco; limits of range as yet undetermined. (See map 10,p. 136).Diagnosis.?Similar in size and general coloration to the raceinterior., but the tints of back and belly much richer in tone.Color.?The color of the fur is nearly " ochraceous tawny " above,and " cinnamon brown " below even to the fur about the anal regionwhich in M. v. interior is usually whitish.Skull.?The skull of the type seems a trifle longer with a narrowerbrain case and longer rostrum than in specimens of M. v. lonfficms,while from typical volans it differs in larger size and less shortenedrostrum.Measurements.?For measurements see tables, pages 146 and 148.Specimens examined.?Total number 3, from the following locali-ties:JALISCO: Los Masos, 2 skins (A. M. N. H.).VERA CRUZ: Cofre de Perote, 1 skin, the type (U. S. N. M.).Remarks.?This Mexican form of Myotis volans is of a richer tonethroughout than those whose ranges lie immediately to the north,but it is not darkened to the extent seen in Myotis volnns longicrus ofthe humid northwest coast. The pecularities in color were noticedby Miller (1897) in his review of the North American Vespertilionidse,but the race was not formally separated until 1914. Specimens fromnorthern Chihuahua and Sonora, Mexico, although approachingMyotis volans amotus., seem to be best referred to M. v. intenor whichthey resemble in the golden hue of the long glossy tips of the hairsabove, and in the paler color below, becoming whitish or buffy aboutthe anal region.
146 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis volans
Locality Number ^3 is
Myotis volans volansLower California:Santa Anita..San JorgeMyotis volans longicrusAlberta: Henry House _Washington:Puget SoundPort TowdSendOregon: Baker County.California:Menlo Park..Do.Nicasio.Do.Do.Do.Do.Do.Do.Do.Do.Do.Myotis volans interiorWyoming: Lake ForkColorado: Grand JunctionNew Mexico: Cantonment...Arizona:Little SpringSan Francisco Mountain.New Mexico: TwiningNevada: Panaca..California:Los AngelesMount LassenNevada City...Owens LakeSan Jacinto MountainSan Emigdio..Walker PassMyotis volans amotusVera Cruz: Cofre de Perote
.
148361
' 146046
? 174133
1642714448214254114254259490594955955759560595625956459566595685956959573
55847548465374203098186943 13342628950920631257945312428951628693156929809
'54437
44.447.2
"49"44.847.845.247.243.05L246.847.843.048.248.851.849.648.048.0
51.047.448.450.050.052.846.447.246.251.04a 648.853.450.0
54.0
40.829.4
'39.5"46.242.842.246.241.048.242.840.442.243.445.84345.639.444.6
47.045.642.649.045.838.245.444.442.846.446.846.044.843.4
35.0
19.2 7.6 35.2
18.8...-19.6; 7.418. 8i 8.018. 8i 8.018.619.219.018.018.018.418.418.219.218.418.818.2
20.019.019.020.018.418.618.219.019.020.0i&o19.420.0
18.2, 8.0
37.039.640.039.039.238.6138. 6i38.0,39.4!39. 4j38.038.639.039.038.237.0
39.040.039.037.439.838.041.040.039.040.638."440.441.2
40.0
34.233.8 32.03L8 11.8
33.4 3L037.0' 35.037.0 35.036.0 34.038.037.037.06.0, 36.07.0 37.26. O! 37.6.4| 36.46.0 36.86. 4| 37. 86.0 36.86.0 36.86.2' 35.0
7.0 38.65.6 37.86.8 36.0
5.0
36.838.036.038.236.837.839.238.836.438.038.0
34.434.834.233.434.434.234.233.434.634.8 __..34.0 11.833.6 12.0
12.013.012.812.fi12.413.011.013.012.811.812.612.013 0
35.634.635.036.235.236.035.034.436.836.034.636.436.2
37.8 35.2
13.814.013.611.813.012.012.412.813.013.413.211.613.2
9.0 7.6
10.010.810.210.410.0U.410.010.211. o;10. 0|10.89.810.4
13.211.410.9.410.2
8.49.28.48.48.28.69.09.09.09.09.09.28.48.28.0
9.08.28.210.08.410.211.011.69.69.0U.O
7.07.28.09.09.27.49.0
* Type of Myotis capitanens Nelson and Qoldman.
' Type of Myotis aUifrons Hollister.
? Type.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 147Cranial measurements of Myotis volans
Locality
148 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis volans?Continued
Locality
AMERICAN BATS OF THE GENEEA MYOTIS AND PIZONTX 149
CDS ; 2. M. CALIFOKNICUS CADRINUS ; 6, M. CALIFOKlNicFOENICUS MBXICANDS
150 BULLETIN 144, UNITED STATES NATIONAL MUSEUMon the upper part of the posterior border, below which it is nearly-straight to the basal shoulder which stands off gradually instead ofprojecting abruptly (as in Myotis yumanensis) . The tragus is nar-row, its inner margin straight, and equaling one-half the entireheight of the ear (6 mm.) ; the region of its greatest width is slightlyabove the level of the inner base, whence the posterior margin isgradually beveled to the acute tip. There is a shallow emarginationopposite the inner base, below which is a small rounded lobe.Wing and meinbranes.?The wing membrane arises from the sideof the foot at the base of the toes, just below the ends of the meta-tarsals. The third to fifth metacarpals are very slightly graduatedor the third and fourth may be almost equal in length. Taking thethird finger as 100, the fourth and fifth fingers are respectively aboutas 86 and 79 (58: 50:46 mm.). When the wing is folded the end ofthe third metacarpal usually comes quite to the elbow (or rarelyexceeds it, or falls 0.5 to 1 mm. short) ; that is, the length of thethird metacarpal about equals that of the forearm. The tip of thetail is free. The fur of the body extends slightly on to the mem-branes, especially on the intertibial part of the uropatagium wherethere are numerous scattered hairs dorsally.Foot.?The foot is small, slender, and weak, its length normallyless than one-half that of tibia (average ratio of foot to tibia in 10specimens from the Northwest Coast, 46.6; in 10 from Point Reyesand Nicasio, Calif., 43.1; in 10 from Lower California, 43.4; and in10 from Patzcuaro, 44.9). The calcar is less than the length of thefree border of the uropatagium (about 13:15 mm.) and ends in aprojecting lobule. Its edge usually bears a well-developed keel whichrises abruptly at about the length of the metatarsus from the heel,and gradually tapers off.Fur amd color.?The pelage is long, full, and of fine texture, thehairs with long silky ends which, however, are dull, rarely burnished.The tips of the hairs are usually brown or yellow in tone, stronglycontrasting with their slaty bases both above and below. The dorsalside of the uropatagium is well though thinly haired between thetibiae above, but not quite to one-half their length below. On theventral side the fur extends sparingly outward as far as a line join-ing the knee and the elbow.Skidl.?The skull (pi. 1, p. 7, fig. 14) is delicate and slender, withrelatively long and tapering rostrum, about 3/7 of the total length ofthe skull. The profile of the rostrum rises rather sharply to theforehead and decidedly flat-topped brain case. Sagittal crest usuallyinconspicuous or absent. Lambdoid ridges distinct though low.Teeth.?In general the teeth resemble those of Myotis evotis re-duced in size proportionately to the smaller skull. Upper molars
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZONYX 151
with well-developed hypocone, the base of which is strongly markedoff from that of the protocone. Protoconule usually less developedthan in M. lucifugus^ rarely absent ; paraloph usually well developed,even when the protoconule is vestigial. Metaloph when presentapparently always confined to the bottom of the valley between basesof hypocone and metacone; not infrequently it is absent in one orboth teeth. Cingulum clearly marked, but not passing aroundantero-lingual base of protocone. The small premolars are littlecrowded ; those of the mandible may be spaced, though usually theyare in contact.Remarks.?Myotis califomicus does not seem to extend its rangebeyond the northern border of the United States except for a shortdistance on the Pacific coast. It is apparently represented in easternAsia by Myotis mouplveiisi-s which it closely resembles in size, inthe relatively small and delicate feet with distinctly keeled calcar,and in the long silky pelage with contrasted brownish tips and darkbases. The third metacarpal, however, is not quite so long in theAsiatic species. Since the northward ranges of this species and ofMyotis volans on the west coast of America are nearly coextensive,it may be expected that the ranges of their Asiatic representatives,M. moupinensis and M. frater respectively, will eventually be foundto be so likewise on the other side of the Pacific. Presumably inTertiary times these areas of distribution were continuous. Someconclusion as to the higher temperature then prevailing to the north-ward when the two continents were united, might therefore be de-duced by a comparison of the temperatures now found at the limitsof these species' range on opposite sides of the Pacific.In its area of dispersal as well as in the number and character ofits local forms Myotis califarnictis closely parallels M. yumanenMs.Each species has a dark race on the humid '' northwest coast," apallid race in the desert interior of the western United States, a formintermediate in color occurring over a wide area where the climaticconditions are neither saturate nor excessively dry, and finally arichly colored form in southern Mexico. Specimens of the twoanimals are sometimes not very distinctive in color, but the largefoot of Myotis yimianensls and the very small foot of M. californicusare always diagnostic in such instances.MYOTIS CALIFORNICUS CALIFORNICUS (Audubon and Bachman)Vespertilio califomicus AnoxiBON and Bachman, Jo'urn. Acad. Nat. Sci. Phila-delphia, ser. 1, vol. 8, pt. 2, p. 285, 1842.Vespertilio nitidus H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1862, p. 247(Monterey, California) ; Monogr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 60, fig. 57-59, June, 1864?Dobson, Catal. ChiropteraBrit. Mus., p. 318, 1878.?H. Allen, Monogr. Bats North Amer., Bull.
152 BULLETIN 144, UNITED STATES NATIONAL MUSEUMU. S. Nat. Mus., No. 43 (1893), p. 94, pi. 12, fig. 1-3, 6-10, March 14,1894.
?
Tbouessabt, Catal. Mamm. viv, foss., p. 130, 1897.
?
Lyon andOsgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus.,No. 62, p. 272, January 28, 1909.VespertiUo oregonensis H. Allen, Monogr. Bats North Amer., Smithsonian Misc.Coll., No. 165, p. 61, June, 1864 (Fort Yuma, California, and Cape St. Lucas,Lower California).
?
Lton and Osgood, Catal. Type-Sp. Mamm. U. S. Nat.Mus., Bull. U. S. Nat. Mus., No. 62, p. 272, January 28, 1909.Vespei'tilio exilis H. Allen, Proc. Acad. Nat. Sci. Philadelphia, p. 283, 1866(Cape St. Lucas, Lower California) ; Monogr. Bats North Amer., Bull.U. S. Nat. Mus., No. 43 (1893), p. 97, footnote, March 14, 1894.VespertiUo tenuidorsalis H. Allen, Proc. Acad. Nat. Sci. Philadelphia, p. 283,1866 (Cape St. Lucas, Lower California) ; Monogr. Bats North Amer.,Bull. U. S. Nat. Mus., No. 43 (1893), p. 97, footnote, March 14, 1894.?Lyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat.Mus., No. 62, pp. 273, 291, January 28, 1909.VespertiUo yumanensis H. Allen, Proc. Acad Nat. Sci. Philadelphia, 1866, p.283 (not of H. Allen, 1864).VespertiUo nigricans H. Allen, Monogr. Bats North Amer., Bull. U. S. Nat.Mus., Nq. 43 (1893), p. 97, footnote, March 14, 1894 (not of Maximilianzu Wied, 1826).Myotis caUfornicus Miller, North Amer. Fauna, No. 13, p. 69, October 16,1897.?Trouessart, Catal. Mamm. viv, foss., p. 1283, 1899.
?
Merriam,North Amer. Fauna, No. 16, p. 89, October 28, 1899.
?
Elliot, Synops. Mamm.North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2, p. 403, March1901 ; List Land and Sea Mamm. North Amer., Field Columb. Mus., publ.57, zool. ser., vol. 2, p. 517, June, 1901.
?
Miller and Rehn, Proc. BostonSoc. Nat. Hist., vol. 30, p. 256, December 27, 1901.?Elliot, Field Columb.Mus., publ. 91, zool. ser., vol. 3, p. 319, March, 1904 ; Land and Sea Mamm.Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 578, 1904.
?
Trouessart, Catal. Mamm. viv, foss., suppl., p. 93, 1904.
?
Elliot, CheckList Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6,p. 476, 1905.?Stephens, California Mammals, p. 266, 1906 (part).?Elliot, Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115,zool. ser., vol. 8, p. 502, 1907.?J. Grinnell, Univ. California Publ. Zool.,vol. 5, p. 158, October 31, 1908.Myotis caUfornicus caUfornicus MiiXEai, List North Amer. Land Mamm. 1911,Bull. U, S. Nat. Mus., No. 79, p. 56, December 31, 1912.?J. Grinnell, Proc,California Acad. Sci., sei'. 4, vol. 3, p. 277, August 28, 1913. Grinnell andSwABTH, Univ. California Publ. Zool., vol. 10, p. 381, October 31, 1913.?H. W.Grinnell, Univ. California Publ. Zool., vol. 12, p. 317, December 4, 1914;vol. 17, p. 279, January 31, 1918.
?
Miller, List North Amer. Recent Mamm.1923, Bull. U. S. Nat. Mus., No. 128, p. 70, April 29, 1924.?Grinnell andStorer, Anim. Life in the Yosemite, p. 21, 1924.
?
Strecker, Check-ListMamm. Texas, The Baylor Bulletin, Baylor University, Waco, Texas, vol,29, No. 3, p. 9, August, 1926.Myotis caUfornicus quercinus H. W. Grinnell, Univ. California Publ. Zool.,vol. 12, p. 317, December 4, 1914 (Seven Oaks, San Bernardino Co., Cali-fornia) ; vol. 17, p. 285, January 31, 1918.?J. Grinnell, Univ. CaliforniaPubl. Zool., vol. 21, p. 314, January 27, 1923.?Miller, List North Amer.Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p, 71, April 29, 1924,Type locality.?" California."
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 153Type.?None specified. The original specimen, on which the de-scription was based, was received from " California." It is probablynot now in existence. By the recognizing of the dark northwest-coastform caurinus and the desert form paUidus the name californicus inits subspecific sense has become restricted to the wide-ranging racewhose color is intermediate between these extremes. This race occursin the Monterey region which, as already explained, has been selectedas the type locality of Harrison Allen's Vespertilio nitidus. No
" selection " seems possible in the case of Vespertilio caUfoimious^because Audubon and Bachman unlike Doctor Allen, mentioned nolocalities among which to choose; but for the sake of conveniencethe one region may be treated as though it were the type localityfor both names.Distrihution.?From about the latitude of the Tropic of Cancer incontinental Mexico, and Cape St. Lucas in Lower California north-ward along the Pacific coast to the region of San Francisco Bay andin the interior to the northern Sierra Nevada, eastern Oregon andextreme southeastern Washington, eastward to western Texas, centralNew Mexico and west-central Colorado ; replaced by a pallid race inthe Great Basin. (See map 11, p. 149.)In the San Bernardino Mountains of California Myotis californi-cus califorriicms has been taken at altitudes of nearly 7,500 feet, andin Placer County at 4,000 feet. On the whole it is an inhabitant ofthe lower altitudes. East of the desert divides of southern Californiatrue californicus is replaced by the paler race, pallidum, and on thecoast of northwestern California it merges into the dark M. o.caurhvus.Diagnosis.?Color above, tawny w^ith a distinct reddish or chestnuttint; tail very long, the ratio of its length to that of head and bodyaveraging about 95.Description.?Dorsal surface of head and body " ochraceous-tawny " (Ridgway, 1912) , becoming slightly paler on the head ; be-low, similar but paler varying from a pale wash of the dorsal color-ation to a " pale buff." The bases of the hairs on both surfaces are
" sooty black," except those on the base of the interfemoral membraneabove and those at the anal region below; these are without darkbases. Several skins from Monterey, Calif., agree in an unusual rich-ness of color, even the entire belly being a bright " ochraceous tawny "like the back.As is often the case with reddish species of bats, melanism is notvery infrequent. Several skins fi-om Lower California are very dark,nearly " mummy brown " above without reddish shades. Othersfrom Placer County, Calif., are slightly melanistic, a condition whichmakes them appear almost as dark as M. californicus oaurvnus.58518?28 11
154 BULLETIN 144, UNITED STATES NATIONAL MUSEUMMeasv/rements.?For measurements see tables, pages 161 and 163.Specvmens examined.?Total number 298, from the followinglocalities
:
ARIZONA: Camp Grant, 2 ale. (U.S.N.M.) ; Chiricahua Mountains, 1 skin,1 ale. (U.S.N.M.) ; Huachuea Mountains, 1 skin (F. M.), 1 skin (U.C.) ;Little Meadow, 3 ale. (U.S.N.M.); Nantan Plateau, 25 miles N. B. ofRice, 5,800 feet, 1 skin (U.S.N.M.) ; Oracle, 5 ale. (U.S.N.M.) ; PinalCounty, 1 skin (A. M. N. H.) ; Santa Catalina Mountains, 1 skin(U.S.N.M.) ; S. B. ranch, 1 skin (F. M.).CALIFORNIA: Calaveras County: San Andreas, 1 skin (U. M.) ; EldoradoCounty: FyfEe, 16 skins (U, C.) ; Fresno County: Dunlop, 1 skin (U. C.) ;no exact locality, 1 skeleton (U.S.N.M.); Kern County: Fort Tejon, 3skins, 1 skull (F. M.), 7 ale., 1 skeleton (U.S.N.M.) ; San Emigdio Creek,1 skin (U. C.) ; San Emigdio Canyon, 1 skin (U.S.N.M.) ; Tejon Pass,1 ale. (U.S.N.M.) ; no exact locality, 1 ale. (U.S.N.M.) ; Los AngelesCounty: Los Angeles, 1 skin (A. M. N. H.) ; Pasadena, 1 skin (U. C.) ;Marin County: Nieasio, 18 ale. (U.S.N.M.) ; Petaluma, 1 ale. (U.S.N.M.) ;Point Reyes, 10 ale. (U.S.N.M.); Mariposa County: Dudley, 9 skins(U. C.) ; Pleasant Valley, 3 skins (U. C.) ; Varain, 1 skin (U. C.) ;Yosemite Valley, 1 skin (U. C.) ; Monterey County: south of Chalk Peak,3 skins (U. C.) ; Monterey, 5 skins (U. C), 1 skin, lectotype of nitidus(U.S.N.M.) ; Paraiso, 1 skin (U.S.N.M.) ; Soledad, 1 skin (U. C) ; SurRiver, 1 skin (U.S.N.M.) ; Placer County: Blue Canyon, 1 skin (U. C.) ;Dutch Flat, 2 skins (U. C.) ; Michigan Bluff, 1 skin (U. C.) ; RiversideCounty: Banning, 1 ale. (U.S.N.M.); Capistrano, 1 skin (U.S.N.M.);Kenworthy, 1 skin (U. C) ; San Bernardino County: Bear Lake, 6,700feet, 1 skin (U. C.) ; Fish Creek, 1 skin (U. C.) ; San Clemente Island,7 ale. (U.S.N.M.) ; Santa Ana River, 1 skin (U. C.) ; Seven Oaks, 2 skins,Including type of queroinus (U. C.) ; San Diego County: CuyamacaMountains, 1 skin (U. C.) ; Dulzura, 2 skins (U.S.N.M.), 3 skins (A. M.N. H.), 8 ale. (A. N. S. P.) ; Julian, 4 skins (U. C.) ; San Diego, 2 skins,melanistic (U. C.) ; Santa Ysabel, 4 ale. (U.S.N.M.) ; Witch Creek, 9 ale.(U.S.N.M.) ; no exact locality, 2 skins (U.S.N.M.) ; Santa Clara County:Pacheco Pass, 2 skins (U.S.N.M.) ; Santa Cruz Island: Friar's Harbor,3 skins, not typical (U. C.) ; Shasta County: Fort Crook, 1 ale.(U.S.N.M.) ; Siskiyou County: Mount Shasta, 1 ale. (U.S.N.M.) ; SolanoCounty: Vacaville, 3 skins (U. C.) ; Sonoma County: 7 miles west ofCazadero, 2 skins (U. C.) ; no exact locality, 1 skin (U. C) ; TrinityCounty: Trinity Mountains, east of Hoopa, 1 skin (U.S.N.M.); TulareCounty; East Fork Kaweah River, 4 ale. (U.S.N.M.); Mount Whitney,4 skins, 1 skull (F. M.) ; Trout Creek, 6,000 feet, 2 skins (U. C.) ; VenturaCounty: Matilija, 1 skin (U. C.) ; Mount Pinos, 2 skins (U. C.) ; SantaBarbara, 1 ale. (U.S.N.M.).CHIHUAHUA: Pacheco, 2 skins (M. C. Z.) ; San Luis Mountains, 2 skins(U.S.N.M.).COAHUILA: Guadalupe, 1 skin (U.S.N.M.).COLORADO: Rifle, Garfield County, 1 skin (U.S.N.M.).LOWER CALIFORNIA: Cape St. Lucas. 2 ale, types of cxilis and tenui-dorsalis H. Allen (A. N. S. P.) ; Comondu, 2 skins (U.S.N.M.) ; Matancita,1 skin (U.S.N.M.) ; San Josg del Cabo, 1 skin (U.S.N.M.) ; San Quintin,1 skin (U.S.N.M.): Santa Anita, 3 skins, 52 ale. (U.S.N.M.), 4 skins(B. M.).
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 155NEW MEXICO: Animas Valley, 1 skin (U.S.N.M.) ; Apache, 2 ale.(U.S.N.M.) ; San Andres Mountains, 5 skins (U.S.N.M.) ; San MateoCanyon, 1 skin, 1 ale. (U.S.N.M.).OREGON: Blue River, 1 skin, 1 ale. (U.S.N.M.) ; Elgin, 1 ale. (U.S.N.M.) ;Eugene, 1 skin, approaching caurinus (U.S.N.M.) ; John Day River (nearCrown Rock), Wheeler County, 3 ale. (U.S.N.M.) ; Mount Hood, 2 skins,approaching caurinus (U.S.N.M.) ; Sisters, 3 skins (U.S.N.M.) ; WallowaLake, 1 ale. (U.S.N.M.).SONORA: Providencia Mines, 1 skin (F.M.) ; San Luis Mountains, 2 skins(U.S.N.M.).TEXAS: Fort Davis, 1 ale. (U. C.) ; Paisano, 1 ale. (U.S.N.M.).WASHINGTON: Almota, 1 ale. (U.S.N.M.) ; Anatone, 1 skin (U.S.N.M.) ;Goldendale, 1 ale. (U.S.N.M.) ; Lyle, 2 skins (U.S.N.M.) ; Orondo, 1 ale.(U.S.N.M.).Remarks.?A series of skins from Lower California, although per-haps on the ayerage a very little paler than the average of those fromsouthern California west of the desert region, can nevertheless beperfectly matched among the latter. The names exilis and tenuidor-salis applied by Harrison Allen to specimens of this species fromCape St. Lucas are therefore placed in the s3^nonymy of Myatis cali-fornicus californicus. The types of both of these are in the collectionof the Academy of Natural Sciences at Philadelphia. We have notbeen able to find sufficient ground for recognizing the Myotis cali-fornicus quercirms described by Mrs. Grinnell.MYOTIS CALIFORNICUS CAURINUS MillerVespertilio nitidus Merriam, Amer. Nat., vol. 29, p. 860, September, 1895.Myotis californicus caurinus Miller, North Amer. Fauna, No. 13, p. 72, October16, 1897.
?
Trouessart, Catal. Mamm. viv. foss., p. 1284, 1899.
?
^Eluot,Synops. Mamm. North Amer., Field Columb. Mus., publ. 45, zool. ser., vol. 2,p. 404, March, 1901; List Land and Sea Mamm. North Amer., FieldColumb. Mus., publ. 57, zool. ser., vol. 2, p. 517, June, 1901.
?
Osgood, NorthAmer. Fauna, No. 21, p. 37, September 26, 1901.
?
Miller and Rehn, Proc.Boston Soe. Nat. Hist., vol. 30, p. 257, December 27, 1901.
?
Trouessart,Catal. Mamm. viv. foss., suppl., p. 93, 1904.^
?
Stone, Proc. Acad. Nat. Sci.Philadelphia (July, 1904), p. 579, November 17, 1904.?Elliot, CheckList Mamm. North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6,p. 477, 1905; Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ.115, zool. ser., vol. 8, p. 503, 1907.
?
Lyon and Osgood, List Type-Sp. Mamm.U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 270, January 28, 1909.?Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79,p. 57, December 31, 1912.?H. W. Grinnell, Univ. California Publ. Zool.,vol. 17, p. 432, April 25, 1918.
?
Miller, List North Amer. Recent Mamm.1923, Bull. U. S. Nat. Mus., No. 128, p. 70, April 29, 1924.Myotis californicus californicus J. Grinnell, Proc. California Acad. Sci., ser.4, vol. 3, p. 277, August 28, 1913 (part).?H. W. Grinnell, Univ. CaliforniaPubl. Zool., vol. 17, p. 279, January 31, 1918 (part).?J. Grinnell, Univ.California Publ. Zool., vol 21, p. 314, January 27, 1923 (part).Type locality.?Massett. Queen Charlotte Islands, British Colum-bia, Canada.
156 BULLETIN 144, UNITED STATES NATIONAL MUSEUMType.?Adult male (in alcohol), No. 72219 United States NationalMuseum (Biological Survey collection), collected at Massett, GrahamIsland, Queen Charlotte Islands, British Columbia, in 1895, byJ. H. Keen.Distribution.?Humid area of the Pacific coast from the extremesouth of the Alaskan archipelago, along the coastal areas of BritishColumbia, Washington, Oregon, and northwestern California, tothe vicinity of San Francisco Bay. (See map 11, p. 149.)Diagnosis.?Similar to the typical race of Myotis califamicus butcolor darker.Color.?General color above nearly "argus brown" of Ridgway(1912); below, similar but slightly paler, about "Sudan brown."Ears, wings and membranes blackish. Area about the eye, upperlip, and chin covered with shorter stiff dark blackish-brown hairs,but their color not in very noticeable contrast. Basal part of thefur everywhere deep " plumbeous black."Measurements.?For measurements see tables, pages 161 and 163.Specimens examined.?Total number 92, from the following lo-calities :ALASKA: Howkan, Long Island, 2 skins (U. C).BRITISH COLUMBIA: Comox, 1 skin (U.S.N.M.) ; Massett, Queen Char-lotte Islands, 18 ale. including type (U.S.N.M.), 2 ale. (B. M.) ; PortMoody, 1 skin (U.S.N.M.) ; Skidegate, Queen Charlotte Islands, 3 skins(U.S.N.M.) ; Sumas, 1 skin (P. M.).CALIFORNIA: Contra Costa County, Walnut Creek, 3 skins (U. C.) ;Mendocino County, Cahto, 1 ale. (U.S.N.M.) ; Lagunitas, 1 skin (U.S.N.M.) ;Willits, 1 skin (U. C.) ; Mount Sanhedrin, 5 skins (A. N. S. P.) ; SanFrancisco County, 1 skin (U. C.) ; San Mateo County, Menlo Park, 1 skin(U. C), 1 ale. (U.S.N.M.) ; Sonoma County, Guerneville, 2 skins (U. C).OREGON: Brownsboro, 1 skin (U.S.N.M.) ; Corvallis, 1 skin (U. C.) ; Look-ingglass, 1 skin, not typical (U.S.N.M.) ; Marmot, 1 skin (U.S.N.M.) ;McKenzie Bridge, 1 skin (U.S.N.M.) ; Philomath, 2 skins (U.S.N.M.) ;Prospect, 1 skin, not typical (U. C.) ; Reston, 1 skin, nearly typical(U.S.N.M.) ; Tillamook, 1 skin (A. M. N. H.), 2 skins (A. Walker).WASHINGTON: Ashford, 1 skin (U.S.N.M.) ; Bartholomew, 7 skins (S. H.Lyman) ; Blue Creek, Stevens County, 1 ale. (U.S.N.M.) ; Carson, 2 skins(U.S.N.M.) ; Castle Rock, 1 skin (U.S.N.M.) ; Chelan, 1 ale. (U.S.N.M.) ;Colville, 1 ale. (U.S.N.M.) ; Fort Steilacoom, 1 skin (B. M.), 2 skins, 1 ale.(U.S.N.M.) ; Godman Springs, 6 skins (S. H. Lyman) ; Hompeg Falls,1 skin (S. H. Lyman) ; Hoodsport, 2 skins (U.S.N.M.) ; Lake Cushman(Mason County), 3 ale. (U. M.) ; Mount Ranier, 1 skin (U.S.N.M.) ;Prescott, 1 skin (U. C.) ; Puget Sound, 2 ale. (U.S.N.M.) ; Tenino, 1 ale.(U.S.N.M.) ; White Salmon, 1 skin (U.S.N.M.).Remarks.?The northwest coast race is not very strongly differ-entiated from true Myotis californicus, and many specimens arefound that are difficult to refer unhesitatingly to one or the other.In general, however, M. c. caurinu^ is darker, with a smoky-brownappearance due to the more uniformly dark tips to the hairs, whereas
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 157in M. c. oalifamicus these brown tips distinctly pale out where theymeet the plumbeous basal portion, giving a lighter, redder effect tothe whole. IttYOTIS CALIFORNICUS PALLmUS StephensMyOtis caUfomicus Millek, North Amer. Fauna, No. 13, p. 69, October 16,1897 (part).?Bailey, North Amer. Fauna, No. 25, p. 208, October 24, 1905(part).?Caby, North Amer. Fauna, No. 33, p. 208, August 17, 1911 (part).?Waeeen, Mammals of Colorado, p. 274. 1910 (part).Myotis caMforniciis paUidJiis Stephens, Proc. Biol. Soc. Washington, vol. 13,p. 153, June 13, 1900.
?
Elliot, Synops. Mamra. North Amer., Field Columb.Mus., publ. 45, zool. ser., vol. 2, p. 405, March, 1901 ; List Land and SeaMamm. North Amer., Field Columb. Mus., publ. 57, zool. ser., vol. 2, p. 517,June, 1901.
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 80,p. 256, December 27, 1901.
?
Elliot, Field Columb. Mus., publ. 91, zool. ser.,vol. 3, p. 319, March, 1904.
?
Tbouesbabt, Catal. Mamm. viv. foss.. suppl.,p. 93, 1904. Elliot, Land and Sea Mamm. Middle Amer., Field Columb.Mus., publ. 95, zool. ser., vol. 4, p. 579, 1904 ; Check List Mamm. NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 477, 1905.
?
Stephens, California Mammals, p. 266. 1900.
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Eu.iot, Catal. Mamm. FieldColumb. Mus., Field Columb. Mus., publ. 115, zool. ser., vol. 8, p. 503, 1907.?Lyon and Osgood, Catal. Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S.Nat. Mus., No. 62, p. 291, January 28, 1909?Milleb, List North Amer.Land Mamm. 1911, Bull. U. S. Nat. Mus., No. 79, p. 57, December 31,1912.?J. Grinnell, Proc. California Acad. Sci., ser. 4, vol. 3, p. 277, August28, 1913; Univ. California Publ. Zool., vol. 12, p. 265, March 20, 1914.?H. W. Grinnell, Univ. California Publ. Zool., vol. 17, p. 288, January 31,1918.?J. Grinnell, Univ. California Publ. Zool., vol. 21, p. 314, January 27,1923.?Miller, List North Amer. Recent Mamm. 1923, Bull. U. S. Nat.Mus., No. 128, p. 70, April 29, 1924.Type JoccMty.?Vallecito, San Diego County, Calif,Type specimen,?Adult male (skin and skull), No. 99829 UnitedStates National Museum (Biological Survey collection), collected atVallecito, San Diego County, Calif., April 1, 1895, by FrankStephens.Distribution.?Desert regions of the Great Basin. (See map 11,p. 149.)Diagnosis.?A pale desert race well characterized by the tricolorpattern of the hairs of the back. These have a dark base, succeededby a whitish portion, and fulvous tip ; tail long.Color.?General color above (topotypes), very pale, nearly "lightochraceous buff" (Ridgway, 1912). The hairs of the body are
" plumbeous black " at the base, with a ring of buffy whitish succeededby a pale ochraceous tip. The short fur at the back of the ears iswhitish to the base, that on the interfemoral membrane pale och-raceous throughout. The short stiff hairs of the lips and about theeyes are slightly darker brownish. Below, the long silky tips ofthe hairs are everywhere clear dull white to " pale buff "; those at thesides on the membrane are of this tint throughout, elsewhere the
158 BULLETIN 144, UNITED STATES NATIONAL MUSEUMbases of the hairs are "plumbeous black." Ears and membranespale brownish, the interfemoral very pale at the base.Measurements.?The measurements of the type were : Total length,80 mm.; tail, 42; tibia, 15; ear, 11; forearm, 30; expanse of wings,208. A topotype measured by the describer: Total length, 81; tail,39; ear, 12; expanse, 221; its forearm is 31.5; knee to end of claw,foot extended, 20. For detailed measurements, see tables, pages 161and 164.Speciviens examined.?Total number 108, from the followinglocalities
:
ARIZONA: Beale Spring, 2 ale. (U.S.N.M.) ; Big Sandy Creek, 1 ale.(U.S.N.M.) ; Bill Williams River, 5 ale. (U.S.N.M.) ; Colorado River,Mellen, 2 skins (U. C.) ; Fort Defiance, 1 ale. (U.S.N.M.) ; Fort Verde,1 ale. (A. M. N. H.) ; Keam Canyon, 1 skin (U.S.N.M.) ; San FranciscoMountain, 1 skin (U. C.) ; Santa Catalina Mills. 2 ale. (A. M. N. H.) ;Tinajas Altas, Yuma County, 3 skins (U.S.N.M.) ; Yuma, 2 ale. (U.S.N.M.).CALIFORNIA: Imperial County: Colorado Desert, 1 ale. (U.S.N.M.) ; FortYuma, 1 ale. (U.S.N.M.) ; Pilot Knob, 3 skins (U.S.N.M.) ; Inyo County:Amargosa River, 1 ale. (U.S.N.M.) ; Bennett's Wells, 1 ale. (U.S.N.M.) ;Death Valley, no exact locality, 10 ale. (U.S.N.M.) ; Funeral Mountains,1 ale. (U.S.N.M.) ; Furnace Creek Ranch, 2 skins (U. C.) ; Lone PineCreek, 1 skin (U. C.) ; Saratoga Springs, 6 ale. (U.S.N.M.) ; MesquiteValley, 2 skins (F. M.) ; Pauamint Mountains, 4 skins (F. M.) ; PanamintValley, 1 skin (U. C.) ; San Bernardino County: Lavie, 2 skins (U. C.) ;opposite Needles, 2 skins (U. C.) ; San Diego County : No exact locality, 3ale. (U.S.N.M.) ; Borego Spring, 1 skin (U.S.N.M.) ; Colorado Desert, 1ale. (U.S.N.M.) ; La Puerta Valley, 4 skins (U. C.) ; Valleeito, 4 skins(U. C).COLORADO : Ashbaugh's Ranch, Montezuma County, 1 skin, 2 ale.(U.S.N.M.).LOWER CALIFORNIA: San Simon, 2 skins (A. M. N. H.) ; ?San PedroMartir Mountains, 20 ale. (A. M. N. H.).NEVADA: Colorado River, 1 ale, (U.S.N.M.); Cottonwood Range, Hum-boldt County, 2 ale. (U.S.N.M.) ; Gold Mountain, 2 ale. (U.S.N.M.) ;Little High Rock Canyon, Washoe County, 2 skins (U. C.) ; PahrumpValley, 1 ale. (U.S.N.M.) ; Rabbit Hole Mountains, 1 skin (U.S.N.M.) ;Vegas Valley, 1 ale. (U.S.N.M.).UTAH: Kanab, 1 skin (U.S.N.M.).Remarks.?In typical specimens of Myotis califomicvs pcMidns thetricolor pattern of the dorsal fur is a striking characteristic due tothe extent of the pale ring on the individual hairs, whose ends arereddish as in true califoriiicus rather than golden as in M. c.mexicamis. Many intermediates occur where the forms meet.It may be questioned whether the name Vespertilio oregonensisH. Allen is not applicable to this race. This name was given in asomewhat ambiguous way to a skin without locality, two skins fromYuma, Arizona, and one from Cape St. Lucas, Lower California. Inview of the vague manner in which it was published and the fact
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 159that one of the specimens included under it (Cape St. Lucas) is refer-able to true Myotis califor^iicits, we prefer to consider the name asynonym of ccdifomicus as has already been done by Miller (1897).MYOTIS CALIFOKNICUS MEXICANUS (SauBsure)Vespertilio mexicanus Saussube, Rev. et Mag. de Zool., ser. 2, vol. 12, p. 282,1860.Vespertilio nigricans Alston, Biol. Centr.-Amer., Mammalia, p. 206, 1881 (notof Wied).Vespertilio agilis H. Allejn, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 282(Mirador, Vera Cruz, Mexico).
?
Lyon and Osgood, List Type-Sp. Mamm.U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 290, January 28, 1909.Vespertilio nitidus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 3, p. 177,December 10, 1890.Myotis califomicus mexicanus Millee, North Amer. Fauna, No. 13, p. 73,October 16, 1897.
?
Miller and Rehn, Proc. Boston Soc. Nat, Hist., vol. 30,p. 257, December 27, 1901.
?
Elliot, Land and Sea Mamm. Middle Amer.,Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 579, 1904.
?
Tbouessabt,Catal. Mamm. viv. foss., suppl., p. 93, 1904.
?
Elliot, Check List Mamm.North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 477, 1905.
?
J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 22, p. 260, July 25, 1906.?Miller, List North Amer. Land Mamm. 1911, Bull. U. S. Nat. Mus., No.79, p. 57, December 31, 1912; List North Amer. Recent Mamm. 1923, Bull.U. S. Nat. Mus., No. 128, p. 71, April 29, 1924.Myotis nigricans J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 22, p. 260,July 25, 1906 (not of Wied; specimens from Jalisco).
?
Nexson, NorthAmer. Fauna, No. 14, p. 18, April 29, 1899 (not of Wied; specimen fromTres Marias Islands).Type locality.?Exact locality unknown, but the type specimenwas collected somewhere in the warmer part of the State of Mexico.Saussure says of the animal : " Habite les parties chaudes du Mexique.J'ai pris ce Vespertilion dans les terres chaudes de la province deMexico."Type specimen.?The description was based on an alcoholic speci-men collected by H. de Saussure. If still in existence it is probablyin the Museum of Natural History at Geneva, Switzerland, wheremost of Saussure's Mexican material is preserved.Distribution.?Southern Mexico, from Oaxaca north to about theTropic of Cancer. (See map 11, p. 149.)Diagnosis.?Slightly larger than typical Myotis coliformcus; gen-eral color tending toward a dull orange brown rather than chestnut
;
tail less elongated than in the other races, its ratio to head and bodyaveraging about 90.Color.?There is a tendency to dichromatism in this race. In thedarker phase the general color is dull brown, of much the same shadeas in Myotis lucifugus., nearly "Brussels brown" (Ridgway, 1912),slightly darker on the head; below, the chin, throat, and chest are
160 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
similar but paler, becoming dull whitish at the anal region. Thehair about the eyes and upper lips is blackish brown; ears andmembranes the same. Bases of the hairs above and below " plum-beous black." In the lighter phase, the upper surface is a lively
" Sudan brown," paling to dull " ochraceous buff " below. Twoimmature specimens from Patzcuaro are very much darker than theadults, sooty above with the tips of the longer hairs on the backreddish; lighter below, clouded with brownish. In this pelage theyare very similar to adults of the northern form of Myotis nigricans,but are distinguishable by the more conspicuous dark underfur onback, smaller foot, and large teeth.Skull.?In general form, the skull is quite like that of true Myotiscalifomicits, with delicate rostrum, brain case high, oval, tapering atthe front end and not flat topped like that of M. subulatus and itsraces. In the Patzcuaro series, it is very distinctly larger than intnie coHifoTnicus, or indeed than in the specimens from Oaxaca.Measurements.?For measurements see tables, pages 162 and 164.Specimens exannined.?Total number 78, from the following
^calities
:
GUANAJUATO: Guanajuato, 1 ale. (A. N. S. P.).JALISCO: Los Masos, 6 skins (A. M. N. H.) ; Santa Rosalia, 1 ale. (B. M.) ;Sierra Nevada de Colima, 9 skins (A. M. N. H.).MICHOACAN: Patzeuaro, 8 skins, 41 ale. (U.S.N.M.).OAXACA: Reyes, 5 skins (U.S.N.M.) ; Cuicatlan, 1 ale. (U.S.N.M.).SAN LUIS POTOSI: La Parada (Hda.), 1 ale. (U.S.N.M.).TAMAULIPAS: Miquihuana, 2 ale. (U.S.N.M.).TRES MARIAS ISLANDS: 1 ale. (B. M.).VERA CRUZ: Mirador, 1 ale., type of agilis (A. N. S. P.).ZACATECAS: San Juan Capistrano (Hda.), 1 ale. (U.S.N.M.).Remarks?Saussure's description of a small Myotis occurring inthe hot part of the Province of Mexico, and having a forearm of33 mm. in length, is taken to refer to the representative of Myotiscalifamicus here described. The color of the type, after immersionin alcohol, was said to be golden brown, grayish below. This ex-cludes the possibility that Saussure had in hand a form of M.nigricans.The series from Patzcuaro, Michoacan, seems typically large anddark. Of similar large size is the type specimen of VespertiUo agilisH. Allen, from Mirador, Vera Cruz ; hence this name is undoubtedlya synonym of mexicanu^. A series of skins from Reyes, Oaxaca, isslightly but distinctly smaller, and the skulls average a little less insize.
AMERICAN BATS OF THE GENEEA MYOTIS AND PIZONYX 161External measurements of Myotis califomicus
Locality
162 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis caWfomtcMS?Continued
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 163Cranial measurements of Myotis californicus
Locality Number -. to =s5!S-? ?a CI 2rt c? "5.
Myotis californicus californ-icusCalifornia:Ni<?sioDoDoDoDoSan Clemente IsDoCapistranoSan Emigdio CanyonSan Bernardino Mountains.Pacheco PassDoMount WhitneyDoDoFort TejonDoDoDoLower California:Santa AnitaDoDoDoDoDoDoDo.DoDoDoDoDoDoDoDoDoDoComonduDoSan QuintinMatancitaSan Jos? del CaboChihuahua:San Luis MountainsDoMyotis californicus caurinusBritish Columbia:MassettDo...Do... ,ComoxPort MoodySumasWashington:Chelan.Mount Rainier..Fort Steilacoom. ,DoEast of ColvilleOregon:McKenzie BridgeMarmot P. O.
'Type.
59497 U.S.N.M.59496594985949960463616836168214990312886115044715064915057813300 F.M.133031330413302133061330738669 U.S.N.M.98.3. 1.25 B.M.26606193556 U.S.N.M.1469451483441483551469461483451483461483491483501483511483521483531483541483571460451460471389451461201466953657336576
1 72219100682100681146765669347214 F.M.303148958916583802238032204387802 IS
12.813.013.613.21212.812.612.813.013.013." 313.313.013.11.3.113.612.613.613.6131.3.413.113.61.3.013.213.113.613.813.014.013.513.213.213.313.41314.013.113.214.0
12.0....12.8....12.8 7.813.0: 8.212.212.412. 2'12! 21 8.12. 6' 8.12. 4i 8.012.613.212.612.412.212.412.412.411.812.812.612.812.612.412.612.212.012.012.412.812.412.212.213.012.212.812.412.012.612.41.3.012.012.412.8
12.8 12.013.0 12.213.212.813.013.31.3.214.012.613.0
12.812.412.212.612.413.012.012.212.213.2 13.013.2 13.0
8.0
7.8
3. 6. 43.0' 6.8
8.0 3.27.6 3.2
4.2 9.04.24.2 9.44.6 0.49.09,'69.09.29.29.010.0
9.'
2
9.29.09.29.28.8
9.210.0
4.85.05.25.05.05.05.04.84.85.05.05.4'..5.0! 5.6.0
5.15.45.05.25.05.05.05.05.05.25.05.05.25.25.25.05.15.5.05.05.15.35.0 5.2
164 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis caUfomicus?CJontinued
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 165Winff and membranes.?The wing membrane arises from the side ofthe foot at the ends of the metatarsals. Metacarpals subequal, thethird usually but very little longer than the fourth and fifth, whichare about equal in length though sometimes slightly graduated. Inone or two cases the fifth just exceeds the fourth, in another the thirdand fifth are equal, exceeding the fourth, while in two other speci-mens the third and fourth are equal, the fifth a trifle shorter, andin another all three are of equal length. In a wing showing slightgradation of these bones, the fourth and fifth fingers are to thethird as 84 and 79 to 100; in a second individual having the thirdand fifth metacarpals equal, the fourth and fifth finger are practi-
MAP 12. DiSTBIBDTION OF MtOTIS SUBDLATUS ; 1, M. SUBULATDS SDBOLATDS ; 2, M.SUEULATDS MELANORHINDS ; 3, M. SUBULATDS LBIBII
cally equal, and stand in the ratio of 84 and 83 respectively to thethird. When the wing is folded, the third metacarpal usually fallsshort of the elbow by 1 mm. or a little more, and individuals inwhich this metacarpal equals the forearm are less frequently metwith than in Myotis califoi^nicus. The extreme tip of the tail is free.The fur of the body extends, as in M. califomicus^ slightly upon themembranes, reaching the level of the distal fourth of the interfemoralabove. Below, it extends to a line joining the middle of the humerusand the knee, and from knee to knee on the lower side of the inter-femoral membrane.Foot.?The foot is small and delicate, essentially like that of Myotiscalifomicv,s. Its length is usually less than one-half that of the tibia
166 BULLETIN 144, UNITED STATES NATIONAL MUSEUM(ratio in 16 specimens, 43.4). The calcar is long and slender, equal-ling or exceeding the free border of the interfemoral, and ends ina small projecting lobule. Its edge has a prominent keel, that riseswith gentle slope at about the length of the metatarsus from thefoot. The length of the leg and foot from knee to end of extendedclaws is usually about 21 to 23.5 mm.Fur and color.?The pelage is even more full and silky than inMyotis califomicus^ the tips of the long hairs toning on flaxen oryellow rather than on the dull chestnut of the latter. The moststriking differences between the two are: (1) the frequently bur-nished tips of the hairs in Myotis subulaius as contrasted with theusually dull lusterless pelage of M. californicus^ and (2) the blackface and ears of M. subulatus giving a masked appearance, whereasin M. californicus^ although the lips and ears may be dark brownthey do not ordinarily show out in sharp contrast with the restof the pelage. The wing membranes of M. subulatvs are usuallymore blackish than those of M. coHifornicus.Skull.?The skull is delicate and slender, proportioned much as inMyotis califomicics, but it is slightly longer and broader (comparepi. 1, p. 7, figs. 15 and 14). It differs notably from that of M. cali-famicus in the breadth and flatness of the brain case, so that viewedfrom behind the occiput (pi. 1, p. 7, fig. 11) appears to be less elevatedthan in the smaller animal, and the summit is actually broaderand more flat ; there is no abrupt step from rostrum to forehead, buta very gradual upward slope.On account of the unusual cranial breadth the temporal ridgesdo not meet until in adult life ; they first come together well forwardof the occiput, cutting off an isosceles triangle with long taperingpoint and sometimes forming a low but sharply defined sagittalcrest. The base of the triangle projects back with convex outlineslightly behind the level of the lambdoid crests.Teeth.?In form the teeth do not differ appreciably from thoseof Myotis californicus. The crown area of the upper molars is,however, appreciably greater than in the small races of M. cali-fornicus occurring north of southern Mexico. Taking specimensof the two species from localities in the United States it is seen thatthe crown of rn? in M. californicus usually measures 1.15 to 1.25 by1.45 to 1.60, while in M. suhulatus the usual dimensions are 1.25 to1.35 by 1.60 to 1.85, a difference which soon becomes obvious to theeye even without direct comparison.Remarks.?This bat, the animal originally described by Say in1823 as Vespertilio suhulatus (not the Vespertilio suhulatus of Har-rison Allen, 1864, and the Myotis subulatus of recent authors), seemsto be uncommon in the eastern United States, though of generaldistribution. It has usually been confused with Myotis lucifugus
AMEBIOAN BATS OP THE GENERA MYOTIS AND PIZONYX 167in the eastern part of its range as it has been with M. caZiformcusin the west. As compared with Myotis lucifugus it is distinguishableby smaller size, small foot, and the distinct keel on the calcar; ascompared with Myotis caZifo7'nicus, by longer thumb (wrist andthumb together usually 8 to 8.5 mm. instead of 6.5 to 7 mm.), larger,more flattened skull, larger teeth (except as compared with M.califomicus mexicanus) and, in a general way, though not invariably,by more glossy dorsal fur.In the east it was first recognized by Audubon and Bachman, who,in 1842 described the animal as Vespertilio leibii on the basis of aspecimen sent by their correspondent Leib from Erie County, Ohio(then Michigan). Their description brings out the characteristicblack ears and wings, long tail, very small feet, and small size ascompared with Myotis lucifugus^ which, under the name Vespertiliovirginianibs, they redescribed in the same paper. So little knownhas the species remained in the region east of the Mississippi Riverthat the name leihii has been either lost sight of entirely or regardedas a synonym of lucifugus. More recently (1913) the name Myotisinnnemana was given to the eastern race of the species on the basisof specimens from Maryland and Vermont which Dr. E. W. Nelsonsaw were not referable to Myotis lucifugus.In 1886 Dr. C. Hart Merriam redescribed the typical arid-plainsform as Vespertilio ciliolahrumi, and four years later he applied thename melanorhinus to another race from Arizona. While ciliolabi^uimhas been currently recognized as a race of califomicus the namemelanorhinus has been placed in the synonymy of Myotis califomicuscalifomicus. In 1893 H. Allen, apparently again noticing the con-trast between Myotis suhulatus Tnelanorhinus and M. califomicus.,suggested the subspecific name henshawii for two specimens fromWingate, N. Mex. Almost exactly the same thing happened 10 yearslater when Elliot described his Myotis orinonvus from Lower Cali-fornia.Not until 1918 were the characters which distinguish Myotis subu-latus from Myotis califomicus clearly recognized. They were thenpointed out in detail, so far as the forms of the two species occurringin California are concerned, by Mrs. Grinnell, who did not, however,realize that the oHnoTnus of Elliot was a race of the species describedas subulatus by Say.This animal bears somewhat the same relation to Myotis califor-nicus that M. lucifugus bears to M. ywnanensis, in being slightlylarger of body, with often distinctly burnished instead of ratherconstantly lusterless pelage, and in having a larger, flatter skull.Furthermore, the contrast in ranges is similar, for while Myotis suhu-latus and Myotis lucifugus extend as species quite across the conti-nent, their lesser counterparts, M. califomicibs and M. yumamensis,
168 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
respectively, are of more restricted distribution, from the humidnorthwest coast eastward across the Great Basin and southward intoMexico.The name suhidattts has been almost universally misapplied to theeastern long-eared Myotis, an animal which should be known asMyOtis keenii septentrionalis. (See p. 107.)MYOTIS SUBULATUS SUBULATUS (Say)Veapertilio suhulatus Say, Long's Exped. to Rocky Mts., vol. 2, p. 65 (footnote),1823 (not of LeConte, 1855, = lucifugus, or Harrison Allen, 1864, = keenii).Vespertilio ciliolahrum Mereiam, Proc. Biol. Soc. Washington, vol. 4, p. 2,December 17, 1886 (near Banner, Trego County, Kans.).Vespertilio nitidus ciliolabrum H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 101, March 14, 1894 (part).?Teouessabt,Catal. Mamm. viv. foss., p. 130, 1897.Myotis caUfomicus ciliolabrum Miller, North Amer. Fauna, No. 13, p. 72,October 16, 1897 (part).
?
Elliot, Synops. Mamm. North Amer., FieldColumb. Mus., publ. 45. zool. ser., vol. 2, p. 404, March, 1901 ; List Landand Sea Mamm. North Amer., Field Columb. Mus., publ. 57, zool. ser.,vol. 2, p. 517, June. 1901.
?
Millek and Rehn, Proc. Boston Soc. Nat. Hist.,vol. 30, p. 257, December 27, 1901.
?
Trouessart, Catal. Mamm. viv. foss.,suppl , p. 93, 1904.
?
Elliot, Check List Mamm. North Amer., Field Columb.Mus., publ. 105, zool, ser., vol. 6, p. 477, 1905 (part).
?
Warren, Mamm.of Colorado, p. 275, 1910 (part).?Cary, North Amer. Fauna, No. 33, p.209, August 17, 1911 (part).
?
Miller, List North Amer. Land Mamm.1911, Bull. U. S. Nat. Mus., No. 79, p. 57, December 31, 1912; List NorthAmer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 71, April29, 1924.Myotls subulatus Warren, Mammals of Colorado, p. 275, 1910.
?
Cary, NorthAmer. Fauna, No. 33, p. 206, August 17, 1911 (not of Miller, 1897, andmost subsequent writers).Type locality.?On the Arkansas Eiver near present town of LaJunta, Otero County, southeastern Colorado.Type specimen.?All trace of the type specimen has been lost.Distribution.?Arid plains and eastern Rocky Mountain regionfrom Kansas and southeastern Colorado north to Montana. (Seemap 12, p. 165.)Diagnosis.?A pale subspecies, flaxen above and often nearly whitebelow.Color.?General color of upper parts ranging from " light buff "to " warm buff " with a slight tricolor effect when the hairs are partedso that their paler intermediate portions are seen in contrast withtheir blackish bases and flaxen tips. The top of the head and thebases of the outer part of the ears are almost whitish, but with abuffy tinge. Muzzle, chin, ears and tragus blackish, the sides ofthe face from muzzle to ears, blackish brown. The under surfaceis a very pale buff, becoming nearly white in some individuals : basesof the hairs blackish except at the extreme posterior region of thebody.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONTX 169Measurements.?For measurements see tables, pages 173 and 174.Specimens examined.?Total number 38, from the followinglocalities
:
COLORADO: Chimney Canyon, Avalo, Weld County, 1 skin, 1 ale.(U.S.N.M.) ; Colorado Springs, 1 mounted (Colorado College).KANSAS : Banner, Trego County, 6 ale, type and paratypes of cUiolabrum(U.S.N.M.) ; near Castle Rock, Trego County, 2 skins (K. U.).MONTANA: Big Timber, 1 skin (U.S.N.M.) ; Miles City, 1 skin (U.S.N.M.).NEBRASKA: Chadron, 1 ale. (U.S.N.M.).SOUTH DAKOTA: Custer, 3 skins (U.S.N.M.), 1 ale. (A. M. N. H.) ;Hermosa, 1 ale. (U.S.N.M.) ; Pine Ridge, 6 skins (A. M. N. H.).WYOMING: Bitter Creek, 2 sldns (A. M. N. H.) ; Bull Lake, ShoshoneIndian Reservation, 1 ale (U.S.N.M.) ; Cody, 1 skin (U.S.N.M.) ; FortBridger, 1 skin, nearly typical (U. C.) ; GreybuU, 4 skins (U.S.N.M.) ;Kinney Ranch, 1 ale. (A. M. N. H.) ; Otto, 2 skins (A. M. N. H.) ; Rattle-snake Mountains, 1 skin (U.S.N.M.).Remxwhs.?While the series from various localities in the dry in-terior from South Dakota and Montana to southeastern Colorado isdecidedl}^ paler and whiter-bellied than that from New Mexico,Arizona, California, and eastern Oregon, the difference is not alwaysvery marked, and the typical race merges into the next by almostimperceptible degrees. This is the only known form of the speciesm which the underparts are normally so pallid as actually toapproach white.MYOTIS SUBULATUS MELANGRHINUS (Merriam)Yespertilio ciliolabrum Meeiriam, Proc. Biol. Soc. Washington, voL 4, p. 4,December 17, 1886 (part; sx)ecimens from Grant County, N. Mex.).Myotis californicus Giliolabrum Caby, North Amer. Fauna, No. 33, p. 209,August 17, 1911 (part; specimen from Snake River, Routt County, Colo.).Vespertilio melanorhinus Merbiam, North Amer. Fauna, No. 3, p. 46, Sep-tember 11, 1890.?Miller, North Amer. Fauna. No. 13, p. 69, October 16, 1897(as synonym of californicus)
.
?Mlller and Rehn, Proc. Boston Soc. Nat.Hist., vol. 30, p. 308, December 27, 1901.?Lton and Osgood, Catal. Type-Sp.Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 271, January 28,1909.Vespertilio albescens melanorhinus H. Allen, Monogr. Bats North Amer., Bull.U. S. Nat. Mus., No. 43 (1893), p. 91, March 14, 1894.?Tbotjessaet., Catal.Mamm. viv. foss., p. 132, 1897.Vespertilio nitidus henshatvii H. Allen, Monogr. Bats North Amer., Bull. U. S.Nat. Mus., No. 43 (1893), p. 103, March 14, 1894 (Wingate, N. Mex.).?Miller, North Amer. Fauna, No. 13, p. 69. October 16, 1897 (as synonym ofcalifornicus).?Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p.308, December 27, 1901.Myotis orinomiis Elliot, Field Columbian Mus., publ. 79, zool. ser., vol. 3, p.228, June, 1903 (La Grulla, San Pedro Martir Mountains, Lower California,Mexico, altitude, 8000 feet) ; Land and Sea Mamm. Middle Amer., FieldColumb. Mus., publ. 95, zool. ser., vol. 4, p. 577, 1904 ; Check List Mamm.North Amer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 476, 1905;Catal. Mamm. Field Columb. Mus., Field Columb. Mus., publ. 115, zool. ser.,58518?28 12
170 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
vol. 8, p. 502, 1907.
?
Gbinnell and Swaeth, Univ. California Publ. Zool., vol.10, p. 138, April 13, 1912.
?
Miller, List North Amer. Land Mamm. 1911,Bull. U. S. Nat. Mus., No. 79, p. 57, December 31, 1912.?J. Geinneill, Proc.California Acad. Sci., ser. 4, vol. 3, p. 278, August 28, 1913.?H. W. Gbinnell,Univ. California Publ. Zool., vol. 17, p. 290, January 31, 1918.?J. GBiNNiax,Univ. California Publ. Zool., vol. 21, p. 314, January 27, 1913.
?
Miller, ListNorth Amer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 71,April 29, 1924.Myotia lucifugus longicrus J. Gkinnell, Univ. California Publ. Zool., vol. 5,p. 158, October 31, 1908 (part).Type locality.?Little Spring, north base of San Francisco Moun-tain, Coconino County, Ariz. ; altitude 8,250 feet.Type specimen.?Adult male. No. 18684, United States NationalMuseum (Biological Survey collection), collected at San FranciscoMountain, Arizona, August 4, 1889, by Dr. C. Hart Merriam andVernon Bailey. The specimen after 20 years' immersion in alcoholwas made into a skin.Distribution.?From southern Colorado, southwestward across NewMexico, Arizona, and northern Mexico to the Pacific coast of southernCalifornia and northern Lower California, and northwestward intoeastern Washington and Oregon. (See map 12, p. 165.)The distribution in California and the northwest has not yetbeen wholly worked out. Specimens have been taken in the south-western part of California, as well as in Nevada, eastern Oregon,and southeastern Washington.Diagnosis.?Color above richer and less pallid than in the typicalsubspecies; underparts apparently always strongly tinged with buffand rarely, if ever, approaching white.Measurements.?For measurements see tables, pages 173 and 174.Specimens etxamdned.?Total number 119, from the followinglocalities
:
ARIZONA: Fort Whipple, 2 skins (U.S.N.M.) ; Huachuca Mountains, 1 skin(F. M.) ; Prescott, 1 skin (A. M. N. H.) ; San Francisco Mountain, 1skin, type (U.S.N.M.) ; Santa Catalina Mountains, 1 ale. (A. M. N. H.) ;White Mountains, 1 skin (A. M. N. H.).CALIFORNIA: 8 miles west of Bakersfield, Kern County, 5 skins (U.C.) ; Carroll Creek, Inyo County, 1 skin (U. C.) ; Dulzura, San DiegoCounty, 1 skin (A. M. N. H.), 3 ale, 1 skull (U.S.N.M.), 8 ale. (A. N.S. P.), 4 skins (U. C.) ; Jacumba, San Diego County, 1 skin (A. M.N. H.) ; Kern River, Tulare County, 1 ale. (U.S.N.M.) ; Owen Valley,Inyo County, 1 ale. (U.S.N.M.) ; San Bernardino Mountains, 1 skin (U. C),1 .skin (U.S.N.M.) ; San Diego County, no exact locality, 1 skull(U.S.N.M.) ; San Jacinto Mountains, Riverside County, 1 skin (U. C), 3ale. (U.S.N.M.) ; Santa Rosa Mountains, Riverside County, 4 skins (U. C.) ;Santa Ysabel, Orange County, 12 ale. (U.S.N.M.) ; Twin Oaks, San DiegoCounty, 1 ale. (U.S.N.M.) ; Walker Pass, Kern County, 1 skin (U. C.) ;Warm Spring, near Vallecito. San Diego County, 1 skin (U. C.) ; WhiteMountains, Inyo County, 1 skin (U. C.) ; Witch Creek, San Diego County,1 ale. (U.S.N.M.) ; Yosemite Valley, 1 skin (U.S.N.M.).
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 171CHIHUAHUA: San Luis Mountains, 1 skin (U.S.N.M.).COLORADO: Antonite, 1 skin (U.S.N.M.) ; Snake River, Routt County,1 skin, 1 ale. (U.S.N.M.).LOWER CALIFORNIA: La Grulla, 3 skins, including type of orinomus(F. M.), 5 skins (U.S.N.M.) ; Hanson Laguna, 1 skin (U.S.N.M.) ; SantaEulalia, 1 skin (U.S.N.M.) ; Vallecitos, 1 skull (F. M.).NEVADA: Cottonwood Range, Humboldt County, 2 ale. (U.S.N.M.) ; LittleHigh Rock Canyon, Washoe County, 2 skins (U. C.) ; Panaca, 1 ale.(U.S.N.M.) ; Rabbit Hole Mountains, 1 skin (U.S.N.M.) ; Pablo, 1 ale.(U.S.N.M.).NEW MEXICO: Cantonment Burgwin, 1 skin (U.S.N.M.) ; Fort Wingate,2 ale. (U.S.N.M.) ; Guadalupe Canyon, 1 skin (U.S.N.M.) ; Las Vegas,Hot Springs, 1 ale. (U.S.N.M.) ; Mount Capitan, 1 skin (U.S.N.M.) ; Pecos,1 skin (U.S.N.M.) ; Santa Rosa, 2 skins (U.S.N.M.) ; Silver City, 1 skin(U.S.N.M.) ; Tres Piedras, 1 skin (U.S.N.M.) ; Wingate, 1 skull (U.S.N.M.) ;Zuni Mountains, 1 skin (U.S.N.M.).OREGON: Barnes, Crook County, 1 skin (U. C.) ; Homestead, 1 skin, ap-proaching ciliolabrum (U.S.N.M,) ; McDermitt, Malheur County, 2 skins(U.S.N.M.) ; Millers, mouth of Deschutes River, 4 skins (U.S.N.M.) ;Riverside, 3 skins (U.S.N.M.) ; Rockville, Malheur County, 1 skin(U.S.N.M.) ; Rome, Malheur County, 1 skin (U.S.N.M.) ; Sheaville, 1 skin(U.S.N.M.) ; Skull Spring, 1 skin (U.S.N.M.) ; Twelve Mile Creek, 2 ale.(U.S.N.M.) ; Warner Valley, 1 skin (U.S.N.M.).TEXAS: Terlingua Creek, 1 skull (U.S.N.M.).WASHINGTON: Bly, 1 skin (U.S.N.M.) ; Goldendale, 2 ale. (U.S.N.M.).Remarks.?Owing to paucity of specimens and the difficulties ofcomparing alcoholic material with skins, previous writers have notrecognized Merriam's Vespertilio melanorhinus. With the betterseries now at hand, however, it becomes clear that this animal is reallya form of Myotic subulatus, and not identical with M. californicus ashad at first seemed to be the case. The name Vespertilio nitidushenshawii of Harrison Allen, based on two females taken by Mr.H. W. Henshaw near Wingate, N, Mex,, probably refers to the sameanimal. These specimens appear to be no longer in the NationalMuseum collection, but two taken at Fort Wingate by Dr. R. W.Shufeldt are referable to inelanorhhvm. Elliot's Myotis orinomiisfrom northern Lower California is also a synonym of M. suhulatttsmelanorhinus. The species has been more often collected in the por-tion of its range occupied by this race than it has been in the easternUnited States.MYOTIS SUBULATUS LEIBII (Aadnbon and Bachman)Vespertilio leibii Audubon and Bachman, Joiirn. Acad. Nat. Sci. Philadelphia,ser. 1, vol. 8, p. 284, 1842.?H. Allen, Monogr. Bats North Amer., Smithson-ian Misc. Coll., No. 165, p. 80, June, 1864 ; Monogr. Bats North Amer.,Bull. U. S. Nat. Mus., No. 43 (1893), p. 190, March 14, 1894.?Miller, NorthAmer. Fauna, No. 13, p. 29, October 16, 1897.Myotis winnemana Nelson, Proc. Biol. Soc. Washington, vol. 26, p. 183, August8, 1913 (Plummer Island, Md.).
?
Elliot, Check-List Mamm. North Amer.,suppl., p. 156, 1917.?MnxEE, List North Amer. Recent Mamm. 1923, Bull.U. S. Nat. Mus., No. 128, p. 71, April 29, 1924.
172 BULLETIN 144, UNITED STATES NATIONAL. MUSEUMType locality.?Erie County, Ohio.Type specimen.?Collected in Erie County, Ohio (formerly Michi-gan), by Dr. George C. Leib, and sent to Audubon and Bachman atPhiladelphia, but not now known to be in existence.Distribution.?From Vermont, New York, and Ohio south to WestVirginia and Kentucky; exact limits of range not yet ascertained.(See map 12, p. 165.)The few specimens at hand probably indicate fairly well the gen-eral range, though the southern limits will doubtless be found toextend to Florida. Intergradation with the pale race of the Plainscountry probably takes place west of the Mississippi.Diagnosis.?General color much darker than in the typical sub-species and closely resembling that of Myotis Jucifugus lucifugusin the olive phase, but slightly more golden above and lacking thedark spot at the shoulder.Color.?Upper surface of body, from forehead to base of tail,nearly " ochraceous tawny " with a golden sheen in some lights dueto the shining tips of the long hairs ; on the sides of the neck the tintis a little brighter. Lower surface " warm buff," the tips of the hairsslightly burnished. Bases of the hairs on the body everywhereblackish, except at the extreme sides. The face from the nose to baseof ears and including the lower lip, black, giving a masked appear-ance, which is heightened by the dull black ears, tragus, nose andchin. Wings and membranes blackish brown.Skull.?Although nearly as long as that of Myotis lucifugus.^ theskull is so flattened that it has a notably less depth ; the brain case isdistinctly narrower.Measurements.?For measurements see tables, pages 173 and 174.SpeciTiiens exartiined.?Total number 8, from the followinglocalities
:
KENTUCKY: Hickman's Cave, 1 ale. (M. C. Z.).MARYLAND: Plummer Island, 2 skins (U.S.N.M.).NEW YORK: Sing Sing, 2 ale. (U.S.N.M.).WEST VIRGINIA: White Sulphur Springs, 1 skin (M. C. Z.).VERMONT: Brandon. 1 skin (U.S.N.M.) ; Proctor, 1 skin (U.S.N.M.).Remarks.?This smallest of the bats known to occur in the easternUnited States seems to be much less common than Myotis lucifuguslucifugus, and the few specimens that have been taken have usuallybeen confused with the better known animal. Its black ears and facialmask, rather golden tint, keeled calcar and shorter forearm (31 to 34-mm. instead of 36 to 40 mm.) will at once distinguish it, however.Though described by Audubon and Bachman eighty years ago, itwas not again recognized until Nelson redescribed it in 1913 asMyotis winnennana.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 173External measurements of Myotis aubulatus
Locality
174 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis subulatus
Locality
AMERICAN" BATS OF THE GENERA MYOTIS AND PIZONTX 175
n. THE SPECIES OF MYOTIS OCCTTRRING IN TROPICAL AND SOUTHAMERICAMYOTIS NIGRICANS (Wied)(Synonymy under subspecies)Distribution.?From southern Mexico (Chiapas and Yucatan)southward over the warmer parts of South America (including thecoastal islands) to Paraguay and southeastern Brazil; islands ofGrenada and Dominica in the Lesser Antilles.Diagnosis?A small bat not very unlike the North AmericanMyotis californicus in size and general characters, but tail less elon-gated (ratio of tail to head and body in series of specimens averagingabout 85), foot relatively larger, the average ratio of its length tothat of tibia usually more than 45 and in some races as great as 53
;
ear, when laid forward, extending about to end of muzzle, usuallynot reaching to nostril but occasionally going beyond it ; metacarpals3 to 5 very slightly graduated, the distal end of the fifth, when foldedback, not reaching the elbow by about 2 mm. ; texture of pelagevarying from rather short and wooly to long and silky, the longerhairs, when well developed, with glossy tips, these, however, rarelyif ever producing a noticeable sheen; color usually dark brown orblackish (rarely grayish or ochraceous), the bases of the hairs notstrongly contrasted with the tips; a tawny dichromatic phase rarelypresent; skull and teeth with no marked peculiarities; forearm 31.6to 39 mm. usually less than 37 mm. ; greatest length of skull rangingfrom 12.6 to 14.7 mm., usually less than 14.5 mm.; maxillary toothrow 4.7 to 5.6 mm., usually less than 5.5 mm. ; mandibular tooth row5.0 to 6.0 mm., usually less than 5.6 mm.; crowns of upper molarssmall, m- usually varying from 1.15 to 1.30 by 1.55 to 1.70 mm.Ears.?The ear is narrow and delicate (usually 11 to 13 mm. inheight from meatus), its anterior border convex to the blunt tip, theposterior border slightly concave below the tip, then convex to theprojecting shoulder of the basal two-fifths. When laid forward thetip of the ear reaches about to the end of the muzzle or a little be-3^ond. Tragus about half the height of the ear. with a small basallobe, marked off by a shallow notch, above which is the broadest partof the tragus. The tip is slightly attenuate and shows a few minutecrenulations on the outer edge.Wing and membranes.?The wing membrane arises from the sideof the foot at the base of the outer toe. Metacarpals 3 to 5 slightlygraduated or sometimes the fourth and fifth are about equal and onlyminutely less than the third. Taking the third finger as 100, thefourth and fifth are respectively as 85 and 78 (59:50.5:46.5 mm.).When the wing is folded the end of the third metacarpal falls shortof the elbow by about 2 mm. Practically the entire tail is included
176 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
within the membrane, though occasionally a minute tip is free. Thefur of the body extends out on the uropatagium, both above andbelow, to or a little beyond a line joining the knees.Foot.?The foot is small and delicate in proportion with the gen-eral small size of the animal. In series of specimens from differentlocalities the average ratio of foot to tibia ranges from about 43 toabout 53. Calcar a little shorter than the free border of the inter-femoral membrane (about 14: 19 mm.), often though not constantlyprovided with a small keel, and usually ending in a minute projectinglobule.Fur and color.?The pelage is soft but not especially full and silkyin quality ; the hairs on the back about 5 mm. in length. The colorof the hairs is everywhere blackish plumbeous at base; tips brownwith a faint gloss and usually so dark as to present no conspicuouscontrast with the under color; entire ventral surface less darkenedand more tinged with yellowish than back. There is a slight tend-ency to dichromatism shown by the rare occurrence of individualswith orange-suffused pelage.Skull.?The skull is small and delicately formed. Profile of theforehead rising rather abruptly; brain case distinctly globular. Thetemporal ridges often form a distinct though low sagittal crest. Therostrum is relatively broad in proportion to its length, with the widthacross the molars slightly exceeding the length of the maxillarytooth row (front of canine to back of last molar).Teeth.?The two minute upper premolars stand usually full inthe tooth row, though rarely the second may be crowded inward fromthe line. The cingulum is distinct and the cusp of the posterior toothreaches about half way to the tip of the anterior. The upper molarsresemble those of the North American Myotis lucifugus in the fulldevelopment of the secondary cusps and ridges and in the breadthand distinctness of the cingulum. The transverse diameter of thecrown in m^ and w^ is less proportionately to the antero-posteriordiameter than in M. lucifugus.Remarks.?Myotis nigricans is a bat peculiar to the warmer partsof America. Its range probably meets that of Myotis californicus insouthern Mexico. Though allied to M. californicus and structurallynot very different from it, Myotis nigricans is undoubtedly a distinctspecies characterized by the almost uniform coloration of the hairsfrom base to tip, the shorter third metacarpal (which does not reachthe elbow as it usually does in M. californicus)., more robust footand shorter tail and ears. The skull has a slightly more elevatedocciput, and the width across the molars is increased so that this dis-tance tends to be perceptibly greater than the maxillary tooth rowinstead of about equal to it.
AMERICAISr BATS OF THE GENERA MYOTIS AND PIZONYX 177Of the technical names aj)plied to this animal before 1850, Ves-pertilio hrasiliensis of Sp:x, 1823, is clearly the oldest, althoughthe exact locality at which the type was obtained is not stated. Itwas preoccupied by Vespertilio hrasiliensis Desmarest, 1822, a namebased on a species of Epteslcus, hence Fischer replaced it by Ves-pertilio spixii in 1829. Meanwhile, however, Wied (1826) haddescribed the species as Y. nigricans. Although no specimens fromthe plains of the Mojos country, eastern Bolivia, have been seen,it seems clear, from the description of Vespertilio hypothrix D'Or-bigny and Gervais, that the form which occurs in this region is thesame as typical nigncan^. Whether or not Temminck's Vespertilioparvulus should be placed in the synonymy of Myotis nigricansis not altogether clear. The description gives little that is diag-nostic. The specimens on which it was based were collected byNatterer in Brazil. Dobson, however, who directly compared analcoholic specimen of nigricans with Temminck's type of parvulusin the Leiden Museum, considered the animals identical. Anotherpossible synonym is Vespertilio nitetis of Wagner, based on Natterer'sdrawing. Wagner's figure, however, shows a short tragus, unlikethat of any known Myotis. Recently the red phase of Myotis nigri-cans has been named as a small form of M. t^her (J. A. Allen, 1914).Several local forms of Myotis nigricans have been described duringthe past forty years. It is possible that some of them will eventuallyprove to be worthy of recognition ; but on the basis of the insufficientmaterial now at hand we have found it impossible to frame diagnosesor to map approximate areas of distribution. We have thereforeplaced the names hondcB^ caucensis, chiriqu?nsis, maripensis, andpunensis in synonymy, where they may safely remain until some oneprovided with adequate series of skins shall be able to show thatthe forms to which they were applied have an actual independentstatus. At present we are forced to regard all the material of thekind hitherto referred to the nigricans group from South Americaand from Central America north to Guatemala as pertaining to thetypical race, an animal which varies noticeably in color withinrather narrow limits, but which is constantly large in size for thespecies. Along the extreme northern border of the range of M.nigHcans three small local forms have been developed, one insouthern Mexico, and one each on the islands of Curasao andDominica. MYOTIS NIGRICANS NIGRICANS (Wied)Vespertilio brasiliensis Spix. Sim. et Vespert. Brasil. Sp. Nov., p. 63, pi. 36,fig. 8, 1823 (not of Desmarest, 1822).Vespertilio nigricmis Wied, Beitriige z. Naturg. Brasil, vol. 2, p. 266, 1826.
?
Fischer, Synopsis Mamm., p. 112, 1829.
?
Temminck, Monogr. de Mamm.,vol. 2, p. 242, 1840.
?
Schinz, Syst. Verzeich. Saugeth. Oder Synopsis Mamm.,
178 BULLETIN 144, UNITED STATES NATIONAL MUSEUM
vol. 1, p. 185, 1844.
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Wagner, Schreber's Saugth., Suppl., vol. 1, p. 533,1844.?DoBSON, Catal. Chiroptera Brit. Mus., p. 319, pi. 19, fig. 9, 1876.?WiNQE, E Mus. Lundli, Zool. Mus. Univ. Copenhagen, vol. 2, art. 1, p. 13,pi. 2, fig. 3, 1892.
?
Allen and Chapman, Bull. Amer. Mus. Nat. Hist., vol.5, p. 231, September 21, 1893.?Tkouessabt, Catal. Mamm., viv. foss., p. 130,1897.?PiBA, Zool. Anzeiger, vol. 28, p. 18, 1904.Vespertilio spixii Fischeb, Synopsis Mamm., p. Ill, 1829 (new name for F.In-asilierms Spix).Vespertilio parvtilus Temminck, Monogr. de Mamm., vol. 2, p. 246, 1840(Brazil).?VoN Pelzeln, Verb. K. K. Zool.-Bot. Ges. Wien, vol. 33, Beiheft,p. 45, 1833.Nyctophylax parvultis Fitzinger, Sitzungsber. Akad. Wiss. Wien, vol. 62, pt. 1,p. 563, 1870.Vespertilio hypothrix D'Orbigny and Gebvais, Voy. dans I'Amer. Merid., vol. 4,pt. 2, mammiferes, pp. 14 (footnote), 16, 1847 (eastern Bolivia).?Gesivais,Exped. Amer. du Sud du Castelnau, zool., mammif&res, p. 83, pi. 15, fig. 5(teeth), 1855.Vespertilio hypothryx D'Orbigny and Gebvais. Voy. dans I'Am^r. Merid., vol.4, pt. 2, mammiferes, p. 16, 1847.Vespertilio concinnus H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 280(San Salvador).
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'Miller, Proc. Biol. Soc. Washington, vol. 13, p. 154, June13, 1900 (part).?J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 384.July 9, 1914.Vespertilio exigiius H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 281(Aspinwall= Colon, Panama).
?
Lyon and Osgood, List Type-Sp. Mamm.U. S. Nat. Mus., Bull. U. S. Nat. Mus., No. 62, p. 290, January 29, 1909.Aeorestes nigricans Fitzingeb, Sitzungsber. Akad. Wiss. Wien. vol. 62, pt. 1,p. 432, 1870.Vesperugo nigricans von Pexzeln, Verb. K. K. Zool.-Bot. Ges. Wien, vol, 33,Beiheft, p. 45, 1883.Vespertilio cincinnus H. Allen, Monogr, Bats North Amer., Bull. U. S. Nat. Mus.,No. 43 (1893), p. 97, footnote, March 14, 1894.Myotis nigricans Miller, North Amer. Fauna, No. 13, p. 74, October 16, 1897(part, synonymy).
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Thomas, Ann. and Mag. Nat. Plist., ser. 7, vol. 8, p. 436,November, 1901. Miller and Rbhn, Proc. Boston Soc. Nat. Hist., vol. 30,p. 257, December 27, 1901 (part).
?
Elliot, Land and Sea Mamm. MiddleAmer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 575, 1904 (part).
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Tbouessabt, Catal. Mamm. viv. foss., suppl., p. 93, 1904 (part).?J. A. Allen,Bull. Amer. Mus. Nat. Hist., vol. 20, p. 77, February 29, 1904 ; p. 343, October8, 1904.
?
Elliot, Check List Mamm. North Amer., Field Columb. Mus., publ.105, zool. ser., vol. 6, p. 475, 1905 (part) ; Catal. Mamm. Field Columb. Mus.,Field Columb. Mus., publ. 115, zool. ser., vol. 8, p. 501, 1907 (part).
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Osgood,Field Mus. Nat. Hist., publ. 176, zool. ser., vol. 10, p. 181, April 20, 1914.?Goldman, Smithsonian Misc. Coll., vol. 69, No. 5, p. 213, April 24, 1920.Myotis chiriquensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 20, p. 77, Feb-ruary 29, 1904 (Boqueron, Chiriqui, Panama).
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Elliot, Land and SeaMamm. Middle Amer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 576,1904 ; Check List Mamm. North Amer., Field Columb. Mus., publ. 105, zool.ser., vol. 6, p. 475, 1905 (part).
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Miller, List North Amer. Land Mamm.1911, Bull. U. S. Nat. Mus., No. 79, p. 58, December 31, 1912 (part).?Gold-man, Smithsonian Misc. Coll., vol. 69, No. 5, p. 213, April 24, 1920.?Anthony, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 373, June 9, 1916.Myotis nigricans concinnus Osgood, Field Mus. Nat. Hist., publ. 155, zool. ser.,vol. 10, p. 65, January 10, 1912.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 179Myotis nigricans mundus Osgood, Field Mus. Nat. Hist., publ. 176, zool. ser.,vol. 10, p. 182, April 20, 1914 (not of H. Allen=ilf. alhescens).Myotis simus Osciood, Field Mus. Nat. Hist., publ. 176, zool. ser., vol. 10, p. 182,April 20, 1914 (not of Thomas).Myotis ruber Tceaysi J. A. Aixen, Bull. Amer. Mus. Nat. Hist, vol, 33, p. 383,July 9, 1914 (Inca Mines, Peru, Lat. 13? 30' S., Long. 70 W., altitude,6,000 feet). Red phase.Myotis punensis J. A. Axlept, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 383, July9, 1914 (Puna Id., Ecuador).Myotic tondce J. A. Axlen", Bull. Amer. Mus. Nat. Hist., vol. 33, p. 384, July9, 1914 (Bonda, Santa Marta, Colombia).Myotis maripensi^ J. A. Axlen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 385,July 9, 1914 (Maripa, Rio Caura, Venezuela).Myotis esmeraldce J. A. Aixen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 385,July 9, 1914 (Esmeraldas, Ecuador).Myotis caucensis J. A. Axlen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 386,October 30, 1914 (Rio Frio, Colombia).Myotis nigricans nigricavs Milleb, List North Amer. Recent Mamm. 1923, Bull.U. S. Nat. Mus., No. 128, p. 72, 1924 (part).Type locality.?Fazenda de Aga, near the Rio Iritiba, province ofEspirito Santo, eastern Brazil,Type specimen.?The type was collected, about 1823, by PrinceMaximilian zu Wied, Its present whereabouts, if it is still inexistence, is not known.Distribution.?Central and South America from Guatemala tosouthern Brazil and from coast to coast.Diagnosis.?Size maximum for the species; greatest length ofskull usually more than 13.5 mm. and not infrequently more than14 mm. (known maximum 14.7) ; foot usually less than half as longas tibia (average ratio of foot to tibia in series of ten examples fromsingle localities ranging from 44.6 to 49.1) ; ear usually not ex-tending beyond nostril when laid forward.Color.?The general color above, when the fur is smoothed sothat the blackish-slate bases of the hairs do not appear on the surface,is a brown near the Mars brown of Ridgway, usually darker andmore blackish but occasionally lighter and more nearly approachingProut's brown or russet. Sometimes there is a decided tinge of grayor ocher, especially on the neck and head. Underparts not so darkas back, and often with a buffy or yellowish cast. In the red phasethe color is suffused with orange, producing a brown very near tothe Sudan brown of Ridgway.Measurements.?For measurements see tables, pages 184 and 187.Specimens exa/tnined.?Total number 648, from the following-localities :BOLIVIA: Balzan, 4 ale. (B. M.) ; Buenavista, 3 skins (A. M. N. H.) ;Caiza, 4 ale. (B. M.) ; Rosario, 4 ale. (U.S.N.M.) ; Rio Surutu, Dept.Santa Cruz, 5 skins (A. M. N. H.) ; Rurreuebaque, Beni. 1 ale. (U.S.N.M.) ;Sania Cruz Province, 8 ale. (B. M.) ; no exact locality, 2 skins (B. M.).
180 BULLETIN 144, UNITED STATES NATIONAL MUSEUMBRAZIL: Bahia, 4 skins, (B. M.) ; Chaco, 1 ale. (B. M.) ; Benevides, 1 ale,(B. M.) ; Lagoa Santa, 6 ale. (M. C. Z.) ; Linhares, 1 ale. (M. C. Z.) ;Macao, 1 ale. (M. C. Z.) ; Minas Geraes, 1 ale. (M. C. Z.) ; Minas Geraes,Rio Jordao, near Araguary, 1 skin (B. M.) ; near Para, 1 skin, 1 ale.(B. M.) ; Parana Prov., Palmeiras. 2 ale. (B. M.) ; Pernambuco. 1 skin,1 ale. (B. M.) ; Rio Grande do Sul, San Lorenzo, 3 ale. (B. M.) ; RioJaneiro, 1 skin, 2 ale. (B. M.) ; Samarao (Bahia), 6 skins (B. M. ), 1 skin(M. C. Z.) ; Santa Catherina, 1 skin (B. M.) ; Sao Paulo, 2 ale. (M. C. Z.) ;Sao Paulo, Cruzeiro, 1 skin, 1 ale. (B. M.) ; Sao Paulo, Sao Sebastian,44 skins (U.S.N.M.). 9 skins (B. M.), 2 skins (F. M.) ; Serra do Itatiaya,1 skin (A. M. N. H.) ; interior, 2 ale. (U.S.N.M.) ; no exaet loeality,2 skins (B. M.).BRITISH GUIANA: Moon Mountains, 14 skins (B. M.) ; UpokaritKanuku Mountains, 6 skins, 3 ale. (B. M.).BRITISH HONDURAS: Stann Creek, 1 ale. (U.S.N.M.).COLOMBIA: Bogota. 1 ale. (B. M. ) ; Bonda, 25 skins, ineludiug type ofhondcB (A. M. N. H.) : Buenavista, 3 skins (A. M. N. H.) ; Cauca River,1 skin (U.S.N.M.); Choco region, 1 skin, (B. M.), 1 ale. (M. C. Z.) ;Jimenez, 1 skin (U.S.N.M.), 3 skins (B. M.) ; Juntas de Tamana, Cauea,3 skins (A. M. N. H.) ; Palomino, 1 skin (M. C. Z.) ; Rio Frio, CaueaRiver, 1 skin, tj'pe of caucensis (A. M. N. H. ) ; Santa Marta, 3 skins,6 ale. (M. C. Z.), 1 skin (B. M.) ; ? Yaldivia, 8 ale. (B. M.).COSTA RICA: Cartago, 2,000 meters, 1 skin (B. M.) ; Rio Frio, 4 ale.(U.S.N.M.) ; Sipurio, 69 ale. (U.S.N.M.) ; Zent, 2 ale. (U.S.N.M.), 1 ale.(M. C. Z.) ; no exact locality, 1 skin (A. M. N. H.).ECUADOR: Ambiyaeu River, 1 ale. (A. N. S. P.); Bahia de Caraquez,Rio Briseho. 2 skins (A. M. N. H.) ; Bucay, 5 skins (A. M. N. H.) ;Caehabi, 1 skin, 5 ale. (B. M.) ; Canelos, 6 skins (A. M. N. H.) ; Casanga,Prov. del Oro, 1 skin (A. M. N. H.) ; Cuaque, El Destine, 2 skins(A. M. N. H.) : Curaray River (Mouth of), 5 skins (A. M. N. H.) ;Daule, 2 skins (A. M. N. H.) ; Esmeraldas, 3 skins, including type ofesmcraldw (A. M. N. H.) ; Gualaquiza, 4 skins (B. M.) ; Manavi, 1 skin(A. M. N. H.) : Mindo, 9 skins (B. M.) : Naranjo, 1 skin (A. M. N. H.) ;Pambilar, 4 skins (U.S.N.M.) ; Paramba, 2 ale. (U.S.N.M.) ; Pto. In-diana, R. Amazonas, 1 skin (A. M. N. H.) ; Puente de Chimbo, 4 skins(A. M. N. H.) ; Puna Island, 2 skins (B. M.), 2 skins, including typeof punensis (A. M. N. H.) ; San Emilio, 1 skin (B. M.) ; San Javier,4 skins (U.S.N.M.) ; San Juan, 15 miles west of Juigra, 1 skin (B. M.) ;San Ramon, 1 skin (A. M. N. H.) ; Santiago River, 2 ale. (U.S.N.M.) ;Sarayaeu, 6 skins (A. M. N. H.) : Suno River, Oriente Province, 11 skins(A. M. N. H.) : Ventura, 14 skins (A. M. N. H.) : Zamora, 1 skin(A. M. N. H.) ; Zaruma, 1 skin (B. M.) ; Zaruma, Loja Trail, 1 skin(A. M. N. H.) ; no exact locality, 1 ale. (B. M.).GRENADA, B. W. L: 1 ale. (M. C. Z.).GUATEMALA: Carolina, 4 skins (A. M. N. H.) ; El Cipres, 1 skin(A. M. N. H.). Both of these localities are near the Volcan Zuuil.PANAMA: Boca de Cupe, 1 skin (U.S.N.M.) : Bogaba, 1 ale. (M. C. Z.) ;Bohio, 1 skin. 3 ale. (U.S.N.M.) ; Boqueron, 2 skins (A. M. N. H.) ; Cana,1 ale. (U.S.N.M.) ; Chepigana, 1 skin (A. M. N. H.) ; Chiriqui, 3 skins,including type of chiriquensis (A. M. N. H.), 1 ale. (U.S.N.M.) ; Cituro,1 skin (A. M. N. H.) ; El Real, 3 skins (A. M. N. H.) ; Culebra, 2 ale.(U.S.N.M.); Gatun, 1 skin, 1 ale. (U.S.N.M.), 1 skin (A. M. N. H.) ;La Chorrera, 1 skin (A. M. N. H.) ; Obispo, 1 ale. (M. C. Z.) ; San
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONTX 181
Pablo, 1 skin (U.S.N.M.), 1 ale. (M. C. Z.) ; Tabernilla, 1 skin, 3 ale.(U.S.N.M.) ; Taboga Id., 1 ale. (U.S.N.M.) ; Tacarcuna, 2 skins (A. M.N. H.) ; Tapalisa, 3 skins (A. M. N. H.).PARAGUAY: Asuneiou, 3 skins (U.S.N.M.) ; Paraguari, 7 skins (U.S.N.M.),4 skins (B. M.), 1 skin (U. C.) ; Puerto Pinasco, 1 skin (U.S.N.M.);Sapucay, 6 skins, 14 ale. (U.S.N.M.), 6 skins, 2 ale. (B. M.) ; Villa Rica,3 skins, 9 ale. (U.S.N.M.), 4 skins (B. M.).PERU: Bellavista, 3 ale. (M. C Z.) ; Cayana, 1 ale. (B. M.) ; Chanchamaya.4 skins (B. M.), 1 skin (M. C. Z.) ; Cuzo, 4 skins (B. M.) ; HaciendaLimon, near Balsas, Maranon River, 1 skin (F. M.) ; Inea Mines, 4skins, including tyi>e of keaysi (A. M. N. H.) ; Junin, 2 ale. (B. M.) ;Masisea, 1 skin (B. M.) ; Moyobamba, 3 skins, 2 ale. (F. M.) ; Perene,2 skins (B. M.) ; Rio Ucayali, Masisea, 8 skins (B. M.) ; San Ignacio.16 skins (A. M. N. H.) ; Santa Ana, 2 ale. (B. M.), 7 ale. (U.S.N.M.) ;Yurimaguas, 1 ale. (F. M.).SAN SALVADOR: 2 ale., cotypes of concinnus (A. N. S. P.).TOBAGO, B. W. I.: Court House, 1 skin (B. M.) ; Waterloo, 1 skin(B. M.), 1 ale. (A. N. S. P.).TRINIDAD: Botanic Gardens, 2 ale. (B. M.).VENEZUELA: Caicara. 1 skin (B. M.) ; Carabobo, 1 skin (B. M.) ;Caracas, 1 ale. (B. M.) ; Maipures, Rio Orinoco, 7 skins (B. M.) ; Maripa,37 skins, including type of niaripensis (A. M. N. H.) ; Merida, Montesdel Escorial, 2500 meters, 1 skin, imm. (M. O. Z.) ; Merida, Montes de laSierra, 1500-2000 meters, 3 skins (B. M.) ; Puerto Cabello. 2 ale, (B. M.) ;Rio Aurare, 1 skin (F. M.) ; San Esteban, 10 skins (B. M.), 10 ale.(U.S.N.M.) ; Taehira, 1 skin (F. M.) ; Valencia, 1 skin (A. M. N. H.)
;
no exact locality, 1 ale. (B. M.).Remarks.?It is with some hesitation that we have placed in thesynonymy of Myotis nigricans nif/ricans the five names published byJ. A. Allen on July 9, 1914. That all were based on representativesof this species there appears to be no doubt; but it is quite possiblethat some of them may eventually be shown to be applicable to localraces. Such races, however, we have been unable to define on thebasis of the material which we have seen,MYOTIS NIGRICANS EXTREMUS. new subspeciesMyotis nigricans Miller, North Amer. Fauna, No. 13, p. 74, October 16, 1897(part, specimens).
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Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol.30, p. 257, December 27, 1901 (part).
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Elliot, Land and Sea Mamm. MiddleAmer., Field Columb. Mus., publ. 95, zool. ser., vol. 4, p. 476, 1904 (part).
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Teouessaet, Catal. Mamm. viv. foss., suppl., p. 93, 1904 (part).
?
Elliot,Cheek List Mamm. North Amer.. Field Columb. Mus., publ. 105, zool. ser.,vol. 6, p. 475. 1905 (part).Vespertilio concinnus Miller, Proc. Biol. Soc. Washington, vol. 13, p. 154, June13, 1900 (part; not Vespertilio concinnus H. Allen).Myotis nigricans concinnus Miller, List North Amer. Land Mamm. 1911, BullU. S. Nat. Mus., No. 79, p. 58, December 31, 1912 (part) ; List North Amer.Recent Mamm, 1923, Bull. U. S. Nat. Mus., No. 128, p. 72, April 29, 1924(part).Type locality.?Huehuetan, Chiapas, Mexico. Altitude, 300 feet.
182 BULLETIN 144, UNITED STATES NATIONAL MUSEUMTy'pe specvmen.?Adult female (skin and skull), No. 77670 UnitedStates National Museum (Biological Survey collection). Collectedat Huehuefean, Chiapas, Mexico, March 1, 1896, by E. W. Nelson andE. A. Goldman. Original number 9455.Distribution.?Tropical zone in southern Mexico (Chiapas,Campeche and Yucatan).Diagnosis.?Size constantly below the maximum for the species;greatest length of skull usually less than 13.5 mm. and never attain-ing as much as 14 mm. (known maximum 13.4 mm.) ; foot usuallymore than half as long as tibia (average ratio of foot to tibia in 10examples about 53.5).Measurements.?For measurements see tables, pages 185 and 188.Sfecvtnens examined.?Forty-one, from the following localities
:
CAMPECHE: La Tuxpana, 1 skin (U.S.N.M.).CHIAPAS: Huehuetan, 10 skins, 26 ale. (U.S.N.M.).YUCATAN: Izamal, 2 skins (U.S.N.M.), 1 skin (A. M. N. H.) ; no exactlocality, 1 skin (B. M.).Remarks.?Superficially this small race of Myotis nigricans re-sembles the Mexican form of M. califomicus in dark immature pelage.Specimens may be recognized, however, by the slight extent of thedark under color on the back, the relatively large foot (more thanhalf as long as tibia), and the small teeth.MYOTIS NIGRICANS NESOPOLUS MUlerMyotis nesopolus Miller, Proc. Biol. Soc. Washington, vol. 13, p. 123, April 6,1900.?TBotJESSABT, Catal. Mamm. viv. foss., suppl., p. 93, 1904.
?
Lyon andOsgood, List Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus., No.62, p. 272, January 28, 1909.Myotis nespotus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 384, July 9,1914 (misprint).Type locality.?Near Willemstad, Curasao, Dutch West Indies.Type specimen.?Adult male in alcohol. No. 101849, United StatesNational Museum, collected near Willemstad, island of Curac:ao,Dutch West Indies, November 4, 1899, by L. B. Smith.Diagnosis.?Similar to Myotis nigricans extremus^ color appar-ently less dark and foot apparently shorter, the ratio of its length tothat of tibia about as in true M. nigHcans.Description.?Color above between raw umber and Front's brown,the bases of the hairs just perceptibly darker. Below, ochraceousbuff, the basal half of the hairs slaty black.Measurements.?For measurements see tables, pages 186 and 188.Specimens exannined.?Total number 2, as follows:CURACAO, D. W. I: Willemstad, 2 ale, including type (U.S.N.M.).Remarks.?Myotis nigricans nesopolus is so similar to the smallrace occurring in southern Mexico that we are unable to give wholly
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 183
satisfactory characters by which to distinguish it. We believe, how-ever, that good material from Curagao will eventually establish thedistinctness of the two forms, and that in the present state of ourknowledge the course which we have adopted in describing theMexican animal as new is less unreasonable than it would be toapply the name nesopolus to both of these small races of Myotislilgrica/ns, separated as they are geographically by Colombia, Pan-ama, and Central America, an area inhabited by the large typicalform. MYOTIS NIGRICANS I>OMINICENSIS MillerMyotis dominicensis Miller. Proc. Biol. Soc. Washington, vol. 15, p. 243, Decem-ber 16, 1902.
?
Miller and Rehn, Proc. Boston Soc. Nat. Hist., vol. 31, p.121, August 27, 1903.?Elliot, Land and Sea Mamm. Middle Amer., FieldColumb. Mus., publ. 95, zool. ser., vol. 4, p. 576, 1904.
?
Trouessart, Catal,Mamm. viv. foss., suppl., p. 95, 1904.
?
Elliot, Check List Mamm, NorthAmer., Field Columb. Mus., publ. 105, zool. ser., vol. 6, p. 475, 1905.
?
Lyonand Osgood, List Type-Sp. Mamm. U. S. Nat. Mus., Bull. U. S. Nat. Mus.,No. 62, p. 270, January 28, 1909.?Miller, List North Amer. Land Mamm.1911, Bull. U. S. Nat. Mus., No. 79, p. 58, December 31. 1912; List NorthAmer. Recent Mamm. 1923, Bull. U. S. Nat. Mus., No. 128, p. 72, April 29,1924.Type locality.?Dominica, British West Indies.Type specimen.?Adult male, in alcohol. No. 113564 United StatesNational Museum, collected on the island of Dominica, British WestIndies, July 20, 1901, by H. Selwyn Branch.Distribution.?So far as known, this bat is found only on the is-land of Dominica, Lesser Antilles. The form occurring on Grenada,to the southward, is apparently true Myotis nigricans. Up to thepresent time no representatives of the species have been recordedfrom the intermediate islands, nor Has any member of the genusMyotis been reported from any other part of the entire Antilleanchain.Diagnosis.?^Like Myotis nigricans extremus but with smallerskull.Description.?A topotype is cinnamon brown both above andbeneath, the membranes blackish brown. Another, probably in asubadult pelage, is fuscous above; chin and belly cinnamon brown,throat fuscous.Shall.?The skull attains the smallest size that is known in anyAmerican Myotis (pi. 1, p. 7, fig. 5). In form it is peculiar in theunusually reduced volume of the brain case.MeasureTiients.?For measurements see tables, pages 186 and 188.SpeciTfieTis examined.?Total number 36, from the following local-ities :DOMINICA, B. W. I: 1 ?kin (B. M.), 1 skin (M. C. Z.) ; 29 ale, includingtype (U.S.N.M.) ; Basin Well, 5 ale. (B. M.).
184 BULLETIN 144, UNITED STATES NATIONAL MUSEUMRemarks.?This is a fairly well-marked island race distinguishedby the small size of the skull and the globular form of the brain case.External measurements of Myotis nigricans
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 185External measurements of Myotis nigricans?Continued
Locality Number o o
Myotis nigricansnigricans?Con.Peru:JuninDoCayanaMoyobambaDoSanta AnaDoInca Mines..Bolivia:Balzan.. ...Do..DoDoRosarioDoDo.-DoSanta Cruz Prov-inceDoDoDoDoDoDoEcuador:CachabiDoDoDo.-.Santiago RiverParambaDoBrazil:PalmeirasDoSan LorenzoDoPara.Rio JaneiroDoInteriorParaguay:SapucayDoDo..Do..Do..Do..Do..Do..Do..Do..Do..Do..Myotis nigricansestremasMexico:Huehuetan, Chia-pasDoDoDoDoDo
0.7.7.63 B.M.0. 7. 7. 64B.M.19955 F. M.19956 F. M.97. 10. 3. 20 B.M.97.10.3.21. B.M.? 15814 A. M. N. H,0.8. 3. 2 B.M.0. 8. 3.
3
0. 8. 3. 40. 8. 3. 5238682 U.S.N.M.23868323868423868521 B. M.28293335252797.11.7.63 B. M.97.11.7.6697.11.7.67e 33239 A.M.N.H.22453113348 U. S. N. M.1133490.6. 29. 25 B.M.0. 6. 29. 2588. 11. 30. 388. 11. 30.
4
1.7.11.392. 11. 24. 1092.11.24.1114689 U.S.N.M.1150851150861150871150881150891150901150941150951214761214772. 4. 7. 50 B.M.2. 4. 7. 52 B.M.
77533 U.S.N.M.7753477535775367753777538
43.0 37.4 16.047.0! 37.0 15.046.21 38.0 15.445.01 34. Ol 15.242. 2; 36. 8| 13. 643.61 37.0: 16.043.0 36.2 15.0 6.8'49", "40" 15.4 8.043.2; 38.2 14.0 7.242. 8i 41. 61 15. 4 7. 647.0' 38. 2i 16.0 7.2
39. Oj 5.8 35.4 33.4 12.835.4 5.2 33.4 31.0 12.034.6 5.4 33.0 31.4 12.035.4 6.0 33.6 31.8 12.434.2 6.0 30.6 20.0 12.636.4 5.6j 34. Oj 32. 4| 12.236.2, 5.4l 33.6. 32. ol 11.838.6; 6. 6j 34.61 32.6
11.410.810.810.410.411.010.2
7.47.88.28.07.28.07.6
35. O: 5. 6 30. 6 29. 4i 12.
4
36.0 5.81 3236.0 5.2! 33. o;44.41 37.6 15.0; 7.21 35.2 6.0: 33.841.81 34.8 13. 8| 7.040. 0| 31.2 14.0: 6.643. 4| 37.4 14.81 6.444.61 34.81 14. 0! 6.241.2 39.245.01 34.443.2: 32.043.4 31.845.0, 35.647.0 37.844.6; 35.0,44.4 37.2'43.6 34.8'44.0 37.243.2 36.241.6 33.240.0; 36.0;39.8 33.040.0 35.241.0 39.040.2 36.044.0; 35.0;41.2 34.044.4: 37.4,41.81 36.441.4;44.044.443.443.041.845.649.040.444.440.643.0.
37.835.235.037.236.037.235,036.435.0,33.6:41.637.0
15.015.014.2;14.4;15. 0|15. 4
1
15.015.813.415.015.014.2!14.6:14.8112.8!13.6114.2'14.8115.014.6:14.8!14.614. 6[14.814.6114.8!15.0:14.0114.214.613.614.614.013.8
32.0 5.433.2 5.235.2 5.233.21 5.035.0 4.634.4 6.033.2 6.032.2 6.034.8 5.436.0 6.035.0 5.435.6; 5.034.0 5.235.8 5.034? 6.034.0 5.034.4 4.835.6 4.4
30. Oi29.831.8
30.4 12.032.0! 12.833. Oi 13.029.01 12.0
12.010.812.0 8.07.88.4
28.630.29.6 28.432.2 30.832.0 31.231.0 30.030.6 29.6!32.6; 31.632.41 31.8!32.01 30.6:
!31.6 30.6!31.2 30. ol34.01 31. 2|33.0; 30.4;29. 4|31.8131.4
12.413.013.012.211.412.212.011.413.012.013.012.413.2
11.2; 8.010.4; 7.09.4' 7.010.4 8.29.4 8.211. 6i 8.69.4 &010. 2I 7.011.01 7.69.8l 6.811. 2I 8.011.2; 7.6
110.41 7.011. 2I 7.212.2 7.2
39.2 35.8 14.443.0 37.4 13.839.8 36.0; 14.041.2 32.4 13.640.0 35.0 14.442.0 37.2 14.0
6.0 345.0 347.0 357.2
5.48; 4.42 5.22 5.42; 5.62 4.88l 5.00; 5.0
I01 5.45.24 5.25.4i0| 5.46 4.44.85.05.45.25.05.0
38.037.035.636.436.036.6
30.833.033.230.632.232.233.031.831.233.233.231.033.031.032.632.032.833.433.029.832.232.233.0
34.033.033.032.832.433.2
12.012.011.029.4 11.030.81 12.831.8! 12.630.8; 11.231.0 12.230.4 11.631. Ol 12.631.2 12.430.030.830.231.830.031.832.032.428.231.231.832.2
32.031.431.031.230.032.0
13.612.812.812.812.412.612.012.812.81111.011.6
13.013.412.812.212.013.4
11.0 7.410.0; 7.29.8j 7.29.8; 6.810.4 7.410. ol 7.210.4: 7.011.0 7.411.4! 7.010.4; 7.29.81 7.411.4 7.811. ol 8.011.8 7.810.6 8.011.0 8.410. ol 6.810.410.410.810.610.010.0
10.09.89.S9.410.0
8.07.07.87.46.67.4
7.88.08.07.68.07.6
? Type of Myotis ruber keaj/si J. A. Allen.58518?28 13 ? Type of Myotis esmeraldx J, Aj Allen.
186 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis nigricans?Continued
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 187Cranial measurements of Myotis nigricans
Locality
Myotis nigricans nigricansCosta Rica:CartagoSipurioDoDo.Do..DoDo....Do....San SalvadorDoPanama:ChiriquiTaboga Island...ChiriquiDo...San PabloColombia:BondaDo...BuenavistaDoChocoDo...Do.Rio Frio, Cauca.Venezuela:MaripaDoDoDoDoDoDo.Do...DoDo....San EstebanDoDoDoDoBritish West Indies:GrenadaTobagoDoEcuador:EsmeraldasDoSan Javier..DoDoDoDoDoDo....DauleDoPuna IslandPeru:Inca MinesDoDo....DoJuninCuzcoDoDoMoyobambaDoMasisea
Number
98. 10. 9. 1 B. M62270622796228262253622556225962283'1114 A.N. S1115 P.17497917512418735A.M.N.H.2 187366763 M. C. Z.
3 14587 A.M.N.H.1514334239A.M.N.H.34240 A.M.N.H.14.5.28.3 B.M.14. 5. 28. 3219467 M. C. Z.'32787 A.M.N.H.17050 A.M.N.H.1705117072170871706517066? 17069170791708117086142568 U.S.N.M.14257214257414256914257614600 M.C.Z.97. 6. 7. 33 B. M.97. 6. 7.34 B. M.
> 33239 A.Ivf.N.H.3324013334313334413334013334113334513334613334736258 A.M.N.H.36259
' 3626315814A.M.N.H.1607316563165640.7.7.3 B.M.98.11.6.598.11.6.498.11.6.319218 FM.19219138 (orig.) B.M.
' Cotypes of Vespertilio concinnus H. Allen.
' Type of Myotis chiriquemis J. A. .\llen.* Type of Myotis bnndx J. A. Allen.* Type of Myotis caucensis J. A. .A.llen.
14.014.214.214.114.114.314.2is.'g14.01,3.614.013.513.714.713.213.313.113.513.2
188 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranml measurements of Myotis nigricans?Continued
Locality
AMEEICAN BATS OP THE GENERA MYOTIS AND PIZONYX 189MYOTIS CHILOENSIS (Waterhouse)(Synonymy under subspecies)Distribution.?South America from northern Ecuador to southernChile ; eastward to the coast of Argentina.Diagnosis,?Like Myotis niffncans but larger (forearm 35 to 43.4mm., usually more than 37 mm.) ; greatest length of skull 14.2 to 16.2mm., usually more than 14.5; maxillary tooth row 5.4 to 6.3 mm,,usually more than 5.5 mm. ; mandibular tooth row 5.5 to 6.5 mm.,usually more than 5.8 mm.; ear longer, extending distinctly be-yond tip of muzzle when laid forward; and crown area of uppermolars obviously greater {m? usually 1.30 to 1.50 by 1.70 to 2.00mm. instead of 1.15 to 1.30 by 1.55 to 1,70 mm.), see Plate 1, Figure2 (p. 7).Exterrud characters.?The general external characters do not differappreciably from those of Myotis nigricans except that the entireanimal is larger and more heavily built, the foot is more robust andthe ear is longer, usually 14 to 15 mm. This last feature is especiallywell developed in one of the local forms occurring in the Andeanregion.Fur and color.?The fur tends to be longer and more silky thanin Myotis nigricans (longest hairs at middle of back about 8 mm.),and in some of the races the glossy tips to the hairs on the dorsalsurface are more conspcuous, though not sufficiently abundant toproduce a definite sheen like that which occurs in M. albescens. Thegeneral color ranges from a rich, dark brown to light grayish oryellowish.Skull and teeth.?Except for their greater size the skull and teethdo not show any special peculiarities as compared with those ofMyotis nigricans. Brain case smooth, with at most a slightlydeveloped sagittal crest,Remarhs.?The differences between Myotis chiloensls and M.nigricans are not very obvious at first sight, but the two animalsappear to be perfectly distinct from each other. In collectionsMyotis nigricans is the commoner of the two, and the material ex-amined appears to show that it has the wider geographical range.The grayish and yellowish forms of Myotis chiloensls are at oncedistinguishable from M. nigricans by their color alone, no corre-sponding phases of the smaller animal having yet been discovered.The same is true of the largest race ; it obviously exceeds all knownforms of M. nigricans in size. True M. chiloensls and a dark formrather widely distributed in the central Andean region are lesseasily recognized, as their size and general color are not verj^ differ-ent from those of the better known species. The longer ears and
190 BULLETIN U4, UNITED STATES NATIONAL MUSEUMgreater area of the molar crowns are, however, sufficiently obviousfeatures to be diagnostic.MYOTIS CHILOENSIS CHILOENSIS (Waterhonse)
Yesperiilio chilocnsis Waterhouse, Zool. Voyage H. M. S. Beagle, pt. '^,Mammalia, p. 5, pi. 3, 1838.
?
Temminck, Mouogr. de Mamm., vol. 2, p. 271,1840.?Gervais, in Gay's Hist. Chile, Zool., vol. 1, p. 42, 1847; Atlas,Mammalogie, pi. 1, figs. 3, 3a ; Exped. dans I'Am^rique du Sud du Castelnau,Zool., Mamm., p. 83, pi. 15, figs. 4-Ad, 1855 (part).
?
Dobson, Catal. Chirop-tera Brit. Mus., p. 322, 1878.
?
Wagner, Schreber'.s Siiugth., Suppl., vol. 1,p. 537, 1844 ; vol. 5, p. 7->3, 1855.
?
Lataste, Actes Soc. Sci. Chile, vol. 1(1891), p. 79, 1892.?Trouessart, Catal. Mamm. viv. foss., p. 131, 1897.Vespertilio gayi Lataste, Actes Soc. Sci. Chile, vol. 1 (1891), pp. 79, 81, 1892(Valdivia, Chile).Myotis diilocnsis Trouessart, Catal. Mamm. viv. foss., suppl., p. 94, 1904.MyOtis gayi Trouessart, Catal. Mamm. viv. foss., suppl., p. 94, 1904.Type locality.?Islets on the eastern side of Chiloe Island, southernChile.Type specimen.?The type was collected in January, 1836, by Lieu-tenant Sullivan. It is not positively known to be extant, bat there isan unregistered specimen in the Tomes collection, British Museum,which may be the one taken by Sullivan. It was received by Tomesfrom the Zoological Society of London and has been labeled byThomas as possibh' the type.Dhtrihution.?Typical Myotis cMloensis is found on the dampcloud-wrapped coast of southwestern Chile, where the " humid andimpervious forests " seemed to Darwin, who brought back the originalspecimen, most uncongenial to a bat. It was during this same voyageof the '? Beagle " that Darwin saw a small bat at Tierra del Fuego,but did not capture it.Diagnosis.?A dark, saturate race ; size small ; back lacking almostentirely the shining tips to the longer hairs; underparts uniformsmoky brown, paling very slightlj^ in the mid-A^entral line.Color.?General color above dark brown, nearly " Vandyke brown "(Ridgway, 1912), the bases of the hairs scarcely darker. Below, thegeneral tint is less dark, becoming brownish and gray in the centerof the abdomen through the presence of pale tips to the long hairs.The dorsal hairs seem to lack the burnished tips almost completely.Mea'Surem?-nts,?For measurements see tables, pages 195 and 196.Specimens examined.?Total number 8, from the followinglocalities
:
CHILE: Temueo, Petal, 2 skins, 4 ale. (B. M.), 1 ale. (U.S.N.M.) ; no exactlocality, 1 ale. (B. M.).Remarks.?Myotis chil&ensis chU'Oensis is the darkest race of thespecies. Between southern Bolivia and south-central Chile (Tenuico)
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 191
no specimens from the higher elevations are available to show thetransition between it and M. c. dinellii of the drier country east ofthe Andes. Passing northward along the coast the dark form soonpales into the desert race atacamensis.There seems little doubt that the Vespertilio gayi Lataste, based onthe description of Vespertilio chlloensis in Gay's Historia de Chile, isthis same animal. The account in Gay's Historia was based on aspecimen from Valdivia, Chile, where the author had found it com-mon in the houses. The specimen he described and figured wasabnormal in having the second upper premolar crowded inwardfrom the tooth row so as to stand in the angle between first and thirdpremolars. The forearm measurement (1 inch and 3 lines?Frenchtranslated into Spanish) was about 35 mm., the same as in smallindividuals of M. chiloensis chUoensls. Apparently no other Myotishas been discovered so far south in Chile. While no specimens areat hand from Valdivia, those from Temuco, only a short distanceto the northward, represent typical M. chiloensis.Lataste mentions a breeding colony of these bats which he foundon November 21 in a disused copper foundry; he captured severaladult females and found that each contained a single embryo orhad just given birth to its young.MYOTIS CHILOENSIS DINELLII ThomasVespertilio chiloensis Dobson, Catal. Chiroptera Brit. Mus., p. 322, 1878 (part).Vespertilio sp. Thomas, Ann. and Mag. Nat. Hist., ser. 6, vol. 20, p. 214, August,1897.Myotis dinellii Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol. 10, p. 493, Decem-ber, 1902.
?
Tkouessakt, Catal. Mamm. viv. foss., suppl., p. 94, 1904.Type locality.?Tucuman, Catamarca Province, Argentina.Type specimen.?Female, skin and skull. No. 0.7.9.4 British Mu-seum (Natural History), from Tucuman, Catamarca Province,Argentina. Collected April 7, 1899, by L. Dinelli.DistHhution.?Argentina, Uruguay, and Bolivia; exact limits ofrange not known.Diagnosis.?A bright yellowish-brown race of Myotis chiloensis;throat and belly grayish buff; color of back uniform, withoutspeckling.Desadption.?Color above varying from deep " ochraceous buff "(Tucuman skins) to light "ochraceous tawny" (Bolivia). Thelong hairs of the back are the usual blackish brown for their basaltwo-thirds, and the light-colored tips are slightly burnished. Below,the hairs are brownish black basally, with fine buffy tips, giving tothe throat, chest, and belly a general dull grayish-buff cast, in somespecimens washed lightly with ochraceous. Ears and membranesbrownish black.
192 BULLETIN 144, UNITED STATES NATIONAL MUSEUMMeasure77ients.?For measurements see tables, pages 195 and 196.Specimens examined.?Total number 40, from the following locali-ties:ARGENTINA: Chubut, Gaimaa, 2 ale. (B. M.) ; Cordoba, 4 ale. (B. M.),8 ale. (U.S.N.M.) ; Mendoza, 1 ale. (B. M.), 1 ale. (M. C. Z.), 3 ale.(U.S.N.M.) ; Merced, Salta Province, 1 ale. (B. M.) ; Neuquen, 1 ale.(B. M.) ; northwestern Argentina, 2 ale. (B. M.) ; Pampa Grande,2 ale. (B. M.) ; Tucuman, Catamarca Province, 5 skins (B. M.) ; UpperCachi, Salta Province, 1 skin, 2 ale. (B. M.) ; Valle de Lerma, SaltaProvince, 1 skin (B. M.).BOLIVIA: Paratani, 1 skin (B. M.).URUGUAY: Dept. of Soriano, 1 ale. (B. M.) ; no exact locality, 4 ale.(U.S.N.M.).Remarks.?East of the Andes the increasing dryness of the climatefinds a response in the brightening hue of the chiloensis bats. A skullfrom Salta Province, Argentina (6.5.8.2 B. M.) lacks the minutemiddle premolar on both sides of the upper jaw, while the first pre-molar stands fully in the tooth row and fills the entire space betweencanine and third premolar {p*).MYOTIS CHILOENSIS ATACAMENSIS (Lataste)Vespertilw atacamensis Lataste, Aetes Soc. Sci. Chile, Santiago, vol. 1 (1891),p. 79, 1892 (Atacama, Chile).
?
Phllippi, Anales Mus. Nac. de Chile, sect. 1,zool., No. 13, p. 5, pi. 1, fig. Or-d, 1896.
?
Teotjessabt, Catal. Mamm. viv.foss., p. 130, 1897 (as synonym of V. nigricans).Myotis atacamensis Trotjessart, Catal. Mamm. viv. foss., suppl., p. 94, 1904.Type locality.?Atacama, Chile.Type specimen.?Mounted specimen, No. 277, National Museum ofChile. Collected February, 1885.Distribution.?This bat seems to be common in the vicinity of San-tiago and Valparaiso, Chile, but how much farther north it occurs atthe lower levels can not yet be said. With increasing dryness north-ward and lack of suitable shelter it no doubt becomes rare or absentaltogether at the lower elevations. A single specimen from Ollantay-tambo, Peru, altitude 9,400 feet, appears to be not distinguishablefrom the Chilean skins.Diagnosis.?Similar in size and proportions to Myotis cMloensiscMloensis but color a light dull (not yellowish) brown above, grayishbelow.Color.?General color above a grayish brown, ranging from nearly"wood brown" (Ridgway, 1912) to drab; the tips of the long hairsfaintly glossy. Below, the general color of the surface is grayishwhite, more or less washed with brownish, especially at the sides andunder the chin. The individual hairs are plumbeous at the basetipped with grayish white or brownish white, except at the extreme
AMEBICAN BATS OF THE GENERA MYOTIS AND PIZONYX 193
sides of the abdomen and in the anal region, where the hairs are clearwhite throughout. Membranes blackish brown.Skull.?The skull is like that of true M. chiloensis, with a rathermarked narrowing of the braincase anteriorly, and a weakly developedsagittal crest. The two minute upper premolars are seldom exactlyin the line of the toothrow, but are usually drawn slightly inward,with a tendency for the longer anterior tooth to stand partly to thepalatal side of the base of the canine.Measurements.?For measurements see tables, pages 195 and 196.Specimens examined.?Total number 12, from the followinglocalities
:
CHILE: Santiago, 6 skins (B. M.) ; Valparaiso, 3 skins (B. M.) ; no exactlocality, 2 skins (B. M.).PERU: Ollantaytambo, 1 skin (U.S.N.M.).Remarks.?It is well known that as one proceeds northward coast-wise, from southern Chile, the country becomes dry and even desertin contrast to the damp cool climate of the southern coast. Thesedrier conditions have had their effect on the local representatives ofMyotis chiloensis, producing a pale-brown race with grayish-whitebelly, in contrast with the excessively dark 31. chiloensis cMloensis ofthe humid coast region, and the yellowish M. c. dinellii of the di"ycountry east of the Andes.MYOTIS CHILOENSIS OXYOTUS (Peters)Vespertilio oxyotus Petees, Monatsber. K. Akad. Wiss. Berlin, 1866, p. 19.
?
DoBSON, Catal. Chiroptera Brit. Mus., 1878, p. 320.
?
Troxjessart, Catal.Mamm. viv. foss., p. 130, 1897.Myotis thoniasi Cabrera, Bol. Soc. Espan. Hist. Nat., Madrid, vol. 1, p. 370,1901; vol. 2, p. 293, 1902 (Archidona, near headwaters of the Rio Napo,Ecuador).
?
Teouessart, Catal. Mamm. viv. foss., suppl., p. 95, 1904.Myotis oxyotus Trouessart, Catal. Mamm. viv. foss., suppl., p. 94, 1904.?J. A.Allen, Bull. Amer. Mus. Nat. Hist., vol. 33, p. 384, July 9, 1914.Myotis sp. Osgood, Field Mus. Nat. Hist., publ. 176, zool. ser., vol. 10, p. 182,April 20, 1914.Type locality.?^Mount Chimborazo, Ecuador, at 9,000 to 10,000feet altitude.Type specimen.?A female ( ? in alcohol) in the Zoological Mu-seum of Munich, Germany, collected at 9,000 to 10,000 feet on MountChimborazo, Ecuador, by Dr. Moritz Wagner.DistHhution.?Andean region from northern Ecuador southwardinto Peru, and probably to northern Argentina.Diagnosis.?Very similar to typical Myotis cMloensis in generalappearance, but color less dark, glistening tips of longer hairs onback more conspicuous, and ear usually longer.
194 BULLETIN 144, UNITED STATES NATIONAL MUSEUMColm\?Upper surface nearly " Mars-brown " (Ridgway, 1912)or occasionally not so dark, gradually darkening to blackish slateat the bases of the hairs. The tips of the longer hairs have a con-spicuous buffy gloss imparting a finely speckled appearance to theentire back. Below, the colors are much the same as on the dorsalsurface, but the buffy tips are paler and more abundant, producing adefinite wash of dull whitish without gloss.Measureriients.?The dimensions of the type as given by Peters,are: Total length, 93 mm.; tail, 46; foot, 6; ear, IT; tragus, 8;forearm, 40; third digit, 63; fourth digit, 55; fifth digit, 52; tibia,16.5; calcar, 14. For measurements of the material examined, seetables, pages 195 and 196.Specimens exainined.?Total number 35, from the followinglocalities
:
BOLIVIA: Cliulumani, 2000 meters, 2 skins, 1 ale. (B. M.).ECUADOR: Loja, 200 meters, 1 skin (B. M.) ; Pallatanga, 1 skin(A. M. N. H.) ; Santa Rosa, Loja, 4 ale. (B. M.) ; Sigsig, 8500 feet,5 skins (B. M.) ; no exact locality, 1 skin, 1 ale. (B. M.).PERU: Chachapoyas, 7600 feet, 1 skin (B. M.) ; Hacienda Limon, 3 mileswest of Balsas, 1 ale. (F. M.) ; Inambiri River, Carabaya, 1 skin(M. C. Z.) ; Inea Mines, 1 skin (A. M. N. H.) ; Iquente, 1 ale. (U.S.N.M.) ;Junin, Rumieruz, 2 skins (A. M. N. H.) ; Rio Negro, 1 skin (A. M. N. H.) ;Santa Ana, 3500 feet, 3 skins, 7 ale. (U.S.N.M.) ; Sureo, near Lima,1 ale. (B. M.).MYOTIS CHILOENSIS ALTER, new subspeciesType locality.?Palmeiras, Parana, Brazil.Type specimen.?Adult female (in alcohol), No. 0.6.29.23, BritishMuseum (Natural History). Collected at Palmeiras, Parana, Bra-zil, by G. Grillo. Keceived from Genoa Museum.Distribution.?Eastern Argentina and southern Brazil.Diagnosis.?Like Myotis cMloen^is oxyotus but general size greater,and skull, in particular, larger, its total length ranging from 15to 16.2 mm. instead of from 14.2 to 15 mm.Measurements.?For measurements see tables, pages 195 and 197.Specimens exommied,.?Total number 21, from the following locali-ties:ARGENTINA: Buenos Aires Province, General Lavalle, 4 skins (M. C. Z.),3 skins, 6 ale. (B. M.), 2 ale. (U.S.N.M.) ; Cliubut, 1 ale. (B. M.).BRAZIL: Palmeiras, Parana, 5 ale. (B. M.).Remarks.?Apart from its larger size Myotis chiloensis alter dif-fers in no appreciable character from the Andean M. c. oxyotus.
AMEBICAN BATS OF THE GENERA MYOTIS AND PIZONYX 195External measurements of Myotis chiloi'nsis
Locality
196 BULLETIN- 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Myotis chiloensis?Continued
Locality Number
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 197Cranial measurements of Myotis chiloensis?Continued
Locality
Myotis chiloensis oxyotus?ContinuedBolivia:ChulumaniDoDo
Number
Myotis chiloSnsis alterBrazil:PalmeirasDoDoDoDoArgentina:General Lavalle.DoDoDoDoDoDoDoDo
1.6.7.1 B.M.1.6.7.22.2.2.125
0.6.29.21 B.M.0.6.29.221 0.6.29.230.6.29.2490.6.20.119509 M.C.Z.195101951119512236236 U.S.N.M.2362379.12.1.5 B.M.9.12.1.69.12.1.7
198 BULLETIN 144, UNITED STATES KATIONAL MUSEUMVespenis iwlythrix Fitzingek, Sitzungsber. Akad. Wissensch. Wien, vol. 62, pt. 1,p. 143, 1870.2ryctop1iylax nubilus Fitzingek, Sitzungsber. Akad. Wissensch. Wien, vol. 62,pt. 1, p. 501, 1870.Myotis ruber Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol. 10, p. 493, Decem-ber, 1902.
?
Trouessakt, Catal. Mamm. viv. foss., suppl., p. 94, 1904.Myotis polythrix Trouessabt, Catal. Mamm. viv. foss., suppl., p. 94, 1904.Type locality.?Paraguay, probably near Asuncion,Type specimen.?None specified; the name is based on Azara's
" Chauve-souris onzieme, ou Chauve-souris cannelle."Distribution.?The range can not at present be accurately definedIt probably includes, however, southern and eastern Brazil, all ofParaguay, northeastern Argentina, and the adjacent part of Peru.Diagnosis.?Like Myotis cMloensis but skull of adult normallyynth. a conspicuous sagittal crest, and color usually reddish. Dis-tinguishable from M. nigricans in the red phase by larger generalsize, more robust teeth, and the normal presence of a sagittal crest.Extti^ial form.?The general external features, except as they aremodified by the animal's larger size, do not differ appreciably fromthose of Myotis chilo'ensis and M. nigricans. The foot, however, isproportionally as well as actually larger than is usual in either of thebetter known species, the ratio of its length to that of tibia aver-aging 54.1 in five specimens from Paragua}'. Calcar, as in M. chilo-
'ensis and M. nigricans, with a low but usually obvious keel. Ear aslong, relatively to muzzle, as in M. chilo'ensis, extending when laidforward about 2 mm. beyond nostril.Fur and color.?The pelage in botR quality and distribution isessentially as in Myotis nigricans. The hairs on the upper part ofthe back are longest, about 6 mm. ; they lack conspicuous burnishedtips.There are, as in Myotis nig^ricans, two well-defined color phases,rufous and brown, but, contrary to the conditions prevailing in thesmaller animal, it is the rufous phase which appears to be dominant.In rufous pelage the entire dorsal surface to the roots of the hairsis a bright " ochraceous tawny" (Ridgway, 1912), darkening on themuzzle and on the chin to maroon or " russet," the tips of the hairswithout evident gloss. The under side of the body is a peculiar tintof brownish yellow, close to "yellow ocher" (Ridgway), well shownin D'Orbign}'^ and Gervais's figure; the bases of the hairs are darker,nearly " Prout's brown." In the brown phase, upper parts " Prout'sbrown," darkening at bases of hairs to " mummy brown," the tips ofthe hairs with a slight gloss ; underparts essentially similar to back,but the color so lightened as to suggest a dirty buff, darker on chinand throat, paler posteriorly.
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 199Skull and teeth.?The cranial and dental characters are essentiallylike those of Myotis chiloensis except that the skull is more heavilybuilt and the sagittal and ISmbdoid crests are better developed. Anoticeable sagittal crest aj)pears to be always normally present inadult skulls imparting to the brain case a characteristic ridged ap-pearance quite different from the nearly smooth surface usually seenin M. chiloensis and in the smaller M. nigricans.Measure7nent8.?For measurements see tables, pages 199 and 200.Specimens examined.?Total number 10, from the folloAvinglocalities
:
BRAZIL: Rio Grande do Sul, San Lorenzo, 2 ale. (B. M.) ; no exactlocality, 1 skin (B. M.).PARAGUAY: Sapiicay, 1 skin, 2 ale. (B. M.) ; 1 skin, 1 ale. (U.S.N.M.)."SOUTH AMERICA": 2 skins (B. M.).
. Remarks.?Although named more than 120 years ago on the basisof Azara's description, Myotis ruber is still rather imperfectly known.The exterior was well represented by D'Orbigny and Gervais, but
? 1 1 5they figured and described the teeth as i ^^ c y* m -^^ thus followingAzara, who says, "Dans la machoire d'en-haut, est une incisive dechaque cote, laissant un espace au milieu; ensuite vient une canine.En-bas, il parait y avoir deux incisives reunies, et tout de suite deuxcanines.'" Temminck had already doubted the correctness of Azara'sstatement regarding the number of incisors, on the basis of the in-dividual (from Corrientes, Argentina) figured by D'Orbignj'^ andGervais. The skull in question, however, was probably that of aTadanda {^^ Nyctino^nus''^) , presumably the specimen later figuredin more detail by Gervais,^^ wherein are seen the minute two-cuspedlower incisors of characteristic form. In 1902 Thomas for the firsttime allocated Geotfroy's name and briefly defined the more salientpeculiarities of the species.External measurements of Myotis ruber
Locality
200 BULLETIN 144, UNITED STATES NATIONAL MUSEUMCranial measurements of Myotis ruber
Locality
Paraguay:SapucayDoDoDoBrazil:No eiact locality.Kio Grande do SulDo"South America"
Number
115097 U.S.N.M.2.11.7.1 B. M.2.11.7.182.11.7.1948.5.6.784.2.8.3188.11.30.57.1.1.529
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 201Type locality.?Paraguay.Type specimen.?None specified. Geoffrey's name is based on the
" chauve-souris douzieme " of Azara/^ hence the type locality is con-sidered to be Paraguay, perhaps near Asuncion where Azara resided.Distribution.?Warmer parts of America from Costa Rica toVenezuela, Paraguay, and Patagonia. Limits of range imperfectlyknown.Diagnosis.?Externally much like Myotis chiloeTisis except thatthe foot is slightly larger (average ratio of foot to tibia about 52instead of about 48 or less), the ear is shorter and the tips to thelonger hairs of the posterior half of back, in fresh pelage, usuallyform a noticeable pale area. Skull with brain case inflated androstrum relatively short and weak. , Cheek teeth much smaller thanthose of Myotis cMloensis (the crown of m^ usually 1.20 to 1.25 by1.35 to 1.50), and smaller relatively to the area of the palate thanin M. cMloensis or any other known American member of the genus.Ears.?The ear is small, narrow and thin; its tip not markedlynarrowed, the anterior outline slightly but evenly convex, the pos-terior outline not noticeably concave below the tip but nearly straightand gradually passing into the convex outer base without making asharply defined shoulder. Tragus narrow and of nearly the samewidth throughout, tapering slightly at the tip, its total height lessthan half the total height from anterior lower margin to tip of ear(about 5.5:12.5 mm.). Laid forward the tip of the ear reaches thenostril.Winff and membranes.?Wing from the side of the foot at the baseof the toes. The metacarpals are slightly and evenly graduated, thethird longest (in an average specimen as 32:31.5:31). The fingersare graduated in the same order. Taking the third finger as 100, thefourth and fifth are respectively as 87 and 81 (58.5:51:47.5 mm.).When the wing is folded, the third metacarpal falls short of theelbow by about 2 mm. The fur extends thinly on the under sideof the wing as far as a line joining the knee and the proximal thirdof the humerus. On the interfemoral membrane above it extendsonly a short distance, about to a line joining the knees. The terminalvertebra of the tail is obviously free.Foot.?The foot is large, both absolutely and proportionally to thelength of tibia. Its length is slightly more than half that of thetibia (average ratio in 7 specimens from Paraguay, 52.5). Thecalcar is long (about 16 mm.) and exceeds the free border of theinterfemoral membrane by about 4 mm. It is usually without awell-defined keel and terminates in a minute projecting lobule.
1' Essals sur rhlst. nat. des quadrupfedes du Paraguay, vol. 2, p. 294, 1801.58518?28 14
202 BULLETIISr 144, UNITED STATES NATIONAL MUSEUMFur aTid color.?The fur is rather thin, of only medium length(4 or 5 mm. at the center of the back), and except imder the wing,it extends very little on to the membranes. Its color is characteristic.The basal three-fourths or thereabouts of the hairs is a uniform lightchocolate, nearly " Rood's brown " (Ridgway, 1912) ; the tips of thehairs above are glistening pale buff giving a finely peppered orfrosted effect anteriorly and forming, in fresh pelage, a noticeablewash posteriorly. Below, the bases of the hairs are of the samelight chocolate, but the terminal portion is clear whitish cartridgeproducing a contrasted pale ventral surface. Posteriorly the paletips increase in length until at the edges of the abdomen the hairsare entirely buff. The interramial area is uniformly dark chocolatebrown. The membranes are light brownish becoming distinctly palebetween the humerus and the legs. The interfemoral membrane isalso pale. In immature specimens the dark portions of the fur arenearly sooty, but the pale tips are characteristic. The membranesalso are darker, blackish brown.Skull.?The skull somewhat resembles that of Myotis chiloensis,with which it agrees in general size, but differs from that of allthe known South American members of the genus in the relativelylarge, noticeably globular, smoothly rounded form of the brain case,and the relatively short, weak rostnini. (PI. 1, p. 7, fig. 8.) Thesagittal crest, when present, is never anything more than a low,inconspicuous ridge.Teeth.?There appear to be no tangible structural characters bywhich the teeth can be distinguished from those of Myotis chiloensirS,except that the transverse diameter of the crown of the upper molarstends to be less in proportion to the diameter along the alveolar linethan in M. cMloensis or any of the other known South Americanspecies. The full complement of secondary ridges and cusps ispresent in the upper molars. Cingulum rather weakly developed.Crown area of cheek teeth unusually small in comparison with areaof intervening palate, this feature alone sufficient to distinguish thespecies from any of the members of the genus with which it mightbe confused.Measurements.?For measurements see tables, pages 204 and 205.SpeciTnens examined.?Total number 98, from the followinglocalities
:
ARGENTINA: Buenos Aires, 1 ale. (B. M.) ; Corrientes Province, Goj-a,2 skins (B. M.) ; La Pampa Territory, Rio Colorado, 1 ale. (B. M.).BOLIVIA: Rosario, 1 ale. (U.S.N.M.).BRAZIL: Amazonas Provinee, Ega (=Teffe), 1 skin, immature (B. M.) ;Manaos, 1 skin (U.S.N.M.) ; Rio Grande do Sul, 1 ale. (B. M.).BRITISH GUIANA: southern part, Ireng Valley, 1 ale. (B. M.).COLOMBIA: Barbacoas, 2 skins (A. M. N. II.); Chooo region, 2 ale.(B. M.).
AMEEICAN BATS OF THE GENERA MYOTIS AND PIZONYX 203COSTA RICA: Bebedero, 2 ale. (B. M.).ECUADOR: Cachavi, 500 ft., 2 skins (B. M.) ; Mouth of Ciuaray River,2 skins (A. M. N. H.) ; no exact locality, 1 ale. (B. M.).NICARAGUA: Escondido River, 1 ale. (U.S.N.M.) ; Prinzapolka River, 2ale. (U.S.N.M.) ; Rio San Juan, 1 ale. (U.S.N.M.).PANAMA: Tabernilla, 2 ale. (U.S.N.M.).PARAGUAY: Asuncion, 3 skins (F. M.) ; Paraguari, 2 skins (A. M. N. H.) ;Tacuaral, 24 skins, 16 ale. (U.S.N.M.), 7 skins (B. M.), 4 skins (M. C. Z.) ;no exact locality, 3 ale. (A. M. N. H.).PERU: Pto. Indiana. R. Amazonas, 10 skins (A. M. N. H.).VENEZUELA : Maracaibo, 1 ale., type of Vespertilio mtindus H. Allen(A. N, S. P.) ; Rio Aurare, 1 skin (F. M.) ; Valencia, 1 skin (B. M.).ReTiKirJcs.?This species is at once recognizable among the SouthAmerican members of the genus by its large foot, relatively short tailand tibia, globular brain case, short rostrum, and relatively weakteeth.Geoffroy's Vespertilio albescens^ based on Azara's " Chauve-sourisdouzieme," is the first name given to this species. The animal isevidently common in Paraguay, where the late William Fostersecured a large series of skins now distributed among variousmuseums. In these skins, as made up by Foster, the necks are sostretched as to present a nearly bare area at the nape where the furis thin, and this bare spot was at first supposed to be a specific char-acter as compared with the condition in other Myotis (see Thomas,1902). Throughout the wide range of the species there is very littlevariation in size or color; but, though the specimens from Paraguayare rather paler than those from farther north, particularly the seriesfrom eastern Ecuador, the latter can be matched by others quite asdark from Goya, Argentina, so that at present there seems to be noground for subdividing the species. The specimens from Barbacoas,Colombia, were referred by J. A, Allen to his Myotis punensis {=M.chilo'ensis atacamensis) , which they superficially somewhat resemble.Of the various names considered as synonyms of Myotis albescens^Wied's F. leucogaster was obviously applied to this species, as indi-cated by the excellent figure showing the characteristic chocolate-brown, pale-tipped fur, as well as the proportions of the forearmand legs. Temminck's Vespertilio cenobarhus, based on a specimenfrom an unknown locality in South America, is undoubtedly thesame. Though the complete tooth formula could not be given, Tem-minck's figure shows the slender head and the long narrow traguscharacteristic of Myotis, while the proportions of tail to total lengthand the description of the color correspond closely with those ofM. albescens. The same author's description of Vespertilio arsinoefrom Surinam, seems to apply also to M. albesceiis. It has a total ofsix molars and premolars in each jaw, the short thin pelage and pale-tipped hairs with clear white belly, and a forearm of about 33 mm.,
204 BULLETIN" 144, UNITED STATES NATIONAL MUSEUMquite as in the latter species. The type was said by Dobson to be inthe Leiden Museum. A fourth synonym is clearly Vespertilio isidoriof D'Orbigny and Gervais (1847), based on a specimen from Cor-rientes, Argentina. It has the characteristic short tail in proportionto body (head and body 40 mm., tail 28) and a forearm of 33 mm.;the color is well described as of a brownish-black at the base of thehairs, which are pale yellow at the tips above, while the belly iswhitish, the hairs brown basally, the membranes bare. To thesesynonyms is now added Vespertilio mundus H. Allen, the type ofwhich, in alcohol, has been examined through the kindness of theAcademy of Natural Sciences of Philadelphia, in whose collection itbears the number 1812. It is immature but clearly shows the dis-tinctive characters of the species. That it is Harrison Allen's actualtype is indicated not only by the locality (Maracaibo, Venezuela)but by the original parchment tag on which the number 5547 of theSmithsonian Institution can still be faintly traced.External measurements of Myotis albescens
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 205Cranial measurements of Myotis albescens
Locality
Paraguay:Asuncion.DoTacuaralDoDo....DoDoParaguariDoArgentina:GoyaDoEcuador: Cachavi...Venezuela: Rio AurareColombia:Barbacoas.--Condoto, ChocoBolivia: RosarioEcuador:CachaviMouth of the Curaray.Peru:Puerto IndianaDoDoDoDoDoDoDoCosta Rica:BebederoDo
Number
18216 P.M.18217 P.M.105562 U.S.N.M.1056611056621056641.8.1.39 B.M.23808 A.M N.H.23809 A.M.N.H.98.3.4.11 B.M.98.3.4.1297.11.7.118710 P.M.
,34241 A.M.N.n.14. 5. 28. 32 B.M.23868697.11.7.1 B.M.71644 A.M.N.n.73233 A.M.N.H.7323473235732377323873239732407324295.8.17.20 B.M.95. 8. 17. 21
13.613.513.014.114.014.213.814.214.214.214.214.014.113.914.213.214.013.813.814.214.013.813.613.613.613.614.414.0
.2^
12.812.512.213.013.213.512.813.113.5; 8.813.113.013.013.1
? a
? oo j- ^ a? o I'.Srt ? >-
13.213.412.613.012.812.813. 2|13. 0|.12.8:12.8112. 8i13.0;.12.813.4113.01.
6.25.05.05.05.4i5.2j5.015.0!5.2;
i5.215.01
10.79.78.910.010.410.310.010.110.310.010.210.410.110.210.09.610.410.09.610.010.09.69.89.69.89.610.010.0
b^ Sa
MYOTIS SIMUS ThomasMyotis simus Thomas, Ann. and Mag. Nat. Hist., ser. 7, vol. 7, p. 541, June,1901.
?
Teouessart, Catal. Mamm. viv. foss., suppl., p. 95, 1904.? (NotOsgood, Field Mus. Nat. Hist., publ. 176, zool. ser., vol. 10, p. 182, April20, 1914, ^Myotis nigricans).Type locality.?Sarayacu, on the Ucayali Kiver, Loreto, easternPeru.Type specimen.?Female in alcohol, No. 81.5.12.2, British Museum(Natural History), collected at Sarayacu, Peru, in 1876 by W. Davis.Distribution.?Amazonian drainage area of Ecuador, Peru, andBrazil. Eange imperfectly known.Diagnosis.?A small species about the size of Myotis chiloensisbut differing from all members of the genus known to occur inAmerica by the following combination of characters: "Wing mem-brane inserted at the ankle ; ear small, its tip usually extending abouthalfway between eye and nostril when laid forward ; fur of a peculiardense woolly texture, the hairs on the back about 3.5 mm. in length
;
skull with unusually well-developed sagittal crest and broad anteriorportion of rostrum; mandible unusually heavy; distance betweenupper canine and large premolar so reduced that the second small
206 BULLETIN 144, UNITED STATES NATIONAL MUSEUMpremolar is normally crowded inward behind cingulum of largepremolar.Eair.?The ear, as compared with that of Myotis nigricans, issmaller and more narrowed, especially in its upper half. Laid for-ward it extends halfway from eye to nostril as described by Thomas.We have examined skins only, and in two of these when thoroughlyrelaxed the conditions in this respect appear to be about as in thetype. Tragus not noticeably different from that of M. nigricans.Wing and membranes.?The wing membrane, so far as can bejudged from relaxed sldns, is unusually narrow at the level of theelbow; it is inserted at the ankle as in the North American 3/>/otisgrisescens. Third metacarpal falling short of elbow by about 4 mm.
;
fourth metacarpal usually a very little shorter than third; fifth ob-viously shorter than fourth and usually about 2 mm. shorter thanthird. Occasionally the fourth equals the third and in a few speci-mens the gradations from third to fourth and from fourth to fifthare equal. Membranes essentially naked throughout; on the basalportion of the uropatagium a few scattered hairs can be detectedwith the aid of a lens.Foot.?The foot resembles that of Myotis nigricans. Calcarusually with a small keel. The ratio of foot to tibia in three speci-mens from Brazil averages 49.4. In the other material examinedthe proximal end of the tibia has been cut off.Fur and color.?Fur everywhere short and woolly, without differ-ences of texture on different parts of the body and with no longhairs on the back or elsewhere. At middle of back the hairsare about 3.5 mm. in length; those of belly are slightly shorter.The fur is strictly confined to the body, not extending out on any ofthe membranes. Two color phases are represented in the series ofskins. The red phase is the more usual (13 out of 16 skins) . In it thecolor is nearly the ochraceous-taAvny of Ridgway, slightly paler,approaching yellow ocher on the underparts, the hairs without darkerbases and without conspicuous gloss. In the brown phase the generalcolor is nearl}^ Mars-brown, the underparts with a slight ochraceouscast, the hairs of the posterior half of back with faintly contrastedpaler tips which give to the fur a slight " watered " appearance.Ears and membranes in both phases blackish.Skull.?The skull is about the same general size as that of Mijotisalbescens, the brain case is similarly high, and the rostrum short (ascompared with M. nigincans and M. chiloensis). But here theresemblance ceases. The brain case of M. albescens is unusuallysmooth, with at most a slightly indicated sagittal crest; that ofM. simus is marked by better developed sagittal and lambdoid creststhan any other known South American member of the genus. InMyotis albescens the rostrum tapers noticeably forward, so that the
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 207
width across bases of canines is obviously less than the interorbitalconstriction; in M. simus the width across canines equals or exceedsthe interorbital constriction (among 12 skulls it equals the constric-tion in 3 and exceeds it in 9). The mandible is conspicuously deeperthan that of Myotis albescens and the angular process is more robust.Teeth.?The teeth are in general more robust than those of Myotisalbescens., in this respect about equaling those of M. chilo'en^is. Thecanines, in particular, are larger than those of other South Americanspecies. In both the maxilla and mandible there is a tendency tocrowding of the small premolars and reduction in the size of thesecond small tooth. This process is carried so far in the upper jawthat the second small tooth is usually hidden by the cingulum of thelarge premolar when the tooth row is viewed from the outer side.Though crowded, the second tooth is present in all of the specimensexamined. Upper molars well characterized by the narrowness ofthe internal segment resulting from the small size and slight distinct-ness of the hypocone from the base of the somewhat unusually highprotocone. Secondary cusps and ridges not well developed, the pro-toconule frequently absent or indistinct.Measureinents.?For measurements see tables, page 208.SfeGvmens examined.?Sixteen, from the following localities:ECUADOR: Mouth of the Curaray River, 13 skins (A. M. N. H.).BRAZIL: Rio Yurua, Amazonas, 3 skins without skulls (B. M.).Remarks.?Myotis simus is so different from all other knownAmerican species of Myotis that it can not be confused with any ofthem. Its shorty woolly fur, small ears, and entirely naked membranesgive it a superficial appearance which recalls some other genus : Pipis-trellus or ScotophiJivs., Thomas suggested in the original description.Among the American members of the genus there is only one, theotherwise very different Myotis grisescens of southeastern UnitedStates, which shares with M. simMs the peculiarity of having the wingmembrane inserted at the ankle. Obviously, however, there is nospecially close relationship between these two species, the cranialand dental characters of which differ widely from each other. Theunusual breadth of the rostrum anteriorly differentiates Myotis simusamong the South American members of the genus ; in North Americathe same character is found in the otherwise very different Myotisvelifer and M. occultus.The peculiar quality of the fur in Myotis simus is a characterwhich is unique among the known American members of the genus.The same may be said of the heavy mandible and unusually highsagittal crest, two characters which are probably correlated witheach other and with the shortening of the tooth row as elements ofa general strengthening of the chewing machine.
208 BULLETIN 144, TJISriTED STATES NATIONAL MUSEUMExternal measurements of Myotis simus
Locality
AMERICAN BATS OF THE GENERA MYOTIS AND PIZONYX 209Wings from the middle of the tibiae, and from the sides of the back almostas close to the spine as in Viespertilio] macrotarsus from the PhilippineIslands. Interfemoral membrane triangular behind, last caudal vertebra free;calcaneum weak, concave behind.Feet remarkably large, proportionally larger than in any other known speciesof the genus ; the outer toe much shorter than the others.Fur long and dense, extending thinly upon the upper surface of the inter-femoral membrane as far as the end of the fourth caudal vertebra, and uponthe legs to the ankles. Above, dark brown, with paler extremities; beneatha lighter shade of the same color.First and second upper premolars minute, internal to the tooth row, butdistinctly visible from without, the second very much smaller than the first;second lower premolar very small and quite internal, in the angle betweenthe closely approximated first and third premolar.Measurements.?The measurements of the type as given in theoriginal description are as follows: Total length, 115; head, 23;height of ear, 19; anterior margin of ear, 17; breadth of ear, 9.5;tragus, 9; tail, 48; forearm, 53; thumb, 14; third metacarpal, 49;fifth metacarpal, 47.5; tibia, 20; foot, 20; calcar, about 22.Remarks.?The type of Vespertilio pilosits, as we were informedby Mr. J. Berlioz of the Museum d'Histoire Naturelle, through thekindness of the late Prof. E. L. Trouessart, is no longer in the ParisMuseum, nor can any mention of it be discovered in the catalogue.The animal was supposed by Peters to have come from Montevideo,Uruguay, but its very obvious similarity to the large-footed OldWorld bats of the " Leuconoe,^'' type and especially to the PhilippineMyotis Tnacrotarsus^ with which it shares the peculiarity of havingthe wings attached high on the sides of the back, at once raises thesuspicion that there was a mistake with regard to the locality, andthat the specimen may have really come from somewhere in the EastIndies, reaching the Paris Museum perhaps through the Verreauxbrothers, notoriously careless in the labeling of the specimens whichthey offered for sale. Since a second specimen of the species hasnot been taken, either in America or in the Old World, it seems forthe present best to include the animal here provisionally, pendingthe confirmation of the alleged locality through the capture of addi-tional examples. That a bat of this kind should exist in South Amer-ica and successfully elude capture for more than half a centurycan not be regarded as an impossibility in view of the remarkablehistory of such a conspicuous animal as Pizonyx vivesi.m. THE SPECIES OF PIZONYXPIZONYX VIVESI (Menegaux)Myotis vwesi Menegaux, Bull. Mus. d'Hist. Nat. Paris, vol. 7, p. 323, 1901.
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Elliot, Land and Sea Mamm. Middle Amer., Field Columb. Mus., publ. 95,zool. ser., vol. 4, p. 574, 1904.
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^Trouessakt, Catal. Mamm. viv. foss., suppl.,p. 94, 1904.
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Elliot, Check List Mamm. North Amer., Field Columb. Mus.,publ. 105, zool. ser., vol. 6, p. 474, 1905.
210 BULLETIN 144, UNITED STATES NATIONAL MUSEUMPieonyx vivesi Miller, Proc. Biol. Soc. Washington, vol. 19, p. 85, June 4,1906; List North Amer. Land Mamra. 1911, Bull. U. S. Nat. Mus., No. 79,p. 59, December 31, 1912 ; List North Amer. Recent Mamm. 1923, Bull. U. S.Nat. Mus., No. 128, p. 73, April 29, 1924.
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McLkllan, Journ. Mamm., vol. 8,p. 243, August 9, 1927.Type locality.?Cardonal Island, Sal si Puedes Archipelago, offSan Rafael Bay, Lower California, Mexico.Cotypes.?Two females in alcohol. Both originally in Paris Mu-seum; one now in British Museum.Distribution.?Islands and coasts of the Gulf of California, Mexico.Diagnosis.?Size and general appearance, apart from the enor-
Map 13.
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Distribution of Pizonyx yivesimously enlarged feet, essentially as in Myotis nvyotis of the palearcticregion. Much larger than any known American Myotis^ the forearmabout 60 mm. in length.Ear.?The ear is rather large ; laid forward it extends about 5 mm.beyond the nostril. In general form it resembles the broad ear ofMyotis thysanodes rather than the more narrow type seen in M. veli-fer. About eight faintly defined cross-ridges are usually visible.As compared Avith the ear of M. thysanodes the antitragus is rela-tively larger and the antero-basal lobe is relatively smaller. Tragusmore bluntly pointed and relatively shorter than in M. thysanodes,its posterior margin finely but distinctly crenulate.
AMERICAISr BATS OF THE GENEEA MYOTIS AND PIZONYX 211Wing and meriihranes.?Metacarpals conspicuously graduated bysteps of about 5 mm., the fourth approximately equal to the forearm,and the third about 3 mm. longer. Wing membrane narrowed in apeculiar mafnner, so that when the wing is half extended the marginof the membrane behind the elbow extends in an approximatelystraight line from the tip of the fifth digit to the region about 15mm. out from the knee ; then it bends backward, rapidly approachingthe tibia and finally arriving at its point of insertion on the dorsalsurface of the outer side of the metatarsus. The result of this narrow-ing is that the lower leg is practically freed from the wing membrane,a condition strongly contrasted with that present in all the Americanspecies of Myotis (and in the large palearctic Myoils miyotis), inwhich the border of the membrane in the half-flexed wing extends asa straight line from the fifth finger to the ankle or foot. The endof the tail projects beyond the general outline of the free border ofthe uropatagium, but a narroAv margin of membrane extends prac-tically to the extreme tip.Foot.?The foot is remarkable for its very large size. In adultsthe claws can almost be hooked over the knee, while in half-grownindividuals they extend considerably beyond the knee when the ankleis strongly flexed. The enlargement is not due to a uniform length-ening of all the elements of the foot, but to changes which chieflyinvolve the phalanges and claws. In an ordinary large-footedMyotis such as M. thysanodes the metatarsus is more than half aslong as the combined phalanges and claws; in Pizonyx vivesi it isbarely one third of this length. The lateral compression of thedigits and claws seen in all species of Myotis is here carried to anextreme development. A few very fine hairs scattered over the sur-face of the foot can be detected with the aid of a lens. Calcar deep-ened and laterally compressed at its base, the area of its attachmentto the heel assuming almost the character of a definite joint. Itslength is about equal to that of the foot or tibia. It terminates in-distinctly and bears no keel.Fur and color.?The fur shows no special peculiarities. Longerhairs at middle of back about 8 mm. in length. It does not definitelyextend out on any of the membranes. Uropatagium practicallynaked below except for a sprinkling of long loose hairs at its baseand near the legs and a fine pubescence of minute silvery or yellowishbrown hairs on its distal third; the upper surface has the sameloosely sprinlded basal area and bare median region, but the distalthird bears a thick growth of appressed backwardly directed hairs3 to 5 mm. in length which form an almost furry coating. Colorabove nearly the wood brown or fawn color of Ridgway (1912), thehairs slaty at base; underparts whitish in strong contrast with the
212 BULLETIN 144, UNITED STATES NATIONAL MUSEUMback, the hairs mostly without darker bases except those near wingmembranes.Skull.?The skull (pi. 1, p. 7, fig. 1) is immediately distinguishablefrom that of any American Myotis by its much greater size (totallength about 22, while no American Myotis is yet known in whichthe skull ever attains a total length of 18 mm.). When comparedwith the skull of Myotis velifer, the species which most nearly ap-proaches it in size (maximum total length IT.6), that of Pizonyxvivesi is seen to present some obvious peculiarities of form. Thebrain case is less elevated behind and less abruptly constricted ante-riorly ; its sagittal crest is low and inconspicuous. The longitudinalmedian groove on the rostrum is relatively deeper and better defined.When the skull is placed on a flat surface the alveolar line rises at anoticeably greater angle than in Myotis velifer^ so that the tips of thehigh canines are elevated conspicuously above the surface on whichthe skull is resting instead of being brought almost in contact with itas is the case with the low canines of the smaller animal.Teeth.?Apart from their greater size the teeth differ from thoseof the American species of Myotis in a general tendency towardheightening and slenderness of the cusps, a tendency which is espe-cially noticeable in the canines and premolars. The height of theupper canine above alveolus is equal to decidedly more than the com-bined crown length of the four large cheek teeth, while in Myotisvelifer it is as decidedly less than half this length. Both of the smallpremolars, mandibular as well as maxillary, are obviously heightened,and the second tooth in each pair exceeds the first both in height asseen from the side and in crown area as seen in apical view, a condi-tion unknown in any American Myotis. In the mandible this height-ening of the second premolar is so great that the profile of the entirerow of cusp summits does not show the abrupt fall in front of thelarge premolar which is so characteristic of the cusp profile in ordi-nary Myotis. The incisors present no marked peculiarities. Molarsnormal in form; m^ not reduced. All of the cusps share in thegeneral tendency to heightening characteristic of the entire dentition,but this tendency is here less pronounced than it is in the canines andpremolars. In m} and in? the small ridge which, in Myotis, extendsfrom the base of the paracone to the protoconule usually runs to theprotocone, leaving the very rudimentary protoconule, when presentat all (usually absent), isolated on the margin of the crown. (Fig.Id, p. 8.)Measurements.?The measurements of the cotypes (both females)as noted by Miller in 1904 are as follows : Total length, 145 and 140
;
tail, 70 and 69; tibia, 24 and 24.6; foot, 23 and 23.8; longest claw,10 and 10; forearm, 62 and 60; thumb, 12.2 and 12.6; claw of thumb.2.4 and 3 ; second finger, 62 and 60 ; third finger, 120 and 120 ; fourth
AMEEICAN BATS OF THE GENERA MYOTIS AND PIZONYX 213finger, 93 and 91 ; fifth finger, 88 and 85 ; ear from meatus, 24.6 and25; ear from crown, 20.4 and 19; width of ear (flattened), 16 and16; tragus, 11.8 and 11. For other measurements see tables, page 214.SpeciTTiens examined.?Fourteen, from the following localities
:
LOWER CALIFORNIA: Cardonal Island, Sal si Puedes Archipelago, 2ale. (Paris; cotypes, examined by Miller in 1904), 3 ale. (A. M. N. H.) ;Isla Partida, about 12 miles north of Cardonal, 7 skinned from ale.(C. A.) ; Islas Eneantadas, 1 skin (Dickey).SONORA: Guaymas, 1 mummy (now in alcohol) (U.S.N.M.)-Remarks.?The striking peculiarities of Pizowyx vivesi render theanimal one of the most easily recognizable of American bats. Thesepeculiarities and more particularly the seemingly anomalous habitatand geographical range make it appear to be possible that this bathas some unusual mode of life. The original specimens were col-lected by Diguet, in December, 1900, under stones heaped up by thewaves along the shore of Cardonal Island. Here the bats lived incompany with a species of petrel {Hdlocy'ptena microsonm Coues).Diguet supposed that they fed on the excrement of the sea birds, buthe presented no evidence in support of this seemingly improbableidea. The specimen in the United States National Museum wasfound February 23, 1904, completely desiccated, on the beach atGuaymas, Sonora, by the late William Palmer. The viscera hadbeen destroyed by insects, so that no trace of the animal's food couldbe discovered. The digestive tract of the cotype in the BritishMuseum was examined for us by Mr. Martin A. C. Hinton, whoreports his inability to discover the slightest trace of food of anykind. That of the adult female topotype in the American Museumof Natural History was similarly explored by Mr. Eemington Kel-logg, of the United States Department of Agriculture. He foundthe stomach and intestines empty except for a few mites, some bathairs, and one small feather. While we are thus left in completeignorance as to the animal's food habits it seems not improbable thatthe enlarged foot and claws, the relative freedom of the leg from thewing membrane, and the elongation of the cusps of the teeth may allbe associated with a diet consisting at least partly of fish. Theresemblance of the leg, foot, and calcar to those of Noctilio., a batwhich is well known to devour small fish,^^ is obvious.
" For a recent account of NocUUo's piscatorialism see Benedict, Journ. Mamm., vol. 7,p. 58, Feb., 1926.
214 BULLETIN 144, UNITED STATES NATIONAL MUSEUMExternal measurements of Pieonyx vivesi
Locality
INDEX TO GENERIC, SPECIFIC, AND SUBSPECIFICNAMES[Figures in bold-face type refer to the descriptions]Pageadversus, Myotis --- 9Vespertilio 30,31aenobarbus, Vespertilio 15,200Aeorestes --- 31albescens 200nigricans 178affinis, VespertUio 15,43agilis, Vespertilio. 15,159alascensis, Myotis 10, 15, 37, 47, 57, 60, 104albescens, Aeorestes 200Myotis -. 6, 7, 11, 34, 200Vespertilio.- 15, 66, 89, 92, 197, 200albicinctus, Myotis 16,50alter, Myotis 11,35,194,195,197altifrons, Myotis 16,140altipetens, Myotis 16,50amotus, Myotis 11, 16, 35, US, 146, 147arsinoS, Myotis 200VespertOio -- 16, 200atacamensis, Myotis 11, 35, 193, 195, 196VespertUio - 16, 192austroriparius, Myotis. 5,10,16,37,76VespertOio 16, 76aztecus, Myotis 10,36,138,129,130baileyi, Myotis 17,97bayleyi, Myotis 17,98bechsteinii, Myotis 10Vespertilio 30blepotis, Vespertilio... 30blythii, Myotis 10bocagei, Myotis 9bondae, Myotis 17,179Brachyotus 30brasiliensis, Vespertilio 17,177brevirostris, Vespertilio 17,43californicus, Myotis 5,6, 11, 35, 148, 151. 155, 157, 161, 103Vespertilio 17, 151capaccinii, Myotis 9Vespertilio 30Capaccinius 30capitaneus, Myotis 17,139capucinus, Vespertilio 17carimonensis, Myotis 9carissima, Myotis 5, 8, 10, 18, 37, 50, 58, 59carolii, Vespertilio 18,43caucensis, Myotis. 18,179caurinus, Myotis 11, 18, 35, 155, 161, 163chiloensis, Myotis 4,6,7,11,34,189,190,195,196Vespertilio 18, 190, 191chinensis, Myotis 10chiriquensis, Myotis 18,178chrysonotus, Myotis 10,36,116,120,121Vespertilio 18, 116
Pageciliolabrum, Myotis 168,169Vespertilio 18, 168, 169cincinnus, Vespertilio 18,178cinnamomeus, Vespertilio 19,197Comastes 31concinnus, Myotis 178,181VespertUio 19, 178, 181crassus, VespertUio 19,43dasycneme, Myotis... 9Vespertilio 30daubentonii, Myotis 9VespertUio 30Dichromyotis 31dineUii, Myotis 11,19,35,191,195,196domesticus, VespertUio 19,43dominicensis, Myotis 6, 11, 19, 34, 183, 186, 188durangae, Myotis 19,66durangoae, Myotis 19,66emarginatus, Myotis 10VespertUio 31Eptesicus 31esmeraldae, Myotis 19,179EuvespertUio 31evotis, Myotis 4, 5, 6, 7, 9, 10, 36, 111, 114, 120, 121VespertUio 19,114,126exiguus, VespertUio 20,178exilis, VespertUio 20,152Exochurus.. 31extremus, Myotis 11,34,181,185,188ferrugineus, VespertUio 20formosus, Vespertilio 31fortidens, Myotis 10,37,54,58,61gayi, Myotis 190Vespertilio 20, 190grisescens, Myotis 4,6,10,20,36,80gryphus, Vespertilio 20,43,76,105hasseltU, VespertUio 30henshawU, Vespertilio 20, 169hUdegardae, Myotis.. 9horsfieldii, Vespertilio 31hypothrix, VespertUio 21,178hypotbryi, VespertUio 21, 178incautus, Myotis 6,10,37,92,95,96Vespertilio 21, 92interior, Myotis 11,21,^5,142,146,147isidori, VespertUio 21,200Isotus - 31jaliscensis, Myotis 21,90keaysi, Myotis 217,21,179keemi, Myotis... 4, 7, 10, 36, 101, 104, 109, 110Vespertilio 21, 104kinnamon, VespertUio 21,197lanceolatus, Vespertilio 22, 43leibu, Myotis 11,35,171,173,174Vespertilio 22, 171215
216 INDEXPageleucogaster, Vespertilio... 22,200Leuconoe 6, 10, 30limnophilus, Vespertilio 31lobipes, Vespertilio - 31longicnis, Myotis 11,35, 68, 97, 140, 142, 145, 146, 147, 170Vespertilio 22, 140
'lucifugus, Myotis.. ? 5,6, 7, 8, 9, 10, 37, 38, 43, 56, 59, 76, 130Vespertilio 22, 43, 80, 105, 140lutosus, Myotis 10,37,72,74macellus, Vespertilio 30macrodactylus, Vespertilio.. 31macropus, Vespertilio 22,66macrotarsus, Myotis. 5,9Vespertilio. 31maripensis, Myotis 23,179megapodius, Vespertilio 31melanorhinus, Myotis 11,23,35,169,173,174Vespertilio 23, 168, 169mexicanus, Myotis 11,35,159,162,164Vespertilio 23, 159micronyx, Myotis 23,117mUleri, Myotis 10,23,36,118,120,121Minlopterus 30mundus, Myotis 179Vespertilio 23,200muralis, Myotis 9muricola, Vespertilio 31murinus, Vespertilio 31Myotis 30adversus 9alascensis 10, 15,37,47,57,60, 104albescens 6,7,11,34,200alblcinctus 16, 50alter. 11, 35, 194, 195, 197altifrons 19, 226altipetens 16, 50amotus 11, 16, 35, 145, 146, 147arsinoe 200atacamensis 11,35, 192, 195, 196austroriparius 10, 16,37,76aztecus 10,36,128,129,130baDeyi 17, 97bayleyi 17,98bechsteinii 10blythii 10bocagei 9bondae 17, 179californicus 5,6,11, 35, 148, 151, 155, 157, 161, 163capaccinii 9capitaneus 17, 139carimonensis 9carissima 5, 8, 10, 18, 37, 50, 58, 59caucensis 18, 179caurinus 11, 18, 35, 155, 161, 163chiloensis 4, 6, 7, 11, 34, 189, 190, 195, 196cbinensis 10cbiriquensis 18, 178chrysonotus 10,36, 116, 120, 121ciliolabrum 168, 169coDcinnus 178, 181dasycneme 9
PageMyotis daubentonii 9dinellii 11, 19, 35, 191, 195, 196dominicensis 6, 11, 19, 34, 183, 186, 188durangae.- 19,66durangoae. 19, 66emarginatus. 10esmeraldae 19, 179evotis 4,5,6,7, 10,36, 111, 114, 120,121extremus 11,34,181,185,188fortidens 10, 37, 54, 58, 61gayi - 190grisescens 4,5,10,20,36,80hUdegardae 9incautus 6,10,37,92,95,96interior. 11, 21, 35, 142, 146, 147jaliscensis.. 21, 90keaysi 21, 179keenii 4, 7, 10, 36, 101, 104, 109, 110leibii 11,35,171,173,174longicrus. . 11, 35, 68, 97, 140, 142, 145, 146, 147lucifugus.. 5,6,7,8,9,10, 37, 38, 43, 56, 59, 76, 130lutosus -. 10. 37, 72, 74macrotarsus 5,9maripensis 23, 179melanorhinus 11, 35, 169, 173, 174mexicanus.. 11, 35, 159, 162, 164micronyx 23, 117milleri 10, 23, 36, 118, 120, 121mundus 179muralis. 9myotis 7,8,9,10mystacinus 10nattereri 10nesopolus 11, 23, 34, 182, 186, 188nesopotus 23,182nigricans 6,7,11, 34, 159, 175, 177, 181, 184, 187occultus 7,10,24,36,97orinomus. 25, 169oxygnathus 10oxyotus 11,35,193,195,196pallidus 11,25,35,157,161,164peninsularis 10, 25, 37, 92, 95, 97pernox. 25, 48phasma 10,37,53,58,61pilosus 11,34,208polythrix 198punensis 26, 179, 200quercinus 26, 152ruber 6, 11, 34, 197rufopictus 10saturatus. 10, 27, 37, 48, 50, 70, 74, 75septentrionaUs 10, 36, 105, 109, 110simus. 4, 5, 7, 11, 27, 34, 179, 205sociabilis 10,27,37,66,68,73,75sodalis 6, 10, 36, 130subulatus (Le Conte) 4,105subulatus (Say) . 6, 7, 11, 35, 164, 168, 173, 174thomasi 28, 193thysanodes 5,7, 8, 9, 10, 28, 36, 122, 126, 128, 129tricolor.. 10ussuriensis 42velifer 7, 10, 37, 42, 80, 86, 89, 94, 96
INDEX 217PageMyotis vivesi 209volans.- 4, 5, 7, 11, 36, IM, 139. 146, 147winnemana 28, 171yumaensis 66yumanensis 10,37,60,53,61,73,75inyotis. Vespertilio 3)mystacinus, Myotis 10Vespertilio 30nattereri, Myotis 10Vespertilio 30nesopolus, Myotis 11,23,34,182,186,188nesopotus, Myotis 23,182nigiieans, Aeorestes 178Myotis 6,7, 11, 34, 159, 175, 177, 181, 184, 187VespertUio 23, 152, 159, 177Vesperugo 178nitens, Vespertilio 23nitidus, Vespertilio 24, 66, 151, 155, 159nubiliis, Nyctophylax 198Vespertilio 24, 197Nyctophylax nubilus 198parvulus 178Nystactes 30obscurus, Vespertilio 24,65occultus, Myotis 7,10,24,36,97oregonensis, Vespertilio 24, 152orinomus, Myotis 25,169oiygnathus, Myotis 10oxyotus, Myotis 11,35,193,195,196VespertUio 25, 193pallidus, Myotis 10,25,35,157,161,164Paramyotis 32parvulus, Nyctophylax 178Vespertilio 25,178peninsularis, Myotis 10,25,37,92,95,97pernox, Myotis 25,48phasma, Myotis 10,37,53,58,61pUosus, Myotis 11,34,208VespertOio 5,25,208Pipistrellus 6Pizonyx-..- 6,31,33,34vivesi. 34,209polythrix, Myotis ^. 198Scotophilus 197Vespertilio 26, 197Vesperus 198Pternopterus 31punensis, Myotis 26,179,200querclnus, Myotis.. 26,152ricketti, Vespertilio 31, 32Rickettia 31,32ruber, Myotis 6,11,34,197Vespertilio 26, 197rufopictus, Myotis 10salarii, VespertUio 26,43saturatus, Myotis 10, 27, 37, 48, 50, 70, 74, 75ScotophUus polythrix 197Selysius 30septentrionalis, Myotis 10, 36, 105,109, 110VespertUio 27,105simus, Myotis 4,5,7,11,27,34,179,205sociabUis, Myotis 10,27,37,66,08,73,75sodalis, Myotis 6,10,36,130spixu, VespertUio 27,17858518?28 15
Pagesubulatus (Le Conte), Myotis 4,105VespertUio 43, 105, 126subulatus (Say), Myotis 6,7,11,35,164,168,173,174Vespertilio 27, 168tenuldorsalis, VespertUio. 28, 152thomasi, Myotis 28,193thysanodes, Myotis 4,5,7,8, 9, 10, 28, 36, 122, 126, 128, 129Tralatitius 31Tralatitus 30tricolor, Myotis 10Trilatitus 30ussuriensis, Myotis 42velifer, Myotis 7, 10,37,42,80,86,89,94,96Vespertilio 28,89,126Vespertilio 30adversus 30,31aenobarbus 15, 200affinis 15,43agilis 15, 159albescens 15,66,89,92, 197,200arsinoS... 16, 200atacamensis 16, 192austroriparius. 16,76becbsteinU 30blepotis 30brasUiensis 17, 177brevirostris... 17,43ealifornicus 17, 151capaccinii 30capucinus 17carolii 18,43chUoensls 18, 190, 191chrysonotus 18, 116eiliolabrum 18, 168, 169cincinnus 18, 178cinnamomeus 19, 197concinnus 19,178,181crassus. 19,43dasyeneme 30daubentonii 30domesticus.. 19,43emarginatus 31evotis-.. 19,114,126exiguus 20, 178exUis 20, 152ferrugineus 20formosus 31gayl 20, 190gryphus 24,43,76,105hasseltu.. 30henshawU 20, 169horsfieldii 31hypothrli 21, 178hypothryx 21,178Incautus 21,92isidori 21,200keenu. 21, 104kinnamon 21, 197lanceolatus 22,43leibU 22, 171leucogaster 22,200limnophUus 31lobipes 31
218 INDEXPageVespertilio longicrus 22,140lucifugus 22,47,80,105,140macellus 30macrodactylus 31macropus 22, 66macrotarsus 31megapodius 31melanorhinus 23, 169meiicanus 23, 159mundus 23, 200muricola 31murinus 31myotis 31mystacinus 30nattereri 30nigricans 23, 152, 159, 177nitens 23nitidus 24,66,141,155,159nubilus 24, 197obscnrus 24, 65oregonensis 24, 152oxyotus 25, 193parvulus 25,178pilosns 5,25,208polythrix 26, 197
Fag*Vespertilio ricketti 31,32raber 26, 197salarii 26,43septentrionalis 27,105spixii 27,178subulatus LeConte 105, 126subulatus Say 27, 163tenuidorsalis 28, 152velifer 28,89,126viUosissimus 31virginianus .. 28,43volans 28,139yumanensis 29,65,152Vesperago nigricans 17SVesperus polythrix 198viUosissimus, Vespertilio. 31virginianus, Vespertilio 28,43viyesi, Myotis 209?'* Pizonyx 8, 34, 209volans", Myotis 4, 5, 7, 11, 36, 135, 139, 146, 147Vespertilio 28, 139wlnnemana, Myotis. ---.-_- -
-::r? - r^r 28,171yumaen"sis7Myotis. ..._...... ... .~1 ~ 66yumanensis^ Myotis? . J . . .10737750,53, 61,'65,'73775Vespertilio 29,65,162
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