THE NORTH AMERICAN BEETLES OF THE GENUSCOCCINELLA By Th. Dobzhansky California Institute 0/ Technology , Pasadena, Calif. The North American species of the genus Coccinella Linnaeus havebeen studied particularly by Casey (1899) and by Leng (1903). Theresults arrived at by these authors are, however, quite different.More than twice as many forms are recognized as separate speciesin the Casey revision as in the more recent survey of Leng. A newsurvey of the genus is, therefore, desirable. As shown by my studies(1925, 1926) on the palaearctic representatives of the genus Coccinella,the structure of the genitalia is a first-class character for the deter-mination of the limits of the species in this genus. Consequently,the description of the genitalia is made the cornerstone of the presentstudy.A thorough comparison of the American species of Coccinella withEurasiatic ones seems also very desirable. The genus Coccinella(hmited as defined by me, 1925) inhabits chiefly the holarctic region.Only a few true Coccinella Kve outside of this region. If the holarcticfauna of this genus is properly understood, the world-wide revisionof the genus may be easily accomplished.The present study is based primarily on the examination of thecollection of the United States National Museum, which was sent to methrough the kindness of Dr. E. A. Chapin. Besides this, collectionsbelonging to the following institutions and individuals were examined : American Museum of Natural History, Cornell University, Universityof Minnesota, Indiana University, Illinois State Natural History Sur-vey, Cahfornia Academy of Sciences (including collections of E. C. VanDyke, F. E. Blaisdell, E. P. Van Duzee, and others), Citrus Experi-ment Station, F. W. Nunenmacher, A. H. Sturtevant, F. T. Scott, andP. H. Timberlake. I wish to express my gratitude to the owners andto the custodians of these collections for the privilege of examiningtheir material. No. 2904.? Proceedings U. S. National Museum, Vol. 80, Art. 468143?31 1 1 Z PROCEEDINGS OF THE NATIONAL MUSEUM vol. 80The geographical distribution of the species of Coccinella is out-lined here on the basis of the material personally studied by me.The localities are grouped in the sequence from east to west, and fromnorth to south. The names of the collectors are indicated only forrare or little-known species and for especially interesting finds.Genus COCCINELLA LinnaeusBody more or less broadly oval, from moderately to very stronglyconvex. Head black, with a yellowish-white spot on each side nearthe eyes, or with a broad transverse white band across the front.Antennae longer than the diameter of the eye, with a compact club.Antennae and mouth parts dark brown or black, the base of themandible frequently with a white spot; labrum brownish. Pronotumblack with quadrangular or triangular white markings in the ante-rior angles, and in some species also with a white stripe along theanterior margin. Underside black, the episterna and the epimera ofthe mesosternum and metasternum white in some species. Prester-num with two carinae, which are slightly convergent anteriorly.Mesosternum not emarginate in the middle of the anterior margin.The coxal lines of the first abdominal sternite divided into two sep-arate branches, one of which runs parallel to the posterior margin ofthe segment and the other directed toward the anterior angles. Legsblack, tibiae with two spurs at the end, tarsal claws with a tooth atbase. Elytra yellow, orange, or red with a variable black pattern.The elytral patterns of all the species and varieties of Coccinella maybe represented as derivatives from the basic pattern consisting of sixspots on each elytron (fig. 30). The first of these spots (^, thescutellar spot) lies on the suture at the scutellum and is common toboth elytra; the humeral spot (1) lies at the humeral angles; thelateral spot (2) lies at one-third of the length of the elytron, near theexternal border; the discal spot (3) at two-fifths of the length ofthe elytron, closer to the suture than to the external border; themarginal spot (4) at two-thirds of the length of the external border,and the apical spot (5) at four-fifths of the length of the elytron,closer to the suture than to the external border. Among the Ameri-can species only Coccinella undecimpunctata Linnaeus, some varietiesof C. johnsoni Casey, and C. transversoguttata Falderman var. nuga-toria Mulsant have the basic elytral pattern of the genus unchanged.In all other species some of the spots are either absent or confluentwith others. Especially frequent is the fusion of spots 4 and 5 intoa common apico-marginal spot (4 + 5) . This fusion is frequently sointimate that the compound nature of the resulting spot may be notat all apparent.Male genitalia.?The terminolog}" of the parts of the genitalia ofCoccinellidae proposed by Verhoeff (1895) seems to me preferable to ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY that used by other authors. The part functioning in Coccinellidaeas a penis is, as shown by its development, homologous to only theproximal part of the penis of other Coleoptera (according to theunpublished data of the author). This part is termed the sipho(penis of other authors). Moreover, the penis, which is homologousto the distal part of the penis of other Coleoptera, is intimately fusedwith the basal plates (basal piece of other authors). The fingerlikeparamera (lateral lobes) are articulated with the basal plates. Thetrabes (tegminal strut of other authors) is an unpaired chitinous rodarticulated to the basal plates, and connected by muscles with theproximal end of the sipho.In the genus Coccinella the sipho (s, fig. 1 and figs. 13-20) is hook-shaped. Its proximal end, inmost species, is strongly chiti-nized and separated from thebody of the sipho to form theso-called siphonal capsule {sc,figs, land 13-20). Onlyin(7oc-cinella undecimpunctata Lin-naeus the siphonal capsule isrudimentary (fig. 20). Thedistal end of the sipho carries bS-..rather strongly developedpraeputial sacs. The penis{p, figs. 1, 2-12) frequentlypossesses complicated proc- 03?esses on its distal end. Theform of the penis is exceed-ingly variable and constitutesthe best specific character.The trabes {tr, fig. 1) is rela-tively short and thick, and itsfree end has no clearly pro-nounced emargination. The basal plates {bp, fig. 1) are stronglydeveloped. Paramera (pa, figs. 1, 2) are fingerlike and slightlycompressed from the sides.Female genitalia.?The receptaculum seminis is large, mostly ratherclearly differentiated into the cornu (c, figs. 22, 28), the nodulus {n),and the ramus (r). The sculpture of the walls of the receptaculum,consisting of chitinous rings, is well developed in all species exceptCoccinella undecimpunctata Linnaeus and its relatives. The infundib-ulum (?', figs. 22, 28) has a funnel-shaped dilatation on its anterior endand, in some species, a similar dilatation on its posterior end. Theductus receptaculi (dr, fig. 28) is very short and nearly hidden in the Figure 1.?Male genitalia of Coccinella novemnotataHerbst (lateral view), bp. Basal plates; p, penis; pa,paramera; s, sipho; sc, siphonal capsule; tr, trabes 4 PROCEEDINGS OF THE NATIONAL MUSEUM vol.80funnel-shaped dilatation of the infundibulum. Some species have anaccessory plate (Verhoeff' s Anhangsplatte, loc cit.) attached to theposterior end of the infundibulum (a/p, fig. 22).COCCINELLA NOVEMNOTATA HerbstCoccinella novemnotata Herbst, Natursystem der Kafer, vol. 5, p. 269, 1793. ? Casey, 1899, p. 88.?Leng, 1903, p. 198; 1920, p. 216.?Johnson, 1910, pp.59-60.Body subhemispherical. Head with a broad, undulate, yellowish-white band across the front, and with yellow anterior margin of theclypeus. Pronotum and pronotal epipleura with white subquadratemarks in the anterior angles, anterior margin of the pronotum moreor less broadly white. Mesepimera and metepimera, the posteriorends of the metepisterna, and in males a spot on the anteriorcoxae and a stripe on the anterior femora, yellowish white. Head,pronotum, and elytra alutaceous, obscurely punctulate with the punc-tures somewhat stronger near the external margin of the elytra.Elytra yellow or orange, with nine black spots (K, 1, 2, 3, 4). Thespot K moderate in size, triangular or rhomboidal, spots 1 and 2 smalland usually rounded, spots 3 and 4 large in size and transverselyelliptical in shape. The spots may fuse together or may be connectedby rather slender black lines. The following patterns have beendescribed: 1+2 (conjuncta Fitch), 3 + 5, K + 3, 2 + 1 + 3, 2 + 1 + 3 + 5(confluenta Fitch). Length of body, 5.3-7 mm.Male genitalia (figs. 2, 13).?Penis rather long and narrow, itsproximal end extended in a triangular process. Paramera muchshorter than penis. Basal plates broader than long. Sipho ratherlong and slender.Female genitalia (fig. 21).?Cornu broad, ramus very small, nodu-lus conic in shape and thick-walled. Infundibulum short and thick,its posterior end dilated and surrounded by a ringlike furrow.This purely American species seems to be related to the palaearcticspecies Coccinella divaricata Olivier ( = distincta Redtenbacher) . Thepoints of similarity are the sculpture of the elytra, the shape of theelytral spots, and the shape of the sipho and the infundibulum. Theform of the penis is, however, very different in these two species.Geographic distribution.?Localities as follows:Quebec: Montreal, Chelsea.Ontario: Britannia.New Hampshire: Lancaster, Mount Washington, Franconia, Wolfeboro.Vermont: Ludlow.Massachusetts: Melrose, Medford, Arlington, Stoughton, Springfield, Plymouth,Middleboro, North Saugus, Woods Hole, Falmouth, Truro, Siasconset, OakBluffs, Edgartown, Nantucket Island, New Bedford.Rhode Island: Watch Hill.Connecticut: Brookfield, Bridgeport. ART. 4 BEETLES OF THE GENUS COCCINELLA?DOBZHANSKYNew York: Black Mountain, Thousand Islands, Kinderhook, White Lake,New York, Farmingdale, Cold Spring Harbor, Riverhead, West Point, Ithaca,Forest Lawn, Honeoye Falls, Batavia, Dansville.New Jersey: Fort Lee, Hackensack, Paterson, Passaic, Ramsey, Boonton, New-ton, Mendham, Milltown, Lakehurst.Pennsylvania: Philadelphia, Ashbourne, Glenside, Reading, Lehigh Gap, Get-tysburg. Figures 2-9.?Male genitalia of the diSerent species of Coccinella: 2, Coccinella novem-notata; 3, C. prolongata; 4, C. californica; 6, C. johnsoni; 6, C. transversoguttata; 7, C.nivicola; 8, C. suluralis; 9, C. difficitis. In all figures the sipho and the trabes are notrepresented, bp. Basal plates; p, penis; pn, parameraDelaware: Newark.Maryland: Baltimore, Hagerstown, Odenton, Glen Echo, Riverdale.District of Columbia: Washington.Virginia: Falls Church, Vienna, Arlington, Fredericksburg, Bowling Green,Cuckoo, Richmond, Norfolk, Blue Ridge Mountains, Staunton, Afton, Peaksof Otter, Pennington Gap. 6 PEOCEEDINGS OF THE NATIONAL MUSEUM vol.80West Virginia: White Sulphur Springs.North Carolina: Areola, Southern Pines, Salisbury, Black Mountain.South Carolina: Oswego, Batesburg, Beaufort.Georgia: La Grange, Baconton, Moultrie, Thomasville.Florida: State record.Michigan: Whitefish Point, Marquette, Port Huron, Detroit, Douglas.Ohio: Newton Falls, Salineville, Columbus.Indiana: Knox, Culver, Nashville, Bloomington, Mineral.Illinois: Chicago, Kankakee, Fulton, Oakwood, Urbana, St. Joseph, Champaign,White Heath, Charleston, Topeca, Havana, Dubois, MetropoHs.Kentucky: Campton.Tennessee: Black Mountains, Coal Creek.Alabama: Longview.Louisiana: Tallulah, Mound.Wisconsin: Waupaca, Madison, Osceola.Minnesota: St. Paul, Minneapolis, High Prairie, Hennepin County, Lake Crystal,Minnehaha Creek, Lake City, Jordan, Shakopee, Rice County, St. Peter,Lesueur Center, Albert Lea, Owatonna, Mora, Brooten, Princeton, New Lon-don, Taylors Falls, Willow River, Houston County, Itasca Lake, Luverne,Ramsey.Iowa: Muscatine, Ames.Missouri: St. Louis, Utica, Willard.Arkansas: Siloam Springs.South Dakota: Madison, Black Hills.Oklahoma: Hobart.Texas: Dallas.Wyoming: Carbon County.Colorado: Boulder, Colorado Springs, Manitou, Rocky Ford.Remarks.?In the individuals coming from the Eastern and South-ern States the elytral spots are distinctly larger than in those fromMinnesota, Missouri, and Iowa. Specimens from Wyoming andColorado are intermediate between the typical novemnotata and thevariety degener Casey (see below). Moreover, the frequency ofspecimens having confluent elytral spots is higher in Eastern andSouthern States, and lower in the Middle West.COCCINELLA NOVEMNOTATA Herbst subspecies DEGENER CaseyCoccinella degener Casey, 1899, p. 88.Coccinella novemnotata Herbst var. degener Casey, Lentg, 1903, p. 198; 1920, p.216.?Johnson, 1910, p. 59.This race differs from the typical form by smaller size of thebody, more polished surface of the elytra, and by smaller, sometimesabsent, elytral spots. The genitalia of both sexes are not signifi-cantly different from those of the typical novemnotata Herbst. Thegeographic distribution of degener Casey gives a convincing evidencein favor of considering it as a subspecies of novemnotata Herbst, andnot as a separate species. Length of the body, 4.8-6.2 mm. AHT. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 7 Geographic distribution.?Localities as follows:Saskatchewan: Carlyle.Nebraska: Mitchell.Kansas: Douglas County.Oklahoma: Summit, Hobart.Montana: Assiniboine, Bear Paw Mountains, Helena, Powderville, BroadwaterCounty, Yellowstone County.Wyoming: Big Horn Mountains, Canyon Camp (Yellowstone Park), Wheat-land, Cheyenne, Carbon County.Colorado: Pine Creek, Fort Collins, Greeley, Dixon, Boulder, Longs Peak (9,000feet), Summit County, Golden, Denver, Florissant, Manitou, ColoradoSprings, Rocky Ford, Buena Vista, Paonia, Salida.New Mexico: Espanola, Santa Fe, Albuquerque, Koehler Junction, Coolidge.Idaho: Moscow, Pocatello, Jerome, Twin Falls, Nampa.Utah: Salt Lake City, American Fork, Saltair, Taylorsville, Murray, Holliday,Fort Douglas, Emigration Canyon, St. George.Nevada: State record.Arizona: Grand Canyon, Bright Angel, Williams, Flagstaff, Clemenceau.Remarks.?The Middle West representatives are clearly intermedi-ate between degener and the typical novemnotata. The variability of all the characters in which these subspecies differ from each other isvery high in this region. On the other hand, many individuals fromthe western part of the distribution of degener have some of thespots on the elytra very small or missing. The subspecies degeneris thus connected with the subspecies oregona Casey and franciscanaMulsant by a series of intergrades.COCCINELLA NOVEMNOTATA Herbst subspecies OREGONA CaseyCoccinella novemnotata Herbst subspecies oregona Casey, 1908, p. 403. ? Leng,1920, p. 216.This subspecies is very close to degener Casey. It differs from itby the larger size of the body (equal to that of the typical novem-notata Herbst), by very dense but obscure punctation, and by verysmall and frequently missing elytral spots. The surface of theelytra is more polished than in either novemnotata novemnotata ornovemnotata degener.Geographic distribution.?Localities as follows:British Columbia: Vernon, Midday Valley, Merritt, Nanaimo, Victoria, Depar-ture Bay.Washington: Fairfield, Pullman, Blue Mountains, Coulee City, Ritzville, Paha,Toppenish, Paradise Inn (Mount Rainier National Park), Olympia, Tenino.Oregon: Wallowa Mountains, La Grande, Portland, McMinnville, Coast Range(Benton County), Corvallis, Klamath County, Amity.California: Modoc County, Klamath Lake, Lassen County, CarrviUe, PlumasCounty, Mono County.Remarks.?This subspecies occupies the northern Pacific States.In California it finds the southern limit of its distribution, and isconnected by numerous intermediates with the more southern sub-species, nsimely franciscana Mulsant. 8 PROCEEDINGS OF THE NATIONAL MUSEUM vol.80COCCINELLA NOVEMNOTATA Herbst subspecies FRANCISCANA Mulsant Coccinella franciscana Mulsant, 1853, p. 19.Coccinella californica Casey (partim), 1899, p. 89; 1908, p. 404.Coccinella novemnotata Herbst var. franciscana Mulsant, Leng, 1903, p. 198;1920, p. 216.?Johnson, 1910, p. 59.This race differs from other subspecies of novemnotata Herbst bythe absence of all or most of the elytral spots, by the bright red of theelytra, and by the polished and sliiny surface of the elytra. Theindividuals from southern California are usually spotless; those fromSan Joaquin Valley usually have a few of the spots characteristic forother subspecies of novemnotata Herbst. The Coccinella living inSan Joaquin Valley may thus be considered intermediate betweenthe subspecies oregona Casey and franciscana Mulsant. Casey(loc. cit.) consideredfranciscana Mulsant as a synonym for Coccinellacalifornica Mannerheim. This view is unfounded. Even withoutconsideration of the structure of the genitalia, the presence of thewhite margin of the pronotum in franciscana Mulsant is evidenceagainst such a supposition.Geographic distribution.?Localities as follows:California: Klamath Springs, Carrville, Oroville, Yuba County, Lake Tahoe,Truckee, Placerville, Mokelumne Hill, Valley Springs, Tuolumne County,Patterson, Fresno, Coalinga, Huntington Lake (7,000 feet). Independence,Visalia, Lemoncove, Lindsay, Bakersfield, San Fernando, Mint Canyon,Mount Lowe, Mount Wilson, Pasadena, Los Angeles, Whittier, Arcadia,Monrovia, Fish Canyon, Riverside, Balboa, San Diego, Coronado, Poway,Imperial County.Utah: St. George.Arizona: Yuma.New Mexico: Espanola.Remarks.?The distribution oifranciscana Mulsant exhibits an inter-esting feature. This form is frequent in regions where Coccinellacalifornica Mannerheim is absent, and is lacking or occurs only seldomin places where C. californica is frequent. Practically the only regionwhere the distribution of the two species overlaps is the vicinityof Los Angeles. In general, californica occupies the region west of theCoast Range, while franciscana lives east of the Coast Range. Such arelationship seems typical for close species which are not far from beingonly subspecies of the same species. This is, however, hardly truein respect of franciscana and californica. Indeed, californica seemsto be related closely to transversoguttata Falderman and not to novem-notata Herbst. On the other hand, franciscana is beyond doubt asubspecies of novemnotata. Perhaps the explanation of this peculi-arity of distribution of the two species lies in some of their ecologicalpeculiarities. AKT. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKYCOCCINELLA PROLONGATA Crotch Coccinella prolongata Crotch, 1873, p. 371. ? Casey, 1899, p. 88. ? Johnson,1910, p. 64.Coccinella transversoguttata Falderman var. prolongata Crotch, Leng, 1903, p. 199.Coccinella monticola Mulsant var. prolongata Crotch, Leng, 1920, p. 216. Figures 10-12.?Male genitalia of Coccinella trifasciata, C. hierogtyphica, and C. undecimpunc-tata, respectively. The sipho and the trabes are not represented.FiGUEES 13-20.?Sipho of the different species of Coccinella: 13, C. novemnotata; 14, C. prolongata;15, C. californica; 16, C. difficilis; 17, C. hieroglyphica; 18, C. nivicola; 19, C. trifasciata; 20, C.undecimpunctata. sc, Siphonal capsuleMore elongately oval than C novemnotata Herbst, moderately con-vex, the sides of the elytra subparallel in the middle of their length,the convexity of the elytra very great in the posterior third of theirlength. Head black with very large white spots near the eyes, leav-es143?31 2 10 PROCEEDIN-QS OF THE NATIONAL MUSEUM vol.80ing only a narrow black bridge between them, Mesepimera andmetepimera, the posterior ends of the metepisterna, and, in the males,a spot on the anterior coxae, white. White markings in the anteriorangles of the pronotum are extended toward the posterior angles andtoward the middle of the disk, and, in most cases, are united by anarrow white stripe on the anterior margin of the pronotum. Ante-rior angles of the pronotal epipleura broadly white. Pronotum andthe elytra alutaceous, rather densely and very finely punctulate, withpunctures somewhat stronger than in novemnotata Herbst. Elytrayellow or orange with a large rhomboidal scutellar spot (K), a smallround spot (2), a large obliquo-transverse spot (3), and a transversespot in the apical third of then- length (4 + 5) , which sometimes isseparated into two spots (4 and 5). Length of the body, 6.2-7 mm.Male genitalia (figs. 3, 14).?Similar to those of C. novemnotataHerbst, but penis shorter, its sides angulate, the proximal endextended into a very broad, rhomboidal process. Basal plates widerthan long. Sipho very large, the siphonal capsule developed morestrongly than in any other species of Coccinella.Female genitalia.?Unknown.Geographic distribution.?Localities as follows:Montana: Helena (W. M. Mann, collector).Wyoming: Yellowstone National Park (W. Robinson, collector).Kansas: State record (National Museum collection).Colorado: Denver, Garland (National Museum collection).Washington: Wenatchee (E. J. Newcomer, collector).Remarks.?This species is rather closely related to C. novemnotataHerbst because of the structure of the genitalia, the punctation ofthe elytra, and the presence of the white spots on the anterior coxaein the males. I can not agree, therefore, with the opinion of Mr.Leng that prolongata Crotch is a variety of nivicola Menetries{monticola Mulsant). I find no characters indicating such a rela-tionship except the pattern of the elytra, which is indeed similarto C. nivicola Menetries var. aluiacea Casey. On the other hand,the genitalia of prolongata Crotch are quite suflSciently differentfrom those of novemnotata Herbst to consider them separate species.The area of habitation of prolongata is completely included in that ofnovemnotata.COCCINELLA PROLONGATA Crotch SEQUOIAE new subspeciesSimilar to Coccinella prolongata Crotch but with the frontal spotssmaller, with the quadrangular white spots in the anterior angles ofthe pronotum not dilated, elytra much more strongly punctate,intervals very finely alutaceous, reddish testaceous, elytral spots3 + 2 + 4 + 5 + 3 + K+3 + 5 + 4 + 2 + 3 confluent.This race differs from the typical form in a rather large series ofcharacters. The white markings on the pronotum are not extended ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 1 1 toward the posterior angles or toward the disk; the white stripe onanterior margin of the pronotiim is missing. The mesepimera andmetepimera are white, and the anterior coxae of the males have awhite spot, as in prolongata. The convexity of the elytra is moreregular than in prolongata, but less regular than in other species ofCoccinella. The elytral spots are strongly increased in size and con-fluent with one another. Spots 2, 3, 4, and 5 form aringlike pattern,which in most specimens is connected with spot ji. The elytralsuture is red, at least in the posterior half of its length. The malegenitalia are slightly different from those of the typical prolongata.The process on the distal end of the penis is somewhat longer andmore pointed at the end; the sides of the penis are more rounded.In spite of all these differences, I consider this form a subspecies ofprolongata rather than a separate species.Geographic distribution.?Localities as follows : California: Sequoia National Park, near Camp Wolverton (7,000 to 9,000 feetJune 24-25, 1929, 51 specimens, E. C. Van Dyke, collector; ibidem, June 29,1930, 2 specimens, T. Dobzhansky, collector; the type in the CaliforniaAcademy of Sciences collection).COCCINELLA PROLONGATA Crotch subspecies BRIDWELLI NunenmacherCoccinella hridicelli Nunenmacher, Ent. News, vol. 24, p, 76, 1913. ? Leng,1920, p. 216.This subspecies is different from the typical form and from thesubspecies sequoiae Dobzhansky by the smaller size and the less con-vex shape of the body, by the more strongly alutaceous surface, andthe color of elytra. Elytra entirely black. The wliite markings onthe head and the pronotum, and the punctation of the elytra, as insequoiae. Mesepimera white, metepimera usually white, but in somespecimens black (the type of bridwelli, according to Nunenmacher'sdescription, has black metepimera). Three out of the five malesstudied had white spots on the anterior coxae. The genitalia notdistinguishable from those of sequoiae. Length of body, 5.5-6.3 mm.This is one of the three entirely black forms known in the genusCoccinella. It is more similar to the subspecies sequoiae than to thetypical form. The subspecies sequoiae may be considered as anintermediate form between the subspecies bridwelli Nunenmacherand the typical prolongata Crotch.Geographic distribution.?LocaUties as follows:California: Tahquitz Valley (type and cotypes, J. C. Bridwell, collector), Tah-quitz Canyon (E. C, Van Dyke, collector), Idyllwild (E, C. Van Dyke,collector), COCCINELLA CALIFORNICA MannerheimCoccinella califarnica Mannerheim, Bull. Soc. Imp. Nat. Moscou, vol. 16, p. 312,1843.?Caset, 1899, pp. 88, 89.?Johnson, 1910, p. 62.Coccinella trasversoguttala Falderman var. californica Mannerheim, Leng, 1903,p. 200; 1920, p. 216. 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 80Body broadly oval, strongly convex. Head with a white spot oneach side near the eyes, pronotum with a quadrangular mark in theanterior angles, pronotal epipleura with a narrow white margin. Thevariety melanocollis Casey has the whole pronotum black. Mesepi-mera wliite, metepimera black. Pronotum and elytra slightlyalutaceous, densely but finely punctulate, the punctures becomingstronger toward the external margin of the elytra. Elytra brightred or orange with a black scutellar spot (]?) and with a narrow blackstripe along the suture. In some specimens the elytra are entirelyred (var. nevadica Casey ?). Very seldom individuals occur havingrudiments of the elytral spot 1, or the spot 3, or both. Length ofthe body, 5.4-6.5 mm. Figures 21-29.?Receptaculum seminis and infundibulum of the diflerent species of the genus Coc- cinella: 21, Coccinella novemnotata: 22, C. californica: 23, C. johnsoni; 24, C. transversoguttata; 25, C.nmcola: 26, C. difficiiis: 27, C. trifasciata: 28, C. hieroglyphica; 29, C. undecimpunctata. ap, Acces-sory plate; c, cornu; dr, ductus receptaculi; i, infundibulum; gr, accessory gland of the recepta-culum seminis; n, nodulus ; r, ramusMale genitalia (figs. 4, 15) .?Penis sUghtly longer than the paramera,broadening distally, deeply emarginated in the distal half of itslength, and extended into a very broad triangular process. Basalplates broader than long. Sipho shorter than in C. transversoguttataFalderman and in G. novemnotata Herbst.Female genitalia (fig. 22).?Receptaculum seminis similar to thatof C. novemnotata and C, transversoguttata. The infundibulum longerand more slender than in the species just mentioned.This species is rather closely related to C. transversoguttata. Thisrelationship is correctly recognized by Leng, but these species mustbe undoubtedly considered separate because of the difference in thestructure of the genitaUa, as well as in the external characters. Iplace Coccinella nevadica Casey as a synonym of californica Man-nerlieim with much hesitancy. I have not examined the Casey type ABT. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 13 specimen, and the description of it is unsatisfactory. C. nevadicamay also be a spotless variety of C. nivicola Menetries, or even maybe synonymous with C. novemnotata Herbst veLT.Jranciscana Mulsant.Geographic distribution.?Localities as follows : British Columbia: Victoria, Nanaimo, Departure Bay.Washington: Whatcom, Port Townsend, Seattle, Forks, Hoquiam.Oregon: Astoria, Cannon Beach, Tillamook, Otter Rocks, Agate Beach.California: Areata, Samoa, Eureka, Scotia, Fortuna, Orick, Klamath, Sisson(J. Bradley collector), Chilcoot (Essig collector), Mendocino, Guerneville,Santa Rosa, Petaluma, Fairfield, San Rafael, Mount Tamalpais, Sausalito,Cazadero, Berkeley, Oakland, Alameda, Sacramento County (Citrus Experi-ment Station collection) , Merced County (F.T. Scott collector) , San Francisco,San Mateo, Redwood City, Palo Alto, Santa Clara, San Jose, Los Gatos,Santa Cruz, Morgan Hill, Salinas, Spreckels, Del Monte, Monterey, PacificGrove, Carmel, Soledad, Pinnacles National Monument, King City, Lindsay(R. Jones collector) , Guadalupe, Betteravia, Los Alamos, Lompoc, Santa Ynez,Santa Barbara, Santa Paula, Oxnard, Santa Monica, Mint Canyon, Saugus,San Fernando, Pasadena, Mount Wilson, Mount Lowe, Los Angeles, Arcadia,Monrovia, Fish Canyon, San Gabriel Canyon, Pomona, Whittier, San Pedro,Santa Ana, Balboa, Laguna Beach, San Juan Capistrano, Fall Brook, LaJoUa, San Diego, El Cajon, Santa Rosa Island, Santa Cruz Island, San NicolasIsland (S. Emerson collector), San Clemente Island.Lower California: Ensenada, Descanso Bay, Guadalupe Island.Arizona: Senator (American Museum of Natural History collection).Remarks.?One may conclude from the data presented above thatC. californica is very common between the Coast Range and thePacific Ocean, but occurs only very seldom between the Coast Rangeand the Sierra Nevada. It is the commonest species of Coccinellain the Los Angeles region, but seems to be entirely absent in theMohave Desert. F. T. Scott informs me that C, californica is notfound in the central part of the San Joaquin Valley, being replacedthere by C. novemnotata Herbst var. franciscana Mulsant, which isnot very common in the Los Angeles region. The finding ofC. californica in Arizona and at Lindsay, Calif, (see above) seems tobe very doubtful. It is probable that this species will be found alsoalong the western coast of the northern part of Mexico.COCCINELLA JOHNSONI CaseyCoccinella johnsoni Casey, 1908, p. 403. ? Johnson, 1910, p. 61.Coccinella novemnotata Herbst. var. johnsoni Casey, Leng, 1920, p. 216.Body more elongate and less convex than in C. californica Man-nerheim. Punctulation of the pronotum and the elytra somewhatstronger than in the latter species. Elytra red with the 11 spots con-stituting the typical pattern of the genus Coccinella (fig. 30), and witha narrow black stripe along the suture. Spots 4 and 5 lie close toeach other, and are frequently confluent. The black sutural marginmaybe absent; likewise spot 2, or spot 4, or both, may be absent. Spot 14 PEOCEEDINGS OF THE NATIONAL MUSEUM vol.80 Yi may be fused with spot 1 to form a transverse black fascia. Lengthof the body, 5.8-6.8 mm.Male genitalia (fig. 5).?Resembling those of C. californica; but theprocess on the distal end of the penis shorter, broader, and less acu-minate at the end.Female genitalia (fig. 23).?Not distinguishable from C. californica.This species is very close to C. californica but not to C. novemno-tata, as suggested by Johnson and Leng. Its specific rank may be,however, a subject of dispute. Indeed, the very small differences inthe structure of the genitalia and in the form of the body might makeit questionable. No intermediates between the two species occur,however, in spite of the fact that the whole known distribution ofC. johnsoni is included in that of C. californica. Notwithstanding thesmallness of the differences between the two species, and the varia-bility of the elytral pattern of C. johnsoni, they may be distinguishedwithout difficulty. The species described by Casey (1908, p. 402)from northern Mexico, namely, Coccinella sonorica, is almost certainlya color variation of C. johnsoni.Geographic distribution.?Localities as follows:British Columbia: Victoria (California Academy of Sciences collection).Washington: Orcas Island (W. M. Mann collector), King County (F. W.Nunenmacher collection).California: Shively (E. O. Essig collector), San Francisco (F. T. Scott collector),Santa Paula, Pasadena, Fish Canyon, Santa Ana, Santa Monica, Costa Mesa,San Diego, Coronado, San Nicolas Island (S. Emerson collector), SanClemente Island (F. Blaisdell collector).Remarks.?The individuals from British Columbia and Washingtonare rather considerably different from those from California in beingless convex, in having the apex of the elytra acuminate, and in havingvery small elytral spots. This fact suggests that the species C.johnsoniis differentiated into two subspecies, one of which is living in BritishColumbia and Washington and the other living in California. Itseems to be wise, however, not to propose Latin names to these sub-species until more material is available,COCCINELLA TRANSVERSOGUTTATA Falderman Coccinella transversoguttata Falderman, Mem. Soc. Imp. Nat. Moscou, vol. 4, p.454, 1835.?Leng, 1903, p. 199; 1920, p. 216.?Johnson, 1910, pp. 61, 62.Coccinella quinquenolata Kirby, Richardson's Fauna Boreali-Americana, p. 230,1837.?Casey, 1899, p. 89; 1908, p. 401.Coccinella transversalis Mulsant, 1850, p. 117.Body broadly oval, very strongly convex. Head with a white spoton each side near the eyes, pronotum with white quadrangular marksin the anterior angles, pronotal epipleura narrowly white in the ante-rior angles. Mesepimera white, metepimera brown or black. Head,pronotum, and elytra densely and rather strongly punctulate, the ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 15punctures becoming stronger toward the external margin of the ely-tra. Elytra yellow or red with a common black subbasal fascia (spotsl + K+l), a very small spot near the external margin (2), which ismostly absent in the American specimens (var, guinquenotaia Kirby),a large transversely oval spot on the disk (3), and a subapical trans-verse fascia (spots 4 + 5) . The subbasal fascia and spot 3 are reducedin the western specimens to narrow black stripes (var. transversalisMulsant). Length of the body, 5.8-7.2 mm.Male genitalia (fig. 6).?Penis longer than the paramera, ratherwide, deeply emarginated in the distal half of its length, the distalend extended into a tonguelike process. Basal plates wider than long.Female genitalia (fig. 24) .?Receptaculum seminis similar to that ofC. novemnotata Herbst, but shorter, wider, and with slightly thinnerwalls. Infundibulum also similar to that of C. novemnotata, but rel-atively longer and more slender.Casey (1899) insisted on calling the American representatives ofthis species quinquenotata Kirby, and on restricting the name trans-versoguttata Falderman to the Asiatic representatives. I see no suffi-cient reason for such a separation, since the only difference betweenthe American and the Asiatic representatives of this species is the fre-quent absence of spot 2 in the former. The absence of this spot is,however, observed also in Asiatic specimens, though as an exception,and its presence is sometimes observed in American individuals. Thegenitaha are completely alike in the American and in the Asiaticspecimens (see Dobzhansky, 1926, fig. 1). Consequently, it seemssuperfluous to preserve the name quinquenotata Kirby even as a sub-specific name.Geographic distribution.?Localities as follows:Nova Scotia: Truro.Quebec: Montreal.Ontario: North Bay, Ottawa, Toronto, Port Credit, HuntsviUe, Ridgeway.Manitoba: Mile 214 on Hudson Bay Railroad, Winnipeg, Aweme.Mackenzie: Great Slave Lake (National Museum collection).Saskatchewan: Prince Albert, Carlyle.Alberta: Calgary, Banff.Alaska: New Rampart House (J. M. Jessup collector), Skagway (Harringtoncollector), Chitina Glacier (30 miles north of Mount St. Elias) (D. W. Eatoncollector).Yukon: Whitehorse (J. A. Kusche collector), Dawson (J. A. Kusche collector),White Pass (J. A. Kusche collector), Carcross (Harrington collector).British Columbia: Vernon, Penticton, Merritt, Fort McLeod, Vancouver.New Hampshire: Mount Washington, White Mountains, Lancaster, Franconia,Wolfeboro, Barnstead.Massachusetts: Arlington, Boston, Cambridge, Forest Hills, Winchester, Truro,Woods Hole.Connecticut: Bridgeport.New York: Thousand Islands, Whiteface Mountain, Batavia, Ithaca, Honeoye ^.Z_ Falls, West Point, Broadalbin. 16 PROCEEDINGS OE THE NATIONAL MUSEUM vol.80New Jersey: Riverton.Delaware: Wilmington.Pennsylvania: State record.Virginia: Fredericksburg.Michigan: Whitefish Point, Marquette, Alpena, Daggett, Douglas.Ohio: Lake County.Illinois: State record.Wisconsin: Waupaca, Winneconne.Minnesota: Two Harbors, Duluth, St. Louis County, Itasca County, BeltramiCounty, Marshal County, Norman County, Otter Lake, North Branch,Taylors Falls, Ramsey County, Anoka, St. Paul, Minneapolis, Lake City,Lake Crystal, New London, Luverne.Iowa: Muscatine, Crawford County.Missouri: Belgrade.North Dakota: Trail County, Binford, Fargo, Valley City, Drake, Oakdale,Bismarck, Mott.South Dakota: Black Hills.Texas: San Antonio, Colorado County.Montana: Moccasin.Wyoming: Big Horn Mountains, Cheyenne, Owl Creek Mountains, CannonCamp (Yellowstone National Park), Paint Creek, Carbon County, GreenRiver.Colorado: Greeley, Pingree Park, Longmont, Boulder, Longs Peak Inn (9,000feet). Golden, Denver, Grant, Rocky Ford, Fowler, Colorado Springs,Manitou, Canon City, Leadville (10,000 feet), Paonia, Grand Junction, VetaPass, Garland, San Luis, Dark Canyon.New Mexico: Chama, Aztec, Espanola, Las Vegas Hot Springs (7,000 feet),Chuska Mountains (8,800 feet).Idaho: Moscow, Lewiston, Blackfoot, Pocatello, Jerome, Twin Falls, Boise.Utah: Logan, Promontory, Huntsville, Kaysville, Coalville, Salt Lake City,Taylorsville, Kamas, Park City, Heber, Silverlake, Alta, Murray, Sandy,Lehi, West Jordan, Prove, Emigration Canyon, Fort Douglas, Price, BeaverMountains (8,000-10,000 feet), St. George.Arizona: Williams, Chiricahui Mountains.Washington: Pullman, Ritzville, Paha, Coulee, Wenatchee, Toppenish, Seattle,Tacoma, Paradise Park (Mount Rainier, 6,000 feet).Oregon: Baker, Cascade Rapids, Portland.Nevada: Shell Canon (Ruby Mountains), Reno, Steamboat Springs, LyonCounty, Esmeralda County.California: Modoc County, Madeline, Truckee, Lake Tahoe, Mono County,Ryan (Dobzhansky collector).This form is apparently lacking in southern California.COCCINELLA TRANSVERSOGUTTATA Falderman subspecies NUGATORIA MulsantCocdnella nugatoria Mulsant, 1850, p. 1021. ? Casey 1908, p. 403.Coccinella transversoguttata Falderman var. nugatoria Mulsant; Lenq, 1903, p.199; 1920, p. 216.?Johnson, 1910, p. 61.This is a western subspecies of C. transversoguttata Falderman, dif-fering from the typical form by smaller size, less convex elytra, finerpunctation, and by the reduction of the pigmentation of the elytra.Spots K and 1 are separate, spot K is transversely oval or pyriform,spot 1 is round and small, spot 2 usually absent, spot 3 is smaller ABT. 4 BEETLES OF THE GENUS COCCIKELLA DOBZHAITSKY 17 and more narrowly oval than in transversoguttata, spots 4 and 5 fre-quently separate. Spot 1, or spot 4, or both, may be missing. Lengthof the body, 5.8-6.5 mm.Casey (1899) insisted on granting a specific rank to C. nugatoriaMulsant. In fact it is only a rather indistinct subspecies of C. tranS'versoguttata Falderman living in the Western States and in Mexico.An enormous area extending from North Dakota and New Mexicoto the Pacific Ocean is inhabited by a mixed population in which allthe intermediates between the typical transversoguttata Faldermanand the typical nugatoria Mulsant may occur. Only in Mexico is thewhole population nugatoria. The genitalia of transversoguttata (fromMassachusetts) and of nugatoria (from California) are alike.Geographic distribution.?Localities as follows:Minnesota: Two Harbors, New London, Lake City.North Dakota: Binford, Bismarck.Wyoming: Clieyenne, Como, Carbon County.Colorado: Pingree Park, Denver, Manitou, Leadville (10,000 feet), Vega.New Mexico: Chama.Idaho: Lewiston, Jerome, Boise, Nampa.Utah: Ogden Canyon, Emery County, Taylorsville, Salt Lake City, MurrayPark City.British Columbia:. Vernon, Penticton, Merritt, Vancouver.Washington: Pullman, Toppenish, Tacoma, Wapato.Oregon: Cascade Rapids.Nevada: Carson City, Esmeralda County, Nixon.California: Modoc County, Madeline, Nevada County, Big Pine.Mexico: Queretaro, Mexico City, Guadalajara.COCCINELLA NIVICOLA Menelries subspecies MONTICOLA Mulsant Coccinella rnonticola Mulsant, 1850, p. 115. ? Casey, 1899, p. 89. ? Leng, 1903,p. 198; 1920, p. 216.?Johnson, 1910, p. 63.Coccinella lacustris LeConte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 131,1852.Coccinella impressa Casey, 1899, p. 89; 1908, p. 402.Broadly oval, strongly convex. Head with white spots near theeyes, pronotum with white quadrangular spots in anterior angles,pronotal epipleura with quadrangular marks or with only a whitestripe in the anterior angles. Mesepimera white, metepimera brownor black. The sides of the pronotum frequently possess one or severalimpressions, which are variable in extent but which may be altogetherlacking. Individuals with very strongly developed impressions on thepronotum were described by Casey (1899) as a separate species, Cocci-nella impressa. In a more recent paper, Casey (1908, p. 402) treatedimpressa as a subspecies of rnonticola Mulsant. The character is, how-ever, only an individual variation, and in my opinion the nameimpressa must be treated simply as a synonym of monticola. Elytra 18 PROCEEDINGS OP THE NATIONAL MUSEUM vol. 80 orange or yellow, alutaceous, punctation dense and strong, becom-ing stronger toward the external margin, with a very large pyriformscutellar spot (}0, an oblique transverse fascia not attaining thesuture or the external margin (spots 2 + 3), and with a large, trans-versally oval spot in the apical third (spots 4 + 5). Length of thebody, 5.8-7 mm.Male genitalia (figs. 7, 18).?Penis considerably longer than the para-mera, slightly wider at the middle of its length than at the base, rapidlynarrowing distally, the distal end extended into rather long fingerlikeprocess. Basal plates longer than wide.Female genitalia (fig. 25).?Receptaculum seminis short and wide,with ringlike sculpture on its walls rather delicate. Infundibulumshort and thick, its posterior end with a funnel-shaped dilatation, thediameter of which is nearly twice as much as the diameter of theanterior dilatation of the infundibulum. Accessory plate absent.I find C. monticola Mulsant not specifically distinct from the Asiaticspecies Coccinella nivicola Menetries, The typical nivicola (fromeastern Siberia) differs from monticola in having the anterior fasciaextended toward the humeral angles and in having a stronger punctu-lation of the elytra. Both characters are, however, variable. Indi-viduals of monticola from Canada and the Northern United Stateshave the elytral fascia very broad, and frequently extended towardthe humeral angles. On the other hand, individuals of nivicola fromsouthern Siberia, Mongolia, and Djungaria have the elytra! patternapproaching that of monticola. Even more important is the fact thatthe genitalia of nivicola (see Dobzhansky, 1926, fig. 6) are identicalwith those of monticola. Thus, monticola must be considered a sub-specific form of nivicola, equivalent to the subspecies alutacea Casey.Geographic distribution.?Localities as follows:Quebec: Quebec, Hudson Bay (University of Minnesota collection).New Hampshire: Mount Washington, Lancaster, Franconia.Michigan: Whitefish Point.Minnesota: Duluth, Itasca Park.Montana: Glacier National Park (T. Ulke collector), Missoula (NationalMuseum collection).Utah: Brighton (National Museum collection).Colorado: Rogers Pass (Bradley collector).Florida: State record (Illinois State Natural History Survey collection; thisrecord is very doubtful).California: Lassen National Park (8,000-10,000 feet, Dobzhansky collector),Tallac (A. Feneys collector). Fallen Leaf (California Academy of Sciencescollection), Eldorado County (Blaisdell collector), Mono (Blaisdell collector):Remarks.?The individuals from Montana, Colorado, and Utahare intermediate between monticola and alutacea. Those from Cali-fornia have the elytral pattern similar to that of monticola from NewEngland, but one individual has the anterior fascia connected withihe scutellar spot. ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 19COCCINELLA NIVICOLA Menetries subspecies ALUTACEA Casey Coccinella alutacea Casey, 1899, p. 89.Coccinella transversoguttata Falderman var. alutacea Casey, Lbng, 1903, p. 200.Coccinnella monticola Mulsant var. alutacea Casey, Johnson, 1910, p. 63. ? Leng,1920, p. 216.Differs from the typical nivicola Menetries and from var. monticolaMulsant by the more strongly alutaceous surface of the elytra; thefiner and sparser punctulation, and by the reduced pigmentation ofthe elytra. The shape of the body is less convex in alutacea than inmonticola. The scutellar spot is rather small, the black fascia nar-row, frequently disintegrating into a small round spot 2, and a largetransversally oval spot 3. The apico-marginal spot (4 + 5) small,round or transversally oval. Spot 2, or spot 4 + 5, or both, may dis-appear (var. biguttata Johnson). Metepimera and the posterior endsof the metepisterna usually brown or yellow, though specimens hav-ing black metepimera also occur. The genitalia of both sexes are iden-tical with those of typical nivicola and of subspecies monticola. Lengand Johnson were correct in not recognizing this variety as a separatespecies.Geographic distribution.?Locahties as follows:Kansas: State record (Illinois State Natural History Survey collection).South Dakota: Ardmore.Montana: Assiniboine, Missoula, Yellowstone County.Wyoming: Newcastle, Big Horn Mountains, Yellowstone National Park, GrandTeton National Park, Cheyenne, Green River (6,000-7,000 feet).Colorado: Pingree Park, Fort Collins, Longniont, Boulder, Denver, Golden,Cimarron, Garden of the Gods, Manitou, Colorado Springs, Rocky Ford,La Veta, Leadville, Buena Vista, Paonia, Saguache.New Mexico: Chama, Espanola, Santa Fe, Las Vegas Hot Springs, Elida,Albuquerque.Idaho: Snake Canyon, Blackfoot, Burley.Utah: Uinta National Forest, Alta, Salt Lake City, Silverlake, Murray, Heber,Logan, Emery County, Beaver Creek Hills, Iron County, St. George.Arizona: San Francisco Mountains (7,900 feet) (Cornell University collection).Nevada: Elko, Ely, Lovelock, Ormsby County, Nixon, Lamoille.British Columbia: Fort McLeod, Vancouver, Victoria.Washington: Sprague Lake, Toppenish, Paradise Park (Mount Rainier, 6,000feet).Oregon: Vale, Huntington, Lake County, Dallas.California: Davis, Modoc County (California Academy of Sciences collection),Siskiyou County, Klamath (F. W. Nunenmacher collection), Lassen County(F. W. Nunenmacher collection), Plumas County (F. W. Nunenmacher collec-tion), Madeline, Piedmont (F. W. Nunenmacher collector), Truckee (Califor-nia Academy of Sciences collection), Bullfrog Lake (10,600 feet) (CaliforniaAcademy of Sciences collection).Remarks.?Individuals from western Utah, Nevada, and Californiaare characterized by an extreme reduction of the pigmentation ofthe elytra. Indeed, most of them have only spot 3 (var. biguttataJohnson), or only spot 5. Besides this they are usually smaller, less 20 PEOCEEDINGS OF THE NATIONAL MUSEUM vol. 80 convex, and more elongate-oval than the typical alutacea from moreeastern localities. Perhaps this race deserves being called a separatesubspecies, but more material is needed before its status can beestablished definitely. It must be noted that this very light race is.recorded in the Sierra Nevada in nearly the same localities in whichmonticola Mulsant (see above) is recorded. In spite of this no inter-grades between these very different races are so far observed. Thegenitalia of the Californian alutacea were studied and found to benot distinguishable from other specimens of alutacea or frommonticola. The whole question certainly deserves further study.COCCINELLA DIFFICILIS Crotch Coccinella difficilis Crotch, 1873, p. 370. ? Leng, 1903, p. 200. ? Johnson 1910,p. 64.Coccinella monticola Mulsant var. difficilis Crotch, Leng, 1920, p. 216.Coccinella vandykei Nunenmacher, Ent. News, vol. 20, p. 161, 1909.Coccinella iransversoguttata Faldernian var. vandykei Nunenmacher, Leng, 1920,p. 216.Broadly oval, rather strongly convex. Head with white spots nearthe eyes, labrum brown, the anterior angles of the pronotum withquadrangular white marks, the anterior angles of the pronotalepipleura with a narrow white margin. Pronotum and elytra non-alutaceous, densely and finely punctulate. Elytra orange or red witha large scutellar spot {%), a small, round, frequently missing spot 2,a large, transversally elliptical spot 3, and a very large transverseapico-marginal spot (4 + 5), tending to break into two separate spots.Mesepimera and metepimera white. Length of the body, 5.4-6.3 mm.Male genitalia (figs. 9, 16).?Penis only slightly longer than theparamera, gradually widening from the base in the distal direction,sharply narrowed at two-thirds of its length, and extended into afingerlike process. Basal plates longer than wide. Sipho very longas compared with the size of the penis. Siphonal capsule angulate.Female genitalia (fig. 26).?Receptaculm seminis short and wide;ramus rudimentary. Infundibulm similar to that of Coccinellanivicola Menetries.This species is surprisingly close to Coccinella tian^hanica Dob-zhansky from middle Asia. The external structures as well as thegenitalia are similar, though not identical, in the two species. Theydiffer in the sculpture of the elytra, in the form of spot 4 + 5, in the-form of the penis, and that of the receptaculum seminis. Among theAmerican species of Coccinella the only close relative of difficilis is C.suturalis Casey. The two species just mentioned, and the Asiaticspecies tianshanica Dobzhansky and iranica Dobzhansky, form a sep-arate section of the genus Coccinella. C. nivicola Menetries belongs, ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 21however, to the group of species headed by the palaearctic species C.sejpteraj)unctata Linnaeus. Though these two sections of the genus arerelated to each other, I can not agree with Leng, who places bothdifficilis Crotch and suturalis Casey as varieties of nivicola Menetries.Coccinella vandyTcei Nunenmacher is a synonym of difficilis Crotch.Geographic distribution.?Localities as follows:Montana: Helena (National Museum collection), Gallatin County (NationalMuseum collection).Wyoming: Cheyenne, Green River.Colorado: Colorado Springs (National Museum collection).Idaho: Bingham County, Pocatello, Jerome, Boise.Utah: Salt Lake City, Beaver Valley, South Creek, Beaver County.Nevada: Elko (H. F. Wickham collector), Lovelock (F. E. Blaisdell collector),Goldfield (Nunenmacher collector, type and two cotypes of C. vandykeiNunenmacher).California: Chilcoot (Essig collector).COCCINELLA SUTURALIS Casey Coccinella suturalis Caset, 1899, p. 89. ? Johnson 1910, p. 64.Coccinella monticola Mulsant var. suturalis Casey, Leng, 1903, p. 198; 1920, p. 216.Elongate-oval, less convex than any other American species of Coc-cinella. Head and pronotum similar to C. difficilis Casey. Anteriorangles of the pronotal epipleura with quadrangular white marks infemales, and only narrowly white in males. Mesepimera white, met-epimera black. Punctulation of the pronotum and of the elytra denseand strong, becoming much stronger toward the external margin ofthe elytra and toward the sides of the pronotum. Elytra orange,with a large, obcordiform scutellar spot (}0, a rather broad black stripealong the suture, an oblique black fascia on the disk (spots 2 + 3), anda tranversal black spot in the posterior third of the length (spots 4 + 5) . Length of the body, 5.5-5.8 mm.Male genitalia (fig. 8) .?Penis similar to that of C. difficilis Crotch,but considerably longer than the paramera, the distal process muchlonger and less pointed at the end. Basal plates wider than long.Siphonal capsule not angulate.Female genitalia.?Unknown.Coccinella suturalis Casey is related to diflcilis Crotch, but differsfrom it in the shape of the body, in the punctulation and the patternof the elytra, and in the structure of the genitalia.Geographic distribution.?Localities as follows : Utah: Alta (Hubbard and Schwarz collection).California (Fresno County): Mount Kaiser (10,000 feet) (California Academyof Sciences collection), Mount Gould (12,000 feet) (California Academy ofSciences collection), Bubbs Creek (9,700 feet) (California Academy ofSciences collection). 22 PEOCEEDINGS OF THE NATIONAL MUSEUM vol.80COCCINELLA TRIFASCIATA LinnaeusCoccinella trijasciata Linnaeus, Systema naturae, p. 365, 1758. ? Leng, 1903, p.200.Coccinella perplexa Mulsant, 1850, p. 1021. ? Casey, 1899, p. 89. ? Johnson,1910, p. 57.?Leng, 1920, p. 216.Broadly oval, strongly convex. Head in females with white spotsnear the eyes, in males with a broad white stripe across the front.Pronotum with triangular or quadrangular white marks in the ante-rior angles, and with white anterior margin, pronotal epipleura withquadrangular white marks in the anterior angles. Mesepimera andmetepimera, and usually also the posterior ends of the metepisterna,white. Pronotum densely and finely punctulate, punctures on theelytra nearly as dense as those on the pronotum but considerablystronger. Elytra yellow or orange with three transverse black fasciae ; the anterior fascia is common to both elytra (it is the result of thefusion of the spots 1 + K+l). The median fascia (spots 2 + 3), andthe posterior fascia (spots 4 + 5) do not reach the suture or the<5xternal margin. Length of the body, 4.5-5.5 mm.Male genitalia (figs. 10, 19).?Penis short and wide, pyriform inshape, its distal end extended into a short process. Paramera onlyslightly, if at all, shorter than penis. Basal plates not wider thanlong. Sipho very short and thick, the siphonal capsule rather smalland rounded.Female genitalia (fig. 27).?Receptaculum seminis long and slender,cornu almost pointed at the end, ramus wide, infundibulum notdilatated at the posterior end.The American representatives of this species were described underthe name perplexa Mulsant. The identity of perplexa Mulsant withthe palaearctic species trifasciata Linnaeus was the subject of a disputebetween Casey and Leng (see Casey, 1908). I find myself in a com-plete agreement with Leng, who regards perplexa as a mere synonymof trifasciata. The only difference between the American and theEurasiatic specimens of this species is the presence of the whitemargin of the pronotum in the former. Johnson (1910) found anindividual of perplexa having no white margin on the pronotum.Furthermore, this character is sometimes found also in Asiatic speci-mens (Dobzhansky, 1926). The genitalia of the American and theAsiatic specimens are quite alike (cf. Dobzhansky, 1926, fig. 10). Itseems that perplexa can not be considered even as a subspecificallydistinct form, and therefore the name should be dropped.Geographic distribution.?Localities as follows:Nova Scotia: Truro.New Brunswick: Hampton.Quebec: St. Hilaire, Lavaltrie, Fort Coulonge.Ontario: Ottavva, Huntsville, North Bay, Toronto, Chatham, Hudson Bay(University of Minnesota collection). ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 23 Manitoba: Aweme, Mile 17 on Hudson Bay Railroad.Alberta: Banff.Alaska: Mountain Sheep Creek (22 miles below Eagle) (Harrington collector).Yukon: Dawson (J. M. Jessup collector), Whitehorse (J. A. Kusche collector).British Columbia: Glacier, Agassiz, Vernon, Victoria.Maine: Orono, Boothbay, Casco Bay.New Hampshire: Mount Washington, Lancaster, Franconia, Center Harbor,Farniington, Barnstead, Nashua.Massachusetts: Boston, Arlington, Melrose Highlands, Princeton, Cambridge,Stoughton, West Medford, Middleboro, North Saugus, Falmouth, W^oodsHole, Oak Bluffs.Rhode Island: Watch Hill.Connecticut: South Windsor, New Haven, Cromwell, Brookfield.New York: Adirondack Mountains, Mount Whiteface, Mount Mclntire, New-comb, Wilmington, Lake Placid, Middletown, West Point, Larchmont, PelhamBay, New York, Staten Island, Ithaca.New Jersey: Fort Lee, Passaic, Ramsey, Great Notch, Newark, Westwood>Paterson, Chester, Boonton.Pennsylvania: State record.Maryland: State record.Michigan: Whitefish Point, Alpena, Detroit.Wisconsin: Madison.Minnesota: Grand Marais, Two Harbors, Duluth, Itasca County, Roseau County|Minneapolis, St. Paul, Lake City, Beaver Dam, Lake Crystal, St. Antony.North Dakota: State record.Montana: Bear Paw Mountains, Columbia Falls.Wyoming: Yellowstone National Park.Colorado: Larimer County, La Veta.New Mexico: Maxwell (National Museum collection).Texas: State record (National Museum collection).Idaho: Moscow, Ashton.Utah: Logan (California Academy of Sciences collection).Washington: Pullman.California: Eureka, Klamath (Dobzhansky collector).Remarks.?The individuals from Montana, Wyoming, Colorado,New Mexico, Idaho, and Utah are similar to those from the North-eastern States. A few individuals from Washington, Oregon, andCalifornia are probably extreme variants of the subspecies subversaLeConte.COCCINELLA TRIFASCIATA Linnaeus subspecies SUBVERSA LeConteCoccinella subversa LeConte, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 19,1854.?Casey, 1899, p. 89.Coccinella perplexa Mulsant var. subversa LeConte, Johnson, 1910, p. 57. ? Leng, 1920, p. 216.Differs from the typical trifasciata by the distinctly smaller size ofthe body and by a strong reduction of the black pigmentation of theelytra. Elytra spotless, or with only the scutellar spot (^) , or withonly a very small discal spot (3), or with both the scutellar and thediscal spots. In any case, the size of the scutellar and the discalspots is very much decreased as compared with typical trifasciata. 24 PEOCEEDIFGS OF THE NATIONAL MUSEUM vol. 80 the pigment is often brownish instead of black, and the outlines ofthe spots are frequently diffuse and indistinct. Length of the body,4.3-5 mm.According to Casey (1899, 1908) this is a species separate fromtrifasciafa {perplexa). The opinion of Leng and Johnson seems to be,however, better founded. An investigation of the genitalia of a largeseries of specimens of C. subversa from Oregon failed to reveal anydifference between it and the typical C. trifasciata from the EasternStates, except, perhaps, an insignificant difference in the absolute size. Thus, C. subversa may be classified as a good example of asubspecific form.Geographic distribution. Localities as follows:British Columbia: Agassiz, Vancouver, Wellington, Victoria.Washington: Orcas Island, Mount Vernon, Everett, Port Angeles, Seattle,Tacoma, Puyallup, North Bend, Easton, Paradise Park (Mount Rainier),Longmire (Mount Rainier), Tenino, Oakville, Vancouver.Oregon: Astoria, Agate Beach, Neskowin, Olney, Portland, Hillsboro, ForestGrove, Cannon Beach, Tillamook. McMinnville, Siuslaw National Forest,Wilsonville, Corvallis, Waldport, North Bend, Josephine County.California: Scotia, Fortuna, Orick, Klamath, Hydesville, Sonoma County,Berkeley, Oakland.Remarks.?As shown by the list of localities, the subspecies isabundant in Washington and Oregon, in the whole region betweenthe Cascade Range and the Pacific Ocean. Its distribution extendsto British Columbia, where it overlaps the distribution of the typi-cal C. trifasciata. In the south the distribution of subversa overlapsthat of the subspecies eugenii Mulsant and Juliana Mulsant. Itmust be noted, however, that the individuals of subversa from Cali-fornia have the form of the body and the punctation of the elytramore similar to that of Juliana than to that of subversa from Oregonand Washington. The parts of Oregon and Washington lying eastof the Cascade Range are apparently inhabited by typical trifasciata.Thus the Cascade Range appears to be the line dividing the twosubspecies.COCCINELLA TRIFASCIATA, Linnaeus subspecies EUGENII Mulsant Coccinella eugenii Mulsant, 1866, p. 95.Coccinella trifasciata Linnaeus var. eugenii Mulsant, Leng, 1903, p. 200.Coccinella perplexa Mulsant var. eugenii Mulsant, Johnson, 1910, p. 57. ? Leng,1920, p. 216.This is a geographical race intermediate between the subspeciessubversa LeConte and the subspecies Juliana Mulsant. The size andthe punctation of the elytra are as in subversa. The pattern of theelytra may consist of the three transverse fasciae similar to those ofthe typical trifasciata but much narrower and showing a clear tend-ency to disintegrate into separate spots. More frequently the inter-mediate fascia is broken into a small round spot 2 and an oblong, ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 25 transverse spot 3. The subapical fascia may also disintegrate intotwo small, usually round spots (4 and 5). Equally frequently theintermediate and the subapical fasciae remain intact, but the hum-eral fascia resolves itself into a round humeral spot (1) and a trans-verse scutellar spot (K). Finally, all the fasciae may disintegrate,resulting in a pattern similar to the typical pattern of the genusCoccinella (fig. 30) . Some spots, especially 2, 4, and 1, may be absent.Geographic distribution.?Localities as follows:Oregon: Cannon Beach, Tillamook, Colestin.California: Modoc County, Mount Shasta, Shively, Slsson, Cayton, Areata,Eureka, Scotia, Fortuna, Orick, Klamath, Hydesville, Mendocino, Guerne- ville, Lagunitas, Fairfax, Albany, Alameda, Piedmont, San Francisco, LosGatos, Napa County, Plumas County, Tallac, Tahoe, Half Moon.Remarks.?The distribution of this subspecies overlaps that of thesubspecies Juliana Mulsant. The subspecies eugenii Mulsant is,however, more frequent in the northern part of California than inthe San Francisco region. The reverse holds true ior Juliana.COCCINELLA TRIFASCIATA Linnaeus subspecies JULIANA Mulsant Coccinella Juliana Mulsant, 1856, p. 135. ? Casey, 1899, p. 89.Coccinella barda LeConte, 1859, p. 286.Coccinella trifasciata Linnaeus var. Juliana Mulsant, Leng, 1903, p. 200.Coccinella perplexa Mulsant var. Juliana Mulsant, Johnson, 1910, p. 57. ? Leng, 1920, p. 216.More oblong but not less convex than other varieties of C. trifas-ciata. Punctation of the pronotum and of the elytra finer andsparser than in typical trifasciata; elytra shining. Pigmentation ofthe elytra strongly reduced. Usually only the humeral fascia ispresent, the two other fasciae being absent completely. The humeralfascia may resolve itself into separate spots (K and 1) . Small blotchesof dark pigment may be present on the places occupied in othervarieties by the spots 3, 4, and 5. Length of the body, 4.6-5.3 mm.The genitalia of the subspecies Juliana are not different from those ofthe typical trifasciata or from those of the subspecies subversa LeConte.This fact, as well as the absence of marked external differences,indicates that Juliana is a geographical race of trifasciata and not aseparate species. The geographic distribution oi Juliana is in accordwith this view.Geographic distribution.?Localities as follows ; California; Areata, Hydesville, Scotia, Fortuna, Orick, Klamath, Dyerville,Shively, Casadero, Felton, Muir Woods, San Anselmo, Point Reyes,Fairfax, Sausalito, Muir Woods, Tiburon, Lagunitas, Mayfield, NapaCounty, Fieldbrook, Berkeley, Alameda, Piedmont, Contra Costa County,San Francisco, Millbrae, San Mateo, Mountain View, Crystal Lake, SanJose, Palo Alto, Los Gatos, Santa Cruz, Santa Cruz Mountains, Monterey,Carmel, Point Sal (T. Dobzhansky collector), Santa Paula, (P. H. Timber-lake collection). 26 PEOCiEEDINGS OF THE NATIONAL MUSEUM vol.80 Remarks.?This subspecies is very common in the San Franciscoregion. To the north it gives numerous intergradations connectingit with the subspecies eugenii Mulsant (see above).COCCINELLA HIEROGLYPHICA Linnaeus subspecies TRICUSPIS KirbyCoccinella tricuspis Kirby, Richardson's Fauna Boreali-Americana, p. 231,1837.?Casey, 1899, p. 90.?Leng, 1903, p. 201; 1920, p. 216.?Johnson,1910, p. 59.Body oval, more elongate than in other species of the genus Cocci-nella, strongly convex. Head with white elongate spots near the eyes,pronotum with quadrangular white marks in the anterior angles, theanterior margin of the pronotum usually white, pronotal epipleurawith white quadrangular marks in the anterior angles. Mesepimeraand metepimera black, mesepimera brownish only in fresh specimens.Pronotum densely and finely, elytra densely and strongly punctu-late, elytra orange, with a common undulate subbasal fascia (spots1+2 + 3 + ^+ 3 + 2 + 1), and with another transverse black fascia inthe posterior half of their length (spots 4 + 5), not continuous acrossthe suture. Length of the body, 3.7-4.3 mm.Male genitalia (figs. 11, 17).?Penis short, sugar-loaf shaped, with-out processes or emarginations. Paramera short and thick. Basalplates wider than long. Sipho short and thick, siphonal capsulerelatively very large.Female genitalia (fig. 28).?Receptaculum seminis slender, cornupointed at the end, ramus strongly developed. Infundibulum notdilatated at its posterior end.I have shown (Dobzhansky, 1926) that the Siberian Coccinella tri-cuspis Kirby subsp. mannerheimi Mulsant is not specifically differentfrom theEuropean Coccinella hieroglyphica Linnaeus. This conclusionmay be extended also to the American tricuspis. The genitalia of C.tricuspis Kirby (figs. 11, 17, 28) are completely like those of C. hiero-glyphica (see Dobzhansky, 1926, fig. 4), and the external differencesbetween these forms, involving the color pattern and the shape of thebody, are by no means more extreme than those that are to beexpected between geographic races. Thus, the species C. hieroglyphicainhabits the entire north of the holarctic region. The subspecies C.hieroglyphica lives in Europe and in western Siberia ; C. mannerheimioccupies eastern Siberia; C. tricuspis occurs in Canada and north-ern United States; and C. humholdtiensis Nunenmacher inhabitsthe Pacific coast area of North America.Geographic distribution.?Localities as follows:Guebec: Quebec, Quinze Lake, Montreal.Ontario: Lake Superior.Manitoba: Aweme, Mile 17 on Hudson Bay Railroad.Saskatchewan: Lebret. ART. 4 BEETLES OP THE GENUS COCCIN-ELLA DOBZHANSKY 27 Alberta: Province record.New Hampshire: Lancaster, Mount Washington, Barnstead.Michigan: Whitefish Point.Minnesota: Two Harbors, Winton, Itasca Park, Minneapolis.Montana: Bear Paw Mountains (National Museum collection).COCCINELLA HIEROGLYPHICA Linnaeus subspecies HUMBOLDTIENSIS Nunenmacher Coccinella humboldtiensis Nunenmacher, Ent. News, vol. 23, p. 448, 1912. ? Leng, 1920, p. 216.Differs from Coccinella hieroglyphica Linnaeus subspecies tricuspisKirby by less elongate form of the body, finer punctulation, and byreduction of the black pigmentation of the elytra. Elytra orangewith a rhomboidal black spot ji and without otherspots, or with spots ji, 2, 3, and 4 + 5, or with K,4, and 5, or 4 + 5, or with two transverse fasciae(2 + 3 and 4 + 5), and the scutellar spot. Lengthof the body, 3.6-4.2 mm.The genitalia of C. humboldtiensis (one male andone female were examined) are identical with thoseof C. hieroglyphica tricuspis. Nunenmacher (loc. cit.) correctly pointed out the closeness of humbold-tiensis to tricuspis, but, in my opinion, overestimat-ed the value of the differences discovered betweenthem. Surprisingly enough, humboldtiensis pos-sesses some similarities to the European hieroglyph-ica, which the geographically closer subspeciestricuspis does not possess. The similarities are theform of the body and the shape of the elytralmarkings.Geographic distribution.?Localities as follows: Figure 30.?Right ely-tron of Coccinellaundecimpunctata,representing the spotsconstituting the typi.cal pattern of the genusCoccinella. H? Spot}A, or the scutellarspot; 1, the first, orhumeral, spot; 2, thesecond, or lateral, spot;3, the third, or discal,spot; 4, the fourth, ormarginal, spot; 5, thefifth, or apical, spotOregon: State record (Koebele collector).California: Crescent City (F. W. Nunenmacher collector, type and cotypes),Plumas County (F. W. Nunenmacher collector), Siskiyou County (Koebelecollector). COCCINELLA UNDECIMPUNCTATA Linnaeus Coccinella undecimpunctata Linnaeus, Systema naturae, p. 366, 1758. ? Leng,1920, p. 216.Oval, not strongly convex. Head with white spots near the eyes,pronotum and the pronotal epipleura with quadrangular or triangu-lar white markings in the anterior angles. Mesepimera and metepi-mera white. Pronotum and elytra very densely and rather finelypunctulate, elytra yellow or brick red with the 11 typical spots(fig. 30). Spots 2 + 3 and 4+5 fuse together forming two transversefasciae (var. boreolittoralis Donisthorpe) . Length of the body,5-5.5 mm. 28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 80Male genitalia (figs. 12, 20).?This species may be recognized atonce by the rudimentary, little chitinized siphonal capsule, and bythe presence of a wide dilatation of the sipho just distal to the siphonalcapsule. This dilatation is provided with an opening on the con-cave side of the sipho. Penis much longer than the paramera, basalplates much longer than wide, extended posteriorly into two diver-gent processes.Female genitalia (fig. 29).?Receptaculum seminis long and slender,cornu pointed at the end, the ringlike sculptor rudimentary. Infun-dibulum short and provided with wide funnel-shaped dilatations atboth ends.Geographic distribution.?Localities as follows:Massachusetts: Stoneham (F. A. Sherriff collector), Nahant (F. E. Blaisdellcollector), Falmouth Heights (T. Dobzhansky collector).Alaska: 4 miles north of New Rampart House (J. M. Jessup collector), 60-75miles north of Rampart House (J. M. Jessup collector).Remarks.?This species lives only along the sea coasts of Europe,on saline soils of Middle and Central Asia, and along the coast of thePolar Ocean from Greenland to the mouth of Yenisei. The findingof this species on the Massachusetts coast may be explained by twodifferent hypotheses. First, it may have been introduced there fromEurope. Second, it may be native there. In the latter case one mayexpect that it will be found also along the coasts of Nova Scotia,Newfoundland, and Labrador. The individuals from Massachusettsare quite similar to those from England and the northern coasts ofEurope. The two individuals from Alaska are different from thosefrom Massachusetts in appearance. They are larger, less convex,and have a stronger punctation. This seems to be a race the distri-bution of which is, as far as known, restricted to Alaska. It seemsprobable that C. undecimpunctata will be found also along the north-ern coast of eastern Siberia.GEOGRAPHIC DISTRIBUTION OF SPECIES OF THE GENUSCOCCINELLAAs shown above, the nearctic region is inhabited by 1 1 species ofthe genus Coccinella. In the palaearctic region there are at least 15species of that genus. Six of the nearctic species are represented inthis region by more than one subspecies. Furthermore, 5 species, asfollows, live in both nearctic and in palaearctic regions: transversa'guttata Falderman, nivicola Menetries, trifasciata Linnaeus, hiero-glyphica Linnaeus, and undecimpunctata Linnaeus. All these species,except perhaps undecimpunctata, are widely distributed in northernand eastern Siberia, in Canada, and in the northern United States.Most of them are found also in Alaska, and it is very probable that all of them will be found there when the country is studied more ART. 4 BEETLES OF THE GENUS COCCINELLA DOBZHANSKY 29 thoroughly. On the other hand, only one of these species?namely,transversoguttata?extends its distribution to the southern limits of bothnearctic and palaearctic regions; that is to say, most of the speciesin common to both regions are distributed only in the northern partof the hoiarctic region. The accompanying table shows the subdivi-sion of the genus into groups, and the correspondence between thenearctic and the palaearctic faunas. Group novemnotata transversoguttata. septempunetata.. . difflcilis trifasciata undecimpunctata. Nearctic species fnovemnotata HerbstIprolongata Crotch.(transversoguttata Falderman.johnsoni Casey Icalifornica Mannerheim InivJcoia subspecies monticola Muisant. [difficilis Crotch.. suturalis Caseytrifasciata Linnaeus.hieroglyphica subspecies tricuspis Kirby.(undecimpunctata Linnaeus1::::::::;:::::::::::: :::::::::::: Palaearctic species divaricata Olivier. itransversoguttata Falderman.magnopunctata Rybakow.[septempunetata Linnaeus.Inivicola Menetries.Ireitteri Weise.[saucerottei Muisant.Jtianshanica Dobzhansky.liranica Dobzhansky.[trifasciata Linnaeus.