eS 8 ed i ee 4 eS Pogeer2 Se, FAS te He. Pe Fee eee 7 a. "i - ion Tae oe i thie Ope RON RT te ad aes one . : : Tonal onan Ta oe” aha tC aE hat a elt 9 nag tal 4 . Cpl a we LMign Tee sant, ~ a \ ba nt i — ee ees — te : = x : a erie 9g Fs at Some ere Se ee. el rt ; i ne = 5° cette ee ee Ain ale Spe SE ne, pee . : ~ ‘ c ae eee fa woe ats : ree P meres rs i sea Ue ie vm “ ee eS aS ate En eae , * - - a ‘ near eee ee = oe Dd Inte RE Pia act! ak ants oe Y . tie ay EN Mate F Mothg te — Ba tee Ree tage a ee swith * a eer an Digitized by the Internet Archive in 2011 with funding from University of Toronto http://www.archive.org/details/reviewoflivingreOOpete ROYAL ONTARIO MUSEUM LIE | | ' ' 3 1761 05013 5938 CONTRIBUTIONS OF THE ROYAL ONTARIO MUSEUM OF ZOOLOGY AND PALAEONTOLOGY RO Va, LIBp ‘ Uy vi ; ‘TA RY SITY GO. MUs No. 34 TOr 100 6, To; a SE UY4 ONTO . atN.S P, mL: O NTARIO Can AD, A REVIEW OF THE LIVING REPRESENTATIVES OF THE GENUS ALCES By Randolph L. Peterson TORONTO OCTOBER 15, 1952 % rf 3 cre. Shes CONTRIBUTIONS OF THE ROYAL ONTARIO MUSEUM OF ZOOLOGY AND PALAEONTOLOGY No. 34 A REVIEW OF THE LIVING REPRESENTATIVES OF THE GENUS ALCES By Randolph L. Peterson TORONTO OCTOBER 15, 1952 “poyulluexad Uddq JIAPY Sustuoeds Yory aM WOW] SIDTRIO] YIM BOLIOULY YIAON ul ISOOUL jo UONNGTAYSIp at} SUIMOYS dey TL “OIA —----- serey jo abuey juasaiy X Ppew useq @BAPY SUO!JINposjUy YUM 4e SiUtog ° poulmexy suawirseds ro) S2i4ujyesoq adAy A REVIEW OF THE LIVING REPRESENTATIVES OF THE GENUS ALCES by RANDOLPH L. PETERSON THE GENUS Alces (North American moose and Old World elk) has never been satisfactorily revised. Taxonomic treatments of the New World moose have been restricted to descriptions of new forms without any serious attempt to evaluate the relation- ships of these races with those of the Old World. Unfortunately, the present attempt to consider all the modern representatives of this genus is seriously handicapped by the lack of adequate series of specimens from the Old World and the paucity of available taxonomic literature dealing with these forms. The most recent revision of the Old World elk or moose available to the author is that of Flerov (1931) who bases his review on a series of some sixty skulls of the genus Alces. The present treat- ment is based on a study of 304 specimens (mostly skulls), seven skulls representing two Old World forms and 297 specimens from North America. ACKNOWLEDGMENTS I am indebted to the following persons and institutions for permission to examine moose specimens in their scientific col- lections and for providing facilities for making critical studies of this material: Drs. F. J. Alcock and R. M. Anderson, National Museum of Canada. Drs. Remington Kellogg, David H. Johnson and Henry W. Setzer, United States National Museum. Dr. H. H. T. Jackson, Mr..S. P. Young and Miss Viola Schantz, Fish and Wildlife Service, United States Department of Interior. Drs. H. E. Anthony, G. H. H. Tate and Mr. H. M. van Deusen, American Museum of Natural History. Mr. H. T. Green, Academy of Natural Sciences of Philadelphia. Drs. A. S. Romer and C. P. Lyman, Museum of Comparative Zoology at Harvard College. Dr. J. K. Doutt and Miss Caroline Heppenstall, Carnegie Museum. Drs. E. R. Hall and R. H. Baker, University of Kansas, Museum of Natural History. Mr. A. M. Bailey, Colorado Museum of Natural History. Mr. Edmund B. Rogers and the Park Naturalist staff of Yellowstone National Park. ] 2 R. O. M. Z. AND P. CONTRIBUTIONS Dr. Don Quimby of the Department of Zoology, Montana State College. Drs. Philip L. Wright and G. F. Weisel, Montana State University. Mr. C. J. Guiguer, Provincial Museum of British Columbia. Drs. Ian McT. Cowan and W. A. Clemens, Department of Zoology, University of British Columbia. Messrs. Karl P. Schmidt, C. C. Sanborn and D. D. Davis, Chicago Natural . History Museum. Drs. W. H. Burt and E. T. Hooper, Museum of Zoology, University of Michigan. Mr. L. A. Walden and my wife, Elizabeth Peterson, were of great assistance while examining material in the above collections. I also wish to express my appreciation to Messrs T. M. Shortt, Eric Thorn and Archie Reid for the preparation of illustrations. Quotations from Allen (1940) have been made with per- mission of the American Museum of Natural History. EXPLANATIONS Measurements. All measurements of specimens are given in millimeters. The following are cranial measurements which have been recorded in table I and found to be the most useful ones from a set of 44 which were recorded for each complete skull examined. Greatest length—Length from anterior tip of premaxillae to the posterior point of the occiput. Median palatal length—Length from anterior tip of premaxillae to the median posterior edge of the palate. Length of rostrum—Length from anterior tip of premaxillae to anterior rim of orbit. Length of nasal aperture—Length from anterior tip of premaxillae to junction of nasal and maxillary bones. Mastoid breadth—Greatest transverse dimension of skull across mastoid process; outside of one to outside of the other. Height of occiput—Vertical dimension from the inion to the lower lip of the foramen magnum. Length of upper toothrow crowns—Greatest distance from the anteriormost edge of the crown of the first premolar to the posteriormost edge of the crown of the last molar. Greatest width of palate between toothrows—Greatest width between third upper molars at the margins of the alveoli. Least width of palate between toothrows—Least width between first upper pre- molars at the margins of the alveoli. “Per cent flare’ of nasal aperture—An expression of the shape of the nasal aperture derived by subtracting the measurement of the width of the nasal aperture at the junctions of the nasal and maxillary bones from the measurement of the greatest inside width of the nasal aperture (along the dorsal rim) and calculating the difference in terms of its percentage of the length of the nasal aperture. PETERSON: THE GENUS ALCES 3 Nature of material studied. Special emphasis has been placed on cranial characters and a majority of the specimens examined are represented by skulls only. Skins were examined where available and direct colour notes were recorded using Ridgway’s “Color Standards and Nomenclature, 1912.’’ Where possible, comparable skins (of the same season) of the various races were compared directly with one another. Intergrading specimens. Specimens which showed evidence of possible intergradation are listed under ‘‘Remarks’’ under A. a. andersont rather than under the “specimens examined”’ section of the race to which they might seem best assignable. Abbreviations of institutions. Institutions containing the speci- mens examined are abbreviated as follows: Academy of Natural Sciences, Philadelphia—A.N.S.P.; American Museum of Natural History—A.M.N.H.; Biological Surveys Collection (U.S. Fish and Wildlife Service)—B.S.C.; British Columbia Provincial Museum—B.C.P.M.; Carnegie Museum—Carn.M.; Chicago Natural History Museum—C.N.H.M.; Denver Museum of Natural History—D.M.N.H.; Montana State College— M.S.C.; Museum of Comparative Zoology, Harvard University—M.C.Z.; National Museum of Canada—N.M.C.; Royal Ontario Museum of Zoology and Palaeontology—R.O.M.Z.P.; United States National Museum—U.S.N.M.; University of British Columbia—U.B.C.; University of Kansas, Museum of Natural History—U.Kan.; University of Michigan, Museum of Zoology— U. Mich.; University of Montana—U. Mont.; Yellowstone National Park— HEN Be ek VARIATION Sexual variation. The variation between sexes of moose is so great that separate treatments must be made. In addition to averaging larger, the males bear antlers which result in certain modifications of the skull. Indiwidual variation. The extent of divergence from the mean exhibited in a series of specimens is quite wide in moose. In order to evaluate the variation found in such a wide ranging species, calculations of the standard errors and the standard deviations have been made for both sexes of the middle-aged specimens studied (see fig. 2). The observed limits of the range of variation are also given for the three age groups of adult males examined (see table I). Variation with age. The range in size variation is so great in skull measurements for animals the size of moose that many criteria must be defined in terms of relative proportions. There are, nevertheless, changes in both direct sizes and _ relative proportions as the animal grows older; for this reason it has PaCS Waa ri We oes Gos 21-243 10'11 16 6 4 15 224 Sill 13 6 4 23259 0. ,11 15.6 4 53 Mmm Aaamericana C—A.a.andersoni A.ashiras’ EA a. gigas Fic. 2. Variation in skull characters of North American moose. Median horizontal bars represent the mean, the central enclosed areas represent two standard errors iS SE = oa VN and the median vertical lines represent two standard deviations on either side of the mean eae a TiN | Figures above indicate number of specimens in each sample. A. Ratio of the least width of the palate (between toothrows) to the length of upper toothrow crowns. B. Ratio of the height of the occiput (inion to the lower lip of the foramen magnum) to the mastoid width. C. Ratio of the difference between the greatest inside width of the nasal aperture- and the width of the aperture at the junction of the nasals, to the length of the nasal aperture. PETERSON: THE GENUS ALCES 5 seemed necessary to separate all adult specimens (above two years of age) into three categories—young adults, middle-aged adults, and old adults. This classification is based on the degree of wearing on the dentition. Specimens showing little or no wear on the teeth, especially the first lower premolar, are re- garded as young adults. Those exhibiting moderate wearing on the dentition are considered middle-aged adults; while well or excessively worn dentitions are classed as old adults. Within these groups, the young adults obviously show the most striking variations. It is extremely difficult to obtain comparative samples containing equal representations of the various stages of development within this age group because of the rapid growth rate during this period. In table I, showing some of the variations found in cranial measurements of the various races of moose, the series of A. a. andersoni being the largest, seems to give the most ade- quate representation of the variations found within the three age groups considered. Here it can be seen that considerable growth takes place during the young adult stage, with little or no increase beyond this middle-aged group. With respect to the dentition, however, there is continued decrease in the length of the upper toothrow crowns as the animals grow older. This seems almost directly attributable to wearing away of the crowns through use. With the decrease in the length of the toothrow, it necessarily follows that there is a change with age of such useful taxonomic characters as the relative widths of the palate compared to the length of the toothrow. The ratio of the least width of the palate, between the tooth- rows, to the length of the upper toothrow crowns, showed a consistent increase with age in all groups examined. A change with age in the shape of the occiput is also evident in the males of all races represented. The height of the occiput increases slowly from the young adult stage onward, while the mastoid width increases markedly between the young and middle-aged adult groups. Undoubtedly, a part of this change is correlated with the development of antlers, and the required modification for their support. Similar trends were also noted in a smaller series of A. a. andersoni females, but in this case the changes were less marked. Geographic variation. There has been considerable disagree- ' ment concerning the number of valid species that should be recognized for this genus. Lydekker (1915) recognized only one R. O. M. Z. AND P. CONTRIBUTIONS Oo A. a. americana | | | | | oS A. a. andersoni TABLE I.—Cranial measurements of moose. The mean is followed by th Young ad. Mid-ad. Old ad. | Young ad, Mid-ad. Olc Greatest length 578.0 (5) 605.9 (21) 599.3 (3) 569.5 (11) 591.7 (24) 593. (554-590) (574-660) (590-610) | (538-586) (559-621) (576 Median palatal! 339.3 (3 360.4 (17) 351.0 (2) | 300.9 (11) | 349.7424) 350. length (321-357) (342-394) (344-358) | (309-350) (331-368) (334 Length of rostrum 351.6 (5) 370.7 (23) ote £4) 347.5 (11). 360.7. (25) 360 (338-370) (348-398 ) (356-387) | (326-363) (335-382) (346 Length of nasal DAS ap AAAS) 275.7. (23) 276.0 (4) | 249.5 (11) 265.0 (25) 267) aperture (238-263) (258-294) (261-286) | (229-260) (248-292) (25574 Mastoid breadth 149.7 (4) 168.6 (21) 167.0 (3) 150.4 (11) 164.0(25) 164.4) (145-155) (156-177) (166-169) | (139-162) (157-176) (152). Height of occiput 118.0 (3) 12057-4415) 122.0 (2) 120.8 (9) 125.2.(22), 124 (115-120) (110-129) (121-123) (115-124) (115-134) (118 Ratio of height of | 78.1 (2) foul Lo) 72.8 (2) 80.0 (9) 76.8 (22) 75.44. occiput to mastoid (76.8-79.4) (70.4-75.8) (71.6-74.0) | (72.5-86.3) (68.8-85.2) (72.0). breadth (%) Length of upper 150.6 (5) 152.6 (24) 148.7 (3) 152.9 (11) 149.8 (24) 146: toothrow crowns’ (145-157) (147-162) (146-153) (149-156) (144-158) (140 i Greatest width of 83.3 (3) 89.1 (21) 87.0 (2) 83.4 (11) 90.3 (25) 90.2}! palate between (80-88) (80-97) (85-89) (78-88) (81-100) (84 toothrows Ratio greatest 55.4 (3) 58.4 (21) 58.0 (2) 54.6 (11) 60.3 (24) 62.04 | width of palate (53.4-57.2) (58.0-63.7) (57.9-58.1) | (51.0-57.5) (55.5-66.4) (57.2) (between tooth- rows) to length of upper toothrow crowns (%) Least width of 52.6 (3) 54.7 (21) 54.0 (2) 54.5 (11) 57.4 (25) 58.1 palate between (52-54) (48-62) (538-55) (51-60) (538-65) (52) toothrows Ratio least width 35.0 (3) 35.6 (21) 36.0 (2) aor CLL) 38.3 (24) 39.91) of palate between (33.8-36.8) (31.0-40.7) aetla (33.3-39.0) (33.6-43.0) (35.6) toothrows to length of upper toothrow crowns (%) ‘‘Per cent flare’ of 8.00 (5) 7.88 (23) 7.02 (4) 8.53: C1) 9.25 (25) 8.71} nasal aperture (6.49-9.49) (6.02-10.64) (4.60-9.10) | (6.42-10.29) (6.24-13.25) (7.6 F(2) Mid-ad. 601.0 (7) (577-634) 344.0 (7) (324-379) 338.4 (9) (334-386) 267.4 (9) (251-293) 160.9 (10) (145-169) 122.8 (4) (118-127) 75.3 (4) (74.7-76.2) 142.9 (11) (136-151) 88.1 (9) (85-91) 61.2 (9) (57.3-66.9) 59.2 (9) (53-63) 41.4 (9) (37.3-44.8) 11.11 (9) PETERSON: THE GENUS ALCES |} specimens examined (in parentheses) with the observed limits given below. Oo A.a. shirasi Old ad. oO A. a. gigas Mid-ad. 579.2 (5) (569-594) 333.8 (5) (322-344) 349.5 (5) (333-359) 257.5 (5) (254-264) 164.7 (6) (160-174) 121.2 (6) (115-127) 73.5 (6) (69.6-76.6) 140.8 (6) (138-145) 90.3 (6) (85-94) 64.2 (6) (60.3-67.6) 61.5 (6) (56-67) 43.7 (6) (40.0—46.2) 11.59 (4) 2.05) (10.10-12.36) (9.26-12.60) 633.8 (11) (601-661) 361.2 (9) (355-382) 381.2 (11) (364-410) 284.4 (11) (263-297) 172.5 (11) (163-181) 134.1 (9) (128-140) 77.7 (9) (72.7-83.2) 147.5 (11) (142-155) 95.9 (10) (92-101) 65.1 (10) (60.0-67.8) 68.0 (10) (62-71) 46.3 (10) (40.0-49.0) 9.70 (10) (7.67-11.50) (9.32-11.35) Oo A.a. pfizenmayeri |S A.a.alces Old ad. | Mid-ad. Oldad. | Mid-ad. 650.0 (4) | 596.0 (2) 621 (1)! 574 (1) (616-686) | (587-605) ayy ash 375.5 (4) | 345.0 (2) 371 (1)| 349 (1) (351-390) | (343-347) chi re 389.0 (5) | 360.5 (2) 379 (1)| 349 (1) (363-408) | (358-363) ac ah 291.6 (4) | 267.5 (2) 287 (1)| 242 (1) (275-300) | (266-269) mee pee! 172.5 (4) | 157.5 (2) 176 (1)| 159 (2) (165-183) | (156-159) ae sie 132.8 (4) | 120.2 (2) 1315(1)| 124.5 (2) (126-144) | (118-122) ex =| (1ES2190) Ve (Bt TON. | eee eek? (74.0-78.6)|(75.6-76.6) .... ahs 145.4 (5) | 145.5 (2) 147 (1) | 149.7 (3) (141-149) | (143-148) See TANG Eee) 97.2 (5) | 92.5 (2)- 102(1)| 94.0 (2) (90-106) | (92-93) ite (90-98) 66.8 (5) | 63.6.(2) 69.5 (1)| 63.5 (2) (62.4—71.6)|(62.8-64.4) .... | (62.5-64.5) 72.4 (5) | 66.5 (2) 69 (1) | 62.0 (2) (61-80) | (65-68) Pash (60-64) 49.9 (5) | 45.8(2) 47.0(1)| 41.9 (2) (40.8-54.1)|(43.9-47.6) ..... | (41.7-42.1) 10.01 (4) | 8.78 (2) 9.05 (1)| 8.25 (1) BiH OAN HT ae ~ 8 R, O. M. Z. AND P. CONTRIBUTIONS species (A. alces). However, most North American authors have continued to recognize from one to three New World species as distinct from the Eurasian forms. Lydekker (1915) listed two races for the Old World and three for North America, not in- cluding A. a. shirast. Flerov (1931) reviewed the Old World forms and supported Lydekker’s theory that both Old and New World forms are conspecific. Further acceptance has been made by Allen (1940), Hall (1946) and others. Flerov (1931) concludes that Lydekker’s description of A. bedfordiae is not diagnostic of any race of moose and that A. a. pfizenmayert should be the name applied to the elk of northern and central Siberia. He describes a small specimen from the Amur River region along the Manchurian border which he regards as different from other Old World races although he does not give it a distinct name. Allen (1940) has shown that Cervus cameloides was applied to elk from Manchuria by Milne-Edwards in 1867 and thus pro- vides the earliest available name for an eastern Asiatic race. Milne-Edwards’ description was again based on one or perhaps more antlers and it is therefore difficult to assign diagnostic features. It therefore seems possible that a third race may occur, or has occurred, in the Amur River region of southern Siberia, northern Mongolia and northern Manchuria which would bear the name Alces alces cameloides (see fig. 5). Unfortunately I have been unable to examine material from this region and therefore cannot determine the validity of its status. If it is not a distinct race the name may prove available for all the elk of eastern Asia. In this treatment I have followed Flerov in considering the northern Asiatic animals as pfizenmayeri with the possibility that the elk of the Amur River region represent a smaller distinct race, cameloides. In a preliminary review of North American moose (Peterson, 1950) it was felt that insufficient Old World specimens were available to properly evaluate the status of the relationships between A. americana and A. alces. A more thorough study of three available skulls of A. a. pfizenmayert and data presented by Flerov (1931) indicate a close relationship between that race and A. a. gigas. Since there seems little justification for sepa- rating the North American moose as a distinct species, one is compelled to agree with Lydekker and Flerov that it should be regarded as conspecific with Alces alces. The North American moose are divided into four geographic races. In fig. 2, showing a statistical treatment of three cranial characters, a general east-west cline is evident in such characters ‘pose1oAr susutioads jo Joquinu 94} 0} Jojor sisoyjueied ur sams8iy *(s]ing y[Npe) $a97p sarap JO soveI dy} Jo sainqiade jeseu ay} Jo adeys pue 9ZIS ISVIOAR BYTE ‘OT ($2) (7) (6) CH") (S) (z) euerizowe' ey 1UOS4apue “ey ISe4lyS “CY sebib'e'y vaakewuaziyd “ery sane “ey L:GSiz 0-597 +192 O-vLZ “uns 0-767 8-SS L' 6 Les ST Alces alces americana eee - Fic. 5. Map showing the world-wide distribution of Alces alces. Characters. The moose (and Old World elk) is readily dis- tinguished by its large size (about the size of a saddle horse; the largest living Cervid), heavy body, long legs, high shoulder 14 R. O. M. Z. AND P. CONTRIBUTIONS region, short tail, broad overhanging muzzle, large ears, short neck and unique dewlap or ‘“‘bell’’. The nose is covered with short hair except for a bare spot between the nostrils which is usually more or less triangular in shape. Adult males develop extremely large antlers which are characteristically broad palmate blades with projecting tines. The upper part of the body is quite dark in new spring pelage, appearing almost black in some individuals, and in others varying towards dark brown, reddish brown and grayish brown. The underparts of the belly and lower legs are distinctly lighter, usually either grayish or brownish in colour, sometimes almost whitish. There is a gradual fading in body colour during summer, fall and winter to produce a more grayish effect by early spring. In addition to general characters given above, Alces may be distinguished by skull greatly lengthened in premaxillary region; nasal region shortened with sizable nasal aperture; distance from front of nasal to front of premaxilla about equal to that from back of nasal to back of occiput; vomer lower posteriorly, not dividing aperture of posterior nares; lacrimal vacuity widely open, the pit well developed, maxillary canines usually absent in both sexes; lower canines incisiform, incisors but little differentiated; molars and premolars rather broad and low crowned. Median metacarpals united as a cannon bone; lateral metacarpals strongly attenuated, rudimentary, with only distal ends developed as styloid vestiges (teleometacarpalian). Meta- tarsal gland absent; front hoofs larger than posterior ones, both long, narrow and pointed. ALCES ALCES ALCES (Linnaeus) European Elk 1758. Cervus alces Linnaeus, Syst. Nat., I, 10th ed., p. 66. 1910. Alces machlis typicus Ward, Records of Big Game, 6th ed., p. 66. 1913. Alces machlis uralensis Matschie, Beroff. Institute Jugdkunde, vol. 2, Pp. 2199. 1913. Alces machlis meridionalis Matschie, op. cit., p. 156. 1914. Alces machlis tymensie Zukowsky, Archiv. f. Naturgeschichte, 80 Jahry., Abt.cAN. 9)-pr42; 1914. Alces machlis angusticephalus Zukowsky, op. ctt., p. 44. 1915. Alces alces alces Lydekker, Cat. Ungulate Mamm.: Brit. Mus. (Nat. Fitst.); volo 4; pi: 232. Type specomen. Unknown. Type locality. Sweden. Geographical distribution. Northern Europe and_ western Fic. 6. Skulls (dorsal view) of six races of Alces alces. A. A.a.americana, adult female, R.O.M.Z.P. no. 19566 from Hayward Twp., Algoma Dist., Ontario, total length, 587 mm.; B. A. a. americana, adult male, R.O.M.Z.P., no. 19551, from Twp. 11-G, Sudbury Dist., Ontario; C. A. a. andersont, adult male, R.O.M.Z.P., no. 19548, from Overflowing River, Manitoba, t.]. 604 mm.; D. A. a. shirasi, adult male, U. Mich., no. 64015, from Wyoming, t.l. 616 mm.; E, A.a. gigas, adult male, U.S.N.M., no. 140368, from Hope City, Alaska, t.1. 629 mm.; F. A. a. pfizenmayeri, adult male, U.S.N.M., no. 155484, from Junction of Aldan and Lena Rivers, Russia, t.1.587 mm.; G. A. a. alces, adult male, U.S.N.M.., no. 196347, from Trondhjem, Norway, t.l. 574 mm. 16 R. O. M. Z. AND P. CONTRIBUTIONS Siberia, eastward probably as far as the Yenisey River and Altai Mountains (Flerov, 1931). Characters. Flerov (1931) gave the following criteria for dis- tinguishing this race from A. a. pfizenmayert (p. 72): (1) Length of skull (adult male) from 550 to 580 mm. (2) Length of the rostrum (from the anterior edge of the orbit, to the end of the intermaxillars) of the adult male from 329 to 347 mm. (3) Pal- atal processes of intermaxillar bone are narrow above, forwards of vomer, and form a spout-shaped concavity; their width is smaller than the vertical diameter of infraorbital foramen. (4) Nasal processes of intermaxillar bone not narrowed up- wards, but are widened spade-like with a blunt end. (5) Colour- ing greyish brown not passing into blackish tints; the limbs of a lighter colour. Remarks. Anexpanded list of synonyms for the species Alces is given by Lydekker (1915). Cranial measurements for two specimens are given by Miller (1912, p. 984). The shape of the nasal process of the premaxillary bone seems distinct enough to distinguish this race from all others. In four specimens examined, the nasal process of the premaxilla extended to within 37.5 mm. (15 to 52) of the nasals. In similar measure- ments of three specimens A. a. pfizenmayer1; the average was 46.0 mm. (42 to 50) while the North American races (mid-adult bulls) averaged as follows: A. a. americana, 72.2; A. a. andersont, 75.6; A. a. shirasi1, 78.6 and A. a. gigas, 81.3 (see fig. 3). Specimens Examined. A total of four as follows: Norway: Trodhiem, 1 (U.S.N.M.); no definite locality, 1 (M.C.Z.). Finland: Near Karstula, 1 (M.C.Z.). Russia: Vicinity of Leningrad, 1 (A.M.N.H.). ALCES ALCES CAMELOIDES (Milne-Edwards) Manchurian Elk 1867. Cervus cameloides Milne-Edwards, Ann. Sci. Nat., Zool. ser. 5, vol. 7, Pe oti. 1868-74. Cervus alces Milne-Edwards, Recherches pour servir a |’Hist. Nat. des Mammiferes, p. 181. ? 1902. Alces bedfordiae Lydekker, Proc. Zool. Soc. London, vol. 1, p. 109 (indefinite locality, possibly applies to this form). ? 1908. Alces machlis bedfordiae Lydekker, A trip to Pilawin, p. 85. ? 1915. Alces alces bedfordiae Lydekker, Cat. Ungulate Mamm., Brit. Mus. (Nat. Hist.), vol. 4, p. 234. 1931. Alces alces subsp. ? Flerov, Comptes Rendus de |’Academie des Sci. de l’URSS., p. 74. 1940. Alces alces cameloides Allen, Mamm. China and Mongolia, Am. Mus. Natidists vol. ily 2p. 1205, Fic. 7. Skulls (ventral view) of six races of Alces alces (same specimens shown in fig. 6). A. A. a. americana, female; B. A. a. americana, male; C. A. a. andersoni; D. A.a. shirasi; E. A. a. gigas; F. A.a. pfizenmayert; G. A.a. alces. 18 R.O. M. Z. AND P. CONTRIBUTIONS Type Specimen. Unknown (?). Supposedly one or more antlers in the Paris Museum. Type Locality. Manchuria (?). Geographical Distribution. Limits of range unknown. Sup- posedly from the Amur River region of extreme southeastern Siberia, northern Manchuria and northeastern Mongolia (see fi.) Characters. From the original description Allen (1940, p. 1206) provides the following: . . . Milne-Edwards (1867) briefly characterizes some antlers brought back by Fontamer, which, though procured in China, came originally, as he was in- formed, from Manchuria. These, writes Milne-Edwards, somewhat recall those of a young elk or moose, but appeared to be from adult animals; they were but slightly elevated, and remarkably heavy; the short beam carried an anterior basal tine, which in adults is almost as stout as the main branch and is bifurcate. The palms are very broad, especially in youth. The main beam turns abruptly backward and gives off a second nearly vertical tine. Concerning a specimen from this same general region, Flerov (1931, p. 74) writes as follows: In the collection of the Zoological Museum there is a skull of a semi adult elk from the environs of the Nicolajcusk1j post on the Amur, brought by L. Schrenk. By its dimensions it does not exceed the size of the European elk, and is con- siderably smaller than all the east Siberian elks. The intermaxillare are like those of the eastern elks. Allen (1940, p. 1205) describes the pelage as follows: The skin of an immature female secured by Dr. R. C. Andrews sixty miles northeast of Urga, Mongolia, is a mixed gray and brownish on the cheeks, neck and sides of the back, but the chest and flanks are clearer darker brown and the chin is blackish brown. The forehead and muzzle are tinged with ochraceous, and the feet and legs are similar but slightly darker, becoming brown near the hoofs. A short mane stands erect along the median line of the neck and shoulders and a very small (one inch) “‘bell’”’ or tuft is present at the throat. lt appears paler than the moose of eastern North America. Remarks. The above references to Alces from this region are all based on what each author regards as apparently immature specimens, although Flerov appears to feel that this specimen actually represented a smaller form than pfizenmayerz. Without recourse to specimens from this area, I have accepted the possible occurrence of this race largely on the basis of Flerov’s recog- nition of its distinctness. He was apparently unaware of Milne- Edwards’ early name available for this race. Adequate material | may well reveal that the above mentioned are merely immature specimens of the Siberian race. In such an event, the name cameloides would apparently stand. Specimens Examined. None. "S9I]D ‘SDIIG "DY “WY LISPAlyS “Dp “| tuosaapud ‘vd Pp “dD taeul ‘vuvILUaWUD “DP D a b ney ‘ Ldak DU UaZyd ‘D * ro) ReAee cod ‘DUDILAIUD OS aes V Vv ‘(9 “SY UI UMOYS SUsUTIDVds BWIRS) $a9JD Saz] pV JO S9IeI XIS JO (MOIA [RI9}L]) STNYAS °g ‘Oly Uy j Hyp 20 R. O, M. Z. AND P,. CONTRIBUTIONS ALCES ALCES PFIZENMAYERI Zukowsky Siberian Elk 1910. Alces pfizenmayert Zukowsky, Wild und Rund, 16 Juhrg., Bd. II, p. 807. 21902. Alces bedfordiae Lydekker, Proc. Zool. Soc. London, vol. 1, p. 109. 21908. Alces machlis bedfordiae Lydekker, A trip to Pilawin, p. 85. 1911. Alces machlis jukutakensis Millais, Field, vol. 118, p. 113. 21915. Alces alces bedfordiae Lydekker, Cat. Ungulate Mamm. Brit. Mus. (Nat. Hist.), vol. 4, p. 234. 1931. Alces alces pfizenmayert Flerov, Comptes Rendus de |’Academie des sci.deVURSS: pr 3 Type Specomen. Unknown (?). Type Locality. Aldan River, Yakutia, Russia. Geographical Distribution. Northern Siberia, eastward from the Yenisey River (Flerov, 1931), probably south to Sayan, Yablonovy and Dzhugdzhur mountain ranges. Characters. Flerov (1931) gives the following: (1) Length of skull (adult male) from 585 to 620 mm. (2) Length of the rostrum from 359 to 373 mm. (8) Palatal processes of inter- maxillar bone are flat and broad, forwards of vomer, and forming no spout-shaped concavity, their width is larger than the vert- ical diameter of infra-orbital foramen. (4) Nasal processes of intermaxillar bone are gradually narrowed and pointed upwards. (5) Colouring blackish-brown, often very dark (almost black) ; the limbs may be of a brownish colour. Remarks. Flerov (1981) suggests the possibility that this form is identical with A. a. gigas. In comparing the cranial measurements of these two races (see table I) it becomes appar- ent that there are several striking similarities. In general A. a. gigas averages larger. The total length of the skull aver- aged 633.8 mm. (601-661) in 11 mid-adult bulls and 650.0 mm. (616-686) in four old-adult males. Similar measurements for A. a. pfizenmayert were 587 and 605 mm. for two mid-adults and 621 for a single old-adult. The length of the rostrum of A.a. gigas also appears comparatively longer, averaging 381.2 mm. (364-410) in 11 mid-adult males and 389.0 mm. (363-408) in five old-adults. Specimens of the Siberian race examined, measured 358, 363, and 379 mm., the last, an old- adult, being six millimeters larger than the upper limit listed by Flerov. On the basis of the material examined, the two races in question appear different with respect to the extension of the nasal process of the premaxilla. In A. a. pfizenmayert this process extended to within 42, 46 and 50 mm. (av. 46.0) of the nasals, PETERSON: THE GENUS ALCES 21 while in A. a. gigas the average was 81.38 for both age groups. When this measurement (which represents the dorsal exposure of the maxilla along the nasal aperture) is represented in terms of its length relative to the length of the dorsally exposed portion of the premaxilla, the average for the A. a. pizenmayert speci- mens is 19.6 (18.0, 20.4 and 20.4) per cent compared to 38.7 (22.5-49.7) per cent for A.a. gigas. The former falls beyond the limits for two standard deviations for the latter (2S = 38.7 + 16.20). A more detailed study of the relationships of these two races must await the availability of adequate series of specimens. In the meantime, however, it seems reasonable to consider them as distinct races. Specimens Examined. A total of three as follows: Russia: Junction of Aldan and Lena Rivers, 1 (U.S.N.M.); Central Siberia near Krasnoyarsk, 1 (A.M.N.H.), and Berosovka R. 200 mi. N. Sredne— Kolymsk, 1 (A.M.N.H.). ALCES ALCES GIGAS Miller Alaskan Moose 1899. Alces gigas Miller, Proc. Biol. Soc. Wash., vol. 13, p. 57. 1901. Alces machlis gigas Lydekker, Great and Small Game of Europe, etc., p. 49. 1902. Paralces gigas Allen, Bull. Am. Mus. Nat. Hist., vol. 16, p. 160. 1904. Alces americanus gigas Osgood, N. Am. Fauna, no. 24, p. 29. 21907. Alces columbae Lydekker, Field, vol. 9, p. 182 (thought at first to be from somewhere in British Columbia). 1915. Alces alces gigas Lydekker, Cat. Ungulate Mamm., Brit. Mus. (Nat. Hist.), vol. 4, p. 237. 1924. Alces gigas Miller, U.S. Natl. Mus., bull. 128, p. 491. 1929. Alces americana gigas Anderson, Natl. Mus. Can., bull. 56, p. 105. 1950. Alces americana gigas Peterson, Roy. Ont. Mus. Zool., Occ. Pap. no. 9, pi, 2. Type Specimen. U.S. National Museum, no. 86166. Type Locality. North side of Tustumena Lake, Kenai Pen- insula, Alaska. Geographical Distribution. Forested areas of Alaska, western Yukon and northwestern British Columbia. Characters. Miller (1899), in his original description, gave the following (pp. 57-58): A larger, more richly colored animal than the eastern moose. Skull with occipital portion narrower, palate broader, and mandible much heavier than in Alces americanus. Color—General color a grizzle of black and woodbrown darkening along spine and changing abruptly to clear black on chest, buttocks, and lower part of sides. Median line of belly hairbrown. Legs hairbrown or broccoli-brown with darker 22 R. O. M. Z. AND P. CONTRIBUTIONS shading. Head like back, but more finely grizzled. Ears yellowish white in- ternally, broccoli-brown externally. Skull and teeth—The skull of Alces gigas differs from that of A. americanus in its larger size and greater massiveness, as well as in certain details of form. Chief among the latter is the great breadth of the palate, relatively to the length of the toothrow. In three males of A. gigas the ratio at least palatal breadth (be- tween anterior premolars) to length of toothrow is respectively 47.1, 47.1, and 44.7. In three males of A. americanus it is only 36, 36, and 39. In this respect Alces gigas resembles Alces alces, though the Alaskan animal shows no approach to the conspicuous deepening of the antorbital portion of the skull, or the peculiar form of the premaxillary characteristic of the European species. The occiput is relatively higher and narrower than in A. americanus. In two males of the latter the ratio of depth between inion and lower lip of foramen magnum to greatest width across paroccipital processes is 68.5 and 72.2, while in three of A. gigas it is 81.8, 84.8 and 87.5. Remarks. Although described and accepted by several authors as a distinct species (Miller, 1924 and others) it appears, on the basis of skulls from northwestern British Columbia and southern Yukon examined by the writer, to be only subspecifically dif- ferent since intergradation seems apparent (Peterson, 1950). Lydekker (1915), Osgood (1904), Anderson (Laing and Ander- son, 1929, Anderson, 1947), Hall (1984 and 1936) and others have suggested this possibility. Presumably, Miller (1899) used A. a. americana material for comparison with the Alaska race, although his chief character for distinguishing gzgas (least width of palate between toothrow relative to length of toothrow crowns) still remains valid when comparing other North Amer- ican races (see fig. 2A and table I). With respect to the shape of the occiput, Miller’s character holds good for separating A.a. gigas from A. a. americana, but not from A. a. andersont and A. a. shiras1, if the mastoid width is used, which appears a more precise measurement than the width across the par- occipital processes (see fig. 2B). If the height of the occiput alone is compared, A.a. gigas appears distinctly greater. ‘Nine middle-aged adult bulls of the latter averaged 134.1 + 1.31 mm., while four cows averaged 127.2 + 1.51 mm. Corre- sponding measurements of 22 A. a. andersont bulls were 125.2 + 0.98 mm., while 14 cows averaged 115.1 + 1.23 mm.”’ (Peter- son 1950, p. 3). Specimens of A. a. americana and A. a. shirasi averaged slightly less than andersont. For a list of the specimens examined which showed apparent intergradation see ‘‘Remarks’”’ under A. a. andersont. Specimens Examined. A total of 38 as follows: Alaska: Funny River, Kenai Peninsula, 4 (A.M.N.H.) and 1 (C.N.H.M.); Rainy Pass, Kenai Peninsula, 3 (A.M.N.H.); Tustumena L., Kenai Pen- PETERSON: THE GENUS ALCES 23 insula, 1 (U.K.) and 1 (U.S.N.M.); Kelly River, Kenai Peninsula, 1 (C.N.H.M.); near Kassilof L., (Tustumena L. ?) Kenai Peninsula, 3 (C.N.H.M.); Kenai Peninsula (no definite loc.), 2 (A.M.N.H.), 2 (B.S.C.), 1 (Carn. M.), 1 (C.N.H.M.), 2 (U.K.); 18 mi. s. McCarthy, 1.(B.S.C.); Coleville R., 1 (B.S.C.); Chugachik Bay, 2 (A.M.N.H.); Mt. McKinley, L(BS:CA. British columbia: Dease R., McDame Post, 1 (B.S.C.); Atlin, 2 (B.C.P.M.). Yukon: Teslin River district, 2 (N.M.C.); N. Teslin Lake, 1 (N.M.C.); Harris Cr., Headwaters White R., 1 (B.S.C.); Kletson Cr. (Trib. White R. ?), 1 (D.M.N.H.). ALCES ALCES SHIRASIT Nelson Yellowstone Moose 1914. Alces americanus shirasi Nelson, Proc. Biol. Soc. Wash., vol. 27, p. 72. 1924. Alces americana shirast Miller, U.S. Natl. Mus., bull. 128, p. 490. 21907. Alces columbae Lydekker, Field, vol. 9, p. 182 (British Columbia ?). Type Specimen. U.S. National Museum, no. 202975. Type Locality. Snake River, Lincoln County, Wyoming. Geographical Distribution. \Western Wyoming, eastern and northern Idaho, western Montana, northward into southwestern Alberta and southeastern British Columbia. Occasional occur- rence in extreme northeastern Utah has been reported by Dur- rant (1952). Characters. “Differs strongly in early winter pelage from typical Alces americanus from Maine, Nova Scotia, and New Brunswick in having the entire top of the back, including upper side of neck rather pale rusty yellowish-brown, slightly washed on tips of hairs with dusky; ears paler and grayer; hoofs much smaller and shorter’’ (Nelson, 1914, p: 72). Hall (1934) was not able to substantiate the smallness of feet. In his original de- scription Nelson (1914) was unable to find any appreciable cranial difference between typical A.a. americana and A.a. shirast, although when considering measurements shown in fig. 2 there can be little doubt of significant difference be- tween these two. It approaches closer to A. a. andersoni but on the basis of specimens examined, including the type of A. a. shirast, it differs in having a greater degree of flare of the nasal aperture. The length of the upper toothrow crowns aver- aged the shortest of all races (see table I). Remarks. The colouring of the pelage along the withers and dorsal portion of the back has been pointed out by Nelson as the most distinctive character of this race. Great difficulty is encountered in making critical colour comparisons because of the impracticability of examining sufficient specimens in series. 24 R. O. M. Z. AND P. CONTRIBUTIONS Although there is much variation in coloration, especially with respect to seasonal changes, this race appears to represent the extreme in light coloration along the back, among North Amer- ican forms. In addition to being more extensive, this light area lacks the rufus tones usually present in A. a. andersoni (based on late fall specimens). The Alaskan race also has a fairly extensive area of paler coloration along the withers and back, but it is much less distinct and usually appears darker with a more brownish or reddish tint than A. a. shirasi. This light area is apparently least extensive in A. a. americana. For a list of specimens showing apparent intergradation see ‘‘Re- marks”’ under A. a. andersont. Specimens Examined. A total of 34 as follows: Alberta: Banff, 1 (U.S.N.M.); Waterton, 1 (U.B.C.). British Columbia: 4 or 5 mi. below Ochre Cr., 1 (U.S.N.M.); Vermilion R.., 1(B5:©))) Praser:R:. 1 (BSC, + Goldens (Ui. ): Colorado: Grand Co., 1 (D.M.N.H.). Idaho: Selway area, 2 (U. Mont.). Montana: Upper Missouri R., 1 (U.S.N.M.); Gallatin Canyon, Yellowstone National Park, 1 (M.S.C.); near Gardiner, Park Co., 1 (U. Mont.); 20 mi. S. Anaconda, Deer Lodge Co., 1 (Carn. M.); 8 mi. S. Wise R., Beaver Head Co., 2 (Carn. M.); 10 mi. N. Polaris, Beaver Head Co., 1 (Carn. M.). Wyoming: Teton Canyon, 1 (U.S.N.M.); Teton Co., 1 (B.S.C.), 1 (U. Mich.); S. Fort Buffalo Cr., Teton Co., 1 (A.N.S.P.); near Jackson, 1 (U.S.N.M.); Bridge L., S. of Yellowstone Park, 3 (A.M.N.H.); near Hawk’s Nest, Upper Yellowstone, 1 (A.M.N.H.); Yellowstone National Park, 4 (Y.N.P.), 1 (R.O.M.Z.P.); S. Ft. Shoshone R., near Cody, 1 (U.S.N.M.); Snake R.., Lincoln Co:,"3:(B.S.G.). ALCES ALCES ANDERSONI Peterson Northwestern Moose 1950. Alces americana andersoni Peterson, Roy. Ont. Mus. Zool., Occ. Pap. NO. 9... bi Way LOS 21907. Alces columbae Lydekker, Field, vol. 9, p. 18a (British Columbia ?). 21907. Alces columbae Lydekker, Zool. Rec., vol. 44, Mamm., p. 69 (Ontario ?). 21915. Alces alces columbae Lydekker, Cat. Ungulate Mamm., Brit. Mus. (Nat. Hist.), vol. 4, p. 236 (Ontario ?). Type Specimen. Adult male (skin and complete skeleton) no. 20068, Royal Ontario Museum of Zoology and Palae- ontology, collected by Mr. F. D. de Delly on December 18, 1949. Type Locality. Section 27, Township 10, Range 16, Spruce- wood Forest Reserve (15 mi. E. Brandon), Manitoba. Geographical Distribution. Northern Michigan and Minnesota, western Ontario, westward to central British Columbia, north PETERSON: THE GENUS ALCES 25 to eastern Yukon Territory and Mackenzie Delta, Northwest Territories. Characters. Differs from A. a. americana in having a greater width of palate (as measured between the upper toothrows) relative to the length of the upper toothrow crowns. In a series of skulls of 24 middle-aged bulls the least width of palate aver- aged 38.3 + 0.46 per cent of the length of the toothrow com- pared to 35.6 + 0.56 in 21 similarly aged bulls of A. a. amer- wana (see fig. 2A). In similar measurements of cows 16 A. a. andersoni specimens averaged 37.7 + 0.56 compared to 33.7 + 1.22 for 11 specimens of A. a. americana. Also differs from the latter by possessing a relatively higher and narrower occiput (as measured from the inion to the lower lip of the foramen magnum) relative to the mastoid width. A series of 22 middle-aged bulls averaged 76.8 + 0.77 per cent compared to 73.1 + 0.63 per cent in 15 males of A.a. americana. In 13 cows of the former the average was 81.5 + 0.71 compared to 76.6 + 1.02 for 11 of the latter (see fig. 2B). In bulls the nasal aperture of 25 specimens showed a distinct difference from 23 A. a. americana by being more widely flaring, although no significant difference between the cows of these two races was found in this case (see fig. 2C). In middle-aged cows, how- ever, a distinct difference in the ratio of the mastoid width rela- tive to the greatest zygomatic breadth was found with 138 specimens averaging 73.0 + 0.76 per cent compared to 76.2 + 0.65 in 11 skulls of A. a. americana. In this character bulls did not appear significantly different. Similarly the cows also differ by having a relatively shorter and wider rostrum when the greatest antorbital breadth is compared with the median palatal length. The former averaged 44.8 + 0.56 per cent of the palatal length in 15 cows compared to 42.6 + 0.39 in 11 A.a. americana females. In bulls similar measurements were 45.0 and 48.5 per cent respectively, but a slight overlap of this character was noted. Differs from A. a. shirasi in having a narrower and less flaring nasal aperture. In the latter the difference between the greatest inside width of the nasal aperture and the width at the junction of nasals averaged 29.9 + 0.64 mm. in 10 middle-aged males and 26.3 + 1.37 mm. in six females of similar age while the corresponding measurement for A. a. andersoni was 24.1 + 0.84 for 25 males and 20.5 + 1.02 for 16 females. When these measurements are compared to the length of the nasal aperture, a significant difference is found between these two races (fig. 2C). 26 R. O. M. Z. AND P. CONTRIBUTIONS In bulls A. a. andersoni differs from A. a. shirasi in having a relatively longer toothrow, averaging 149.8 + 0.73 in 24 middle-aged specimens of the former and 142.9 + 1.83 in 11 of the latter. Further difference is found in bulls in the least width of the palate between the toothrow relative to the length of upper toothrow crowns (fig. 2A) although in both tooth char- acters, overlapping was found in the smaller number of cows examined. A.a. anderson differs from A.a. gigas in having a much narrower least width of palate between toothrows relative to the length of the upper toothrow crowns (fig. 2A) and in having a lower occiput as measured from the inion to the lower lip of the foramen magnum. Nine middle-aged adult bulls of the latter averaged 1384.1 + 1.31 mm. while four cows averaged 127.2 + 1.51 mm. Corresponding measurements of 22 A. a. andersont bulls were 125.2 + 0.98 while 14 cows averaged 115.1 + F.238 mm: In colour, A.a.andersont appears intermediate between A. a. americana and A. a. shirasit with usually more light rufus tones appearing on the head and back region in fall specimens, although much variation is found with the seasons; light area of shoulder hump region usually more extensive than the former and less extensive than the latter. Hall (1936) came to similar conclusions regarding a specimen from Bowron Lake, British Columbia. Observations in the field give the impression that A. a. americana has a heavier and shorter neck. A check on the cervical vertebrae of one specimen from eastern Ontario and two from western Ontario (all with comparably sized skulls) indi- cated no significant difference in the length of the centra although each vertebra of the eastern form was consistently the heavier and wider. A further correlation of this condition is suggested in the relatively wider occiput (fig. 2B) of A. a. amer- icana. Remarks. Hall (1934) found that in moose from Bowron Lake, British Columbia, the least width of the palate amounted to 40 (39-42) per cent of the crown length of the upper toothrows in bulls and 36 (32-87) per cent in cows, while in two specimens from Minnesota the bull was 37 per cent and the cow 34 per cent. These measurements fall within the limits for A. a. andersont and thus seem referable to that race. Specimens showing varying degrees of apparent intergradation with other races were ex- amined as follows: PETERSON: THE GENUS ALCES 27 With A. a. gigas: British Columbia: Cassiar Mts., 1 (Carn. M.); Telegraph Creek. 3 (B.C.P.M.) and 1 (D.M.N.H.). Yukon: MacMillan R., 2 (B.S.C.); Sheep Mt., E. of Atlin L., 1 (C.N.H.M.); N. of Leslin L., 3 (N.M.C.); Teslin R., 3 (N.M.C.). With A. a. shirast: Alberta: Saskatchewan R., mi. 109, 1 (U.B.C.); Southesk L., Jasper Park, 1 (U.B.C.). British Columbia: Field, 1 (U.B.C.); Golden, 1 (U.B.C.). With A. a. americana: Ontario: Although no mature specimens are presently available from the centre of the zone of overlap of these two races, there seems little doubt that intergradation is taking place along a belt from the north shore of Lake Superior, northward. One yearling male from Flint Lake, 35 mi. E. Longlac exhibits a mixture of characters between A. a. americana and A. a. andersont, Specimens Examined. A total of 132 as follows: Alberta: Entrance, 1 (A.M.N.H.); Little Smoky R., N. of Jasper Park, 2 (A.M.N.H.); Thoral Cr., 1 (A.M.N.H.); Hay R. Flats, 1 (U. Mich.); Eagle Nest Pass, 1 (Carn. M.); Brazeau R., 175 mi. N.W. Calgary, 1 (Carn. M.); Elk Island Park, 1 (N.M.C.). British Columbia: Jarvis Pass, 1 (U.S.N.M.); McDame Post, Dease R., 1 (B:S.C.); Takla Landing, 2.(B:C:P:M)); Athn, t (B-C.P.M.); Tupper Cr;, 2 (B.C.P.M.); Chezacut, 1 (B.C.P.M.); Ootsa L., 2 (U.B.C.); Caribou Dist., 1~(U:B.C.); Quick, 6 (U.B.C.); Quesnel, 1 (U.B.C.); Telkwa, 1 (U.B.C.); Shesley Mts., 2 (A.M.N.H.); Level Mt., 1 (A.M.N.H.); no definite locality, 5 (U. Kan.) and 1 (A.M.N.H.). Manitoba: Portage la Prairie, 4 (U.S.N.M.); Hudson Bay R.R. (no definite locality), 1 (U.S.N.M.); Overflowing R., N. of Lat. 53, S. of the Pas, 1 (R.O.M.Z.P.); Sprucewood Forest Reserve, 15 mi. E. of Brandon (type specimen), 1 (R.O.M.Z.P.). Michigan: Chippewa Harbor, Isle Royale, 3 (U. Mich.); Lake Eva, Isle Royale, 1 (U. Mich.); Houghton Cove, Isle Royale, 5 (U. Mich.); McCargoe Cove, Isle Royale, 3 (U. Mich.) and 2 (A.M.N.H.); no definite locality, Isle Royale, 8 (U. Mich.). Minnesota: Temperance R., Cook Co., 1 (B.S.C.); Moose R. (?), 2 (U. Kan.); Thief L., Marshal Co., 1 (U. Kan.); Lake of the Woods, 19 (U. Kan.); no definite locality, Northern Minn., 1 (U.S.N.M.) and 1 (M.C.Z.). Ontario: (Unless otherwise designated specimens are in the Royal Ontario Museum of Zoology and Palaeontology). Whitefish L., Thunder Bay Dist., 1 (A.M.N.H.); St. Ignace Is., L. Superior, 18; Simpson Is., L. Superior, 2; Bead Is., L. Superior, 1; 40 mi. N.W. Port Arthur, 1; Long Bay, Lake of the Woods, 1; Ear Falls, Kenora Dist., 1; Gull Rock L., Kenora Dist., 1; Wabaskank L., Kenora Dist., 1; Nungessor L., Kenora Dist., 1; Narrow L., 20 mi. E. Kenora, Kenora Dist., 1; Sioux Narrows, Kenora Dist., 1; no definite locality, Kenora Dist., 1. Saskatchewan: Birch L., 1 (N.M.C.); Harper L., 1 (N.M.C.). Yukon Territory: Ross R., Canol Rd., mi. 95, 1 (N.M.C.); Lapie R. (Canol Rd. ?) mi. 120, 1 (N.M.C.); Teslin Dist., 3. (N.M.C.). 28 R. O. M. Z. AND. P. CONTRIBUTIONS ALCES ALCES AMERICANA (Clinton) Eastern Moose 1822. Cervus.americanus (Clinton), Letters on Nat. Hist. and Int. Resources of New York, p. 193. 1835. Alces americanus Jardine, Nat. Library, vol. 21 (Mammalia; deer, antelope, camels, etc.), p. 125. 1846. Cervus lobatus Agassiz, Proc. Boston Soc., vol. 2, p. 188. 1852. Alces muswa Richardson, Zool. Herald, Mamm., p. 66. 1873. Alces lobata Fitzinger, Sitzber. K. Ak. Wiss. Wien, vol. 68, pt. 1, p. 348. 1884. Alce americanus Merriam, Mammals of Adirondacks, p. 138. 1885. Alces machlis True, U.S. Natl. Mus., vol. 7, p. 592. 1901. Alces machlis americanus Lydekker, Great and Small Game of Europe, etc., p. 46. 1902. Paralces americanus Allen, Bull. Am. Mus. Nat. Hist., vol. 16, p. 160. 21907. Alces columbae Lydekker, Zool. Rec., Vol. 44, Mamm., p. 69 (Ontario 2). 21915. Alces alces columbae Lydekker, Cat. Ungulate Mamm., Brit. Mus. (Nat. Hist.), vol. 4, p. 236 (Ontario ?). 1915. Alces alces americanus Lydekker; op. cit., p. 234. 1924. Alces americana americana Miller, U.S. Natl. Mus., bull. 128, p. 490. Type Specimen. Unknown. Type Locality. ‘‘Country north of Whitestown”’ (probably in the western Adirondack region), New York. Geographical Distribution. From Maine and Nova Scotia west- ward through Quebec to central northern Ontario where it apparently intergrades with A. a. andersont. Characters. Differs from other moose in having a narrower palate relative to length of toothrow. Differs from A. a. ander- sont by possessing a relatively lower and wider occiput (as measured from the inion to the lower lip of the foramen magnum, relative to the mastoid width). In colour it usually appears darker than the latter in late fall coat with deeper brown tones instead of the lighter rufus of A. a. andersont. The extent of light coloration on the withers apparently averages the least of all North American races. Specimens Examined. A total of 64 as follows: Maine: Eustis, 1 (U.S.N.M.); Umbagog L., 1 (M‘C.Z.); Megantic, 1 (M.C.Z.); no definite locality, 4 (M.C.Z.) and 1 (D.M.N.H.). New Brunswick: Fredericton, 1 (U.S.N.M.); Nepisiquit, 1 (U.S.N.M.); Chip- man, 1 (U.S.N.M.); Bolt Mt. near Newcastle, 1 (U.S.N.M.); Gloucester Co., 1 (N.M.C.); Bald Mt., 1:(B.S.G.); Trousiers L., 1 (A-M.N.H.); no definite locality; 2 (B.S:C:); (A MCN) ame 4 ( Carn: i.) New York: Adirondacks, 1 (U.S.N.M.). Nova. Scotia: Bear R:, 1 (B.S:C. and 1 CV.CiZ,); Guysbero-Co,,. 4.08 se 2)e Head of Roseway R., 1 (B.S.C.); Kedgemakwoge L., 3 (B.S.C.); no definite locality, 1{ME.G.Z))3 7s PETERSON: THE GENUS ALCES 29 Ontario: (Unless designated otherwise, specimens are in the Royal Ontario Museum of Zoology and Palaeontology). Mackay, Renfrew Co., 1 (U.S.N.M.); Stonecliff, Renfrew Co., 1 (U.S.N.M.); Mattawa, Nipissing Dist., 1 (M.C.Z.); 250 mi. N.W.. Sault Ste Marie, 1 (Carn. M.); S.W. boundary Algonquin Park, 1; S.W. sect. of Algonquin Park, 1; Long L., Algonquin Park, 1; Upper Ottawa R., 1; Cavendish Twp., Peterborough Co., 1; Twp. 11B, Sudbury Dist., 1; Nemagos, Halsey Twp., Sudbury Dist., 1; Twp. 13H, Sudbury Dist., 1; Sadler Twp., Sudbury Dist., 1; Twp. 11G, Sudbury Dist., 1; Raney Twp., Sudbury Dist., 1; Twp. 12G, Sudbury Dist., 1; 15 mi. S. Kapuskasing, Cochrane Dist., 1; Kapuskasing Twp., Cochrane Dist., 1; Mowbray Twp., Cochrane Dist., 1; Twp. 46, Algoma Dist., 1; Hayward Twp., Algoma Dist., 1; Walls Twp., Algoma Dist., 1, and Cobden Twp., Algoma Dist., 1. Quebec: Teiton Clut (?), 1 (U.S.N.M.); Two Rivers (?), 1 (U.S.N.M.); Sandy Creek, 4 (N.M.C.); Spruce Grove, Pontiac Co., 2 (N.M.C.); Temiskasing L., 1 (R.O.M.Z.P.); Fumerton L., 50 mi. N.W. Amos, 1 (R.O.M.Z.P.). LITERATURE CITED ALLEN. GLOVER M. 1940. The mammals of China and Mongolia. Natural History of central Asia. Am. Mus. Nat. Hist.,; vol. 11, pt. 2, sec. 4, pp. 621-1350. ANDERSON, R. M. 1947. Catalogue of Canadian Recent mammals. Natl. Mus. Can. bull. 102, biol. ser. 31, pp. 1-238. DURRANT, STEPHEN D. 1952. Mammals.of Utah. Univ. Kans. Pubs., Mus. Nat. Hist., vol. 6, pp. 1-549. FLEROV, C. 1931. A review of the elks or moose (Alces Gray) of the Old World. Comptes Rendus de |’Academie des Sciences de |’U.R.S.S., pp. 71- 74, FrIcK, CHILDS 1937. Horned ruminants of North America. Bull. Am. Mus. Nat. Hist., vol. 69, pp. 1-669. HALL, E. RAYMOND 1934. Mammals collected by T. T. and E. B. McCabe in the Bowron Lake region of British Columbia. Univ. Calif. Pub. in Zool., vol. 40, no. 9, pp. 363-386. 1936. Identity of the Bowron Lake moose of British Columbia. Murrelet, vou. It, tans p71. 1946. Zoological subspecies of man at the peace table. Journ. Mamm., vol. 27, no. 4, pp. 358-364. Hay, 0.-P. 1923. The Pleistocene of North America and its vertebrated animals from the states east of the Mississippi River and from the Canadian provinces east of longitude 95°. Carnegie Inst. of Wash., pp. 1-499. 1924, The Pleistocene of the middle region of North America and its vertebrated animals. Carnegie Inst. of Wash., pp. 1-385. 1927. The Pleistocene of the western region of North America and its vertebrated animals. Carnegie Inst. of Wash., pp. 1-346. 30 R. O. M. Z. AND P. CONTRIBUTIONS LAInNG, H. M. and R. M. ANDERSON 1929. Notes on mammals of upper Chitina River region, Alaska, in Birds and mammals of the Mount Logan Expedition, 1925. Natl. Mus. Can. bull. 56, pp. 96-107. LYDEKKER, R. 1915. Catalogue of the ungulate mammals in the British Museum (Natural History). Brit. Mus. (Nat. Hist.), vol. 4, pp. 1-438. MILLER, GERRIT S., JR. 1899. Anew moose from Alaska. Proc. Biol. Soc. Wash., vol. 13, pp. 57-59. 1912. Catalogue of the mammals of western Europe (Europe exclusive of Russia). Brit. Mus. (Nat. Hist.), pp. 1-1019. 1924. List of North American recent mammals, 1923. U.S. Natl. Mus., bull. 128, pp. 1-673. MILNE-EDWARDS, ALPHONSE 1867. Observations sur quelques mammiféres du nord de la Chine (Extract) Ann. des Sci. Nat., Zool., ser. 5, vol. 7, pp. 375-377. NELSON, E. W. 1914. Description of a new subspecies of moose from Wyoming. Proc. Biol. Soc. Wash., vol. 27, pp. 71-74. OsGoop, WILFRED H. 1904. A biological reconnaissance of the base of the Alaska Peninsula. N. Am. Fauna No. 24, U.S. Dept. Agri., pp. 1-86. PETERSON, RANDOLPH L. 1950. A new subspecies of moose from North America. Roy. Ont. Mus. Zool., Occ. Pap. no. 9, pp. 1-7. REYNOLDs, S. H. 1934. A monograph on the British Pleistocene Mammalia. Alces. Paleo. Soc. Monogr., London, vol. 87 (suppl. pp. 1-16). Scott, W. B. 1937. A history of land mammals in the Western Hemisphere. Revised Ed., MacMillan Co., New York, pp. 1-786. * des tat 54 , “ —— LIBRARY R@YAL @NTARIO MUSEUM pe Ee So ee, ica She . See ee ~~ ee ee Co ehh ne =e . 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