H355 1934 f GEOGRAPHIC VARIATION AMONG BROWN AND GRIZZLY BEARS {URSUS ARCTOS) IN NORTH AMERICA E. RAYMOND HALL Museum of Natural History The University of Kansas Lawrence, Kansas 66045 SPECIAL PUBLICATION OF THE MUSEUM OF NATURAL HISTORY, UNIVERSITY OF KANSAS NUMBER 13 1984 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Copies of publications may be obtained from the Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. A list of available Special Publications is provided on the inside back cover. « HARVARD UNIVERSITY Library of the Museum of Comparative Zoology University of Kansas Museum of Natural History Special Publication No. 13 August 10, 1984 Geographic Variation Among Brown and Grizzly Bears (Ursus arctos) in North America By E. Raymond Hall Museum of Natural History The University of Kansas Lawrence, Kansas 66045 The University of Kansas Lawrence 1984 f 7 0 7 University of Kansas Publications, Museum of Natural History Editor: Joseph T. Collins Special Publication No. 13 Pp. i-ii; 1-16; 6 figures 1 table PubUshed August 10, 1984 Copyrighted By Museum of Natural History University of Kansas Lawrence, Kansas 66045 U.S.A. MCZ LIBRARY OCT 15 1987 HARVARD univhrsity Printed By University of Kansas Printing Service Lawrence GEOGRAPHIC VARIATION AMONG BROWN AND GRIZZLY BEARS {URSUS ARCTOS) IN NORTH AMERICA E. Raymond Hall Three species of bears of the genus Ursus inhabit the North American continent (Fig. 1), and up to the present (1983) at least 96 names have been proposed for American bears of the Ursus arctos group. These names are to be found in Merriam (1918 and 1929), 84 having been proposed by Merriam himself. All 96 names were proposed for material of Recent geologic age (material of post-Pleistocene age). Rausch (1963) commented meaningfully on geographic variation in Ursus arctos in North America and introduced (pg. 43) the term "local demes" to account for local variations, many of which Merriam had named as subspecies or even species. Rausch did not, however, formally synony- mize these names. Kurten (1973) explained the morphologi- cal variation in skulls of Ursus arctos and color variation in its pelage as inheritance from an ancestral population in eastern Asia and western North America and in the con- necting land (now under water) that joined the two continents at an earlier time. Kurten expressed variation in terms of "isophenes" at right angles to clines showing gradual changes in characters from one place to another. Hall and Kelson (1959) and Hall (1981) listed all names based on North American specimens of Ursus arctos (without formally using that name for a single inclusive species) and showed on a map (no. 498, 1981) the type localities and some marginal records of occurrence. The ninety-some names proposed for supposed kinds of these bears in North America is only about a third as many as the number (271) that Kurten (1973:2) wrote have been proposed for the corresponding bears in the Old World— a number he charac- terized as a waste of systematic effort. The time of arrival in North America of the ancestral stock of the modem Grizzly- Big Brown Bear (Ursus arctos Linnaeus) is not certainly known at this time. However, the species now occurs on both sides of the Bering Strait that narrowly separates (pres- ently less than 80 miles of water) the New and Old World continents (Figs. 3, 4). The Diomede Islands near the center of the Strait reduce the land-to-land distance to less than 50 miles. Snow-covered ice may in season completely bridge the water in the Strait. Against the preceding background of in- formation, nine subspecies of Ursus arctos are here recognized as having occurred in North America in Recent time (Fig. 2). Were it not for uncertainty about the provenience of some specimens from certain islands and the adjoining mainland of southern Alaska, it might be possible to recognize in North America a few more than the nine subspecies listed below. In the period 1962-1968 I measured cer- tain dimensions of the crania of adults and some subadults of Ursus arctos, and dimen- sions of the upper permanent teeth of those and younger specimens. Most measure- ments were recorded by my wife, Mary F. Hall, or by Charles Long as a Research Assistant, or by some other assistant. For rearrangement and other help with the meas- urement forms, I acknowledge the invaluable skill of Mrs. Eleanor Lohmann. Of the 2,476 skulls examined, 912 were from adult or old animals. The remainder were: juveniles (per- manent dentition not all in place); young (certain bones less than full size as shown by open cranial sutures); and subadults (sagittal crest short or wanting in males and in fe- SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 males temporal lines separated or barely touching). Anyone especially interested in cranial change with age should consult Zavatsky (1976) who listed dental and cranial features characterizing individuals of the "Brown Bear' ' according to eleven age groups rang- ing from group 1 ("Cubs to 5 months of age") to group 11 ("Both sexes— older than 18 years"). Any person who works with variation in American grizzly and big brown bears and wishes to understand why Merriam named as "species" many specimens that subse- quent taxonomists would have identified as "subspecies," or "local demes" (to use Rausch's 1963:43, terms), should read Mer- riam's two articles "Criteria for the Recogni- tion of Species and Genera," and "Why Should Every Specimen be Named?", in volume one, number one of the Journal of Mammalogy. The kinds recognized by Merriam are here regarded as belonging to nine sub- species of one species. Most of the mor- phological variants are regarded as "local demes" or individual variants of one or another of the nine subspecies. The precise assignment of each name and of the minor variants it represents is shown in the follow- ing synonymies. Detailed measurements and notes on which the following summary is based are on file at the Museum of Natural History, Uni- versity of Kansas, and are available for study by later students interested in geographic variation of these bears. Three facts that complicated such study are an unusual amount of variation, the unavailability of ade- quate material from some areas, and the impossibility of obtaining specimens now that the bears have been exterminated from much of their original range. Ursus arctos alascensis Merriam Grizzly or Brown Bear 1896. Ursus horribilis alascensis Merriam, Proc. Biol. Soc. Washington, 10:74, April 13, type from Unalaklik River, Alaska. 1902. Ursus kidderi Merriam, Proc. Biol. Soc. Washington, 15:78, March 22, type from Chinitna Bay, Cook Inlet, Alaska. 1904. Ursus kenaiensis Merriam, Proc. Biol. Soc. Washington, 17:154, October 6, type from Cape Elizabeth, extreme W end Kenai Penin- sula, Alaska. 1904. Ursus horribilis phaeonyx Merriam, Proc. Biol. Soc. Washington, 17:154, October 6, type from Glacier Mtn., Tanana Mts., Alaska (about 2 mi. below source Comet Creek, near Fortymile Creek, between Yukon and Tanana rivers). 1910. Ursus sheldoni Merriam, Proc. Biol. Soc. Washington, 23:127, September 2, type from Montague Island, Prince William Sound, Alaska. 1914. Ursus alexandrae Merriam, Proc. Biol. Soc. Washington, 27:174, August 13, type from Kasilof Lake, Kenai Peninsula, Alaska. 1914. Ursus innuitus Merriam, Proc. Biol. Soc. Washington, 27:177, August 13, type from Golofnin Bay, S side Seward Peninsula, west- em Alaska. 1914. Ursus internationalis Merriam, Proc. Biol. Soc. Washington, 27:177, August 13, type from Alaska- Yukon boundary, about 50 mi. S of Arctic Coast. 1914. Ursus toklat Merriam, Proc. Biol. Soc. Washington, 27:182, August 13, type from head Toklat River, N base Alaska Range, near Mt. McKinley, Alaska. 1914. Ursus kidderi tundrensis Merriam, Proc. Biol. Soc. Washington, 27:196, August 13, type from Shaktolik River, Norton Sound, Alaska. 1916. Ursus cressonus Merriam, Proc. Biol. Soc. Washington, 29:137, September 6, type from Lakina River, S slope Wrangell Range, Alaska. 1916. Ursus eximius Merriam, Proc. Biol. Soc. Washington, 29:139, September 6, type from head of Knik, Cook Inlet, Alaska. 1916. Ursus nuchek Merriam, Proc. Biol. Soc. Washington, 29:146, September 6, type from head Nuchek Bay, Hinchinbrook Island, Prince William Sound, Alaska. 1929. Ursus holzworthi Merriam, Proc. Biol. Soc. Washington, 42:173, June 15, type from last slope of Talkeetna Mts., near headwaters of Oshetna or Black River, Alaska. 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS In the northern third of Alaska, males average 16 percent smaller in condylobasal length than in gyas of the Alaska Peninsula, and the one female available from northern Alaska is 18 percent smaller than the average in gyas on the Alaska Peninsula. The average difference in zygomatic breadth for males is 17 percent. Zygomatic breadth is not avail- able for the female from northern Alaska. The M2 is 8 percent shorter and 10 percent narrower in the males from northern Alaska. The length of M2 of the one female from northern Alaska (33.2) is near the minimum for the Alaska Peninsula (32.0) and width is 17.1, which is less than the minimum (17.7) for the Alaska Peninsula. The difference in measurements between specimens from northern Alaska and those of alascensis from the southern mainland of Alaska (including the Kenai Peninsula) east of the range of gyas is much less than between the northern sample and gyas, being nine percent smaller for males in condylobasal length, 11 smaller in zygomatic breadth, 4 shorter for M2, and 6 narrower for M2. The female from northern Alaska is 13 percent smaller in condylobasal length. The length of M2 of the female from northern Alaska (33.2) is near the minimum (28.5) for the southern mainland, and the width (17.1) is near the minimum (16.7) for the southern mainland. The change in size in alascensis appears to be gradual from north to south instead of abrupt at any particular latitude. Comparisons with U. a. dalli and U. a. horribilis are made in the accounts of those subspecies. MARGINAL RECORDS.-Alaska. Vi- cinity of Barrow, 1 MVZ; Colville River, 1 MVZ; Barter Island, 1; British Mts. between Barter Id. and Demarcation Pt., 1 NMC; Alaska- Yukon Boundary, about 50 mi. S of Arctic Coast (lat. 69°00'30"), 1 NMC; Kan- dik R., 40 mi. from junction with Yukon R., 1 MVZ; Chitina Glacier, 1; Mt. St. EUas, 1; Malaspina Glacier, 2, thence westward along coast, including Montague and Hinchinbrook islands, to Cape Douglas, 3; Bristol Bay, Kogguing, 1, thence westward along coast to Good News Bay, Kuskokwim Bay, 2, north- ward along coast to South Coast Range, Norton Sound, 1; Port Clarence, 1; Pit- megea River, Cape Sabine, 2 MVZ. Ursus arctos beringianus Middendorff Grizzly Bear 1853. Ursus arctos var. beringiana Middendorff, Sibir. Reise, 2, 2:4, pi. 1, figs. 1-6, type from Great Shantar Island, Sea of Okhotsk. Color pale brown (Geist, 1934:317): for cranial characters of this Asiatic subspecies, see Heptner and Naumov (1967). MARGINAL RECORD. - Alaska. St. Lawrence Island (Geist, 1934, Jour. Mamm., 15:316, November 15; A. H. HoweU, 1940, Jour. Mamm., 21:216, May 16; R. [L.] Rausch, 1953, The Murrelet, 34:19, Octo- ber 5). Ursus arctos californicus Merriam California Golden Bear 1896. [Ursus horribilis] subspecies californicus Merriam, Proc. Biol. Soc. Washington, 10:76, April 13, type from Monterey, Mon- terey Co., California. 1914. Ursus klamathensis Merriam, Proc. Biol. Soc. Washington, 27:185, August 13, type from Beswick, near mouth Shovel Creek, Klamath River, Siskiyou Co., California. 1914. Ursus colusus Merriam, Proc. Biol. Soc. Washington, 27:187, August 13, type from Sacramento River, probably between Colusa and Sacramento, California. 1914. Ursus californicus tularensis Merriam, Proc. Biol. Soc. Washington, 27:188, August 13, type from Fort Tejon, Tehachapi Mts., Kern Co., California. 1914. Ursus magister Merriam, Proc. Biol. Soc. Washington, 27:189, August 13, type from Los Biacitos, head San Onofre Canyon, Santa Ana Mts., San Diego Co., California. 1914. Ursus henshawi Merriam, Proc. Biol Soc. Washington, 27:190, August 13, type from southern Sierra Nevada, near Havilah, Kern Co., California. 1916. Ursus mendocinensis Merriam, Proc. Biol. Soc. Washington, 29:145, September 6, type SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 from Long Valley, N of Sherwood, Mendocino Co., California. Compared with U. a. stikeenensis directly to the north, the skull of calif ornicus aver- ages larger in condylobasal length, zygomatic breadth, depth of skull, and length and breadth of M^ in males. The same is true of females except that the depth of the skull is less. Comparison of californicus with horribilis of Utah (the geographic ranges of which may not ever have met in historic time in Nevada nor south thereof), reveals that males of californicus average larger in condylobasal length, average smaller in zygomatic breadth and depth of skuU, and average larger in length and breadth of M2. These average differences might disappear if more speci- mens were available. MARGINAL RECORDS. -California. E end Siskiyou, near Beswick, 1; Baird, Shasta County, 1; Los Biacitos, head San Onofre Canyon, Santa Ana Mts., San Diego County, L Baja California. Sierra Juarez near Santa Catarina Mission (Hall, 1981:953), thence up coast to California. 10 mi. from Blocksburg, Humboldt Co., Lassen Peak Canyon, 1. Ursus arctos dalli Merriam Brown Bear 1896. Ursus dalli Merriam, Proc. Biol. Soc. Washington, 10:71, April 13, type from Ya- kutat Bay (NW side), Alaska. 1914. Ursus nortoni Merriam, Proc. Biol. Soc. Washington, 27:179, August 13, type from Yakutat, Alaska. 1916. Ursus townsendi Merridim, Proc. Biol. Soc. Washington, 29:151, September 6, type from mainland of southeastern Alaska, probably between Cross Sound and Alsek River Delta, but exact locality unknown. 1918. Ursus orgiloides Merriam, N. Amer. Fauna, 41:46, February 9, type from Italio River, Alaska. Compared with males of U. a. alascensis of the southeastern mainland of Alaska, males of dalli average larger in condylobasal length and smaller in zygomatic breadth and are essentially the same in depth of skull; M^ averages shorter and narrower. Females of dalli average larger in con- dylobasal length and in zygomatic breadth but smaller in depth of skull; M^ averages shorter and broader. Compared with U. a. sitkensis, males of dalli average essentially the same in con- dylobasal length, zygomatic breadth, depth of skull, and length and breadth of M2. Compared with sitkensis, females of dalli average larger in condylobasal length and zygomatic breadth, and less in depth of skull; M2 is longer and broader than in sitkensis. Compared with U. a. horribilis from the Yukon, dalli differs as follows: males average larger in cranial measurements and smaller in length and breadth of M^; females average larger in all cranial measurements and in length and breadth of M^. MARGINAL RECORDS. -Alaska. Chaiks Hills, between Mt. St. Elias and Yakutat Bay, 1; N side Yakutat Bay, be- tween Dalton and Hubbard glaciers, 1; Hd. Disenchantment Bay, 2; Alsek River, near forks, 3; Dry Bay, mainland, 3 USNM, 1 MVZ; Alsek River, near coast, 1, thence up coast to Yakutat Bay region (Ankow River), 1 MVZ; Point Manby, N. Yakutat Bay, 1.— Italicized type for place names denotes omis- sion of symbols on the map in order to prevent overcrowding. Ursus arctos gyas Merriam Brown Bear 1902. Ursus dalli gyas Merriam, Proc. Biol. Soc. Washington, 15:78, March 22, type from Pavlof Bay, Alaska Peninsula, Alaska. 1902. Ursus merriami J. A. Allen, Bull. Amer. Mus. Nat. Hist., 16:141, April 12, type from Portage Bay, opposite Port Muller, Alaska Peninsula, Alaska. Skull averaging largest (Fig. 6) in con- dylobasal length and zygomatic breadth of the seven American mainland subspecies, and M2 averaging 10 percent longer in males and 8 percent longer in females than in the 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS population of alascensis from the northern third of Alasl^a. Comparison with middendorffi is made in the account of that subspecies. MARGINAL RECORDS. -Alaska (Alaska Peninsula). Kukak Bay, near Mt. Katmai, 1, southwestward along coast to Wide Bay, 1 MCZ; Morzhovoi Bay, 3; Un- imak Id., 2 skins only, collection in which preserved not recorded; Eagle Bay [Un- alaska Island], 2, thence northeastward to Isenbek Bay, 3; Port Heiden, 1 MCZ; Be- charofFLake, 1. Ursus arctos horribilis Ord Grizzly Bear or Silver Tip 1815. Ursus horribilis Ord, in Guthrie, A new geog., hist., coml. grammar . . . , Philadel- phia, 2nd Amer. ed., 2:291 (described on p. 299). Type locality, Missouri River, a little above mouth Poplar River, northeastern Montana. 1820. Ursus cinereus Desmarest, Mammalogie . . . , p. 164, in Encyclopedie methodique .... Type locality, not designated. Re- garded as a synonym of U. horribilis by Merriam, N. Amer. Fauna, 41:17, February 9, 1918. 1822. Ursus griseus Choris, Voyage pittoresque autour du monde, Paris (unpaged). Name applied by Choris to the bear of the interior of North America, but Choris identifies his ani- mal with Ursus griseus Cuv. Regarded as a synonym of U. horribilis by Merriam, N. Amer. Fauna, 41:17, February 9, 1918. 1827. Ursus candescens Hamilton-Smith, in Griffith's Cuvier, The Animal Kingdom . . . , 2:229 (fide Griffith, loc. cit.). Regarded as a synonym of U. horribilis by Merriam, N. Amer. Fauna, 41:17, February 9, 1918. 1838. Ursus richardsoni Swainson, Animals in menageries . . . , p. 54. Type locality as- sumed to be shore of the Arctic Ocean, on W side Bathurst Inlet about 8 mi. from mouth Hood River, Mackenzie. 1858. UfrsusJ. horribilis var. horriaeus Baird, Mammals, in Repts. Expl. Surv. . . . , 8(1): 224, July 14, type from old copper mines near present town of Santa Rita, Grant Co., New Mexico (see V. Bailey, N. Amer. Fauna, 53:357, March 1, 1932). 1904. Ursus hylodromus Elliot, Field Columb. Mus., Publ. 87, Zool. Ser., 3:257, January 7, type from Alberta. 1914. Ursus russelli Merriam, Proc. Biol. Soc. Washington, 27:178, August 13, type from Mackenzie Delta, Mackenzie. 1914. Ursus imperator Merriam, Proc. Biol. Soc. Washington, 27:180, August 13, type from Yellowstone National Park, Wyoming. 1914. Ursus absarokus Merriam, Proc. Biol. Soc. Washington, 27:181, August 13, type from head Little Bighorn River, northern part Big- horn Mts., Carbon Co., Montana. 1914. Ursus phaeonyx latifrons Merriam, Proc. Biol. Soc. Washington, 27:183, August 13, type from Jasper House, Alberta. 1914. Ursus Shoshone Merriam, Proc. Biol. Soc. Washington, 27:184, August 13, type from Estes Park, Larimer Co., Colorado. 1914. Ursus shoshone canadensis Merriam, Proc. Biol. Soc. Washington, 27:184, August 13, type from Moose Pass, near Mt. Robson, British Columbia. 1914. Ursus nelsoni Merriam, Proc. Biol. Soc. Washington, 27:190, August 13, type from Colonia Garcia, Chihuahua. 1914. Ursus horriaeus texensis Merriam, Proc. Biol. Soc. Washington, 27:191, August 13, type from Davis Mts., Jeff Davis Co., Texas. 1914. Ursus navaho Merriam, Proc. Biol. Soc. Washington, 27:191, August 13, type from Navajo country near Fort Defiance (Moll- hausen), Arizona, type probably killed in 1856 in Chuska Mountains, on boundary between northeastern Arizona and northwestern New Mexico. 1914. Ursus bairdi Merriam, Proc. Biol. Soc. Washington, 27:192, August 13, type fi-om Blue River, Summit Co., Colorado. 1914. Ursus utahensis Merriam, Proc. Biol. Soc. Washington, 27:193, August 13, type from N fork Salina Creek, about 10 mi. SE Mayfield, Sanpete Co., Utah. 1914. Ursus kennerleyi Merriam, Proc. Biol. Soc. Washington, 27:194, August 13, type from mts. near Los Nogales, Sonora. 1916. Ursus apache Merriam, Proc. Biol. Soc. Washington, 29:134, September 6, type from Whorton Creek, S slope White Mts., a few miles W Blue, Greenlee Co., Arizona. 1916. Ursus arizonae Merriam, Proc. Biol. Soc. SPECIAL PUBLICATION MUSEUM OF NATUPIAL HISTORY NO. 13 Washington, 29:135, September 6, type from east side Escudilla Mts., Apache Co., Ari- zona. 1916. Ursus kluane Merriam, Proc. Biol. Soc. Washington, 29:141, September 6, type from McConnell River, Yukon. 1916. Ursus ophrus Merriam, Proc. Biol. Soc. Washington, 29:148, September 6, type from eastern British Columbia; exact locality un- known. 1916. Ursus pallasi Merriam, Proc. Biol. Soc. Washington, 29:149, September 6, type from Donjek River, southwestern Yukon. 1916. Ursus Selkirk! Merriam, Proc. Biol. Soc. Washington, 29:150, September 6, type from Selkirk Mts., Upper Columbia River, British Columbia. 1916. Ursus washake Merriam, Proc. Biol. Soc. Washington, 29:152, September 6, type from N. Fork Shoshone River, Absaroka Mts., between Bighorn Basin and Yellowstone Na- tional Park, Wyoming. 1918. Ursus dusorgus Merriam, N. Amer. Fauna, 41:33, February 9, type from head Jackpine River, near Mt. Bess, close to British Colum- bian boundary. Alberta. 1918. Ursus planiceps Merriam, N. Amer. Fauna, 41:37, February 9, type from Colorado, exact locality unknown, but probably in foothills or on western edge of plains. 1918. Ursus macrodon Merriam, N. Amer. Fauna, 41:38, February 9, type from Twin Lakes, Lake Co., Colorado. 1918. Ursus mirus Merriam, N. Amer. Fauna, 41:40, February 9, type from Slough Creek, Yellowstone National Park, Wyoming. 1918. Ursus rungiusi rungiusi Merriam, N. Amer. Fauna, 41:49, February 9, type from Rocky Mts. on headwaters of Athabaska River, Alberta. 1918. Ursus rungiusi sagittalis Merriam, N. Amer. Fauna, 41:50, February 9, type from Champagne Landing, southwestern Yukon. 1918. Ursus macfarlani Merriam, N. Amer. Fauna, 41:51, February 9, type from Ander- son River, 50 mi. below Fort Anderson, Mac- kenzie. 1918. Ursus idahoensis Merriam, N. Amer. Fauna, 41:54, February 9, type from N fork Teton River, Fremont Co., Idaho. 1918. Ursus pulchellus pulchellus Merriam, N. Amer. Fauna, 41:55, February 9, type from Ross River, Yukon. 1918. Ursus pulchellus ereunetes Merriam, N. Amer. Fauna, 41:56, February 9, type from Beaverfoot Range, Kootenai Dist., British Columbia. 1918. Ursus oribasus Merriam, N. Amer. Fauna, 41:56, February 9, type from upper Liard River, Yukon, near British Columbian bound- ary. 1918. Ursus perturbans Merriam, N. Amer. Fauna, 41:64, February 9, type from a canyon on Mt. Taylor, 12 mi. E San Mateo, Valencia Co., New Mexico. 1918. Ursus rogersi rogersi Merriam, N. Amer. Fauna, 41:65, February 9, type from high up on Greybull River, Absaroka Mts., Yellow- stone National Park, Wyoming. 1918. Ursus rogersi bisonophagus Merriam, N. Amer. Fauna, 41:66, February 9, type from Bear Lodge, Sundance National Forest, Black Hills, Crook Co., Wyoming. 1918. Ursus kluane impiger Merriam, N. Amer. Fauna, 41:81, February 9, type from Colum- bia Valley, British Columbia. 1918. Ursus pellyensis Merriam, N. Amer. Fauna, 41:82, February 9, type from Ketza Divide, Pelly Mts., Yukon. 1918. Ursus andersoni Merriam, N. Amer. Fauna, 41:83, February 9, type from E branch Dease River, near Great Bear Lake, Mackenzie. 1918. Ursus crassus Merriam, N. Amer. Fauna, 41:90, February 9, type from upper Mac- millan River, Yukon. 1918. Vetularctos inopinatus Merriam, N. Amer. Fauna, 41:132, February 9, type from Ren- dezvous Lake, NE of Fort Anderson, Mac- kenzie. Skulls of males of horribilis from the Yukon, compared v^ith those of alascensis from east-central Alaska, averaging smaller in condylobasal length, zygomatic breadth and depth of skull, but M^ is longer and wider. In females the three cranial measure- ments average smaller, as in males, but the length and breadth of M^ average less. Skulls of three males of horribilis from northeastern British Columbia, compared with those of 26 specimens of U. a. stikeen- ensis from northwestern British Columbia, average larger in the five measurements mentioned above except for condylobasal 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS length, which averages smaller. The three females from northeastern British Columbia, compared with 23 specimens from north- western British Columbia, average smaller in all of the five measurements. Complete intergradation between sti- keenensis and horribilis is evident in south- central British Columbia, as shown by the adult male (USNM 215440) from Kamloops. Its dark but not brown color provides evi- dence of intergradation, as do the cranial measurements. Comparison with U. a. calif ornicus is made in the account of that subspecies. More specimens of both subspecies might reveal that the pelage of horribilis always was less brownish. Comparison with U. a. dalli is made in the account of that subspecies. MARGINAL RECORDS . -Mackenzie . Baillie's Cove, S end of Arctic Sound, Bathurst Inlet, 1 NMC; Stapylton Bay, Union Strait, 1 NMC; Kugaryuak River, Coronation Gulf, 4 NMC; "type locality [of U. richardsont] assumed to be shore of the Arctic Ocean, on W side Bathurst Inlet about 8 mi. from mouth Hood River" (Hall, 1981 956). Keewatin. Baker Lake (HaU, 1981 953). Mackenzie. Beaverhill Lake, 1 NMC Barren Grounds E of Great Slave Lake, 1 NMC; W tip of Aylmer Lake (109n3'W, 64°08'N), 3 NMC; Contwoyto Lake, 1 NMC; Copper Mines Valley between Great Bear and Dismal lakes, 1 MCZ; East Branch Dease River, near Great Bear Lake, 1 AMNH; Sekwi River, E side of McKenzie Mt., Canol Road, mi. 174E, 1 NMC. Al- berta. Smoky River, 5; N of Slave Lake, 1 U. Alberta; Red WiUow Creek, 1 U. Alberta. Saskatchewan. Found in Crane Lake when it dried up, pick up in May of 1937, 1 Old Timers Museum at Maple Creek; Dollard, 1 NMC. Minnesota. Sandhill River in southern Polk Co. (Hall, 1981:953). North Dakota. Ft. Clark, 1; near Middle Butte, now generally Bullion Butte, 1. Wyoming. Bearlodge of Sundance Natl. Forest, 1; Fort Laramie, 1; Lone Tree Canyon, between Hawk Springs and Chugwater, 1 Scotts Bluff Nat. Monu- ment Museum. Kansas. Trego Co. (Hall, 1981:953). Colorado, type locality of Ursus planiceps, 1; 12 mi. NE Saguache, 1. New Mexico. 25 mi. NE Taos Saw Mill Park, 1; Taos, 20 mi. SE on Rio Chiquito, 1; Magda- lena Baldy, 1. Texas. Davis Mts., 1. Coa- huila. vie. Cuatro Cienegas (Hall, 1981:953). Durango. Southern Durango (ibid.). Chihua- hua. Arroyo del Nido, 600 ft., 25 mi. SW Gallego, 1 MVZ; Colonia Garcia, 6. Sonora. Los Nogales, 1. Arizona. SW slope of Baldy Peak, 10,000 ft., near head Hurricane Creek, Apache Co., 1 MVZ; 8 mi. N Payson, near Green Valley, 1; 30 mi. S William, 1; San Francisco Mts., 1. Utah. Pine Valley Mts., 1; N Fork Salina Creek, about 10 mi. SE Mayfield, 2; Logan Canon, 1. Idaho. Minidoka, 1. Oregon. From dry bed of Mal- heur Lake, 1; South Ice Cave, 40 mi. S Bend, 1. British Columbia. Rossland, 1 MVZ; near Vernon, 1; Okanagan, 1; Shus- wap, 4; Canim Lake, 1; Slough Creek Mts., above timber, Barkerville District, 1 MVZ; Omineca River, 1 ANSP; Gundahoo Pass, 5000 ft., 1 MCZ. Yukon. Upper Liard River, near B. C. Boundary, 1; 50 mi. S White- horse, 2 ANSP. British Columbia. Rainey Hollow, 2. Yukon. Dalton House, 5; hd. Alsek River, 1; Duke River, Kluane, 4; Caldem Cr., 30 mi. E Mt. Natazat, hd. White River, 1; headwaters. White River, 1; Divide, White, Glacier and Tanana, Yukon side, 1; Ogilvie Range, Klondyke headwater, 1. Mackenzie. Foothills W of McKenzie River Delta, 1 NMC; Richards Island, Mac- kenzie Delta, 1 NMC. Ursus arctos middendorjfi Merriam Kodiak Brown Bear 1896. Ursus middendorffi Merriam, Proc. Biol. Soc. Washington, 10:69, April 13, type from Kodiak Island, Alaska. 1911. Ursus kadiakiYAemschmidi, Outdoor Life, 27:3, January. Name applied to the big brown bear of "Kadiak Island, Alaska Peninsula, Montague Island, and Yacutat," all in Alaska. Zygomatic breadth and depth of skull (Fig. 5) in both sexes from Kodiak Island average larger than in any other subspecies. 8 SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 U. a. middendorffi specimens from Afognak average smaller in both sexes (two adults of each sex) than those from Kodiak Island in the three cranial measurements except for condylobasal length which averages more. The M2 averages longer and narrower in middendorffi (Kodiak and Afognak islands) than in g^^as in both sexes. MARGINAL RE CORDS. -Alaska. Afog- nak Island, 7 (specimens of all ages including 3 KU); Kodiak Island, 116 (specimens of all ages including 44 AMNH, 4 ANSP, 3 Boone and Crockett specimens at Carnegie Mus., 7 CAS, 4 KU, 4 MCZ, 12 MVZ). Ursus arctos sitkensis Merriam Big Brown Bear 1896. Ursus sitkensis Merriam, Proc. Biol. Soc. Washington, 10:73, April 13, type from near Sitka, Alaska. 1904. Ursus eulophus Merriam, Proc. Biol. Soc. Washington, 17:153, October 6, type from Admiralty Island, Alaska. 1914. Ursus eltonclarki Merriam, Proc. Biol. Soc. Washington, 27:175, August 13, type from Freshwater Bay, Chichagof Island, Alaska. 1914. Ursus orgilos Merriam, Proc. Biol. Soc. Washington, 27:176, August 13, type from Bartlett Bay, E side Glacier Bay, south- eastern Alaska. 1914. Ursus caurinus Merriam, Proc. Biol. Soc. Washington, 27:187, August 13, type from Berners Bay, E side Lynn CanaJ, south- eastern Alaska. 1914. Ursus shirasi Merriam, Proc. Biol. Soc. Washington, 27:195, August 13, type from Bybus Bay, Admiralty Island, Alaska. 1916. Ursus eltonclarki insulans Merriam, Proc. Biol. Soc. Washington, 29:141, September 6, type from Admiralty Island, southeastern Alaska. 1916. Ursus kwakiutl neglectus Merriam, Proc. Biol. Soc. Washington, 29:144, September 6, type from near Hawk Inlet, Admiralty Island, Alaska. 1916. Ursus mirabilis Merriam, Proc. Biol. Soc. Washington, 29:146, September 6, type from Admiralty Island, Alaska. Compared with males of U. a. stikeenen- sis from immediately east in northwestern British Columbia, males of sitkensis average larger in condylobasal length, zygomatic breadth and depth of skull; M^ is shorter but broader. Females of sitkensis likewise average larger in condylobasal length, zygomatic breadth and depth of skuU; M^ is both shorter and narrower. Compared with males of stikeenensis from the area to the south (bounded by Bella Coola, Lagoon Lake, Kleena Kleene, and Ashlulm Creek), males of sitkensis average larger in condylobasal length, zygomatic breadth and depth of skull, but M^ averages shorter and narrower. Females of sitkensis are larger in all measurements except con- dylobasal length. U. a. sitkensis, when compared with all specimens referred to stikeenensis, averages larger in all measurements of the males, and in females averages larger in the three cra- nial measurements but smaller in length and breadth of M2. Comparison with U. a. dalli has been made in the account of that subspecies. MARGINAL RECORDS. -Alaska. Headwaters Bear Creek, 40 mi. from Haynes, 1 CAS; head of Chilkoot Lake, 1; Bemer's Bay, 5; Taku Inlet, mainland, 1; Sumdum, mainland, 1; Admiralty Island (al- legedly), 261 (including 3 AMNH, 5 ANSP, 1 CAS, 1 CM, 6 MCZ, 25 MVZ); Baranof Island, 78 (including 7 MVZ); Kruzof Island, 9; Chichagof Island, 170 (including 1 AMNH, 1 ANSP, 4 CM, 1 MCZ, 3 MVZ); NW side Lituya Bay, 2; Fairweather Glacier, about 15 mi. NW Lituya, 1; Porcupine, Chilkat, 1. From my (E. Raymond Hall's) Diary for August 1, 1962, written at the Division of Mammals, U.S. National Museum, Washing- ton, D.C., I quote: "At Museum E. P. Walker (Phone W06-5358) phoned having heard from his sister Winifred Deering of U. S. Fish and Wildlife Service, that I was in Washington D. C. He hoped I could nominate someone to get vol. 1 of his 4 vol. work on genera of Mammals ready for the printer. When I told him that I was here examining 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS skulls of bears with the aim of arriving at a better classification than the current one, Walker said let me tell you something that will help you 'Ignore all specimens obtained by Merriam from fur dealers and persons who sold skulls to Merriam. Dr. Merriam sent word, for example, to Mr. X in Alaska that skulls were wanted from Admiralty Is- land. Mr. X told the Indians that bear skulls were wanted from Admiralty Island. The Indian hunters brought the skulls to Mr. X and told him that the skulls were from Admi- ralty Island. No wonder that four kinds of bears were recorded from Admiralty Island that had close relatives on the mainland. After skulls from Admiralty Island had been obtained. Dr. Merriam sent out word that skulls were wanted from the mainland. The fur dealers told the Indians and hunters and the skulls that came in were all from the mainland, according to the hunters, regard- less of where the bears were shot.' " Ursus arctos stikeenensis Merriam Big Brown Bear 1914. Ursus stikeenensis Merriam, Proc. Biol. Soc. Washington, 27:178, August 13, type from Tatletuey Lake, near head Skeena River, northern British Columbia. 1914. Ursus tahltanicus Merriam, Proc. Biol. Soc. Washington, 27:181, August 13, type from Klappan Creek (" = 3d So. Fk. Stikine River"), British Columbia. 1914. Ursus pervagor Merriam, Proc. Biol. Soc. Washington, 27:186, August 13, type from Pemberton [ = Lillooet] Lake, British Colum- bia. 1916. Ursus chelan Merriam, Proc. Biol. Soc. Washington, 29:136, September 6, type from T.30N, R.16E, Willamette Meridian, Wenat- chee National Forest, E slope Cascade Mts., northern Chelan Co., Washington. 1916. Ursus hoots Merriam, Proc. Biol. Soc. Washington, 29:140, September 6, type from Clearwater Creek, a N branch Stikine River, British Columbia. 1916. Ursus kwakiutl Merriam, Proc. Biol. Soc. Washington, 29:143, September 6, type from Jervis Inlet, coast of southern British Colum- bia. 1916. Ursus kwakiutl warburtoni Merriam, Proc. Biol. Soc. Washington, 29:145, September 6, type from Atnarko River, British Columbia. 1918. Ursus chelidonias Merriam, N. Amer. Fauna, 41:21, February 9, type from head Jervis Inlet, British Columbia. 1918. Ursus atnarko Merriam, N. Amer. Fauna, 41:22, February 9, type from Lonesome Lake, Atnarko River, one of upper forks of Bella Coola, British Columbia. 1918. Ursus crassodon Merriam, N. Amer. Fauna, 41:90, February 9, type from Klappan Creek (Third South Fork Stikine River), Brit- ish Columbia. Comparison of stikeenensis has been made with three geographically adjoining subspecies, californicus, horribilis, and sit- kensis in the accounts of those three sub- species. MARGINAL RECORDS. -British Co- lumbia. Dease River, Cassiar Range, 2; Tat- letuey Lake, near head Skeena River, 1; Bear River, Bear Lake, 1 ANSP; Tacla Lake, 6; "Big Creek, B.C." but printed on front of label is "Williams Lake, Cariboo BC," 1 NMC; Kamloops, 1. Washington. Holman Pass, Hd. Holman Creek, trib. of W fk. Pasayton River, 1; Twp. 30N, range 16E, WM, Chelan Co., 1. Presumably to coast of Oregon and presumably up coast of Washing- ton to British Columbia. Nass River, 1 U. Alberta. Alaska. Burroughs Bay, Unuk River, 1; Bradfield Canal, 1; Groundhog Basin, 8 mi. S mouth Stikine River, 3. British Columbia. Stikine River, 1 mi. above bound- ary, 1; Stikine River, 50 mi. N boundary, 1; Shesley River, 45 mi. N Telegraph Creek, 4; 12 mi. NE Tulsequah on Taku River at entrance to Zohini Creek, 1 Boone and Crockett specimen at Carnegie Mus.; Ben My Chree, Tagish Lake, 1 CAS; Atlin, 1; 60 mi. E Atlin, 1. 10 SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 Table l. Average, minimum, and maximum cranial and dental measurements of Ursus ardos. No. of specimens examined Condylobasal Length Zygomatic Breadth Depth of Skull* Length M2 Breadth M2 No. Specimens and Institution** 11 a 333 220 124 35.0 18.5 3 MVZ: 2 NMC; 284 182 109 29.8 17.2 1 KU 372 239 144 37.5 20.1 1 5 2a,'> — 106 33.2 17.1 1 NMC U. A. alascensis from east-central Alaska 22 ' ' 346 232 129 36.7 19.2 2 AMNH; 1 ANSP; 313 205 112 32.6 17.7 1 MVZ 382 260 158 43.4 22.0 17 .- : 298 192 109 36.1 17.9 2 AMNH; 1 ANSP; 280 182 103 32.4 16.0 1 MCZ 311 210 129 39.5 18.9 U. a. califomicus 14 J ' 365 227 129 38.0 20.2 1 MCZ; 1 MVZ; 339 203 115 35.5 19.6 1 ANSP 388 240 149 40.5 21.7 5; . 319 194 108 36.3 19.1 1 MVZ; 1 CAS 312 190 103 35.5 18.0 330 197 111 37.0 20.1 U. a. dalli 11 ' ' 359 238 130 35.4 18.7 1 MCZ 327 203 114 31.4 16.6 396 270 156 40.0 21.4 699 328 206 110 34.2 18.2 1 MVZ 309 192 103 31.1 17.8 342 217 113 36.2 18.8 U. a. gyas 47 iS 397 264 152 38.1 20.6 7 AMNH; 3 MVZ; 362 201 126 33.0 18.8 3 MCZ; 3 B&C 424 296 173 42.3 22.1 25 ■■ 346 217 123 34.9 19.3 5 AMNH; 1 MVZ; 322 206 110 32.0 17.7 1 MCZ 366 240 133 37.4 21.6 U. a. horribilis from Yukon 56 ' ' 332 216 123 37.2 19.4 3 KU; 2 NMC; 303 197 110 33.3 17.3 1 ANSP 360 241 143 45.9 23.0 35 = . 291 184 106 33.7 17.7 1 NMC; 1 MCZ; 270 165 98 29.0 15.7 1 ANSP 311 212 119 37.7 19.8 U. a. horribtlts from NE British Columbia 3 ii 338 223 128 37.9 19.0 332 215 126 36.9 18.2 343 228 130 39.9 19.8 3 99 303 182 108 33.2 17.5 1 MCZ 297 178 104 31.6 17.1 315 188 113 35.2 17.9 U. a. middendotjfi from Kodiak Island 31 (5 i 393 284 164 38.2 20.0 8 AMNH; 3 B&C; 350 243 134 34.7 17.9 3 CAS; 2 KU; 415 319 189 42.5 21.8 1 T. Dolan, III 16 vv 335 227 128 35.2 18.3 6 AMNH: 1 MVZ; 314 209 119 33.3 17.6 1 CAS; 1 KU; 363 243 142 37.9 19.4 1 MCZ U. a. sttkensis from the area circumscribed by marginal records provisionally mapped as sttkensis 113' ' 359 240 131 35.7 18.8 5 CM; 5 MVZ; 318 198 110 30.5 16.5 1 AMNH; 1 ANSP 398 270 160 40.3 20.5 94 99 313 200 116 33.6 17.5 6 MVZ; 2 AMNH; 283 173 101 28.8 15.0 2 MCZ 341 223 126 36.6 20.8 U. a. stikeenensts from NW British Columbia 26 i ' 340 222 123 36.2 18.7 2 CAS; 1 AMNH; 302 193 111 31.6 16.5 1 MVZ 372 259 137 42.6 21.7 23 99 304 190 111 34.4 17.8 2 AMNH; 1 NMC 285 175 102 31.3 16.6 319 205 120 38.0 19.0 Measured from sphenoid floor of braincase above posterior margin of palate to highest point on braincase, *• Remaining speamens are from USNM. LITERATURE CITED AND SOME OTHER PERTINENT LITERATURE NOT IN THE PRECEDING SYNONYMIES BOBRINSKII, N. A., B. A. KUZNETSOV, and A. P. KUZYAKIN. 1965. Opredelitel' mlekopitayushchikh SSSR. [Key to the mammals of the USSR.] "Prosveshchenie," Moscow, 382 pp., 40 color plates. 111 maps. Couturier, M. A. J. 1954. L'ours brun, Ursus arctos L. Printed by the author— Rue Thiers, Grenoble (Isere, France), pp. xiii + 907, 82 pis., 49 figs. Geist, 0. W. 1934. Brown bear seen on St. Lawrence Island. Jour. Mamm., 15:316-317, November 15. Hall, E. R. 1981. The mammals of North America (2nd ed.). John Wiley & Sons , Inc . , Ne w York , 1 : xvii + 600 + 90, and 2:vii + 601-1181 + 5>C, 1367 illustrations, April 3. Hall, E. R., and K. R. KELSON. 1959. The mammals of North America. Ronald Press Co., New York, l:xxx + 546+79, and 2:ix + 547-1083+ Z9, 1231 illustrations, March 31. Harrison, D. L. 1968. The mammals of Arabia. Vol. 2. Camivora, Artiodactyla, Hyracoidea. Ernest Benn Lim- ited, London, pp. xiv + 193-381, pis. 60-128, figs. 95-169, tables 107-174, April 24. Hassinger, J. D. 1973. A survey of the mammals of Afghanistan. . . . Fieldiana: Zoology, 60:xi + 195, illus- trated, April 6. Hatt, R. T. 1959. The mammals of Iraq. Miscl. Publ. Mus. Zool., Univ. Michigan, 106:1-113, 6 pis., 1 map, 6 tables, February 12. Heptner, V. G., and N. P. Naumov (eds.). 1967. Mlekopitayushchie Sovetskovo Soyuza [Mammals of the Soviet Union], vol. 2, pt. 1, "Vysshaya Shkola," 1003 pp. Howell, A. H. 1940. Brown bear killed on St. Lawrence Island. Jour. Mamm., 21:216, May 16. KURT£n, B. 1973. Transberingian relationships of Ursus arctos Linne (brown and grizzly bears). Commentationes Biologicae, 65:1-10, 2 figs., 4 tables, June. Lay, D. M. 1967. A study of the mammals of Iran, resulting from the Street Expedition of 1962-63. Fieldiana: Zool- ogy, 54:1-282, 32 figs., 1 table, October 31. Merriam, C. H. 1918. Review of the grizzly and big brown bears of North America (genus Ursus) with description of a new genus, Vetularctos. North Amer. Fauna, 41:1-136, 16 pis., February 9. Merriam, C. H. 1919. Criteria for the recognition of species and genera. Jour. Mamm., 1:6-9, November 28. Merriam, C. H. 1919. Why should every specimen be named? Jour. Mamm., 1:41-42, November 28. Merriam, C. H. 1929. Ursus holzworthi, a new grizzly from the Talkeetna Mountains, Alaska. Proc. Biol. Soc. Wash- ington, 42:173-174 + 4 pis., June 15. Rausch, R. 1953. On the land mammals of St. Lawrence Island, Alaska. The Murrelet, 34(2):18-26, 4 tables, October 5. Rausch, R. L. 1963. Geographic variation in size in North American brown bears, Ursus arctos L., as indicated by condylobasal length. Canadian Jour. Zool., 41:33-45, 2 figs., 3 tables. Shou, Zhen-HUANG (editor). 1964. Zhong Guo Jing Ji Dong Wu Zhi. [Records of Economic Mammals of China.] Scientific Publications Oflice, pp. xiii + 554, 153 figs., 138 maps, plus 72 pis. Simpson, G. G. 1980. Splendid isolation. The curious history of South American mammals. Yale University Press, pp. ix + 266, 43 figs., 14 tables. SOKOLOV, V. E., and V. N. Orlov. 1980. Opredelitel' mlekopitayushchikh Mongol'skoi Narodnoi Respubliki. [Key to the mammals of the Mongolian Peoples Republic] "Nauka," Moscow, 351 pp., 51 maps, 137 figs. Zavatsky, B. p. 1976. The use of the skull in age determina- tion of the brown bear. Pp. 275-279 in M. R. Pelton, et al., editors. Bears— their biology and management. lUCN Publications new series No. 40. 11 12 SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 Ursus maritimus Polar Bear X »♦•. /•;-/: ■ 1 .'.•» ■'^ /'J ■/..'.; '///,;7,:'. •■'■■ y/^ Ursus arctos-group Big Brown and Grizzly Bear ..ifl^m-:m^'~-^;r-^^ x'is 1^ Ursus americanus Black Bear Figure l. The Holarctic and Nearctic bears (genus Ursus). 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS 13 U. a beringianus- FlGURE 2. Subspecies of Ursus arctos in North America. 14 SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 60 30 60 0 30 60 90 120 150 Figure 3. Geographic range of Ursus ardos in Eurasia at about 1400 B. C. 1984 NORTH AMERICAN BROWN AND GRIZZLY BEARS 15 Figure 4. Geographic range of Ursus arctos in North America and adjoining part of Asia at about 1400 B.C. 16 SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 13 Figure 5. Ursus arctos middendorffi Merriam, C. H., 1896:69. Collected by Ross Beach. ' ad. No. 5355 Fort Hays Kansas State College Collection of Vertebrates. From Upper Spiridon Lake, Kodiak Island, Alaska, x Va natural size (condylobasal length of skull is 395 mm). Figure 6. Ursus arctos gyas Merriam, C. H., 1902:78. Collected by Colonel Colby. ^ ad. No. 135502 Amer. Mus. Natural Hist. From Canoe Bay, Alaska Peninsula, x Vs natural size (condylobasal length of skull is 398 mm). QL737.C27 H355 1984 ('.io..r..plMc uin.ih.m .inunm Incwil ■'" Harvard MCZ Library \(,l i-(. 3 2044 062 384 714 Date Due AVAILABLE SPECIAL PUBLICATIONS MUSEUM OF NATURAL HISTORY, UNIVERSITY OF KANSAS 1. Catalogue of publications in herpetology published by the University of Kansas Museum of Natural History. By Linda Trueb. Pp. 1-15. December 1976. 2. Catalogue of publications in mammalogy published by the University of Kansas Museum of Natural History. By Robert S. Hoffinann. Pp. 1-19. 15 February 1977. 3. Maintenance of rattlesnakes in captivity. By James B, Murphy and Barry L. Armstrong. Pp. 1-40. 29 December 1978. 5. The natural history of Mexican rattlesnakes. By Barry L. Armstrong and James B. Murphy. Pp. 1-88. 14 December 1979. 7. A diapsid reptile from the Pennsylvanian of Kansas. By Robert R. Reisz. Pp. 1-74. 18 February 1981. 8. 1982 Catalog of publications of the University of Kansas Museum of Natural History. Pp. 1-28. November 1982. 9. The ecological impact of man on the South Florida herpetofauna. By Larry David Wilson and Louis Porras. Pp. 1-89. 8 August 1983. 10. Vertebrate ecology and systematics: A tribute to Henry S. Fitch. Edited by Richard A. Seigel, Lawrence E. Hunt, James L. Knight, Luis Malaret and Nancy L. Zuschlag. Pp. 1-277. 21 June 1984. 11. Museum of Natural History, The University of Kansas, Annual Report 1983. By Gary McGrath. Pp. 1-48. June 1984. 12. Principles and methods of phylogenetic systematics: A cladistics workbook. By Daniel R. Brooks, Janine N. Caira, Thomas R. Piatt and Mary R. Pritchard. Pp. 1-92. 20 April 1984. ^