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Full text of "Geographic variation among brown and grizzly bears (Ursus arctos) in North America"

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 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- 



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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 



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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. 



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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. 



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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. 



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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 







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 

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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. 



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