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with the 


Special Publication No. 11 

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O very creature is better alive than dead, 
men and moose and pine-trees, and he 
who understands it aright will rather 
preserve its life than destroy it. 

The Maine Woods 















W. REID BLAIR, Secretary 
New York Zoological Park 


HAROLD J. COOLIDGE, Jr., Chairman 





San Diego Society of Natural History Museum of Zoology, Univ. of Michigan 


Museum of Comparative Zoology Carnegie Museum 


California Academy of Sciences Washington University, St. Louis 


Acad. Nat. Sciences of Philadelphia Chicago Zoological Society 


Camp Fire Club of America California Institute of Technology 


American Society of Mammalogists New York Zoological Society 


Field Museum of Natural History National Audubon Society 

Philadelphia Zoological Society Wilderness Club 


WlL * IAM 

American Museum of Natural History 

Boone and Crockett Club 


Museum of Vertebrate Zoology FREDERIC C. WALCOTT 

American Wildlife Institute 


Allan Hancock Foundation, ALEXANDER WETMORE 

University of Southern California Smithsonian Institution 

* Deceased 


AY 1, 1942, is likely to be a significant date in the history of con- 
servation, for on that day the "Convention on Nature Protec- 
tion and Wildlife Preservation in the American Republics " came into 
effect for the seven American countries, including the United States, 
that had deposited ratifications with the Pan American Union. 
Twelve other Republics have now signed the Treaty, and it is hoped 
that their ratifications are soon to follow. 

Publication of the present volume, devoted to the extinct and 
vanishing mammals of the New World, including certain marine 
mammals of all the oceans, is therefore particularly timely. Its 
author, Dr. Glover M. Allen, of the Museum of Comparative Zoology 
at Harvard University, who died on February 14, 1942, was one of 
the great naturalists of his day. His profound knowledge of mammals 
and birds was the result not only of a life-time study in museums but 
also of intimate observation of animals in their native habitats in 
Africa, Australia, the Caribbean, and South America, as well as in 
many parts of his own country. The final preparation of this report 
was one of the last projects that Dr. Allen completed before his un- 
timely death. We know that he thought of it as one might think of 
the clearing of a new trail into a virgin forest. It will, indeed, be the 
opening of the way to a wider understanding of the genuine need for 
international cooperation, if the vanishing wildlife of the Americas 
is to be preserved for the benefit of future generations. 

This volume is dedicated to Dr. Allen's friend Dr. John C. Phillips, 
founder and first chairman of the American Committee for Interna- 
tional Wildlife Protection. It was largely due to Dr. Phillips's vision 
that in May 1936 a project for a comprehensive study of the recently 
extinct and vanishing mammals of the world was undertaken under 
the auspices of the American Committee. Information on this sub- 
ject had never been assembled, and it was felt that such a report, if 
carefully prepared, would be invaluable to the work of the Committee 
in helping to determine those species of mammals most urgently in 
need of protection and, at the same time, to estimate factors that 
might have caused the extinction of species. Basic information was 


needed to substantiate proposals for the protection of vanishing 
species in their native habitat through the establishment of adequate 
national parks and reserves. The material for the preparation of 
such a report was widely scattered, and assembling and evaluating it 
involved a great deal of correspondence with zoologists, game 
wardens, nature protection societies, and governments in many parts 
of the world, as well as a survey of widely scattered articles in popular 
magazines, books on sport and travel, and scientific literature. 

Dr. Francis Harper, of Philadelphia, under arrangements made by 
Dr. Phillips, undertook this work and devoted many months of care- 
ful research to the preparation of material on the Old World mammals, 
treating in his report (as yet unpublished) about 400 forms. Dr. 
Allen made use of a special bibliography of references to literature 
collected by Dr. Harper, and with this as a basis prepared the present 
work on the New World and marine mammals, assembling much 
additional material to complete it. 

The Harper-Allen study could never have been brought to its 
present state of completion had it not been for the most generous 
financial assistance received in the form of gifts from many individuals 
over a period of seven years, a special grant from the Penrose Fund 
of the American Philosophical Society (1937, No. 195), and a publi- 
cation grant from the Boone and Crockett Club and the Conservation 
Committee of the New York Zoological Society. The American 
Committee takes this opportunity to express its gratitude to Dr. 
Allen and to Dr. Harper for their months of work in the preparation 
of material for reports on recently extinct and vanishing mammals 
of the world. We wish also to thank the forty financial contributors 
of information from many parts of the world. We are indebted 
further to the Academy of Natural Sciences of Philadelphia, whose 
famous library proved of inestimable assistance, and to the Museum 
of Comparative Zoology, at Cambridge, for the use of its library and 
other facilities. Thanks are due also to Dr. John Eric Hill, of the 
American Museum of Natural History, for his contributions to Dr. 
Allen's volume, and to Mr. Paul H. Oehser, editor of the United 
States National Museum, for the editorial supervision of this volume 
through the press and for the preparation of the index. The draw- 
ings for the illustrations were made especially for this work by Mr. 
Earl L. Poole, of the Reading Public Museum and Art Gallery. 

In these dark days, when even man is fighting to save himself from 
extinction, let us hope that this book, inspired by a great sportsman 
and conservationist and written by a great naturalist "whose gentle- 


ness and purity of spirit were beyond all praise, " will in some measure 
be helpful in promoting a wider understanding of the need for pre- 
serving the wildlife heritage of the American republics for the enjoy- 
ment and wise use of generations to come. "Natural beauty," in 
the words of Trevelyan, "is the highest common denominator in the 
spiritual life of today. " And what a great destiny lies ahead for the 
peoples of the Americas if, in winning their struggle for world free- 
dom, they keep faith with their birthright of natural beauty, as well 
as with the wild creatures over which they have dominion and with 
the earth that sustains them! 


(for the Committee) 
Washington, D. C. 
June 12, 1942 

Publication Committee: 







Order INSECTIVORA : Moles, shrews, and their relatives 5 

Family Nesophontidae : Extinct Antillean insectivores 5 

Family Solenodontidae : Solenodons 10 

Family Soricidae : Shrews 14 

Order CHIROPTERA: Bats 15 

Family Phyllostomidae : Leaf-nosed bats 16 

Family Natalidae : Long-legged bats 29 

Family Vespertilionidae : Simple-rosed bats 30 

Order EDENTATA: Edentates 34 

Family Dasypodidae: Armadillos 34 

Family Megalochnidae : Ground sloths 36 

Order RODENTIA: Gnawing mammals 41 

Family Sciuridae: Squirrels 42 

Family Castoridae : Beavers 49 

Family Cricetidae : Hamsterlike rats and mice 87 

Family Echimyidae : Spiny rats and their relatives 99 

Family Heptaxodontidae 125 

Family Dinomyidae: Giant rodents 126 

Family Dasyproctidae : Agoutis 128 

Order CARNIVORA : Dogs, cats, and their relatives 134 

Family Ursidae : Bears 135 

The black bears 135 

The grizzly bears 139 

Family Mustelidae : Weasels and martens 165 

The pine martens 165 

Family Canidae: Wolves, dogs, foxes 194 

The kit foxes 194 

The American wolves 199 

Family Felidae: Cats 233 

The pumas 234 

The jaguars 252 




Order ARTIODACTYLA : Even-toed ungulates 256 

Family Cervidae: Deer 257 

The wapiti, or American elk 257 

The caribou 296 

Family Antilocapridae : Pronghorns 322 

The pronghorn antelope 322 

Family Bovidae : Sheep, goats, cattle 329 

The muskoxen 329 

The American bison 337 

The bighorn, or mountain sheep 349 


Order EDENTATA: Edentates 375 

Family Megatheriidae : Giant ground sloths 375 

Order RODENTIA : Gnawing mammals 377 

Family Cricetidae : Hamsterlike rodents 377 

The Galapagos rice rats 377 

Family Myocastoridae : Coypus 383 

Family Dinomyidae: Giant rats 386 

Family Chinchillidae : Chinchillas 389 

Order CARNIVORA: Dogs, cats, and their relatives 396 

Family Ursidae: Bears 396 

Family Canidae: Wolves, dogs, foxes 400 

Order PERISSODACTYLA : Odd-toed ungulates 403 

Family Tapiridae : Tapirs 404 

Order ARTIODACTYLA : Even-toed ungulates 406 

Family Camelidae: Camels, llamas 406 

Family Cervidae : Deer 412 


Order CARNIVORA: Dogs, cats, and their relatives 417 

Family Mustelidae: Weasels, martens, otters 417 

Order PINNIPEDIA: Seals, sea-lions, walruses 424 

Family Otariidae: Sea-lions, fur seals 425 

Family Phocidae : Hair seals 447 

Family Odobenidae : Walruses 469 

Order CETACEA: Whales, porpoises, dolphins 477 

Family Iniidae: Fresh-water dolphins 479 

Family Delphinidae : Dolphins and porpoises 480 

Family Ziphiidae: Ziphioid whales 481 

Whaling 484 

Family Physeteridae : Sperm whales 492 

Family Rhachianectidae : Gray whales 495 


Order CETACEA Continued. 

Family Balaenidae : Right whales 499 

Family Balaenopteridae : Fin whales 513 

Order SIRENIA : Sea-cows 528 

Family Dugongidae: Dugongs 528 

Family Hydrodamalidae 535 

Family Trichechidae : Manatees 538 


INDEX . .589 



Hispaniolan solenodon (Solenodon paradoxus] 11 

Jamaican hutia (Geocapromys brownii) 108 

St. Vincent agouti (Dasyprocta albida) 131 

Kermode's bear (Euarctos americanus kermodei) 138 

Barren-ground bear (Ursus richardsoni) 164 

Eastern pine marten (Maries americana americana) 173 

Fisher (Maries pennanti) 180 

Black-footed ferret (Mustela nigripes) 185 

Long-eared kit fox (Vulpes macrotis macrotis) 198 

Eastern wapiti (Cervus canadensis canadensis) 264 

Newfoundland caribou (Rangifer terraenovae) 321 

White-faced muskox (Ovibos moschatus wardi) 335 

Wood bison (Bison bison athabascae) 348 

Badlands bighorn (Ovis canadensis auduboni) 360 

Peruvian chinchilla (Chinchilla lanigera brevicaudata) 395 

Antarctic wolf (Dusicyon australis) 402 

Vicuna (Vicugna vicugna) 411 

Sea otter (Enhydra lutris) 423 

West Indian seal (Monachus tropicalis) 455 

Northern elephant seal (Mirounga angustirostris) 461 

Lesser rorqual (Balaenoptera borealis); humpback whale (Megapiera 
novaeangliae) ; Greenland whale (Balaena mysticetus) ; California 

gray whale (Rhachianectes glaucus) 512 

Australian dugong (Dugong australis) 531 

Steller's sea-cow (Hydrodamalis stelleri) 537 

Manatee (Trichechus manatus) 543 


THE accounts that follow are intended to cover, first, those 
species of New World mammals that have been extermi- 
nated or seriously depleted by human agency during the past 
four and a half centuries; and second, the oceanic species of 
world-wide distribution, mainly cetaceans, seals, and sea-lions, 
that within the historic period have suffered similar reduction, 
some of them over a longer term. The method followed is to 
give the common name, the current Latin name, reference to 
original description with type locality, the important syno- 
nyms, and references to figures of the exterior or the skull, or 
both. Following a brief description there are given the geo- 
graphic range past and present said a short description of each 
species from the viewpoint mainly of its economic or com- 
mercial use, present status, protective measures, and other 
facts pertinent to a better understanding of its control or 
encouragement or other special needs. The introductory para- 
graphs to each of the major groups were prepared by Dr. J. 
Eric Hill, of the American Museum of Natural History, and 
are signed with his initials. 

Full acknowledgment is due Dr. Francis Harper for the large 
share of the bibliographic work for the present volume that he 
had already accomplished in connection with his work on the 
Old World species preliminary to the preparation of the final 
accounts, and which he has handed on to me. In laying out 
the work Dr. Harper selected for inclusion those mammals 
that have been wholly or in part extirpated or have been greatly 
restricted in range or numbers by clearing and settlement or 
by intensive hunting for food, fur, or other economic and com- 
mercial purposes. In a search through literature and by ex- 
tensive correspondence Dr. Harper had already accumulated 
a considerable mass of information concerning the past and 
present status of many of these species, and of this I have made 


full use. To the list of species he selected I have added a few 
others that, as the work progressed, seemed to require treat- 
ment. Possibly others should have been included, such as the 
three-banded armadillo, which though still found in parts of 
Brazil, seems now to have an interrupted range and may have 
been locally exterminated for food. Possibly, too, various 
species of mammals have been in part exterminated by pre- 
historic peoples, if the evidence now accumulating is to be 
trusted that primitive weapons found in association with 
skeletons of extinct species of bison and mammoth in parts of 
our West are proof of the killing of some of the survivors of 
these species by human contemporaries. There is some evi- 
dence, though of a rather uncertain nature, that in southern 
South America prehistoric man was contemporary with species 
of ungulates, now extinct, the remains of which date from the 
Pampean time of late Pleistocene. Such animals have been 
omitted, however, since the present inquiry does not extend so 
far back into the past. 

Although few modern species of New World or marine 
mammals have been directly exterminated by man, it is never- 
theless true that almost any of those having a considerable 
value for food, fur, oil, or other products, as well as those 
having a less tangible interest as game animals for trophies, 
are likely to be endangered increasingly with the more intensive 
methods of modern times and with the growth of populations 
and consequent reduction of areas available for their support. 
With the settlement of this hemisphere by Europeans, the 
control measures they imposed on the native mammals were 
naturally directed first against the larger predators wolves, 
pumas, bears that menaced their meager flocks or even at 
times endangered their own lives. Clearing of forests for 
agricultural purposes had an immediate effect in reducing the 
available food and shelter for sylvan species and thus indirectly 
in driving out or eliminating some of these. Probably this 
factor has been a potent one in the extinction of various fruit- 
eating bats of the West Indian islands. The growing demands 
of the fur trade, with its increasing inroads on the fur bearers, 
have resulted in great reduction of native stocks in many re- 
gions, even those remote from civilization where professional 
trappers have for long periods pursued the more valuable 
species unremittingly. The large game animals, which fur- 


nished dependable supplies of meat and hides, were drawn 
heavily upon during pioneer stages and later were intensively 
pursued, in part for sport and in part for incidental food pur- 
poses. Finally the oceanic species, particularly the larger 
cetaceans and seals, have been hunted with increasing vigor 
and improved devices the world over, until at the present day 
many of the regions formerly productive now no longer repay 
the effort or expense of pursuit. 

Protective legislation in aid of certain species of commercial 
or economic importance, began early, as in colonial New 
England, but in most cases did not come till much later, some- 
times too late to be of much avail. On the other hand, there 
have been some mammals that have proved more adaptive or 
have responded more quickly to a release of the pressure put 
upon them, and with proper management may prove a de- 
pendable asset for a long time to come. Any facts that will 
serve to bring out the needs of those species worth conserva- 
tional treatment should be of value as a guide to future manage- 
ment of these natural resources. The last decade has seen much 
progress in these matters and a wider awareness that the time 
has come when immediate steps <are necessary to preserve such 
assets before their depletion has gone too far. 

A long step forward in the preservation and protection of 
American wildlife was made when, on October 12, 1940, 
through the agency of the Pan American Union, representa- 
tives of the nations of the Western Hemisphere affixed their 
signatures to the Convention on Nature Protection and Wild- 
life Preservation in the American Republics, thus pledging 
their countries "to adopt measures for the protection of useful, 
harmless and ornamental species of plant and animal life. 
They have thus given formal recognition to the fact that many 
such species know no national boundaries, and that true con- 
servation of the gifts of Nature should begin before these 
resources have been dissipated by thoughtless or selfish de- 
struction" (from the statement of Dr. Hector David Castro, 
Minister of El Salvador). The Senate of the United States 
ratified this convention on April 7, 1941. In the "Annex" to 
the Convention, three mammals are included the protection 
of which is declared to be of special urgency and importance in 
the United States and is to be as nearly complete as possible, 
namely, the woodland caribou, the sea otter, and the manatee 


(see Congressional Record Senate, for April 7, 1941). Similar 
lists are to be submitted by the other countries. 1 

February, 191+2. 

1 For a full account of this and other related matters see HAYDEN, SHERMAN STRONG, 
The International Protection of Wild Life, an examination of treaties and other agree- 
ments for the preservation of birds and mammals, 246 pp. Columbia University Press, 


Order INSECTIVORA: Moles, Shrews, and Their Relatives 

T NSECTIVORES are primitive mammals of moderate or 
1 small size, with long pointed muzzles and sharp-cusped 
chewing teeth. They feed chiefly on insects and other inverte- 
brates, but some prey on small mammals or fish. The New 
World insectivores belong to the suborder Lipotyphla and 
include two groups differing in important dental characters. 

The archaic insectivores, with V-shaped molar teeth, are 
represented in the West Indies by two families; all are either 
extinct or threatened with extinction: 

(1) Nesophontidae, five primitive species from the islands of 
Puerto Rico, Haiti, and Cuba. These are extinct, so far as 

(2) Solenodontidae, two species of long-snouted heavy -bodied 
insectivores, found in the islands of Haiti and Cuba. 

The more modern insectivores, with W-shaped molar teeth, 
are represented by two families, both of which are widely 
distributed in the Old World and North America, but only one 
extends into Central or South America: 

(3) Talpidae, moles. None of these are discussed here. 

(4) Soricidae, shrews. A single species is considered to be 
in danger of extinction because of rarity. J. E. H. 

Family NESOPHONTIDAE: Extinct Antillean Insectivores 

Nesophontes edithae Anthony, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 725, 1916 

("Cueva Catedral, near Morovis, Porto Rico"). 
FIGS.: Anthony, H. E., 1916, pi. 23 (cranium); 1918, pp. 366-375, figs. 17-23 (skull 

and larger bones). 

Our first knowledge of this remarkable genus and species is 
due to Dr. H. E. Anthony, who, during his investigation of cave 
deposits in Puerto Rico, brought to light abundant remains of 



the species that he named Nesophontes edithae, including many 
nearly complete skulls and other parts of the skeleton from 
caves at Morovis and Utuado. It apparently lived on the 
island until comparatively late, perhaps post-Columbian, 

This is the largest of the species at present known from the 
Greater Antilles, but since no living examples have been found, 
only the skeletal characters have been described. It was 
probably about the size of a chipmunk. In life, the snout was 
probably elongate and flexible, the head long and slender, the 
eye small, the limbs only moderately long and probably with 
five clawed toes on each; the thorax seems to have been narrow 
instead of widened as in Solenodon, and the tail was probably 
about as long as the body. The tubular skull lacked the jugal 
bone. The anterior pair of incisors is not greatly enlarged as 
it is in that genus; the canine is double-rooted and retains its 
primitive large size, while the upper molar teeth have high 
cusps, forming a V-shaped pattern. Only one of the original 
four premolars is missing so that the tooth formula is: if, CT, 
pmf , ml = 40. The humerus shows the entepicondylar fora- 
men, the pubic bones are not united below, and the tibia and 
fibula are nearly separated, becoming slightly united distally 
in adults. "No living insectivore has such an assemblage of 
generalized characters," a fact that makes the group of unusual 
interest. It was made by Anthony the type of a separate 
family, Nesophontidae, and is regarded by Winge as somewhat 
ancestral to the moles. The skull length is about 40-44 mm. 
The abundant remains found by Anthony represent a larger 
and a smaller size, the former of which are taken to be of males, 
the latter of females, an unusual difference among the insecti- 

Among the rugged and tree-covered hills of Puerto Rico, this 
mammal must once have been a common species and doubtless 
was preyed upon by owls, through the agency of which its 
remains were probably brought into the limestone caves where 
they have been found. Why this large species should be found 
on this island alone, while on Cuba and Hispaniola only small 
forms occur, or how it originally reached these islands at all, are 
questions still unsolved. Nor is it clear that the introduction 
of rats from Europe, or the burning of forests with intensive 
cultivation, is alone responsible for its disappearance. Pre- 


sumably it lived up till the earlier post-Columbian times. 
Hitherto no representative of the genus is known from Jamaica 
or other West Indian Islands except Cuba and Hispaniola. 
That so primitive and interesting a type should have wholly 
died out within Recent times is to be regretted. 


? Nesophontes micrus G. M. Allen, Bull. Mus. Comp. Zool., vol. 61, p. 5, pi., fig. 14, 
Jan. 1917 ("Cavern in the Sierra of Hato-Nuevo, Province of Matanzas, Cuba"). 

FIGS.: Allen, G. M., 1917, pi., fig. 14 (jaw); 1918, pi., figs. 7-10; Anthony, H. E., 1919, 
pi. 37, figs. 8, 9, 11, 12 (crania). 


Nesophontes longirostris Anthony, Bull. Amer. Mus. Nat. Hist., vol. 41, p. 633, 1919 

("Cave near the beach at Daiquiri, Cuba"). 
FIGS.: Anthony, 1919, pi. 37, figs. 10, 13 (cranium). 

These two Cuban Nesophontes are at first glance very similar, 
though N. longirostris has a slightly longer rostrum, and the 
upper premolars are spaced instead of being in contact. They 
are both known from skeletal remains only, the former from 
both western and eastern Cuba, the latter from the type locality 
in the eastern part of the island. 

These closely similar species differ remarkably from N. 
edithae of Puerto Rico in their very much smaller size. Other- 
wise they are much alike, although a number of minor charac- 
ters separating them have been pointed out by Anthony (op. cit. 9 
p. 633), such as the greater size of the first as compared with 
the second upper premolar, whereas in N. edithae these are 
equal; again, in the lower jaw, the second premolar is only 
about half as large as the two others instead of equaling them. 
Length of palate from postpalatal notch to alveolus of first 
incisor, about 14.1-15.3 mm. in N. micrus; 14.7 in N. longi- 
rostris, contrasted with 20.5 in N. edithae. Cranial length of 
N. micrus, 28 mm. 

Nothing is known of N. longirostris except that it occurred 
in eastern Cuba up to relatively recent times. Since this end 
of the island still harbors various species that are absent from 
the western end, it is likely that this was a local species, differ- 
ing only slightly from N. micrus, the remains of which are now 
known from recent cave deposits on the Isle of Pines as well as 
from both eastern and western Cuba itself. No doubt, as 


Dr. Anthony has suggested, the remains of these insectivores 
in the cavern deposits are due to the agency of owls, such as 
the Cuban barn owl, which even at the present time is building 
up in some of these caves deposits of bones of introduced rats 
as well as of bats. These bones, especially skulls, are more or 
less broken so that complete crania are rare among the deposits. 
Anthony adduces some evidence that the Nesophontes were 
abundantly represented in the deposits prior to the deposition 
of rat bones, so that the introduction of the rats may have 
been the decisive factor in extermination of the insectivores. 
Thus the latter may have been extinct only since white occupa- 


Nesophontes paramicrus Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 3, 1929 

("Large cave near St. Michel, Haiti"). 
FIGS.: Miller, 1929a, pi. 1, fig. 1 (skeletal remains); 1930, pi. 2, figs. 2, 3 (rostrum). 


Nesophontes hypomicrus Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 4, 1929 

("Deep cave near the Atalaye plantation [near St. Michel], Haiti"). 
FIGS.: Miller, 1929a, pi. 1, fig. 2 (skeletal remains); 1930, pi. 2, fig. 1 (rostrum). 


Nesophontes zamicrus Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 7, 1929 ("Large 

cave near St. Michel, Haiti"). 
FIG.: Miller, 1929a, pi. 1, fig. 3 (skeletal remains). 

These three insectivores, apparently extinct and known only 
from skeletal remains found in Hispaniolan caves, may be dis- 
cussed together. They have no common names. 

They are differentiated from one another largely on size 
characters. In body dimensions they approximated mice or 
small rats. 

"Insectivores of the genus Nesophontes are abundantly 
represented in the Haitian caves. They have not previously 
been recorded from the island of Hispaniola. In the superficial 
layer of the cave floors the bones of these animals occur in un- 
disturbed material along with remains of Epimys rattus [roof 
rat] and Mus musculus [house mouse]. This association is so 
intimate that there appears to be no reason to doubt the 
simultaneous occurrence of the insectivores and the introduced 
rodents. Some of the jaws of Nesophontes are more fresh in 


appearance than some of the jaws of Epimys near which they 
were found. Whether or not Nesophontes now exists alive is 
a question which for the present cannot be answered. No bones 
of insectivores have been found in any of the numerous fresh 
owl pellets which I have examined. It seems not improbable, 
however, that if any part of the island remains uninvaded by 
the roof rat, the native animal might now be found to exist 
there" (Miller, 1929a, p. 3). 

Wetmore and Swales write (1931, p. 238) : "It seems probable 
that the deposits [made by the extinct owl in the Haitian Caves] 
were accumulated over a long period of years extending perhaps 
from four hundred to two thousand or more years ago." 
They record the finding of skulls of Nesophontes in a supposed 
nesting site of this owl in a sink-hole on La Selle, Haiti. 

Miller also records (1929c, p. 4) remains of N. paramicrus 
and N. hypomicrus from a deposit made by the extinct owl, 
Tyto ostologa, in a cave on the islet San Gabriel in the Samana 
Bay region of the Dominican Republic. 

The occurrence of remains of these insectivores with those 
of the introduced roof rat and house mouse indicates their 
survival to post-Columbian times, while their usual absence 
from fresh owl pellets suggests their extinction. 

In the following year, Miller (1930) found bones of both 
these species in material collected earlier in 1930 about 10 
kilometers southwest of Constanza, Dominican Republic. 
This consisted of a mass of owl pellets, apparently those of the 
small living barn owl, found "in a shelter under an overhanging 
ledge about 100 feet up the northern flank of Monte Culo de 
Maco" in the rain-forest region where disintegration may take 
place much sooner than in the drier parts of the island. Some 
of the bones were so fresh as to retain bits of dried tissue while 
the brain case of one of the skulls was found to be "packed full 
of hair by the action of the owl's stomach." From this evidence 
Miller concludes that these insectivores as well as two associ- 
ated genera of rodents may still exist in that region. It is 
hoped that future exploration may confirm this suggestion. 


Family SOLENODONTIDAE : Solenodons 


Solenodon paradoxus Brandt, Mem. Acad. Imp. Sci. St. Petersbourg, ser. 6, vol. 2, p. 

459, 1833 (Haiti). 
FIGS.: Brandt, 1833, pis. 1, 2; Allen, G. M., 1910, pis. 1-3 (colored), 4-9 (anatomy). 

First made known by the Russian zoologist Brandt, a little 
over a century ago, this large insectivore was supposed to be 
nearly or quite extinct in its island habitat until 1907, when A. 
Hyatt Verrill found that it was still living in the interior of 
northeastern Hispaniola. 

It is a stoutly built animal, with a total length of about 22 
inches, of which the strong tapering tail constitutes about 10 
inches. Males and females are alike in size and in color, which 
varies from a mixed blackish and buff dorsally, with clearer 
yellowish sides, to a deep ferruginous tint, darkest and clearest 
on the lower throat. Characteristic is a small squarish spot 
of white in the middle of the nape. The tail is nearly naked 
and dusky in color except at the base, which is flesh-color. The 
feet and distal part of the limbs are nearly hairless and pale- 
tinted. The muzzle is provided with a long cartilaginous 
snout, supported at its base by a small round bone (the os 
proboscidis). The feet are armed with strong, slightly curved 
claws for digging, five on each foot. In its long tubular skull 
it somewhat resembles Nesophontes and the Madagascar tenrec, 
but the teeth show wide divergence from the former, in that 
the anterior upper pair of incisors and the second lower incisors 
are remarkably enlarged, the latter deeply grooved on the 
inner side. This enlargement is suggested by the generic name 
("sword tooth"). The upper and lower canines are somewhat 
reduced, the former about equaling the second incisor. Except 
for the loss of one premolar, the full placental tooth formula is 
present: it, ci, pml, ml = 40. 

Although this solenodon is said by Verrill to go under various 
names in Hispaniola, as orso, hormigero, juron, these are merely 
Spanish equivalents for "bear," "anteater," and "ferret." 
Hearne (1835, p. 105), however, in presenting a specimen to the 
Zoological Society of London in 1835, declared that in Haiti it 
was known to the people as agouta. Verrill writes that in the 
Dominican Republic it is called milqui. 




Following its rediscovery by Verrill in 1907, five living 
animals were sent to the zoological gardens at Washington and 
New York from the Dominican Republic, and others including 
four living adults were received by the Museum of Comparative 
Zoology at Cambridge. The last came from a locality in the 
interior of the northeastern part of Hispaniola known as La 
Vega. Since that time a German collector, Paul Thumb, by 
the aid of a well-trained dog, succeeded in 1935 and 1936 in 
securing a number, about 20 in all, which he shipped alive to 
the zoological garden at Hamburg, Germany. Of these, 
several survived the voyage and formed the subject of three 
short papers by Dr. Erna Mohr (1936-37) concerning their 
ways in captivity, with a number of photographic illustrations. 
Herr Thumb secured his solenodons mainly in rocky and 
wooded country in the vicinity of San Jose de las Matas. In 
life the solenodon is a rather slow-moving creature, constantly 
on the move with a shuffling gait, sniffing everywhere with its 
long nose, and scraping and scratching here and there with its 
long claws, exploring for food. It is capable of delivering a 
sharp bite if too much disturbed, however. 

According to Dr. Mohr (1937 A), Herr Thumb found soleno- 
dons in dens among broken rock for the most part, or occa- 
sionally in hollow logs. In the former case the depth varied 
according to conditions of the rocks. The holes he dug out in 
1935 were packed with the empty shells of a snail, but this was 
not the case with those examined in the next year; while at the 

Hispaniolan solenodon (Solenodon paradoxus) 


Hamburg gardens, Dr. Mohr found the solenodons quite 
indifferent to live snails presented to them. Thumb found as 
many as eight solenodons in a single burrow. In captivity it 
was ascertained that the female may have one to three, usually 
two, young at a birth and may bring forth twice in a single 
year, though showing no particular breeding season. The 
group of eight in a den may thus be accounted for as the two 
parents with their young of two successive litters. Three of 
the families dug out by Thumb consisted of the adult pair and 
their single young. A family secured for the Museum of Com- 
parative Zoology in 1937 by W. J. Clench was of similar consti- 
tution. These were from southwest of Sabana, Dominican 

At the present time it appears that this solenodon is in no 
immediate danger of extinction, but is still present in areas of 
stony forest in northeastern Dominican Republic. If, however, 
such areas are extensively encroached upon through clearing 
and cultivation, it will no doubt be reduced in numbers. 



Solenodon cubanus Peters, Monatsb. Konigl. Preuss. Akad. Wiss. Berlin, 1861, p. 169, 

1862 (mountains near Trinidad and Bayamo, Cuba). 
FIGS.: Peters, 1863, pis. 1-3 (exterior and anatomy); Mohr, 1937a, figs, on pp. 7, 8. 

The Cuban solenodon was probably the animal first made 
known by Oviedo as the ay re in his famous "Historia General y 
Natural de las Indias," 1535, but it remained for the Cuban 
naturalist Felipe Poey to bring it to the notice of modern 
scientists, in a newspaper report in El Plantel in 1838, a 
century ago. It was not till nearly 25 years later, however, 
that Peters was able to compare a specimen with the Hispanio- 
lan species and point out its distinctive characters. 

In size this is slightly smaller than the species S. paradoxus, 
about 20 inches long, tail about 7 inches, and is quite different 
in color. In place of the blackish and buffy to ferruginous back, 
the Cuban solenodon is blackish brown in color, with a varying 
amount of white, or buffy, which may include only the base of 
the muzzle, the cheeks, and a mark on each shoulder, as in a 
specimen in the Museum of Comparative Zoology, or may be 


more extensive, to include the entire head and lower surfaces, 
and extend back along the side of the neck to the shoulder, 
whence it continues in scattered long white hairs to the 
haunches. There is no distinct white mark on the nape. 

The first definite knowledge of this species is due to Poey, 
who obtained specimens in the mountains east of Bayamo, 
where it was said to be well known, and he published an account 
of it with a colored plate in his "Memorias sobre la Historia 
Natural de Cuba" in 1851. He, however, supposed it to be the 
same as the species of Hispaniola, and after search in the earlier 
accounts of Cuban animals, failed to identify it with any known 
to the early historians of the country. For this reason, he 
proposed that it be called the almiqui, a name derived from 
that of one of the mountains near where his specimens were 
taken, in the region east of Bayamo. Gundlach subsequently 
obtained it near Trinidad, about halfway of the length of the 
island of Cuba, to the westward. In 1886, he sent three to the 
United States National Museum that he secured from the high 
mountains about 30 miles from Bayamo. One of these was 
exhibited alive at a meeting of the Biological Society of Wash- 
ington (Proc. Biol. Soc. Washington, vol. 4, p. x, 1886). It was 
the surviving adult male of a family of three, of which the adult 
female and her young one succumbed during the journey from 
Cuba, and was said by Dr. F. W. True (1886) to be destined 
for the zoological garden at Philadelphia. Even at that time 
the animal seems to have become rare, for Gundlach wrote 
that it had been secured only after "the promise of many 
years" by the same person who had supplied specimens pre- 
viously to himself and Poey. No more are known to have been 
captured until 1909, when two Swedish engineers secured a 
live one in the mountains of eastern Cuba. This animal died 
soon after, and its remains were sent to the Riksmuseum at 
Stockholm. Two photographs taken before its death are repro- 
duced by Dr. Erna Mohr (1937a) and are apparently the only 
ones that show the living animal. At about the same time a 
second specimen (or possibly the same one) was in captivity 
in the possession of M. Bofell, director of the Municipal 
Museum of Santiago. According to Paul Serre (1910) it had 
been captured by laborers near Baracoa, in extreme eastern 
Cuba, in the course of road construction. Its owner is said to 
have refused various offers to buy it, but its subsequent fate 


is unknown. In the 30 years that have since elapsed I have 
no knowledge of the capture of any others. Efforts of various 
naturalists to discover it in eastern Cuba have been unavailing, 
and even Herr Thumb, who was so successful in finding S. 
paradoxus in the Dominican Republic with the aid of his dog, 
failed to find it when in 1937 he hunted for it in that region. 

Very little is known of its habits. It was believed to have 
lived in caves, but, curiously, the researches of Dr. H. E. 
Anthony and others in cave excavations have revealed very 
little even in the way of recent remains, although bones of the 
much smaller Nesophontes are common. The Museum of 
Comparative Zoology, however, has a lower jaw from a cave 
at San Lucas, near Maisi, eastern Cuba. This paucity of cave 
fragments may indicate that the animal was little preyed upon 
by owls, perhaps on account of its size and strength, but was 
rather secure in holes among rocks such as the Hispaniolan 
species makes use of. Captive animals are said to have eaten 
the meat of chickens, either raw or cooked, while the one 
captured by the Swedish engineers ate freely of raw beef given 
in small pieces. With the introduction and spread of the Bur- 
mese mongoose in Cuba, it is very possible that this voracious 
carnivore may already have exterminated the last of the 
solenodons. The loss of either of the two species is greatly to 
be deplored, since they are in the New World the last living 
representatives of the group of insectivores with triangular 
instead of squarish molars, the Zalambdodonta, found else- 
where at the present time only in Africa (the potamogale) and 
on the island of Madagascar (the Madagascar hedgehog and 
its relatives). 

Family SORICIDAE: Shrews 



ftorex teneUus myops Merriam, Proc. Biol. Soc. Washington, vol. 15, p. 76, Mar. 22, 
1902 (Pipers Creek [Cottonwood Creek], near main peak of White Mountains, 
altitude 9,500 feet, Mono County, California). 

FIGS.: Jackson, H. H. T., 1928, pi. 3, fig. B'; pi. 6, tig. R (skull). 

Few specimens of this tiny shrew are known. In color the 
back is drab, the lower parts pale smoke gray tinged with pale 


olive-buff (Jackson); tail buffy brown to tawny olive above, 
below nearly "tilleul buff, darkening toward tip." It is said 
to be thus paler than its near relatives of the ornatus group, and 
its skull is smaller than in any except Sorex tenellus and S. 
nanus of Owens Valley, California, and Colorado, respectively. 
Total length, 98 mm., tail, 41; hind foot, 12; length of skull, 

This small shrew is regarded by Dr. H. H. T. Jackson (1928) 
as a distinct species, though its relationship to S. tenellus is 
doubtless close. For many years it was represented only by 
the two specimens in the collection of the U. S. Biological 
Survey. Dr. Jackson believed it was probably confined to the 
White Mountains of California, "where it is seemingly rare 
and may even be exterminated by the destruction of its habitat 
through sheep grazing." More recently, however, Dr. William 
H. Burt (1934) has discovered it in the Charleston Mountains 
of southern Nevada, where he considers it common "along the 
small streams at altitudes of 8,000 feet or above" up to 10,000 
feet, not far from water. The possibility that grazing has 
already much restricted its habitat in the White Mountains 
makes its inclusion here excusable. 


The bats form a large complex order; they are the only 
mammals that truly fly. There are two suborders : 

Megachiroptera, fruit bats or flying-foxes. There is a single 
family, found in southern Asia, Africa, Australia, and many of 
the oceanic islands. They are characterized by blunt cheek 
teeth and a second claw in the expanded wing-hand. 

Microchiroptera, insectivorous bats (several are secondarily 
fruit-eating or carnivorous and one family is specialized for a 
diet of blood). Sixteen families are usually recognized, of 
which six are restricted to the New World and two are com- 
mon to both hemispheres. The New World families follow: 

(1) Noctilionidae, primitive bats, found in tropical America. 
They are related to the Old World Rhinopomidae and Embal- 
lonuridae. Two genera are recognized; neither is here con- 
sidered endangered. 

(2) Phyllostomidae, New World leaf-nosed bats. These do 
not appear to be closely related to any Old World group, but 


probably developed in South America. Most of the species of 
this large family have lancetlike nose-leaves; those that do not 
show this character have cutaneous growths on the lower jaw. 
Fifty-two genera are recognized, of which seven (including 13 
species) are discussed in this volume, as either extinct or 
extremely rare. 

(3) Desmodontidae, vampire bats. The blood-drinking bats 
are restricted to the American tropics. They are related to the 
preceding family and have a rudimentary nose-leaf. None is 
discussed in this work. 

(4) Natalidae, long-legged bats. These small, delicately 
built bats are found in the American tropics, north to the 
Bahama Islands and central Mexico. One species is believed 
to have become extinct in historical times. 

(5) Furipteridae. Two genera, closely related to the long- 
legged bats, are found in tropical South America. Neither 
genus is discussed here. 

(6) Thyropteridae. A single genus, with large suction disk 
on the thumb, is restricted to tropical America. This bat is not 
considered endangered. 

(7) Vespertilionidae, common bats. They are found in both 
hemispheres to the limit of tree growth and on the oceanic 
islands to Samoa and Hawaii. Three species are here con- 
sidered in danger of extinction. 

(8) Molossidae, free-tailed bats. Found in the warmer parts 
of both hemispheres. None of these bats is thought to be a 
vanishing form. J. E. H. 

Family PHYLLOSTOMID AE : Leaf-nosed Bats 


MonophyUus frater Anthony, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 565, 1917 (cave 

near Morovis, Puerto Rico). 
FIGS.: Anthony, H. E., 1917c, pi. 56, figs. 5, 6; 1918, p. 349, fig. 8 (skull). 

Of the various genera of bats confined to the West Indies 
and unknown from the neighboring mainland, the long-tongued 
members of MonophyUus are remarkable, for they are found in 
almost all the islands of the entire Antillean chain. Because 
those on the several islands can usually be distinguished from 


their neighbors by minute characters, they have been given 
distinctive names. Thus we have as the oldest named form, 
M . redmani of Jamaica; others are M . cubanus of Cuba, M . 
cubanus ferreus of Haiti, M. portoricensis of Puerto Rico, an 
allied form on Antigua, M. luciae on Santa Lucia, M. plethodon 
on Barbados, and M. clinedaphus from an unknown locality. 
All are living, and no doubt vary in abundance on the several 
islands. On Puerto Rico, however, in recent years, Dr. H. E. 
Anthony has discovered a second and larger species, which he 
has distinguished as M. f rater; on each of the other islands only 
one form is known to occur. 

Members of this genus of the subfamily Glossophaginae are 
likely to be confused with Glossophaga of the mainland (and 
some of the Antillean islands). They are small species with 
long snouts surmounted by a small lancet-shaped nose-leaf 
and are provided with a long extensible tongue for feeding on 
fruit juices and perhaps in part upon nectar or pollen of 
flowers. The teeth are peculiar in that the lower incisors are 
minute and the cheek teeth are slightly elongate and spaced. 
The cusps of the latter are low and the outer ones are placed 
close to the edge of the crown "-so that the usual W-shaped 
pattern of the cusps is obscured. The dentition includes, on 
each side, two incisors above and below, a canine, two upper 
and three lower premolars, and three molars above and below. 
Externally it is easily distinguished from Glossophaga by the 
longer tail, which is about half as long as the femur and pro- 
jects beyond the edge of the narrow interfemoral membrane, 
whereas in the latter genus the tail extends barely to the middle 
of the wide interfemoral membrane. 

The species M. f rater is known only from five fragmentary 
skulls excavated in the large Cathedral Cave near Morovis, 
Puerto Rico, by Dr. H. E. Anthony some years ago. With 
these was associated a skull of the living species still found in 
Puerto Rico, M. portoricensis, showing that the two were con- 
temporaries and represented distinct species. M . f rater is 
considerably larger than the latter, with a narrow elongate 
rostrum. This larger size is obvious from the comparative 
measurements: Length of the upper cheek teeth (alveoli), 
6.8-7.1 mm. as against 5.3; in the smaller species, length of 
palate 12.4-12.7 mm. as against 9.3. In spite of much collect- 
ing done in Puerto Rico, no living examples of this larger 


species have yet come to light. While it is quite possible that 
it may eventually be found, it seems more likely that it is now 
extinct, perhaps as a result in part of extensive clearing and 
agriculture. As Anthony points out, the interesting thing 
about this species is that it affords another instance of two 
closely related species from the same region, differing most 
obviously in little but size. A similar case is afforded by the 
bats of the genus Chilonycteris, of which three species occur in 



Reithronycteris aphyUa Miller, Proc. Acad. Nat. Sci. Philadelphia, 1898, p. 334 

FIGS.: Miller, 1898, figs. 2b (head), 3 (skull), 4 (section of palate), 5a (tongue). 

The present species seems most closely related to Phyllonyc- 
teris, which it may represent in Jamaica. Externally and in 
its dentition it resembles Phyllonycteris poeyi but is at once 
distinguished by the vestigial nose-leaf, which instead of 
forming an erect lancet is reduced to almost nothing and ap- 
pears as a short piglike snout, set off by an encircling groove. 
The ears, too, are shorter, and the tragus shows four notches 
on its outer margin; the hind foot is large, three-fourths as long 
as the tibia, and with strong claws; calcar absent, as in Phyllo- 
nycteris; and tail shorter than tibia. Color light yellowish 
brown (in alcohol) . The genus differs remarkably from Phyllo- 
nycteris, however, in that the floor of the brain case is elevated 
out of its usual position in such a way that the roof of the 
posterior nares is formed "by two longitudinal folds given off 
probably by the pterygoids and nearly meeting in the median 
line." The skull has the rostral portion relatively broader, 
and there is less contrast in both diameter and height of the 
upper incisors. Total length, 88 mm.; tail, 12; tibia, 17; 
forearm, 48; skull length, 26. 

The only known specimen of this genus and species is the 
type, a male preserved in the Museum of the Institute of 
Jamaica. Its further history is not recorded, but it was taken 
in that island some time prior to 1898; no fragments were found 
by Dr. H. E. Anthony in the course of his explorations for 
cave fossils in Jamaica, so that nothing is known of its haunts, 
habits, or present status. Very likely, however, it is on the 
way to extinction if it has not already gone. 




Stenoderma nichollsi Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 7, p. 529, 1891 (Island 
of Dominica, Lesser Antilles). 


Stenoderma montserratensis Thomas, Proc. Zool. Soc. London, 1894, p. 133 (Island of 
Montserrat, Lesser Antilles). 


Stenoderma luciae Miller, Proc. Acad. Nat. Sci. Philadelphia, 1902, p. 407 (Island of 
St. Lucia, Lesser Antilles). 



Ardops annedens Miller, Proc. Biol. Soc. Washington, vol. 26, p. 33, Feb. 8, 1913 
("Island of Guadeloupe, Lesser Antilles'^. 

So far as known, the bats of this genus are confined to the 
islands of the Lesser Antilles, where they appear to be rare, for 
few specimens seem to be preserved in collections. They are at 
present known to inhabit St. Lucia, Dominica, Guadeloupe, 
and Montserrat only, and on each seem to be represented by a 
slightly differing race. In time, with clearing of wooded areas, 
their habitat is likely to be more and more restricted, wtth the 
possibility of their final extermination; hence they may be 
considered here collectively. 

All the four known "species" are of similar appearance and 
of medium size, about 3 inches long from nose to tip of tail, 
with rather long loose pelage of a light-brown color. The fur 
of the body extends thickly over the hind limbs to the ankles 
and out along the border of the lateral membrane for a short 
distance; on the arms the hair extends thickly out to the wrist. 
The interfemoral membrane is very narrow, and there is a 
short calcaneum. The head is short and blunt; the ears are 
not much longer than the distance from their base to the end 
of the nose. The genus is closely related to Phyllops and 
Ariteus, of Cuba and Jamaica, respectively. It is smaller than 


the species of Artibeus found in the same islands and differs 
conspicuously in the skull by having the palate deeply emar- 
ginate posteriorly and in the great development of the post- 
glenoid process. In each jaw are two incisors, a canine, two 
premolars, and three molars. The crown of the inner upper 
incisor is short and thick, scarcely or not higher than long. 
Additional details of the teeth are given fully by Miller (1907, 
p. 151). 

The first-described species, A. nichollsi, of Dominica, is 
small, with a forearm of about 46 mm.; A. montserratensis, of 
Montserrat, is slightly larger, with a forearm of 51.5 mm.; the 
Guadeloupe form, A. annectens, is of intermediate size, with a 
forearm of 48 mm.; while the most southerly one, A. luciae, 
of St. Lucia, is very little larger than A. nichollsi, forearm 47 
mm., and has a small white spot on the shoulder and distinctly 
bifid inner upper incisors. If, as seems likely, the genus once 
occurred on Martinique, it has probably now been extirpated 
with clearing and volcanic destruction. 

These bats are evidently tree-living, rather than cave- 
haunting, which implies not only that they are somewhat 
solitary and so easily escape observation, but also are more 
difficult to find on account of living among the leaves by day. 
Of the Montserrat species, Thomas records that it is said to 
hang by day underneath branches of trees, and to do much 
damage to the cocoa plantations. It seems likely, however, 
that it can hardly be common enough to be very destructive, 
and that the larger Artibeus is the real culprit. Of the Guade- 
loupe form, Miller mentions five specimens, implying a small 
group captured. Dr. G. K. Noble, who secured an adult 
female and well-grown young in Guadeloupe in 1914, writes 
me that he found these hanging together directly over the 
path he was following through the woods near Sainte Rose. 
Later his guide caught another one, so that they must occur in 
some numbers still. Ardops is a member of the fruit-eating 
group of leaf -nosed bats; hence it must depend for a living 
upon the soft fruits of various forest trees, and these in turn 
must occur in sufficient variety to provide a continuous supply 
throughout the year. It is evident, therefore, that any impor- 
tant change in the forest cover, whereby such trees are de- 
stroyed or replaced by other kinds that do not produce the 
desired fruit, must inevitably affect these local species. One 


may surmise that as already indicated, such changes have 
taken place on Martinique and possibly too on St. Vincent, 
where otherwise one might expect the genus to occur. Of its 
status on the other Lesser Antillean islands practically nothing 
is known. 



Arctibeus falcatus Gray, Ann. Nat. Hist., vol. 4, p. 1, 1839 ("Cuba"). 

SYNONYM: Stenoderma albomaculatum Gundlach, Monatsber. Konigl. Preuss. Akad. 

Wiss. Berlin, 1861, p. 155. 
FIGS.: Dobson, 1878, pi. 28, figs. 3, 3a (dentition); Anthony, H. E., 1917b, pi. 34, fig. 

3 (skull). 


Phyllops veins Anthony, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 337, 1917 ("Cave at 

Daiquiri, Province of Oriente, Cuba"). 
FIGS.: Anthony, 1917b, pi. 34, figs. 4-6 (skull). 



Ardops haitiensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 24, p. 581, Sept. 11, 

1908 (Cafta Honda, Dominican Republic). 
FIGS.: Anthony, H. E., 1917b, pi. 34, fig. 2 (skull). 

The small fruit-eating bats of this genus much resemble 
those of the genus Ardops of the Lesser Antilles, but so far as 
known they are confined to Cuba and Hispaniola. Their 
superficial resemblance to the larger Artibeus, common in the 
same islands, may at times have caused them to be overlooked, 
yet from the paucity of known specimens they are probably 
actually fewer in numbers, or even, in the case of P. veins, are 
already extinct. 

Similar in size to the members of the genus Ardops, and like 
them with a short blunt rostrum, hairy limbs and wing-border, 
they may be at once distinguished by the fact that the deep 
emargination at the posterior part of the palate is continued 
forward in a converging V-shaped outline, which extends about 
to the level of the middle of the second molar, whereas in 
Ardops this indentation is a narrow, parallel-sided arch. In 
Phyllops, also, the crown of the inner upper incisor is slender, 


noticeably higher than long, whereas in Ardops and in Ariteus 
the crown of this tooth is short and thick, scarcely or not higher 
than long (Miller, 1907). In contrast to the related genus 
Stenoderma, the rostrum of the skull rises above the level of the 
low supraorbital ridges and the nasal opening extends much 
less than halfway back to the point of union of these ridges. In 
addition, the third upper molar is still more minute, and the 
second upper molar is scarcely three-fourths the size of the 
first instead of nearly equaling it in crown area. In P. falcatus 
the head and body measure about 1.9 inches; the tail is absent; 
forearm, 1.65 inches; tibia, 0.6 inch; hind foot, 0.4 inch. The 
fur is described as gray-brown, with dark tips; the lower side is 
slightly paler. 

The species described from Cuba as P. vetus is smaller than 
P. falcatus in its skull, the palatal emargination is narrower, 
and the small third upper molar is nearly circular instead of oval. 
The Hispaniolan species, P. haitiensis, is of about the same 
size as P. vetus but differs in the sudden contraction of the V-- 
shaped palatal emargination near its tip and in the apparent 
lack of the shallow pits in the basioccipital. 

Very little is known of the habits or status of these three 
bats. Both P. falcatus and P. vetus are Cuban, while P. 
haitiensis is at present known from Hispaniola only. The 
first was sent by MacLeay to the British Museum, in 1838 or 
1839, with the note, "Killed in my bedroom." Gundlach 
mentions specimens from Matanzas and Cardenas, and in 1914 
I examined two skulls taken from owl pellets in eastern Cuba 
by Dr. Charles T. Ramsden. In 1917, Dr. H. E. Anthony, in 
the course of excavations in a cave at Daiquiri, eastern Cuba, 
secured seven fragmentary skulls and four mandibles. Al- 
though he did not find the species alive, he mentions "an almost 
perfect skull taken from an owl pellet collected by Mr. Barnum 
Brown ... at the 'Cueva de los Machos' near Cienfue- 
gos." "The material," he writes (1919), "is, for the most part, 
fresh in appearance and not deeply discolored, some of it very 
recent in fact." In addition, there is a specimen in alcohol in 
the U. S. National Museum from Santiago, Cuba. Of P. vetus, 
discovered by Dr. Anthony in the cave deposits at Daiquiri, 
eastern Cuba, about 40 skulls were obtained. He states that 
it was "found as a fossil and judging from the condition of the 
specimens evidently has not been frequenting this region since 


the more recent animal remains, bats of the present day, were 
deposited. Therefore, while it is quite possible that this bat 
may be discovered living on some part of the island I am led to 
believe that it is truly extinct, a fossil of an earlier period than 
the very recent." Since, like its living relative, this was in all 
probability a tree-dwelling and fruit-eating species, it may have 
had a rather precarious foothold in this drier eastern part of 
Cuba. In a note in his paper of 1919, Anthony adds that a 
further comparison of the specimens of the two species, P. 
falcatus and P. veins, confirms his belief that the latter was an 
older inhabitant of the region for the bones "are all more 
ancient in appearance, more deeply stained and discolored"; 
while the fact that the owls that prey upon these bats have not 
brought them in to add to the more recent deposits is indicative 
of the same thing. 

Of the Hispaniolan species still less is known, for beyond 
the original specimen from Cana Honda, Haiti, it has elsewhere 
been recorded only from owl deposits. These are: near Con- 
stanza, in the mountainous interior of the Dominican Republic, 
where "in a shelter under an overhanging ledge about 100 feet 
up the northern flank of Monte* Culo de Maco," Herbert W. 
Krieger secured a broken skull and a mandible (Miller, 1930, 
p. 6); in a large cave near St. Michel, ten skulls and several 
mandibles; one skull from the deep cave and a mandible from 
the crooked cave near the same place; and a skull from owl 
pellets in the cave at Diquini, Haiti. These were found at all 
levels from the surface to a depth of two feet (Miller, 1929a). 

These small, heavy -bodied, fruit-eating bats may have been 
easily taken by their enemy the barn owl. Probably the 
species still persists in wooded regions of this island. 



Artibeus achradophilus P. H. Gosse, Naturalist's Sojourn in Jamaica, p. 271, footnote 

1851 (Content, Jamaica). 
SYNONYMS: Artibeus sulphureus P. H. Gosse, op. cit., p. 271, footnote, 1851; Adieus 

flavescens Gray, Proc. Zool. Soc. London, 1866, p. 117. 
FIGS.: Gosse and Hill, 1851, pi. 6, fig. 4 (nose leaf); Peters, 1867, pp. 433-434, pi. 2. 

For practically all we know of this bat, we are indebted to 
P. H. Gosse who first described it in his "Naturalist's Sojourn 
in Jamaica," the island to which it is confined. 


The species is one of the group of West Indian fruit-eating 
bats, of which the genera Ardops, Phyllops, and Stenoderma 
are related members. Of about the same size and appearance 
as these, it may at once be distinguished by the possession 
of only two instead of three upper molars (the minute third 
molar of the other genera having been lost in Ariteus) and by 
the presence of a "minute though evident metaconid" in the 
first lower molar. As in the related genera, the nose-leaf is 
lanceolate with a distinct midrib; the short interfemoral mem- 
brane is concave behind and like the legs is covered with 
rather long hair, forming a fringe along the posterior margin. 
It is said to be light reddish brown in color, paler beneath, 
and with a small white patch on each shoulder. There are no 
facial stripes, and the tail is lacking. Length of head and body, 
2.2 inches; forearm, 1.6 inches; tibia, 0.6 inch; hind foot, 0.4 

This genus may be regarded as the representative in Jamaica 
of the genus Phyllops, from which the above characters dis- 
tinguish it. Gosse (Gosse and Hill, 1851) writes that the first 
specimen he secured came in at the open window of a house at 
the Vineyard, near Black River, where he was staying. He 
supposed that, like other bats that flew in and out, it was in 
search of insects attracted by the lights. Three others he 
succeeded in shooting in the early evening at Content, and 
these are presumably the specimens listed by Dobson as in the 
collection of the British Museum. Here a large and fruitful 
naseberry tree (the nispero of the Spanish colonists) Achras 
sapota attracted many bats by its large fruits, resembling 
"a very rough russet apple, firm and fleshy, of a rich sugary 
sweetness." At about a quarter of an hour after sunset the 
bats began to visit the tree. "First one comes, takes a rapid 
flight around the tree, darts once or twice through the dense 
foliage, and winging away is lost in the light of the sky. Another 
and another comes immediately, and performs the same evolu- 
tions . . . By carefully following the flight of an indi- 
vidual with the eye, we perceive that now and then he alights 
for a moment on some object at the extremity of a bunch of 
leaves ; but no sooner has the eye rested on the spot than the 
sooty wings are again spread, and he is pursuing his giddy 
course with his fellows. The object of his visit is a ripe nase- 
berry, nestled in the midst of that rosette of leaves. Occa- 


sionally the weight of the suspended Bat dislodges the ripe 
fruit, and it falls to the ground, splitting with the shock. On 
picking it up we see that it has been just bitten, not gnawed as 
by the rodent incisors of a mouse, but nibbled in a ragged 
manner." No doubt several species of fruit-eating bats visit 
these trees for the feast. 

What may be the present status of this bat does not seem to 
be known. However, it is evidently rare in collections, whether 
from actual scarcity or through failure of visitors to capture it. 
On account of its island habitat and the intensive agriculture 
carried on in Jamaica, it is likely to become reduced in numbers 
as time goes on. 


Stenoderma rufum E. Geoffroy, Description de 1'Egypte, vol. 2, p. 114, 1818 (locality 

SYNONYM: Artibaeus undatus Gervais, in Exped. Amer. du Sud de Castelnau, Zool., 

Mamm., p. 35, pi. 9, fig. 3 (teeth), 1855. 
FIGS.: Peters, 1876, pi. 1, figs. 1-3; Anthony, H. E., 1918, fig. 10 (skull and teeth). 

This small fruit-eating bat was described 125 years ago from 
a specimen in the Paris Museum, but unfortunately there was 
no record of its place of origin. Since that time no other living 
specimen has been taken, and its habitat has remained un- 
known. In 1918, however, Dr. H. E. Anthony recorded the 
discovery of over two dozen "good-sized fragments of skulls, 
some nearly complete, " during his excavations in the Cathedral 
Cave near Morovis, Puerto Rico. The presumption is thus 
strengthened that the real home of the species was the Greater 
Antilles, perhaps Puerto Rico. 

In life this bat doubtless resembled somewhat the larger 
Artibeus, found in the same region, but the skull differs mark- 
edly in its short, broad rostrum, which, with its parallel and 
short tooth rows, is nearly square as seen from below. The 
median notch of the palate extends far forward to the level of 
the middle of the first molar. The incisors and upper canines 
form together a nearly transverse row, while the last molar, 
both above and below, is very small. The first two upper 
molars are broad for crushing and show little trace of the four 
primitive cusps. Total length of skull, exclusive of incisors, 
23.1 mm.; zygomatic width, 15.7. 

Since no other living examples of this small fruit bat have 
been found since it was first made known by Geoffroy, it is 


believed now to have become extinct during the last century. 
Anthony's interesting discovery of its remains in the Cathedral 
Cave in Puerto Rico indicates not only that this island may 
have been its home but also that it was fairly common there. 
The curious shape of the rostrum, so short and square, with 
heavy anterior molars, may indicate that it was adapted for 
feeding on some special kind of fruit, the abundance of which 
has been seriously affected through the clearing and burning 
of the original forest cover with cultivation on the island. 
Coincident with such restriction of food, the bat may have been 
much reduced in numbers until its final extinction. Anthony 
mentions the possibility that it may still be found on other 
islands but has been overlooked on account of a certain resem- 
blance to the common Artibeus. 


Phyllonyderis major Anthony, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 567, Sept. 7, 

1917 (cave near Morovis, Puerto Rico). 
FIGS.: Anthony, 1917c, pi. 56, figs. 1, 2; 1918, p. 356, figs. 12, a-d (skull). 


Phyllonyderis obtusa Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 10, Mar. 30, 
1929 ("Cave near the Atalaye plantation, about 4 miles east of St. Michel, 

The bats of this genus are known only from Cuba, Puerto 
Rico, and Hispaniola of the Greater Antilles, where slightly 
differing local forms have developed on each island. On Cuba 
the living species, P. poeyi, though apparently not very com- 
mon, nevertheless occurs in numbers in suitable caves, as at 
Guana jay, where Palmer secured a large series in 1900 (Miller, 
1904), Baracoa, or in the Sierra de Hato Nuevo, where in 1917 
Dr. Thomas Barbour found a large colony. On the other hand, 
in Puerto Rico it is unknown in the living state, though found 
in the superficial cave deposits, and the same is true of the 
Hispaniolan form; both are therefore included here among 
species nearly or quite gone. 

These are small bats with long narrow skulls and long pro- 
trusible tongues, which are useful in licking up fruit pulp and 


juices on which they largely feed. Probably pollen and flower 
nectar are also eaten. A small hastate nose-leaf is present at 
the tip of the snout, and the interfemoral membrane is narrow, 
extending only to the middle of the tibia. The calcar is absent 
altogether. In each jaw, both upper and lower, there are two 
incisors, a canine, two premolars, and three molars. The last 
are rather weak and rounded, the second and third lower 
molars without cusps, a character contrasting with the condi- 
tion in the related genus Erophylla, occurring in the same 
regions. The Puerto Rican Phyllonycteris major is decribed as 
closely related to the Cuban P. poeyi, but "noticeably larger, 
with wider brain case and heavier dentition"; greatest length 
of skull, 27-28 mm.; breadth of brain case, 11.4 mm. The 
form from Haiti, P. obtusa, is described as very similar to the 
Cuban species "but incisive foramina smaller and anterior 
border of premaxillaries as viewed in palatine aspect less 
narrowly curved." Greatest length of skull, 22.2 mm.; breadth 
of brain case, 10.2 mm. 

During the course of his investigations in Puerto Rico caves, 
Dr. H. E. Anthony found no evidence that P. major is still 
living in that island. In only one cave, that near Morovis, did 
he find its remains, including about 60 skulls and many jaws. 
This may indicate not only that it was a cave-dweller, like its 
Cuban relative, but also that its choice of caves and perhaps 
even its numbers were limited. "It was contemporaneous 
with Nesophontes as well as with Stenoderma ruficm and other 
bats to be found living today." Concerning the Hispaniolan 
P. obtusa, Miller (1929a) writes that remains were found in 
three distinct places: "The crooked cave near the Atalaye 
plantation, St. Michel, Haiti," a skull and mandible; a skull 
from a cave near Port-de-Paix; and a skull from owl pellets 
found in a cave at Diquini. The last implies that the skulls 
were fairly recent, and very likely the species may still be 
found there living, although none of the various collections of 
bats made in Hispaniola has hitherto yielded specimens. It is, 
as in other cases, difficult to assign an exact reason for the 
decline of such a species. Nevertheless, since it depends on 
a constant supply the year around of various fruits and berries 
in their season, it is obvious that any material changes in the 
forest or shrub covering, if this eliminated or reduced any one 
species of fruit even for a short critical season, might have a 
disastrous effect. 


Concerning the Cuban species, P. poeyi, Palmer (in Miller, 
1904) found it "very abundant in a wet, ill-ventilated cavern 
on Guanajay Mountain. On entering this cave, the vertical 
opening of which, about 12 feet across, was concealed by 
bushes, we descended about 25 feet, and were then standing 
some 20 feet above the lowest level. The slight noise which we 
made disturbed the bats in the inner chambers, and we could 
distinctly hear the rumbling made by their wings. As we 
proceeded this sound increased, until, when we reached the 
inner and thickly populated chambers, it became a grand, 
rushing roar of thousands on thousands of wildly flying animals. 
To reach the inner chamber it was necessary for us to descend 
from the first landing to the real floor of the cavern, and there 
light our candles, for not a ray of light and very little fresh air 
penetrated so far. From the floor we worked our way over 
many guano-covered, damp bowlders and through arches and 
narrow passages up to a sloping shelf, where, owing to the low 
roof, a man could not stand upright. By this time the bad 
air and excessive warmth was [were] telling on us, and we were 
in a most profuse perspiration. The bats were now thoroughly 
aroused, and the noise of their wings was astounding. Many 
were darting out through the passage by which we had entered 
. . . We began swinging a dip net in every direction, 
trusting to chance to secure specimens. About fifteen minutes 
of such work usually resulted in the capture of 20 to 30 bats, 
nearly all of this species . . . Before June 7, all the females 
were big with a single young, but after this date we found the 
pink, almost hairless little ones of different sizes hanging to the 
roof and scattered over much of its surface. On our last visit, 
late in June, the cave was so hot as to be unbearable . 
Among the specimens captured at the mouth of the damp cave 
near Baracoa . . . were many of this species . . . On 
one side of the vertical opening of this cave grew a large tree 
whose roots descended like a stream into the cavity. The 
people of the neighborhood assured me that the majas (the 
Cuban boa, Epicrates angulifer) coil themselves among these 
roots and grab at the bats as they fly out. I was told that a 
snake frequently secures a bat in this manner." 


Family NATALIDAE: Long-legged Bats 


Natahis primus Anthony, Bull. Amer. Mus. Nat. Hist., vol. 41, p. 642, Dec. 30, 1919 
("Daiquiri, Cuba," in the Cueva de los Indies). 

The bats of the family Natalidae are small and delicately 
formed and have noticeably long and slender legs and distinctly 
funnel-shaped ears. The ample interfemoral membrane is 
supported in part by the tail, which extends to its border. In 
distribution the members of this family are confined to tropical 
America and occur on some of the West Indies, to which three 
of the genera are confined: Chilonatalus (on the Bahamas, 
Greater Antilles, and Old Providence Island), Nyctiellus (the 
Bahamas and Cuba), and Phodotes (island of Curagao). The 
genus Natalus is the only one occurring on the mainland from 
tropical South America north to central Mexico, as well as in 
both the Greater and the Lesser Antilles. In the latter it is 
known living from the island of Dominica (N. dominicensis 
Shamel) and Antigua; in the former, from the Dominican 
Republic (Natalus major Miller). It is thus especially inter- 
esting that in the course of his cave investigations in eastern 
Cuba, Dr. H. E. Anthony should have found the present species 
represented by mandibles associated with Nesophontes and 
Boromys in the Cueva de los Indios at Daiquiri, "buried but a 
short distance under the surface. The bone is stained a very 
dark brown and probably represents an extinct form." From 
N. major, of the Dominican Republic, hitherto the largest known 
member of the genus, "it may be known by its even greater 
length of mandible and noticeably heavier teeth. The first 
lower molar is especially 'plump' in contour and the tooth 
extends externally considerably beyond the alveolar border." 
The mandible on which the description is based measured 14.4 
mm. in greatest length or about a millimeter more than in the 
large form of the Dominican Republic, of which it may be 
regarded as the Cuban representative. The color in life was 
perhaps yellowish, as in some of its near relatives. 

Nothing further is known of the species, but since it has not 
been found living by any of the various collectors who have 
searched for bats in Cuba, it is probably, as Anthony says, 


In this connection it may be surmised that the present 
sporadic distribution of this genus in the West Indies, with one 
living form on Dominica, one in Hispaniola, and a third prob- 
ably recently extinct in Cuba, implies that it may once have 
been more widespread over this area and has already died out 
on other islands. Of this, however, there is at present no 
evidence. The other related genera, too, may be thought of as 
in a restricted situation. The little Nyctiellus, one of the 
smallest of bats, and formerly believed to be confined to Cuba 
and the Isle of Pines, has in late years been found in numbers 
in some of the Bahamas (G. M. Allen and Sanborn, 1937) and 
is probably in no present danger. The somewhat larger 
Chilonatalus is found, in several closely allied races, on Cuba, 
Jamaica, the Bahamas, and on Old Providence Island, dwelling 
in caves. Wherever its small colonies exist it does not seem 
especially difficult to find and may be fairly safe for the present, 
since it is insectivorous like the others of the family, and there 
would seem to be no lack of sustenance for it. Of the genus 
Phodotes, a close relative of Natalus, practically nothing is 
known beyond Miller's original description based on a speci- 
men from the island of Curagao, off the coast of Venezuela. 
If, as supposed, it is peculiar to that island, its future status 
may well be somewhat precarious on account of changes taking 
place with intensive cultivation. 

Family VESPERTILIONIDAE : Simple-nosed Bats 


Histiotus maculaius J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 3, p. 195, 1891 
(near Piru, Ventura County, California, probably at mouth of Castac Creek). 

FIGS.: Miller, 1897a, pi. 1, fig. 11 (ear); pi. 3, fig. 3 (wing); 1907, p. 226, fig. 37 (skull); 
Bailey, V., 1931, pi. 21, B; Grinnell, H. W., 1918, pi. 16, fig. 9 (photograph). 

For 12 years after its first discovery in Ventura County, 
Calif., this bat remained unique; then a second one was found 
in the biological laboratory of the College of Agriculture and 
Mechanic Arts at Mesilla Park, N. Mex. Since that time only 
six additional specimens have been discovered, all in the South- 
western United States. This apparent rarity has led to the 
belief that the species is actually a waning one, and for this 
reason it is here included. 


This is a fairly large bat with a total length of about 110 min. ; 
tail, 50; forearm, 50. In color the fur is deep black, with three 
large white spots: one at each shoulder and one on the lower 
part of the back, giving a striking contrast. The lower side is 
washed with white. The very long ears measured 34 mm. in 
one specimen. The two upper and two lower premolars, the 
nostrils simple instead of opening upward or in connection with 
nasal swellings, and the distinctive color and long ears will 
serve to identify it easily. 

An account of the skeleton with figures of the important 
bones has been given by E. R. Hall (1934), who in a later 
note (1939) adds an eighth to the list of known individuals 
taken. The localities are: Piru, Ventura County, Calif.; 
Mesilla Park, N. Mex.; Yuma, Ariz.; Mecca, Kern County, and 
Yosemite Valley, Calif.; Reno, Nev.; and Salt Lake City, 
Utah (see Durrant, 1935, p. 226). In every instance only a 
solitary individual has appeared, and in such unexpected 
places as clinging to a fence rail, or the under side of a rock, 
under the eaves of a schoolhouse, in a biological laboratory (!), 
or on the side of a building. One was found dead in the overflow 
of a railway water tank. From these few instances it has not 
been possible to deduce what is the natural resting place of 
the species by day. Possibly the one found clinging to the 
lower side of a rock indicates more nearly the sort of place they 
normally seek for shelter. The area outlined by these eight 
known instances takes in roughly the arid region of the Great 
Basin of the Southwestern United States, to which probably 
the species is confined. Other than the scanty information 
supplied by the eight specimens hitherto discovered, nothing 
whatever is known of its habits. It is, however, not at all 
impossible that it may somewhere be found in greater numbers. 
Its evidently solitary disposition and its liking for outside 
resting places make it seem unlikely that it is a colonial and 
cave-dwelling bat. 

Whether this is a species in danger of eventual extermination 
can not now be told with certainty, nor does it appear that 
anything can at present be done to foster it. 



Atalapha brachyotis J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 4, p. 47, Mar. 25, 
1892 ("Chatham Island," Galapagos Islands). 

This, the only bat at present known from the Galapagos 
Islands, is one of the red-bat group, found on the American 
continents from north-temperate to south-temperate latitudes. 
It is a smaller relative of the hoary bats of the American 
continents and Hawaii and like them has a hairy tail-membrane, 
short rounded ears, and only a single upper incisor. 

The Galapagos bat is said to resemble closely its mainland 
representative in its rusty-red coloring and minutely frosted 
tipping to the hairs, but the ears are smaller and the wings 
slenderer, the dentition lighter, with shorter canines. Forearm 
length, 39 mm. 

Of this bat almost nothing is known. Dr. George Baur, 
who collected the type specimen about 50 years ago, wrote 
at the time of its capture (Allen, 1892): "On Chatham Island, 
at an elevation of about 1700 feet, where the hacienda is placed, 
we observed bats nearly every evening." The specimen was the 
only one he succeeded in capturing, and it is at present in the 
Museum of Comparative Zoology (its skull, however, unfor- 
tunately lost). Dr. Baur adds: "Bats have been observed on 
Indefatigable Island by Dr. Habel, and I observed one on 
South Albemarle." It is, of course, uncertain whether all 
these represented the same species of bat. Red bats are 
known to reach Bermuda in their migrations in eastern North 
America, so that it is not surprising that this strong-flying 
species should have reached the Galapagos at some distant 
period. Whether it will continue to survive the arid conditions 
and restriction of forest growth on the islands can not at present 
be told. 



Atalapha semota H. Allen, Proo. U. S. Nat. Mus., vol. 13, p. 173, Sept. , 1890 (Kauai, 

Hawaiian Islands). 
SYNONYM: Lasiurus grayii in part, J. E. Gray, Proc. Zool. Soc. London, 1862, p. 143. 

This is the Hawaiian representative of the continental 
hoary bat, which as a species ranges from north-temperate and 
sub-boreal latitudes to south-temperate climates, thus covering 


a wide latitudinal extent. It is migratory in northern United 
States and southern Canada; hence perhaps one may assume 
that the ancestor of the Hawaiian form reached that group of 
islands by flight over sea at some time in the distant past, 
coming from the American continent, Although at one time 
believed to be the same as the continental L. grayii of Chile, 
the Hawaiian form is darker, and in the red phase lacks to a 
large extent the hoary tips to the hairs of the upper surface 
found in that race and in the North American L. cinereus. 

In the Hawaiian hoary bat the color of the fur above may 
be either gray as in L. cinereus or dark red, almost blackish 
brown, with the middle portions of the hairs dull whitish, 
showing through if the fur is disturbed. On the lower flanks, 
legs, and interfemoral membrane above, the color of the back 
in the red phase passes gradually into a bright chestnut or 
mahogany red, with practically no white tipping to the hairs. 
Below, the color is paler, with chin and upper throat buffy 
white, and an indistinct half-collar of darker, set off by white 
tips to the hairs, from the dull-brown belly. Fur along the 
under side of forearm and at base of fingers clear buffy. A 
minute tuft of short white hairs #t base of thumb and of digit 
5, and a third white tuft on the inner side of the forearm just 
beyond the elbow. Forearm measures about 50 mm. (in the 
original description it is given as 40, but G. S. Miller, Jr. 
(1939) shows that this is a misprint for 50 as shown by an 
examination of the lectotype in the U. S. National Museum). 

This bat was first reported from the Hawaiian Islands by 
J. E. Gray in 1862, who, however, believed it identical with 
the Chilean race, which Tomes had earlier named Lasiurus 
grayii. Harrison Allen in naming it as a distinct form, had 
eight specimens from the "Sandwich Islands," only two of 
which had the more precise locality of Kauai. Perkins (1903, 
p. 465) writes that it chiefly frequents the mountains where, 
presumably, more forested areas remain. He adds that it is 
locally common in the uplands of Hawaii, and he has seen it, 
though rarely, on the islands of Kauai and Oahu. Bryan, 
writing in 1915, says: "They have always been rare, but are 
apparently still to be seen in the uplands of Hawaii." There 
are four specimens in the museum at Tring, England, from 
Hawaii taken in 1891, and the Museum of Comparative 
Zoology has a specimen taken in 1937 at Waimea, island of 


Hawaii, by Miss Barbara Lawrence, who says that the boys 
sometimes find them hanging among the fronds of tree ferns 
in the lowland valleys. 

While at present the species may be in no particular danger, 
it is likely that any considerable changes, such as reduction in 
the amount of sheltering tree growth, will affect the species 
adversely. As the only bat at present known to inhabit this 
group of islands, this species is of particular interest, and its 
obvious derivation from the American Continent makes it 
further noteworthy from the distributional viewpoint. 

Order EDENTATA: Edentates 

The New World edentates belong to the suborder Xenarthra. 
Two groups are recognized: 

(1) Cingulata, the armadillos and their relatives. A race of 
the widely distributed nine-banded armadillo is restricted in 
its range to a small West Indian island and may be easily 

(2) Pilosa, sloths, ground sloths, and American anteaters. 
Of these the ground sloths have become entirely extinct; five 
species are thought to have persisted into historic times in 
Puerto Rico, Haiti, and Cuba. J. E. H. 

Family DASYPODIDAE: Armadillos 


Dasypus novemcinctus hoplites G. M. Allen, Bull. Mus. Comp.. Zool., vol. 54, p. 195, 
July, 1911 ("Hills back of Gouyave, island of Grenada," West Indies). 

This is a small race of the continental nine-banded armadillo, 
but it may be included in this account since any island race 
represented by a relatively small population is liable to further 
reduction if in the course of years the conditions change 
greatly through fire, settlement, or other causes that may 
further restrict the available habitat. 

Externally this is a somewhat smaller replica of the ordinary 
nine-banded armadillo, with the usual external bony cuirass 
consisting of a shoulder shield, a rump shield, and nine inter- 
mediate half-rings on the body. The tail is covered with a 


series of 12 complete bony rings, and the color in life is light 
flesh-color. From the posterior free edges of the transverse 
body-rings project three or four short bristles from each scale 
and similar but smaller bristles are present at the posterior 
margins of the scales on the shields of body and tail. On the 
under side of the body there are transverse rows of small round 
scutes, each in the center of a cluster of yellowish bristles. The 
skull in addition to being smaller than that of the mainland 
representative has one or two less teeth in each series, usually 
six or seven instead of eight. Total length, up to 678 mm.; 
tail, 82; hind foot, 37; greatest length of skull, 85.5. 

At the present day this armadillo is confined to the rough 
country covered with rain forest on the hills in the central 
part of the island of Grenada. Probably, too, the armadillo 
found on the island of Tobago is very similar if not identical. 
It is more or less hunted by the local Negroes who esteem its 
flesh. They not only hunt the armadillos with dogs but catch 
them in deadfalls set in the runways. Whether the armadillo 
naturally occurred in these islands or whether it was introduced 
there long ago is uncertain, but in either case it has been native 
for a long enough time to allow its development into a distinct 
dwarf race. De Rochefort, writing in 1658, mentions the 
"Tatou" as a native of Tobago, especially calling attention 
to the small size. Soon after, Du Tertre in the 1667 edition of 
his work on the history of the French Islands, was the first to 
record it from Grenada, where he says it was then common. 
It was unknown on any of the other French islands, and 
apparently all attempts to introduce it in Martinique from 
Grenada had ended in failure. Labat in 1742, however, 
mentions having eaten the flesh of armadillos that had been 
brought to Martinique from Grenada, and in 1700 had several 
times eaten it on the latter island. 

At the present time I know of no reason to suppose that 
this armadillo is in any danger of extinction. When I visited 
the island in 1910, I found it well known to the people and a 
Mr. John Branch assured me that often the Negroes might 
capture several in a single night, hunting with dogs, in the 
forest back of Victoria. The animal is fairly prolific and should 
survive without difficulty if conditions remain substantially 


Family MEGALOCHNIDAE : Ground Sloths 


Acratocnus odontrigonus Anthony, Ann. New York Acad. Sci., vol. 27, p. 195, Aug. 9, 
1916 ("Cueva de la Ceiba, near Utuado, Porto Rico"). 

FIGS.: Anthony, H. E., 1916, pis. 7-10, pi. 11, fig. 1; 1918, figs. 43-48, 50-54 (except 
those of A. major'); pi. 69, figs, la-c; pi. 70, figs. 2-6; pi. 71, figs. 1, 3-6; pi. 72, 
figs. 1, 3, 4; pi. 73, figs. 1-3, 5, 7; pi. 74, figs. 1-4 (skeletal parts). 


Acratocnus major Anthony, Mem. Amer. Mus. Nat. Hist., ser. 2, vol. 2, p. 412, 1918 

("Cave . . . near Utuado, Porto Rico"). 
FIGS.: Anthony, H. E., 1918, figs. 42, 43, 48, 49, 51, 53 (parts pertaining to No. 17169); 

pi. 69, figs. 2a^c; pi. 70, figs. 1, 7, 8; pi. 71, fig. 2; pi. 72, figs. 2a-b, 5; pi. 73, figs. 



Acratocnus (?) comes Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 26, Mar. 30, 

1929 ("Large cave near St. Michel, Haiti"). 
FIGS.: Miller, 1929a, pi. 5, fig. 2; pi. 6, fig. 2; pi. 8, fig. 1; pi. 10, fig. 1 (limb bones and 


Among the most interesting finds of recent years is the dis- 
covery of bones in the caves of Puerto Rico and Haiti repre- 
senting a small ground sloth, which was evidently contem- 
porary with aboriginal culture. It was approximately the 
size of a two-toed sloth but represented the ground-living 
group, known from the mainland of North America by the 
gigantic mylodons of the Pleistocene era. The nearest rela- 
tives seem to be the small ground sloths of the genus Hapalops 
from Miocene formations of Patagonia, although there are 
various differences. Their presence on both Puerto Rico and 
Hispaniola is additional evidence of the faunal affinities of 
these two islands, and since the three forms hitherto described 
seem closely allied, they may be considered under a single head. 

The smaller ground sloth of Puerto Rico was the first 
described and is the best known, from abundant skeletal 
remains found in the cave earth during excavations originally 
undertaken in the course of archeological research. As in the 
living tree sloths, the premaxillae were toothless, and the 


incisors and canines of the lower series are also lacking. The 
skull is relatively long and narrow instead of globular as it is in 
the tree-living types, the rostrum is short and broad, and there 
is a well-developed median crest extending forward from the 
wide occipital ridge to the frontal region, where a lower branch 
goes to each of the short postorbital projections. The large 
tusklike upper canines are strongly triangular in section, 
followed by a diastema of nearly the same length as the diam- 
eter of the tooth or somewhat longer, then four cheek teeth in 
two parallel rows, each tooth approximately oval in section, 
with the broader end on the lingual side. In the lower jaw, 
there is a large tusklike first premolar corresponding to the 
upper canine, then a space, followed by three cheek teeth 
much like those of the upper series. The humerus has a 
slender shaft, a wide distal expansion, and a large entepicondy- 
lar foramen. The femur is wide in front view, but not thick, 
and shows a low third or outer trochanteric ridge. The tibia is 
short and stout and the fibula is free. The calcaneum or heel 
bone is broadly hatchet-shaped, and the terminal phalanges 
were evidently provided with stout claws. The tail is believed 
to have been short but rather stout. Dr. Anthony gives the 
following dimensions of the major skeletal elements: Skull 
length, tip of rostrum to occipital condyles, 136.2 mm.; breadth 
of brain case, 28.5; greatest breadth across postorbital proc- 
esses, 47.5; alveolar length of upper molar series, 35.5; longest 
humerus, 145; ulna, 160; radius, 128.8; longest femur, 163.3; 
longest tibia, 128. 

Among the remains of this species recovered in caves in 
Puerto Rico were a few indicating a similar sloth of slightly 
larger size, which Dr. Anthony in a later communication 
decided must represent a second and larger species of the same 
genus. This he named Acratocnus major, and defined it as 
being very similar to A. odontrigonus but "larger and heavier 
and with different skull characters"; it has "proportionally a 
much broader muzzle and an elevated basioccipital region," 
and a larger upper canine. No complete measurements of the 
skull are available, but the breadth across postorbital processes 
is 66 mm.; length of ulna, 171; length of tibia, 133. By com- 
parison with the corresponding measurements given above for 
the smaller species, the obvious size difference may be appre- 


The Haitian species, A. (?) comes, was based on a femur from 
a large cave and represents an animal much like A. odontrigo- 
nus, weighing perhaps 50 pounds. The bone differs from the 
corresponding one of the latter in having the neck of the 
articular condyle shorter and less bent outward and forward 
so that it diverges less noticeably from the general contour of 
the shaft. In describing the species, Miller refers to it a few 
other fragments, including several caniniform teeth from 
Gonave Island, off the coast of Port-au-Prince, Haiti, but the 
complete skull is as yet unknown. 

In Puerto Rico, remains of this type of ground sloth occur 
in dry caves in the limestone of the mountainous parts of the 
island. " Caves up on the sides of small hills yielded the most 
bones and the size of the cave was immaterial. The cave that 
contained the greatest amount of Ground Sloth material was a 
small one with the entrance on a rather steep hillside and 
opening out on a sheer front of limestone. Inside, the cave 
did not open out very wide but had a deep fissure at the left 
toward which the floor sloped abruptly. This fissure was richly 
packed with bones of the sloth and the large rodent [Elasmo- 
dontomys], the bones beginning at near the surface and con- 
tinuing down some nine feet when excavations had to be given 
up because of the impossibility of reaching any deeper. This 
cave had the appearance of a trap for any animal that wandered 
into it and certainly would have proved so for any old or sick 
animal that had strength enough to crawl up through the 
cave entrance." 

Concerning the remains of this genus and a larger sloth 
found in Hispaniola by Miller, the latter states (1929a) : 
"That one or both of these sloths continued to exist on the 
island until after the advent of man I have no doubt. The 
facts that have led me to this conclusion are as follows : (a) In 
the two caves near St. Michel most of the sloth remains were 
found within two feet of the surface; and human bones and 
pottery occurred to the same depth without any indication 
that they had been dug in. (b) Near the entrance to the 
smaller of the two main caves bones of ground sloths (certainly 
two and perhaps more individuals) were inextricably mixed 
with bones of man (adult and infant) and domestic pig. The 
remains were scattered among the small fragments of limestone 
which made up the greater part of the floor material, and I was 


unable to determine any definite level-relationship among 
them, (c) Near the entrance to the large cave I unearthed 
with a trowel, in fine, soft, undisturbed material at the bottom 
of a trench two feet deep, the femur of a ground sloth, and, 
about 18 inches from it, a fragment of coarse dark pottery . . 
the bone and pottery had every appearance of having been 
deposited on the former surface of the cave floor and subse- 
quently covered by the gradual accumulation of detritus . . 
In general the ground sloth bones were associated with the 
human remains in exactly the same manner as the bones of 
Isolobodon and Plagiodontia, rodents which are positively 
known to have been contemporary with man." The external 
appearance of these animals may only be conjectured, for no 
fragments of hide or hair are known to be preserved. 


Parocnus serus Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 29, Mar. 30, 1929 

("In a large cave near St. Michel, Haiti"). 
FIGS.: Miller, 1929a, pi. 7; pi. 8, fig. 2; pi. 9; yl. 10, figs. 2, 3. 


Megalocnus rodens Leidy, Proc. Acad. Nat. Sci. Philadelphia, 1868, p. 180 (Cuba). 

Miller, in describing the larger ground sloth from Haiti, be- 
lieved there was good evidence that it lived to be the contem- 
porary of man, since its remains were found in close association 
with pottery and in layers of cave earth at no great depth. 
Judging from the size of the few bones found, he surmised it to 
have been an animal weighing upward of 150 pounds and more 
heavily built that Acratocnus. The femur, on which the de- 
scription mainly rests, is at once distinguishable by the absence 
of the lesser trochanter, "as well as by its greater size and the 
much more noticeable antero-posterior flattening of the upper 
portion of the shaft." No skulls have yet been found suffi- 
ciently well preserved to give much idea of the cranial characters. 
An important feature of the humerus seems to be the lack of an 
entepicondylar foramen, of which no trace is visible in the 
figure published by Miller. The humerus measures 200 mm. 


in greatest length. Three calcanea are essentially similar to 
the calcaneum of Mylodon. 

While Acratocnus and Parocnus did not perhaps become ex- 
tinct till fairly recent times, possibly not until after the dis- 
covery of the West Indies by Europeans, the Cuban ground 
sloths may have died out at an earlier time. Their remains are 
known in some abundance in deposits around a warm spring 
at Ciego Montero, associated with bones of a crocodile and a 
giant tortoise. Another important locality is the Casimba 
in the Sierra de Jatibonico, in central Cuba, a fissure spring 
at the bottom of a ravine, where many bones have been 
found. Although a full account of these remains was in 
preparation by the late Dr. W. D. Matthew and Prof. Carlos 
de la Torre, this seems to have been indefinitely postponed 
through the death of the former. Nevertheless, in pre- 
liminary papers (Torre and Matthew, 1915; Matthew, 1918, 
1931) they distinguish no less that four genera, to which names 
are given. Matthew (1918, p. 660) restricts Megalocnus to the 
largest of these, "about the size of a black bear"; the smallest 
is Microcnus, "about the size of a cat, and there are two of 
intermediate size, Mesocnus, with a rather long, narrow muzzle, 
and Miocnus, with a broad, square muzzle." To this last 
genus Acratocnus of Puerto Rico and Hispaniola is said to be 
related. Possibly, too, Parocnus may be found to be closely 
related to one of the two intermediate genera or even identical 
with it, when better material is assembled and restudied. The 
three other genera, Megalocnus, Microcnus, and Mesocnus, 
have large tusks of a "peculiar dished shape, with a tendency 
to approach toward each other like the incisors of rodents. " 
Matthew regards the Cuban ground sloths and the Puerto 
Rican Acratocnus odontrigonus as descendants of a common 
ancestral type of Upper Miocene or Lower Pliocene age, related 
to that of Megalonyx. Matthew adds that "there is so much 
individual and age variation in ground sloths that it is difficult 
to say how many species of these genera are present. Mega- 
locnus occurs abundantly both at Ciego Montero and the 
Casimba, but apparently only one species at Ciego Montero, 
while at the Casimba there may be two or three. The species 
found at the eastern end of the island agree better with the 
Casimba forms than with the Ciego Montero species. Of the 
smaller forms there are clearly two species of Mesocnus, but I 


see no proof of more than one of Miocnus or Microcnus." So 
far nothing seems to have been discovered that would indicate 
that ground sloths in Cuba were contemporaneous with early 

Order RODENTIA: Gnawing Mammals 

The rodents, here considered to include the rabbits and 
pikas, are the largest order of mammals, both in number of 
species and of individuals. They are characterized by chisel- 
like incisor teeth, usually a single pair above and below, but 
with a second pair of upper incisors in the rabbits. They are 
mostly small mammals, with large populations and rapid rates 
of reproduction. There are four suborders: 

(1) Lagomorpha (Duplicidentata) : The rabbits and hares, 
found in all zoogeographical regions, except the Australian, 
and pikas of Asia and western North America. Rabbits have 
been introduced into Australia and elsewhere. 

(2) Sciuromorpha : This includes the squirrel family, which 
is world wide, except for the oceanic islands and the Australian 
Region; the scaly -tailed African "flying squirrels," the African 
springhaas; the beaver, found in Europe, northern Asia, and 
North America; and the sewellel of the Pacific coast of North 

(3) Myomorpha: The rats, mice, jerboas, and their relatives; 
representatives of this group occur in all regions, although 
human agency is doubtless responsible for their presence on 
some oceanic islands. 

(4) Hy stricomorpha : The porcupines, cavies, chinchillas, 
and their relatives; representatives of this group are found in 
all regions, except the Australian and the oceanic islands. 

The vanishing or extinct North American rodents are the 
following : 

(1) Sciuridae, squirrels, four forms of a single genus. 

(2) Castoridae, beavers, eighteen races of the single Ameri- 
can species. 

(3) Cricetidae, native rats and mice, eight species of three 

(4) Echimyidae, hystricoid spiny rats, twenty species and 
subspecies of nine genera. 

(5) Heptaxodontidae, the unique species, thought to have 
become extinct about the time of Columbus. 


(6) Dinomyidae, giant hystricoid rats, two species of two 

(7) Dasyproctidae, agoutis, three insular species of the 
typical genus. 

The last four families belong to the New World Hystrico- 
morpha and have their relatives in South America, where they 
probably originated. J. E. H. 

Family SCIURIDAE: Squirrels 



Sciurus kaibabensis Merriam, Proc. Biol. Soc. Washington, vol. 17, p. 129, 1904 ("Head 
of Bright Angel Creek, top of Kaibab Plateau, north side of Grand Canyon of 
Colorado, Arizona"). 

FIGS.: Nelson, 1918, p. 448, lower fig. (col.); Goldman, 1928, p. 127, pi. 16 (photo- 

This is perhaps the handsomest of the North American 
squirrels, a close relative of Abert's squirrel, with long-tufted 
ears, which ranges from Colorado into Mexico. The white- 
tailed squirrel has become isolated on the north side of the 
Grand Canyon in the high Kaibab Plateau and has developed 
striking color characters, such especially as its large white tail. 
Because of its restricted habitat and conspicuousness it may be 
included here. 

In its general appearance it resembles Abert's squirrel, but 
the under parts are mainly black instead of white, and the tail 
is practically all white instead of on the under side only. The 
back is dark grizzled gray, with a wash of ferruginous from 
shoulders to rump. The ears are blackish in summer but 
rimmed anteriorly with gray in winter; the ear tufts are black, 
the face, fore feet, and toes mixed gray and black; hind feet in 
summer mainly gray but in winter mainly black. The tail is 
ample and bushy, white, with an indistinct buffy-gray stripe 
down the middle of the upper side. Size about that of a gray 

The Kaibab squirrel is found in a restricted area about 40 
miles in length and 20 miles in greatest width, at the broadest 
(southern) part of the Kaibab Plateau. Isolated here from its 
nearest allies of the south side of the Grand Canyon, it is con- 
fined to the portions of the plateau where the yellow pine 


(Pinus ponder osa) is found. It is largely dependent on this 
tree for food and shelter as are its relatives of the Abert's 
squirrel type, which, wherever they occur, "are rarely or never 
seen beyond the local range of this tree. On the Kaibab 
Plateau the altitudinal range is mainly from 7,000 to 8,500 
feet" (Goldman, 1928). Major Goldman (1928) writes: 
"Yellow pines are rarely hollow and I have never seen one of 
these squirrels enter a hole. Dome-shaped nests, about 2 feet 
in diameter, made of cut pine branches, are usually well con- 
cealed among crowded limbs in the upper part of a tree. Like 
most squirrels, the white-tail is most active during the early 
morning and late afternoon, but may be found abroad at any 
time of day. Practically nothing seems to be known of its 
breeding habits." Although it depends partly for food upon 
the seeds of the yellow pine, which it extracts by gnawing away 
the scales, this supply is more or less precarious in years when 
the crop is poor. Its main reliance is therefore the cambium 
beneath the bark of the newer twigs of the yellow pine, which 
provides an unfailing food supply. "In feeding, the leaf- 
bearing branch tip is commonly severed and allowed to drop 
to the ground. A subterminal section of tender stem 2 to 4 
inches in length is cut off, scaled, and when the cambium sought 
as food has been neatly removed, the peeled wood remaining 
is also dropped ... In summer, however, they evidently 
indulge in a more varied diet. Much time is spent in foraging 
upon the ground and I have at various times observed them 
feeding upon mushrooms and other funguses, and they doubt- 
less eat many other things. " 

Although it is known that these squirrels are occasionally 
taken by hawks, Major Goldman believes that their numbers 
are no more seriously affected by these enemies than are those 
of other forms of Abert's squirrel. "The destruction of much 
ground cover through overbrowsing by deer may, however, 
expose the squirrels to some additional danger from hawks. 
Squirrels everywhere vary in numbers from year to year owing 
to causes imperfectly understood, but at present there seems 
to be no great danger of the extinction of the white-tail. These 
beautiful squirrels are objects of great interest to tourists along 
the Grand Canyon highway that bisects the Kaibab plateau, 
and efforts should be made to establish them in other yellow 
pine forested areas. " He adds the caution that care should be 


taken in such an enterprise, not to introduce them into other 
areas already occupied by races of Abert's squirrel, since hy- 
bridization would inevitably tend to the loss of the distinctive 
and conspicuous characters of this particular form. At the 
present time the Kaibab squirrel is completely protected by 
law and should be fairly well safeguarded, but any influences 
that might tend to impair the growths of yellow pine to which 
it is restricted would react unfavorably on its welfare in the 
limited area in which it is found. 



Sciurus niger avicennia A. H. Howell, Journ. Mammalogy, vol. 1, p. 37, Nov. 1919 
("Everglade, Lee County, Florida"). 

There are few less-inviting places for mammals than a man- 
grove swamp. The clean trunks and branches offer little in the 
way of food or shelter; the soft ooze from which the trees grow 
is daily washed by the tides and thus is unsuitable for ground- 
livers ; while the suckerlike stems and rooting tips form a tangle 
almost impossible for the larger species to penetrate. It is 
therefore the more remarkable that a depauperate form of the 
southern fox squirrel has made the mangrove belt of the south- 
west coast of Florida its home and become adapted to a life in 
this apparently unfavorable environment. 

The mangrove squirrel is described as smaller and darker 
(more tawny) both above and below than the typical race, 
with the feet clearer white and less tinged with buff. The type 
is in the so-called buff phase or at least is not of the entirely 
black sort, with white face and ears. The head and back are 
black sprinkled with cinnamon but more abundantly on lower 
back; sides shading to orange-cinnamon; nose, lips, front of 
face, and ears white; fore legs blackish washed with orange- 
cinnamon, the feet and toes white; hind legs with more orange- 
cinnamon than black, the feet edged with white. Tail orange- 
cinnamon above, mixed with black, and shading on the sides 
to hazel; under side rich tawny, with a submarginal band of 
black. Under parts dull orange-cinnamon washed on throat 
and breast with black and white. A black phase also occurs, 
in which the nose, ears, paws and sometimes the end of the 
tail are white, while elsewhere the pelage is deep shining black, 


with a few pale hairs on the under side. Total length of an 
adult male (the type), 535 mm. (21.25 inches); tail vertebrae, 
260 (10.25 inches); hind foot, 75 (about 3 inches); skull, occip- 
itonasal length, 65.5. 

This squirrel is confined to "the damp, dark forests of black 
and red mangrove which extend practically without a break 
from Marco Pass to Cape Sable and around the southern end 
of the peninsula" of Florida, "to the shores of Biscay ne Bay 
on the east coast." Here it is apparently not common, for 
Ho well writes that " several days spent in hunting through these 
mosquito-infested forests resulted" in only a brief glimpse of 
one, while the type specimen he secured through an Indian boy 
who knew where its home tree was located. A small series of 
specimens, however, was secured by W. S. Brooks in the spring 
of 1920, for the Museum of Comparative Zoology, mostly in the 
black phase. I am indebted to Dr. Thomas Barbour for a few 
notes on this squirrel. He tells me that it is well known to the 
residents of the region, though not regarded as common, and 
that it may be found on even some of the isolated keys grown 
up to mangroves. Its feeding habits are remarkable for, in- 
stead of depending on nuts and other seeds, it lives on the buds 
and bark of the mangroves, gnawing the latter from twigs 
which it cuts. Dr. Barbour reports that after the hurricane 
that passed over this region a few years ago its numbers seem 
to have been lessened. Evidently it is very difficult to secure 
satisfactory estimates of the size and extent of this squirrel 
population, since the swamps can hardly be penetrated except 
by canoes along small waterways. While this race of fox 
squirrel may be in no very immediate danger at the present 
time, it might easily be affected by any large-scale attempts to 
alter the local ecologic conditions. 


Sciurus niger bryanti H. H. Bailey, Bull. Bailey Mus. and Library of Nat. Hist., no. 
1, p. 1, Aug. 1, 1920 ("Cambridge, Dorchester County, Maryland"). 

Dorchester County, Md., lies on the southwestern side of 
the large peninsula, nearly cut off from the mainland on the 
east by Delaware Bay and Delaware River and on the west by 
the long estuary formed by Chesapeake Bay. Isolated in the 


area near the tip of this peninsula is a small population of fox 
squirrels, which differ from the eastern fox squirrels in colora- 
tion by lacking the buff and russet tints. They form a local 
race, named by H. H. Bailey in honor of the ornithologist 
Walter E. Bryant. 

In a small series of specimens of this squirrel the coloration is 
remarkably uniform. The entire dorsal surface of the body is a 
hoary gray, the result of abundant white-tipped hairs mixed 
with black hairs. On the forehead the black hairs slightly pre- 
dominate. The under surfaces from chin to root of tail are 
white. Feet and ears dull white, sometimes with a faint tinge 
of buffy. Tail mixed black and white both above and below, 
the black forming a narrow and continuous submarginal 
border on the under side. 

Although, according to Bailey, the local gunners secure a 
few each season in Dorchester County, this race is so localized, 
with little or no likelihood of its increasing its range, that it 
must be thought of as like an island form threatened with 
gradual encroachment on its available habitat. For this reason 
it is likely in time to become more and more restricted, with 
little possibility of long survival unless given special protection. 

Concerning the present status of this squirrel, I am indebted 
for a few details to David V. Black, manager of the Blackwater 
National Wildlife Refuge, Cambridge, Md., who in response to 
inquiries writes, under date of March 21, 1941: "I have talked 
with several persons in Dorchester County about the fox 
squirrels found here, and they all seem to be of the opinion 
that they are fairly abundant and are found in all parts of the 
county. There is no particular protection given to Bryant's 
fox squirrel. It is hunted indiscriminately with other squirrels 
during the open seasons in the State. I do not think that it is 
threatened with extermination as yet. Most of the timber 
has been cut out in the county, and what little remains seems 
to be in the process of cutting with small portable sawmills, so 
that the natural habitat for squirrels and other woods dwellers 
is continually diminishing. The squirrels do occur on the 
refuge, where, of course, no hunting is allowed." 




Macroxus neglectus Gray, Ann. Mag. Nat. Hist., scr. 3, vol. 20, p. 425, 1867 (Wilming- 
ton, Delaware). 

SYNONYMS: Sciurus vulpinus Schreber, Saugthiere, vol. 4, p. 772, 1792 (preoccupied 
by Sciurus vulpinus Gmelin) (Baltimore, Maryland); Sciurus ludovicianus vicinus 
Bangs, Proc. Biol. Soc. Washington, vol. 10, p. 150, 1896 (White Sulphur Springs, 
West Virginia). 

FIGS.: Audubon and Bachman, 1849, Quadrupeds of North America, vol. 1, pi. 17 
(col.) (as Sciurus cinereus); Bangs, 1896, pi. 8, fig. 3 (skull). 

The fox squirrels are typical of open heavy forests in the 
eastern half of the United States and extend in various sub- 
species from New England west to the plains region in South 
Dakota, and south to southern Florida, and to Texas and 
northeastern Mexico. In the more thickly settled areas of the 
East their numbers have been greatly reduced locally, and 
especially in the Northeast they have reached nearly the point 
of complete extirpation. The typical form, Sciurus niger niger, 
is still fairly common in the southeastern States, from southern 
South Carolina to Florida, frequenting chiefly the open "piney 
woods." The northeastern race*, S. niger neglectus, however, 
is gone from its more northern outposts and is in danger of 
complete wiping out east of the Alleghenies. 

The eastern fox squirrel is slightly larger than that of the 
Southeast and somewhat resembles a large gray squirrel. The 
upper parts of the head and body are a grizzled buffy and black, 
slightly darker on the forehead; cheeks, ears, forearms and 
feet, the lower hind legs and hind feet pale rufous; under side 
of body white; tail about as long as body, mixed black and 
buffy above, rufous below and on the sides. Total length, 
about 2 feet; tail, 11 inches; hind foot, 2.75 inches (73 mm.). 
Distinguished from the more western race, rufiventer, by its 
white instead of rufous belly, and from the southern typical 
form by its rufous ears instead of white, and the usual lack of 
black color, though rarely entirely black individuals occur. 

The eastern fox squirrel probably in former times just 
reached southwestern Connecticut but has long since gone. 
Linsley (1842), in his list of the mammals of that State, knew 
of it only from Northford, a century ago. In New York State 
it was formerly found in the southern parts in some numbers, 
as attested by Bachman in 1839. He mentions several lately 


received from Orange County but adds that in northern New 
York it is exceedingly rare, as he "only saw two pair during 
fifteen years of close observation." Merriam (1884) called it a 
rare or accidental straggler in the Adirondacks region and 
mentions a specimen or two at Lake George in 1872 or 1873 as 
the only recent instance he knew of, but there seems even 
there to be some doubt whether or not it had been brought in 
and escaped. In Rensselaer County he records two killed in 
1854. At about the same time Dr. A. K. Fisher knew of its 
having been killed in Westchester County, but adds that in 
1896 none had been known "of late even in that wild region." 
At the present time the fox squirrel is practically gone from 
the State. In New Jersey it was present locally at least up to 
about 1865 but was apparently gone by the nineties. In 
Pennsylvania the species was formerly common in the lower 
altitudes but is now much reduced. Rhoads (1903) says of its 
status in the early years of this century that it was "probably 
always rarer in Chester and Delaware counties and in southern 
N. J. than in south central Pa. and northern N. J. Now 
exterminated in N. J. but found occasionally in the Pa. counties 
bordering the lower Susquehanna, also yet recorded from the 
northwestern part of Pa." He quotes Todd that it was shot 
"at rare intervals in some of the northern counties" of the 
western border of Pennsylvania; and believed that it was 
destined to extermination within the entire limits of the State 
unless large areas of country in middle Pennsylvania "revert 
to a wilderness condition or become game reservations under 
state protection." Even in 1896 Bangs wrote that Dr. B. H. 
Warren informed him that "the northern fox squirrel is 
practically extinct in Pennsylvania except in the counties of 
Dauphin and Cumberland" near the south-central part of the 
State. In West Virginia, Kellogg (1937) reports that "it now 
survives in the heavily wooded and sparsely settled higher 
altitudes of the Allegheny Mountains" and that for many 
years numbers were shipped to Center Market in Washington, 
"from points in western Virginia and from eastern West 
Virginia." Apparently in all these regions the numbers have 
constantly and slowly dwindled in recent decades, for not only 
is the species constantly in demand for food, but it is less 
adaptable to the changing conditions of clearing and settle- 
ment than its congener, the gray squirrel. It prefers primeval 


stands of old timber and is much given to foraging and traveling 
on the ground. Hunters often employ dogs to tree the squirrels, 
after which they may be shot with little difficulty. At the 
present time it may be called locally uncommon in less settled 
areas from south-central Pennsylvania to the mountains of 
central Virginia and West Virginia. 

Family CASTORIDAE: Beavers 

Beavers of the genus Castor are the only living members of 
this family of rodents; they are of circumboreal distribution 
in the watered and forested regions of the north-temperate 
zone. Since prehistoric times they have been of great impor- 
tance in the economy of the human race, chiefly for their fur 
and their meat. The Old World beavers (Castor fiber and 
races) are regarded as a species distinct from the North Ameri- 
can species (C. canadensis), but the actual differences are slight, 
as in the proportionate depth of the rostrum (less in the Old 
World animal) and the length of the nasal bones (extending 
back of the lacrimal level more in the Old World animal). 
Of the New World beaver, a nurfber of slightly marked races 
have been described, depending on minor differences in color 
or in the size or the relative shapes of some of the cranial bones. 
Everywhere as the country has been opened and settled the 
beaver has been so reduced or extirpated that its present num- 
bers and distribution are but a fraction of their original extent. 
In various eastern localities reintroductions have taken place, 
with varying success. The literature on the American beaver 
is extensive, for probably few of our native mammals have been 
more often written about. In the following account of the 
various races only an outline of the main points in the history 
can be given, and in many cases it is hardly possible to obtain 
accurate details of recent date. The various races may be 
taken up in alphabetical sequence. 



Castor canadensis Kuhl, Beitrage zur Zoologie, 1820, p. 64 (Hudson Bay). 
FIGS.: Radford, 1908, 10 unnumbered pis. (col. exterior, photos, skull, works); Nelson, 
1916, p. 443 (col.); Elliot, 1901, p. 115, fig. 27 (skull). 


Beavers are stoutly built, the northeastern form weighing up 
to 60 pounds. The short, rounded ears and broadly webbed 
hind feet are adaptations to aquatic life; the coarse, brown 
overfur sheds water and protects the short, dense, and plush- 
like underfur. The broad, flattened tail is practically bare and 
covered with scales. Swimming is done with the powerful hind 
feet, while the small fore feet are held against the breast or 
used in transporting material. The skull is stoutly built, 
rather triangular in dorsal aspect, with broad nasals that end 
slightly behind the level of the upward branch of the premaxil- 
lary. In general color the eastern beaver in fresh pelage is 
dark chestnut-brown above, the base of the tail and thighs 
clearer tawny; below, the throat is buff, the rest of the under 
side drab. In this, the typical race, the tail is over twice as 
long as broad, a character distinguishing it from the Carolina 
beaver, which has a relatively broader tail. Total length, about 
1,100 mm. (about 35 inches); tail, 410; hind foot, 175. 

The range of this race formerly extended from about New 
Jersey northward to southern Labrador and Hudson Bay 
(Churchill region) and westward across Canada and Alaska, 
as far northward as the limit of trees, in well- watered country. 
Over all this area its numbers are now greatly reduced, or in 
places it is quite gone. Of the progress of this destruction only 
the merest outline can be given here. At the time when the 
first white settlers arrived in Massachusetts Bay and the Hud- 
son River region, beavers were plentiful along the streams, even 
near the coast. The earlier accounts contain many references 
to them and their works. Thus, Governor John Winthrop, in 
his "History of New England," tells how on January 27, 1631, 
he set out with a small company from Boston and "went up by 
Charles River about eight miles above Watertown, and named 
the first brook, on the north side of the river, (being a fair 
stream, and coming from a pond a mile from the river,) Beaver 
Brook, because the beavers had shorn down divers great 
trees there, and made divers dams across the brook." The 
name still persists, though the beavers have long since gone. 
Remains of beavers are common in the Indian kitchen middens 
all along our eastern coast, as in Connecticut, on Block Island, 
and the coast of Maine. In the earlier years of the Colonies 
beaver skins were one of the most important trade products 
of the country. These were obtained largely from the Indians 


who exchanged them for various trifles. Traders gathered 
together large quantities of beaver pelts, which were sent back 
to England. The magnitude of this industry may be gathered 
when we read in Winthrop's account that in April 1633 a 
vessel from Massachusetts, bound out for London, was wrecked 
on the Virginia coast with the loss among other things of four 
hogsheads of beaver weighing 900 pounds; and again, in the 
same year, he tells of a Mr. Graves sailing with between five 
and six thousand weight of beaver. Bradford's "History of 
the Plimmoth Plantation" lists seven sailings between 1631 
and 1636, in which a total of over 12,500 pounds of beaver was 
exported from the colony, with a total value of about 10,000 
sterling. Soon after this time, John Pynchon, of what is now 
Springfield, Mass., was the chief one to handle the furs brought 
in by the Indians from the surrounding region of the Connecti- 
cut Valley. His old account books show that in the six years 
between 1652 and 1657 he packed for England 47 hogsheads 
containing 8,992 beaver skins, weighing over 13,000 pounds, 
with an additional 663 pounds sent in bundles. From 1658 to 
1674 he packed 6,480 beaver skins. Many of these skins were 
from the country to the north add west of Springfield. Other 
furs also appear in the inventories, but in lesser quantities. 
At that time a beaver skin was sold at a standard value per 
pound, about 8 shillings; and in lieu of currency beaver skins 
were accepted at standard rates, which varied somewhat from 
time to time. The traditional "beaver hats" were made from 
the felted fur of this animal, and although these were mostly 
manufactured in England there were in later years some among 
the colonists who were skilled at this trade. For example, 
Deacon Ebenezer Hunt, of Northampton, Mass., manu- 
factured hats extensively for 40 years following 1734, buying 
his furs chiefly in Boston or Albany. After 1750, beaver hats 
sold at from 20 to 42 shillings each (Judd, S., "History of 
Hadley, Mass.," p. 355, 1863). At this time beavers were 
already becoming scarce in the southern part of New England. 
Albany, N. Y., then known as Beaverwyck, and New York 
City itself, then New Amsterdam, were the centers from which 
the Dutch East India Company's posts gathered in rich 
harvests of furs, while in the north the Hudson's Bay Com- 
pany, chartered in 1670, formed the outlet for the fur catch in 
that region. 


After a century and a half of constant trapping and with the 
spread of settlements into the eastern part of the country, the 
beaver was extirpated from the coastal regions of New England 
and greatly reduced in numbers elsewhere in the eastern 
United States. It is said that the Indians were careful to leave 
a certain proportion of the beavers to continue the stock, or to 
kill adult animals and leave the younger ones to mature, but 
the insatiate white trappers killed old and young indiscrimi- 
nately and took as many as possible for their immediate gain, 
with the result that the animals, bit by bit, were killed out from 
the more accessible parts of their eastern range. Yet the re- 
mains of their dams lasted for many years after the beavers 
had gone and often were mentioned as monuments in land 
conveyances, while the old beaver ponds silted up and became 
grassy meadows, where frequently the earlier settlers, pushing 
their way up the streams into the interior, found the only 
available pasturage for their cattle. In southern New Hamp- 
shire the last beaver in the town of Rindge is said to have been 
killed about 1780 (Stearns, E. S., "History of the Town of 
Rindge, N. H.," 1875); and the last one in the town of Peter- 
borough about 1790 (Smith, A., "History of the Town of 
Peterborough, N. H.," 1876). In the town of Pawlet, southern 
Vermont, the "last" beaver was killed about 1800 by one 
Ansel Whedon, who came upon it in his cornfield and slew it 
with a hoe (Hollister, H., Vermont Hist. Mag., vol. 3, p. 890, 
1877). On the Maine coast near Wells, the local, historian 
records that a trapper took 17 beavers there in 1755, but they 
seem to have quite gone before many years after. In northern 
New England, the beaver, though greatly reduced, has never 
been quite exterminated. Zadock Thompson, writing in 1842 
("History of Vermont," p. 39, 1842), believed that they were 
then nearly if not quite gone and mentions that the last one 
that he knew of was killed in Essex County, about 1830. 
Probably a few lingered in this northeastern corner of the 
State until much later, for at that time this region was nearly 
a primeval wilderness. Indeed, F. S. Hoag in 1909, after 
having made many inquiries among the trappers of Vermont, 
was credibly informed by a W. E. Balch that the last beaver he 
knew of in the State lived at Neale's Pond, north of Lunenburg, 
Essex County, about 20 years before (about 1890). He be- 
lieved that it was completely extinct in Vermont by 1909. 


In the adjacent part of New Hampshire there were beavers in 
the Connecticut lakes at least down to 1884, according to Ned 
Norton (Forest and Stream, vol. 24, p. 457, 1885). Beavers in 
small numbers continued in the remoter parts of Maine till the 
middle decades of the last century. At about that time the 
fashion for beaver hats abated in favor of silk hats, and the 
demand for fur was also lessened by the substitution of nutria 
fur from South America. Thoreau, in 1853, on his memorable 
journey to the Maine woods, relates that they were getting to 
be numerous again in the Lake Chesuncook region, but their 
skins brought so little that it hardly paid then to hunt them. 
J. G. Rich, an old trapper of those days and a correspondent of 
Agassiz, wrote ten years later that the fur of beavers com- 
manded only $2.50 a skin, whereas formerly as much as a 
dollar an ounce was paid. In 1866, legal protection was first 
given the beaver in Maine. Its numbers slowly increased 
until during the first decade of the present century it seems to 
have become locally common in the remoter townships, where 
it was not allowed to be killed at any time. Increasing com- 
plaints of timber owners that beavers were damaging their 
trees by cutting and flooding resulted in 1910 in the establish- 
ment of a short open season for two years in Somerset and 
Franklin Counties. In 1911 the area on which beavers could 
be trapped was extended to other counties. At the present 
time there seems to be no reason why a fair number of beavers 
can not be maintained in the wilder areas of the State. 

The history of the beaver in New York State has been re- 
viewed by Radford (1908), particularly its status in the 
Adirondacks. "As early as 1623 the importance of the beaver 
to the Dutch colony was so well recognized as to lead to its 
incorporation in the seal of New Netherlands." In those times 
Indians from the St. Lawrence Valley trapped beavers in 
northern New York and traded them to the French companies 
at Quebec. Merriam adduces evidence to show that in the 
1830's beavers in the Adirondacks were at a very low ebb, and 
DeKay, who traversed the height of land between the sources 
of the Hudson and the St. Lawrence Rivers in 1840, believed 
them nearly extirpated, though reported in 1841 "on Indian 
and Cedar Rivers, and at Paskungameh or Tupper's lake" 
and in scattered families in parts of Hamilton, St. Lawrence, 
and Essex Counties. Radford believes that by 1860 there 


could have been hardly more than 50 beavers remaining in New 
York State, chiefly along the Raquette and St. Regis Rivers 
and in the well-watered region northwest of Upper Saranac 
Lake in Franklin County. Here a slowly dwindling remnant 
remained for the next 30 years, their numbers gradually re- 
duced by occasional trappers in spite of efforts to protect them 
by a local inn-keeper on or near whose lands they lived. In 
the winter of 1894-95 probably not more than ten remained. 
In the latter year a law was passed removing an open season 
for beavers in the State, and in 1904 this was given added force 
by increasing the fine for taking beavers to $100 and making it 
an offense to set traps for them or to molest their dams or 
houses. From the beginning of this century for a number of 
years beavers showed a slight increase, and at the same time 
small numbers were introduced at favorable localities. Some 
of this stock came apparently from Canada; 30 or more others 
were obtained in 1906 from Yellowstone Park. Under protec- 
tion, and through this addition of new breeding stock, beavers 
have so greatly increased in New York State that during the 
season of 1924, when trapping was allowed, no less than 2,478 
pelts were taken, valued at $39,548, and the following year 
yielded 3,573 pelts, which brought in all $71,460 (Couch, 1937). 
Under the present law, the Conservation Department of New 
York may declare an open season in the month of March only. 
Of late years a two-week period has been given in counties that 
seem to have a good supply of the animals. The population, 
however, is so unevenly distributed that in some regions more 
beavers are desirable, while in others, which they are invading 
rapidly, it is doubtful if a beaver population can be maintained. 
Since 1924 to 1939, seven open seasons of two weeks each have 
been permitted. Each trapper is limited to six pelts to be 
taken in the two weeks, and these must be tagged by a game 
protector. In its practical application it is found that the 
protective law can be best enforced when blocks of counties 
are governed by like seasons, so that granting open seasons 
becomes in good part a problem of law enforcement. In 1939 
"there are not more than a half-dozen counties in the State, 
aside from those comprising Greater New York City, in which 
beaver colonies were not established. Even in Rockland 
County, whose border is but a half-hour's motor trip from 
Broadway, an open trapping season has been permitted for 


several years" (Dr. G. Bump, in Hit.). The following sta- 
tistics have been obligingly furnished by Dr. Gardiner Bump, 
Superintendent of the Bureau of Game, concerning recent 
beaver catches in New York State. In 1935, 2,498 pelts were 
taken in eleven counties; in 1936 there was no open season; in 
1937, a total of 2,014 pelts was taken in thirteen counties; and 
in 1938, in twelve counties, 2,629. It is thus clear that under 
wise management and intelligent supervision the bearver may 
become a considerable asset in regions where conditions are 
suitable for the maintenance of a breeding population. 

The story in Pennsylvania is much like that in New York 
State. Rhoads (1903), after giving in detail such older records 
as he could find concerning their former abundance and gradual 
extirpation in the State, summarizes these as follows: "It is 
evident that this interesting animal was practically extermi- 
nated in the eastern half of its Canadian habitat in Pa. about 
1830; that some remained in the headwaters of the west branch 
of the Susquehanna till about 1840, and that almost the last 
stragglers of their race were killed in Elk, Clarion, and Centre 
Cos., between the years 1850 and 1865." Beavers killed in 
Clinton County in 1884 and on* seen in Cambria County in 
1899 may have been escapes. It seems safe to say that beavers 
had been practically exterminated in Pennsylvania for 60 years 
when in 1917 a pair was imported from Wisconsin. From then 
until 1924, 94 beavers "were imported and set free in certain 
sections of the State at a cost of about $50 each. The animals 
increased so rapidly that it soon became necessary to transfer 
some of them to other sections. A survey by the State Board 
of Game Commissioners of the streams in Pennsylvania in 
1931 revealed 899 beaver dams with an estimated beaver 
population of 4,377, which by 1934 had increased to 15,000. 
During the 1934 trapping season, 6,455 beavers were legally 
taken, which, at the average price of $15 a pelt, brought the 
trappers a total of $96,825. In 1936, under increased trapping 
restrictions, 2,261 pelts were taken which brought a total 
return to the trappers of $22,610, an average price of $10 each 
. . . These financial returns demonstrate that the beaver 
constitutes a natural resource of great importance" (Couch, 

Profiting in part by the experience of New York and Penn- 
sylvania, beavers have in recent years been introduced into 


Vermont, central New Hampshire, and Mount Desert, Maine, 
and at the present time they seem well established in several 

From west of the Alleghenies to the edge of the Plains, 
beavers seem at the present time to be quite gone over most of 
their former range. For Ohio, Bray ton (1882) mentions their 
former abundance but implies that they had practically gone 
by the middle of the last century. In Indiana, Hahn (1909) 
gives 1840 as about the last authentic record, and both he and 
Lyon (1936) believe that the beaver of this State may have 
represented the southeastern race, while the latter adduces 
what seems a reliable recent record of the animal from Wells 
County in the northeastern part of the State and implies that 
it may have come in from adjacent parts of Michigan, where a 
few beavers still exist. Possibly, however, it was introduced, 
for in 1935 small "plants" of beavers were made in three 
counties, Jasper, Pulaski, and Starke, on game preserves, and 
these have so multiplied that in 1939 they had spread into ten 
other counties of northern Indiana, and additional "plants" 
have been made in Jennings and Clark Counties in the southern 
portion, where, however, the conditions of sufficient food and 
water are less favorable for them. Strict protective laws are in 
force, but in general land owners are interested in the encour- 
agement of the beaver, in part from the benefit to water con- 
trol and in part on account of the interesting habits of the 
animals (from Outdoor Indiana, Nov. 1939). 

Still farther west, Cory (1912) writes: "Beavers were for- 
merly common throughout Illinois and Wisconsin but at the 
present time they are practically exterminated in the former 
State," although "it is probable that a very few individuals 
may exist in the extreme southern portion," in Alexander 
County, whence fresh cuttings were obtained in 1900. In 1854, 
Kennicott wrote of beaver dams still existing at various points 
in Illinois, as if the animals had practically gone even by that 
day. In northern Wisconsin, however, they are now common. 

North of the United States, beavers still occur plentifully 
in the more remote parts of western Canada but have been 
largely reduced in the more accessible regions and, of course, 
extirpated in the settled areas. G. G. Goodwin (1924), writing 
of the mammals of the Gaspe Peninsula, Quebec, tells of finding 
a skull at the forks of the Ste. Anne River and adds that 


according to his guide "there were a few beavers left above the 
forks of the river and that they live in holes in the river bank." 
Dr. R. M. Anderson (1939a) states that though formerly 
common in the wooded parts of the Province of Quebec it is 
trapped out in most localities. However, "some projects for 
restoration of the beaver by setting aside preserves have been 
gratify ingly successful, particularly in the regions south and 
east of James Bay." Although beavers have been trapped in 
this region for nearly three centuries, since the establishment 
of the posts of the Hudson's Bay Company, they appear to 
exist in reduced numbers to this day. Preble (1902) writes, 
however, that owing to persistent trapping they were "becom- 
ing scarce throughout the region" over 30 years ago, although 
skins were annually traded at the Company's posts on the 
coasts of Hudson Bay. He noted "the remains of a beaver 
house between Pine and Windy lakes and a comparatively 
recent dam on a small stream which empties into Hayes River 
about 15 miles above York Factory. A number of skins were 
seen at Fort Churchill. These had been taken on the Lower 
Churchill River. Several black pelts were among the furs at 
Norway House." He mentions further that "Dr. Bell reports 
that a family of beavers was found by Indians on North 
River, a stream that flows into the Bay about 15 miles above 
Fort Churchill"; and that according to Hearne's account of 
his explorations in this region, published in 1795, the Indians 
accompanying him killed beavers on Seal River, the mouth of 
which is about 40 miles north of Churchill. This marks perhaps 
nearly the northern limit of the beaver on the west side of Hud- 
son Bay; of its northward limits on the east side of the Bay no 
recent information is at hand. Low, writing in 1895, after 
traversing parts of northern Labrador, says that it is common 
in the wooded regions and extends into the semibarrens where 
food is available. 

As to the recent status of beavers in the Athabaska-Macken- 
zie region still farther west, Preble (1908) has given a brief 
review. He quotes J. Alden Loring that in 1896 evidence was 
obtained of the former abundance of the beaver in suitable 
localities in western Alberta, but at that time it was nearly 
exterminated; tracks were seen on a small stream between 
Jasper House and Smoky River, but no other recent traces of 
the animals were found. Farther north it was formerly abun- 


dant nearly to the limit of trees; but on his journey in 1903-4, 
to the Great Slave Lake and Mackenzie region, he found it 
largely gone in many parts, and nowhere common, "though 
skins are received annually by all the posts throughout the 
region . . . The vast region which stretches from Great 
Slave Lake to the Rocky Mountains at present seems to be the 
best beaver country in the north. Many skins are brought 
from the upper reaches of Hay River by the Beaver Indians, 
and from Trout Lake by the Indians who frequent that 
locality. The Horn Mountain country also furnishes many 
skins. Along my route between Great Slave and Great Bear 
Lakes, the beaver has now become scarce, owing to constant 
hunting, but my guide intimated that in certain localities off 
the main route which we were following he knew of small 
colonies of beavers. About Great Bear Lake the best beaver 
ground seemed to be to the northward of Fort Franklin, and 
I saw several skins, some quite dark, just brought from the 
hunting grounds about two days' travel to the northward. 
During the winter of 1903-4 several beavers were killed by 
Indians in the region about Fort Simpson. In the spring the 
animals often descend the smaller streams to the main river 
and follow it to the mouth of the next tributary. A young one 
was shot near the mouth of the Liard in May, and several 
adults and young ones have been killed in recent years near 
the mouth of Bluefish Creek, opposite Fort Simpson, as a re- 
sult of this habit. 

"While descending the Mackenzie in the summer of 1904 I 
saw no beavers, but obtained information regarding the traffic 
in skins. About 700 skins were said to have been traded during 
the preceding winter at Fort Norman, which receives the fur of 
a very large extent of country. Skins from the country toward 
the Barren Grounds, according to the testimony of C. P. 
Gaudet, of that place, are smaller and average darker than 
those from the vicinity of the post. Fort Anderson, according 
to the fur returns, never received more than five skins annually 
during the few years of its existence." 

In the Yukon River Valley beavers are nearly extirpated. 
Dr. W. H. Osgood, who traversed the region in 1899, wrote 
(1900): "It hardly seems possible that half a million or more 
beaver skins have been secured in the Territory of Alaska. 
The animal is now almost as rare as it is in the United States, 


the inevitable result of continued pursuit by both whites and 
natives, which has so many parallels that it is useless to 
emphasize it here. At Fort Selkirk I saw several beaver skins 
taken on a small tributary of Stewart River and at St. Michael 
I found a very few in the warehouses of the trading companies. 
Beyond this I saw or heard nothing of them." 

Dr. Francis Harper states that under recent protective laws 
beavers are increasing in Alberta at the present time, and it 
seems likely that proper restrictions, if they can be enforced in 
all this region, will result in an increase of beavers, so that they 
may in time prove a regular and profitable source of revenue. 

Concerning the habits of the beaver a great deal has been 
written. Their general activities center around, first, the dams 
constructed solidly of sticks, branches, and larger pieces of 
wood, together with stones, mud, and other material from the 
pond bottom; then with the resulting formation of a pond, the 
construction of a house which serves as a shelter for young and 
adults, and has entrances under water into the pond. In the 
pond, aspen logs are sunk for a winter food supply, for the 
bark of this tree is a favorite food. An older colony will in 
time connect various pools with? canals and the dams may be 
extended to works of great magnitude. A single litter of from 
two to six young is produced in the course of a year, so that 
the increase of a colony may be fairly rapid and the young 
when of adult size move off to found new colonies. Extensive 
accounts of the habits may be found in the following works: 


ca. 1913. The romance of the beaver: Being the history of the beaver in the 
western hemisphere, xvi -f- 225 pp., illustr. Philadelphia and London. 

1922. Beaver habits, beaver control, and possibilities in beaver farming. U. S. 

Dept. Agric. Dept. Bull. 1078, 29 pp., 7 figs., 7 pis. 
1927. How beavers build their houses. Smithsonian Inst. Kept, for 1926, pp. 

357-360, pis. 1-6. 

1937. Beaver pioneers, xiv -f- 153 pp., illustr. New York and London. 

1892. Castorologia, or the history and traditions of the Canadian beaver, xvi + 

238 pp., illustr. Montreal and London. 

1913. In beaver world, xiv + 228 pp., illustr. Boston and New York. 

1922. The life of the Yellowstone beaver. Roosevelt Wild Life Bull., vol. 1, 
pp. 187-221. 



1926. A study of the beaver in the Yancey region of Yellowstone National 
Park. Roosevelt Wild Life Annals, vol. 1, pp. 13-191, illustr., maps. 

1926. Notes on the beaver colonies in the Longs Peak region of Estes Park, 
Colorado. Roosevelt Wild Life Annals, vol. 1, pp. 193-234, illustr. 

1927. The beaver: Its work and its ways. Amer. Soc. Mammal. Monogr. 2, 
177 pp., illustr. Baltimore. 


Castor canadensis baileyi Nelson, Proc. Biol. Soc. Washington, vol. 40, p. 125, Sept. 26, 
1927 (Humboldt River, 4 miles above Winnemucca, Nevada). 

This race of the beaver is darker than frondator and has a 
slenderer skull. From C. c. pacificus from the Columbia River 
drainage in eastern Washington and Oregon it differs in its 
distinctly paler color and slenderer skull. The narrow skull 
and especially the rostrum of the Humboldt River beaver 
contrast strongly with the massive skull and broad heavy 
rostrum of subauratus (Nelson, loc. cit.). The upper parts 
are described as "dull rusty chestnut, brightest on crown 
with a dull yellowish shade on the cheeks; ears dark brown; 
base of tail all around uniform with adjacent parts of body; 
tops of hind feet dark chestnut; underparts of body dull drab 
brown." Total length of type, 1,064 mm.; tail, 254 by 135; 
hind foot, 183. 

Concerning this race of the beaver little seems to be in 
print. Dr. Nelson had specimens from Winnemucca, Iron 
Point, Golconda, and Deeth in Nevada. It is presumably con- 
fined to the region of the Humboldt River Basin of Nevada. 
Borell and Ellis (1934) wrote of it a few years ago: "Beavers 
are found in limited numbers along the main branches of the 
Humboldt River, but have been almost completely extermi- 
nated along the smaller streams which flow out of the Ruby 
Mountains. Mr. August Rohwer and Mr. William Toyn re- 
ported that there were still a few beaver in one or two of the 
canyons on the west slope of the Ruby Mountains. Character- 
istic beaver gnawings found by us on some partly decayed aspen 
stumps at about 7,000 feet altitude along Toyn Creek, near the 
summit of Harrison Pass, indicated the former presence of 
beaver there." 

Still more recently, Vernon Bailey (1936) has reported that 
"these desert-valley beavers are still found in the Great Basin 


drainage of northern Nevada along the Humboldt River, and 
its tributaries and in the Malheur Lake and Steens Mountains 
drainage of southeastern Oregon . . . There are a few 
beaver all along the Blitzen from its headwaters down to near 
Malheur Lake, and George Benson reports one shot at the edge 
of Malheur Lake in 1909. In 1916 they were found in Big 
Fish Creek, McCoy Creek, and Kiger Creek. It is fair to 
assume in the absence of specimens that the beaver in the 
Silvies River and its branches on the north of Malheur Lake 
are also of this subspecies. In 1920 they were still common in 
many places along the Silvies and Blitzen Rivers, and while in 
places they were doing some damage by flooding the meadows, 
more often they were merely holding up the water to a better 
depth and improving the meadows by subirrigation." Con- 
cerning their former abundance in this last region, Bailey 
quotes from older records to show that a little over a hundred 
years ago, in 1826 and 1828, beavers were so numerous that a 
party of trappers took over 300 in a month. 

On occasion, Bailey adds, they may dam up irrigation 
ditches or locate in the banks of streams near fields and or- 
chards where they do serious mischief and have to be removed, 
driven away, or destroyed. With intelligent handling, "how- 
ever, they could become a valuable asset on many of the 
eastern Oregon ranches." 


Castor canadensis belugae Taylor, Univ. California Publ. Zool., vol. 12, p. 429, Mar. 
20, 1916 ("Beluga River, Cook Inlet region, Alaska"). 

According to its describer, this race of beaver is the one 
found from central British Columbia northward along the 
coast of the mainland to the mountains of Alaska. Specimens 
referred to the race are mentioned from Stuart Lake, British 
Columbia, and from various localities on the shores of Cook 
Inlet, Alaska. In color the single skin available to the describer 
was said to be slightly paler than in the neighboring races, 
phaeus and leucodonta. The skull, in comparison with that of 
the latter, is "immediately distinguishable through the nar- 
rower blades of the hamular processes of the pterygoids," with 
a tendency for the maxillary tooth row and ratio of this to the 
basilar length to be greater. 


In the region of the type locality, beavers were becoming 
scarce 40 years ago. Osgood (1901) mentions three secured by 
trappers near the mouth of Turnagain Arm in 1899 and adds 
that a few were taken every season along streams in the moun- 
tains about 60 miles inland from Tyonek. "A trading station 
on the lower Sushitna River also obtains a small quota annually. 
Compared with former receipts, however, the number now 
obtained is lamentably small." In a later report Osgood (1904) 
adds that a little farther to the westward on the Chulitna River 
he found evidence of a few scattered small colonies of beavers. 
"The extensive area of low land about the sources of the 
Chulitna River is covered with hundreds of small lakes and 
ponds connected in most cases by small, sluggish streams 
eminently suitable for beavers, and no doubt a great many are 
still scattered throughout this area." A small number of skins 
were annually brought in to the trader at Nushagak at the 
mouth of the river of the same name. Farther south, in the 
Atlin region of northwestern British Columbia, Swarth (1936) 
writes that "as everywhere, beavers hereabout are trapped to 
the verge of extinction. They in all likelihood were originally 
abundant and generally distributed throughout the lowland 



Castor caecator Bangs, Bull. Mus. Comp. Zool., vol. 54, p. 513, July, 1913 ("Near 

Bay St. George, Newfoundland"). 
FIGS.: Bangs, 1913, figure on p. 514 (skull). 

Although the Newfoundland beaver was first described as a 
species distinct from typical C. canadensis of the neighboring 
mainland, it can scarcely be considered as more than the local 
representative of that animal, which through a long period of 
isolation has become slightly different in certain cranial char- 
acters. The color has not been accurately described on account 
of the lack of specimens in museum collections, but Bangs, in 
naming it, mentions that he had seen two in the flesh, which, 
though showing nothing distinctive in external appearance, 
nevertheless appeared "rather small" and were of an "excep- 
tionally fine rich color." The distinctive characters of the 
skull are: the slightly smaller size as compared with the main- 
land beaver, typical canadensis; the wider, "more roundish" 


interparietal; and the lighter, less flaring zygomata, the outer 
side of which is more nearly straight, thus giving a much more 
triangular outline to the skull as seen from above. The nasals 
are also said to be shorter and wider. 

Concerning the present status of the beaver in Newfound- 
land, little accurate information is at hand. Bangs, in 1913, 
wrote that "though much trapped for its fur, it still occurs in 
fair numbers in the remoter parts of the island," and probably 
this still holds true. It doubtless finds the more barren portions 
of the country less suitable to its needs and hence can be 
expected to thrive best in the western and central parts of the 
island. Dugmore, in his "Romance of the Beaver," published 
about 1913, writes that when he first visited Newfoundland 
about 1900 and in annual visits during the three succeeding 
years, he never saw but one beaver colony, and that a very 
small one in a remote and rather inaccessible part of the 
country. Under governmental protection, however, beavers 
multiplied in following years so that on visiting the same 
region in 1912 he counted no less than 27 lodges within a short 
day of walking and canoeing. As elsewhere, this illustrates 
how readily beavers will breed up to the carrying capacity of 
the land if free from persecution, for their natural enemies 
are few. 

For a comment on the recent status of beavers in Newfound- 
land I am indebted to Frank Strickland, who is familiar with 
conditions there. He says that previous to the declaration of 
a close time on beavers, about 1917, they were plentiful in 
Grand Lake and Red Indian Lake, but today they are very 
scarce in that section. What beavers are left in Newfoundland 
are mostly to be found between Port Saunders north to White 
Bay and between Gander River east to the bottom of Bay 
d'Espoir, including thus the central districts of the island. 



Castor canadensis carolinensis Rhoads, Trans. Amer. Phil. Soc., new ser., vol. 19, p. 
420, Sept. 1898 ("Dan River, near Danbury, Stokes County, North Carolina"). 
FIGS.: Rhoads, 1898, pi. 23, figs. 1, 2 (skull). 

The beaver of the southeastern United States is broader- 
tailed than typical canadensis., and in this respect it resembles 


the southwestern race frondator, which, however, is paler in 
color. Rhoads describes the type as bright hazel above, with 
seal-brown underfur, the hinder back shading into bright hazel 
to cinnamon-rufous and tawny-olive; ears blackish, feet bister. 
Below, dark broccoli brown, the tips of the longer hairs wood 
brown. Total length, 1,130 mm.; tail, 279 by 158; hind foot, 
184. The skull is characterized by its short, broad nasals and 
short rostrum, with the former scarcely or not extending back 
of the level of the orbits. 

The limits of the range of this race, though in general from 
northern Florida to the Mississippi and northward to the low- 
lands of New Jersey and Ohio, Indiana, and Illinois, will 
probably never be exactly definable since it is now extinct 
over the greater part of the area and but few complete speci- 
mens are extant. Rhoads believed that it intergraded with 
typical C. canadensis in southern New Jersey and the lower 
parts of Pennsylvania, and Cory (1912), in his map of the 
range, included southern Ohio, Indiana, and Illinois in the 
area of presumed intergradation. Lyon (1936), who figures an 
imperfect skull as the only one extant from Indiana, assumes 
that it was probably the form occurring in that State. 

Rhoads believed that by 1700 beavers had been practically 
exterminated from the lowlands of the Delaware, Schuylkill, 
and Susquehanna Valleys, which were the regions first settled ; 
while by the time of the American Revolution, they were 
practically wiped out from all the lowland valleys of Pennsyl- 
vania except the northern tributaries of the Ohio; but of this 
there is almost no definite record. In New Jersey, owing to 
the inaccessible and unproductive character of the lands in 
the southern part of the State, beavers persisted longer, in the 
most retired swamps of Atlantic and Cape May Counties, but 
were probably quite gone by 1830, or earlier. Traces of their 
old dams persisted for many years later, and Dr. Witmer 
Stone, writing in 1908, mentions remains of a very large one 
that he had visited on the Nescochaque branch of Mullica 
River. In 1900, according to Dr. Stone (1908), there was a 
colony of beavers in Sussex County, N. J., due apparently to 
individuals that had escaped from a private preserve where 
they had been introduced. 

Atwater's " History of Ohio," published in 1838, speaks of 
beavers as formerly common on the headwaters of the larger 


streams in that State but adds that "they have long since 
disappeared." For Indiana much the same story is true; Dr. 
Lyon (1936) quotes the Prince of Wied's statement, in 1832 
and 1833, that beavers were then abundant along the lower 
Wabash, but with the opening up of the country to settlement 
in succeeding decades, they must have disappeared rapidly. 
Nevertheless, he adduces some evidence for the belief that in 
Wells County, Ind., beavers were present only a few years 
since, possibly introduced; while similarly, Cory (1912), al- 
though he writes that beavers are practically extinct in Illinois, 
believes it "probable that a very few individuals may exist in 
the extreme southern portion of the state." 

To the southward of this tier of States that may have marked 
the northern limits of this beaver's range, information is 
meager, but it indicates that the animals still hold on in a few 
widely separated localities, where with real protection there is 
some hope that they may continue and even increase con- 

Within the memory of men now living beavers were present 
in small numbers in the counties a short distance south of 
Richmond, Va. A writer in Forest and Stream (vol. 7, p. 197) 
for November 2, 1876, states that for the two years previous 
to that time trappers had been taking beavers in Dinwiddie, 
Nottoway, Brunswick, Cumberland, and other counties, some 
making good returns on these and other furs. At that time 
the pelts of beavers sold for a dollar apiece. Here, too, the 
correspondent mentions having seen in some of these localities 
places where beavers had totally destroyed acres of corn, caus- 
ing serious loss. At the present time, I have no information of 
any beavers remaining in the State. Possibly, however, a few 
may have persisted in the mountainous parts of this State 
and of West Virginia. F. E. Brooks (1911) mentions what he 
believes to be a well-authenticated case of a beaver having 
been killed in Pocahontas County, W. Va., about 1907, al- 
though this was possibly brought in from elsewhere. Still 
later, A. B. Brooks (1923) published a note on the reappearance 
of beavers in West Virginia. He writes: "A few of the older 
trappers and hunters still living state that beaver work was 
observed by them many years ago on the headwaters of the 
Williams and Greenbrier Rivers," but they had been "counted 
as extinct in West Virginia for fifty years or more" until the 


early fall of 1922, when a farmer in Hampshire County re- 
ported finding fresh beaver work on a creek tributary to North 
River. Subsequent investigation disclosed a colony of beavers 
here. They had "built a dam and done much cutting along 
both banks of the stream." All efforts to learn of the origin 
of this colony were unsuccessful. 

Beavers persisted in North Carolina until recent years. 
The series of specimens from Dan River, near Danbury, Stokes 
County, on which the race carolinensis was based, was col- 
lected between 1897 and 1899. C. S. Brimley, writing in 
1905, says that it occurs sparingly there and is also reported 
from Bertie County. A specimen taken 15 or 20 years pre- 
viously at Weldon is in the State Museum at Raleigh. These 
localities are all on the Roanoke River or its tributaries. He 
adds that "Mr. J. H. Armfield reports a few occurring in 
Beaver Swamp in the northern part of Guilford County and 
southern part of Rockingham, and Mr. K. E. Shore reports 
them from the Yadkin River, between Yadkin and Forsyth 
Counties." At the present time, 1939, the beavers seem to 
have quite disappeared. 

No recent record of the beaver in South Carolina is known 
to me. In Georgia, however, a few still remain and are care- 
fully protected, in the Flint River section in the southwestern 
part of the State and at the headwaters of the Chattahoochee. 
Dr. Francis Harper (1927) in his account of the mammals of 
Okefenokee Swamp, Ga., adduces evidence that they were 
formerly present there. He was shown an incisor tooth taken 
from a beaver killed about 1890 by an old Negro woman a 
mile or so east of Lloyd's Island. "The story goes that the 
woman came upon the animal in the road and clubbed it to 
death." Dr. Harper suggests that the southward range of the 
beaver may have been to a large extent limited by the presence 
of the alligator. He doubts if it ever regularly inhabited the 
interior of Okefenokee Swamp, else it would probably have 
survived there or left some tradition of its earlier presence. 
In former times there were beavers in extreme northern 
Florida (Bartram mentions them), but little or nothing is 
known of the time of their disappearance. An unmistakable 
molar tooth was lately found in an Indian mound on Indian 
River by J. H. Rowe. 

To the westward of the Alleghenies, the Carolina beaver is 


believed to have extended to the Mississippi, perhaps as far 
north as Iowa. In this State, according to J. A. Allen (1870a), 
they had been nearly or quite exterminated in most of the 
eastern and southern portions by 1870, although at that time 
a few were reported still to exist on the South Raccoon River. 
Kellogg (1939) has lately summarized its history in Tennessee, 
where it was originally well distributed in former times and 
apparently did not begin to show much depletion until after 
the middle of the eighteenth century. About 1788 the salaries 
of the county clerks and others were paid in beaver skins, 300 
for the county clerk, 200 for the clerk of the House of Com- 
mons, and three for members of the Assembly. The latest 
records for beavers seem to be those of Rhoads (1896) for the 
extreme western part of the State. He examined a beaver 
house in the cypress swamp bordering Reelfoot Lake, Obion 
County, about 1895 and was told that there were then about 
20 beavers in that district. Another colony was reported to 
him from Hay wood County. Beavers still exist in Alabama, 
and according to A. H. Howell (1921) have held their own in 
some localities remarkably well, even in settled farming regions. 
He says that "apparently they are more numerous at present 
[1920] in the central part of the State, in Montgomery and 
Lowndes Counties, than in either the wild hill country of the 
northern part or the big swamps of the south." Old residents 
of Montgomery County assured him that although abundant 
there in early times, they had entirely disappeared from the 
region, but about 1908 reappeared, coming up from the lower 
stretches of Catoma Creek. In Lowndes County, L. J. Gold- 
man found beavers in numbers along Big Swamp Creek and in 
Jones Lakes, where they do considerable damage to the corn 
patches. They are little trapped for fur, although their meat 
is much sought after. They now have protection either locally 
by the planters or in general by State law, from March to 
November. "In the northern part of the State beavers seem 
almost to have been exterminated; they formerly occurred in 
small numbers in the Tennessee River at Muscle Shoals, but 
disappeared about 1895; they lived about the 'towheads' 
(small islands) in the river and burrowed into the banks, but 
did not build dams. A few were reported in 1916 in Big Wills 
Creek, near Collinsville. They disappeared from Talladega 
Creek, near Dean, about 1896." 


These southern beavers attain a large size; Howell speaks of 
one killed on Pintlala Creek that weighed 65 pounds. Another 
from Catoma Creek weighed 38.5 pounds and measured 1,035 
mm. in total length; tail, 290 by 163; hind foot, 170. Compared 
with a specimen of the typical race from Canada, its skull is 
shorter and relatively broader, with wider brain case and 
interorbital region. 

In Kentucky, which should come within the western range of 
this race, Funkhouser (1925) writes: "We can find no record 
of its having been seen . . . within the past twenty 
years." In earlier times it was occasionally mentioned by the 
pioneers, but "was probably never very abundant in Ken- 



Castor canadensis concisor Warren and Hall, Journ. Mamm., vol. 20, p. 358, Aug. 14, 
1939 ("Monument Creek, southwest of Monument, El Paso County, Colorado"). 
FIGS.: Warren, 1910, p. 141, figs. 46-48 (photographs; skull). 

Until recently the beaver of Colorado was believed to be the 
same as the race frondator, typical on the Mexican boundary of 
Sonora. Now, however, the above authors, after a minute 
study of a series of skulls, believe that it represents a recog- 
nizable subspecies, of dark color, but with a skull resembling 
that of mexicanus, yet on the average differing in the size and 
shape of the angular process of the mandible, which is longer 
and more produced posteriorly and sharp-pointed rather than 
short and posteriorly rounded. Other slight differences seem 
to separate the Colorado beavers from adjacent races. Greatest 
occipitonasal length of skull, to 145.2 mm.; length of nasals, 

Warren (1910) writes that although the beaver "was no 
doubt at one time found in every county in Colorado which 
contained streams with sufficient water for its needs, and to- 
day, in spite of the persecution to which it was at one time 
subjected from the trappers, nearly resulting in its extinction, 
it is found over a large area of the State, and thanks to the 
protection accorded it by law, is on the increase. We have 
records of it from Larimer, Weld, Morgan, Grand, Routt, 
Arapahoe, Gunnison, Delta, Garfield, Eagle, El Paso, Teller, 
and Mineral counties," thus covering much of the west-central 


mountainous region of the State. The same author (Warren, 
1926) has given a detailed account of beaver colonies in Estes 
Park, Colo., with descriptions and photographs of their haunts 
and works, as well as maps illustrating their engineering enter- 
prises. Under protection the beaver seems for the present 
fairly safe in Colorado. 



Castor canadensis frondator Mearns, Preliminary Diagnoses of New Mammals of the 
Genera Sciurus, Castor, Neotoma and Sigmodon, p. 2, 1897; reprinted in Proc. U. S. 
Nat. Mus., vol. 20, p. 502, Jan. 19, 1898 (San Pedro River, Sonora, Mexico, near 
monument No. 98 of the Mexican boundary line). 

FIG.: Mearns, 1907, p. 351, fig. 57 (skull). 

Like the southeastern beaver (C. c. carolinensis) , this is a 
broad-tailed race, but its color is paler and brighter. Vernon 
Bailey (1931) describes it as light chestnut above; middle of 
the belly reddish chestnut; hind feet dark chestnut; colors 
slightly darker in winter. Total length, 1,070 mm.; tail from 
anus, 360; hind foot, 185; skull length, 133, its width 99. 
Weight of type (a female), 62 pounds. Old males are slightly 
larger than females and may measure up to 1,130 mm. in length. 

This race of beaver extends slightly to the south of the 
Mexico-United States boundary in northern Sonora and is 
found in Arizona along the Colorado River and in the extreme 
western part of New Mexico on branches of the Gila and San 
Juan Rivers. According to Bailey (1931) it is the form found 
from all points whence specimens were examined along the 
Colorado River system, but although formerly numerous, it is 
now everywhere much reduced in numbers. "There are still a 
few beavers in the headwaters of the Gila . . . also some 
in the San Juan River and its tributaries" in western New 
Mexico. Mearns (1907) has also given many details of its 
presence during the latter part of the last century along the 
Mexican boundary of Arizona. Within the rather restricted 
range at present assigned to this race, it is evidently rare and 
likely to be further reduced in the future. 




Castor canadensis leucodonta Gray, Ann. Mag. Nat. Hist., ser. 4, vol. 4, p. 293, Oct. 

1869 (Vancouver Island, British Columbia). 
FIGS.: Taylor, 1916, p. 450, fig. J, a; p. 451, fig. K, a; p. 452, fig. L, a (views of skull). 

Dr. W. P. Taylor (1916) has briefly summarized some of the 
characters that he believes distinguish the Vancouver beaver. 
Externally it closely resembles typical C. canadensis from New 
Brunswick in color above, but the under side is "about hair 
brown in leucodonta, near bone brown, dark grayish brown, 
dark vinaceous-drab or natal brown in canadensis. " Cranially, 
the external outline of the nasals is different, "tending to be 
more parallel in leucodonta than in canadensis, in which there 
is a dilation in the outline anteriorly; foramen magnum slightly 
broader . . . ; hamular processes . . . definitely 
broader bladed. " There are other slight cranial differences 
between this and neighboring races. 

In the middle of the last century beavers were common on 
Vancouver Island, to which this race is now regarded as re- 
stricted. Robert Brown (in Green, A. H., 1869), who collected 
the original specimens for the British Museum, spoke of them 
near Victoria, and especially in the interior, where "they are 
almost everywhere abundant and on the increase. In a swampy 
lake near the mouth of the Cowichan Lake we found many; 
and an extensive swamp near the entrance of the Puntledge 
Lake was a great stronghold ... In the spring of 1866, 
when crossing the island from Fort Rupert to the head of 
Quatseeno Sound with some Indians, a great portion of our 
route lay among these beaver ponds and dams. All through 
this district beavers swarm. " 

Swarth (1912) has presented some measurements and cranial 
dimensions from a series of this beaver trapped in 1910 on 
Vancouver Island. The external appearance, he says, differs 
from that of beavers from southeastern Alaska principally in 
slightly paler color. He found that "as a result of a number 
of years of protection beavers have multiplied on Vancouver 
Island so as to be really abundant in many places . . . 
There was a small colony at a point near Errington, but we 
found them in far greater numbers at Beaver Creek, near 
Alberni . . . There was old beaver work along the streams 


throughout this valley, and from the size of some of the old 
dams they must have existed here in immense numbers at one 
time, but at present, though there are a good many left still, 
they are more scattered, apparently not more than one or two 
families at any one point, and such groups separated by inter- 
vals, sometimes of several miles. Nearly all the smaller streams 
were obstructed by their dams, and most of the adjoining low 
lying land flooded as a consequence . . . We saw no 
beavers at Nootka Sound, but were told that there were many 
at Vernon Lake, some eight or ten miles inland from the head 
of Tahsis Canal" and a few at Central Lake. 

In response to inquiry, Maj. Allan Brooks writes me under 
date of July 8, 1939: "Beavers still exist in some numbers on 
Vancouver Island. There are few localities where the trapping 
is not in the hands of licensed trappers, who pay $10 a year for 
a denned area where no other trappers are allowed. By this 
method they can conserve their furbearers. There are locally 
closed areas also, as at Comox, Vancouver Island, where no 
beavers are allowed to be trapped and they are numerous. 
Over the whole northern third of this island the beavers of late 
years have been decimated by co\igars; it will take them many 
years to recover. " 



Castor canadensis mexicanus Bailey, Proc. Biol. Soc. Washington, vol. 26, p. 191, 
Oct. 23, 1913 ("Ruidoso Creek, 6 miles below Ruidoso, New Mexico"). 

The Rio Grande beaver is closely related to the race frondator 
with which it had been associated until Bailey distinguished it 
in 1913 as slightly different in its duller and paler colors, with 
very little chestnut at any season. He gives the weight of an 
adult as 47 pounds and the total length of an adult female as 
1,070 mm. 

The geographic range of this race is believed to be only the 
Pecos and Rio Grande drainage in New Mexico and Texas. In 
northern New Mexico it shows a tendency to intergrade with 
the darker frondator. The most comprehensive account of it is 
that of Bailey (1931), who has gathered what information is 
available as to its habits and status. In earlier times the beaver 
was a source of food and fur for the Indians in the region and 
its remains may be found in their kitchen middens. Bailey 


mentions that in 1826 Pattie and a party of trappers took 
beavers all along the Pecos River from 20 miles above its 
mouth to the first Spanish settlement (probably Anton Chico). 
From 1881 to 1884' L. L. Dyche collected on the headwaters of 
the Pecos and found at that time "a considerable number of 
beavers about 15 miles from the head of the Pecos River" ; there 
were many beaver dams and, he adds, "in the pools which 
were caused by these dams I found the best trout fishing of any 
locality I have ever visited in the Rocky Mountains," an 
interesting comment on the effect of such dams in making 
pools where trout may live. In 1898, C. M. Barber found a 
large colony on Ruidoso Creek, below the town of the same 
name, and there he secured the series of specimens on which 
Bailey later founded the name mexicanus. At that time the 
beavers were feeding on corn, which they cut and dragged to 
the stream and floated to their dens. Very few were being 
killed by the Mexicans on the ranches, since they lacked the 
technique for trapping. In 1902 Bailey visited Ruidoso Creek 
" and found that there were still a few beavers along this stream, 
which, with its sections of deep water and steep banks, is 
peculiarly adapted to the habits of beavers. " In the following 
year he also found some of the streams inhabited by them on 
the headwaters of the Pecos. "There were old cuttings along 
many of the other streams, but in most cases the beavers had 
been entirely trapped out ... In the deep water of the 
larger valley streams they are not so easily caught, and fortu- 
nately enough have escaped in spite of persistent trapping to 
prevent the complete extermination of the species . 

"In many places along the canyons of the Rio Grande above 
Santa Fe there were still some of the animals in 1903-4, and 
trappers were then catching them in considerable numbers 
. . . and in 1909, E. A. Goldman saw some old beaver 
cuttings near Socorro and was told that there were still a few 
along the river. He also saw signs of them at Garfield and 
found them common in the Rio Grande near Las Palomas, and 
they were reported near Las Cruces ... In 1915, J. S. 
Ligon reported them as becoming abundant in places along the 
Rio Grande above and below San Marcial, where there were 
some complaints of their felling trees across the fences. 

"In such localities beavers live entirely in the banks of the 
rivers and select the deepest water for their operations. They 


are not easily trapped and usually remain the longest where they 
do real damage, while in the higher mountains where they can 
do no damage they are easily caught and quickly destroyed. " 
While from an economic standpoint beavers may do some 
damage to forest growth and even to crops, as in the case just 
noted, by feeding on cornstalks, nevertheless this is easily con- 
trolled. Bailey mentions that in 1910 the water company of 
Santa Fe was offering $50 a pair for live beavers to be placed 
in the upper part of the Santa Fe Canyon to aid in conserving 
the city's water supply. He adds : "On almost all the mountain 
streams they should be protected and encouraged. A series of 
beaver ponds and dams along the headwaters of a mountain 
stream would hold back large quantities of mountain water 
during the dangerous flood season and equalize the flow of the 
streams so that during the driest seasons the water supply 
would be greatly increased in the valleys. Beaver ponds not 
only hold water but distribute it through the surrounding soil 
for long distances, acting as enormous sponges as well as 
reservoirs. A series of ponds also increases the fishing capacity 
and furnishes a safe retreat for the smaller trout and protec- 
tion from their enemies . . * . A legitimate amount of 
trapping should eventually yield large annual returns over 
extensive areas of the country from which they have been 
almost exterminated." 


Castor canadensis michiganensis Bailey, Proc. Biol. Soc. Washington, vol. 26, p. 192, 
Oct. 23, 1913 (Tahquamenaw River, 5 miles above falls, Luce County, Michigan). 

This is a very dark race of beaver, with a short, broad skull. 
According to its describer, it is dark umber-brown above, 
brighter on head and cheeks; ears, feet, and nose black. Under 
side darker than back, with blackish on the breast and flanks. 
The skull has the rostrum shorter and the nasals more quadrate 
than in typical canadensis, the zygomata abruptly spreading, 
the occiput with an upright crest, giving a short, "sawed-off " 
appearance. Though said to be of small or medium size, the 
measurements given do not show this: Total length, 1,170 mm.; 
tail, 470; hind foot, 185; skull, condylobasal length, 129; 
zygomatic width, 96.4; nasals, 46 by 24 (type skull). 


Although described from the northeastern (or "upper") 
peninsula of Michigan, between Lake Superior and Lake 
Michigan, the range of this race is believed to include much of 
the "Lower Peninsula," now or in the recent past. While at 
the present time beavers are regarded as extinct in southern 
Michigan, they still occur locally in northern parts of the State. 
According to Dice, they were found up to about 1920 along 
Boyne River in Charlevoix County but are now rare or extinct 
there except that beavers have been reintroduced on Spring 
Branch. In the neighboring counties, Cheboygan and Mont- 
morency of the Lower Peninsula, they are scarce but seem to 
be increasing. Presumably intergradation of this race with 
neighboring races takes place in Wisconsin or to the northward, 
but the exact limits of the range of this form remain to be 
worked out. 

Because of the rich dark color of its fur, this race of beaver is 
especially recommended by Vernon Bailey for introduction 
into other and now depleted localities. It has been much used 
in restocking areas in Michigan that seemed suitable, and Ruhl 
and Lovejoy (1930) have published a summary of such intro- 
ductions carried out in various counties in southwestern, 
central, and northern parts of the Lower Peninsula in recent 
years. In most cases only one or two animals at a time were 
available for this purpose, yet these authors conclude that 
"there seems to be no difficulty in getting beaver to settle and 
winter even if planted as late as November 1," but there is no 
assurance that they will remain in any particular locality nor 
was the number necessary for efficient colonization determined. 
Where large streams are available, these introduced beavers 
were found usually to make burrows in banks rather than to 
construct lodges. According to Vernon Bailey (1922), choice 
skins of this dark form from northern Wisconsin have in recent 
years brought as much as $50 each. He writes further: "So 
far as at present known, the darkest, richest-colored, and 
handsomest beaver fur is found native along the south shore of 
Lake Superior, in northern Michigan and Wisconsin [and is of 
the race here under discussion] . In this region of heavy forest 
and deep snows the outer hairs of the animals are very dark 
brown and the under fur is almost black. When tanned and 
plucked the skins are very beautiful, and when made up into 
wearing apparel they almost equal sea otter in depth of fur 


and richness of color. They are decidedly superior to the 
Canadian and Alaskan skins . . . For many years the 
beavers of the region south of Lake Superior have been care- 
fully protected . . . They are now fairly abundant in the 
region. " 



Castor canadensis missouriensis Bailey, Journ. Mammalogy, vol. 1, p. 32, Nov. 28, 
1919 ("Apple Creek, 7 miles east of Bismarck, North Dakota"). 

This race is said to be "slightly smaller than canadensis and 
much paler and duller brown. Skull more triangular in outline, 
not so massive and heavy ; much like that of mexicanus, shorter 
and heavier than that offrondator. From mexicanus the colors 
differ in being noticeably duller and darker; from frondator, 
duller, and not so rusty. " The total length of the type (prob- 
ably not fully grown) was 900 mm. "Specimens show closer 
affinity with those of the Rio Grande drainage, than with those 
in the same State [North Dakota] in the streams flowing into 
Hudson Bay." 

The range of this form of beaver is believed to be the drain- 
age system of the Missouri River, typically in western North 
Dakota, but further study is needed to settle this. Provision- 
ally, South Dakota, Nebraska, and possibly Kansas may be 
included in its present or former range. Vernon Bailey (1926) 
has given an excellent account of the recent status of the beaver 
in North Dakota. He writes that in 1804-5, Lewis and Clark 
found beavers abundant along the Missouri, even near long- 
established Indian settlements. At that time trappers were 
just beginning to find the region a source of pelts, and in the 
three decades following they reaped a large harvest. In 1833 
Maximilian reported that 25,000 beaver skins had been bought 
during the year at Fort Union (now Buford); by the next 
decade, however, Audubon found them already scarce. Never- 
theless, Bailey writes, where "the beavers had the protection 
of the deep water and high banks of the larger rivers" they 
persisted and "with characteristic tenacity they still cling to 
their old haunts or merely scatter out to establish new colonies 
in tributary streams." As late as 1913, he and others found 
small numbers as at Buford, Fort Clark, and near Williston. 
At this time also they were present on the Little Missouri and 


its tributaries. In 1919, after two years of open season on 
beavers, many of the small colonies were reduced or wiped out; 
"a few traces of their work were found along the Missouri 
River at Sanish and Bismarck, and there were said to be a few 
beavers still in Apple Creek and Burnt Creek. Near the mouth 
of the Cannonball River they were very scarce, although they 
had been fairly common up to 1916." Where cutting of trees 
seems too evident, the local residents often become exasperated 
and demand the killing-off of the animals, although the loss of 
the trees and at times that of the grain growing near at hand 
is less than the value of the beavers. 

It is perhaps this form of beaver that was formerly common 
on some of the rivers of northern and western Kansas. Hibbard 
(1933, p. 241) writes of its recent status, that although once 
common throughout the State, along the large streams, it was 
exterminated from many parts by early trappers; a few, re- 
ferred to this race, are still "found in small scattered colonies 
along the Republican, and the Kaw east into Douglas County, " 
of eastern Kansas, while a few colonies are reported along the 
Arkansas River in the western part of the State. 


Castor canadensis pacificus Rhoads, Trans. Amer. Philos. Soc., ser. 2, vol. 19, p. 422, 
Sept. 1898 (Lake Keechelus, Cascade Mountains, Kittitas County, Washington). 


Castor canadensis idoneus Jewett and Hall, Journ. Mammalogy, vol. 21, p. 87, fig. la, 
Feb. 1940 (Foley Creek, Nehalem River, Tillamook County, Oregon). 

The beaver of the northwest coast from most of Oregon 
northward into southern British Columbia was described by 
Rhoads as a distinct race in 1898, but for many years thereafter 
this name was considered a synonym of leucodonta, the Van- 
couver Island race. In 1916, however, Taylor in his review of 
the western beavers, reestablished it as a distinct subspecies. 
Its characters are slight and relate principally to the form of 
the nasal bones, which "in pacificus have their outlines some- 
what more invaded laterally by the backward-extending 
tongues of the premaxillaries, " beyond which their lateral out- 
lines again become somewhat more parallel. This posterior 


parallel part is longer than in leucodonta. Externally, the color 
of the fur is "remarkably close" to that of the Californian 
race, subauratus. 

Bailey (1936) includes in the range of this beaver in Oregon 
the coastal area west of the Cascade divide and the whole 
drainage of the lower Columbia River basin and (in eastern 
Oregon) the basin of the Snake River. Specimens from the 
headwaters of the Deschutes, La Grande, Pine Creek near Pine 
in Baker County, Oreg., and from 5 miles south of Walla Walla, 
Wash., and from Boise River west of Boise, Idaho, are, ac- 
cording to this author, "typical pacificus, " but to the southeast 
intergradation takes place with the Nevada race, bailey i; while 
in south-central Oregon, in the Klamath section, it is believed 
that the beaver is identical with the form shastensis of north- 
eastern California. 

Very recently Jewett and Hall have further divided this 
coastal beaver by describing as a separate race Castor c. 
idoneus, a smaller darker subspecies than its immediate 
neighbors, with a relatively broad skull and short nasals that 
do not extend much, if any, back of the premaxillae. Although 
definitely known only from the* type locality (Foley Creek) 
and Blaine, both in Tillamook County, Oreg., the range is 
believed to include probably the humid coastal area in Oregon, 
west of the Willamette drainage. For convenience the two 
may be treated together here. 

Quoting again from Bailey's account, "a century ago beavers 
were abundant in almost every lake and stream in Oregon, so 
abundant that the first trapping expeditions brought back rich 
returns in fur from the least-inhabited parts of the State. In 
the vicinity of extensive Indian settlements beavers were less 
numerous, or even scarce, in those days, and in a comparatively 
few years of vigorous trapping they became scarce over the 
whole State, and later while unrestricted trapping was allowed, 
they were reduced to the verge of extermination." As an 
example of their abundance, Ogden's trappers from October 
1826 till the end of the following March took over 2,200 
beavers in the State. West of the Cascades, the Willamette 
Valley was considered at one time the finest hunting ground 
for beavers in the United States west of the Rocky Mountains, 
but by 1824 they had already become scarce, and Ogden in 
1827, when he reached the Rogue River, was informed that 


Indian trappers from the Willamette had already visited the 
river and taken all the beavers. Even down to 1860, Lord, 
traveling along the headwaters of the Deschutes River, re- 
ported beavers abundant there. Beavers were gradually re- 
duced in number over all this region. Even "at their lowest 
ebb a few remained in the larger rivers, however, and during 
the past quarter century under the awakened interest in wild- 
life and the most rigid legal protection that could be given 
them in an area of extensive wilderness, they have come back 
to some of their old haunts and increased locally until they 
now may be found in many of the streams and lakes of the 
State. In fact, they are apparently more common now in the 
Grand Ronde and Walla Walla Valleys than they were a 
hundred years ago when these valleys were occupied by settle- 
ments of Indians who depended largely on the native animals 
for food and clothing." In recent years they are reported as 
more or less common in a number of counties both east and 
west of the Cascade Mountains. 

The district forester for Oregon reported in 1930 that the 
law providing an open season on beavers had proved to be a 
mistake. The eagerness of trappers to be first in the field had 
resulted in the greater part of the catch being taken before 
the fur was prime, while at the same time the number of 
beavers had been very greatly reduced. The effect of their 
dams in storing and holding back water during dry seasons was 
gone, with much resulting loss. Another observer in 1931 
states that "the removal of beaver has been a large factor in 
the shortage of water during the drought through which we 
are passing. Streams have dried up below former beaver dams 
to an alarming extent and water for stock has been reduced." 
On a stream where previous to the advent of beavers in 1920 
even a bridge was unnecessary, their dams had since increased 
so that in the 1931 drought, the driest season on record, "water 
was plentiful for a distance of a quarter of a mile below the 
beaver dams . . . and at least 20 acres of land that were 
dry in the very wet season of 1914 are kept fairly moist." 
Such observations indicate the value of beavers in water 
storage at the heads of streams during critical seasons. 

In southern British Columbia beavers, though "fast disap- 
pearing" 40 years ago, are still found in small numbers in 
remoter districts. 




Castor canadensis phaeus Heller, Univ. California Publ. Zool., vol. 5, p. 250, Feb. 18, 

1909 (Pleasant Bay, Admiralty Island, Alaska). 
FIG.: Taylor, 1916, p. 431, fig. c (tail outline). 

The beaver of Admiralty Island was at first believed to be a 
very dark race, but Taylor (1916) shows that of six specimens 
from Admiralty Island two are hardly different in color from 
two New Brunswick examples of typical canadensis, although 
others are very dark. The type is described as having the long 
hair of the upper parts almost black, with a slight chestnut 
tint at the tips; shoulders deep russet, lightening to chestnut- 
brown on the head ; underfur very dark seal brown or blackish ; 
ears black. On the under side, the longer hairs are seal brown, 
becoming slightly reddish at the base of the tail. The ratio of 
length to width of tail is even greater than in the southern 
broad-tailed beaver, frondator, but the widest part is located 
more proximally. The skull is chiefly notable for its narrower 
interorbital region, longer and narrower nasals, broader fora- 
men magnum, and longer tooth row as compared with typical 

This is an island race of beaver, hitherto known only from 
Admiralty Island, Alaska, although probably it was present 
also at one time on some of the adjacent islands. Heller re- 
garded a specimen from Prince of Wales Island as intermediate 
toward the race pacificus of the mainland to the south. He 
quotes Dixon that there were said to be no beavers on any of 
the islands between Sitka and Juneau, so that their discovery 
on Admiralty Island was of particular interest. Here they 
lived in the lakes, which with their irregular shore line and 
quiet little bays made an ideal home. They were seldom 
disturbed, although at intervals Indians from the mainland 
would come over and take a few. Their main food supply was 
found to be spruce bark, although that of willow where obtain- 
able was preferred. Four miles to the south of Wide Harbor 
it was reported that beavers had been lately exterminated. 
On the adjacent Chichagof Island, some old and dilapidated 
beaver dams were found, but the beavers seemed to have been 
cleared out. Presumably they were of this race. 

Concerning the status of the beaver on Admiralty Island in 


the years since its description by Heller in 1909, no recent 
information is at hand. 


Castor canadensis repentinus Goldman, Journ. Mammalogy, vol. 13, p. 266, Aug., 1932 
("Bright Angel Creek, Grand Canyon of the Colorado River, Arizona"). 

This is said to be a "light-colored subspecies closely allied 
to Castor canadensis frondator, of southeastern Arizona and 
northeastern Sonora, but upper parts paler, yellowish cinna- 
mon, instead of near pecan brown (Ridgway, 1912) as in the 
type; cranial characters, especially the long nasals, distinctive. 
Similar to C. c. baileyi, of Nevada, but slightly paler, upper 
parts more yellowish, less rufescent; skull decidedly broader. 
Differing from C. c. mexicanus, of New Mexico, in slightly 
yellower coloration (duller brownish in mexicanus) and in 
cranial details, especially the longer, less expanded nasals." 

Goldman lists specimens from Bright Angel Creek and Yuma, 
Ariz.; those from the latter locality with shorter nasals "prob- 
ably grade toward frondator" but in the massive jugal and 
other respects seem "nearer to repentinus" He adds: "At the 
type locality the beavers inhabit Bright Angel Creek which in 
short reaches descends the terraced north side, below the outer 
rim, of the Grand Canyon. None are known from the Colorado 
River in that vicinity, and a measure of isolation would seem 
to be due to the rapids and heavy current through rock-bound 
gorges extending for many miles, and affording very few places 
suitable for beavers to establish homes." According to Dr. 
Joseph Grinnell (1933) it was "originally and recurrently 
numerous along those parts of this river's course where willow 
and cotton wood grow abundantly . . . Since 1911, bea- 
vers have invaded Imperial Valley north of Mexican line 
following Alamo River and larger distributary canals." 


Castor canadensis sagitiatus Benson, Journ. Mammalogy, vol. 14, p. 320, Nov., 1933 
("Indianpoint Creek, 3200 feet, 16 miles northeast of Barkerville, British Colum- 

FIG.: Benson, 1938, fig. c, p. 321 (skulD. 

This local race of beaver is described as dark colored, with 
the ventral underfur dark grayish; "skull with narrow, pointed 


rostrum; angle between side of rostrum and zygomatic arch 
obtuse, so much so as closely to approach a straight line; 
nasals anteriorly narrow and pointed." The slight cranial 
peculiarities on which this form is mainly based are not very 
striking, but are shown in comparison with skulls of five other 
races in the figures accompanying the paper by Benson cited 
above; he also makes brief comparisons of its characters with 
those of the surrounding western races, and presents tables of 
cranial measurements. 

This race is "known only from the Cariboo Range, in east- 
central British Columbia." It is supposed, however, that it 
meets the range of typical C. canadensis in the mountains, 
while in south-central British Columbia it probably intergrades 
with the race pacificus, and Davis has lately identified as of 
this race a skull from northern Idaho. Presumably, too, its 
range meets that of being ae "in the northern portion of the 
Eraser River drainage system." Concerning its general status 
in the east-central part of British Columbia little information 
is at hand, but according to Benson it apparently exists in 
considerable numbers in the mountains of this region. Cowan 
(1939) in his recent account of the vertebrates of the Peace 
River district, British Columbia, writes of the beaver, which 
evidently represents this race, that it is "rare over most of the 
district but according to reports of trappers increasing on 
certain areas thanks to the combined efforts of the game 
wardens and certain of the more intelligent trappers." He 
mentions that a large female from South Pine, British Colum- 
bia, and a yearling male from Pine River are typical of cana- 
densis and exhibit no trace of intergradation with sagittatus, 
the race to the west of the mountains. 


Castor subauratus shastensis Taylor, Univ. California Publ. Zool., vol. 12, p. 433, 
Mar. 20, 1916 ("Cassel [Hat Creek], Pit River, Shasta County, California"). 

This race is based on cranial characters of relatively slight 
significance. It is closely similar to the race subauratus but 
although occurring in the same drainage basin, is found on the 
eastern or Great Basin side of the Sierra Nevada. The distinc- 
tion is based mainly on the outline of the nasal bones, which 


show an indentation in the outer margin posteriorly, due to the 
premaxillary bone; the interorbital constriction is less than in 
subauratus, while the temporal ridges tend to unite to form a 
distinct sagittal crest posteriorly and show a "higher degree of 
approximation anteriorly." The interparietal also is somewhat 

The type locality "is situated on Hat Creek, a tributary of 
the Pit River, which is in turn a tributary of the Sacramento 
River." Thus, although living in the same hydrographic basin 
as the form subauratus, its range on the eastern side of the 
Sierra Nevada is probably bounded by the Pit River Narrows, 
"a barrier not regularly crossed by beavers." It is possible, 
according to Benson, that the beavers of the Great Basin may 
be identical with this form, but the eastern range is not defi- 
nitely worked out. Grinnell (1933) gives its area of distribu- 
tion as the Pit River basin in the northeastern corner of 
California and adds that it is recorded from "Pit River, above 
Narrows, north to Willow Creek and Steele Meadow, near 
Clear Lake, and east to Lassen Creek, east of Goose Lake, and 
to North Fork of Pit River above Alturas these localities 
being in Modoc County." Its present status is not altogether 
clear, but Grinnell, Dixon, and Linsdale (1937) have sum- 
marized all the information they could obtain as to the status 
of this beaver in northeastern California and the adjacent 
region up to late 1931, as follows: In September of that year 
three colonies were known on Fletcher Creek, one consisting of 
6 animals at Willow Creek Ranch, another of 10 or 12 indi- 
viduals somewhat farther up Fletcher Creek, and the third 
and largest in the entire section comprising between 20 and 25 
beavers near the "Mulkey place." Another colony, consisting 
of about 15 beavers, lived on South Willow Creek, but a colony 
formerly living on the North Fork of Willow Creek had ap- 
parently been destroyed by trapping. About 8 miles north of 
Canby lived a colony of half a dozen beavers. The largest 
colony in the county was on Lassen Creek and comprised about 
50 animals. It extended from a point about 2 miles above 
where the creek enters Goose Lake for a distance of 2J/2 miles. 
Four or five individuals lived on Davis Creek where it runs 
into Goose Lake, and another group of the same size was 
known on the north fork of the Pit River 16 miles north of 
Alturas. They formerly (1898) occurred in the Shasta River 


and Scott River, Siskiyou County, flowing into the Klamath 
River, and there is evidence that beavers "were present in the 
Klamath River above Requa at least during the years 1915, 
1916, and 1917" and were probably of this race. From the 
figures given, the approximate population of this beaver in 
California in 1931 must have been about 120 individuals. 

The range extends into extreme south-central Oregon along 
the Klamath River drainage to Goose Lake, a region to the 
east of the Cascade Mountains, and Bailey (1936) believes 
that the beavers of the Klamath section, Lost River, Sprague 
River, and the Yamsay Mountains, are also of this form. As 
late as 1860 beavers were very abundant in this district, 
There are still a few here, where they seem to be holding their 
own or even increasing in recent years (Bailey, 1936). 


Castor subauratus Taylor, Univ. California Publ. Zool., vol. 10, p. 167, May 21, 1912 
("Grayson, Stanislaus County, San Joaquin River, California"). 

FIGS.: Taylor, 1916, fig. H, a, p. 431 (tail outline); fig. J, b, p. 450; fig. K, 6, p. 451; 
fig. L, 6, p. 452 (skull); Grinnell, Dixon, and Linsdale, 1937, vol. 2, pi. 8 (colored 
fig. of exterior) . 

Fur with a golden sheen dorsally and ventrally; size largest 
of the western beavers. Skull with the nasal bones more 
expanded and foramen magnum wider in proportion to its 
height than in any other western beaver. The average propor- 
tion of width of tail to length is about 42 percent, in the race 
leucodonta about the same, but in frondator greater, nearly half 
(49.1 percent). The type measured: Total length, 1,171 mm.; 
scaled part of tail, 320; hind foot, 196. 

This is the beaver of the "lower courses of San Joaquin and 
Sacramento Rivers and lower portions of larger tributaries of 
these main rivers" below 1,000 feet altitude in western Cali- 
fornia. Grinnell (1933) gives its former range as "from Tulare 
Lake (formerly), Kings County, and from Kings River, near 
Sanger (formerly), and at Mendota in Fresno County, north 
to Sacramento River and Butte Creek, north of Marysville 
Buttes, and at one time to McCloud and upper Sacramento 
rivers, in Shasta County." In 1912 Taylor wrote that this 
beaver was then "approaching extinction, although the en- 
forcement of the present protective law may enable it to 


regain a foothold." Twelve years later Grinnell and Storer 
(1924) reported that the protection thus accorded was effec- 
tive "and the prospects for their perpetuation are now bright." 
Indeed, by 1920 they had so increased as to become troublesome 
at Snelling, through interfering with irrigation works. In the 
river basins of western California, these beavers feed princi- 
pally on the bark of willow and cotton wood. At the present 
time they are found in some numbers in the extreme western 
end of the Yosemite section, along the lower courses of the 
Merced and Tuolumne Rivers west of the foothills. With 
proper management it is expected that this population may be 
maintained in localities where it does not interfere with human 
activities. This seems especially desirable, since the golden 
beaver is one of the better-marked races of North American 

A very thorough account of this beaver in California has 
been published by Grinnell, Dixon, and Linsdale (1937, vol. 2, 
pp. 629-721) in their recent work on the "Fur-bearing Mam- 
mals of California." They find that it is more a "bank" 
beaver than most of its relatives, adapting its habits to life 
along sluggish lowland streams. The largest specimen of 
which there is authentic record weighed no less than 82 pounds. 
They indicate that because of individual variation the shape 
of the tail is of less diagnostic value in the discrimination of 
races than had hitherto been supposed, and further it appears 
from measurements of a series of fresh specimens that in males 
its width is proportionally greater than in females. The 
period of gestation is about three months, and two to four 
young is the usual number per litter. Beavers in California 
are now "just about free from any natural enmy," with the 
reduction of bears, mountain lions, and river otters. Although 
the number of beavers in a given locality is often overestimated, 
these authors believe that in 1920 there were about 250 inhab- 
iting 15 square miles of river bottom in the vicinity of Snelling. 
Trapping on the San Joaquin River near Mendota in the same 
year indicated about 12 animals to the square mile. On this 
basis it was calculated that in 1921 there were about a thousand 
golden beavers in California. Much of the area formerly 
inhabited by beavers is now reclaimed and in use for agricul- 
tural purposes in the delta region, so that the beaver's range 
has become more and more restricted, and by 1911 very few 


were left in that district. With total legal protection given 
them in 1911, however, "they had bred up and had become 
numerous enough to endanger the levees along Cache Slough, 
and in that year seventeen beavers were trapped there, under 
special permit." Ten years later beavers had greatly increased 
in the Merced River bottom in the region about Snelling and 
became something of a nuisance to ranchers by stopping up 
irrigation canals. In 1925 legal protection of the golden 
beaver was removed, and during the two years following the 
population was severely reduced. From that time until 1931 
the "decrease has continued until it seems possible that 
eventually this species may be eliminated entirely from the 
Great Valley of California ... if protection is not 

In agricultural areas in California the damage done by 
beavers is summed up by the authors quoted under six heads: 
Burrowing into levees or dikes of reclamation work; burrowing 
into banks of irrigation canals; obstruction of drainage canals 
by dams; flooding and waterlogging of land; gnawing at head 
gates of irrigation canals; and cutting down of fence posts and 
fruit trees. They also mention the complaints of certain 
ranchers that beavers damage alfalfa fields and fruit trees in 
their use of these products as food. On the other hand they 
may, in regions not under immediate use, be of much value in 
the activities of storing water, and instances are mentioned in 
which the water held in beaver ponds has saved crops from 
drought, or their ponds may be utilized by installing pumps to 
distribute water for irrigation. "One rancher states that 
beavers have saved him several hundred dollars each season in 
this way ... In several known instances, beavers have 
lived in streams and ponds of pasture lands among cattle and 
hogs . . . with little or no competition with" the stock. 
"There appears to be no good reason why beavers, through 
their fur value, could not be made to yield a regular income on 
such ranches." The authors of the "Fur-bearing Mammals 
of California" "believe that much of the waste, dredged-over 
land along the Merced, Tuolomne, and American rivers, 
wherever these animals are not already reestablished, could be 
used to support beavers and thus be made an asset to the 
entire State." 




Castor canadensis taylori Davis, The Recent Mammals of Idaho, p. 273, Apr. 5, 1939 
("Big Wood River, near Belle vue, Blaine County, Idaho"). 

The beaver of the Snake River drainage basin in southern 
Idaho and northern Nevada has lately been distinguished on 
the ground of its "averaging darker" than the race bailey i; 
"nasals long and narrow (breadth averaging 46 percent of 
their length); anterolateral rim of orbit narrow (near 7.0 mm.); 
occiput nearly vertical." 

No details are at hand concerning the past or present status 
of this beaver. Davis based his description on 11 specimens 
from Ada, Lemhi, Blaine, -and Bannock Counties in Idaho and 
four from Goose Creek, Elko County, Nev. 



Castor canadensis texensis Bailey, North Amer. Fauna, no. 25, p. 122, Oct. 24, 1905 

("Cummings Creek, Colorado County, Texas"). 
FIGS.: Davis, 1940, pp. 85-86, figs. 1, A, and 2, D. 

This seems to be a slightly marked form, described as re- 
sembling frondator in its pallid coloring, but distinguished by 
the skull, in which the sagittal crest is short and the lateral 
ridges lyrate or spreading even in old age; supraoccipital crest 
doubly curved, nasals long, spatulate, and tapering to a 
narrow point posteriorly. These characters shown in the three 
specimens studied by the describer "are so well marked and 
uniform as to justify describing the subspecies, even on so 
scanty material." The localities given are on the Colorado 
River of east-central Texas, but "whether the beaver of other 
streams north and south" of this valley are the same "can be 
settled only by specimens." More recently the matter was 
reviewed by Dr. W. B. Davis (1940) with additional specimens, 
and he concludes that the distinctive characters of texensis 
are its "relatively darker color; posteriorly pointed nasals, 
with an expanded area near middle of rostrum; dorsal outline 
of skull nearly flat, except for the distal portion of the rostrum 
which is abruptly depressed; tail relatively long and narrow, 
ratio of width to length, 48 (39-55)" as compared with the 


race mexicanus. His specimens were from Blanco, Kimble, 
and Edwards Counties. 

To the eastward, intergradation with the race carolinensis 
may be expected, if specimens become available; while to the 
westward the beavers of the Rio Grande and Pecos Rivers 
probably represent the race mexicanus. 

Writing in 1905, Bailey states that "beaver are still found 
in many of the streams of eastern Texas, especially in the 
larger rivers, where deep water and steep banks afford protec- 
tion against relentless trapping. In 1892, at Arthur, in north- 
eastern Texas, I was informed that they were fairly common 
along the Red River and that trappers caught a few each year. 
In 1902, at Texarkana, Oberholser was told that a few were 
still found in the Red River, and in 1901, at Mobeetie, was 
informed that they were common in Sweetwater Creek, a 
branch of the North Fork of the Red River." A few were 
found at this time in scattered colonies along the Colorado 
River and in the Brazos and Trinity Rivers. In 1904, Bailey 
was told that in Polk County, in eastern Texas, beavers were 
abundant only a few years before. In northeastern Texas, on 
the upper Sabine River, trappers annually obtained them. 

A. H. Cook (1940), in a study of live Texas beavers captured 
in the summer of 1939, describes an infestation, of three out of 
40 animals examined, by the screw- worm fly (Cochliomyia 
americana), which had deposited eggs in wounds on sides, 
back, or hind limbs. He suggests that such wounds were due 
to fighting as a result of overcrowding. One of the infected 
animals seemed unable to control movements of the hind legs 
and died; while another, treated with "bone oil" (a fly repel- 
lent), recovered. 

Family CRICETIDAE: Hamsterlike Rats and Mice 


Oryzomys aniillarum Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 1, p. 177, 1898 

The Jamaican rice rat is believed now to be extinct, although 
it lived up till about the 1880's. Probably the introduced 
mongoose had a large share in its disappearance, for the last 


known examples were taken about 1877, only five years after 
the introduction of that carnivore into the island. These 
specimens are in the United States National Museum. An 
earlier example preserved in the British Museum was made, in 
1898, the type of the present species by Thomas, who describes 
its color briefly as dull rufous above, rather richer on the rump 
and grayer on the head, with a slight lining of black on the 
back; under side dull yellowish, not sharply defined, the hairs 
slaty gray basally; tail about as long as head and body, nearly 
naked, hands and feet dull whitish. Total length about 10 
inches (252 mm.) of which the tail is slightly more than half: 
hind foot, 29.2; length of skull, 30.5 (Goldman, 1918). 

This rice rat, as Thomas pointed out in his original descrip- 
tion, is very similar to the mainland 0. couesi or some of its 
Central American races. Indeed, it seems doubtful if it 
should not be regarded as identical with some one of them, for, 
as the only cricetine rodent known from the Greater Antillean 
islands, it is obviously a recent intrusion in a rodent fauna 
otherwise of a hystricoid type. In his review of the North 
American rice rats, Goldman (1918) compares the skull with 
that of typical couesi of Yucatan and Honduras but finds 
relatively few points of difference, namely, that the nasals 
reach slightly farther back beyond the premaxillaries, the 
maxillary arm of the zygoma is heavier, and the anterior 
palatine foramina are shorter than usual in couesi. Since, 
however, only the two specimens in the U. S. National Museum 
(from Metcalfe Parish and Spanish Town, respectively) were 
available for comparison, it seems likely that these differences 
may not be very significant. Without further evidence to the 
contrary the name may for the present be retained, but at the 
same time the possibility is recognized that the animal may 
have been a rather recent introduction through human agency. 
That it nevertheless thrived and multiplied is evident from the 
fact that Dr. H. E. Anthony (1920a) in exploring caves along 
the seacoast of Jamaica found none of the subfossil types of 
hystricoid rodents in them, though they often did yield "re- 
mains of the extinct rice rat, thus showing that at no very 
remote period this small rodent had a widespread distribution 
and was so common that it formed an important part of the 
diet of the barn owl," by which evidently they were brought 
in to the caves. "The reason for its disappearance obviously 


lies in the advent of the ubiquitous Norway rat and the blood- 
thirsty mongoose." 

Doubtless this is the field mouse described by P. H. Gosse 
in his "Naturalist's Sojourn in Jamaica" (Gosse and Hill, 1851) 
as of a "beautiful reddish colour, with a milk-white belly." 
"It takes up its habitation chiefly about the hollow roots of 
large trees, and the rocky acclivities of gullies and river banks. 
It is far from numerous." The type specimen in the British 
Museum was collected by Gosse in 1845. 


Oryzomys victus Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 1, p. 178, Feb., 1898 ("St. 
Vincent, Lesser Antilles"). 

Thomas, in describing this species, wrote: "It is difficult to 
say to what species this mouse is most nearly allied." At that 
time he compared it with the South American Oryzomys longi- 
caudatus, giving it the following characters: Size and propor- 
tions about as in 0. longicaudatus; color dark rufous, but this 
probably affected by the alcohol in which the type is preserved ; 
below, buffy white, the hairs with slate-colored bases. Length 
of head and body, 96 mm. ; tail, 121 ; hind foot without claw, 25 ; 
basal length of skull, 23.8. 

The type and only known specimen was presented to the 
British Museum by F. DuCane Godman in 1897. Since that 
time no further attempt to determine the nearer affinities of 
the specimen has been made, nor have any additional examples 
been taken. It therefore still remains problematical whether 
it is a native or an introduced animal, what its continental 
relatives are, and whether it still exists on the island. Since 
no representative of the genus is known from any other of the 
Lesser Antilles, it may prove to have been introduced in St. 
Vincent through local trade from some adjacent part of South 
America; on the other hand, the type specimen collected by 
H. H. Smith was marked by him "forest rat," implying its 
existence in wooded areas on the island. Goldman (1918, p. 87) 
in speaking of it, says: "This rice rat seems likely to be en- 
dangered by the presence of the mongoose, if it has not already 
been exterminated since the introduction of that indiscrimi- 
nately destructive animal." 



Miis desmarestii J. B. Fischer, Synopsis Mammalium, p. 316, 1829 ("In Insula Mar- 

SYNONYMS: Mus pilorides Desmarest, Diet. Sci. Nat., vol. 44, p. 483, 1826 (not of 

Pallas, 1778); Hesperomys (Megalomys) pilorides Trouessart, Le Naturaliste, no. 

45, p. 5, 1881; Oryzomys piloris Forsyth Major, Ann. Mag. Nat. Hist., ser. 7, vol. 

7, p. 205, 1901 (based on Castor Piloris of Zimmermann, not a technical name). 
FIGS.: Geoffrey and Cuvier, 1830, vol. 4, pi. 258 ("Le Pilori"); Trouessart, 1885. 


Oryzomys luciae Forsyth Major, Ann. Mag. Nat. Hist., ser. 7, vol. 7, p. 206, Feb., 1901 
("Santa Lucia," Lesser Antilles). 


Megalomys audreyae Hopwood, Ann. Mag. Nat. Hist., ser. 9, vol. 17, pp. 328-330, Mar., 

1926 (Barbuda, Lesser Antilles). 
SYNONYM: Megalomys majori Trouessart, Cat. Mamm. Viv. Foss., ed. 2, pt. 2, p. 415, 

1904 (nomen nudum). 
FIG.: Hopwood, 1926, pi. 12. 

The so-called "Musk-rat of the Antilles" may formerly 
have occurred on most of the islands of the Lesser Antilles, 
but it has by now probably been quite exterminated. Although 
its range may once have been more extensive, it is certainly 
known only from the islands of Martinique (the typical form, 
M. desmarestii), St. Lucia (the smaller form, M. luciae), and 
from the little island of Barbuda, where a sub fossil jaw was 
found. These were rather large rodents, with a head and body 
length up to 360 mm. (14.5 inches) and tail only a little shorter, 
and belong to the group of sigmodont species, abundantly 
represented in South and Central America. 

Th Martinique form was the one first named and is the 
larger of the two known from complete specimens. Trouessart 
describes a specimen in the Paris Museum as having the head 
and body 360 mm. long, the tail 330 mm. The head was 
rather short in appearance and rounded, the muzzle more 
obtuse than in a house rat, and the upper lip with a deep 
vertical furrow. The ears were well developed and nearly 


naked. The fur was long and harsh but not spiny and of a 
dark reddish brown, both above and below. The tail was 
black at the base, with a white intermediate area and a black 
tip. Desmarest, however, speaks of the color as lustrous black 
except for the chin, throat, and base of the tail, which are 
white. The skull as figured by Trouessart (1885) is provided 
with a conspicuous bony ridge on each side from the upper 
anterior edge of the orbit back over the brain case to the lateral 
corner of the cranium behind. The skull is rather squarely 
truncate posteriorly; 70 mm. long. 

The St. Lucia musk-rat was slightly smaller, with the greatest 
length of the skull fron tip of nasals to foramen magnum about 
49 mm. The color was apparently much the same as in the St. 
Vincent form, but the belly was nearly wholly brown instead of 
largely white. 

That the musk-rat occurred on Barbuda to the northward is 
interesting, for in this extension to the northern Lesser Antilles, 
there is a certain parallelism in distribution with the agoutis. 
The Barbuda species, M . audreyae, is based on a lower jaw and 
an upper incisor tooth found in a cave breccia, which, though 
presumed to be of Pleistocene age, may not be of any great 
antiquity. The jaw seems slightly smaller still than that of 
the St. Lucia musk-rat, and in certain slight details of the last 
two molars that are preserved shows a less development of the 
small accessory cusps, or paraconids. 

The Martinique musk-rat was first mentioned in literature 
by Du Tertre in 1654, in his "Histoire Generale des Isles de S. 
Christophe, de la Guadeloupe, de la Martinique, et Autres 
dans I'Amerique." He did not know of it from any of the 
French islands except Martinique, where, he relates, it was 
commonly eaten by the people, who, after first singeing off the 
hair, exposed the body overnight to the air, and then boiled it, 
throwing off the first water in order to get rid of the strong 
musky odor. It was said to live in burrows in the ground and 
against it the colonists waged war on account of its destructive 
habits in their plantations. In addition to human enemies, 
the large serpents of Martinique also attacked it. Du Tertre 
mentions killing a large snake in the stomach of which was one 
of these rats "almost as big as a cat." Of preserved specimens, 
there appear to be scarcely above half a dozen. The specimen 
figured by Geoffrey and Cuvier is a mounted one in the Paris 


Museum, one of two or three collected by M. Plee, who lived 
at Martinique from 1821-26. Another from the same collector 
is said to be in the Leiden Museum. The Paris Museum has 
another mounted one brought back by a M. Le Prieur (Pdate) 
with no other locality but "des Antilles." The teeth of this 
last specimen, but little worn, and the skull and worn dentition 
of another are figured by Trouessart (1885). Exactly when the 
Martinique musk-rat became extinct is difficult to say, but 
that it may have existed up to the end of the nineteenth 
century is indicated by the following statement communicated 
to me by the late Dr. G. Kinsley Noble, who in 1914 visited 
Guadeloupe and was told by a Mr. Delphin Duchamp, a 
former resident of Martinique, that "about five years before 
the eruption of Mount Pelee [1902] there used to exist in great 
numbers among the cocoanut plantations along the Riviere 
Blanche, close to St. Pierre, a species of rat which was black 
as coal on the back and white as milk below. When adult 
this creature was some 40 cm. long without the tail. I killed 
many of them, for their flesh is very delicate. The negroes 
call this rat the pilorie. It lives almost entirely in [Pamong] 
the cocoanut trees but will take to water when driven from 
shelter. I never heard of it occurring in any other part of the 
island except on these plantations, and since this whole region 
was destroyed by the great eruption of 1902, it is very probable 
that the rat is now extinct." Inquiries among the hunters on 
Guadeloupe showed that it was unknown to any of them as 
occurring there, though they knew of its former presence on 
Martinique. No doubt the devastating eruption of Pelee 
accompanied by clouds of poisonous gas completed its destruc- 

Of the St. Lucia musk-rat, Trouessart (1885) records a speci- 
men preserved in the Paris Museum and calls attention to 
its smaller size and less amount of white below as compared 
with the "pilorie" of Martinique. This individual was 
brought back by Bonnecour some time previous to 1881. The 
only other specimen extant seems to be the one now in the 
British Museum, which served as the type of Forsyth Major's 
"Oryzomys" luciae. This in turn was doubtless the one men- 
tioned as received from St. Lucia by the Zoological Society 
of London in 1849 (see Proc. Zool. Soc. London, 1849, p. 105). 
It was sent by Lieutenant Tyler and may have lived in London 


for three years, since the accession date as shown by its number 
in the register of the British Museum was 1852. Possibly this 
species was exterminated prior to the final destruction of the 
Martinique species. 

There is nothing to indicate when the Barbuda musk-rat 
was living, but probably it disappeared soon after the occupa- 
tion of the island by Europeans and consequent destruction of 
cover. Even in Du Tertre's time the musk-rat was not known 
from any of the other islands, for De Rochefort's assertion 
that it was one of the five species of mammals native to Tobago 
may be received with caution. 



Arvicola breweri Baird, Mammals North Amer., Pacific Railway Repts., vol. 8, p. 525, 

1857 (Muskeget Island, Massachusetts). 
FIGS.: Miller, 1896, pi. 1, figs. 1, la (skull). 

The Brewer's beach mouse of Muskeget Island, lying be- 
tween Marthas Vineyard and Nantucket Island off southeast- 
ern Massachusetts, is of special interest on account of the 
characters it has developed of large size and pallid coloration, 
differing in the latter respect particularly from all the other 
island meadow mice of our coast. It is generally supposed that 
these traits have resulted from isolation and inbreeding on the 
confines of this small sandy island. On the other hand, it may 
be that the beach mouse is a relict species that was formerly 
an inhabitant of the now- vanished sandplain, which extended 
coastwise from New Jersey to Newfoundland, and forms a 
case parallel with that of the Ipswich sparrow of Sable Island, 
Nova Scotia, a pallid form of that sandy spot, not found 
elsewhere at the present day. 

In addition to its somewhat larger size when adult, as com- 
pared with the common Microtus pennsylvanicus of the neigh- 
boring mainland and the island of Nantucket, the beach mouse 
is strikingly pale, a "light gray throughout, purest (almost 
white) on the belly and tinged with wood-brown on the sides 
and back, the latter somewhat darkened by a sprinkling of 
longer blackish hairs . . . Tail indistinctly bicolor, brown- 
ish dorsally, whitish ventrally" (Miller, 1896). A large per- 
centage of these mice have a white mark in the forehead. The 


skull in adults is larger than that of typical M. pennsylvanicus, 
with somewhat longer and narrower brain case, more abruptly 
flaring zygomatic arches, and the outline of the interparietal 
is longer and narrower. Average adult length, about 200 mm. 
(8 inches). 

Muskeget is distant only 6 miles from the western end of 
Nantucket. The intervening waters are shallow and on the 
south side it is partly protected by some small islands and long 
beaches. In his account of this mouse and its island, Miller 
(1896) writes that in 1887 the United States Coast Survey 
found the southern point extending out much farther to the 
southwest than in later years, during which the sea washed 
away all but the tip of the point, which remained as South 
Point Island. He found about a dozen species of birds breeding 
there and the white-footed mouse occurred in the clumps of 
beach plum, or about the numerous small houses along the 
south shore of the island. The beach mice seem to have had 
various ups and downs. In 1869 they were recorded as "exces- 
sively abundant"; in 1890, however, only a few were found by 
two naturalists visiting the island, while in 1891, William 
Brewster was unable to find any trace of them. Miller records 
that the warden, Mr. Sandsbury, who was stationed there in 
summer to protect the tern colony and who had been familiar 
with the mice for many years, believes that they had actually 
been exterminated by cats that ran wild on the island after the 
burning of the Life Saving Station a few years before. Since 
the mice can burrow but little in the loose sand, they may 
have been the more easily exterminated in this way. When 
Miller visited the locality in 1892, he found, however, that 
there were large and thriving colonies on South Point Island 
and the adjacent Adams Island, which had been cut off from 
Muskeget previous to the burning of the Life Saving Station. 
This accident had therefore saved the mice from total extinc- 
tion. Of the further history of this interesting population, 
Miller (1896) writes: "The 28th of December, 1892, I spent 
on Muskeget and the neighboring islands. The mice were as 
numerous as before on South Point Island, but the colony on 
Adams Island had greatly diminished. In June, 1893, I again 
visited Muskeget, this time in company with Mr. Outram 
Bangs and Mr. Chas. F. Batchelder. We found that the 
Microtus colony on Adams Island had entirely disappeared. 


On South Point Island, however, the mice were so abundant 
that in less than two hours we caught forty -three. After 
selecting as many as we wanted for specimens, we turned out 
twenty -six on Muskeget. Although I have not been at Muske- 
get since 1893, I have heard that the mice on the main island 
are increasing rapidly. Mr. Sandsbury has written several 
times to this effect, and Mr. W. K. Fisher collected two dozen 
specimens for the U. S. Department of Agriculture during 
July, 1895. Mr. Fisher found two colonies both in clumps 
of beach plum bushes one at the east end of the island, near 
the place where we liberated the mice in 1893, the other about a 
mile farther west. He also found that the species was extinct 
on South Point Island." The Muskeget colonies have ap- 
parently continued to flourish since their reintroduction, for as 
lately as the summer of 1937 Donald R. Griffin obtained a few 
specimens there. The island is now a reservation for nesting 
terns and other birds and is provided with warden service 
through part of the year. To this end, also, the chance intro- 
duction of cats is especially guarded against, so that for the 
present the outlook is bright that the mice will continue to 
survive. But it is obvious that any important change in the 
living conditions, the introduction of parasites or disease, or 
the lessening of human guardianship may again imperil them. 
Concerning the special habits of these mice, Miller (1896) 
writes further that they have been "modified to meet the needs 
of a life among coarse, loose sand, in which no extensive bur- 
rows can be made except during the brief and irregular periods 
in winter, when the surface is frozen. Throughout the greater 
part of the year the animals are exposed to the full force of the 
elements, their only natural protection being that furnished 
by the scant beach vegetation or by fragments of drift wood 
and wreckage. Where the mice are abundant, a labyrinth of 
well beaten paths crosses the sand in every direction, and 
when one of the animals is chased, he follows these runways 
aimlessly and helplessly, until exhausted or until he finds some 
place of refuge, perhaps in a tuft of Ammophila, or beneath the 
stalks of a beach golden rod. Occasionally a mouse tries to 
escape by entering a burrow among the roots of the beach 
grass, but these tunnels always afford a very insecure protec- 
tion as they are close to the surface and seldom extend more 
than a few inches, or at most a few feet. The mice apparently 


make these short tunnels with the sole object of reaching the 
soft parts of the beach grass stems. They are in no way con- 
nected with the nesting burrows. Another means of shelter is, 
so far as I know, peculiar to the Muskeget mice. They con- 
struct frail nests or forms which may be seen scattered about 
everywhere. The forms are usually open at the top and made 
of the finer shreds torn off from the beach grass. Each is 
large enough to contain one animal only. The walls are much 
thinner than those of the breeding nests, being a mere film of 
grass fiber, through which the occupant can be distinctly seen. 
The forms may be built on the bare sand or under some shelter 
indifferently. Often one partly arches a beaten path. 

"The breeding nests, which resemble those of Microtus 
pennsylvanicus, are sometimes made under the protecting 
stalks of the luxuriant Solidago sempervirens or under cover 
of a fragment of wreckage. When no such convenient shelter 
can be found the mice construct short nesting burrows. These 
are from one to two feet in length and penetrate the sand at an 
angle of about 45. In this way the tunnel descends quickly 
through the dry crumbling sand at the surface to the more 
compact layers beneath, where the walls are less likely to cave 
in and smother the helpless young. At the end of the tunnel is 
a hollow, completely filled by the bulky nest . . . Four or 
five young is the usual number in a litter . . . The food 
of Microtus breweri consists chiefly of the tender bases of the 
beach grass stalks ... As the supply of Ammophila on 
Muskeget is practically unlimited the beach mice never suffer 
from hunger, but the struggle for existence must be fierce, 
notwithstanding, for few small mammals live in as exposed 



Microtus scirpensis Bailey, North Amer. Fauna, no. 17, p. 38, 1900 ("Amargosa 
River, near Nevada line, Inyo County, California"). 

This is a desert-living race of the widespread M. calif ornicus, 
from which it differs in being slightly brighter in color, a pallid 
neutral gray above, but not so black as in the neighboring race 
mariposa of the western foothill belt of the Sierra Nevada; 
belly smoky gray; tail indistinctly bicolor, brown above, gray- 


ish below; feet brownish gray, not dusky. Six adults averaged 
in total length, 203 mm., about 8 inches; tail, 65; hind foot, 
25.1. Skull, basal length, 31 mm.; zygomatic breadth, 19; 
upper molar series, 8.7. 

When first described by Bailey this race was known only 
from the type locality, where it lived in wet ground under tall 
tules (Scirpus olneyi), in a little marsh surrounding a warm 
spring. Here the mice had made extensive runways through 
the mud and water. Kellogg (1918), in reviewing the meadow 
mice of the calif ornicus group, points out that it is an outlying, 
isolated form, a relict of what was doubtless in former times 
a more or less continuously distributed species. "It is a re- 
markable race because of its occurrence away from the main 
mountain axes and in an area of extremely high summer 
temperature" in southeastern California. Since assiduous 
trapping at the original locality in May, 1917, failed to yield a 
single specimen, Kellogg believed that it had "been extermi- 
nated within recent years, as the type locality, a small tule 
marsh near Shoshone, Inyo County, has been burnt over for 
several consecutive years and is now being used as a hog 
pasture." Nevertheless, it seems that a remnant persists, for 
Dr. Joseph Grinnell writes in response to Dr. Francis Harper's 
inquiries, in 1937, that it has been "rediscovered within the 
last few years, not to be sure, at the type locality, but within a 
few miles' radius, as a result of intensive search. This is to be 
credited to Miss Annie M. Alexander, who found them in 
exceedingly small numbers at two desert seepages, where, 
however, the favoring conditions might at any time be wiped 
out." On account of its living in such very localized swamps, 
where a series of dry years or drainage and fire might at any 
time destroy its habitat, the race is in a precarious state much 
like that of an animal living on a few small islands where the 
environment may become unsuitable through slight changes. 



Microtus nesophilus Bailey, Science, new ser., vol. 8, p. 782, Dec. 2, 1898 (Great Gull 

Island, at entrance to Long Island Sound, New York). 
SYNONYM: Microtus insularis Bailey, Proc. Biol. Soc. Washington, vol. 12, p. 86, Apr. 

30, 1898 [not Lemmus insularis Nilsson = Microtus agrestis (Linnaeus)]. 


This meadow mouse is an island race of the common Micro- 
tus pennsylvanicus of eastern United States, from which it 
differs only in minor characters. At the present time it is 
believed to be extinct. 

Bailey describes this as darker than typical M . pennsylvani- 
cus, the upper parts dark yellowish bister, heavily mixed with 
black hairs, darkest on nose and face; belly dusky, washed 
with cinnamon; feet blackish; tail blackish above, dark brown 
below. The skull is characterized by its shorter, wider brain 
case and its wider and more abruptly spreading zygomatic 
arches. In the last upper molar the anterior inner and outer 
triangles are confluent and nearly opposite instead of being 
separated by enamel walls. Basal length of skull, 26 mm.; 
zygomatic breadth, 16.2. 

Beyond the original and supplementary descriptions of this 
meadow mouse by Bailey (1900, p. 26), nothing seems to be 
known of it. That author remarks: "Microtus nesophilus 
needs no comparison with breweri or terraenovae, the other two 
insular forms from the Atlantic coast. In general appearance 
it more nearly resembles pennsylvanicus, but in cranial char- 
acters it is as distinctly different as either of the other island 
species. During the month of August, 1898, Mr. A. H. Howell 
visited Great Gull Island for the purpose of getting specimens 
of Microtus, but he found their old haunts covered by the 
earth moved in grading the island for fortifications, while no 
trace of the animals remained. He thinks they are completely 
exterminated." Fifteen specimens are listed in the collection 
of the United States Fish and Wildlife Service, but apart from 
these and Bailey's account there seems to be no further record 
of the race. It is a good example of the precariousness of the 
status of an island form, which with some unusual modification 
of its habitat is exposed to new dangers, 

Since Bailey's account of this race (for such it may best be 
considered), the meadow-mouse of Block Island, R. I., has 
been described as an insular race by Bangs (1908, p. 20) under 
the name Microtus provectus. The differential characters seem 
slight but perhaps sufficient to warrant this action. There is 
nothing to indicate that it is in any present danger, though 
conceivably changing conditions on Block Island might alter 
its habitat sufficiently to imperil it. Of the supposed race M. 
pennsylvanicus shattucki Howe, of Tumbledown Dick Island, 


Penobscot Bay, Maine, it has been shown by Wyman (1922, p. 
163) that it is identical with the mainland species. Meadow 
mice are excellent swimmers and may be expected to occur on 
most of the small coastwise islands where conditions are 
suitable and where the distances between such islands and the 
nearest land are not too great to be crossed at intervals by them. 

Family ECHIMYIDAE: Spiny Rats and Their Relatives 



Brotomys voratus Miller, Smithsonian Misc. Coll., vol. 66, no. 12, p. 7, 1916 ("Kitchen 

midden at San Pedro de Macoris, Santo Domingo"). 
FIGS. : Miller, 1916b, pi., fig. 1 (four views of type rostrum) ; 1929c, pi. 2, fig. 1 (rostrum) ; 

1930, pi. 1, figs. 2, 2a, 2b; pi. 2, fig. 5 (femur). 


Brotomys (?) contractus Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 13, Mar. 

30, 1929 ("Small cave near St. Michel, Haiti"). 
FIG.: Miller, 1929a, pi. 2, fig. 2 (palate). 

These two species of Hispaniola, the second with peculiarly 
narrowed palate, may, as Miller remarks, represent even dis- 
tinct genera, but with the fragmentary material at present 
known, this is still uncertain, though they evidently are related 
rather closely. 

Brotomys represents a spiny rat, about the size of the South 
American Proechimys canicollis, remarkable for its "robust 
skull with weak teeth." The antorbital opening is very large 
and lacks a secondary canal at the lower corner for the passage 
of the facial branch of the fifth nerve, a character readily dis- 
tinguishing it from Boromys. The upper cheek teeth are four 
on each side, with three short roots each. There is a deep 
enamel infolding from about the middle of each side of each 
cheek tooth, meeting or slightly overlapping at the center. 
Probably a shallower infold is present in the crown of a fresh 
tooth, which soon wears down to leave a small island of enamel 
in at least the posterior half of each upper molar. Length of 
four upper alveoli, 10.8 mm. The palate is slightly emarginate. 
In the toothless palate representing B. contractus the very 
narrow space between the molar rows is a most striking 


This spiny-rat genus is known only from Hispaniola, where 
Miller (1916b, 1929a, 1929c, 1930) has recorded it as relatively 
common not only in the cave deposits, doubtless made in large 
part by owls, but also in kitchen-middens of the aborigines. 
Thus he secured two imperfect skulls and more than 50 mandi- 
bles from caves near St. Michel, in central Haiti, while in the 
region about Samana Bay in northeastern Dominican Republic 
it occurred in every one of the various Indian sites investigated. 
He writes (1929c) : "The frequency with which the bones of this 
animal occur in the Indian deposits indicates that Brotomys 
must have been abundant and generally distributed in pre- 
Columbian days. It was probably much like the living South 
American spiny-rats in size and general form, but with heavier, 
less elongated head. I have little doubt that this animal was 
the Mohuy described by Oviedo as ... somewhat 
smaller than the hutia [Plagiodontia], its color is paler and 
likewise gray. This was the food most valued and esteemed 
by the caciques and chief of this island; and the character of 
the animal was much like the hutia except that the hair was 
denser and coarser (or more stiff), and very pointed and 
standing erect or straight above." 

Nothing is known of B. contractus except the type palate. 
No doubt the two were contemporaneous and were early ex- 
terminated after the discovery of the island. It seems unlikely, 
however, that hunting for food was the main cause of extinc- 
tion. Possibly the introduced rats became a menacing factor, 
or the actual extinction may have been the result of several 



Boromys o/ella Miller, Smithsonian Misc. Coll., vol. 66, no. 12, p. 8, 1916 ("Village 

site at Maisi, Baracoa, Cuba"). 
FIG.: Allen, G. M., 1918, pi. 1, fig. 6 (mandible). 



Boromys torrei G. M. Allen, Bull. Mus. Comp. Zool., vol. 61, no. 1, p. 6, Jan., 1917 

("Cavern in the Sierra of Hato-Nuevo, Province of Matanzas, Cuba"). 
FIGS.: Allen, G. M., 1917, pi., figs. 10-13; 1918, pi. 1, figs. 11-13 (skull and teeth). 


Remains of these two supposed spiny rats differ chiefly in 
size, and both occur together in cave deposits; hence they may 
be considered under a common grouping. The genus is related 
to Brotomys of Hispaniola, but the skulls differ in that the 
latter shows no external swelling of the bone over the posterior 
termination of the root of the upper incisor, nor is there a 
definitely marked channel for the passage of the fifth nerve on 
the floor of the antorbital foramen; in Boromys, on the other 
hand, the posterior termination of the incisor root is marked 
by an obvious external swelling, and the floor of the antorbital 
canal is provided with a groove for the nerve. 

So far as known from parts of the skeleton, the two species 
were alike in structure. There were four cheek teeth in each of 
the tooth rows. Each tooth had a deep inner and outer re- 
entrant of enamel extending to the center of the tooth, the 
inner slightly in advance of the outer in each jaw. A smaller 
reentrant of less vertical depth was present in both the anterior 
and the posterior halves of the premolar and in the posterior 
half only of the three molars, penetrating from the outer side. 
With wear these smaller inlets are soon cut off and appear as 
little circular islands of enamel in the center of their respective 
sections and with still further wearing down, disappear. The 
enamel wall at this stage has nearly the outline of a figure 8. 
The alveoli of the molars are squarish in outline with shallow 
cavities for short roots. The incisors were orange-yellow in 
color. In the larger species, B. offella, the alveoli of the upper 
teeth have a combined length of 11.5 mm., against 7.6 mm. in 
the smaller animal, B. torrei; while the crown area of a single 
molar in the former is about three times that in the latter. 

In former times both these species seem to have been well 
distributed not only in Cuba, but in the Isle of Pines as well. 
The larger animal was first discovered in excavations of an old 
village site at Maisi, Baracoa; hence it was probably used as a 
food animal by the Indians. Shortly after this discovery, a 
number of maxillae and mandibles were found in a cave at 
Limones, Cuba, and in similar situations in the Sierra de 
Casas, Isle of Pines, by Dr. Thomas Barbour and his associates. 
These remains may have been brought in by the now extinct 
giant barn owl. Other but "scanty material" representing the 
same animal was found at about the same time by Dr. H. E. 
Anthony (1919) in the course of excavations in caves at 
Daiquiri, in eastern Cuba, brought in probably by owls. 


Of the lesser Cuban spiny rat, remains seem more numerous 
in the caves, perhaps because the animals were more easily 
taken by the owls. The species was first found in a breccia 
solidly cemented by infiltrated lime deposits in a cave in the 
Sierra de Hato Nuevo, Matanzas Province. Anthony found 
it abundantly represented in the cave deposits at Daiquiri, 
while on the Isle of Pines additional remains were unearthed 
in caves in the Sierra de Casas by Messrs. Barbour, Brooks, 
and Warner and at another cave deposit in the same island by 
the Messrs. Link (Peterson, 1917). Nothing further is known 
of either species, which must both have become extinct within 
the past century from causes not altogether clear. Neither 
Gundlach nor Poey seems to have known of the animals, yet 
some of the bones recovered, such as the nearly perfect skull 
figured in my paper of 1918, are so fresh that they can be of no 
very great age. Anthony (1919) writes concerning the bones 
he collected in the Cueva de los Indios, at Daiquiri: "Some 
idea of the abundance of the members of this genus is shown 
by the fact that I have cleaned and examined over five hundred 
mandibular rami, while nearly as many fragments have been 
ignored as too badly broken up. The small torrei seems to have 
been the dominant form, as only about five percent of this 
series represents the larger" one. That the smaller species was 
ignored as a food animal by the natives may be indicated by 
the absence of its remains in kitchen middens. 

Although in the general structure of the teeth the genus is 
strongly reminiscent of the continental spiny rats of South 
America, it has relatively "broader nasals. Miller points out a 
close resemblance of the enamel pattern to that of the extinct 
Stichomys of the Santa Cruz formation of the South American 
Miocene. The fact that the related Brotomys of Hispaniola is 
nevertheless unknown elsewhere, and the lack of similar forms 
on the mainland, point to long isolation. Miller regards the 
two genera as relatives of the Puerto Rican Heteropsomys, 
which, if this surmise proves to be substantiated, would imply 
some common origin of the three types now regarded as 
generically distinct. Anthony, however, believes that Hetero- 
psomys is a near ally of the agoutis. 

True (1885) records a large spiny rat taken on Martinique 
by Ober in 1878. It was sent to the United States National 
Museum and there identified as "Loncheres" (= Echimys) 


armatus of South America, whence it may have been intro- 
duced. There seems to be no recent evidence of its continued 
presence on the island, although Austin H. Clark in 1904 was 
told by the natives that it still occurred. 


Caprornys melanura Poey, Monatsber. preuss. Akad. Wiss. Berlin, 1864, p. 384 

(Manzanillo, southeastern Cuba). 
FIGS.: Dobson, 1884, pis. 18-21 (colored figure of exterior; anatomy); Bucher, 1937, 

pis. 10-12 (photographs of captives). 

So far as known, the black-tailed hutia is confined to ex- 
treme eastern Cuba in the wooded mountains. Its circum- 
scribed range places it in a somewhat dangerous position, and 
although it may continue to exist there for years to come, 
changes incident to human influence, the clearing of forest, 
the introduction of the mongoose, and hunting for food are 
likely to reduce its numbers more and more. 

This is a slightly smaller animal than the common hutia of 
Cuba, Capromys pilorides, less clumsily built, and of more 
arboreal habits. With a body slightly larger than that of a 
muskrat, and with a bushy tail about as long as the body 
without the head, the general color is a mixed blackish or 
brownish and buffy, due to the presence of scattered all-black 
hairs and more abundant hairs having smoky-brown bases 
and pale-buff y or ochraceous tips. The forehead and cheeks 
are nearly clear dark brown or clearer brown, as are the hands 
and the central area of the hind feet. The under side is similar 
but paler, mixed grayish and brown, clearer brown on the 
throat. There is a varying amount of clear white, which may 
include only the axillae and inside of the thighs, or, as in one 
specimen examined, the white may include the entire upper 
throat, the under side of the arms, and continue over the 
entire mid ventral surface to the inner side of the thighs, the 
ankles, and the outer margins of the hind feet. The tail is 
thickly haired, tapering, with abundant somewhat-stiff black- 
ish to maroon hairs from one-half to three-quarters of an inch 
long. Total length about 26 inches, of which the tail consti- 
tutes about 11 or 12 inches. The claws are slender and sharply 
curved for climbing, and the soles are granular for better 


Although somewhat resembling Capromys prehensilis, the 
longer-tailed and prehensile-tailed species of western Cuba, 
this seems to be a quite distinct species. Although its tail is 
prehensile, the hair is not worn away through use of the tail in 
holding as it is in the former. Moreover, as is well known to 
those accustomed to hunt it, the tail readily breaks away near 
the base if strain is placed upon it. For this reason the hunter 
does not attempt to drag the animal out from a hole by the 
tail. As with spiny rats, the point of weakness is close to the 
base of the tail and is marked externally by a sudden change in 
the character of the hair from that of the body to the more 
shaggy type of the rest of that member. It comes away easily 
without bleeding, leaving a rosette of muscle fibers. The de- 
tails of the anatomy and external appearance are illustrated in 
the paper by Dobson (1884). 

First clearly described by Poey in 1864, the species continues 
to be little known and rare in collections. Those in the Museum 
of Comparative Zoology were secured in the foothills of the 
Sierra Maestra of extreme southeastern Cuba in 1913, and again 
others were obtained in the mountains north of Imias in 1936. 
This species replaces C. prehensilis in the eastern tip of Cuba 
and is closely related to it. The only good account of its habits 
is the recent one of Bucher (1937), who kept several of this and 
the other larger Cuban species in captivity and found them 
interesting pets. Of C. melanurus he writes that they are 
active and feed late in the evening and at night. In a wild 
state they apparently live in pairs together, and the females 
produce one or two young at a birth. The males fight one 
another for possession of a female but otherwise seem good- 
natured. Unlike C. pilorides, they do not enjoy being petted 
and can not be trusted not to snap at one's finger at times. 
They chatter their teeth when approached and at mating time 
use a short, low birdlike whistle. In captivity the gestation 
period seems to be about 17 or 18 weeks. In actions they are 
said by Bucher to be very squirrellike, running along branches 
or leaping from tree to tree, using the tail as a balance or in a 
prehensile manner. Bucher fed his C. melanurus at first on 
fruit of "yagruma" (Cecropia), and later they became adapted 
to a diet of sweetpotato vine, prunings of guava, any citrus 
fruit and grapevine leaves, buds, and bark. Unlike C. pilorides, 
they will not eat bread or cracker. Probably their arboreal 


life and nocturnal habits will help to keep them from exter- 
mination for a considerable time to come. 

So far as known the genus Capromys is confined to Cuba and 
the Isle of Pines. No fossil species are to be found on any of 
the other West Indian Islands, which seems a significant fact 
in the distribution. The two other large species, C. prehensilis 
and C. pilorides, are represented by only slightly differing 
races on the Isle of Pines, relictus and gundlachi, respectively. 
Bucher (1937) has brought out some interesting details in 
which their habits differ. Since, however, neither species 
seems in immediate danger of extermination, they may be 
omitted here. 



Capromys nana G. M. Allen, Proc. New England Zool. Club, vol. 6, p. 54, 1917 ("Cave 

deposit in the Sierra de Hato Nuevo, Province of Matanzas, Cuba"). 
FIGS. : Allen, G. M., 1918, pi., figs. 1-5; Morrison-Scott, 1939, pis. 5-7 (skull and teeth). 

This small Capromys was first made known from jaws found 
in recent cave deposits at two localities in Cuba. Subsequently 
it was found living in the great Zapata Swamp, on the southern 
coast of middle-western Cuba, south of the original locality. 

About the size of a Norway rat, this is typical Capromys, 
with a rather coarse pelage of all-black hairs mixed with others 
having a buffy or ochraceous tip, except that on the muzzle 
and in front of the ear the particolored hairs have whitish tips 
instead. The fingers and sides of the hind feet are whitish, 
but the latter have the central area darkened. The tail, 
which is slightly shorter than the combined length of head and 
body, has the long hair of the back continued to its basal 
three-quarters of an inch, beyond which the hairs become 
suddenly shorter and somewhat sparse, so that the scaling of 
the tail shows through. The upper side of the tail and the 
terminal inch of its under side are black; the lower side is 
sharply marked off and ochraceous-buff. Total length, about 
375 mm.; tail, 176; hind foot, 45. 

When first described this dwarf hutia was supposedly 
extinct. It was described on the basis of ten small jaws from 
cave deposits probably made by barn owls, in the Sierra de 
Hato Nuevo and near Limones in western-central Cuba. 
Soon after, Dr. H. E. Anthony (1919) procured no less than 


300 mandibles and 60 cranial fragments from caves at Daiquiri, 
on the eastern tip of Cuba, indicating that in former times it 
inhabited suitable areas in the length and breadth of the island 
and probably was regularly preyed upon by barn owls, which 
had brought these remains into the caverns where the birds 
rested by day. None of these fragments appeared to be "at all 
recent and since this mammal formed such a large part of the 
owl diet in times gone by, it would almost certainly appear in 
recent owl pellets" were it still living in eastern Cuba. In the 
meantime, through the efforts of Dr. Thomas Barbour, living 
specimens had been secured in 1917 and sent to the Museum of 
Comparative Zoology by Don Jose Garcia. These were ob- 
tained in the great Zapata Swamp, on the southern coast of 
west-central Cuba, an area that is ordinarily nearly impene- 
trable but can be entered at times of unusual drought. So 
far as known, no other examples have since been obtained, 
until November 1937, when Dr. Hans Boker collected a live 
pair in the same swamp, south of the town of Jaguey. As pre- 
viously stated by Senor Garcia, these had been found living 
on small, dry, bush-covered islands in the swamp, where they 
are probably at most times fairly well protected against invad- 
ing mongooses or other enemies. Senor Garcia told Dr. 
Barbour that the dwarf hutias are sometimes routed out when 
the sawgrass of these marshes is burned. Morrison -Scott 
(1939) records that the pair secured by Dr. Boker were 
brought alive to the Berlin zoo, where on December 1, 1937, 
the female gave birth to a single young one, but it as well as 
its mother died within the next two weeks. These two are 
now preserved in the British Museum, while the adult male, 
which survived until the following February, is now in the 
Berlin Museum. The dentition of the young one, which 
Morrison-Scott figures, already consisted of a premolar and 
the first molar. 

While it seems likely that the species may survive for a 
good many years in the Zapata refuge, it must elsewhere in 
Cuba have succumbed before the introduced mongoose and 
probably the roof rats. 



Capromys brownii J. B. Fischer, Synopsis Mamm., Addenda, p. 389 [= 589], 1830 

("In Jamaica"). 
SYNONYM: Capromys brachyurus Hill, in Gosse and Hill's Naturalist in Jamaica, p. 

471, 1857. 
FIG.: Anthony, H. E., 1920a, p. 163 (photograph). 

This is the first of the short-tailed hutias to be described, 
based by Fischer on the account of the "Large brown Indian 
coney" in Patrick Browne's "Civil and natural history of 
Jamaica," 1789. Originally placed in the same genus as the 
long-tailed hutias of Cuba, it was later included in a separate 
subgenus, Geocapromys Chapman, now regarded as a group of 
generic rank. The characters of the genus in comparison with 
Capromys have been defined by Miller (1929b), of which the 
more important are: First lower cheek tooth with a small 
additional reentrant of the enamel wall on the inner side of the 
first space, making three instead of two indentations; upper 
tooth rows convergent anteriorly, so that the bases of the two 
premolars of opposite sides are in contact instead of widely 
separated; root capsule of the upper incisors passing slightly 
outside of and beyond the premolar instead of stopping in 
contact with its base; bony bar in front of the eye nearly 
vertical or sloping backward instead of forward. Externally, 
it is distinguished by the very short tail. 

This hutia is about the size of a cottontail rabbit but more 
stoutly built, with short legs and a stumpy tail. Above, the 
color is a general dark brown, resulting from abundant parti- 
colored hairs with dark-gray bases and ochraceous tips, mixed 
with all-black hairs, which are most abundant in the middle of 
the back. The lower surface is paler, with a buffy-gray throat 
and lower belly, while the chest and upper part of the belly are 
more thickly clothed with ochraceous-tipped hairs. The tail 
and backs of the feet are nearly clear brown. The ears are very 
short, rounded, and clothed with minute hairs. Total length, 
425 mm.; tail, 50; hind foot, 60. 

This was doubtless a common species on Jamaica in former 
times and was confined to that island. It formed an important 
article of food for the aborigines as attested by the fact that its 
bones were commonly found with ashes of their camp sites. 
It is said that the name Indian cony is from this association. 


Though evidently now much restricted in habitat and abun- 
dance, through hunting and probably through ravages of the 
Burmese mongoose, introduced in 1872, it nevertheless still 
occurs in some numbers in the John Crow Mountains at the 
eastern end of the island where Dr. H. E. Anthony found it in 
1919. All his efforts to discover a trace of it in the Blue 
Mountains, however, were unsuccessful. In the former region 
the native Negro population often hunt them with small dogs, 
specially trained for the purpose. These dogs find the dens 
of the hutias by scent, under large tree roots, or among loose 
rocks. They at once give tongue to their excitement and try 
to enlarge the opening and force their way in a few feet to the 
chamber where the quarry lives by day. Dr. Anthony (1920a) 
recounts a day's hunt of this nature in the course of which a 
few specimens were secured. When the dog has penetrated 
to the den, the hutia voices its alarm in "faint birdlike chirp- 
ings" but is soon dragged out by the dog and at once seized by 
the hunters lest the dogs devour it. "The cony possesses 
sufficient vitality to withstand a great deal of mauling, and 
can put up a good fight." One of them bit a small piece out of 
a dog's nose. "Not infrequently several animals are taken 
from the one hole," perhaps, as with other hutias, representing 
an adult pair and their grown young. In his day's hunt, 
however, Dr. Anthony found but a single animal in each den. 

Jamaican hutia (Geocapromys brownii) 


It is said to live on grasses and the leaves of low shrubs, 
whence conies its local name of "grazee" in use among the 
Negroes. It would be worth while to restrict, if possible, the 
hunting of the species in its mountain habitat in order to 
protect it from extinction as long as may be. Since 1922 it has 
been accorded full legal protection (Journ. Soc. Pres. Fauna 
Empire, pt. 2, p. 15, 1922). 

Whether any other species of this genus or of the allied 
Capromys ever occurred on Jamaica is uncertain. However, 
Miller (1916a) reports that, in examining a few bones recovered 
from ancient burial sites near Salt River, jaws of two sizes of 
hutia were found. Although without teeth, the larger of these 
seemed identical with the living Geocapromys brownii, while 
the smaller, with obviously reduced last molar, was indis- 
tinguishable from the mandible of Geocapromys thoracatus of 
Swan Island. 



Capromys brachyurus thoracatus True, Proc. U. S. Nat. Mus., vol. 11, p. 469, 1888 
(" Little Swan Island . . . entrance of the Gulf of Honduras"). 

This is a slightly smaller and grayer species than the Ja- 
maican animal and was first discovered by Dr. C. H. Townsend, 
when as naturalist on the U. S. Fish Commission steamer 
Albatross, more than 50 years ago he secured specimens on 
Little Swan Island, which lies about 110 miles off northeastern 
Honduras, and at about a quarter the distance separating that 
country from Jamaica. 

While resembling in general appearance the Geocapromys 
brownii of Jamaica, to which it is probably closest allied of the 
known species, this is somewhat smaller and in correlation 
with its drier island habitat is less darkened. The general 
color of the entire upper side is a very even mixture of blackish 
and pale ochraceous, resulting in a rather yellowish-gray 
appearance. The backs of the hands and feet are a nearly 
uniform dusky, minutely punctate with ochraceous. On the 
under side, the most conspicuous feature is a dull whitish half- 
collar extending across the lower throat between the arm pits. 
Elsewhere the color is like that of the upper side but with very 
little darkening, producing a smoky -buff shade. Size less than 
in the Jamaican animal, with a total length of some 15.5 


inches (390 mm.) of which the tail is about 2.25 (57 mm.); 
hind foot, 2.50 (65 mm.). In both these species, the auditory 
bullae do not extend below the level of the basioccipital bones. 

There is very little on record as to the habits of this hutia. 
Lowe (1911), who visited the Swan Islands in 1908, gives an 
excellent picture of the conditions there in his book "A Nat- 
uralist on Desert Islands." The two islands comprise a larger 
one, which is inhabited and serves as a signal station, and a 
smaller lying about a quarter of a mile to the eastward, which 
is untenanted except by wildlife and seldom visited on account 
of the difficulty of making a landing. The latter island is the 
home of many sea birds, which nest on the tops of the scrubby 
bush growth or among its rocks, and is the only place where 
this hutia is found. It is about a mile and a quarter long and 
rises abruptly from the sea with low cliffs on all but a part of 
its northwesterly side. "The plateau formed by the top of the 
island is densely overgrown with trees and bushes, which form 
a closely matted dome of vegetation above its wind-swept 
surface." On the occasion of Lowe's visit it was entirely un- 
disturbed and could be reached from Big Swan Island only on 
very calm days. It appears to be a "big 'stack' of solid coral 
limestone which has been thrust bodily up from the sea bot- 
tom." In the central portion the surface is somewhat more 
even, with vegetable mould and a dense canopy of trees. Here 
Lowe and his party captured several of the hutias. They are 
diurnal in habits and were seen running about or "bolting into 
the big crevasses with which the island is seamed." One 
individual ran over some rocks and tried to escape under the 
spreading roots of a large undermined tree but allowed itself 
to be carefully drawn out and placed alive in a fishing creel. 
It was surprisingly mild-mannered and showed not the slightest 
fear or suspicion, when later it was taken alive to England and 
brought face to face with King Edward's favorite dog, which 
it quietly inspected. The two taken captive became very tame 
and were allowed the freedom of the yacht's deck, "drank 
milk with avidity, and ate any form of vegetable or fruit that 
was offered them." In 1912 George Nelson visited Little 
Swan Island and secured specimens for the Museum of Com- 
parative Zoology. 

At the present time the species seems safe enough in the 
narrow limits of its island home, but should any important 


changes take place there, such as clearing, or the introduction 
of goats or mongooses, its future would at once be in danger. 
How it originally reached this isolated little spot, whether 
through introduction by aborigines or by rafting at some pre- 
historic time from elsewhere, is still an unanswered question. 


Capromys columbianus Chapman, Bull. Amer. Mus. Nat. Hist., vol. 4, p. 314, 1892 

("Cave near Trinidad," Cuba). 
SYNONYMS: Synodontomys columbianus G. M. Allen, Bull. Mus. Comp. Zool., vol. 61, 

p. 5, 1917; Geocapromys cubanus G. M. Allen, ibid., p. 9. 
FIGS.: Chapman, 1892, fig. 3 (type, maxillary bone, and tooth); Allen, G. M., 1917, 

pi., figs. 7-9 (teeth). 

This short-tailed hutia is not known from living specimens, 
yet it must have been common in Cuba up till relatively 
recent times. It was first made known by Dr. Frank M. Chap- 
man from a half-palate lacking the teeth and from a few other 
fragments discovered in a cave near Trinidad, Cuba, "asso- 
ciated with the remains of birds^and bats, and also fragmentary 
pieces of the bones of Capromys." " The cave is situated in the 
southern slope of the coral limestone coast range at an altitude 
of about seven hundred feet, and within two hundred feet of 
the summit of this part of the range . . . The floor of 
the cave was covered to the depth of several feet with a red, 
ferruginous earth, and on this was a layer four or five inches in 
depth of a dark earth in which the bones" were found. 

On account of the fragmentary nature of the original speci- 
mens and of other later-discovered remains, satisfactory com- 
parisons of this Cuban hutia with other forms of the genus are 
yet to be made. It would be important to know the relations 
of the audital bullae to the level of the basioccipital bone, 
which might indicate a relationship with the Bahaman Geo- 
capromys on the one hand or to the Jamaican and Swan 
Island species on the other. Dr. Chapman notices the strongly 
converging upper tooth rows, which, however, as later finds 
showed, do not actually touch in front on the palatal aspect. 
The Cuban animal was evidently as large as the Jamaican G. 
brownii. A median bony ridge on the roof of the palate ends 
abruptly at the palatal margin in the latter and in G. thoracatus 
is continued as a short median projection at that point. In 


both G. columbianus and G. ingrahami, however, the ridge 
fades out before reaching the palatal margin, while the opening 
of the posterior nares seems more narrowed. In the former, 
too, the enamel folds are more nearly at right angles to the 
long axis of the tooth than in Capromys. Toothless jaws may 
often be identified by the indication of the extra small reentrant 
on the inner side of the tip of the anteriormost cheek tooth. 
It is obvious that until better-preserved skulls of this species 
are found, its distinctive characters in comparison with other 
forms of the genus are not very well ascertained. 

This was evidently a common and well-distributed species 
not only throughout Cuba in former days but also on the 
adjacent Isle of Pines. On Cuba, in addition to the original 
locality, Dr. Thomas Barbour secured numerous remains from 
caves in Matanzas Province, as well as from the Isle of Pines, 
and Dr. H. E. Anthony obtained an abundance of jaws and 
palates from owl deposits in a cave at Daiquiri, in eastern 
Cuba. In the latter instance the fragments seemed to be 
mostly of immature and hence small animals, indicating that 
the adults were too large to be managed by the barn owls that 
evidently were the cause of the deposits. Why a species ap- 
parently so common should have died out before any modern 
specimens were obtained, and in so well populated an island 
as Cuba, is not at all clear, for apparently they disappeared 
before mongooses were introduced. Possibly the larger size 
and arboreal habits of Capromys were factors in their survival, 
whereas Geocapromys was ground-living and less able to escape. 


Capromys ingrahami J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 3, p. 329, Dec., 

1891 ("Plana [or Frenchman's] Keys, Bahamas"). 
FIGS.: Allen, J. A., 1891, p. 335, figs. 1, 3, 5-9 (skull and teeth). 

Except for the Jamaican and Swan Island hutias, no living 
member of this short-tailed genus is known other than this 
small species now confined to the Plana Keys between the 
island of Mariguana and Acklins Island in the southern part 
of the Bahama Archipelago. 

The general color is a mixed "yellowish brown, gray and 
black, giving the general effect of grayish brown . 


Below, nearly uniform pale yellowish brown, barely a little 
more yellowish posteriorly . . . some are quite strongly 
suffused with yellowish and have less black; others are faintly 
suffused, giving the general effect above of pale yellowish gray 
mixed with black." A notable feature of the skull is the great 
development of the audital bullae, which in this and the other 
Bahaman races project conspicuously beyond the level of the 
basioccipital bone instead of ending at the same level with 
this bone. In the lower molars, the two reentrant loops of 
enamel from the inner side are relatively short and straight, 
barely reaching the center of the tooth. In size this is slightly 
smaller than the other living species. 

The discovery of this hutia still living in one of the Plana 
Keys is due to D. P. Ingraham, who in mid-February, 1891, 
landed there on several occasions while weatherbound under 
the lee of the island. This key he describes as a small rocky 
islet, the highest point probably not more than 50 feet above 
sea level, "with crevices and caves worn in the rocks by the 
action of water, and in many places broken strata of rocks, 
piled upon each other." It is about half a mile wide and 4 or 5 
miles long, quite without permanent fresh water. In the course 
of two weeks he secured a small series of specimens for the 
American Museum of Natural History. In none of these were 
fetuses found. There appeared to be a concentration of 
individuals at places where conditions were favorable in afford- 
ing many hiding places. He once saw a number of them to- 
gether at a distance from the rocks among palmettos, but later 
concluded that this was a result of the beginning of the rutting 
season. They fed mostly at night, though occasionally foraging 
by day, were not shy, and with caution could be approached 
within 25 or 30 feet before dashing to the shelter of the rocks. 
They fed on the leaves, twigs, and bark of the bushes, "espe- 
cially the black button wood, and the succulent growth of the 
cactus plants," which doubtless supplied the necessary mois- 
ture. They seemed " very fond of the fruit of the paw-paw, and 
even of the body of the tree itself," for in some instances 
trunks were found nearly as large as a man's body "so nearly 
eaten off that they would not sustain their own weight. A 
sweet potato left on the shore was eaten up, while the body of a 
bird, left to tempt them, was untouched." 

So far as the writer knows, no other naturalist visited East 


Plana Key after 1891 until James C. Green way, Jr., landed 
there in February 1933. He found the hutias still present in 
some numbers and brought back a few specimens for the 
Museum of Comparative Zoology. They frequented especially 
small clumps of bushes and would dash out if disturbed, seek- 
ing the shelter of an adjacent clump. 

It seems evident that at some former time these or similar 
hutias were more widely distributed in the Bahama group. 
Catesby mentions a Cuniculus bahamensis and presents a 
figure of what may have been one of the Cuban species of 
Capromys, though no locality is given. According to J. A. 
Allen (1891), the narrators of Columbus's voyage "make 
frequent mention of their abundance not only in the Bahamas 
and at Jamaica, but also in Cuba and Hispaniola." In recent 
years remains of this genus have been brought to light in the 
investigation of caves and aboriginal sites in some of the 
Bahamas, and these prove to be allied but slightly distinct 
forms, a brief account of which may be included here. 


Geocapromys ingrahami irrectus Lawrence, Occas. Papers Boston Soc. Nat. Hist., vol. 

8, p. 190, Nov. 7, 1934 ("Gordon Hill Caves, Crooked Island, Bahamas"). 
FIG.: Lawrence, 1934, fig. 2 (rostrum). 


Geocapromys ingrahami abaconis Lawrence, op. cit., p. 191, Nov. 7, 1934 ("Imperial 

Lighthouse Caves, Hole in the Wall, Great Abaco Island, Bahamas"). 
FIG.: Lawrence, 1934, fig. 3 (rostrum). 

These two forms are very nearly related and may be treated 
together. Both are known from skeletal remains, including 
rostra and some other parts of the skull, secured in the course 
of archeological investigations in the Bahamas in 1933-34, by 
Froelich Rainey. The points of distinction thus far made out 
lie mainly in the details of the rostral bones and in the teeth. 
G. ingrahami irrectus seems to have been slightly larger than 
the Plana Keys animal, still living, with longer tooth row, 
wider individual teeth, and longer zygomatic arch. The two 
enamel loops on the inner side of the three lower molars are 


much longer than in the latter animal, extending as long finger- 
like folds nearly to the enamel wall of the opposite side of the 
tooth instead of about to the center of the tooth. The audit al 
bullae, however, were large, as in typical G. ingrahami, ex- 
tending well below the level of the basioccipital. The lower 
tooth row attained a maximum length of up to 17.5 mm. 
(alveoli); upper tooth row, 16.5; humerus, 50; femur, 67; tibia, 
64. Remains of this form are known from four of the central 
islands of the Bahama group: Crooked Island, Eleuthera, Long 
Island, and Exuma. Those from the first three islands men- 
tioned were in more or less close association with aboriginal 
sites in caves and hence may have been contemporaneous 
with early Indian occupation. Those from Exuma Island, 
however, though undoubtedly of the same form, may have 
been earlier, for the deposit in which they occurred abundantly 
(see G. M. Allen, 1937) showed no sign of human influence, 
and furthermore contained remains of three genera of extinct 
hawks and owls. To what extent the aborigines may have 
utilized these animals as food, or transported them from one 
island to another, is not now possible to make out. 

The Abaco hutia, described' from palates and other cranial 
fragments, was closely similar to that on the Crooked Island 
group, but with the anterior lower premolar "slightly more 
tapering anteriorly with the anterior lobe somewhat smaller 
and more slender" and with longer frontal bones, which ended 
in a pointed projection "reaching anteriorly between the nasals 
and the premaxillaries." The postorbital processes in available 
specimens are more reduced than in the two other forms of the 

Nothing further is known about these Bahaman hutias. 
Possibly they were still living in Catesby's day, but the species 
he figures as a Bahaman cony, about 1750, appears to be a 
Cuban Capromys. Their extinction can not be wholly due to 
the extinct eagle and giant barn owl, which must have preyed 
upon them, but more likely was a result of clearing and burning 
the cover during the earlier centuries of white occupation, of 
which unfortunately there is practically no record. 

Hexolobodon phenax Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 19, Mar. 30, 

1929 ("Small cave near St. Michel," central plain of Haiti). 
FIGS.: Miller, 1929a, pi. 3, figs. 1, la, Ib (palate and jaw). 


This is an interesting genus, closely related to Geocapromys 
and apparently taking its place on Hispaniola, where it was 
contemporary with other species of the recent cave fauna, such 
as Isolobodon, Aphaetreus, and the still-living Plagiodontia. 
At first glance the tooth pattern in the lower jaw is very 
similar to that in Geocapromys ingrahami irrectus in having the 
long enamel reentrants of the two last molars extend well be- 
yond the center of the tooth. The anteriormost tooth, the 
premolar, however, differs from that of Geocapromys and 
agrees with Capromys in lacking the additional small inner 
reentrant to the first section. In the palate the convergence 
of the tooth rows anteriorly is similar to their appearance in 
Geocapromys, and the relation of the incisor roots to the first 
premolar is likewise the same, but, as Miller points out, the 
root of the premolar is "thrown conspicuously forward away 
from that of the first molar." Other characters are the less 
specialized roots of the cheek teeth, which appear to close with 
age instead of continually growing, and "the extension of the 
lower incisor root to the outer side of the mandibular tooth 
row." Size probably about as in the Cuban Capromys pilorides 
but perhaps with a shorter rostrum. Maxillary tooth row 
(alveoli), 22 mm. 

This animal is known from a palate, six mandibles, and four 
isolated cheek teeth, collected by Miller from caves in central 
Haiti near St. Michel and 1'Atalaye. At first sight it might 
easily be taken for a Geocapromys, which is as yet unknown 
from Hispaniola. The characters pointed out by Miller, 
however, appear to be distinctive. It was presumably ex- 
terminated soon after white occupation. 



Plagiodontia aedium F. Cuvier, Ann. Sci. Nat., zool., ser. 2, vol. 6, p. 347, 1836 ("Saint- 

Domingue," taken as Haiti by Miller). 
FIGS.: Cuvier, 1836, pi. 17 (exterior and skull); Miller, 1927, pi. 1, fig. 2. 


Plagiodontia hylaeum Miller, Proc. U. S. Nat. Mus., vol. 72, art. 16, p. 4, 1927 ("Gua- 

rabo, 10 miles east of Jovero, Samana Province, Dominican Republic"). 
FIGS.: Miller, 1927, pi. 1, figs. 1, la-lc (skull). 



Described and figured by Cuvier over a century ago, from a 
specimen brought by Alexander Ricord from Hispaniola in 
about 1826, no additional specimens of the genus had since 
been found living until Dr. W. L. Abbott secured a series of 
ten adults and three young at Guarabo, Dominican Republic, 
about 10 miles east of Jovero, Samana Bay. These he sent to 
the IT. S. National Museum, where Miller (1927), on subse- 
quent study, decided that they represented a species distinct 
from aedium and proposed to call it P. hylaeum. 

Both these species were of similar appearance in life. Cuvier 
describes P. aedium as of rather stout build, the head and body 
about a foot in length, the tail 5 inches and naked. In a general 
way these animals resemble a medium-sized Capromys pilorides, 
but the teeth in both species are very different, with the enamel 
folds extending diagonally instead of transversely across each 
molar. In this respect also they recall the teeth of Isolobodon, 
the Puerto Rican hutia, but the enamel reentrants are much 
more nearly longitudinal in direction, and the upper tooth rows 
are more nearly parallel instead of converging strongly in 
front. There were five claws qn all four feet, although that of 
the thumb was a short, blunt nail. The pelage in P. aedium 
is described as thick, of fine silky hairs above, which were gray 
in the basal three-quarters, tipped with tawny and intermixed 
with longer all-black hairs. The under surfaces were paler 
and clearer, probably nearly buff. The mustachial hairs, as 
usual in this group, were long and abundant. Miller, who ex- 
amined the type in Paris, adds that the tail was naked and 
smooth, its small scales not overlapping but tending to be 
rounded-pentagonal and scarcely a millimeter in diameter at 
30 mm. from the base of the tail. "Ears naked internally, 
thickly furred along edge and apparently on outer side also 
. . . hind foot with claws, 74 mm." 

The distinguishing features of P. hylaeum are given by 
Miller (1927, p. 4) as follows: Anteroposterior diameter of the 
first and second true molars in the lower jaw, each less than the 
transverse diameter instead of equaling or exceeding it, as in 
P. aedium; reentrant enamel folds relatively narrower and 
longer; and the size smaller with the length of the mandibular 
tooth row (alveoli) in adults less than 21 mm., instead of about 
24 mm. as in P. aedium. The color throughout is "nearly 
wood-brown" darkening to buffy brown on chest and belly. 


On the under parts the light brown is uniform, but, above, it 
is "finely intermingled with dark brown." It is thus probably 
a somewhat darker-colored animal than P. aedium, the original 
specimen of which is, however, much faded at the present time. 
The largest specimen of Dr. Abbott's series measured 405 mm. 
in length of head and body; the tail 130; foot, 74. 

It seems likely that at the present time P. aedium, the older- 
known species of Hispaniola, is quite gone and that the smaller 
P. hylaeum is now confined to forested areas in the Samana Bay 
region of northeastern Dominican Republic. Dr. W. L. 
Abbott, who discovered it here in 1923, wrote that his speci- 
mens were obtained for him by an old man who "caught them 
with dogs in hollow trees down near a lagoon near sea shore." 
The four adult females were all pregnant, each containing, in 
early December, a single young, indicating a slow rate of 
breeding, which he observes, "will probably help their extinc- 
tion." He supposes that they may still have been abundant 
in the region and at that time were little hunted for food. They 
can climb to some extent, but their short tail and general 
stout appearance indicate that terrestrial habits are more 
typical. For this reason they are the more liable to attack by 
the introduced Burmese mongoose. Since Dr. Abbott's visit 
in 1923, William J. Clench, of the Museum of Comparative 
Zoology, secured two other specimens for that institution, in 
the same general region, in August 1937. These were an 
adult male and a half-grown animal of the same sex, doubtless 
born earlier in the same year. 

Nothing is known of the habits and haunts of P. aedium 
beyond the few statements of Cuvier, to whom Ricord reported 
that in "Santo Domingo" the animal was known a century 
ago as " Rat-Cay es," or house rat, because it frequented settled 
areas. It was active only by night, hiding during the day. 
The male and female of a pair remained usually together. Its 
principal food consisted of roots and fruits; hence its flesh 
was good to eat, and the Haitians hunted it with such zeal 
that even at that time it had become very rare. From the fact 
that Miller unearthed six lower jaws from a cave in the interior 
of Haiti in 1925 and de Booy had previously secured a jaw 
from old kitchen middens at San Pedro de Macoris on the 
south coast of the island about 60 kilometers east of San 
Domingo City, it may have been that P. aedium was the 


animal of western and southern Hispaniola, and P. hylaeum 
that of the northeastern parts of the island. Even so, the 
ranges could hardly have been entirely separate, for in a later 
paper, Miller (1930), reporting on mammal remains from 
Indian kitchen middens on the valley floor near Constanza in 
the mountainous interior of Dominican Republic, identifies 
both species as present. Evidently both were used as food by 
the aborigines. 

It is significant that this well-distinguished genus is as yet 
unknown in either fossil or recent state from any other Antillean 
islands or from the mainland. 


Plagiodontia spelaeum Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 18, Mar. 10, 
1929 ("The crooked cave near the Atalaye plantation, Haiti"). 

Nothing is known of this supposed third species of Hispanio- 
lan Plagiodontia beyond the account given by Miller, who 
found 15 mandibles, four in the group of caves near St. Michel 
in north-central Haiti, and the>others in the "crooked cave" 
near the Atalaye plantation in the same region. Its main 
point of difference from P. aedium, remains of which occurred 
in the same caves, lies in its smaller size, as indicated by the 
mandibular tooth row of 15.6-16.2 mm., as against 23.6 in the 
former. Compared with P. hylaeum, the discrepancy is less, 
for the latter has a tooth row of 18.6 mm. 



Isolobodon portoricensis J. A. Allen, Ann. New York Acad. Sci., vol. 27, p. 19, 1916 

("Cueva de la Ceiba, near Utuado, Puerto Rico"). 
FIGS.: Allen, J. A., 1916a, pis. 1-5; Anthony, H. E., 1918, pi. 62, figs. la-6b; pi. 63, 

figs, la-b, 4-12c; pi. 74, figs. 6a-c; text-fig. 39, A-H (skeletal parts). 

This well-marked genus is as yet unknown in the living 
state and is undoubtedly extinct, although abundantly repre- 
sented in kitchen middens of the Indians in Puerto Rico, 
the Dominican Republic, and the Virgin Islands. 

The skeletal characters, which alone are available, show that 
it was an animal of about the size of the Hispaniolan hutias, to 
which it is somewhat closely related. Its teeth, however, are 


very different. The upper tooth rows converge strongly 
forward instead of being nearly parallel, while the pattern of 
the enamel folding is simpler. Each of the upper molars has 
a shallow outer and a much deeper inner reentrant loop, the 
latter of which extends inward and forward, diagonally, to 
touch the smaller outer one. The first tooth of the four, the 
premolar, has in addition a second small outer reentrant in 
advance of the one that meets the inner loop. In the lower 
tooth row there are two shallow reentrants of enamel on the 
inner side of each molar, the posterior of which meets the 
single outer reentrant nearly at the center of the tooth. In 
Plagiodontia the inner reentrant is much deeper, runs more 
diagonally forward, and does not meet the smaller fold of the 
outer side. The teeth of young animals have the same pattern 
as do those of adults, for they are continuously growing; but 
the size of the individual teeth increases with age and wear. 

This must have been a common species on Puerto Rico up 
till the period of white occupation. Both Dr. H. E. Anthony 
and I found its bones on the surface in undisturbed caves on 
the island, and they are abundant in kitchen middens, mingled 
with broken pottery and ashes, indicating that they formed a 
staple article of diet for the Indian aborigines. Shortly after 
the discovery of its bones in cave deposits probably of human 
origin in Puerto Rico, Miller (1916b) announced the finding of 
additional jaws and other parts of the skeleton in the course of 
excavating Indian sites at Macoris and San Lorenzo, in the 
Dominican Republic. He was unable, however, to distinguish 
these remains from those found on Puerto Rico. Again, in a 
later paper he (Miller, 1918a), reports the discovery of many 
other bones of this hutia, secured by Theodoor de Booy in the 
course of excavating two Indian sites in the Virgin Islands, one 
at Magens Bay, St. Thomas, the other at Salt River, St. Croix. 
The wide distribution of the species Puerto Rico, Hispaniola, 
and the Virgin Islands the complete lack of differentiation on 
these three areas, and finally the frequent occurrence of the 
remains in deposits of human origin, all point to the fact that 
it was a regular source of food for the aborigines and that they 
carried it about by canoe from island to island, to serve their 
needs. Whether its original home was on Puerto Rico or on 
Hispaniola must remain uncertain, but Anthony (1918) in- 
clines to the view that judged from its apparently wide distribu- 


tion on the former it was indigenous there and that "it is un- 
likely that its primitive range included both Santo Domingo 
and Porto Rico because no differentiation between specimens 
from the two places is seen." The fact that its bones may here 
be found near or at the surface in some deposits is evidence for 
his further belief that it was the last native mammal to become 
extinct on Puerto Rico. This must have been soon after the 
discovery by whites. 

If we assume that the Indians used this as a staple food 
animal, the implication of its presence on these three different 
island habitats is that they kept it in a state of semidomestica- 
tion and perhaps bred it in captivity for food. This they 
apparently did with the guineapig (the cori), remains of which 
also occur in shell heaps in the Dominican Republic and Cuba. 
Possibly, too, this was a particularly docile and tractable 
species, as some of the hystricomorphs are known to be. 



Ithydontia levir Miller, Smithsonian MisA Coll., vol. 74, no. 3, p. 5, Oct. 16, 1922 
(Cave "northeast of St. Michel de 1'Atalaye, northwest end of the central plain of 

Isolobodon levir Miller, Smithsonian Misc, Coll., vol. 81, no. 9, p. 14, Mar. 30, 1929. 

FIGS.: Miller, 1929a, pi. 2, figs. 3, 3a. 

Originally based on a single molar tooth from cave deposits 
in Haiti, further excavations in the same region disclosed that 
this species was after all an Isolobodon and that the original 
tooth was an upper premolar and not a lower molar as was at 
first supposed. 

From the abundant remains secured in the later work in the 
St. Michel area, which lies at the northwest end of the central 
plain of Haiti, Miller (1929a) was able to show that this is the 
most abundantly represented of the vertebrates found in the 
bone-bearing deposits. The fragments of palates and jaws 
are constantly smaller than are those of the Puerto Rican 
isolobodon, the eight largest among 600 mandibles having 
tooth rows 16.2-17.2 mm. in length, while in the latter animal 
the range is from 17.6 to 19.2 mm. Such other measurements 
as can be obtained from crania of the Haitian animal confirm 
its slightly smaller size. 

The evidence now available, as Miller brings out (1929a, 


1929c), suggests that the remains of this genus identified as 
I. portoricensis, occurring abundantly in the Indian cultural 
deposits in the region of Samana Bay, Dominican Republic, 
were probably brought in originally by the Indians, if not 
directly from Puerto Rico, at least as domesticated stock from 
that island, for food. The smaller form native to Haiti and 
doubtless extending to other parts of Hispaniola was found 
sparingly in kitchen middens but is the sole one represented in 
the great number of remains from the caves near St. Michel in 
the central part of Haiti. Imperfect skulls also were found in 
a small deposit at San Gabriel near Samana Bay, a deposit 
which, like that at St. Michel, was probably due to the giant 
barn owl, Tyto ostolaga, now extinct, which must have sub- 
sisted largely on these animals and died out when they be- 
came too few to sustain it. At what time the animal vanished 
is, of course, indeterminate, but the implication is that the 
aborigines about Samana Bay, at least, found it easier to 
subsist on /. portoricensis, which they may have domesticated, 
than to hunt the smaller Haitian animal, which may already 
have become scarce or restricted in distribution at about the 

time of the discovery. 




Aphaetreus montanus Miller, Smithsonian Misc. Coll., vol. 74, no. 3, pp. 3, 4, Oct. 1C, 
1922; also vol. 81, no. 9, p. 16, 1929 ("Larger of the two caves northeast of St. 
Michel tie 1'Atalaye, northwest end of Central Plain," Haiti). 

Fios.: Miller, 1929a, pi. 2, figs. 4, 4a, 4b. 

This is another remarkable octodont rodent, of about the 
size of Isolobodon and closely related to it, but so far as known 
confined to the island of Hispaniola and now apparently too 

It was first described by Miller from mandibles from the 
cave deposit northeast of St. Michel, Haiti, among the moun- 
tains. The teeth differ conspicuously from those of Isolobodon 
and Plagiodontia, which they somewhat resemble in their 
diagonally placed enamel plates, through a further develop- 
ment of these folds, so that the posterior of the two inner 
folds extends across and fuses with the outer, thus cutting off 
a posterior transverse space. In addition the teeth are re- 


markably compressed anteroposteriorly, so that they are 
much narrower than wide. In these characters it represents 
a more advanced stage of development than Plagiodontia and 
Isolobodon, in which the reentrant folds are still separate. 

In the excavations at the type locality carried on after the 
preliminary work had disclosed the two jaws on which the 
original account was based, a more extended search brought 
to light abundant remains of this animal, permitting the more 
detailed description in Miller's (1929a) second paper. Judged 
from the material recovered this was the second most common 
species brought in to the caves supposedly by the now extinct 
giant barn owl. In a later report Miller (1929c) remarks that 
no remains of this animal have as yet been discovered in the 
various kitchen middens and cultural deposits in the Samana 
Bay region; hence there is no real evidence that the species was 
utilized by the aborigines as a food animal, though they may 
have hunted it. In addition to the collection of 17 skulls 
and more than 200 mandibles recovered from the cave deposit 
at St. Michel, other fragments were secured at San Gabriel 
near Samana Bay, from a deposit evidently made by owls, as 
in the case of the former. Equally interesting is the discovery 
of six mandibles, all of immature individuals, in cave deposits 
on Gonave Island off southern Haiti, indicating its wide 
dispersal in Hispaniola in older times (Miller, 1930). The 
animal was apparently unknown to the early Spanish explorers, 
nor is it possible to tell whether it survived later than the early 
years of white occcupation. 


Hcteropsomys insulans Anthony, Ann. New York Acad. Sci., vol. 27, p. 202, Aug. 9, 

1916 ("Cueva de la Ceiba, near Utuado, Porto Rico"). 

FIGS.: Anthony, 1916, pi. 11, figs. 2-3; pi. 12, figs. 1-5 (skull); 1918, p. 407, fig. 40, 
A-D (skull), p. 410, fig. 41 (vertebrae). 


Homopsomys antillensis Anthony, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 187, Jan. 29, 

1917 ("Cave . . . near Utuado, Porto Rico"). 

FIGS.: Anthony, H. E., 1917a, pi. 5, fig. 8 (rostrum); 1918, p. 407, fig. 40, E (skull). 

These two species are known from cranial parts chiefly 
excavated, with remains of other animals, in caves of Puerto 
Rico. In all likelihood they were contemporaneous with 


these latter and may have survived up till relatively recent 
times. They are believed to represent genera closely akin to 
the agoutis of South and Central America but present certain 
differences in cranial and dental characters. They are in- 
cluded here for the sake of completeness. 

Both were perhaps slightly smaller than the typical agouti 
of Trinidad or Brazil, representing the genus Dasyprocta. 
Anthony (1916, 1918) describes Heteropsomys as having a 
skull smaller than that of an agouti, with a total length of about 
70 mm. There are four molars in each jaw, of nearly equal 
size, each with a prominent infolding of enamel on the middle 
of the inner side in the upper teeth and on the outer side in 
the lower teeth. In the few known specimens there are three 
small transverse islands of enamel in the first upper tooth, 
and two in the second (and probably two succeeding true 
molars). An unworn tooth would doubtless show that these 
are the result of wearing away the summits of shallow trans- 
verse depressions in the enamel covering. The molars are each 
provided with two short conical roots. The posterior narial 
passage or interpterygoid fossa extends forward in a V-shape 
to the level of the second molar. In general the skull is less 
elongate and relatively wider than in typical agoutis. Anthony 
writes that judged from the position of the remains discovered 
"this rodent is doubtless of a later age" than either the small 
ground sloth Acratocnus or the large rodent Elasmodontomys 
found in the same cave deposits. 

Concerning Homopsomys, only a few broken crania are 
known from which to determine its relationships. The denti- 
tion is missing from these, except for a few fragments, but the 
skull shows a similar broad, flattened, frontal area, with 
bluntly pointed postorbital projections. In his later paper 
Anthony (1918) writes: "Homopsomys is very closely related 
to Heteropsomys. Indeed it now appears that the two forms 
are probably congeneric, and I would make them so but for 
the fact that the material at hand is rather inadequate." The 
differences between the two, such as the length of palate, 
smaller size of Heteropsomys, its lighter incisors, and more 
convex frontals, may be characters due to age rather than 
specific or generic characters, so that future specimens, if 
obtained, may throw additional light on the relationship of 
the supposed species. Although Miller compares his Brotomys 


with these, Anthony insists that they resemble agoutis rather 
than spiny rats and would even make a separate subfamily, 
Heteropsomyinae, for their inclusion. 

Hitherto these genera have not been found in any of the 
West Indies except Puerto Rico. 



Heptaxodon bidens Anthony, Bull. Amer. Mus. Nat. Hist., vol. 37, p. 183, Jan. 29, 

1917 ("Cave . . . near Utuado, Porto Rico"). 
FIGS.: Anthony, 1917a, pi. 5, figs. 1-6 (palate, jaw, teeth). 

This must have been a remarkable hystricomorph, and it 
adds another to the peculiar mammals of this type known from 
Puerto Rico. It is known only from fragments of the jaws 
secured in cave excavation at Utuado and near Morovis and 
Ciales and hence was probably well distributed in the forested 
parts of the island at no very distant period. Its few remains 
seem to have occurred in much the same manner as those of 
other mammals in these deposits; hence it may be assumed 
that it lived up till about the time of discovery by whites. 

Of about the size of a woodchuck, this animal was notable 
not only for the laminated pattern of its upper and lower 
grinding teeth, but also for the fact that these seem to have 
been reduced to but two in each series. The upper premolar 
was of seven "distinct and separate parallel laminae of strong 
enamel . . . oblique to the main axis of the toothrow," 
while the lower premolar was similar but reversed. In the form- 
er, however, the last enamel space is not quite completely cut 
off from the one in front. In the type fragment a space is 
present for a molar, and the palatal notch extends forward to 
the level of the hind margin of the premolar. In a lower 
mandible there are apparently only the premolar and first 
molar present. The length of the premolar is 11.5 mm. 

According to Dr. Anthony, its discoverer, the affinities of 
Heptaxodon are with the Puerto Rican Elasmodontomys, but 
on the other hand it is so different that both may be regarded 
as representing distinct subfamilies of the Chinchillidae, or it 
may be relegated to a distinct family of its own (Miller and 
Gidley). What relationship if any it may have to the genera 
discovered by the same investigator in Jamaica, and named 


by him Clidomys, Speoxenus, and Spirodontomys, is still to be 
made out. These three are represented by teeth and a few 
other fragments almost all of which were in a hard breccia and 
may be of greater age than the cave material of Puerto Rico. 
The teeth indicate stages of development that may be ancestral 
to such a condition as found in Heptaxodon, but as yet only a 
preliminary account has been published (Anthony, 1920b). 

Family DINOMYIDAE: Giant Rodents 


Quemisia gravis Miller, Smithsonian Misc. Coll., vol. 81, no. 9, p. 23, Mar. 30, 1929 

("The crooked cave near the Atalaye plantation," St. Michel, Haiti). 
FIGS.: Miller, 1929a, pi. 4, figs. 2, 3. 


Elasmodontomys obliquus Anthony, Ann. New York Acad. Sci., vol. 27, p. 199, Aug. 9, 

1916 ("Cueva de la Ceiba, near Utuado, Porto Rico"). 
FIGS. : Anthony, 1916, pis. 13, 14 (skull and teeth); 1918, figs. 25-37 (skull and other 

skeletal parts). 

Since almost nothing beyond skeletal remains of these two 
species is known, and because they stand evidently in close 
relationship, they may be considered together. In general 
size they were about alike and probably were nearly as big as a 
paca, Quemisia, inhabiting Hispaniola, and Elasmodontomys, 
the adjacent island of Puerto Rico. Miller, in describing the 
former, expresses his conviction that it is the animal called 
"Quemi" by Oviedo in his account of the animals of Hispaniola, 
published slightly more than a quarter of a century after the 

These two mammals, the largest of the extinct rodents known 
from their respective islands, are hystricomorphs with rela- 
tionships to the South American Dinomyidae and Chinchilli- 
dae, but since there is no evidence that they had developed 
leaping modifications, it may be better to regard them as 
members of the former, as proposed by Miller and Gidley. 
In both, the enamel pattern of the cheek teeth consists of 
diagonally transverse folds running nearly across each tooth, 
so as to present a laminate appearance, with five enamel 
ridges to each. In Quemisia, according to Miller (1929a), the 


main points of difference in contrast to Elasmodontomys are: 
Reentrant enamel folds less nearly transverse, slanting for- 
ward at an angle of about 21 instead of about 50; the man- 
dibular symphysis longer, extending back beyond the middle 
of the first molar instead of barely to the middle of the pre- 
molar; shaft of the lower incisor not extending behind the 
symphysis instead of far beyond it to terminate beneath the 
middle of the second molar. The incisors show a shallow 
depression on their front face, slightly more marked in the 
lower than in the upper pair. The skull of Elasmodontomys 
was about 5 inches long and proportionately broader than in 
the Cuban Capromys pilorides. Anthony (1918) has well 
illustrated the principal bones of the skeleton. He shows that 
the sacrum consists of four well-fused vertebrae and that the 
tail was probably short. It seems to have been a heavy- 
bodied animal, and its short phalanges indicate terrestrial 
rather than climbing habits. 

According to the brief description of Oviedo, the "Quemi" 
resembled the hutia in color but was larger. That it was util- 
ized by the natives of Hispaniola as a food animal is indicated 
by the presence of its limb bones at a depth of about 4 feet in 
a kitchen midden near the entrance of a cave at Boca del 
Infierno, in the Samana Bay region, Dominican Republic 
(Miller, 1929c). It must have become extinct soon after the 
coming of the Spaniards, probably about the first half of the 
sixteenth century. As to its representative on Puerto Rico, 
Elasmodontomys, nothing is known. Anthony found its bones 
"well bedded in the red stalactite formation, the deeper 
layers of the cave deposit," so that it may have died out at an 
earlier time. 

Here may be mentioned appropriately a third but much 
larger member of the same group, a rodent almost as large as 
a black bear, to which Cope (1868) gave the name Amblyrhiza 
inundata and later synonyms based on single teeth of the same 
animal. These teeth were discovered in a cargo of cave earth 
sent to Philadelphia from the island of Anguilla at the extreme 
northeastern point of the Antillean chain. Additional frag- 
ments were later discovered in a cave on the adjacent island 
of St. Martin. More recently several fragments, including a 
large part of a skull and portions of the palate and incisors, 
have come to light in the museums at Leiden and Delft, 


collected over 50 years ago. These supply additional facts 
concerning the structure and indicate that the animal was 
closely allied to the "Quemi" of Santo Domingo, with essen- 
tially the same enamel pattern of the molars but with a rela- 
tively longer rostrum. A full account of these has been pub- 
lished by Schreuder (1933) with figures. The complete skull 
is estimated to have been about 400 mm. (16 inches) in length, 
and hence the animal must have been a giant in comparison 
with the "Quemi". How long ago this species lived, how it 
reached the two neighboring small islands on which its remains 
are found, and why it should have perished there are questions 
still impossible to answer. That it lived well after glacial 
times is probable, but whether it was a contemporary of the 
aboriginal Indians there is uncertain. 

If Miller is correct in identifying his Quemisia grams with 
the "Quemi" of Oviedo, the name should stand as Quemisia 
quemi (J. B. Fischer), since this name was given in 1830 by 
Fischer [Synopsis Mamm., Addenda, p. 389 ( = 589)], who 
quotes Oviedo and Latinizes his description. He includes as 
synonymous, however, the account of the "Spanish Racoon" 
from Browne's "Civil and Natural History of Jamaica," an 
unidentified animal that was probably the larger Cuban hutia, 
Capromys pilorides. 

Family DASYPROCTIDAE : Agoutis 


Dasyprocta albida Gray, Ann. Nat. Hist., ser. 1, vol. 10, p. 264, 1842 ("St. Vincent's," 
West Indies). 

In former times agoutis were doubtless present in most of 
the Lesser Antilles, and they still occur or did so until modern 
times on at least the following (named in order from south to 
north): Tobago, Grenada, St. Vincent, Barbados, St. Lucia, 
Dominica, Guadeloupe, Montserrat, and St. Kitts. This 
carries them all the way, practically, from Trinidad and South 
America, north throughout the chain to the northeast corner 
of the Antilles, a point where the distribution of some other 
Lesser Antillean animals stops, and beyond which to the 
westward the distribution of certain Greater Antillean species 


begins (such as that of Amblyrhiza on Anguilla and its relatives 
on Puerto Rico) . It is difficult to say whether this is a natural 
distribution, owing to the species having succeeded in crossing 
from island to island northward, or whether it is due to human 
agency whereby they were introduced and successfully colon- 
ized. Their flesh is considered excellent food, and they are not 
difficult to keep in captivity; hence it is reasonable to believe 
that the aborigines may have been responsible for their intro- 
duction from the continent or Trinidad to the various islands 
to the northward. Du Tertre, writing in 1654, says that 
among the French islands implying St. Kitts, Guadeloupe, 
and Martinique they were well known at that time and were 
much hunted for their flesh, dogs being trained to run them. 
When pursued, they seek shelter in a hollow log or tree whence 
the hunters smoke them out. The female was said to bring 
forth two young at a time, in a nest made on the ground under a 
bush. In early days, too, it was said that their sharp incisor 
teeth were used as blades, probably set in some kind of a 
handle, by the Caribs for cutting their skin all over their 
bodies to draw blood in their religious ceremonies. A later 
French missionary, Labat, writing in 1742, said that he be- 
lieved it was found on all the islands, although he acknowledged 
that he did not know of its presence on Martinique. It was 
then common on Dominica, Guadeloupe, and St. Kitts. Its 
absence, even at the present day, on Martinique is interesting 
and requires explanation. Possibly the introduction of so 
poisonous a snake as the bushmaster in early times may have 
exterminated it there if it previously had been present. Per- 
haps, too, volcanic eruptions, with their discharge of poison 
gas, may have had an occasional disastrous effect in the past, 
as when Mount Pelee went into action a generation ago. 
Agoutis are more or less diurnal in habit, feeding on vegetable 
matter, and are especially fond of certain forest fruits, so that 
hunters sometimes resort to the expedient of sitting in such a 
tree and attracting the agoutis by throwing a stone from time 
to time through the leafy branches. The agoutis evidently 
mistaking the noise of the stone crashing through the leaves 
to the ground for the fall of one of the fruits may sometimes be 
seen cautiously coming from the thicket where they have hid- 
den, to seek the fruit, and thus expose themselves to a shot. 
In external appearance this agouti does not differ essentially 


from the Dasyprocta rubatra of Trinidad. The forehead, back, 
flanks, chest, and belly are a mixture of blackish with small 
tickings, which vary in color from ochraceous or buff on fore- 
head and shoulders to orange-rufous on the lower back, 
ochraceous-buff on the flanks and belly, and whitish on the 
lower throat. The chin is nearly clear whitish to buffy; the 
inner sides of the thighs are bright buff; the nape, limbs, and 
feet are uniform brownish black, and the central part of the 
back may vary to a darker blackish tone. The hind feet have 
but three toes, the front feet four, with stout short claws for 
digging. There is a short, naked, stumplike tail. Ears short 
and rounded. Length of head and body up to 18 inches. 

It is perhaps uncertain how far the agoutis on the different 
Leeward Islands have become differentiated into island races. 
No comparisons have yet been made with sufficient series of 
adults from them to be very significant. Nevertheless, as long 
ago as 1842, J. E. Gray named as a new species an agouti 
from St. Vincent, calling it Dasyprocta albida, on account of 
its being "whitish grey, nearly uniform, the hair of the back 
elongated, white at the base." This specimen must have been 
somewhat albinistic, and probably young, for it is said to have 
been the size of a guineapig. On this basis alone the St. 
Vincent agouti is not distinguishable, for other specimens from 
there prove to be of normal coloring. Nevertheless, scanty 
material seems to indicate a smaller size as compared with D. 
rubatra of Trinidad, with a shorter hind foot (about 95 mm. 
instead of 110), and a smaller skull, having a shorter and 
more sharply tapering rostrum and shorter nasals. In view of 
these apparent differences the St. Vincent agouti may be re- 
garded as distinct and Gray's name will apply to it. 

As to the present status of this race, for such it seems best 
regarded, little is known. Sclater, in 1874, recorded two 
specimens sent to the Zoological Society of London in August, 
1868. In February, 1904, Austin H. Clark procured an adult 
female there in the forest behind Barrouallie, where probably 
it may still be found in small numbers. A similar dearth of 
recent information exists in regard to the status of agoutis on 
the other Leeward Islands. However, where large stretches of 
forest still remain, there the animals may be expected to survive 
until further clearing or the introduction of such predacious 
animals as mongooses takes place. That on some of these 



islands apparently local races differing in slight characters of 
size or proportion are recognizable, points to an introduction 
of agoutis at a long-distant period or else they reached their 
present localities by occasional accident at equally remote 
times. On a visit to Grenada in 1910, I inquired of native 
hunters, who agreed that an agouti still occurred in the moun- 
tain forests of the interior, but I was unable to secure a speci- 
men. In the nearby group of small islands known as the 
Grenadines, Austin H. Clark tells me that an agouti has been 
introduced at the southeastern end of the island of Bequia, but 
apparently it has not thrived, perhaps on account of the lack 
of water. The Museum of Comparative Zoology has a skin of 
the Antillean agouti from Grenada that was received about 
1879. Another specimen, collected on Barbados about 1870, 
came from Governor Rawson and presumably was of local 
origin, though the agouti is not mentioned in Hughes 's account 
of the natural history of Barbados in 1750. With the gradual 
clearing of wooded areas on any of these islands and the intro- 
duction of the mongoose (already common for a number of 
years past on Grenada) it is tp be expected that agoutis will 
eventually be extirpated from all the Lesser Antilles, perhaps 
before specimens can be had sufficient to determine the extent 
of their local variation. 

St. Vincent agouti (Dasyprocta albida) 




Dasyprocta antillcnsis Sclater, Proc. Zool. Soc. London, 1874, p. 666 (St. Lucia, West 

FIG.: Sclater, 1874, pi. 82 (colored). 

In November 1874, two agoutis from the island of St. Lucia, 
West Indies, were presented alive to the Zoological Society of 
London and were shortly afterward described as a new species 
by Sclater, who, however, compared them only with the very 
different Dasyprocta punctata of Central America. His colored 
plate shows an animal of closely similar appearance to the D. 
rubatra of Trinidad or the D. albida of St. Vincent, so that 
there can be no doubt of the near relationship of these two. 
That the name may stand for the island animal as a valid race, 
however, is indicated by a comparison of two skulls from skins 
in the Museum of Comparative Zoology, taken on St. Lucia 
probably about 1880. 

This agouti is nearly the same in size and appearance as D. 
rubatra, but in the skulls available it has shorter nasals, which 
seem more narrow and tapering distally. The rostrum is 
shorter, and the incisive foramina are prolonged posteriorly 
to the meeting of the premaxillary with the maxillary, instead 
of ending in advance of that point. How far these characters 
will hold with a larger series remains to be seen. Nor is it 
certain whether agoutis from other islands of the Lesser 
Antilles are locally different. 

No recent information is at hand as to the status of the 
agouti in St. Lucia at the present time. On the neighboring 
island of Dominica Allen and Chapman (1897) recorded a speci- 
men from the forested part, and it was said to be still common 
in the interior. On Montserrat it occurred also until at least 
very recent years, for the United States National Museum has 
a specimen received in 1902 from that island. The skull of this 
individual shows a number of peculiarities, which if borne out 
by additional skulls might warrant naming the agouti of this 
island as a distinct local race. Agoutis of this type must once 
have been common on St. Kitts, for Du Tertre (1654) and 
Labat (1742) both mention them two or three centuries ago. 
Two skins from this island are recorded in the catalogue of 
mammals in the Museum of Comparative Zoology as having 


been received about 1881, but they have since been lost sight 
of. There is another in the United States National Museum. 
Presumably the species may still be present there. St. Kitts 
is thus the most northern of the Lesser Antilles from which it 
has been reported. It may be added that the South American 
Dasyprocta aguti, characterized by its bright ochraceous or 
rufous rump, is the one introduced on St. Thomas. Miller 
(1911) reported the identity of this agouti through a specimen 
received by the National Museum, while in 1917 the Museum 
of Comparative Zoology received one from the same island. 
In the island of Guadeloupe an agouti has long been known, 
and in 1914 I named it on the basis of two specimens secured 
by Dr. G. K. Noble. 



Dasyprocta noblei G. M. Allen, Proc. New England Zool. Club, vol. 5, p. 69, 1914 
("Goyave, Guadeloupe Island, West Indies"). 

Two specimens secured in ?914 by Dr. G. K. Noble differ 
notably from available skins of the neighboring islands in 
their darker appearance, due to extensive blackening of the 
head, neck, shoulders, and back, with a suppression of the 
reddish or ochraceous ticking, which becomes confined to the 
lower flanks. In a third specimen, later obtained, however, 
the opposite effect is seen, through the concentration of rusty 
tips to the hairs of the entire back, perhaps an individual and 
erythristic condition. If the agouti of Guadeloupe thus proves 
to be distinct from those on the other islands, it may eventually 
be better to regard it as merely an insular race of Dasyprocta 

As to the status of this animal on Guadeloupe, Dr. Noble 
reported that at the time of his visit in 1914 it was difficult to 
secure and was confined to a limited area of uncleared country. 
The animals make runways and are occasionally obtained by 
waiting nearby for a shot as one may chance to appear. 


Order CARNIVORA: Dogs, Cats, and Their Relatives 

The larger carnivorous mammals that compete with man for 
food, destroy domestic animals, or threaten man's security, 
have been exterminated or greatly reduced in settled areas. 
Seven families of true carnivores are recognized : 

(1) Canidae, the dog family, includes wolves, jackals, foxes, 
and some less familiar types. Representatives of this family 
are found throughout the world, except in New Zealand and 
some of the oceanic islands. 

(2) Ursidae, the bear family, formerly occurred throughout 
Europe, Asia, and North America and in the East Indies, the 
Atlas Mountains of North Africa, and in the South American 

(3) Procyonidae, the raccoon family, includes the pandas, 
the coati, the kinkajou, and the cacomistle or ring-tailed cat. 
Except for the Asiatic pandas this group is confined to the 
New World. 

(4) Mustelidae, the weasel family, includes the small fur 
bearers weasels, minks, otters forms that suffer persecution 
chiefly for the sake of their valuable pelts. The weasels and 
their relatives occur in all regions except the Australian and 
the oceanic islands. 

(5) Hyaenidae, the hyaenas and the aard-wolf, are restricted 
in Recent times to Africa and southern Asia. The African 
aard-wolf is a termite-eating mammal with degenerate teeth 
and jaws; it is rare everywhere in its range. 

(6) Viverridae, the genets, civets, and their allies, are catlike 
or weasellike carnivores with partially retractile claws and with 
numerous cheek teeth. Representatives are found in the Ethi- 
opian, Oriental, and parts of the Palearctic regions. 

(7) Felidae, the cat family, is world wide in distribution, 
except for the Australian and polar Regions and the oceanic 
islands. Except for the fossane (Cryptoprocta) of Madagascar, 
which is genetlike, the cats are a very uniform group, differing 
in little except size, length of tail, and color pattern. 

Four families are considered in the present volume : 

(1) Ursidae: Three races of the black bear, all the grizzlies, 
and Alaskan brown bears. 

(2) Mustelidae, weasels, martens, mink, and wolverene: 20 
species and subspecies, belonging to three genera, are discussed 
here; all are valuable fur bearers. 


(3) Canidae, foxes and wolves: 22 forms, representing two 
genera, have been exterminated or endangered. 

(4) Felidae, the cat family: 10 races of the puma and two of 
the jaguar are treated here. J. E. H. 

Family URSIDAE: Bears 


The black bears are much smaller than the grizzlies, with 
shorter claws and usually with black coloration, although in 
western North America several local color phases occur. The 
skull is characterized by its relatively short snout and short 
nasals in comparison with the grizzly bears; the length of 
nasals is equal to, instead of being more than, the width across 
the front of the first upper molars, and the two posterior cusps 
of the first lower molar instead of being in approximately the 
same transverse plane are arranged so that the outer is slightly 
in advance of the inner. The black bears were formerly found 
all over the wooded parts of North America south into northern 
Mexico, but at the present tmie this range has been locally 
restricted through killing them out in the more settled areas. 
Nevertheless so wary and intelligent are these animals that 
they have in many regions been able to survive in spite of per- 
secution, and hence only a brief consideration is here needed. 
Hitherto, in addition to the typical form of the eastern parts of 
North America, no less than 13 local forms have been described. 
The value of these can not be fully estimated until a general 
study is made of the entire group with adequate material. 
However, in a recent paper, Dr. E. R. Hall (1928) has reviewed 
the bears of this group inhabiting the Northwest coast and 
concludes that the seven forms described from the region are 
all subspecies of the typical black bear. In the western United 
States and on the Pacific coast, a cinnamon phase occurs as 
far north as Taku River, Alaska, and young in the same litter 
may be either black or cinnamon in color. A gray or bluish 
phase is found in the region between Lynn Canal and Cape St. 
Elias, and this has been described as Ursus emmonsii, the 
glacier bear. Hall (1928) points out cranial characters that 
may serve to distinguish these bears as a local race apart from 
color, for black individuals occur within the same range. 


Finally, on Gribble and Princess Royal Islands, British 
Columbia, three color phases occur, the lightest of which is 
nearly white and has been named Ursus kermodei. But "the 
light phase is not always pure white and may include yellowish 
rufous and bright golden rufous bands and spots." Since the 
color characters of Kermode's bear offer but slight grounds for 
distinction, Hall recognizes this as a valid race chiefly on the 
basis of minor cranial characters. 

In the following account only a bare outline of the present 
status of some of the forms seems necessary. 



Ursus americanus Pallas, Spicilegia Zoologica, pt. 14, p. 5, 1780 (eastern North 

America) . 
FIGS.: Elliot, 1901, pi. 34 (three views of skull); Nelson, 1916, p. 440 (col. figs.). 

Since the settlement of eastern North America by whites, 
the black bear has retreated from the more populous or cleared 
sections, but, given a reasonable area of forest, it will readily 
adapt itself to contact and persist in wild areas. In eastern 
United States a few bears still exist in western Massachusetts, 
they are common in the less settled parts of Maine, and are 
present in small numbers in the White Mountains of New 
Hampshire and the mountainous parts of Vermont. They no 
longer are found in Rhode Island or Connecticut but remain in 
some numbers in the Adirondacks of New York State and in the 
Pennsylvania mountains. According to recent census esti- 
mates of the U. S. Fish and Wildlife Service, the approximate 
bear population of these States is: Maine, 1,400; New Hamp- 
shire, 650; Vermont, 400; Massachusetts, 10; New York 1,000. 
Rhoads in 1903 said that at that time it was almost extermi- 
nated in New Jersey; however, a few seem to have persisted in 
the cedar swamps of the southern part of the State and in 
the adjacent parts of Delaware. A bear was reported to have 
been killed in Johnsonburg, N. J., in 1920. Pennsylvania, 
with its large areas of mountain forests, is estimated to have 
a bear population of 3,300; Maryland, 150; and West Virginia, 
2,100, in the mountainous regions. Virginia has fewer, an 
estimated 600; North Carolina nearly twice that number, 
South Carolina 230, and Georgia 400. West of the Alleghenies 


bears are still present in numbers in the wilder parts of Michi- 
gan and Wisconsin, but they were exterminated from Illinois 
by 1860 and from Indiana by the middle eighties. Tennessee 
still harbors a good number in the mountains, but Kentucky, 
which never was very good bear country, holds only a few at 
the present time. There are a few in Mississippi, and formerly 
they were no doubt present in Missouri, but the census quoted 
includes none from the latter State. Hibbard (1933) writes that 
though once common in eastern Kansas, the black bear is now 
extinct in that State. 

Northward of the United States, black bears range through 
New Brunswick into southern Canada and as far northward as 
Ungava. They occur also in Newfoundland but are said now 
to be scarce. In the wilder regions they extend across to Alaska 
in timbered country. Bears of this species avoided the open 
plains country but formerly extended into the valleys of 
North Dakota. At present they are probably very scarce or 
nearly gone in that State. 

Whether the black bear of Florida and Alabama, known as 
Euarctos floridanus, and the so-called yellow bear of Louisiana, 
Euarctos luteolus, are to be Regarded as races of Euarctos 
americanus or as separate species is still uncertain. At all 
events, the former is now rare in Florida, though in some 
sections it still occurs and the Wildlife Census credits the State 
with 350 and Georgia to the north with 400. In Alabama, 
according to A. H. Ho well (1921), bears are at present ex- 
terminated everywhere except in the swamps of the southern 
part. Skulls from this region are intermediate between the 
two forms noted above. The census mentioned gives the 
State a population of 300 bears. Westward bears are present 
over suitable country to central California. In this State they 
seem to have increased in abundance with the disappearance 
of the grizzly. They formerly extended to northern Mexico 
and no doubt still occur there, but no estimates of status are at 



[Ursus americanus] var. emmonsii Dall, Science, new ser., vol. 2, p. 87, 1895 (St. Elias 

Alps, near Yakutat Bay, Alaska). 
FIG.: Nelson, 1916, p. 440 (left col. fig.). 



This is a race of the black bear, distinguished from the 
typical form by slight cranial characters. As might be ex- 
pected it is in these characters most like the race perniger, of 
the Kenai Peninsula, but "has the rostrum decidedly longer 
and inflated anteriorly over the canines. Also the upper 
molars are smaller and the mastoid and zygomatic breadths 
are greater" (Hall, 1928). The name will apply to the black 
bears of the mainland of southern Alaska. In this race a dilute 
color phase occurs, which is blue-gray in appearance, but not 
all the bears of the region are thus colored, so that this peculi- 
arity, which first led to its recognition in nomenclature, is not 
characteristic of all the individuals. 

The blue bear, or perhaps in particular those individuals 
showing the "blue" phase, is supposed to be "threatened with 
extinction" because of over-hunting and because this phase has 
a circumscribed distribution, in the vicinity of the Glacier Bay 
National Monument. However, since this latter is now a 
reserve, adequate patrolling of the region to prevent poaching 
should insure its perpetuation. 



Ursus kermodei Hornaday, 9th Ann. Kept. New York Zool. Soc., for 1904, pp. 81-86, 

Jan., 1905 (Gribble Island, British Columbia). 
FIGS.: Hornaday, 1905, 2 pis.; Allen, J. A., 1909, p. 234, fig. 1 (skin); p. 236, figs. 2, 3 

(skull); Anthony, H. E., 1928, pi. 4, opp. p. 76 (colored). 

Kermode's bear (Euarctos americamis kermodei) 


Originally believed to be a species distinct from the black 
bear and characterized as clear creamy white to the roots of 
the hairs, further specimens of this bear from the type locality 
show that this coloring is not constant but skins may have the 
top of the head "yellowish rufous, with the back . . . con- 
spicuously varied with bands and irregular patches of bright 
golden rufous" (J. A. Allen, 1909). The skull presents dis- 
tinctive characters that warrant its recognition as "a strongly 
marked form"; however, according to Hall (1928), the latest 
writer to consider its status, it is best regarded as a local race 
of the black bear. The known range extends "from the lower 
part of South Bentinck Arm, Bella Coola River (lat. 52), and 
from Aristazable, Princess Royal, Gribble, and Pitt Islands, on 
the coast, to a considerable distance into the interior" of 
British Columbia. A list of specimens and localities up to 
1909 is given in the paper of J. A. Allen (1909). 

Because of its circumscribed range, and its relative con- 
spicuousness, it might be thought that this type of bear would 
be a special object of pursuit and thus be in danger of early 
extermination. However, I. M. Cowan, in a verbal communi- 
cation to Dr. Francis Harper in 1937, regards it for the present 
as "safe enough." It may be thought of as a geographical 
race of the black bear, with in some cases a white color phase. 


In the early settlement of western North America, the 
grizzly bear was about the only really dangerous carnivore to 
be found; its size and strength as well as its rather truculent 
nature combined to make it a formidable beast against which 
the rifles of earlier days were often ineffective, and many a 
hunter has owed his life to the fact that the grizzly is too large 
and heavy to climb a tree that afforded the hunter a refuge. 
Because of these traits and its uncertain temper, as well as on 
account of its depredations among the rancher's cattle and 
sheep, the grizzly bear has been persistently hunted and killed 
wherever possible for upward of a century, with the result 
that at the present day it has been largely exterminated over a 
great part of the former range. 

Merriam (1896) in his preliminary review of the North 
American bears wrote: "The Grizzly bears (including the 


Barren Ground bear) may be separated into 4 more or less well- 
marked forms, as follows: (a) the true Grizzly, Ursus horribilis 
Ord, from the northern Rocky Mountains; (b) the Sonoran 
Grizzly, 'var. horriaeus' Baird, probably only a subspecies; (c) 
the Norton Sound, Alaska, grizzly, probably another sub- 
species; (d) the very distinct Barren Ground bear, Ursus 
richardsoni Mayne Reid. Whether or not the large Grizzly 
from southern California deserves subspecific separation from 
the Sonoran animal (horriaeus) has not been determined." 
Following this preliminary statement of the case, Dr. Merriam 
in succeeding years brought together an enormous series of 
specimens from the larger part of the range of the grizzly, and 
after intensive study of this material he came to the conclusion 
that no less than 86 species of grizzly and big brown bears 
could be distinguished as inhabiting western North America! 
The main points of difference lie in cranial peculiarities, and 
it is uncertain how far the species may be distinguished by 
external characters. To many, Dr. Merriam writes, this num- 
ber of species of grizzly bears will appear "preposterous," yet 
the differences in cranial details are evident on close study. 
It becomes then a matter of interpretation as to their signifi- 
cance. Thus we do not yet know whether the five species of 
grizzly supposed to live together on Admiralty Island will 
actually interbreed, or whether the 15 that Dr. Merriam has 
described as occurring in British Columbia maintain pure lines 
of descent or hybridize. He adds: "Cranial and dental char- 
acters among the big bears are very subtle. As a rule com- 
parison of any two skulls of essentially the same size brings to 
light so many resemblances that one is likely to infer a far 
closer relationship than actually exists. This is because the 
big bears of the genus Ursus are such a closely interrelated 
group that the resemblances far outnumber the differences. 
Hence the greatest caution is necessary to avoid misleading 
conclusions." Under these circumstances a conservative 
course of treatment may be adopted and Dr. Merriam's 
original inclusive groups of grizzly bears be considered first in 
a very general sense; then the various forms that he since dis- 
tinguished may be listed with a brief statement of supposed 
range and status. 

In a general way the more typical grizzlies inhabited the 
West from the edge of the Plains to the Pacific, south into 


northern Mexico, and north into Alaska and Canada. The 
barren-ground grizzly is the type of subarctic western Canada; 
the big brown bears are found along the coast from northern 
British Columbia to and including the entire Alaskan Penin- 
sula, but not to the exclusion of the grizzly -bear forms. From 
an economic point of view the grizzly bear is regarded as a 
destroyer of game and more especially is disliked on account 
of its depredations against livestock, or, recently in northern 
Alaska, against the reindeer herds. On the other hand, it 
helps keep down the increase of certain burrowing ground 
squirrels, and in parts of the North its intestines are used in 
making clothing. Its present value to human beings may well 
be largely an esthetic and recreational one. To an increasing 
number of persons who enjoy the sight of large wild animals or 
the thrill of photographing them, these bears can not fail to 
give much pleasure, while to the naturalist and to the sports- 
man they are a source of interest. The money brought in to 
the parts of the country where they are pursued is a substan- 
tial help to the local population. 



Ursus horribilis Ord, Guthrie's Geography, 2d Amer. ed., pp. 291, 300, 1815 ("Mis- 
souri River, a little above mouth of Poplar River, northeastern Montana, " fide 

SYNONYMS: For a full list of the names applied to grizzly bears see Merriam, 1918. 

FIGS.: Audubon and Bachman, 1854, Quadrupeds of North America, vol. 3, pi. 131; 
Baird, 1857, pis. 41, 42 (skull and teeth) ; Nelson, 1916, p. 442 (col. fig.) ; Merriam, 
1918, pis. 1-16 (living animal and skulls). 

Merriam (1918) writes that "the differences between the 
grizzlies on the one hand and the big brown bears on the other 
are neither so great nor so constant as at one time believed 
. . . The typical brown bears differ from the typical 
grizzlies in peculiarities of color, claws, skull, and teeth. The 
color of the former is more uniform, with less of the surface 
grizzling due to admixture of pale-tipped hairs; the claws are 
shorter, more curved, darker, and scurfy instead of smooth; 
the skull is more massive; the fourth lower premolar is conical, 
lacking the sulca^e heel of the true grizzlies. But these are 
average differences, not one of which holds true throughout the 


Baird describes the general color of grizzly bears as varying 
from a nearly uniform dark brown with the tips of the hairs 
lighter, or brownish yellow, to a condition in which there is a 
dark blackish dorsal stripe, and the sides of the body are pale 
brownish yellow, bordered by a dark stripe along the flanks. 
The limbs are generally black or dark brown; head and belly 
yellowish, and a tinge of hoary over the shoulders and sides. 
Claws long and nearly straight, those on the fore feet about 4 
inches or more long. Skull long, the length of nasals exceeding 
the width across last premolars. First lower molar with the two 
posterior cusps on about the same transverse plane, and with 
a small accessory cusp or two on the inner side between the 
anterior and posterior cusps. Weight upward of 1,200 pounds, 
but males are larger than females; "in some the disparity in 
size is very remarkable ... in a few cases the difference 
is slight" (Merriam). Bear skulls undergo a series of changes 
from early life to old age and in most species do not attain 
their mature form until seven or more years of age (Merriam) . 
Baird states that the largest specimens are "from six to seven 
feet" in total length. The total length of skull runs to 14 or 
15 inches. 

In a general way the former range of the grizzly extended 
from the eastern edge of the Great Plains in about the longi- 
tude of the one-hundredth meridian, westward to the Pacific 
coast in northern Lower California, thence south to Durango, 
Mexico, and northward to the southern and central parts of 
the Alaskan Peninsula and the Arctic coast. In the barren- 
ground region west of Hudson Bay it is represented by the 
so-called barren-ground grizzly. Baird, in 1857, wrote that 
"it appears first to occur on the Missouri, above Fort Pierre 
[in South Dakota], and becomes more and more abundant 
higher up on the Missouri, and especially on the Yellowstone; 
thence to the Rocky Mountains, which it inhabits throughout 
its entire extent in the United States . . . To the north 
it extends far into the British possessions, and southward into 
Mexico." It is safe to say that at the present time the grizzly 
bear is extinct over much of this area and survives only in 
some of the national parks or in wilderness regions. 

In their general habits grizzly bears like somewhat open, 
rugged country with scattering thick growth where they may 
take shelter. Their food includes a wide range of both animal 


and vegetable matter. In early times they preyed partly upon 
the larger game and animals, such as deer, wapiti, or, on the 
plains country, the bison, and later found an easier prey in the 
introduced livestock of the settlers. In the Southwest, Bailey 
mentions their fondness for acorns, pinyons, cedar berries, 
various succulent roots, the sweet mesquite berries, bulbs, and 
tubers of various kinds. They also turn over logs and stones 
for the insects to be found underneath. In more northern 
parts of the range they dig out the burrows of small rodents, 
especially of ground squirrels, for their occupants. Fish too are 
very much relished, and when salmon are running in the 
Alaskan waters the grizzlies and brown bears feast upon them, 
wading in and scooping out the fish. 

The grizzly bear hibernates in the northern part of its range 
from late September to April, more or less, seeking a shelter 
among rock crevices or in the hollow base of a large tree or 
under its upturned roots. In the southern latitudes, as in New 
Mexico, however, Bailey says that hibernation is less regular, 
and the bears may be found active at various times in winter. 
The young are born in these winter dens and may be two or 
three in number, more commonly twins. 

In various papers Vernon Bailey has much to say concerning 
the grizzly's former range and abundance. In his "Mammals 
of North Dakota" (1926) he presents a number of early records 
of its occurrence along the eastern border of its range here. 
"At the coming of the white man these large grizzlies were 
apparently common over practically all of North Dakota." 
Alexander Henry while in the Red River Valley in 1800 found 
them not so numerous as in the Hair Hills and Devils Lake 
region, where they were said to be "very malicious." He also 
reported one skin in the catch from the Sandhill River, Minn., 
in 1807, and one from Portage la Prairie, Manitoba, localities 
from near the limit of its eastward range. In 1833, Maxi- 
milian found them more common the farther up the Missouri 
he went. Generally bears were found feeding here on buffalo 
carcasses, which were often plentifully distributed in the quick- 
sands or along the river banks by floods and breaking ice. "Ap- 
parently the Missouri River Valley with its great abundance 
and variety of large game, wild fruit, and berries, bulbs, tubers, 
roots, and underground beans, was a paradise for these bears 
before the days of the rifle." Here, too, Audubon, in 1843, 


found them common, and in 1856 F. V. Hay den of the trans- 
continental surveys collected specimens. Bailey quotes an old 
resident that in 1867 there were many grizzlies in the river 
bottoms about Fort Buford and farther west in Montana. 
Old hunters told of killing grizzlies in the Killdeer Mountains 
between 1864 and 1870. In the late eighties they were still 
to be found in smaller numbers along the Little Missouri, but 
they were constantly hunted down, so that "at the present 
time there is certainly not a grizzly bear left in the State of 
North Dakota, and it is doubtful if there is anywhere a living 
representative of tnis original species of the grizzly group that 
was first given a scientific name and status in literature . . . 
Like some of the savage tribes with which it was associated, it 
has in passing left behind a thrilling record of savage bravery 
of surpassing interest to red-blooded Americans." 

In the middle of the last century grizzly bears were found in 
some numbers in Nebraska and Kansas. Baird (1857) lists 
specimens in the LTnited States National Museum from several 
localities in the former State, and Hibbard (1933) notes that 
in earlier years it was common in Kansas west of the Flint Hills. 
It has long been extinct in these States and probably in Texas 
as well, where, Vernon Bailey (1905) writes, "The only speci- 
men of grizzly bear that I have seen or heard of from Texas 
was killed in the Davis Mountains in October, 1890" and was 
found to agree in all essential characters of the skull with the 
so-called Sonoran grizzly. Merriam, however, made it the 
type of a special subspecies, Ursus horriaeus texensis, in 1914, 
later according it specific rank; he believed it identical with 
bears from the adjacent mountains of southern Colorado, 
a region where Gary (1911) found grizzlies rare in the wilder 
parts over 30 years ago, though occasionally seen in the San 
Juan, San Miguel, and La Plata Ranges. In northern Colo- 
rado, according to the best information he could obtain, in 
1905-6 it was already rare. Along the eastern slopes of the 
Front Range, however, where grizzlies were still found in the 
early seventies, they were by 1907 apparently quite gone. 
He gives a number of recent records of the grizzly in the 
southern mountains of Colorado up to 1907, and no doubt a 
few still remain in the more inaccessible mountainous regions 
of the central and western parts of the State, for a 1939 census 
issued by the U. S. Fish and Wildlife Service placed the 


estimated number of grizzlies in Colorado at only 10 indi- 

In recent decades grizzlies have become all but extinct in 
New Mexico and Arizona and probably in the adjacent parts of 
northern Mexico as well, where a century ago they were 
common. A skull collected about 1852 near Coppermines 
(now the site of Santa Rita, Grant County, N. Mex.) was 
regarded by Baird (1857) as representing a smaller brown race, 
which he named horriaeus, but Bailey (1931) says that speci- 
mens collected in that region in later years are not identical 
with it, so that an immigration of grizzlies into this area from 
elsewhere is assumed. He quotes the journals of Pattie who 
hunted in this region in 1824 and 1825 and wrote that it 
"abounds with these fierce and terrible animals." Bailey 
(1931) writes that in 1894 Dr. A. K. Fisher was told that 
grizzly bears were then common in the mountains about 80 
miles north of Silver City, N. Mex. In 1905 a large female was 
killed in the Tularosa Mountains and a large male was said to 
be still at large in that range. In 1908, there were said to be a 
few still in the Mimbres, Mogollon, and San Francisco Ranges. 
Since then the numbers seem to have decreased still further 
for in 1939 the U. S. Biological Survey estimated that only 
three grizzlies remained in New Mexico, so that the species 
soon no doubt will no longer exist there. Much the same story 
is probably true of Arizona, for which the same report fails to 
include any bears at all. It may be doubted, too, if any remain 
in the adjacent parts of Mexico, although Merriam lists speci- 
mens in the National collection as follows: Arizona, 1856, 1913; 
Chihuahua, 1899. A grizzly killed on Mount Taylor, northern 
New Mexico, in 1916, may well be among the last from that 

A century and more ago grizzly bears were common in Cali- 
fornia and extended a short distance farther south into the 
mountains of northern Lower California. From the latter 
region, however, it must soon have been exterminated, for Dr. 
E. W. Nelson (1921) states that the early account by Pattie 
of his journey in these regions gives the only Lower Californian 
record, namely, that there were grizzlies in the Sierra Juarez 
near Santa Catarina Mission. In their work on the "Fur- 
bearing Mammals of California," Grinnell, Dixon, and Lins- 
dale (1937) have given a very full account of the grizzly bears 


in that State. Dr. Merriam from a study of available skulls 
believes that seven specific types are represented amqng them. 
In the coastal counties, Humboldt and Mendocino, grizzlies 
were commoner in the river valleys where settlements were 
first made, so that on account of their depredations upon 
stock and the threat to the lives of children and others passing 
along roads, they were soon exterminated, within a short time 
after the first settlers arrived, about 1850. According to one 
informant the last grizzly taken in Humboldt County was 
killed in the Mattole section in 1868. The same person tells 
that his father in the early days had once counted 40 bears in 
sight at one time from a high point in the Mattole section 
that afforded a wide outlook over open country. In Mendocino 
County the authors quoted found on inquiry among old resi- 
dents that the last grizzlies there were killed in the fall of 1875, 
when an old female, a yearling, and a large male were shot. 
Various fragments concerning the killing of these bears or 
incidents of narrow escapes from them are brought together 
in the work mentioned, from which one may gather an impres- 
sion of the abundance of grizzlies in the third quarter of the 
last century, their boldness and occasional depredations, as 
well as the continual warfare waged against them by the 
settlers. As a result of constant hunting, bears became 
scarcer by the beginning of the last quarter of the century, 
but they still persisted in some localities. In the nineties there 
were still grizzlies "in the near ranges of the San Gabriel 
Mountains due north of Pasadena," even as late as 1897. 
"In the heavily chaparral-covered Santa Ana (or Trabuco) 
Mountains of Orange and extreme northwestern San Diego 
counties" they persisted "until well along in the 1900's," and 
the latest record of one being killed there is given as January 5, 
1908. One of the last grizzlies killed in the Yosemite region 
was in 1887. "The last known southern California grizzly" 
was killed October 28, 1916, near Sunland, Los Angeles County, 
while so far as the writers above quoted are able to ascertain, 
the last certain record of a grizzly killed in California was of 
one shot in Tulare County in August, 1922. A few later 
reports were un verifiable. 

In the State of Oregon these bears are perhaps already gone. 
Bailey (1936) has brought together many notes on their earlier 
abundance and gradual diminution. In the days of muzzle- 


loading rifles, grizzlies were bolder, and hunters might well 
hesitate to risk coming to close quarters with them. In the 
Lake of the Woods region the last grizzly is said to have been 
shot in 1868. In the last quarter century these bears have 
slowly diminished in numbers even in national forests. Bailey 
writes that in 1910 the supervisor of the Siskiyou National 
Forest reported grizzlies as very scarce; one was said to have 
been killed in the Yamsay Mountains about 1911, and there 
were possibly a few in the Mount Pitt section at about that 
time. In 1924 and 1925, the United States Forest Service 
reported one grizzly on each of the Cascade and Siskiyou 
National Forests, and in 1931 two and in 1932 one on the 
Wallowa Forest, while in 1933 it reported one on the Willam- 
ette. The last (1939) report of the U. S. Fish and Wildlife 
Service does not include it in a game census of the State ; in the 
State of Washington, however, it gives a surviving remnant 
of nine. 

At the present time the main population of grizzly bears in 
the United States outside of Alaska centers in the Rocky 
Mountain region in the States of Wyoming and Montana. 
This is partly a result of their protection from usual hunting 
in Yellowstone and Glacier National Parks, though others 
apparently frequent the wilder parts of these States. In the 
Yellowstone Park some grizzlies even come to the garbage 
piles near the hotel, but in Glacier Park, writes Vernon Bailey 
(1918), few have learned to do this, but most of them are shy 
and live in the less frequented portions of the park. A few 
are killed around the edges of the park annually. At times, 
too, the forest ranger in these parks finds it necessary to elimi- 
nate a particular grizzly that has developed troublesome ways. 
According to the 1939 census issued by the U. S. Fish and 
Wildlife Service (in December, 1940) the following estimates 
of the grizzly-bear population by States give at least an ap- 
proximation of its present status: Arizona, 7; Colorado, 10; 
Idaho, 44; Montana, 620; New Mexico, 3; Washington, 9; 
Wyoming, 480. There is thus an estimated total of approxi- 
mately 1,100 grizzlies in the United States, excluding Alaska. 

Northward of the United States, grizzly bears are not un- 
common in the Rocky Mountain region of British Columbia 
and Alaska. Cowan (1939) writes that they are now restricted 
to the mountainous parts of the country and are fairly com- 


mon in the mountains bordering the Pine River in British 
Columbia and in the Rockies north and south of the Peace 
River. Still farther north, grizzly bears are well known to 
occur in some numbers in the wilder parts of these regions to 
central Alaska. Preble (1908), writing of the Athabasca- 
Mackenzie area, says that they are found "throughout the 
Rocky Mountain range and its eastern spurs west of the Mac- 
kenzie, north to the Arctic .coast." Although annually some 
are killed, it seems likely they will long hold out here. A report 
of the U. S. Biological Survey in December 1938 states that 
"the large brown and grizzly bears in Alaska are holding their 
own as an outstanding wildlife resource of the Territory." 
In parts of Alaska they appear to be more plentiful than for 
many years, owing, it is believed, to the favorable attitude of 
Territorial residents, to protective regulations under the 
Alaska Game Law of 1925, and to sanctuary areas that total 
more than 8,500,000 acres. Some three-fourths of these 
sanctuary areas are comprised in Mount McKinley National 
Park and Katmai and Glacier Bay National Monuments. 
In addition, certain areas are now closed to hunting and else- 
where regulations afford a closed season from June 20 to 
September 1. The bag limit for large brown and grizzly bears 
is two a year, except on Admiralty Island, where a limit of one 
has been made, to induce nonresident hunters to visit other 
areas. The sale of bear hides is now prohibited in Alaska, a 
regulation considered as an outstanding factor in permitting 
the increase of these big bears, while of equal importance is 
the development of a better feeling by the residents, who now 
"thoroughly appreciate this source of income and have no 
desire to jeopardize it by killing off this attraction at no profit 
to themselves." Along the Arctic coast bears may still be 
regarded as a nuisance by certain interests, as miners and 
trappers, but the Survey's agent at Nome reported that in his 
district grizzlies are fairly numerous with no definite increase 
or decrease. The few animals that are killed are those slain 
by natives working with reindeer herds, and he believes their 
actual damage is slight. In these regions that may long serve 
as hunting areas it seems likely that these interesting and 
remarkable carnivores will continue indefinitely under wise 




Ursus richardsoni Swainson, Animals in Menageries, p. 54, 1838 (shore of the Arctic 
Ocean, on west side of Bathurst Inlet near mouth of Hood River, Mackenzie, 

FIGS.: Merriam, 1896, pi. 4, fig. 6; pi. 5, fig. 5; pi. 6, fig. 3 (skull). 

This grizzly is believed to range over the barren-grounds 
from west of Hudson Bay to the Mackenzie River, and north- 
ward to the shores of the Arctic Ocean. The color varies from 
yellowish to grizzly brown. Size rather small for a grizzly, 
and the skull distinguished by its broadly spreading zygomata 
and the sagittal crest, which runs far forward and divides 
dichotomously with a branch running to each postorbital 
process so far anteriorly that the two ridges are nearly trans- 
verse. Dr. Merriam (1918) regards the animal from the mouth 
of the Mackenzie as a related but distinct species, which he has 
called Ursus russelli, and in his paper he gives a series of 
cranial measurements of the two. 

Preble (1908) has summarized various bits of information 
concerning this little-known type of grizzly. Apparently the 
first white man to come in contact with it was Samuel Hearne, 
who in his famous journey (1770-1772) to the mouth of the 
Coppermine River saw a skin of an enormous grizzled bear 
at the Eskimo tents there. Franklin also observed it several 
times along the Arctic coast from near Fort Enterprise to 
Bathurst Inlet and mentions seeing a female with three cubs. 
The stomach of a bear killed at Bathurst Inlet contained the 
remains of a seal, a marmot, roots, berries, and grass. In 
another case a bear had been feeding on a caribou. Bell met 
with the species in 1900 "quite often" along the west and north 
shores of Great Bear Lake. 

As to its present status, Dr. R. M. Anderson (1939b) writes: 
"A few grizzlies of the Barren Ground group, a typical form 
being Ursus richardsoni, ranging eastward in the Arctic Zone 
as far as Bathurst Inlet, and perhaps a little beyond, have 
until the past few years escaped hunting by the Eskimos, who 
formerly kept off the bear's range in summer. As these inter- 
esting bears do not have the shelter of mountains, it is feared 
that with the recent spread of trapping operations and mining 
developments in the Far North, and particularly the extension 


of domestic reindeer herding into the Arctic portion of the 
Northwest Territories with the effect of patrolling the range 
throughout the year, some of these rare species will become 
extinct within a few years." 

The persistent reports of a grizzly bear existing in the barrens 
of northern Labrador have from time to time been investigated 
as far as such tales may, but no real evidence that such an 
animal is found there has been elicited. Even though the tale 
were true that a skin of a grizzly had once been brought in, 
this might have been traded from still farther west. In that 
way one might account for the origin of the reports. Dr. E. 
Stirling (1884) sums up the evidence in a brief statement to the 
effect that John McLean in his notes of 25 years' service in 
Hudson Bay Territory reports that skins from Ungava had 
several times been sent to Europe and that a factor named 
Mittleberger said he had known of the animal in Labrador. 



Ursus middendorffi Merriam, Proc. Biol. Soc. Washington, vol. 10, p. 69, Apr. 13, 1896 

("Kadiak Island, Alaska"). 
FIGS.: Merriam, 1896, pi. 4, figs. 2, 3; pi. 5, fig. 2; pi. 6, fig. 2 (skull); 1918, pi. 3 

(skull); Holzworth, 1930 (frontispiece in color and other photographic figures). 

This animal is rated by Merriam as the "largest of living 
bears, though only slightly larger than Ursus beringiana Mid- 
dendorff, from Kamschatka; frontal region in male enormously 
elevated, highly arched, and relatively narrow; zygomata 
bowed outward to an extraordinary degree ; postzygomatic part 
of skull very short." Distinctive features are the contact of 
the forepart of the jugal with the lachrymal bone and the 
small size of the first upper and last lower molars. The claws 
are shorter and more curved than in the usual type of grizzly, 
measuring about 100 mm. on the curve. 

The Kodiak bear is found on Kodiak Island and the adjacent 
Afognak and Shuyak Islands, but it may be taken as a type 
of the so-called big brown bears, of which closely allied forms 
are found on the nearby mainland, such as Ursus gyas from 
Cook Inlet to the entire length of the Alaska Peninsula. 
Merriam (1918) writes that "the differences between the 
grizzlies on the one hand and the big brown bears on the other 


are neither so great nor so constant as at one time believed. 
And there are species which in the present state of knowledge 
can not be positively referred to either group." The color of 
the brown bears is more uniform with less of the surface 
grizzling, the shorter claws are scurfy instead of smooth, the 
skull more massive. The brown bears are in general confined 
to the southern Alaskan coast and peninsula and the adjacent 
islands. (See Dufresne, 1942, for distribution map.) 

In 1912 Sheldon wrote that "the grand bear of Kadiak 
Island ... is rarely seen." Osgood (1904), writing of 
the big brown bears of the base of the Alaska Peninsula, says 
that though formerly abundant persistent hunting by the 
natives has much reduced their numbers. They still occur in 
many localities but have become extremely shy. Fifteen 
years previously it was not uncommon to see "from eight to 
fifteen bears scattered about on one mountain side." Their 
period of activity varies, but they usually come out from hiber- 
nation late in March or early in April, depending on the season, 
and are not seen after early November. The young are brought 
forth usually in January and may number as many as four at 
a birth, though, as with the other grizzlies, two is the usual 
number and three not uncommon. Ground squirrels are much 
sought after by digging up their burrows, but when salmon are 
entering the rivers in spring they become a main object of 
pursuit. Small fruits, roots, and grasses also form part of 
their fare. 

Although these bears a few years ago seemed in much danger 
of extirpation, present reports indicate that under the game 
laws that went into effect in 1925, and with the creation of 
sanctuaries, the outlook is bright that they will remain an 
asset to Alaska for a long while to come. Dufresne, the execu- 
tive officer of the Alaska Game Commission, reported in 1938 
that they are holding their own and that "information from 
Kodiak Island and the Alaska peninsula . . . leaves no 
question as to the plenitude of the huge brown bears." A 
favorable attitude on the part of Alaskan residents, who are 
beginning to realize the value of the bears to themselves 
through bringing in a certain number of hunters and photogra- 
phers, is doing much to insure proper protection and regulation 
of the numbers killed. For the present it appears that these 
splendid bears are no longer in danger of being exterminated. 


In order to coordinate the account of the grizzly and brown 
bears with the nomenclature proposed by Merriam (1918) in 
his review of these animals, the following synopsis of the names 
used in his paper and their grouping is added, with a few words 
on the status of these forms where information is at hand. 


Ursus horribilis horribilis Ord. BIG PLAINS GRIZZLY 

Range: Plains of Montana, Wyoming, Colorado, Dakotas. 
Nearly or quite extinct. 

Ursus horribilis bairdi Merriam. BAIRD'S GRIZZLY 

Range: "Southern Rocky Mountain region from San Juan 
Mountains, southwestern Colorado, northward through Wyo- 
ming to Montana, and perhaps to southeastern British Colum- 
bia." Supposed to be a mountain form; nearly extinct. 

Ursus horribilis imperator Merriam. YELLOWSTONE PARK 


Range: "Yellowstone National Park, Wyoming; limits un- 
known." A small number live in the Park and perhaps the 
nearby country. 

Ursus chelidonias Merriam. JERVIS INLET GRIZZLY 

Range: "Head of Jervis Inlet, British Columbia." Status 

Ursus atnarko Merriam. ATNARKO GRIZZLY 

Range : Region of Atnarko River, upper Bella Coola, British 
Columbia. Status unknown. 

Ursus kwakiutl Merriam. KWAKIUTL GRIZZLY 

Range: "Coast region of British Columbia from south- 
western corner (Burrard Inlet, Howe Sound, Jervis Inlet) 
northwesterly to or beyond the lower Bella Coola." Probably 
a few left. 

Ursus nortoni Merriam. YAKUT AT GRIZZLY 

Range: "Limited apparently to coastal plain on southeastern 
side of Yakutat Bay." Probably still found; specimens col- 
lected in 1910. 

Ursus warburtoni Merriam. WARBURTON PIKE GRIZZLY 
Range :" Coast region , . . of southeastern Alaska and 


adjacent parts of British Columbia from Chilkat River south- 
easterly to Atnarko River." Probably still occurs; specimen 
collected 1915. 

Ursus neglectus Merriam. ADMIRALTY ISLAND GRIZZLY 

Range: Admiralty Island. Restricted but probably not in 
immediate danger. 

Ursus californicus Merriam. CALIFORNIA COAST GRIZZLY 

Range: "Humid coast region of California from San Fran- 
cisco Bay south about to San Luis Obispo." Probably extinct 
since 1908. 

Ursus tularensis Merriam. TEJON GRIZZLY 

Range: "Dry chaparral hills of interior coast ranges between 
the San Joaquin Valley and Los Angeles plain," California. 
Extinct in 1916. 


Range: "Sacramento (and perhaps also San Joaquin) Valley 
and adjacent foothills; westerly ... to Dobbins Creek 
canyon on the boundary between southeastern Humboldt and 
southwestern Trinity Counties." Extinct. 

Ursus dusorgus Merriam. RINDSFOOS'S GRIZZLY 

Range: "Head of Jack Pine River near Mount Bess, Al- 
berta." Status unknown; type collected in 1916. 


Ursus nelsoni Merriam. NELSON'S GRIZZLY 

Range: "Sierra Madre of Mexico from northwestern Chi- 
huahua and northeastern Sonora south to southern Durango." 
Probably a few remain. 

Ursus texensis texensis Merriam. TEXAS GRIZZLY 

Range: Davis Mountains, Texas, and mountains of southern 
Colorado. Probably nearly or quite extinct. 

Ursus texensis navaho Merriam. NAVAHO GRIZZLY 

Range: "Probably restricted to the isolated Chuska Moun- 
tains" between northeastern Arizona and northwestern New 
Mexico. Extinct. 


Ursus planiceps Merriam. FLAT-HEADED GRIZZLY 

Range: Foothills along the western edge of the plains in 
Colorado and Wyoming. Nearly if not quite extinct. 

Ursus macrodon Merriam. TWIN LAKES GRIZZLY 

Range: Twin Lakes region of Colorado. Probably nearly 


Range : Yellowstone Park region. Probably now uncommon ; 
specimen (the type) killed in 1915. 

Ursus eltonclarki Merriam. SITKA GRIZZLY 

Range: "The Sitka Islands, Baranof and Chichagof," 
Alaska. Probably still occurs. 

Ursus tahltanicus Merriam. TAHLTAN GRIZZLY 

Range: "Middle and upper Stikine-Skeena region, limits 
uncertain/' in British Columbia. Probably still present in 
small numbers. 

Ursus insular is Merriam. ISLAND BEAR 

Range: Admiralty Island, Alaska. Probably not now in 
immediate danger. 

Ursus orgilos Merriam. GLACIER BAY GRIZZLY 

Range: Bartlett Bay and east side of Glacier Bay, Alaska. 
Probably still occurs; collected by Hasselborg in 1912. 

Ursus orgiloides Merriam. ALSEK GRIZZLY 

Range: Italic River region, Alaska; in coast strip southeast 
of Yakutat Bay. Probably present in small numbers; type 
collected in 1916. 

Ursus pallasi Merriam. PALLAS'S GRIZZLY 

Range: "Southwest corner of Yukon Territory, east of the 
St. Elias Range . . . and adjacent eastern border of 
Alaska; easterly to McConnell River and Teslin Lake and 
south into northern British Columbia." Probably not in 
immediate danger. 

Ursus rungiusi rungiusi Merriam. RUNGIUS'S GRIZZLY 

Range: Region about the head of Athabaska River and 
Kootenay Pass, Alberta, and adjacent parts of British Colum- 
bia (Indian Point Creek). Probably still in small numbers. 


Ursus rungiusi sagittalis Merriam. CRESTED GRIZZLY 

Range : Southwestern Yukon, Canada. Probably still occurs 
in small numbers and is in no immediate danger. Type col- 
lected in 1915. 

Ursus macfarlani Merriam. MACFARLANE'S BEAR 

Range: Region of Anderson River, Mackenzie; limits un- 
known. Specimens from Franklin Bay, Stapylton Bay. 
Status unknown; probably still to be found in small numbers. 

Ursus canadensis Merriam. CANADA GRIZZLY 

Range: "Eastern British Columbia; limits unknown (type 
from near Mount Robson; and an adult female from Kootenay 
Lake)." Not known to be in danger. 


Ursus arizonae Merriam. ARIZONA GRIZZLY 

Range: Known from Escudilla Mountains, Apache County, 
Arizona. Status unknown, perhaps extinct. 

Ursus idahoensis Merriam. IDAHO GRIZZLY 

Range: Eastern Idaho (North -Fork Teton River) to Wallowa 
Mountains, Oreg. Probably still present in small numbers. 

Ursus pulchellus pulchellus Merriam. UPPER YUKON GRIZZLY 
Range: Ross River, Ross Mountains and McConnell River, 
Yukon, Canada. Probably still present in small numbers; 
type collected in 1916. 

Ursus pulchellus ereunetes Merriam. KOOTENAY GRIZZLY 

Range: Kootenay District, British Columbia. Status un- 
known; collected in 1916. 

Ursus oribasus Merriam. LIARD RIVER GRIZZLY 

Range: Upper Liard River, Yukon, near British Columbia 
boundary. Collected in 1916; probably present in small 

Ursus chelan Merriam. CHELAN GRIZZLY 

Range: "Cascade and Cassiar Mountains from northern 
Washington to upper Stikine River and Dease Lake, British 
Columbia." Nearly extinct. 

Ursus shoshone Merriam. SHOSHONE GRIZZLY 

Range: "Mountains of Colorado and Wyoming." Probably 
very few left in Colorado but more in Wyoming. 


Ursus kennerlyi Merriam. SONORA GRIZZLY 

Range: "Nothing is known of the range of kennerlyi except 
that the type specimen came from mountains near Nogales, 
Sonora," Mexico. "Its affinities with utahensis suggest that 
formerly it may have had a disconnected distribution north- 
ward in the mountains of central Arizona." Extinct. Type 
skull dated 1855. 

Ursus utahensis Merriam. UTAH GRIZZLY 

Range: "Southern Wasatch and Pine Valley Mountains 
[Utah]; limits unknown." Must be now nearly gone, though 
specimens have been collected as lately as 1911. 

Ursus perturbans Merriam. MOUNT TAYLOR GRIZZLY 

Range: Mount Taylor and Datil Mountains, New Mexico; 
limits unknown. Now practically extinct. 

Ursus rogersi rogersi Merriam. ROGERS'S GRIZZLY 

Range: Absaroka Mountains, Wyoming; limits unknown. 
The considerable grizzly-bear population of Wyoming must 
include a number of this form. 

Ursus rogersi bisonophagus Merriam. BLACK HILLS GRIZZLY 
Range: "Black Hills of South Dakota and adjacent north- 
east corner of Wyoming." Probably extinct; type collected in 

Ursus pervagor Merriam. LILLOOET GRIZZLY 

Range: "Interior of southwestern British Columbia; known 
only from Lillooet Lake and Bridge River." Probably extinct; 
type taken in 1883. 

Ursus caurinus Merriam. LYNN CANAL GRIZZLY 

Range: "Coast of mainland of southeastern Alaska from 
Chilkat River Valley and Lynn Canal south an unknown 
distance." Probably rare; type collected in 1911. 

Range: Admiralty Island, Alaska. Probably not now in 
immediate danger. 

Ursus klamathensis Merriam. KLAMATH GRIZZLY 

Range: "Siskiyou Mountains of northern California and 
southern Oregon, ranging north in recent times to Fort Kla- 
math region and Rogue River valley; in early days to lower 


Willamette Valley (presumably same species); south in Sierra 
Nevada an unknown distance." Now about extinct; last ones 
killed perhaps about 1911. Bailey (1936) states that according 
to Forest Service reports, there were in 1924 and 1925 one 
grizzly bear on each of the Cascade and Siskiyou National 
Forests, in 1931 two and in 1932 one on the Wallowa Forest, 
and in 1933 one on the Willamette. 

Ursus mendocinensis Merriam. MENDOCINO GRIZZLY 

Range: Known only from Mendocino County, California, 
along the northwest coast of the State. Extinct. The last 
grizzlies were killed in the county in the autumn of 1875, a 
female, a yearling, and a large male, all killed at the same 
place on Eel River south of Covelo (Grinnell, Dixon, and 
Linsdale, 1937, p. 70). 


Range: Santa Ana or Trabuco Mountains, Cuyamaca and 
Santa Rosa Mountains, and probably San Jacinto Mountains 
of California. Now extinct; last one killed October 28, 1916, 
near Sunland, Los Angeles County, California (Grinnell, 
Dixon, and Linsdale, 1937). 


Ursus hylodromus Elliot. FOREST GRIZZLY 

Range: "Rocky Mountain region of western Alberta and 
eastern British Columbia, including Selkirk Range." Prob- 
ably a few still remain. 

Ursus kluane kluane Merriam. KLUANE GRIZZLY 

Range: "Southwest corner of Yukon Territory east of the 
St. Elias Range, extending northwesterly in Alaska to Mount 
McKinley region (head of Toklat), easterly in Yukon Territory 
to McConnell River . . . and probably south into north- 
west corner of British Columbia" (Merriam). Not believed 
to be in present danger. 

Ursus kluane impiger Merriam. INDUSTRIOUS GRIZZLY 

Range: Merriam gives the following localities as indicative 
of the range: Brisco, Columbia Valley, in British Columbia; 
Morley and Jasper, Alberta; and headwaters of North Fork 
Blackfoot River, western Montana. Hollister (1912a) wrote 
that "the grizzly bear is still a common animal throughout the 


Robson region" of Alberta, a statement which doubtless still 
applies to this animal. 

Ursus pellyensis Merriam. PELLY GRIZZLY 

Range: Pelly Mountains, Yukon. Grizzlies are still found 
in small numbers in the Yukon region, but since it is not 
possible to identify animals without killing them and examin- 
ing their skulls, it is unknown whether this form is represented 
or not. 

Ursus andersoni Merriam. ANDERSON'S GRIZZLY 

Range: The type specimen is from Dease River, near Great 
Bear Lake, Mackenzie; limits of range unknown. Grizzly 
bears, according to Treble's (1908) summary, are not uncom- 
mon in the region. 


Ursus apache Merriam. APACHE GRIZZLY 

Range : White Mountains of eastern Arizona. The type was 
collected in 1913, but at the present time the predatory- 
animal campaigns have resulted in reducing the numbers to so 
low an ebb that already this form may have been extirpated. 

Ursus horriaeus Baird. NEW MEXICO GRIZZLY 

Range: "Parts of New Mexico, south to Casas Grandes, 
Chihuahua, Mexico; probably extending into eastern Arizona" 
(Merriam). Probably now extinct. 

Ursus henshawi Merriam. HENSHAW'S GRIZZLY 

Range: "Lower slopes of southern part of Sierra Nevada; 
limits unknown" (Merriam). The type specimen was from 
near Havilah, Kern County, California, and Grinnell, Dixon, 
and Linsdale (1937) mention as representing probably the 
same sort of animal, skins from Madera County. They pro- 
visionally refer to this form the grizzly population of the 
Yosemite region and the central and northern Sierra Nevada 
generally. In the Yosemite region, according to Grinnell and 
Storer (1924) the grizzly bear was rare already by 1887, when 
the capture of one was considered "an unusual event." The 
last one known to have been killed in the Yosemite Park was 
shot "'about 1895' at Crescent Lake." Possibly a few indi- 
viduals persisted longer, for the authors mentioned tell of one 
whose tracks were frequently seen on Bullion Mountain 


between the years 1908 until 1911. Undoubtedly this form 
of bear is now extinct. 


Ursus stikeenensis Merriam. STIKINE GRIZZLY 

Range : Region about head of Finlay River and Dease Lake 
region, northern British Columbia (Merriam). Grizzlies in 
this region seem in no immediate danger. 

Ursus crassodon Merriam. BIG-TOOTH GRIZZLY 

Range: Merriam refers to this form, skulls from Klappan 
Creek (third south fork of Stikine River) ; White River, Yukon; 
Wolf Lake, Yukon; Tatletuey Lake, upper Finlay River; Glen- 
lyon Mountains and Quiet Lake, southern Yukon. Since other 
supposed forms of grizzlies also inhabit this general region, 
they may only be identified by killing them and studying the 
skull. At present the grizzly bear is in no danger in this region, 
for no cattle are raised, human population is small, and the 
occasional hunter is their only enemy. 

Ursus crassus Merriam. THICK-SET GRIZZLY 

Range: Merriam records skulls from upper Macniillan 
River, Yukon; and Anderson River, Horton River, and Lang- 
ton Bay, on the Barren-grounds. The type specimen was 
collected in 1916. No immediate danger for it is seen. 

Ursus mirabilis Merriam. STRANGE GRIZZLY 

Range: Admiralty Island, Alaska. With the present re- 
ported favorable attitude of the local population and the 
recent game regulations, it would seem that this grizzly will 
not be hunted beyond a reasonable limit. 

Ursus absarokus Merriam. ABSAROKA GRIZZLY 

Range: "Laramie and Bighorn Mountains, eastern Wyo- 
ming, Black Hills region, South Dakota, and northward along 
Little Missouri to Missouri and Yellowstone Rivers." Prob- 
ably extinct in the Black Hills and eastern Wyoming. The 
latter State still has a relatively large population of grizzly 
bears but these center in the Yellowstone Park region, and are 
regarded by Merriam as a special type. The type specimen 
from near head of Little Bighorn River, northern part of Big- 
horn Mountains, Montana, was killed in 1893, and the animal 
may still occur in that region. 



Ursus alascensis Merriam. ALASKA GRIZZLY 

Range: Norton Sound region, Alaska, south to Chinitna, on 
Cook Inlet; limits unknown. Osgood (1901, p. 68) at the 
beginning of this century wrote that "large bears are still very 
often seen both on the Alaska Peninsula side of Cook Inlet and 
on the mountainous Kenai Peninsula. According to report 
they were very abundant about ten years ago, but in the short 
time since have been so constantly pursued that their numbers 
have been greatly reduced." Probably this local form requires 
protection if it is to survive, but no recent report is at hand. 

Ursus toklat Merriam. TOKLAT GRIZZLY 

Range: "So far as known, restricted to Alaska Range" 
(Merriam). The type specimen came from the head of 
Toklat River, north base of Alaska Range, near Mount Mc- 
Kinley, Alaska. Since this region is now a national part, 
there seems to be no immediate cause for anxiety for the 
continued existence of these bears in their range. 

Ursus latifrons Merriam. BROAD-FRONTED GRIZZLY 

Range: Rocky Mountains of western Alberta and eastern 
British Columbia from Jasper House northwesterly to head- 
waters of Stikine River. Probably the same remarks apply to 
the status of this bear as to that of Ursus crassodon. Some 
numbers may still occur. 


Ursus richardsoni Swainson. BARREN-GROUND BEAR 

Range: The type came from the shore of the Arctic Ocean 
on the west side of Bathurst Inlet, near the mouth of Hood 
River, and probably is representative of the animal of northern 
and northeastern Mackenzie. Anderson notes that of late 
years with the extension of reindeer herding by the natives, 
these regions, which previously were but little hunted, may 
now be more often patrolled and these bears are likely to be 
in a much more precarious situation than formerly. 

Ursus russelli Merriam. MACKENZIE DELTA GRIZZLY 

Range: Lower Mackenzie region; limits unknown. Of the 
barren-ground bear in general, Preble (1908) states that it is 
rare over the greater part of its range. Probably it is not 


likely to persist about the mouth of the Mackenzie for any 
great length of time. 

Ursus phaeonyx Merriam. TANANA GRIZZLY 

Range: "Tanana Mountains between Tanana and Yukon 
Rivers," Alaska. Dr. W. H. Osgood, who obtained the type 
specimen in 1903, wrote that "very little accurate information 
is obtainable in regard to the grizzly in the Yukon region," 
and this state of affairs seems to continue. However, with the 
development of mining interests and settlement, it is not likely 
that this form of the middle Yukon will resist encroachment 
for many years. 

Ursus internationalis Merriam. ALASKA BOUNDARY GRIZZLY 

Range: "Region bordering Arctic coast along international 
boundary, and doubtless adjacent mountains, between the 
coast and the Yukon-Porcupine; limits unknown." Status 
not determined. 

Ursus ophrus Merriam. HIGH-BROWED GRIZZLY 

Range: The type, collected in> 1915, came from an unknown 
locality in eastern British Columbia. Nothing further is 
known of its status. 

Ursus washake Merriam. WASHAKIE GRIZZLY 

Range : The type came from the north fork of the Shoshone 
River, Absaroka Mountains, in western Wyoming, between 
Bighorn Basin and Yellowstone National Park. Since the 
characters distinguishing the form are to be found in the skull 
and teeth it is not possible to say what portion of the many 
grizzlies found living in western Wyoming are like it. 


Ursus kidderi kidderi Merriam. KJDDER'S BEAR 

Range: "Alaska peninsula for its entire length." According 
to the report on Alaska bear sanctuaries issued by the U. S. 
Department of Agriculture in December 1939, "information 
from . . . the Alaska Peninsula . . . leaves no question 
as to the plenitude of the huge brown bears," so that with the 
continuance of protective legislation no fears are felt for them. 


Ursus kidderi tundrensis Merriam. TUNDRA BEAR 

Range: "Tundra region of northwestern Alaska from Shak- 
tolik River on Norton Sound, southerly across the lower 
Yukon, Kuskokwim, and Nushagak Rivers to Bristol Bay and 
north side of base of Alaska Peninsula" (Merriam). The 
Government report above quoted states that "the grizzlies of 
the interior seemed to be on the increase during the last year" 
(1938). The agent at Nome reports that "the Alaska grizzly 
is fairly numerous all through my district, and no increase or 
decrease is definitely known. Very few animals are killed and 
these by natives working with reindeer herds. Bears are 
reported by all reindeer men as destructive to deer, but I 
believe that the extent of damage is not important." 

Ursus eximius Merriam. KNIK BEAR 

Range : Nothing seems to be known of the range of this bear 
beyond its occurrence at the head of Knik Arm, Cook Inlet, 
Alaska. Probably, like Kidder's bear of the Alaska Peninsula, 
it is "holding its own." 


Ursus innuitus Merriam. INNUIT BEAR 

Range: "Coastal region of Norton Sound, Alaska, from 
Unalaklik northward and westward; limits unknown" (Mer- 
riam). The range of the grizzly bear probably does not reach 
the sea to the northward of the region inhabited by this form. 
The report of the Nome agent, previously quoted, seems to 
indicate that grizzly bears are in no immediate danger of 
extermination in this area. 

Ursus cressonus Merriam. CHITINA BEAR 

Range: "Chitina River valley and adjacent slopes of Skolai 
and Wrangell Mountains, westerly doubtless through the 
Chugach Mountains to the west side of Cook Inlet; occurs as 
far south as the Iliamna region" (Merriam). This local type 
may be thought of as present in small numbers, but exact 
information is not at hand. 

Ursus alexandrae Merriam. Miss ALEXANDER'S GRIZZLY 

Range: Kenai Peninsula, Alaska. This is notable as the 
largest of the grizzly bears. No recent information of a definite 
nature is at hand as to its present numbers. 



Ursus townsendi Merriam. TOWNSEND'S BEAR 

Range: Mainland of southeastern Alaska. The type came 
from Sitka in 1889. No information as to status is at hand. 


Ursus dalli Merriam. D ALL'S BROWN BEAR 

Range : Malaspina Glacier and region northwest of Yakutat 
Bay, Alaska. Osgood (1904) writes of the general region about 
the base of the Alaska Peninsula that "Brown Bears were 
formerly abundant in much of the country . . . but the 
persistent hunting by the natives since the introduction of 
modern repeating rifles has reduced their numbers greatly. 
They still occur in many localities, but have become extremely 
shy and are seldom obtained unless a special campaign for 
them is conducted." 

Ursus hoots Merriam. STIKINE BROWN BEAR 

Range: Stikine River region, British Columbia. This is 
believed to be a small race of tj^e brown bear, perhaps closely 
related to the form sitkensis. Merriam mentions but three 
specimens from the Stikine River. Its present status is un- 
known but is perhaps little likely to have altered much recently. 

Ursus sitkensis Merriam. SITKA BROWN BEAR 

Range: Sitka Islands (Baranof and Chichagof), Alaska. In 
the press notice of the U. S. Department of Agriculture for 
December 18, 1938, Wildlife Agent Douglas Gray of Juneau 
reports that the large brown bear continues to be "the greatest 
attraction to the trophy hunter that this district offers. Their 
numbers seem to be as plentiful as ever, notably on the Baranof- 
Chichagof-Admiralty Island group." With proper regulation 
of hunting, therefore, this form seems in no danger. 

Ursus shirasi Merriam. SHIRAS'S BROWN BEAR 

Range: Restricted to Admiralty Island. This is "a very 
large member of the brown bear group," but whether it is 
always black like the type specimen is unknown. Its "excep- 
tionally broad skull with broad short rostrum, excessively 
broad and short frontal shield, and huge massive postorbital 
processes," though obviously different in comparison with the 
"long narrow skull with slender elongate rostrum, long and 


narrow frontal shield, and insignificant postorbital processes" 
of the form eulophus living on the same island, may not be as 
distinctive in life. Hence its status on Admiralty Island is 
not made out. With the closing of two areas totaling 52,000 
acres on Admiralty Island, it is presumed that the brown 
bears present in some numbers there must include a proportion 
of this type. 

Ursus nuchek Merriam. NUCHEK BROWN BEAR 

Range: Prince William Sound easterly to Mount St. Elias; 
limits unknown. The range continues that of the Yakutat 
bear on the Alaskan coast. No late information is at hand as 
to its status there, but under present regulations the existing 
conditions are at least favorable. 


Ursus gyas Merriam. PENINSULA GIANT BEAR 

Range: "Entire length of Alaska peninsula from Cook Inlet 
to Isanotski Strait and adjacent Unimak Island." This is 
"either the largest living bear or second only to the great 
Kadiak bear (middendorffi)," according to Merriam. Skulls 
of adult males show "a surprisingly wide range in size and 
form," so that Merriam distinguishes four types among them. 
On account of its large size this bear might form an especial 
attraction to hunters and thus be in more danger from this 


Barren-ground bear (Ursus richardsoni) 


source than other "species" having the same range. Present 
information from the Alaska Peninsula, issued by the U. S. 
Department of Agriculture, "leaves no question as to the 
plenitude of the huge brown bears." 

Ursus middendorffi, Merriam. KODIAK BEAR 

Range : Confined to Kodiak and the adjacent islands Afognak 
and Shuyak. This bear shares with gyas the distinction of 
being the largest of living bears. The two are closely allied. 
Present information indicates that its numbers are being 


Ursus kenaiensis Merriam. KENAI GIANT BEAR 

Range: Kenai Peninsula, Alaska. No recent statistics are 
at hand. With the passage of the Alaska game law in 1925, 
however, the conditions are probably as favorable as may be 
expected. According to the 1938 press information of the U. S. 
Department of Agriculture, "there has been a perceptible 
change for the better in the brown and grizzly bear population." 

Ursus sheldoni Merriam. MONTAGUE ISLAND BEAR 

Range: Believed to be confined to Montague Island, Prince 
William Sound, Alaska. The name is based on specimens of 
the brown bear type taken in 1905. No recent information as 
to the bear of this island is at hand. 

Vetularctos inopinatus Merriam. PATRIARCHAL BEAR 

This genus and species were based on a skull from Rendez- 
vous Lake, northeast of Fort Anderson, Mackenzie, but the 
genus is currently regarded as inseparable from Ursus, while 
the species is not further known beyond the original specimen 
collected in 1864. Its peculiarities would ordinarily be as- 
cribed to individual variation. 

Family MUSTELIDAE: Weasels and Martens 

Among the northern fur bearers the martens, because of 
their soft, handsome fur, are much sought after, and their 
pelts bring high prices in the markets. They are characteristic 
of the evergreen forest belt of the northern United States 
northward into Canada and extend their range southward 


along the higher parts of the Alleghenies in the East and the 
Rocky and Sierra Nevada Mountains of western North 
America. Like many carnivorous animals they occur in small 
numbers relative to the abundance of the rodents and other 
animals on which they prey and are given to traveling over 
rather wide circuits in their search for food. 

In general appearance pine martens are about the size of a 
large slender squirrel, with a rather short cylindrical and well- 
furred tail, short legs, and erect, rather prominent ears. 
Males are larger than females and when adult develop more 
prominent median crests on the skull. The general coloring is 
yellowish to light brown on the head and forepart of the body, 
darkening to a warm brown posteriorly and on the tail and 
legs. The under side is cinnamon, darkening posteriorly, and 
often with a tinge of brighter ochraceous-orange on the chest. 
Adult males weigh about 2.5 pounds, females about a quarter 
less. Total length, about 25 inches, of which the tail is 7 or 8 

Over their wide range across the northern part of the conti- 
nent pine martens show a slight degree of geographic variation, 
both in shades of color and in details of the skull and teeth. 
These have been made a basis for recognizing at least two 
species, the typical eastern marten and the darker\animal of 
the Northwest coast, each with sundry races. While at; present 
it seems still uncertain whether only a single species is Actually 
represented, with more or less well-marked subspecies, these 
may be listed with their type localities in accordance with 
current usage, as follows : 

Maries americana americana (Turton). EASTERN PINE MARTEN 

Mustela americanus Turton, Turton's ed. of Linnaeus's Systema Naturae, vol. 1, 
p. 60, 1806 (eastern North America). 

Maries americana abieticola (Preble). KEEWATIN MARTEN 

Mustela americana abieticola Preble, North Amer. Fauna, no. 22, p. 68, Oct. 31, 
1902 (Cumberland House, Saskatchewan, Canada). 

Maries americana abietinoides Gray. NORTHWESTERN MARTEN 

Maries americana var. abietinoides Gray, Proc. Zool. Soc. London, 1865, p. 106 
(Rocky Mountains between Kicking Horse Pass and Columbia River). 

Maries americana actuosa (Osgood). ALASKAN PINE MARTEN 

Mustela americana actuosa Osgood, North Amer. Fauna, no. 19, p. 43, 1900 (Fort 
Yukon, Alaska). 

Maries americana atrata (Bangs). NEWFOUNDLAND MARTEN 

Mustela atrata Bangs, Amer. Nat., vol. 31, p. 162, Feb., 1897 (Bay St. George, 


Maries americana boria (Elliot). MACKENZIE MARTEN 

Mustela boria Elliot, Proc. Biol. Soc. Washington, vol. 18, p. 139, Apr. 18, 1905 
(Lower Mackenzie River district; probably same as M. a. actuosd). 

Maries americana brumalis (Bangs). LABRADOR MARTEN 

Mustela brumalis Bangs, Amer. Nat., vol. 32, p. 502, July, 1898 (Okak, Labrador) 

Maries caurina caurina (Merriam). PACIFIC MARTEN 

Mustela caurina Merriam, North Amer. Fauna, no. 4, p. 27, Oct. 8, 1890 (near 
Grays Harbor, Chehalis County, Washington). 

Maries caurina humboldtensis Grinnell and Dixon. HUMBOLDT MAR- 

Maries caurina humboldtensis Grinnell and Dixon, Univ. California Publ. Zool., 
vol. 21, p. 411, Mar. 17, 1926 (5 miles northeast of Cuddeback = Carlotta, Hum- 
boldt County, California). 

Maries caurina origenes (Rhoads). ROCKY MOUNTAIN MARTEN 

Mustela caurina origenes Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1902, p. 
458, Sept. 30 (Marvine Mountain, Garfield County, Colorado). 

Maries caurina nesophila (Osgood). QUEEN CHARLOTTE ISLANDS 

Mustela nesophila Osgood, North Amer. Fauna, no. 21, p. 33, Sept. 26, 1901 
(Massett, Graham Island, Queen Charlotte Islands, British Columbia). 
Maries caurina sierrae Grinnell and Storer. SIERRA MARTEN 

Maries caurina sierrae Grinnell and Storer, Univ. California Publ. Zool., vol. 17, 
p. 2, Aug. 23, 1916 (Lyell Canyon, Yosimite National Park, California). 
Maries caurina vancouverensis Grinnell and Dixon. VANCOUVER 


Maries caurina vancouverensis Grinnell and Dixon, Univ. California Publ. Zool., 
vol. 21, p. 414, Mar. 17, 1926 (20 miles south of Alberni, Vancouver Island, 
British Columbia). 

FIGS.: Grinnell, Dixon, and Linsdale, 1937, vol. 2, pi. 4 (col.), of M. c. sierrae; Nelson, 
1918, p. 453, upper fig. (col.), of M. americana; Elliot, 1901, p. 334, fig. 66 (skull 
of M. americana); Elliott, C., 1942, col. pi. facing p. 75, of M. americana. 

A brief survey of the status of the pine marten shows that 
although once common in Nova Scotia, New Brunswick, the 
northern New England States, New York, Pennsylvania, and 
even New Jersey, it is now through much trapping greatly 
reduced, though in places where there are large tracts of 
forested country small numbers may remain. In New Bruns- 
wick, Chamberlain reported it as common 60 years ago, and 
probably it is still to be found in wilder areas. Anderson 
(1939a) writes that it was fairly common in the wooded parts 
of Quebec in earlier days but has been overtrapped and is 
"now rare, found only in areas remote from settlement." Of 
the large dark Labrador race no recent statistics are at hand, 
but there are many skulls in the Museum of Comparative 


Zoology secured from trappers by J. D. Sornborger about 1900. 
The Labrador wilderness is not much penetrated by outside 
trappers, and the supply of fur is likely to hold out for a long 
time to come. The Newfoundland race, however, is said now 
to be very much reduced. In northern New England martens 
were formerly much trapped in the Maine Woods and the 
White Mountains and are still found but in smaller numbers. 
A century ago Emmons (1840) wrote that the pine marten was 
to be found in the mountainous parts of the Berkshires, in 
Massachusetts, especially in beech woods, and describes the 
method of trapping it in deadfalls. At that time, he states, 
the pelts were worth 90 cents to $1.1^. Probably martens 
have been extinct in Massachusetts for many years, but they 
still are found in the Vermont forests. Merriam regarded them 
as common in the Adirondacks in 1882 and said that hundreds 
were trapped for fur every winter; they still persist in smaller 
numbers. The fur is prime early in November. Rhoads, 
writing in 1903, makes the general statement that the marten 
was "once abundant over all the mountain regions" of Penn- 
sylvania and New Jersey, invading even the Nfoothills of the 
Alleghenies, but was early exterminated fro^the latter 
regions, was probably extinct in New Jersey aboutl&>0, and 
at the beginning of this century was to be found only in the 
forested parts of the mountains in Pennsylvania. South of 
this area there seems to be no recent report of the marten's 
presence. Rhoads (1903) in his account of the species in Penn- 
sylvania states that there it was partial to hardwood forests 
rather than evergreens. Reports from various of the counties 
are nearly unanimous in telling of its former occurrence and its 
later decline, and in most cases tell of none having been found 
for a number of years. Rhoads points to the effect of forest 
fires as bearing on the depletion of the stock. In Ohio, Brayton 
(1882) mentions the marten as already extinct, and there is 
apparently no evidence of its occurrence in Indiana within 
recent years. Dr. H. H. T. Jackson (1908) regarded it as 
practically extinct in Wisconsin 30 years ago and mentions one 
taken in Vilas County as lately as 1904-5, but Cory (1912) 
wrote that although "steadily declining in numbers" and then 
"a comparatively rare animal" in Wisconsin it was still to be 
found in the northern part of that State. It seems to be gone 
from Illinois, though earlier recorded from Cook County by 


Kennicott. Northern Minnesota was once good marten 
country, and Herrick (1892) mentions that many dark pelts, 
especially prized, formerly came from this area. As to its 
recent status along the northern border of Minnesota and in 
the adjacent part of Canada, areas now forming national 
reservations, Cahn (1937) writes: "The marten is now prac- 
tically if not completely extinct in the Quetico country. It has 
been exterminated from the Superior National Forest on the 
American side of the boundary. Trappers report seeing an 
occasional track in winter in the larger timber north of Saga- 
naga and Saganagonse lakes, and I found one skull at a trap- 
per's cabin up the Wawiag River in 1930. With the removal 
of the larger timber and their constant pursuit by trappers, the 
marten has retreated to the north." Still farther west, toward 
the edge of the Plains, martens were common in northeastern 
North Dakota a century and more ago. Bailey (1926) quotes 
Wied's early account of the number of furs brought in from 
local and more western points, including 500 or 600 marten 
skins. More particular numbers of martens taken in this 
region are quoted from records of Alexander Henry in 1801-7, 
and range from three to as many as 75 in a season at the Pem- 
bina River. "At present," says Bailey, "there are probably 
no martens in North Dakota," nor is it probable that they 
would return "even with careful protection" to the limited 
forest areas of the State. 

The vast region north arid west of the Great Lakes in Canada 
is probably now the main producing ground for marten fur. 
Concerning the species there Preble (1908) writes: "The marten 
is rather common throughout the forest belt of the Athabaska 
and Mackenzie regions. It varies greatly in color, being more 
or less subject to melanism throughout this area. The 'dark' 
martens are more highly prized than the lighter ones, and 
bring a higher price, sometimes as much as four or five times 
the value of ordinary skins . . . Martens are common 
along the Athabaska, Slave, and Peace Rivers, and large 
numbers are traded at all the posts on their banks. Skins 
taken in these valleys average rather dark in color and furnish 
a good grade of fur. The valley of the Athabaska below Grand 
Rapid is said to be especially good trapping ground. The 
number annually taken by each trapper varies from a few to 
a hundred or more. James MacKinlay informed me that 


three hunters working in the Caribou Mountains, southwest 
of Great Slave Lake, trapped in one season nearly 500 martens, 
an unusually large catch. The lower Liard River region and the 
Horn Mountains also are good trapping grounds. Upward 
of 3,000 skins are said to be usually traded at Fort Norman. 
Fort Good Hope also receives a large number, and as many as 
6,000 have been collected at Fort McPherson during an unusu- 
ally good season . . . Martens vary greatly in abundance 
in the same locality in different years, and to some extent this 
increase and decrease are periodic. The winter of 1903-4 was 
marked by a great scarcity of martens over much of the upper 
Mackenzie Valley, though in other sections the catch was 
about normal." At Fort McPherson, A. J. Stone was told in 
1900 that martens were then as numerous as they had been 
50 years before when the post was first established. 

In Keewatin and the Hudson Bay region Preble (1902) 
found the marten (subspecies abieticdta^l^Tly common" 
north to the tree limit but most abundant in the heavy spruce 
forests of the southern part. Many skins come in to the posts 
at Norway House, Oxford House, and York Factory, and a 
few at Fort Churchill that come from the lower Churchill 

Osgood (1900, p. 44), in describing the large marten (M. a. 
actuosa) of the Yukon region, Alaska, wrote at the time that 
it "is still the commonest fur-bearing animal of Alaska, not- 
withstanding the hundreds of thousands that have already 
been taken. Trappers are always confident of a harvest of 
martens whether other animals are abundant or not." To the 
westward, however, in the Cook Inlet region at the base of the 
Alaskan Peninsula, Dr. Osgood (1901) found it only moderately 
common. One trapper had taken but 15 martens in a season 
of two and a half months in 1899 near the mouth of Turnagain 
Arm. The martens of this area seem to be the typical sub- 
species. The latest report (July 16, 1939) of the Biological 
Survey credits Alaska with having shipped in 1938 no less than 
9,200 pelts of this animal, surely a large number. 

In the western United States and along the Pacific coast 
the numbers are much less, though it is still fairly plentiful in 
British Columbia. Of the insular race, nesophila, of the Queen 
Charlotte Islands, British Columbia, Dr. Osgood (1901) 
reports that it evidently is not abundant. The Haida Indians 


trap more or less every winter for them, but the annual catch 
seldom exceeds 40 skins. According to the fur traders these 
martens are short-haired and light-colored and do not com- 
mand so high a price as do those from the mainland. On 
Vancouver Island they seem to be "fairly common in the 
higher mountains " (Swarth, 1912). 

While martens are absent from the treeless parts of western 
United States, they are common in many areas of forest, in 
spite of much trapping. Vernon Bailey (1918) wrote two 
decades ago that in the Glacier National Park region of north- 
western Montana "they are probably as common in the park 
as anywhere in the country, but no animal with the price on 
its skin that they have long maintained could well be numerous 
or very common. For at least half a century the park region 
has been famous for the number of martens caught each year 
by trappers . . . The animals are reported to be more 
common on the west slope of the mountains than on the east, 
but this is probably because the timber there is more dense and 
extensive and it has not been possible to trap them out so 
thoroughly . . . Steel traps are generally employed, but 
many deadfalls of the ordinary type are used." He recom- 
mends that "special permits to reliable parties for trapping 
them in the park during a limited season when they become 
too numerous would probably control their numbers here, 
while outside the park there is no danger of their ever becoming 
too abundant." They were reported common in the nineties 
in the Salmon River and Saw Tooth Mountains of Idaho by 
Merriam, who says that prospectors even complained that 
martens would carry off their meat from camp. Southward, 
the Rocky Mountain marten (subspecies origenes) seems to be 
still fairly common throughout the forested areas into northern 
Colorado and in smaller numbers into the higher mountain 
forests of the northern part of New Mexico. Gary (1911) 
writes that from 1900 to 1905 the best regions for martens in 
Colorado were the mountain ranges surrounding Middle and 
North Parks, the Williams River Mountains, and the moun- 
tains south of Aspen, Pitkin County. They were reported 
common on the Park Range but very rare on the Medicine 
Bow Range. "A conservative estimate of the annual catch 
in the Middle and North Park region would be 100 skins." 
Fur buyers at Aspen estimated about the same for their dis- 


trict. Cary was informed that 30 years before (about 1875) 
a trapper had taken a great many martens in the mountains of 
Gunnison County, central Colorado. Farther south the marten 
is much less common and reaches the southern limits of its 
range in northern New Mexico. Bailey (1931) adduces a few 
older records for this region and concludes that "the animals 
occur, but are by no means common in the Canadian Zone 
forests of the Sangre de Cristo and San Juan Mountains. It is 
very doubtful if they occur in any of the ranges farther south." 
According to this author (Bailey, 1936, p. 296) it is this race 
of the marten that extends into the extreme northeastern part 
of Oregon in the Blue Mountains, where, however, they seem 
to be uncommon and decreasing in numbers. In the Wallowa 
Mountains a few are still taken, but recent reports indicate 
that the numbers are small. \ 

In the Coast Ranges of OregoA the resident animal is re- 
garded as typical M . caurina, grading in the southern districts 
into the race sierrae of California^/ Bailey (1936) quotes re- 
turns for the season November 1, 1913, to February 28, 1914, 
of 518 martens taken in that State by registered trappers, 
indicating that the species Js still well represented there. 
These at prices then current were worth in the neighborhood 
of $13,000. In California two races occur, sierrae of the Sierra 
Nevada, and humboldtensis of the northern Coast Range. 
They are confined to the Boreal Zone and are regularly trapped, 
especially in the northern parts of the State. Grinnell, Dixon, 
and Linsdale (1937) have published an extended account of the 
species in California and show that particularly in summer it 
haunts slide-rock areas among the crannies of which it pursues 
small rodents, especially mouse-hares and white-footed mice. 
The southern limit of range seems to be Tulare County, in the 
southern Sierra Nevada; on the coast, however, the marten 
extends only to the northwestern part of the State in the higher 
mountains. The authors quoted say that "at least three- 
fourths of the martens trapped in California are taken by a few 
professional trappers who specialize in this animal." One of 
these men took as many as 96 in a single season. However, 
reports from the trappers show "a marked decline, amounting 
to fully 75 per cent, in the number of martens trapped in a 4- 
year period," from 452 in 1920 to 121 in 1924, a decrease be- 
lieved to be independent of any cyclic decline. The authors 



believe this is due to over-trapping and that unless more ade- 
quate protection is given the species it will in a short time be- 
come scarce or absent except in such protected areas as the 
national parks. A year-round closed season is recommended. 
Concerning the marten of the Coast Range (race humboldtensis) 
the authors above quoted regard it now as of "rather sparse 
occurrence" in the redwood belt, "though in earlier years it 
was more generally distributed and fairly numerous." Its 
southernmost occurrence was in Sonoma County. In 1926 
and 1927 two trappers were said to have worked a somewhat 
inaccessible pocket within ten miles of Carlotta and to have 
caught in all 25 martens, which is considered a remarkable 

In general it may be said that while the pine marten is still 
plentiful in the most favorable parts of its range, as in the 
northwest of Canada and in parts of the Rocky Mountains and 
Alaska, its outpost areas in the east and south have been so 
very much depleted in recent decades that it is in need of 
careful protection if it is to maintain anything like former 
numbers or to survive at all. They require a large stretch of 
territory to range over and a sufficient food supply in the way 
of small rodents. Factors that seriously affect them are the 
burning by forest fires of parts of their forest range, clearing 
through extensive lumbering operations, overtrapping, and 

Eastern pine marten (Maries americana americana) 


killing of females. The fact that they are more or less subject 
to a periodic cycle of decline, probably due to disease or failure 
of food supply through dying off of certain rodents that form 
a staple diet, or to both factors combined, probably would 
have an influence on the numbers of a small population, and 
when these factors coincide with other unfavorable circum- 
stances a danger point is reached. Martens are relatively easy 
to trap, so that this trait adds to the hazard of their lives. 
Finally, an important factor in their decline from a peak of 
numbers lies in the fact that their gestation period is long, so 
that killing of females often involves the death of a potential 
litter. At the present time there have been various attempts 
made to raise martens in captivity, but with only moderate 
success. Young are born in March and April and are independ- 
ent by some three months later. Further experiments being 
carried on by the U. S. Fish and Wildlife Service will doubtless 
provide a better knowledge of the best methods of management. 
In 1936 a five-year closed season on martens in the United 
States was urged by the Chief of the Biological Survey and en- 
dorsed by the American Wildlife Conference. Some idea of the 
number annually killed in western Canada may be gained from 
the statement of Cameron (1936, p. 625) that the fur yield of 
the Northwest Territories for the year 1933-34 included no 
less than 5,580 marten pelts. 


Mustela pennanti Erxleben, Syst. Regni Anim., vol. 1, p. 470, 1777 (eastern Canada). 

SYNONYMS: Mustela canadensis Schreber, Saugthiere, vol. 3, p. 492, pi. 134, 1778 
(eastern Canada); Mustela melanorhyncha Boddaert, Elenchus Anim., p. 88, 1784 
(eastern Canada, based on Pennant); Viverra piscator Shaw, General Zool., vol. 
1, p. 414, 1800 (eastern Canada, based on Pennant); Mustela nigra Turton, 
Tiirton's ed. Linnaeus's Systema Naturae, vol. 1, p. 60, 1806 (eastern Canada, 
based on Pennant); Mustela godmani J. B. Fischer, Synopsis Mamm., p. 217, 1829 
(Pennsylvania); Mustela canadensis pacifica Rhoads, Trans. Amer. Phil. Soc., 
new ser., vol. 19, p. 435, Sept. 1898 (Lake Keechelus, Kittitas County, Washing- 

A possibly valid race lately distinguished is 

Maries pennanti columbiana Goldman, Proc. Biol. Soc. Washington, vol. 48, p. 176, 
Nov. 15, 1935 (Stuart Lake, near headwaters of Fraser River, British Columbia). 

FIGS.: Nelson, 1916, p. 446. upper fig. (col.); Grinnell, Dixon, and Linsdale, 1937, 
vol. 2, pi. 5 (col.); figs. 72-81 (details, skull, habitat). 


This larger relative of the pine marten has a range in North 
America nearly coextensive with that of its smaller cousin and 
covers mainly the coniferous forests from eastern Canada and 
Maine across to Hudson Bay, southeastern Alaska and British 
Columbia, and southward on the higher parts of the mountain 
country to Tennessee in the East, the Rocky Mountains to 
Wyoming, and the Sierra Nevada in California. Although 
Rhoads in 1898 described as a separate race the fisher of the 
Pacific coast, Grinnell, Dixon, and Linsdale (1937) have lately 
demonstrated that there are no clear distinctions setting off the 
animals of that region, but that only a single recognizable form 
covers the immense range indicated. Goldman, however, 
believes the British Columbian animal is distinguishable. The 
fisher is much larger than the pine marten, measuring about 
40 inches in length, of which the tail vertebrae comprise about 
16 inches. The ears are shorter, hardly projecting above the 
head. Females are smaller than males. Males may weigh up 
to about 10 pounds, females to about 5.5 pounds. The general 
color is ashy brown on the head, gradually darkening posteri- 
orly to black on the legs and tail. There may be irregular 
white areas on the throat and chest or on the abdomen. 

This large marten is highly prized for its handsome and 
durable fur and is much trapped and sought after. The pelts 
bring a high price. Not many years ago a prime skin might 
fetch as much as $100. Fishers, like martens, inhabit the 
heavy woods, subsisting on large and small mammals, such as 
hares and even the porcupines, which they seem to attack 
successfully; squirrels, wood rats, and mice of various kinds 
are also taken. Though hunting and traveling much on the 
ground, they are also expert climbers and possessed of great 
strength and agility; like the marten they have the habit of 
passing at intervals over a somewhat definite course, and are 
said to go often in pairs. Two or three young are the usual 
number, born late in spring (late in May in California). 

On account of the value of its fur, the fisher has everywhere 
been relentlessly hunted and trapped, yet it is remarkable that 
it has persisted in some areas in spite of persecution. In the 
East there are still a few fishers left in northern and central 
Maine, northern New Hampshire and the White Mountains, 
and in northern Vermont and the Green Mountains. Probably 
a few are to be found in southwestern New Hampshire in the 


Mount Monadnock region; at all events, the late Abbott 
Thayer obtained one or two specimens there 20 or more years 
ago. Formerly there were fishers in western Massachusetts, as 
attested by Emmons (1840) a century ago, but it is safe to say 
that they are no longer to be found in that State. Nevertheless 
they are still taken in some numbers in the Adirondack region 
of New York, and according to the 1939 report of the Biological 
Survey that State leads in the total number of fishers taken in 
1938, no less than 117, exceeding even Alaska! Dr. Francis 
Harper (1929) quotes from the reports of the New York State 
Conservation Department the numbers of fishers caught, pre- 
sumably all in the Adirondack counties, for the years 1920-25, 
as follows: for 1920, 132; for 1921, 186; for 1922, 563; for 1923, 
112; for 1924, 144; for 1925, 61. The total for 1922 of 563 
animals seems very remarkable. Apparently in spite of these 
large catches the animals are still holding their own. This 
area must be one of the most favorable for them in the country. 
In Pennsylvania the fisher was formerly found in some num- 
bers, especially in the beech forests of the more mountainous 
portions, but it had much declined by the middle of the last 
century. Rhoads (1903) has assembled many notes on its 
former occurrence, most of which attest its presence in the 
fifties and seventies, with a few instances of its more recent 
capture, as in Lancaster County in 1896; but most of the re- 
ports speak of it as gone by the beginning of the present cen- 
tury. In summary he says: "At the present time [1903] about 
the only counties where these animals are to be found are 
Clearfield, Cameron, Elk and probably Clinton, Potter, and 
Sullivan, and in all of these they are reported to be very rare." 
Reports of fur dealers, Rhoads further states, indicate that 
probably not over half a dozen fishers were annually killed in 
Pennsylvania at that time. In a more recent survey of the 
mammals of Pennsylvania by Williams (1930) the animal is 
not mentioned at all. Southward of Pennsylvania the fisher in 
former times extended along the Alleghenies of West Virginia 
to North Carolina and Tennessee, but seems to have long ago 
been extirpated from these regions. Dr. Remington Kellogg 
(1937, 1939) states that the authentic records for West Vir- 
ginia are few; according to Surber (1909) it formerly occurred 
in the black-spruce region, but F. E. Brooks (1911) gives no 
specific records later than 1873. In 1888 Dr. C. Hart Merriam, 


after a journey of several hundred miles through "the Great 
Smoky Mountains of Tennessee and North Carolina, reported 
that the pekan was unknown" to local residents, but "reliable 
information exists that this animal formerly occurred in that 
area" (Kellogg, 1939). Audubon and Bachman (1846) mention 
skins from eastern Tennessee and the capture of an individual 
near Flat Rock. 

West of the Alleghenies the fisher seems to have been rare 
even in early days, south of the Great Lakes. Bray ton (1882) 
includes it in his list of Ohio mammals but cites no specific 
records. For Indiana, Dr. Lyon (1936) states that there is 
considerable evidence of its former presence, "either as a regu- 
lar resident or as a wanderer from the north," but specific 
instances are perhaps three, which he gives as (1) the statement 
of Plummer in 1844 that it was not uncommon at an earlier 
period, but had not been seen since 1820; (2) the listing of a 
skin purchased of local trappers near Noblesville in 1859; and 
(3) the testimony of Wied as to its former occurrence at New 
Harmony. Evidently it has been gone from this region for 
well nigh a century. Cory in ^912 believed it by no means 
improbable "that a few individuals may still exist in some of the 
extreme northern counties" of Wisconsin, and he was informed 
"that it is occasionally taken in the wilder portions of the 
Michigan peninsula." He instanced three from the latter 
region, taken in Iron County in 1898 and 1900. The latest 
record for this region that he gives is of one killed in November 
1900, between Iron Mountain and Pembine, Wis. Kennicott, 
half a century before, mentioned it as rather frequently seen 
in the heavy timber along Lake Michigan. Evidently it is now 
practically .gone from the lake region. As to its present status 
still farther northwest, along the border of Minnesota and the 
adjacent Quetico Park, Canada, Cahn (1937) writes: "The re- 
moval of the large timber has profoundly affected the distribu- 
tion of the fisher, as it has that of the marten, and it is now to 
be regarded as extremely rare in the Quetico, although some 
still are to be found there ... I have in my collection 
17 skulls of fisher which I collected from about the camps of 
trappers up the Wawiag River. " In former days the range of 
the fisher extended to northeastern North Dakota, where 
judged from the data presented by Bailey (1926) it was fairly 
common at the beginning of the last century, for Alexander 


Henry records a total of 1,118 skins brought in from the sur- 
rounding region during the years 1801-7. In 1833 Maximilian 
zu Wied stated that the approximate number of skins annually 
brought in to Fort Union (now Buford), N. Dak., was 500 or 
600, but a large part of these doubtless came from still farther 
west. "At the present time, " Bailey says, "there are certainly 
no fishers within the State and there seem to be no authentic 
records of their occurrence since the early trapping days. " 

In eastern Canada the numbers of fishers have been sadly 
depleted in recent years. Although formerly found in Nova 
Scotia, there seems to be no evidence of its presence there since 
Gilpin's time (1868) when he included it among the mammals 
of that province. In New Brunswick a few are found in re- 
moter parts, as in the mountains of the Gaspe region, whence 
Goodwin in 1924 records a few taken by local trappers. In 
Quebec it is now "one of the most rare and valuable fur-bearing 
mammals of Canada . . . According to Low (1889) the 
fisher does not occur east of Mingan nor north of Mistassini 
[central Labrador]. It is still found in small numbers around 
the southern end of James Bay but is extremely rare near any 
white settlements. With a consistently high price paid for its 
fur, the fisher is pursued relentlessly and has steadily decreased 
in numbers in all parts of its former range" (Anderson, 1939a). 
West of Hudson Bay Preble (1902) found it "sparingly through- 
out the southern part" of Keewatin and saw many skins at 
Norway House and Oxford House. Near York Factory it was 
rare, but "farther south more are taken; about thirty or forty 
are annually traded at Trout Lake and a few at Severn River. " 

The same author (Preble, 1908) states that in the Athabaska 
and Mackenzie region it is found throughout the country north 
to Great Slave Lake and Liard River. "It seems to be no- 
where abundant, and becomes rare toward the northern limits 
of its range. Along the Athabaska and Slave rivers a limited 
number of skins are collected at all the posts north to Fort Reso- 
lution, and I was informed that the animal was rather common 
in the region about the mouth of Peace River." Although 
according to Osgood (1900), "there is little doubt that the 
animal occurs along the upper Yukon" no Alaskan specimens 
could be adduced in proof, nor does Coues (1877) mention any 
localities in his statement that its range includes Alaska. 

Fishers apparently are present in some numbers in the forests 


of British Columbia, and Goldman (1935) has recently distin- 
guished as Maries pennanti columbiana the animal from the 
headwaters of Fraser River. The species seems to be absent 
from Vancouver Island but is found in small numbers in the 
States of Washington and Oregon, particularly in the national 
forest reserves. Scheffer (1938), in some recent notes on the 
species in the State of Washington, says that it occurs in the 
Cascade and Olympic Mountains and on the Olympic peninsula 
north to Skagit Valley and Cascade River. In 1937, however, 
the U. S. Forest Service estimated that the total number in 
the national forests of Washington was about 230 individuals. 
In Oregon fishers are found in the cool humid coast ranges, and 
probably also in the Blue Mountains section (Bailey, 1936), 
but the number taken for fur seems to be small. Thus in the 
trapping season of 1913-14, nine fishers were reported to the 
State Game Commission by registered trappers as taken in 
Washington. At that time skins were quoted at $25 each, but 
in 1920-25 the price advanced to as much as $100 to $150 
apiece! Fewer numbers are found in the northern Rocky 
Mountain region. Bailey (191$), writing of the mammals of 
Glacier National Park in Montana, speaks of it as rare, but the 
few remaining in the park are now, with proper protection, 
"likely to hold their own" for some time to come. "In many 
years of trapping in the park region in the early days, Walter S. 
Gibb has caught three of these animals, but some of the old 
trappers have not secured a skin. Donald Stevenson reports 
two skins that were taken by trappers on the Upper Swift cur- 
rent in 1910, and tracks which he saw on Swan River and 
South Fork as late as 1912. " Merriam in 1891 reported fishers 
as already rare in Idaho and mentions a large adult male caught 
the previous year near Alturas Lake that weighed 10 pounds 2 
ounces. Formerly at least it extended its range to Wyoming, 
but no recent report of its presence there is available, nor is 
there according to Cary (1911) any evidence of its being found 
in Colorado, although in 1874 it was said to occur (J. A. Allen). 
Grinnell, Dixon, and Linsdale (1937) have given an excellent 
account of the fisher, its habits and range, as known at the 
present time in California. It is found still "in the north- 
western part of the State south from the Oregon line to Lake 
and Marin Counties and east to and including Mount Shasta; 
not often in the immediate coastal region (redwood belt) nor, 



so far as known at present, in the Warner Mountains, Modoc 
County; south from Mount Shasta and Lassen Peak through- 
out the main Sierra Nevada to Greenhorn Mountain in north 
central Kern County. " It is found mainly at middle altitudes, 
from 2,000 to 5,000 feet in northern California and 4,000 to 
8,000 feet in the Mount Whitney region to the southward. At 
the present time the fisher is found in the Sierra Nevada mainly 
on the west side of the range. On the coast range it formerly 
reached Marin County as the southward extension of its area 
of distribution. Fishers are nowhere abundant in California, 
and at best "it is unusual to find more than one or two to the 
township" in suitable country. These authors believe "it is 
doubtful if there is one fisher to each 100 square miles" of its 
range in California, and in 1926 estimated the entire population 
in the State at about 300. Forty years before 1909 they were 
found all along the ridges of Mendocino County, but owing to 
excessive trapping very few were then left. From 1920 to 1924 
the annual catch dropped from 102 to 34, or a loss of 67 percent 
in five years. This decrease has involved the entire State and 
is not merely local. Since 1925 the fisher has received total 
protection in the national parks of California. The authors 
mentioned believe that a long-term closed season is much need- 

Fisher (Martes pennanti] 


ed if the species is to continue in the State. They put forth as 
major causes in the decrease of the species the following facts : 
(1) The fisher is by nature a solitary animal; (2) it requires a 
large area of forage territory in order to live ; (3) the areas suit- 
able for it are limited ; (4) the rate of reproduction is relatively 
low; (5) the forests on which it depends are being constantly 
reduced. To these factors may be added the hazard of forest 
fires, and, finally, the fact that the gestation period is 11 
months, so that trapping operations in winter, through de- 
stroying pregnant females, inevitably double or triple the 
potential loss of individuals. 

The beauty and rarity of fisher fur as well as the high price 
it brings make it an attractive proposition for breeding under 
artificial conditions. Ashbrook (1928) points out, however, 
that the breeding of fishers in captivity for fur has not expanded 
to any great extent since 1912, when there were but two fisher 
"farms " on this continent. The difficulty seems to lie in getting 
them to breed under these conditions even with seemingly 
proper food. In the case of an animal naturally so shy and 
nervous, breeding seems to be accomplished with difficulty 
where there is the least disturbance. At present "little authen- 
tic information is available on handling fishers in captivity," 
but no doubt the difficulties will in time be solved. 

Dr. Francis Harper has contributed notes as to the number 
of fisher pelts yielded in recent years. Thus in the London fur 
market in 1927 one dealer offered 1,700 skins; at another sale 
2,729 were offered (Journ. Soc. Preservation Fauna Emp., pt. 
9, p. 75, 1929). According to Cameron (1936) the fur yield of 
the Northwest Territories, Canada, included in 1934 the pelts 
of 21 fishers. 


Lutreola macrodon Prentiss, Proc. U. S. Nat. Mus., vol. 26, p. 887, July 6, 1903 
("Brooklin, Hancock County, Maine," from Indian shell-heaps). 

SYNONYM: Lutreola vison antiquus Loomis, Amer. Journ. Sci., ser. 4, vol. 31, p. 228, 
Mar., 1911 ("Flagg Island, Casco Bay, Maine"). 

FIGS.: Prentiss, 1903, fig. a (rostrum); Loomis, 1911, figs. 1, 2 (rostrum and jaws). 

Remains of a very large mink occur commonly in Indian 
shell-heaps along the shores of the mainland and islands of the 
coast of Maine, as independently described by Prentiss in 1903 


and by Loomis eight years later. The upper tooth row (front 
of middle incisor to back of molar) in the type specimen meas- 
ured 30 mm., and in Loomis 's type 29.5, which is larger even 
than in the large Alaskan mink. Rostra are infrequent among 
the shell-heap fragments obtained, but jaws are fairly common 
in kitchen middens along the coast and islands of Penobscot 
Bay and Mount Desert Island. Two sizes, evidently repre- 
senting males and females, occur. 

No other type of mink is found in these Indian kitchen 
middens, and since there is an absence of European artifacts 
it is believed that these shell-heaps date from previous to, or 
perhaps up to, the time of European occupation. Remains are 
at present known from such sites as far as Casco Bay in the 
south, and northeastward to Mount Desert and Frenchmans 
Bay, and Roques Island, Washington County, Maine. Soon 
after the description of these bony fragments, the late Manly 
Hardy published a note (Forest and Stream, vol. 61, p. 125, 
1903) recalling that his father, himself, and other fur-buyers 
had in former years "recognized as a distinct form a very large 
mink, skins of which were received from islands and coast of 
Maine until about 1860. By reason of their large size they com- 
manded a higher price than did the skins of other minks . . . 
To this animal the name of 'sea mink' was given by the fur buy- 
ers" of Maine. "According to Hardy, the animal was large, 
very fat, and possessed of an odor entirely different from that 
of" the smaller inland animal (see Norton, 1930). Furthermore, 
the fur was said to be coarser and of a more reddish color than 
that of the latter. There seems no reason to doubt that the 
sea mink is the one that inhabited the Maine coasts in Indian 
days and persisted up to the latter part of the last century. 
With a limited range and on account of eager pursuit by the 
trappers, it probably became extinct about the sixties or seven- 
ties. Norton (1930) tells of a very large mink skin handled 
about 1880 by fur buyers at Jonesport, Maine, which without 
the tail measured over 26 inches long and may have been one 
of the last of this race. It was taken on one of the islands in the 
township of Jonesport. Norton mentions also a large mounted 
specimen of a mink taken at Campobello Island, New Bruns- 
wick, about 1894, and then in the collection of Clarence H. 
Clark, of Lubec, Maine. He speaks of its light color (possibly 
a result of fading) and regards it as "probably of this species. " 
No measurements are given. 


It seems probable that this big animal was the only form of 
mink to be found in the eastern part of the Gulf of Maine in 
earlier times. That it also ranged to Nova Scotia seems likely 
from the account by J. B. Gilpin (1867) of large skins from that 
region that measured as much as 32.5 inches in total length, 
though perhaps slightly stretched. At the present day the 
mink occurring along the Maine coast represents Mustela 
vison mink, the more southern race, of which a large male will 
seldom measure over 23 inches. Evidently this latter race, 
which has somewhat of a liking for seacoasts, has taken the 
place formerly held by the now extinct sea mink. Possibly, 
too, circumstances favorable to this eastward spread within 
the last century contributed to the driving out of the larger 


MUSTELA NIGRIPES (Audubon and Bachman) 

Putorius nigripes Audubon and Bachman, Quadrupeds of North America, vol. 2, p. 

297, 1851 (Fort Laramie, Laramie County, Wyoming). 
FIGS.: Audubon and Bachman, 1851, pi. 93; Coues, 1877, pi. 7 (skull) ; Nelson, 1918, p. 

449, lower fig., col. 

This ferret is of special interest on account of its being in 
North America the only representative of the Old World group 
of black-bellied weasels to which the polecats and their rela- 
tives of eastern and northern Asia belong. On account of its 
angular heavy skull and robust premolar teeth, as well as the 
color pattern, it is placed in a special subgenus with these, 
Putorius, sometimes regarded as a distinct genus. The Ameri- 
can representative, however, has only the feet and facial mask 
and the end of the tail blackish ; the upper parts are elsewhere 
creamy, with a wash of brown over the back, the lower surfaces 
whitish. An adult male is about 20.5 inches (529 mm.) long; 
the tail relatively short, about 5 inches (130 mm.) (Osgood). 

The black-footed ferret is an animal of the interior plains of 
North America, east of the Rocky Mountain foothills, from 
western North Dakota to northern Montana and Alberta (?) 
and thence southward to Texas and central New Mexico. Its 
range is practically coextensive with that of the prairie-dogs, 
in the colonies of which it lives and upon which it preys. Its 
relations with these little burrowers is thus a close one and is 
like that obtaining in Mongolia between its relative the black- 


bellied weasel, Mustela eversmanni, and the bobac marmot in 
the "towns" of which it lives, preying on these rodents. The 
black-footed ferret is believed to rely chiefly upon the prairie- 
dogs for sustenance as an abundant and dependable source of 
food, yet it doubtless captures other small animals, and 
Roosevelt has even recorded a case of one killed in the act of 
attacking a small antelope fawn. It is surprising that in view 
of the abundance and accessibility of its natural prey, the 
prairie-dogs, this ferret should be nevertheless relatively rare, 
for specimens are rather uncommon in collections, and indi- 
viduals are seldom seen in life even by those living in its habitat. 
Bailey (1926) has even suggested "that this very abundance 
[of the prairie-dogs] has in some way pauperized the species 
until reproduction is restricted," but adds that "apparently 
nothing is known of the breeding habits or of the number of 
young at a birth. " Since there are three pairs of mammae the 
size of a litter probably does not exceed this number and may 
normally be less. It is obvious that in a case of such specialized 
food preference it would be disastrous for the species to become 
so common as seriously to deplete the numbers of its host, so 
that undoubtedly some sort of adjustment has been evolved, 
but its nature remains uncertain. Possibly enemies as yet un- 
recognized are a factor, such as rattlesnakes that might devour 
the young, or the hibernating habits of the prairie-dogs, al- 
though not well marked in the southern parts of the range, may 
affect the ferret's abundance. In its habits it is believed to be 
in large part nocturnal, but it has been seen active by day. 

In view of the damage done to range vegetation or to culti- 
vated crops, the hand of the agriculturist is against the prairie- 
dog, and of late years poisoning campaigns on a large scale 
have been carried on that in some localities have largely 
exterminated these rodents. Undoubtedly a further result of 
these activities will be to eliminate or at least to reduce con- 
siderably the numbers of the weasel so that the reason for its 
inclusion among the vanishing mammals is clear. 

Writing of North Dakota, the eastward extension of its 
range, Bailey (1926) says that a few have been taken in the 
western part of the State and adduces four instances in the 
years 1910 to 1915; he quotes Jewett, who killed a specimen 
near Quinion in 1913, that none of the old residents to whom he 
showed it were acquainted with the species. Coues (1877) in 


his account of the history of this animal mentions single speci- 
mens obtained after special enquiries, in Wyoming, Montana, 
Colorado, and Kansas. Hibbard (1933) speaks of it as of 
former occurrence in the western part of Kansas, from which 
probably it is now largely gone. In eastern Colorado it seems 
to be best known, though as elsewhere it is rare. Cary (1911) 
mentions that it was reported to Streator in 1894 from three 
localities in the Arkansas Valley; Lantz heard of a few in 1905 
about Hugo, where they are known as prairie-dog ferrets, and 
he had himself heard of it in Baca and Prowers Counties, 
" where it seems to be more generally known . . . than in 
other sections on the plains." Southward of Colorado black- 
footed ferrets "are found over the Staked Plains country of 
western Texas and undoubtedly occupy the plains region of all 
eastern and northern New Mexico, but few specimens have 
been taken" (V. Bailey, 1931) ; a number of specific records are 
given for this part of the State, to which Bailey (1905) adds a 
few more from east and south of the Staked Plains of Texas. 
Although the range may be expected to overlap slightly the 
borders of southeastern Arizona and the adjacent parts of 
Mexico, where prairie-dogs occur, specific records are not at 

Nelson (1918) is inclined to attribute to its activities the 
occasional deserted prairie-dog "towns," which "are not infre- 
quently observed on the plains with no apparent reason for the 
absence of the habitants." He mentions one case, however, 
where a black-footed ferret lived for several days under a 
wooden sidewalk in the border town of Hays, Kansas, "where 

Black-footed ferret (Mustela nigripes) 


it killed the rats harboring there. " As to its prospects for the 
future he writes: "With the occupation of the country and the 
inevitable extinction of the prairie-dog over nearly or quite all 
of its range, the black-footed ferret is practically certain to 
disappear with its host species," a sad prophecy for this re- 
markable and useful animal! 


Ursus luscus Linnaeus, Systema Naturae, ed. 12, vol. 1, p. 71, 1766 (Hudson Bay). 
Gulo auduboni Matschie, Sitzb. Ges. Naturf. Freunde Berlin, 1918, p. 153 (Rensselaer 

County, New York). 

Gulo bairdi Matschie, op. cit., p. 153, 1918 (Fort Union, North Dakota). 
FIGS.: Elliot, 1901, pi. 36 (photographs of skull); Nelson, 1916, p. 428 (col.); 

Anderson, 1935, p. 87 (photograph). 

The wolverene is the largest of the weasel family and an 
animal of boreal evergreen forests and barrens. It is repre- 
sented by closely similar races in the northern parts of the Old 
World, and although several forms have been named from 
North America it is very doubtful if their validity can be main- 
tained, or if at best some of them are more than slight sub- 
species of the Hudson Bay animal. 

Audubon and Bachman describe a specimen from New York 
State as blackish brown, with a pale reddish brown band ex- 
tending from behind the shoulder to the rump on each side. 
These two bands join across the rump, and form a contrasting 
though not sharply defined light stripe. There is another pale 
brownish- white band from the eye to the ear on each side. 
The fur is long and rather shaggy. The head and body measure 
some two feet nine inches in length, the short tail eight inches. 
The animal stands about a foot high at the shoulders, and 
weighs 25 to 36 pounds. 

Much has been written of the wolverene in the North, of its 
great strength in proportion to its size, and of its habit of 
breaking into the caches of hunters or trappers and spoiling 
what part of the provisions it can not eat. Vernon Bailey 
(1926) writes: "Wolverenes are found mainly within timbered 
sections of the country, but are great wanderers and at times 
may strike out over open country in search of new hunting 
grounds. They are omnivorous hunters and scavengers and 
have the reputation of being gorging gluttons, a fact which 


has given them one of their common names . . . They are 
said to have from two to four young." 

The fur of the wolverene is much in demand among the 
Eskimo and other northern people for trimming their parkas, 
especially the sides of the hood, for it has the peculiar charac- 
teristic that the moisture from one's breath does not readily 
freeze and cake on it. Bailey (1936) writes that the fur has 
never brought high prices; the quotation for prime skins in 
1923 was but $6 to $8, yet at this same time California trappers 
received an average of $25 each for wolverene pelts. 

The wolverene is an animal of solitudes, usually found singly 
and shunning the vicinity of human occupation, so that it 
quickly vanishes with the spread of "civilization. " It is there- 
fore a species that has already gone from a large part of its 
former range in the United States and may be expected to sur- 
vive only in the more remote wilderness areas of the North. 
Of its occurrence in the northeastern States, very little record 
remains. Audubon and Bachman (1846) nearly a century ago, 
wrote that they had heard of its existence, "although very 
sparingly," in Maine, and abou^t 1810 "we" (probably Bach- 
man) had examined three skins in the possession of a merchant 
in Lansingburg, N. Y., that were said to have come from the 
Green Mountains of Vermont. While these do not constitute 
certain evidence of the occurrence of the wolverene in New 
England, it is possible that it was formerly found, as these 
authors say, "very sparingly. " It should be recalled, however, 
that the Canada lynx was sometimes called "wolverene" in 
this region, so that hearsay reports have questionable value. 
Nor can much credence be placed in a recent (1918) report of 
wolverenes in northern New Hampshire (C. F. Jackson, 1922). 
Even in New York State the wolverene must have been rare 
over a century ago. The only records seem to be those of 
Audubon and Bachman (based probably on the latter's obser- 
vations) , who give a circumstantial account of the pursuit and 
capture of one in Rensselaer County in 1810, the description 
and measurements of which are detailed and form the basis of 
Matschie's (1918) Gulo " auduboni"', and of a second examined 
in 1827 that had been killed near Sacketts Harbor, Jefferson 
County, near Lake Ontario. The most southerly record in the 
East is that given by Rhoads, of a wolverene killed about 1858 
near Great Salt Lick in Portage Township, Pennsylvania. 


Surprising as this seems, Rhoads (1903, p. 165), from his 
knowledge of the men who told it to him, had faith in the truth 
of this occurrence. The animal was said to have been caught 
in a wolf trap. Wolverenes formerly were found in the forests 
of New Brunswick but seem to have disappeared quickly. 
Chamberlain (1884) mentions in his "Mammals of New 
Brunswick" that about 1850 "it was occasionally met with; 
but no recent instance of its occurrence is known." In the 
Province of Quebec the wolverene "has become extinct or 
very rare in the more southern districts, but is found in small 
numbers in northern Quebec. It is more commonly found in 
wooded districts, but frequently wanders far into the Barren 
Grounds of the north and even occasionally to the Arctic 
islands" where, Dr. Anderson (1935, 1939) writes, it has been re- 
corded from Victoria, King William, and Melville Islands, and 
it occurs fairly commonly in the region of Repulse Bay. 
Several times it has been taken within 30 miles of Pond Inlet. 
"The comparative scarcity of the Wolverene in the southern 
half of Baffin Island, and the relative frequency at the northern 
end of the island, is good evidence that they come to Baffin 
island by way of Melville peninsula. " It still is found in small 
numbers in the Labrador Peninsula but probably seldom now 
in the southern part where Cartwright found it a century and 
a half ago. Audubon and Bachman (1846) mention having 
seen specimens "procured at Newfoundland," but it may well 
be doubted if they were trapped there. At all events there 
seems to be no evidence that wolverenes occurred on that 
island within the memory of living men. 

Westward along the northern border of the United States 
wolverenes were formerly present in small numbers. Cory 
(1912) writes that "old trappers living in the vicinity of 
Champion, Michigan, claim that wolverines were occasionally 
killed in that locality, 30 or 35 years ago" but quotes Edward 
G. Kingsford, of Iron Mountain, Mich., who was much in the 
woods of northern Michigan, Wisconsin, and Minnesota be- 
tween 1880 and 1900, to the effect that the Rainey Lake dis- 
trict in the last-named State was the nearest to Michigan he 
had ever known of one being killed. Schorger (1939) has 
lately unearthed, however, what seems a "satisfactory record" 
of one trapped on February 15, 1860, by a man living at 
Marquette, Lake Superior, in the northern peninsula of Michi- 


gan. In earlier times there were a few in Wisconsin, and Hoy 
mentions one killed in La Crosse County in 1870. Rather 
surprising, however, is the evidence adduced by Hahn (1909) 
of a wolverene having been killed in 1840 in Noble County, 
Ind., and a second in 1852 in Knox County of the same State. 
The records probably mark nearly the southern limit of range 
in the east in historic times. In Minnesota, Herrick, writing in 
1892, says he has no recent knowledge of it, so that it doubtless 
was rare there at that time, but Kingsford, quoted by Cory 
(1912), says that "about 1895 to 1897 they were quite plentiful 
in northern Minnesota. " More recent records indicate that a 
few individuals were found in Minnesota up to a much later 
time, for Dr. H. H. T. Jackson (1922) has recorded a skull in 
the collection of the U. S. Biological Survey from an animal 
killed in Itasca County, January 11, 1899; and C. E. Johnson 
(1923), who made special efforts to learn of the wolverene's 
former status, records that a fur trader of long experience in 
northern Minnesota told him in 1922 that in 30 years of buying 
furs in the region he would not average more than one wolver- 
ene pelt a year and had bought none during the previous three 
years along the Canadian border. He specifically mentions a 
pelt bought by this trader that was taken in 1918 in the 
northern part of St. Louis County, Minn. It seems to be the 
last definite record for that State. 

In the early years of the last century these animals were 
found at least in the northeastern part of North Dakota and 
were taken in small numbers with the fur catch of those days 
(see V. Bailey, 1926). It seems to have been present in the 
State till about 1850, but there are no later records. According 
to Baird (1857) it occurred in "the Black Hills of the Missouri" 
at about the same time. 

While in the eastern United States and southeastern Canada 
the wolverene is doubtless now to be regarded as extinct, it 
still is found in some numbers in the wilder and more moun- 
tainous parts of the western United States and more frequently 
in the Canadian Northwest, although on account of its solitary 
habits it can not be accounted a "common" species at any 
time. Writing of the mammals of the Athabaska-Mackenzie 
district Preble (1908) states that it is found throughout that 
region "but is nowhere common, though a few skins are 
collected at all the posts we visited. During our trip in 1901 


more skins were seen at Fort Rae than at any other post, and 
these had been brought mainly from the Barren Grounds, 
where the animal occurs more commonly than in the wooded 
country. . . In the semibarren country along the southern 
shore of Great Bear Lake, tracks of wolverenes were common. 
. . . A large proportion of the wolverene skins which are 
obtained from the Indians of the Mackenzie are shipped to 
Fort McPherson and traded to the Eskimo, who prize this fur 
highly for trimming their clothing." Pike and other travelers 
mention it as being met with across the Barren Grounds. 
Trappers in the Yukon region secure a few annually. A. J. 
Stone (1900) in the Canadian Northwest found them in timber 
and on the barrens and mentions seeing them "far out on the 
ice of the deep bays along the coast." Osgood (1901) speaks of 
them as "apparently rather common" in the Yukon and also 
in the Cook Inlet region of Alaska, whence a number of skins 
annually are sent via St. Michael to trading posts on the 

In British Columbia, on the other hand, wolverenes seem to 
be diminishing in numbers. Cowan, in 1939, writes that they 
are "rare in the Peace River .district, but according to certain 
reports are more plentiful in the Rocky Mountains." He 
mentions one trapped on Pine River in 1936. On Vancouver 
Island a few wolverenes still exist, according to Swarth (1912), 
but are confined to the higher mountains and are seldom 
trapped, except by the Indians, who bring in "one or two every 

In the Rocky Mountains south of the Canadian border a 
small number of these mammals remain. Fifty years or more 
ago Merriam (1891) wrote that wolverenes were "tolerably 
common in the Saw Tooth Mountains" of Idaho, where in the 
previous winter a trapper had caught five. Another had been 
taken a few years before 1891 in the Blackfeet Mountains. 
No doubt their numbers have since diminished, for Vernon 
Bailey in 1918, writing of the mammals of Glacier National 
Park in northwestern Montana, says that although in 1895 he 
was told by trappers in the St. Mary Lake region that a few 
existed there and one was occasionally caught, and that a few 
were trapped each year before the park was made a reserve, 
there were thought to be none left in the park at the time of 
writing. In the high forests of the wilder parts of Colorado 


wolverenes were still to be found in 1911, but although in 
former times generally distributed, they were then apparently 
restricted "largely to the San Juan and La Plata Mountains, 
the mountains of northern Gunnison County, and the ranges 
surrounding North and Middle Parks" (Cary, 1911). A num- 
ber of specific records are given by Cary, which prove that the 
species was much commoner in the latter part of the last 
century. To the southward there is some evidence that 
wolverenes extended into the higher mountain forests of 
northern New Mexico. Of this Coues (1877) was assured by 
hunters there in 1864, but no definite records exist (Bailey, 
1931), and it must have soon been extirpated from this southern 
outpost of its range. Confirmatory evidence, however, is 
given by Seton (1931), who was told by an Acoma Indian in 
1930 of an animal known formerly to the people of that part 
of New Mexico as the Ho-ho-an or Ko-ko-an, somewhat like a 
small bear with a bushy tail, and the Indian then produced from 
his medicine bag a small carving of the creature's head, an 
excellent likeness of a wolverene's. He said it "was formerly 
found in all these mountains but had disappeared. None of 
the present generation had seen one." Baird mentions a 
specimen from the basin of the Great Salt Lake in Utah 
previous to 1857, but it has long been gone from this region. 

The status of wolverenes on the Pacific coast still requires 
investigation as to numbers and racial relationships. Elliot 
in 1905 distinguished specimens from Mount McKinley, 
Alaska (type from Sushitna River), as Gulo hylaeus on the 
basis of their darker instead of gray heads and larger audital 
bullae as compared with specimens from eastern Canada. It 
is perhaps a recognizable race. In 1918, Matschie named as a 
distinct species on very slight basis, Gulo katschemakensis, from 
the Kenai Peninsula, Alaska, but it is doubtful if this is really 
different. In the same paper he described Gulo niediecki from 
a skull from Dease Lake, British Columbia, but here again the 
validity of the race is questionable. Very likely, however, the 
animal of the coastal ranges is a recognizable form, described 
by Elliot as Gulo luteus, and is accorded subspecific rank by 
Dr. Grinnell (see postea). 

Recent information gathered by Dr. Francis Harper indicates 
that wolverenes are still taken in some numbers in northwestern 
Canada and a few in northeastern Canada. In 1931-32, 536 


wolverene skins were produced, according to returns from 
Canada (Journ. Soc. Preserv. Fauna Empire, p. 75, pi. 21, 
1934). In 1934 the fur yield of the Northwest Territories in- 
cluded 98 wolverenes (Cameron, 1936, p. 625). 

In 1936, the Chief of the U. S. Biological Survey urged a 
five-year closed season on wolverenes in the United States and 
a resolution to the same effect was adopted by the North 
American Wildlife Conference at Washington, D. C., shortly 
after, in the hope of saving the species from complete extermi- 
nation within the United States. That Alaska is nearly the 
center of the wolverene's abundance at the present time is 
indicated by the fact that according to the report of the 
Biological Survey, 248 skins were taken in Alaska in 1938. 
No reports of it are given for other States. 



Gulo luteus Elliot, Field Columbian Mus. Publ., zool. ser., vol. 3, p. 260, Dec., 1903 

(27 miles south of Mount Whitney, Tulare County, California). 
FIGS.: Grinnell, Dixon, and Linsdale, 1937, vol. 1, pi. 6 (col.), text-figs. 91-97. 

An excellent account of the characters and habits of this race 
is that given by Grinnell, Dixon, and Linsdale (1937), summing 
up present knowledge of the wolverene in California. It was 
named by Elliot in 1903 as a separate species differing in its 
supposedly paler coloration as compared with eastern animals. 
The authors quoted show, however, that this was "merely a 
feature of a young stage of pelage" and that in the original 
description not a single character given is diagnostic of the 
race, which may only be distinguished by skull characters, 
namely, "skulls from California are slightly smaller than 
skulls of corresponding ages from Hudson Bay, Alberta, 
British Columbia, and Alaska, and the dentition is noticeably 
and uniformly lighter in particular the carnassials are de- 
cidedly less massive. Three examples from Idaho and one 
from Utah, in the National Museum, are small-toothed like 
the California skulls." A Montana and a Minnesota skull are 
said to be of intermediate size, while one from Chelan, Wash., 
is "nearer the average of Sierran luteus" 

The range of this race may thus be taken to include the 
Sierras of California northward to Washington, merging by 


imperceptible degrees with that of the typical form in Utah 
and Idaho. Although formerly the range was more extensive, 
over the higher parts of the California mountains, it has, since 
at least about 1900, become restricted to the central and 
southern Sierra Nevada above 8,000 feet "from the vicinity of 
Lake Tahoe south through the Mount Whitney region" to 
Kern County. Though the altitudinal range varies from 5,000 
to 13,000 feet, the animals are most frequent at about timber- 
line. Probably this restricted range has resulted in making 
the population of Californian wolverenes never very large. 
The first to mention the species in this State was Cooper, 
who in 1868 wrote that at that time a few were killed every 
winter in the northern Sierras. There is some evidence, ad- 
duced by the above authors, that in the middle of the last 
century wolverenes occurred in the coastal mountains of Marin 
and Sonoma Counties, but no specimens are extant. Slightly 
farther north, however, these animals seem to have occurred 
down to about 1873, as on the higher peaks of Mount San- 
hedrin, Mount Linn, and Yolla Bolly. Merriam is quoted 
concerning a specimen killed ^Jbout 1893 between Mounts 
Shasta and Lassen. 

At the present time there are probably few wolverenes left in 
California. "From 1920 through 1924, the average number 
reported captured by the trappers of California was less than 
three a year. After considering all the available data, the 
authors [Grinnell, Dixon, and Linsdale, 1937] believe that at 
the present time (1933) there are at most not more than 15 
pairs of wolverenes left in the State. This animal is thus one 
of the rarest furbearers now living in California." "At present 
the main hope" of protecting the species from extinction 
within the State is its preservation within the guarded areas of 
Yosemite and Sequoia National Parks, but the animals are 
likely at times to wander outside the limits of these areas and 
are caught or killed by trappers who may legally take fur out- 
side the park boundaries. 

How far north the characters of the southern wolverene may 
hold is tentatively intimated by the above authors as perhaps 
extending to the State of Washington. Bailey, in 1936, be- 
lieved that they are not yet extinct in the Cascades and 
Sierra Nevada and mentions one taken in the upper McKenzie 
Valley west of the Three Sisters Peaks, Oregon, in 1912; how- 


ever, at the present time their numbers must be few indeed. 
Still later, Scheffer (1938) notes that the U. S. Forest Service 
estimates that in the national-forest areas of the State of 
Washington, the wolverene population is about 20. 

Family CANIDAE: Wolves, Dogs, Foxes 

These small foxes are about half the size of a red fox and 
live in the open plains country or sandy deserts of western 
North America. The upper side of head and body is some 
shade of buff overlain with gray; the sides of neck, shoulders, 
and body are clear buff, the ears and legs usually brighter buff, 
the tail gray with black-tipped hairs, which terminally con- 
centrate to form a black tip. Two species are found, the more 
eastern, ranging over the Great Plains, Vulpes velox, and the 
more western, centering in the Great Basin and southern 
California, V. macrotis. The former is a slightly larger animal 
with shorter ears than the latter, and comprises two races; the 
latter is more an animal of sandy desert regions and is at 
present divided into eight races. 



Cam's velox Say, Long's Exped. to Rocky Mts., vol. 1, p. 487, 1823 (South Platte River 

(? Logan County), Colorado). 
SYNONYM: Canis microtus Reichenbach, Regnum animale . . . , vol. 1, pt. 1, p. 10, 

1836, as quoted by Wiegmann in Wiegmann's Archiv fUr Naturg., Jahrg. 3, Bd. 

2, p. 162, 1837. 
FIGS.: Audubon and Bachman, 1851, The Viviparous Quadrupeds of North America, 

vol. 2, pi. 13. 


Vulpes velox hebes Merriam, Proc. Biol. Soc. Washington, vol. 15, p. 73, Mar. 22, 1902 
(Calgary, Alberta, Canada). 

The range of the typical race is from the Staked Plains of 
northwestern Texas northward over the Great Plains to prob- 
ably South Dakota. The northern race, V. v. hebes, is larger 
and slightly grayer, "dark patches on sides of nose darker, 
skull larger and heavier; palate much longer; under jaw longer, 
heavier, more bellied under sectorial tooth" (Merriam). 


Average of four males: Total length, 844 mm., tail, 280; foot, 
131; ear, about 50. Weight, 4.5 pounds. 

"These little foxes," Bailey (1931) writes, "live in burrows 
in the open plains country, usually selecting side hills or the 
sunny slope of a bank in which to make their dens. They are 
mainly nocturnal, are wonderfully swift and graceful in their 
motions, but, unlike the red foxes, are so unsuspecting in their 
natures as to be readily caught in traps . . . and are so 
unable to cope with the advanced civilization that they are 
rapidly disappearing from the face of the earth." Gary in 1902 
was told that in the Staked Plains of Texas they were at that 
time scarce in comparison with former years, for in the winter 
they come readily to poisoned bait put out for coyotes and 
wolves. This bait the kit foxes will take the first night it is 
set out, but coyotes wait till later (V. Bailey, 1905). Formerly 
this was a common species over the plains of eastern Colorado, 
but by 1911 had "become rare in most sections" (Gary, 1911). 
It is found still in the western part of Kansas and has been 
taken as far east as Douglas County (Hibbard, 1933). In 
1870 Dr. J. A. Allen reported its occurrence "more or less 
frequently" in western Iowa, but at the present time it is 
probably gone from this eastern outpost of its range. 

It is probable that the range of the northern race may be con- 
sidered as from about North Dakota north to the Saskatche- 
wan River. Bailey (1926) refers those of North Dakota to this 
race and states that "apparently the kit foxes of the northern 
Plains at one time covered the whole of the prairies of North 
Dakota, but at present they are restricted to the western part 
of the State, and even there they have become very scarce, 
although a century ago they were one of the common fur 
animals of the Red River Valley." With the settling of the 
country they have easily succumbed to trapping, poisoning, 
and capture by dogs, and reports from various localities in that 
State indicate that of recent years they have been reduced 
nearly to the point of extermination. In northwestern Mon- 
tana, Bailey (1918) speaks of this fox as common over the 
plains along the eastern edge of Glacier National Park, which 
it probably enters in the open area at the lower end of St. 
Mary Lake. There is thus apparently no immediate danger 
of its extermination in this region, yet on account of its less 
wary nature it is likely to suffer gradual reduction with the 


encroachment of settlements and especially with trapping and 
poisoning campaigns carried on against coyotes or wolves. 
Baird records the usual stomach contents as "fragments of skin, 
leather with hair, berries, remnants of mice, and grasshoppers" 
so that economically the kit fox is an asset through its destruc- 
tion of small rodents and grasshoppers and deserves whatever 
protection may be accorded it. 

This race of the kit fox formerly extended in range as far 
north as the Saskatchewan River in western Canada but is 
now gone from many parts of this area. Specimens in the 
Museum of Comparative Zoology from Calgary and Buffalo 
Lake, Alberta, were received in 1897, and others from Assini- 
boia were obtained in the following year. Fowler (1937), 
however, writing of changes in the fauna of the High River 
district, Alberta, states that it became extinct there shortly 
after those dates; "in fact, only those people who were here 
before 1900 can recall having seen a kit-fox. My father states 
that the last bunch of kit-foxes he saw was in the summer of 
1897. These had their den in a sandy knoll about a mile 
north of High River." 



Vulpes macrotis Merriam, Proc. Biol. Soc. Washington, vol. 4, p. 136, Feb. 18, 1888 
("Riverside, Riverside County, California," vicinity of Box Springs, western 
margin of San Jacinto Plain, about 10 miles southeast of Riverside). 

FIGS. : Grinnell, Dixon, and Linsdale, 1937, pi. 9, bottom fig. in color; and (other races) 
figs. 154-162 (skull, live animal, habitat). 

The long-eared kit fox resembles V. velox but has longer ears 
(about 80 mm.), smaller hind foot, and slightly brighter color- 
ing, but otherwise it has a similar appearance. As a species its 
range extends from Riverside County in southern California 
eastward across the Great Basin into extreme western Texas 
and south into Lower California and northern Mexico. Over 
this range it divides into a number of slightly marked local 
races, of which no less than nine, including the typical form, 
are described. These with their type localities are: arizonensis, 
from 2 miles south of Tule Tanks near the Mexican border of 
Yuma County, Ariz.; arsipus, from Daggett, San Bernardino 
County, Calif.; devia, from Llano de Yrais, opposite Magdalena 
Island, Lower California; mutica, from Tracy, San Joaquin 


County, Calif.; neomexicana, from San Andreas Range, Dona 
Ana County, N. Mex., about 50 miles north of El Paso; 
nevadensis, from Willow Creek Ranch, near Jungo, Humboldt 
County, Nev.; tenuirostris, from Trinidad Valley, northwest 
base of San Pedro Martir Mountains, Lower California, 
Mexico; and zinseri, from San Antonio de Jaral, southeastern 
Coahuila, Mexico. In a recent study of the group, Benson 
(1938) shows that arizonensis is synonymous with the older- 
described race arsipus, so that the number of recognized races 
is now eight. This author writes that "kit foxes are now as a 
rule few in number even in the areas which seem suitable to 
them. They are so unsuspicious that they are easily trapped 
and even more easily poisoned. Consequently, wherever 
trappers are active, and especially wherever control campaigns 
involving the use of poison have been carried out against 
predatory animals on areas inhabited by kit foxes, the foxes 
have been greatly reduced in number or entirely eliminated. 
Unfortunately, there are few kit fox habitats in the United 
States that escape visitation by these agencies." Benson 
comments that the race mutica pf the San Joaquin Valley and 
Walker Basin in California is a well-marked form, while the 
race arsipus, ranging east of the Pacific slope drainage, though 
also easily distinguishable, clearly extends into southern 
Nevada and may have to include nevadensis as a synonym. 

The numbers of all these various races seem to be rapidly 
decreasing, but the only one which is rather certainly now 
extinct is the typical one, V. macrotis macrotis. This form 
once ranged over parts of California in the "San Diegan sub- 
faunal district, from Riverside County, northwest ... to 
Los Angeles County" in the Lower Sonoran Life Zone. It 
formerly occurred in the Allessandro, Perris, and San Jacinto 
Valleys, westward along the plain at the southwest of the San 
Gabriel Range to San Fernando Valley, where in the early 
nineties local residents trapped them. The last one known to 
have been taken in the San Jacinto Plain was trapped near 
Moreno in 1903 (Grinnell, Dixon, and Linsdale, 1937). At the 
present time the race mutica is still to be found in the San 
Joaquin Valley of California, but the authors just quoted 
indicate that its range has shrunk to about half of what it was 
not so many years ago, so that it is now restricted to the driest 
plains of the southern and western parts of the valley. 


"Open level sandy ground is the preferred habitat of the 
kit fox," for here it can easily excavate its burrows, and this 
type of ground is likewise preferred by numerous small rodents, 
such as kangaroo rats and pocket mice, which form the chief 
food of this little fox. " These special adaptations may account 
for the rather sharp limitation observable in the general dis- 
tribution of this type of fox, which man's presence further 
restricts." Their shallow burrows are dug in open ground and 
may have one to four entrances. Grinnell, Dixon, and Lins- 
dale (1937), from whose account these details are taken, found 
that the breeding season is early. Young are found newly born 
late in February, and the number to the litter may average four 
or five but in one case was as many as seven. It seems to have 
few natural enemies. There is a close correlation between the 
areas inhabited by the kit fox and the range of the large kanga- 
roo rats Dipodomys deserti and D. spectabilis, on which with 
other smaller rodents the fox largely feeds, although insects, 
occasionally rabbits and snakes, and rarely poultry are also 
items in its diet. 

Of the race mutica of the San Joaquin Valley, Calif., the 
authors quoted say that of recent years large numbers have 
been killed for fur. Thus, in 1919, one trapper caught 100 
foxes in a single week on an area 20 miles long by 2 miles wide 
on the west side of the valley. But "probably the greatest 
number of kit foxes has been destroyed by poison campaigns 
directed against coyotes . . . Hundreds of kit foxes were 

Long-eared kit fox (Vulpes macrotis macrotis) 


doubtless thus destroyed in a single season." Some unusual 
condition added to these efforts at "control" might conceivably 
wipe out the remnant of these foxes in the San Joaquin Valley. 
However, "in the Mohave and Colorado deserts, which are now 
remote from human activity and where the conditions of the 
soil and the lack of water will long prevent future settlement, 
it is probable that" the race arsipus will endure indefinitely. 

The most northerly ranging race of the desert fox is neva- 
densis, concerning which Vernon Bailey (1936) writes: "These 
graceful little foxes have been found in the valleys of northern 
Nevada, southwestern Idaho, and southeastern Oregon, not 
far beyond the limits of Lower Sonoran Zone valleys, where 
the species generally ranges." Specimens are mentioned from 
the Owyhee Valley as the only ones available from Oregon, 
but it "may be looked for in the Alvord and adjoining valleys 
that open out into northern Nevada." He says nothing of 
their present status there but surmises that they will soon be 
gone. He further testifies to their adaptability as pets, on 
account of their gentleness, their intelligence and affectionate 
behavior, and their ready confidence and alertness. "Why 
not," he asks, "keep such animals instead of cats and dogs and 
save a few from extermination?" 


Wolves are of boreal distribution in both the Old World and 
the New, originally ranging without interruption from western 
Europe eastward to the Pacific Ocean in the former and from 
Alaska to the Atlantic coast of eastern North America in the 
latter area. Pocock (1935) in a recent paper regards them as 
of a single species with local races, which appears to be a 
logical course, since all are much alike. Even the narrow 
Bering Straits, closed by ice during part of the year, probably 
form no insuperable barrier to an occasional crossing. In 
North America wolves are found from the Arctic regions 
southward to the Gulf of Mexico and the Mexican tableland, 
but in a recent review Goldman (1937) regards the most 
southern or red wolf as a separate species. Exclusive of this 
and its two subspecies, there are at present recognized no less 
than 20 local races in this area, distinguished for the most part 
by minor characters of the skull and teeth; all these are treated 
as subspecies of Canis lupus of Europe. It is often said that 


the European wolf is the ancestor of the domestic dog, but the 
evidence for this is merely inferential. Usually wolves of any 
race may be distinguished from even the largest dogs by their 
proportionally larger teeth, shorter ears, and less elevated 
forehead due to the smaller development of the frontal sinuses. 
In dogs the length of the lower first molar or carnassial tooth 
is usually less (and in medium-sized and smaller breeds much 
less) than 22 mm., whereas in wolves it is in excess of this 
measurement. In their general proportions and a certain 
"plumpness," however, the teeth of wolves are exceedingly 
like those of domestic dogs and differ from the narrower more 
bladelike teeth of coyotes, which some have thought ancestral 
to dogs. 

Everywhere in North America that wolves have come into 
competition with white men, in settlements or in agricultural 
communities, they have been regarded as a common enemy 
and killed whenever possible. The inevitable result has been 
their greater or less reduction in the thinly settled regions and 
complete extermination from larger areas where white popula- 
tion is denser or agricultural and pastoral use of the land makes 
it impossible to tolerate their presence on account of their 
depredations against livestock. While this is more or less to be 
deplored, it is unavoidable, for the rancher can not be expected 
to view with complacency the resulting loss. The outlook for 
the preservation of a remnant of this interesting species is 
therefore bright only in the wilder and more inaccessible 
regions or in large public reservations where the numbers can 
be controlled and where, conceivably, a small number of 
wolves might serve a useful purpose not only in preventing 
too great an increase in wild deer or mountain sheep but also 
in actually improving or keeping up a standard of quality by 
eliminating sickly or stupid individuals. It is remarkable that 
even in parts of Europe to this day as in Spain and France a 
few wolves have remained in spite of long occupation by 
white races. Furthermore, if one may believe the mass of 
published accounts, wolves under the stress of hunger have 
been known to pursue and kill human beings in Europe and 
Asia, whereas at least in eastern North America there seem to 
be few or no authentic cases of such bold actions on the part of 
wolves here in early days, although there are instances of 
wolves having followed and frightened many of our early 


Like other large carnivores, wolves require food in larger 
amounts, so that in addition to what smaller game they may 
pick up, it is the larger ungulates that they specially hunt. 
In the North the caribou, and on the plains the mule deer, and 
even the young or sickly bison were objects of pursuit, while in 
the wooded regions of eastern North America the white-tailed 
deer was a staple food species. In the case of the last-named 
the wolf in areas of deep winter snows was doubtless a main 
factor in preventing the spread of this deer to the northward 
into regions where now it is abundant. 

In recent years a more intensive campaign has been carried 
on under the U. S. Biological Survey to eliminate the wolf 
entirely from stock-raising regions of the West. This has 
resulted in the accumulation by the Survey of sufficient speci- 
mens to "afford a fairly satisfactory basis for determining 
specific and subspecific relationships" of the American races, 
as worked out in Goldman's (1937) paper. Some of these are 
already extinct, others nearly so. In the following brief survey, 
the races are taken up alphabetically, giving a condensed 
statement of the general status so far as at present known. A 
final account of the genus and its natural history is to be 

In a general way, the various races of wolves are much alike, 
about the size of a modern "police dog" but shaggier, with 
short ears and bushy tail, carried usually with a slight hump 
near the root. The usual color is a mixed black, white, and 
gray, giving a grizzled appearance. The long hairs of the coat 
are usually white for the greater part of their length, with 
more or less black tipping or white, producing this mixed 
appearance. Often the white portion of the hairs is greater 
than the black tipping, resulting in a pale coat, and in the 
Arctic parts of the range the black may be eliminated almost 
altogether, so that the pelage is white. On the other hand, 
black may predominate and the pelage in that case is blackish 
to entirely black. This wide variation may occur within the 
same family group of animals, so that color characters are 
often of little avail for the separation of geographic races. 
These are therefore in large part based on cranial characters 
and on size differences where they are significant. While 
wolves must often travel over considerable areas in their 
hunting, and thus tend to segregate less than more sedentary 


species, they must nevertheless keep to more or less definite 
regions where under varying conditions of environment slight 
differences have developed. Intergradation through individual 
variation often makes the exact definition of races difficult, and 
series are needed in working out the status of animals from 
different parts of the range. 

According to a useful pamphlet by Vernon Bailey (1907) 
wolves enter very little into the national forests of the western 
United States but favor foothill regions for breeding and 
hunting. In some areas there is evidence of a certain amount of 
seasonal migration, in the course of which the wolves of a dis- 
trict follow the cattle herds into the mountains in spring, and 
again in autumn accompany them into lowland areas, much as 
in former times they followed the bison. The same paper has 
some account of the abundance of wolves in the West and 
South up to the early years of this century, with instructions 
for trapping and poisoning. 



Canis lupus arctos Pocock, Proc. Zool. Soc. London, 1935, p. 682, Sept. 12 ("Melville 

Island, Arctic America"). 
FIG.: Nelson, 1916, p. 421. 

According to Pocock the skull of a specimen of Arctic wolf 
from Melville Island "may be distinguished at a glance by the 
great elevation of the frontal region and the resulting much 
more strongly convex curvature of the frontorostral line. It is 
also broader . . . in the muzzle and palate . . . The 
bulla also is noticeably more inflated" as compared with a 
skull regarded as representing C. I. tundrarum. This is a 
white wolf. 

In addition to the skull of the type from Melville Island, 
Pocock lists a second in the British Museum from Discovery 
Bay, Ellesmere Land. The former was secured in 1853-54, the 
second about 1878. Parry seems to have been the first to men- 
tion this wolf, which he noticed was smaller than the white 
wolf of the mainland (tundrarum) . 

The present status of this wolf in Ellesmere Land and ad- 
joining islands to the west is uncertain. There are few human 
inhabitants, either Eskimo or white. However, since it would 


be expected that the numbers of caribou in the region would 
attract a small permanent population of wolves, it seems 
strange that they should be so rarely observed. Peary, who 
was familiar with the region from his various polar expeditions, 
wrote ("The North Pole," p. 61, 1910) that in the northern 
part of the island there might be "perhaps once in a generation 
a stray wolf," implying that it was almost unknown a genera- 
tion ago. Had it been of regular occurrence following the 
caribou herds, the Eskimo would surely have reported it. 
Possibly these stray Arctic wolves may have come from farther 
south on rare occasions. However, the more recent testimony 
of Macmillan is rather at variance with Peary's statements. 
In his book "Four Years in the White North" (1918) he has a 
good deal to say about the Arctic wolves met with in Ellesmere 
Land and Axel Heiberg Land. Late in March, 1916, two were 
started at Bay Fiord, near Etah, but they dashed away and 
finally disappeared over the rough ice of Smith Sound; later 
one was killed near Eureka Sound, and another was started 
from the ice foot and also ran off over the ice. "Axel Heiberg 
Land is infested with them," he^wrote, after seeing a pack of 12, 
all white. In April of that year two others appeared and when 
pursued by the sledge dogs ran off over the sea ice toward 
North Cornwall. In October, 1916, the local Eskimos returned 
from their annual caribou hunt, which covers the region from 
Etah to the Humboldt Glacier on the Greenland coast. They 
reported no young caribou, but tracks of wolves everywhere, 
implying that the young caribou find it difficult to survive on 
account of the wolves. Macmillan believes that a large band of 
white wolves had crossed Smith Sound and was following the 
herds on the North Greenland coast. They also probably 
capture a few seals along the ice foot, which would account for 
their presence in that place, while muskoxen too must fre- 
quently fall a prey. If this explorer is correct in his belief that 
wolves cross Smith Sound, from Ellesmere Land to North 
Greenland, as seems likely, there appears no reason to suppose 
that their wanderings in search of game might not take them 
across the northern part of that country or even to the east 
coast, so that the supposed distinction of the wolves of the 
two regions may prove to be unfounded. 


CANIS LUPUS BAILEYI Nelson and Goldman 

Canis nubilus baileyi Nelson and Goldman, Journ. Mamm., vol. 10, p. 165, May, 1929 
("Colonia Garcia (about 60 miles southwest of Casas Grandes), Chihuahua, 

FIG.: Bailey, 1931, pi. 18 (photograph). 

This is the wolf of "southern and western Arizona, southern 
New Mexico, and the Sierra Madre and adjoining tableland of 
Mexico as far south, at least, as southern Durango," and ac- 
cording to its describers it is most nearly allied to the plains 
wolf (C. I. nubilus) but is smaller and darker, more brownish or 
rufescent, with a more slender and depressed rostrum. Its 
greater size and other cranial characters indicate "no close 
relationship" to C. rufus. Total length, 1,570 mm.; tail, 410; 
skull, condylobasal length, 262. 

The authors quoted list various specimens from Chihuahua, 
one from Sonora, and one from Durango, Mexico, as well as a 
number from Arizona and New Mexico. "According to old 
residents wolves that formerly inhabited the southern end of 
the tableland, near the valley of Mexico, became extinct many 
years ago. Wolves are still numerous, however, in the Sierra 
Madre as far south at least as Durango." Specimens from 
southwestern New Mexico show indications of intergradation 
toward the plains wolf (Bailey, 1931, p. 303). "Their greatest 
abundance has long been in the open grazing country of the 
Gila National Forest" in the open yellow-pine forests and the 
orchardlike growth of juniper, nut pine, and oak. "Apparently 
they are not known in the Lower Sonoran valleys of this 
region." Bailey (1931) has given an excellent account of the 
habits of wolves in this area. He states that they persistently 
cling to certain localities in their range for raising their young. 
"Dens are always placed in the most out-of-the-way places, 
with seemingly little regard for convenience as to water or 
food." In rough country the dens are generally in a crevice or 
natural cavity "usually in the rim of a mesa"; badger holes 
are often enlarged, or the burrow may be worked out beneath 
a mesquite or other bush that serves as a blind. 

Bailey (1931) tells further of trapping wolves in this region 
and records that J. S. Ligon with a corps of trappers began a 
trap line in 1916 for the purpose of reducing the number of 


wolves. He estimated that in May, 1917, there were 103 
adult wolves in New Mexico and 45 in July, 1918. "In 1927 
he said that they were practically eliminated from New Mexico 
and that he was then concerned only with reinfestation from 
Mexico." Thus it appears that continued and skillful effort 
will at length succeed in exterminating even these wily animals, 
but that with their tendency to range wide and far, there is a 
constant opportunity for new individuals to come in from 


CANIS LUPUS BEOTHUCUS G. M. Allen and T. Barbour 

Cam's lupus beothucus G. M. Allen and T. Barbour, Journ. Mamm., vol. 18, p. 230, 

May, 1937 ("Newfoundland"). 
FIG.: Allen and Barbour, 1937, p. 231, fig. (upper carnassial). 

In the Newfoundland wolf the skull is relatively slender, 
and the nasal bones are long in contrast to the short ascending 
branches of the premaxillaries ; the upper carnassial is char- 
acteristic in its shortness and in the size and sharp inward turn 
of the anterointernal cusp in contrast to the condition in other 
wolves. The color is white, but occasional black individuals 
are said to have been known. The largest of four skulls 
measured in greatest length, 276 mm.; outer length of nasals, 
103; tip of nasal to point of premaxillary, 47; length of upper 
carnassial, 25.5; length of first lower molar, 28.7. 

In the large size and slender proportions of the skull, as well 
as in its white coat, this wolf resembled the white Arctic wolf. 
At the present time it is apparently extinct in Newfoundland, 
where in former days it followed the herds of Newfoundland 
caribou. Bonnycastle in 1842 spoke of it as large in size and 
as frequently seen in the neighborhood of St. John's. On 
account of its depredations against young cattle, a bounty was 
declared on wolves. A writer in Forest and Stream (vol. 4, p. 
390) in 1875 spoke of these wolves as then common in the 
island, often prowling near the houses of settlers or seen near 
settlements in chase. In winter when the caribou were feeding 
in the bogs, the wolves cooperated in hunting them. While 
some lay in wait along the border of coniferous trees, others 
went around to windward, when the caribou, scenting them, 
turned and ran directly into the ambush. The same writer 
mentions an instance where a Micmac Indian in April, 1866, 


was standing by the side of a small lake formed by the River 
Exploits, when he saw an old wolf coming toward him on the 
ice, to be followed presently by five or six more. The Indian 
ran for his "tilt" and his gun, but the wolves, following him, 
gained so rapidly that he sought safety by climbing into a tree 
beyond their reach; otherwise, he felt certain, they would have 
attacked and killed him. But after remaining about for nearly 
an hour they departed. The skin of a white wolf killed north 
of Grand Lake, Newfoundland, about 1896, by Dr. El wood 
Worcester, was presented by him to the Museum of Compara- 
tive Zoology. It was shot as it came over the side of a hill 
where Dr. Worcester was waiting, gun in hand, for a chance at 
a caribou. Two others, which he saw at about the same time, 
were running along the shore of a lake above River of Ponds 
in pursuit of a caribou. One of these looked very dark in 
color. Bangs (1913) wrote that when Doane was collecting for 
him about 1894 one was killed, but neither Millais, writing 
about 1906, nor Dugmore in his book on the Newfoundland 
caribou in 1913, mentions seeing anything of the wolf, which 
by that time must have been greatly reduced in numbers. 
The last record known to me is of one reported to have been 
killed about 1911. The late Dr. John C. Phillips, who con- 
tributed this note, endeavored to obtain the specimen but was 

While Newfoundland at its nearest point is hardly 10 miles 
distant from the mainland of Labrador, it seems that this wolf, 
like so many other animals of the island, must have been 
isolated here for a long period and gradually developed the 
slight cranial differences that distinguish it. At the present 
time the Newfoundland wolf is believed to be extinct. 


Canis occidentalis crassodon Hall, Univ. California Publ. Zool., vol. 38, p. 420, Nov. 
8, 1932 ("Tahsis Canal, Nootka Sound, Vancouver Island, British Columbia"). 

Skulls of the Vancouver Island wolf differ from those of the 
adjacent mainland and southern Alaska in their smaller size, 
shortened preorbital region, less elevated tip of rostrum, and 
greater shallowness of the lower jaw through the coronoid 
process. The color is gray to black. Condylobasal length of 
skull, 240 mm. 


Five specimens are mentioned by Hall in his original descrip- 
tion of this wolf. Concerning the status of the race, Swarth 
(1912) writes that a quarter of a century ago wolves were still 
fairly common "in the wilder parts of at least the northern 
two-thirds of Vancouver Island, sufficiently so as to be a serious 
menace to the deer in many places." At Beaver Creek he 
heard one howl and was told that they were occasionally seen 
in the valley during the winter but very seldom in summer. 
At Nootka they were said to be abundant. "On the Tahsis 
Canal we stayed at the camp of a trapper who made it his 
principal occupation during the winter to hunt wolves and 
panthers." With the bounty of $15 a head added to the value 
of the fur, the trapper evidently secured enough to make it pay. 
This trapper used poison exclusively and seemed to have had 
no trouble in killing the animals, but in several years' hunting 
he had actually seen but two or three alive, so cunning are they 
in keeping out of sight. Swarth stated that wolves had been 
so common at Friendly Cove during the years immediately 
preceding 1912 that deer had been almost completely driven 
out of the neighborhood. Of the status of this wolf at the 
present day no further information is at hand. 


Canis lupus var. fusca Richardson, Zool. Beechey's Voyage of the Blossom, Mammals, 
p. 5, 1839 ("California and the Banks of the Columbia River"). 

SYNONYM: Lupus gigas Townsend, Journ. Acad. Nat. ScS., Philadelphia, ser. 1, vol. 2, 
p. 75, Nov., 1850 (near Vancouver, Columbia River, Washington). 

The wolf formerly occurring from northern California to the 
Puget Sound region, and probably southwestern British 
Columbia west of the Cascades, was regarded as a valid form 
by Miller (1912) in notes on the names of the large wolves of 
North America. He states that a skull from the Puget Sound 
region "indicates an animal differing from the timber- wolf of 
the interior region in less great size and in less enlarged teeth." 
Hall (1932) contrasts specimens from Oregon with the race 
crassodon of Vancouver Island and says that they differ 
noticeably in "the markedly less inflated tympanic bullae, 
much smaller teeth," in the more convex lower profile of the 
jaw and other minor skull characters. The color is "much 
darker" than in the Plains wolf (Bailey). 


This race, which was perhaps characteristic of the saturate 
region of forest along the Northwest coast, is doubtless now 
extinct in California (Grinnell, 1933). In Oregon, Bailey 
(1936) writes, "a few of these large, dark gray wolves are still 
found in the timbered country west of the Cascades . 
and locally northward to British Columbia and Alaska. In 
recent years they have been found mainly along the west 
slope of the Cascade Range, but before extensive white settle- 
ments were made in the State they seem to have been common 
in the Willamette Valley and west to the coast ... In 
1897 Captain Applegate reported them as formerly common, 
but at that time extremely rare in the southern Cascade 
region." In 1913-14 bounties were paid on 30 wolves taken in 
Oregon, but in later years they seem to have been largely 
exterminated within the State. Bailey mentions one taken in 
1930 on the Umpqua National Forest, where it had killed 
several sheep. It was very old with much worn teeth. A 
second old male was killed the same winter in Klamath County, 
while in Douglas County and Lane County one each was killed 
in 1930 and 1931. At the present time, he writes, their num- 
bers are so nearly under control that their damage is negligible. 
This probably applies also to Washington, but no recent 
statistics are available. In the remoter regions of western 
British Columbia they are probably still present in some 



Canis lupus irremotus Goldman, Journ. Mamm., vol. 18, p. 41, Feb., 1937 ("Red 
Lodge, Carbon County, southwestern Montana"). 

This is the wolf of the northern Rocky Mountain region and 
high adjoining plains, from northwestern Wyoming northward 
through western Montana and eastern Idaho at least to Leth- 
bridge in southern Alberta. Intergradation with neighboring 
races is presumed and makes the precise definition of the 
range difficult. It is a light-colored race with narrow but 
flattened frontal region, in these characters differing from the 
race youngi of the southern Rocky Mountains. It is larger 
and paler than nubilus of Nebraska or fuscus of Washington, 
and smaller than the form occidentalis to the northward of its 


The race is based on more than 30 specimens from Montana 
and Idaho; hence its characters seem well established. Prob- 
ably this is the form of eastern Oregon concerning which 
Bailey (1936) has a few interesting notes (under Canis lycaon 
nubilus). In this region wolves were numerous in the middle 
of the last century and were plentiful even up till the nineties. 
At the present time they are largely gone from all this region, 
but doubtless still remain in small numbers in Montana and 



Canis lupus labradorius Goldman, Journ. Mamm., vol. 18, p. 38, Feb., 1937 ("Vicinity 
of Fort Chimo, Quebec" northern Labrador). 

In color this wolf of northern Labrador varies from "dark 
somewhat grizzly gray to almost white." It is larger than the 
eastern wolf, with heavier dentition and minor cranial differ- 
ences, and seems to be only slightly differentiated. Its geo- 
graphic range is not at present determined but is presumed to 
be the Labrador Peninsula north of the forest region. 

Concerning the status of these wolves in Labrador but little 
information is available. They are, however, not uncommon 
and follow the herds of barren-ground caribou. William B. 
Cabot, in his account of travels into the eastern interior of the 
peninsula, speaks of them as occasionally to be heard, and in 
1905 he saw tracks of perhaps as many as 200. They are 
occasionally killed by hunters near the coastal settlements in 
winter, but since the interior of the country is rarely pene- 
trated by white men it seems unlikely that they will be much 
disturbed there for a long period to come. 


Canis lupus ligoni Goldman, Journ. Mamm., vol. 18, p. 39, Feb., 1937 ("Head of 
Duncan Canal, Kupreanof Island, Alexander Archipelago, Alaska"). 

This wolf is believed to be the race characteristic of the 
Alexander Archipelago and adjacent mainland of southeastern 
Alaska. It is a race of medium size, "much smaller than Canis 
lupus pambasileus of the Mount McKinley region, Alaska, or 
Canis lupus occidentalis of the Fort Simpson region, . . . 


Mackenzie." It is similar to the race C. I. fuscus of the coast 
districts to the south, but a larger percentage of black or 
blackish individuals is normal, while it is less suffused with 
buffy in the gray phase. The small auditory bullae distinguish 
it from the race crassodon of Vancouver Island, while its longer 
rostrum and palate and the longer narrower nasals separate it 
from fuscus. None of these characteristics seems very trench- 
ant, taken alone, but Goldman considers it a "well-marked 
form," closest related to the last. 

Swarth (1936), commenting on the distribution of this wolf, 
says that it is "found upon the same islands as the Red Squirrel 
[i. e., Mitkof, Kupreanof, and Kuiu], and also upon the large 
Prince of Wales and Dall Islands with some others of this 
southern group that the squirrel has never reached. Evidently 
the wolf did not come directly from the north" but reached 
those of the inner islands that formed a series of stepping- 
stones most easily crossed from the adjacent southeastern 
coast. I have no recent information concerning its present 
status, but apparently it still occurs on these islands in small 



Canis lycaon Schreber, Saugthiere, vol. 3, p. 353, pi. 89, 1776 ("Vicinity of Quebec, 

Canada," as selected by Goldman, 1937, p. 37). 
FIGS.: Stone and Cram, 1902, pi. opposite p. 278; Nelson, 1916, p. 423, upper figs. 

(color phases). 

The eastern timber wolf is slightly smaller than the wolf of 
northern Labrador, and usually of a grizzled black, white, and 
gray. As long ago pointed out by Baird, it differs from the 
Plains wolf and the neighboring northern races in its weaker 
rostrum. Weight upwards of 100 pounds. 

Three hundred years ago this wolf ranged over the wooded 
and open areas of eastern North America from southern Que- 
bec to probably the southern States and westward to the 
Plains. With the coming of white men and the occupation of 
the country, it has been gradually extirpated from the settled 
regions until at the present time wolves are gone from practi- 
cally all the eastern United States and probably none are now 
to be found south of the St. Lawrence and east of Michigan. 

When the first English colonists landed on the shores of New 
England they found wolves plentiful and troublesome and were 


obliged at once to take active measures to prevent them from 
seriously depleting their precious cattle and sheep brought 
with so much labor from England. No doubt too they feared 
the attack of wolves in packs against themselves, particularly 
if traveling alone or at night, yet there seems to be no authentic 
instance of anyone actually being killed by wolves, though 
many a person going after dark to a neighbor's hurried his 
footsteps on hearing a wolf howl. Apparently the Indians 
along these coasts were little afraid of wolves (for they had no 
livestock), but occasionally they caught them in traps, and 
wolf bones in small amount occur in the Indian shell heaps. 
The early settlers often kept their sheep on the smaller outer 
islands where they were comparatively safe and unable to 
stray. But flocks or cattle in pasture were in constant danger 
even when in pens near the houses. The early records of 
Plymouth and Massachusetts Bay colonies show that a de- 
termined campaign was waged against wolves. Bounties were 
offered for adult wolves and lesser ones for whelps, and the 
heads of wolves brought in for the reward were publicly ex- 
posed on hooks at the meetinghouse. The Indians were active 
in this campaign and secureVl their share of the bounties. 
Yet for nearly a century wolves continued to be a menace to 
stock. In the "History of Cape Cod" Freeman relates that 
in 1717, the early settlers discussed a project for erecting a 
high fence of palisades or boards between Sandwich and Ware- 
ham to shut off the outer Cape completely as a safe pasture 
for the livestock against the wolves. But the scheme finally 
was abandoned, since the several towns involved could not 
agree on their proportionate share of the expense, while those 
outside the Cape did not see why the wolves should be retained 
on their side of the fence! Many of the towns maintained 
" wolf pits," dug deep enough so that a wolf could not jump out, 
the inwardly sloping sides sometimes smoothly faced with 
stone, and the whole baited with fresh meat. Wolves jumping 
in for the bait were unable to leap out and were later killed. 
The remains of such a trap are said still to exist in the Lynn 
Woods park in Massachusetts. The idea was doubtless taken 
from the Indians. 

Even down to the time of the American Revolution wolves 
were fairly frequent in or near the settled parts of New Eng- 
land, but soon after, the unremitting fight against them had so 


reduced their numbers that they were no longer a serious 
menace. Occasional small packs or single cunning survivors 
were systematically hunted down by companies of men who 
would track them to deep woods or swamps which were sur- 
rounded and the animals driven out and often shot. In 1774, 
Connecticut repealed the law that formerly offered a bounty 
on wolves. But if wolves were then no longer dangerous to 
stock in Connecticut, they continued to be so in the less settled 
regions of southern New Hampshire and Maine. Bounties 
were increased and hunting wolves became a regular occupa- 
tion of some of the Indians or white hunters. The constant 
war of extermination at length began to tell, and by the early 
years of the last century few wolves were left in southern New 
Hampshire and Vermont. In his history of the town of Gilsum, 
near the present Keene, N. H., Hay ward in 1879 tells that in 
March or April, 1828, took place what was then still remem- 
bered as "the wolf hunt," when a lone individual that had 
killed a number of sheep was finally hunted down, twice sur- 
rounded by a party of determined farmers, and shot as it slunk 
out from an old spruce top. Many similar tales may be found 
in the local histories. The "last wolf" in Lancaster, N. H., 
was trapped about 1840 according to local account, but I have 
reason to believe that an occasional one or two came into the 
White Mountain region till a much later date. Zadock Thomp- 
son mentions a specimen in the University of Vermont that 
was killed in Addison County about 1830, but probably they 
occasionally were found well after that time. Merriam in his 
"Mammals of the Adirondacks" (in 1882) lists wolves on 
which bounties had been paid that were taken in St. Lawrence 
and Washington Counties, N. Y., even up to the latter date. 
When the last wolf was taken in Maine is perhaps uncertain. 
The only specimen known to me from that State is a skin now 
in the Boston Museum of Natural History, killed near Moose- 
head Lake in 1863. The late Manly Hardy, who traveled all 
over Maine as a trapper and fur buyer, and whose authority 
is unquestioned, wrote in 1884 (Forest and Stream, vol. 22, p. 
141) that "in 1853 wolves were very plenty, and for the next 
five years were not scarce; plenty could be found within sixteen 
miles of Bangor in 1857 and 1858. They seemed to leave 
quite suddenly. The last I know of positively being taken was 
. . . in 1860, at Munsengun . . . There were rumors 


of occasional wolves being seen from 1875 to '80, but the first 
real proof I can give is that in 1880 I bought the skin of a 
freshly killed wolf, taken at Union River. The one who 
brought it said it had a mate, and they had been heard at 
times for several years in that vicinity." A few later reports 
of wolves or their tracks seen in northern Maine bordering 
New Brunswick may have been authentic, but evidently they 
were practically gone by that time, not only from Maine but 
from New Brunswick as well. Indeed, Chamberlain (1884) 
writes that although common until about 1860 in the latter 
region they had since "entirely disappeared." 

In the wilder regions of the Adirondacks of New York, 
wolves in small numbers continued till a much later time. 
Miller (1899) has published a table showing by counties the 
bounties paid on wolves by New York State, incorporating 
and continuing the earlier one of Merriam (1882). From this 
it appears that between 1871, when a bounty was declared, and 
1897, no less than 98 wolves were killed on which a total of 
$2,910 in bounties was paid. Further, in the decade preceding 
1880, six was the most killed in any year, but 12 were brought 
in in 1881 and no less than $1 in 1882! In 1883 nine were 
killed, in the next three years two in each, one in 1887, and 
two in 1888. Thereafter appears a blank in the record, and 
some have believed the animal was then extinct in New York, 
but in each of the years 1895, 1896, 1897, bounties were paid 
on six wolves! One can not help the thought that something 
is strange about this. Of the 98 wolves on which the State paid 
bounties between 1871 and 1897, 45 or nearly half came from 
St. Lawrence County, which borders the river of that name, 
suggesting that there may have been occasional immigrants 
from the Ontario side. For the Toronto region, however, 
Fleming, in 1913, writes as if they had long since been extinct 
and states that "according to the late Dr. Brodie, wolves 
occasionally drove deer into Markham as late as 1840." 

The status of the wolf in Pennsylvania and New Jersey has 
been carefully investigated by S. N. Rhoads, and the results 
of his search for records are summarized in his book on the 
"Mammals of Pennsylvania and New Jersey" (1903). From 
the latter State they were so early extirpated that little record 
remains to indicate for how long they were present there, but 
perhaps till early in the nineteenth century. Pennsylvania, 


however, with its mountains and extensive wilderness areas, 
continued to harbor them up to comparatively recent years. 
The last record for New Jersey is given as of one killed in 
Wayne County in 1887, which was driven in from New York 
State by dogs, but its ultimate source may be questionable. 
Wolves seem to have remained common in the less settled and 
wilder parts of Pennsylvania until about the middle of the last 
century. Thus, Rhoads quotes Warren as authority for the 
statement that "about the year 1845 wolves were abundant in 
Tomhickon Valley" while between 1808 and 1820 Lucerne 
County paid $2,872 in bounties for wolf scalps at about $5 
each. As many as 273 wolves were killed in one of these years. 
Another correspondent informed him that he saw many as late 
as 1857 on the head waters of Pine Creek and Sinnemahonig 
in Potter County, where the last one he knew of was killed 
about 1875. Tomkins (1931) records a "last" wolf near 
Williamsport in 1867 and one near Ralston, in 1872, in the 
north-central part of the State. Many records are quoted for 
the middle part of the nineteenth century, and Rhoads makes 
the summary statement that they finally became "apparently 
exterminated in Pennsylvania, within the last 10 or 15 years," 
that is, in the middle or late eighties. More recent reports 
lack authenticity, while of various cases investigated in which 
bounties had been paid in later years some proved to be 
deliberate frauds. In another it turned out that a wolf had 
been brought from the West and had later escaped. In Elk 
County, Pa., the last wolf is said to have been shot in 1891. 
Probably the actual extinction of native eastern wolves in the 
State may be set at about that date. A few seem to have re- 
mained still in West Virginia wildernesses, where, according to 
Fred E. Brooks, "what is supposed to have been the last gray 
wolf in West Virginia was killed in Randolph county by 
Stofer Hamrick in January 1900." 

Apparently wolves were found in North Carolina up to 
about the beginning of the present century. At all events, 
C. S. Brimley, writing in 1905, states that it was then "found 
sparingly throughout the mountains. Dr. Donald Wilson re- 
ports it from Graham County, Mr. R. W. Collett says it is 
growing very scarce in Cherokee, Mr. Fain says there are a 
very few in Buncombe [County]. Mr. H. H. Brimley, Curator 
of the State Museum, informs me it has been taken in recent 


years in Yancey, Caldwell and Watauga. " Dr. C. Hart Mer- 
riam states that during one of his visits to Roan Mountain, in 
1887 or 1892, a den of wolves was discovered and the young 
were captured. In the last quarter of the eighteenth century 
wolves were spoken of as abundant in South Carolina, but by 
Audubon's day they were much fewer. The last one known to 
have been killed in the State was shot in Berkeley County, 
near St. Stephens, between 1856 and 1860. E. B. Chamberlain, 
of the Charleston Museum, who kindly supplied this record, 
had reports of other wolves in the Santee section at about the 
same time. 

How far to the south the gray or timber wolf extended its 
range may never be certainly made out, for the black or 
Florida wolf of extreme southern United States is now regarded 
as a race of the species Canis rufus by Goldman in his recent 
review. Dr. Francis Harper (1927) refers to this form the 
wolves formerly found in Okefenokee Swamp, southern Georgia, 
where according to his investigations the last one killed was 
about 1908. 

West of New York State and the Allegheny Mountains, the 
timber wolf was formerly common and is believed to have 
intergraded with the Plains wolf in the more open region west 
of the Mississippi. The story of its original abundance and 
gradual extermination is much the same as elsewhere in the 
East, except that in the more remote and wilder regions, as in 
northern Wisconsin, some still remain. In northern Minnesota 
wolves were common till at least the late years of the last 
century. Herrick (1892) has an interesting note in his "Mam- 
mals of Minnesota" indicating, as in some other instances, an 
occasional influx of numbers in certain districts: "During the 
winter of 1884-85, wolves became very abundant and insolent 
in Wright county, and were seen about the outskirts of Monti- 
cello in broad daylight almost daily, though they were suffi- 
ciently wary to escape capture." Cory (1912) writes that this 
wolf was still "common in northern Wisconsin and possibly 
occurs occasionally in other parts of the State," but although 
stragglers may at times appear in Illinois, all efforts to secure a 
specimen have failed, and most of the reports of wolves prove 
on investigation to be based on coyotes. In Wisconsin, recent 
records are available from as far south as Buffalo County. 

In the region of the Ohio Valley wolves were formerly plenti- 


ful but with the settling of the country a century ago were 
soon reduced to innocuous numbers, and in Ohio had become 
very rare by 1838 and "nearly extinct" a decade later. Audu- 
bon, while living at Henderson, Ky., early in the second decade 
of that period found black wolves abundant. He recounts that 
one morning he found a black wolf in one of his wild-turkey 
pens and shot it in the act of devouring one of the birds of 
which it had already killed several. Whether these wolves 
were of the timber wolf type or the red wolf type is indetermi- 
nable. In Tennessee wolves were exterminated many years 
ago (Kellogg, 1939), and but few records exist. Merriam in 
1887 found that a few still were present in the Smoky Moun- 
tains and in middle Tennessee they were reported as occasion- 
ally found in Van Buren County. Late records of wolves 
killed, Kellogg gives as: A female and her pups at Waynesboro, 
about 1917; and one in 1919 on North Fork River, Cumberland 
County. In western Tennessee the last available record is 
given as about December 10, 1895, when two were killed near 
Brownsville, Haywood County. 

In the region to the southward of South Carolina and Ten- 
nessee, the black race of the red wolf is believed to be the form 

At the present time the timber wolf is still to be found in 
limited numbers in eastern Canada, as in the less settled por- 
tions of Quebec along the north shore of the Gulf of St. Law- 
rence, where near Matamek I was shown in 1929 the skin of a 
large one killed the winter before. Doubtless there are many 
more between the Gulf and James Bay. At least until recent 
years, wolves were present in the Algonquin National Park, 
while farther to the west, in Quetico National Park, along the 
international boundary from Lake of the Woods to the western 
end of Lake Superior, and in the adjacent Superior National 
Forest contiguous with it on the United States side in north- 
eastern Minnesota, Cahn (1937) writes that "wolves are still 
common" and cross back and forth freely between the two areas. 
Here numerous and spasmodic drives have been made to rid 
the area of these predators but have resulted mainly "in the 
killing of about everything except wolves, which usually elude 
most cleverly the clumsy efforts to trap them. Late in fall 
their call is to be heard frequently all over the Quetico, but in 
all my travels I have seen but 2 live wolves Their 


persecution is to be regretted, for they form a perfectly natural 
and normal check on other species." In this last statement is 
to be found the key to the preservation of the species in regions 
where it comes in contact with white men. Under the nearly 
natural conditions of a large national preserve for game, wolves 
in these northern regions will be useful in preventing too great 
an increase of deer, and to a less extent of moose, which in the 
absence of such natural predators may easily become unduly 
abundant to the impairment of their range as in the case of the 
Kaibab deer or those of the Murderers Creek Game Refuge in 
eastern Oregon. Even in such situations, however, the game 
ranger has a special problem in determining how many wolves 
may safely be allowed on refuge areas, and how best this num- 
ber may be maintained through occasional destruction of 
young or of adults that become too bold or range outside the 
preserve to kill cattle or sheep in adjacent districts. 


Canis lupus mogollonensis Goldman, Jouri?. Mamm., vol. 18, p. 43, Feb., 1937 ("S. A. 
Creek, 10 miles southwest of Luna, Catron County, New Mexico"). 

This is a "rather small, usually dark-colored subspecies. 
Similar in general to Canis lupus youngi of the Rocky Moun- 
tain region . . . but smaller and usually darker in color. 
Decidedly larger and usually lighter in color than Canis lupus 
bailey i of the Sierra Madre of Mexico. Closely allied to Canis 
lupus monslrabilis of Texas, but smaller" and the skull with 
"zygomata more widely spreading; frontal region less elevated 
and inflated; auditory bullae smaller." 

The distribution of this form is said by Goldman to include 
the Mogollon Plateau region of central Arizona, extending 
eastward through the Mogollon Mountains to the Sacramento 
Mountains of New Mexico, and though rather circumscribed 
this seems nevertheless to be well borne out by a series of 120 
specimens studied by its describer. Vernon Bailey (1931) 
writes that "in 1908 he was again over the Gila National Forest 
and Mogollon Mountain region and found the wolves still 
common." They were feeding on nothing but fresh meat of 
their own killing, for "stock was abundant, and they had no 
trouble in finding cattle of any age or condition they preferred, 


but they seemed to show little choice in their selection of beef. 
Cows, steers, or calves seemed to be killed indiscriminately as 
the wolves happened to come upon them when hungry. Cattle 
are killed throughout the year and seem to be preferred to 
deer, which are more nimble and not so easily caught. In the 
wolf droppings along the trails cattle hair was almost the only 
recognizable constituent, but occasionally some jack-rabbit fur 
could be detected. " 

On account of their constant depredations against stock, a 
determined warfare has been carried on in this region during 
recent years with the result that in 1937, Goldman writes that 
this form is now "nearly if not quite extinct." There is some 
evidence, however, that if the campaign of extermination is 
relaxed for a time, other wolves keep coming across the border 
from the adjacent parts of Mexico into southern New Mexico 
and Arizona, so that if in later years specimens are taken in the 
former range of the Mogollon Mountain wolf, they may be of 
some other race. 


Canis lupus monstrabilis Goldman, Journ. Mamm., vol. 18, p. 42, Feb., 1937 (" 10 miles 
south of Rankin, Upton County, Texas"). 

This is a slightly marked race averaging darker than the 
wolves to the north and with a more highly arched frontal 
region. Its range formerly included southern and western 
Texas and northeastern Mexico, but because of systematic 
persecution its numbers are at present greatly reduced. 

Vernon Bailey's (1905) account of this wolf furnishes much 
valuable information. At that time they were abundant in 
and about the Davis and Guadalupe Mountains and over the 
Staked Plains and open country east of the Pecos River. He 
writes: "When opportunity offers, the 'loafer' not only kills 
sheep but often kills a large number, apparently for the 
pleasure of killing. His regular and most serious depredations, 
however, are on the scattered and unguarded cattle of the 
range. Two or three wolves usually hunt together and some- 
times pull down a steer, but most of their meat is procured 
from yearlings or cows. Occasionally a colt is killed but not 
often. Where two or three wolves take up their residence on 


a ranch and kill one or more head of cattle almost every day, 
the ranchmen become so seriously alarmed that they frequently 
offer a reward of $50 or $100 apiece for scalps. " Bounties have 
been tried, and in some cases several smaller ranches combine 
to offer a large reward in addition to that paid by the county, 
but the danger is that the hunters are tempted "to save the 
breeding females and dig out the young each year for the 
bounty, thus making their business not only profitable but 
permanent. " A better method seems to be to employ profes- 
sional hunters who are well paid by the month to keep down 
wolves or other noxious animals. 



Canis nubilus Say, Long's Exped. Rocky Mountains, vol. 1, p. 160, 1823 (Engineer 

Cantonment, near present town of Blair, Nebraska). 
SYNONYM: Canis variabilis Wied, Reise in das Innere Nord-Amer., vol. 2, p. 95, 1841 

(Fort Clark, near Stanton, Mercer County, North Dakota). 

This was the wolf found on the Great Plains of the United 
States from the Dakotas, Nebraska, and Kansas west to the 
Rocky Mountains and eastward possibly to the western parts 
of Iowa and Missouri. Though not so pale as the races to the 
north, it was less dark than C. I. lycaon to the east. The skull 
was similar in size to that of the adjacent races monstrabilis 
and youngi, but the frontal region was more flattened than in 
the former, with wider rostrum. 

In the middle of the last century and earlier these wolves 
were common on the Central Plains, following the herds of 
bison and picking off young or disabled individuals, but at the 
present time they are extinct over the greater part of the area 
once occupied. With the pushing of the railroads out across the 
country, the coming of settlers, and the development of stock 
grazing, the buffalo were reduced or exterminated and the 
wolves for a time devoted their attention to cattle, with the 
result that the ranchers and hunters pursued them relentlessly 
until now only a few remain in the remoter regions, where the 
roughness or barrenness of the country makes agriculture im- 

Vernon Bailey (1926) has gathered together many interesting 
notes on their former presence in the Dakotas, where in the 


days of the early explorers they were abundant. With the 
passing of the buffalo, however, they were poisoned and 
trapped in such numbers that they rapidly disappeared from 
their old haunts. They remained abundant up to the time of 
Wied (1833) and of Audubon (1843), but by 1870 J. A. Allen 
wrote that in Iowa "although rather common twenty years 
since, they are now scarce, especially in the more settled dis- 
tricts." Coues says that in 1873 they did not appear to be 
numerous in the Dakotas, while by 1887 they were practically 
gone from most of the country across North Dakota. Bailey 
states that in 1913 they were numerous enough to be trouble- 
some in the Badlands along the Little Missouri, though rare 
elsewhere. A few were reported in 1916 west of Cannon Ball, 
and in 1922 one was killed near Fargo. At the time of writing, 
in 1926, Bailey believed there were still a few wolves left in the 
least-settled parts of the rough Badlands region west of the 

Dr. M. W. Lyon, Jr. (1936), records the former abundance of 
this wolf in Indiana. "About the middle of the last century 
they were practically exterminated although a few fairly au- 
thentic records date back as recently as 25 years ago," i. e., 
about 1911. The pioneer's method of hunting them in the 
Wabash Valley about 1830 was a sort of round-up, with lines 
of hunters converging on a central point. Often two to ten 
wolves would be taken in a day at these hunts, while the bounty 
provided an added stimulus for their destruction. 

In Kentucky this type of wolf may once have existed, but 
Funkhauser (1925) believes it doubtful if at the present time 
any are to be found in the State. On the other hand, it is 
surprising that in Missouri it still occurs in some numbers. 
In their recent survey of the game and fur animals of that 
State, Bennitt and Nagel (1937) believe that on the basis of 
careful investigation lately made "the maximum density of 
wolves is about one per 10 square miles and the minimum about 
one per township (36 square miles)." They estimate the total 
wolf population of Missouri as in the neighborhood of 3,500 
wolves, with the maximum concentration in the ten west- 
central counties south of the Missouri River (a portion may 
be C. rufus gregoryi). The State paid bounties on 249 wolves 
between July 1, 1923, and July 1, 1925, but it is likely that a 
good part of these were for large feral dogs of which there are 


many, perhaps exceeding the wolf population. Wolves and 
coyotes probably furnish a most effective check on the undue 
increase of woodchucks and ground squirrels, but the part 
played by each is uncertain. Wolves also kill some deer. 
With the diminution of good wolf cover in recent years, there 
has probably been a concentration of the animals proportion- 
ately where cover is good, and it is even believed that their 
numbers have shown an actual increase in later times. If they 
become really numerous, these authors observe, they must be 
controlled, but under usual wild conditions they are a natural 
element of the fauna and their presence may even have a bene- 
ficial effect on the species that they prey upon in improving 
the quality of the race through their elimination of the less 
able animals. 

Farther west, the Plains wolf is apparently quite gone from 
Kansas (Hibbard, 1933), though it was common in that State 
in the sixties or later, and the same is probably true for Ne- 
braska where they remained in some numbers till about 1890. 
In Montana, Bailey (1918) writing of the fauna of Glacier 
National Park, says: "Along the eastern edge of the park, in 
the open country, are still founS the large, light-colored plains 
wolves . . . and a few occasionally range up through the 
valleys and over the high parts of the mountains . . . 
Along the trails in the Belly River valley in August, 1917, I 
saw wolf signs that were not very old. The wolves had evi- 
dently been down in the cattle country as the sign was com- 
posed mostly of cattle hair. Don Stevenson reports them in 
the country about Chief Mountain for the past 20 years, where 
they have ranged up and down the edge of the Plains killing 
cattle and some horses, and in 1914 he saw their tracks on St. 
Mary Ridge at the park line. There are said to be some in the 
North Fork valley, where it is probable they are attracted by 
the abundance of deer, as they are on the eastern border by the 
abundance of stock. In 1895 they seemed to be no more 
common than at the present time, as I saw then only a few 
tracks on the prairie below St. Mary Lake, and some fine skins 
among the Indians on the Blackfeet Reservation. As the 
valleys settle up, more vigorous hunting and trapping is likely 
to crowd the wolves back into the park at any time and make 
them more numerous than they are at present. If so, their 
destruction of game will be correspondingly increased * . . 


If unmolested they seem to prefer the domestic stock where it 
is abundant and easily accessible. " With the demands of the 
cattle interests for their destruction, their only possibility of 
long surviving is in large preserved areas such as the national 



Canis lupus occidentalis Richardson, Fauna Boreali-Americana, vol. 1, p. 60, 1829 

(Simpson, near mouth of Liard River, Mackenzie). 
SYNONYM: Canis lupus griseus Sabine, Franklin's Narr. Journ. Polar Sea, p. 654, 1823 

(Cumberland House, Keewatin, Canada). 

This is the wolf of the northern interior plains east of the 
Rocky Mountains and north of the United States, northward 
to the Arctic tundra. It is still paler than the Plains wolf, 
C. I. nubilus, and larger. 

Writing of the Athabaska-Mackenzie region in 1908, Preble 
says: "Gray or Timber wolves are found throughout the 
wooded parts of the region, and are fairly abundant and ap- 
parently increasing in some sections. In 1901 we saw numerous 
skins at nearly all the posts visited, and found a skull at a 
trapper's cabin on Slave River, 10 miles below the mouth of 
the Peace. Among a number of skins seen at Fort Rae, most 
of which were in the normal or gray phase, was one the color of 
which was mainly dark bluish gray, the throat and back were 
nearly black, the latter flecked with a few white hairs; the chest 
had a white patch; the belly and tail were bluish gray, the 
latter blackish toward the tip. 

"During the season of 1903 we heard that wolves had been 
rather abundant for several years past in the region west of 
Smith Landing, in the Birch Mountains, and in the vicinity of 
Athabasca Landing." In the same winter a black one was 
killed at Fort Simpson. Tyrrell says that this wolf occurs in 
the country between Athabaska Lake and Churchill River, 
but not plentifully. Preble quotes the record of skins of this 
wolf received by the Hudson's Bay Company as follows : From 
1858 to 1884, from the Athabaska district, 2,119; from 1885 to 
1889, a further 339 skins. For the district of Mackenzie, 
between 1863 and 1884, a total of 1,880 skins; in 1889 only 49 
skins; Fort Resolution and Great Slave Lake, between 1862 


and 1887, 193 skins. Dr. Francis Harper (1932) traversing 
this region in 1914 noted seeing a number of skins and other 
evidences of wolves. No doubt they will continue in the general 
region in slowly decreasing numbers for a long time, though 
annually trapped and poisoned. 



Canis lupus orion Pocock, Proc. Zool. Soc. London, 1935, p. 683, Sept. 12 ("Cape 

York, on Baffin Bay, Greenland"). 
FIGS.: Manniche, 1910, figs. 18, 19 (animal), fig. 20 (skull) ; Jensen, 1928, p. 322, fig. 2 

(mounted specimen). 

The Greenland wolf is described as of "a uniform whitish 
grey, except for the tolerably extensive marbling and streaking 
of the black-tipped contour hairs of the upper side, but the 
ears are pale buff, turning to drab towards the point; the top 
of the head as far as the angle of the eye, and of the muzzle, 
apart from its white tip, is very pale brownish grey . . 
the tail has a black tip, but the rest of its upper side is not so 
conspicuously black and white as the back." The skull is said 
to be small and with smaller teeth than those of the races arctos 
of Ellesmere Land or tundrarum of the Arctic coast of Canada. 
"In the fore feet, the digital pads are greatly reduced in size 
and there is no trace of the pollical or carpal pads. " 

Professor Jensen (1928) states that this wolf "has at the 
present time the same distribution as the musk-ox, in that it 
occurs on the north coast and the northern part of the east 
coast, as far as the region round Scoresby Sound . . . 
Even in the regions where the wolf now occurs, it is by no 
means frequent; it is only found scattered and is comparatively 
rarely seen. Before 1899, when Nathorst, on his expedition to 
northern East Greenland, observed several polar wolves, this 
animal had not been seen in those parts, although they had 
been visited by various travellers; this caused Nathorst to set 
forth the view that the wolf had only quite recently immigrated 
to the east coast from the north. " The caribou or reindeer on 
which these wolves must have preyed are now gone from the 
east coast, but the muskox still remains in small numbers, and 
presumably furnishes a major part of the wolfs sustenance. 
Manniche (1910) could find no evidence that the Arctic hare 


or the fox formed any part of the prey. In his account of the 
mammals observed by the Denmark Expedition to East 
Greenland, during which the ship of the expedition wintered 
near Hvalrossodden, he states that the party found evidence 
of wolves between latitudes 75 and 83 10' N. He believes 
that they have been derived from the North American regions 
by way of northern Greenland. At all events they are few in 
number and, curiously, seemed very wary, though one would 
have expected the contrary from their very slight contact with 
man. In the last of August, 1906, he saw a large white wolf 
and at length succeeded in shooting it by lying in ambush 
wrapped in the skin of a muskox he had killed, the meat of 
which served as bait. This female was in excellent condition, 
"almost fat," and weighed 35 kilograms. Its total length was 
about 1500 mm.; tail, 420; ear, 100 long by 100 wide. Two 
others were seen and tracks were found among the mountains 
inland from Hvalrossodden and slightly to the west and north. 
During the winter of 1906-7, while the expedition was frozen 
in near this place, a group of three Arctic wolves came about 
the ship from time to time, but were very wary, and in the dim 
polar night offered no good chance for a shot. Several sledge 
dogs were once set upon them, but they were driven back by 
the wolves, which eventually succeeded in killing four of their 
animals and wounding others. If a sledge journey were made 
to the meteorological station or elsewhere in the neighborhood 
one or more of the wolves would follow at a distance, but even- 
tually they became bolder, so that by the end of the winter all 
three had been shot or trapped. Of these the largest was a 
male weighing 29 kilograms, very thin and emaciated. 

It thus appears that the Greenland wolf at the present time 
is confined to the narrow strip of country between the inland 
ice and the sea from northern Greenland to about the region of 
Scoresby Sound, following the muskox. That in former times 
it ranged much farther south can hardly be doubted, and it 
probably followed the caribou now largely extinct in Greenland. 
It seems probable that the wolf, like the lemming and the 
muskox, spread across northern Greenland and down the east 
coast rather than the west coast, on account of the great ice 
barriers in the region of Inglefield Gulf. The wolf seems to 
have been gone from southern and western Greenland long ago, 
though mentioned by early historians. Even Egede's "De- 


scription of Greenland" in 1745 mentions them as a thing of 
the past. Hence it was a notable circumstance that in the 
winter of 1868-69 one was killed near Unamak and is now 
mounted in the Museum at Copenhagen. It was one of a pair 
seen together, but where they could have come from is diffi- 
cult to tell. At the present time the wolf population of north- 
eastern Greenland must be limited, and whether there is oc- 
casional immigration from Ellesmere Land over the ice is also 
uncertain. The slight basis on which orion was described 
makes necessary a further comparison between wolves of the 
two areas when a sufficient number of skulls can be assembled. 


Canis pambasileus Elliot, Proc. Biol. Soc. Washington, vol. 18, p. 79, Feb. 21, 1905 
("Sushitna River, region of Mt. McKinley, Alaska"). 

Skull and teeth larger than in the neighboring races; "ridge 
of sagittal and occipital crest nearly on a level with frontal and 
with only a very slight descent at occiput and very deep at 
that point; maxillae very b?oad . . . premaxillae ex- 
tending considerably over one-half the length of the nasals 
Color from nearly uniform black to white and black in 
various mixtures. " The total length of the type skull is 263 
mm.; length of carnassial tooth, 27. 

Elliot describes these wolves from the upper waters of the 
Sushitna River in the region of Mount McKinley as "remark- 
able for their large size and black color." He believes their 
skull characters are distinctive, and Goldman in his paper of 
1937 lists this as a valid race, without attempting to define its 
range. Evidently, however, it is larger than the race ligoni of 
the Alexander Archipelago and somewhere intergrades to the 
south with the northern Plains wolf, C. I. occidentalis. Its very 
dark color is in contrast with the usual paleness of the latter. 

The late Charles Sheldon (1930) has left a vivid account of 
his hunting experiences during a year spent in the "Wilderness 
of Denali" (the Indian name for Mount McKinley). He saw 
little of these wolves, though on nine occasions he found in the 
snow the tracks of wolves that had attempted to catch moun- 
tain sheep by dashing upon them from above from a distance 
of a hundred yards or so. In each case the wolf was unsuccess- 


ful; nevertheless this danger to the sheep caused them to be 
very wary and to keep to the high ridges. The wolves seemed 
to find the caribou at lower levels a more dependable source of 
food. Just to the north of Mount McKinley, Sheldon found 
wolves along the upper Toklat River very abundant, always 
hovering about the feeding herds of caribou and following them 
as they roamed, usually in a fairly well-defined circuit. Late 
in September the caribou frequent ridges more than the low 
country and would be fairly conspicuous to a wolf. He re- 
marks also on the innate caution of the wolf, which, though 
rarely pursued in this region, is so wary that he never saw or 
trapped one. Living as they do, in McKinley National Park, 
it seems likely that the wolves of this area will continue for a 
long time to harry the caribou of the outer ranges and less 
often penetrate the upper slopes for sheep. 


Canis tundrarum Miller, Smithsonian Misc. Coll., vol. 59, no. 15, p. 1, June 8, 1912 

("Point Barrow, Alaska"). 
SYNONYM: Canis lupus albus Sabine, Franklin's Narr. Journ. Polar Sea, p. 655, 1823 

(Fort Enterprise, Mackenzie, Canada). (Not Canis lupus albus Kerr, from 


The tundra wolf has a narrower palate and slender rostrum 
in comparison with the Plains wolves and the northern timber 
wolves. It is decidedly larger than the eastern timber wolf 
and in color is "said to be frequently white or whitish." It is 
the form found in the barren-grounds of the Arctic coast region 
from near Point Barrow eastward toward Hudson Bay, and 
perhaps too in the Arctic Archipelago north of those shores. 

Little recent information concerning this wolf is at hand. 
In the middle of the last century and earlier, explorers found 
them in some numbers about the Isthmus of Boothia, and 
Back's expedition saw white wolves near Artillery Lake. 
Preble (1908) has summarized some of these later observations 
as follows: "Armstrong states that a wolf was seen near Prin- 
cess Royal Islands in February, 1851 ; and that many were seen 
at Mercy Bay, Banks Land, during the winter of 1851-52. 
McDougall states that a pack of wolves was seen on Melville 
Island near Cape Russell in June, 1853; one was seen on May 
27, 1854, near Cape Hotham, Cornwallis Island; McClintock 


reports that wolves were observed in October, 1858, at Port 
Kennedy, and in May, 1859, by Lieutenant Hobson on King 
William Land. Kennedy records one seen in the central 
part of Prince of Wales Land in April, 1852. While in the 
Barren Grounds to the northeast of Fort Rae in the early 
spring of 1894, Russell found wolves rather common. Of a 
band of six, two were snow white, the others a light gray. 
During his exploring trip between Great Slave Lake and Hud- 
son Bay in 1900, J. W. Tyrrell found large wolves on the east 
side of Artillery Lake. J. M. Bell informs me that during the 
same season he occasionally saw wolves near . . . the eastern 
end of Great Bear Lake. Hanbury, while traveling overland 
between Baker Lake and the Arctic coast in the early spring of 
1902, noted an occasional wolf. On April 30, when the party 
was near latitude 67 between Lake Garry and Ogden Bay, 
Darrell, his companion, encountered a band of 16 large wolves. 
Darrell writes me that of this band 13 were of the ordinary 
dirty white color, 2 were nearly black, and 1 pied. He states 
that though these wolves live largely on caribou they are not 
very successful in killing these animals unless they can separate 
one from the herd, and that they always seem to be starving 
. . . The band of 16 was the largest pack seen, the animals 
usually being found singly or in pairs, though occasionally half 
a dozen were observed together." 

It seems clear that this race of wolf is nowhere very common 
but is found in small companies or more often singly or in 
pairs scattered over a large area and is not likely to become 
exterminated for a long time. How great an area the animals 
may cover in their search for food may not be possible to find 
out, but it must be considerable, extending as it does to the 
Arctic Archipelago. With the accumulation of specimens in 
later years, a critical comparison is needed to demonstrate the 
validity of the Ellesmere Land and Greenland races in com- 
parison with that of this area. 



Canis lupus youngi Goldman, Journ. Mamm., vol. 18, p. 40, Feb., 1937 ("Harts Draw, 
north slope of Blue Mountains, 20 miles northwest of Monticello, San Juan 
County, Utah"). 

FIGS.: Grinnell, Dixon, and Linsdale, 1937, vol. 2, figs. 208-214 (as C. I. nubilus). 


The wolf of the southern Rocky Mountain region is described 
as a rather light-colored race of medium size, resembling the 
Plains wolf, C. I. nubilus, but larger, its upper side more black- 
ish and suffused with buff and its frontal region rising less 
steeply. It is larger and paler than C. I. baileyi of Chihuahua, 
with a broader, less depressed rostrum. 

This wolf takes the place of C. I. nubilus, the Plains wolf, 
west of the prairie region of Nebraska and Kansas and was 
formerly found over "southeastern Idaho, southwestern Wyo- 
ming, northeastern Nevada, Utah, western and central Colo- 
rado, northwestern Arizona (north of Grand Canyon), and 
northwestern New Mexico. " Over most of this range it has 
been exterminated within recent decades by persecution of 
hunters and of ranchers with whose cattle interests it interfered. 
Bailey (1931) writes that through such efforts wolves were 
practically eliminated from New Mexico by 1927. Goldman 
(1937) states that at the present time it is mainly restricted to 
northwestern Colorado. Warren (1910), in his account of the 
mammals of the latter State, tells of one wolf, whose track was 
identifiable through its having lost two toes in a trap, that be- 
came so bold that special measures had to be taken for its 
destruction. Its tracks with those of others were in a short 
period found about the carcasses of 75 head of cattle and horses. 
He quotes the figures of wolves killed in 1907 (from U. S. 
Biological Survey Circular no. 63) as follows: In Colorado, 69; 
in Wyoming, 1,009; in Idaho, 14. In 1910, Warren believed 
there was not a county in the State of Colorado that did not 
harbor some wolves. Cary (1911) wrote at about the same 
time that wolves were still found in considerable numbers in 
North Park and in Routt and Rio Blanco Counties, where they 
kill a great many range cattle, and they were similarly destruc- 
tive in Baca and eastern Las Animas Counties, in spite of much 
trapping. Cary was told by professional trappers that so per- 
sistently have wolves been hunted, trapped, and poisoned that 
they would rarely come to a scent of any kind and seldom to a 
baited trap, while poisoning was no longer successful. Traps 
set blind in trails or near water holes were most successful. In 
the Lily Park region on the lower Bear River wolves were 
numerous until 1902, but in the two years following a trapper 
killed 61, which nearly put a stop to their local depredations. 
By 1907 they had become uncommon over most of southern 


Colorado. Such constant persecution is gradually reducing the 
wolf population in all this region. 

Whether this form of wolf originally occurred in the extreme 
eastern parts of California may never be known. Grinnell, 
Dixon, and Linsdale (1937) report the wolf as having been 
practically exterminated from California for many years. No 
specimen of wolf from the State had been preserved in any 
museum up to 1922, while the two since taken on the eastern 
border these authors identify as "clearly of the plains wolf 
type" referring them to the race nubilus. 

CANIS RTJFUS RUFUS Audubon and Bachman 

Canis lupus var. rufus Audubon and Bachman, Quadrupeds of North America, vol. 2, 

p. 240, 1851 (Texas; fixed as 15 miles west of Austin). 
FIG.: Audubon and Bachman, 1851, pi. 82. 



Canis floridanus Miller, Proc. Biol. Soc. Washington, vol. 25, p. 95, May 4, 1912 

("Horse Landing, St. Johns River, Florida"). 

SYNONYMS: Canis lycaon $ americana Hamilton Smith, Griffith's Cuvier, Anim. King- 
dom, vol. 5, p. 144, 1827 (not Canis alopex americanus Kerr, 1795) ; Canis ater 
Richardson, Fauna Bor.-Amer., pp. 70-72, 1829 (in part). 


Canis rufus gregoryi Goldman, Journ. Mamm., vol. 18, p. 44, Feb., 1937 ("Macks 
Bayou, 3 miles east of Tensas River, 18 miles southwest of Tallulah, Madison 
County, Louisiana"). 

FIG.: Arthur, 1931, fig., p. 147 (photograph of captive animal). 

In his review of the American wolves, Goldman (1937) states 
that the red wolf, Canis rufns, "usually regarded as a wolf 
. . . , exhibits a departure from the true wolves, and in 
cranial and dental characters approaches the coyotes." He 
therefore gives it full specific rank and relegates to it as sub- 
species the Florida black wolf and the wolf of the lower Mis- 
sissippi Valley which he describes as C. r. gregoryi. This course 
may be tentatively accepted and the three treated together. 

The typical form of red wolf was slightly the smallest of the 


three races and the most southwestern, with a range extending 
from central Texas south west ward to the Mexican tableland. 
In central Texas its range meets that of the Texas gray wolf, 
Canis lupus monstrabilis, "but the two are not closely allied." 
The latter has a much more massive and highly arched skull, 
its dentition is simpler, the upper carnassial with a smaller 
(often obsolete) anterointernal cusp, and the large upper molar 
with less prominent cusp development on the internal lobe; 
the posterior upper molars are relatively smaller than in rufus. 
The color is reddish mixed with gray. 

Bailey, writing in 1905, believed that the ranges of the red 
and the gray wolves, though contiguous, did not overlap, but 
whether there is overlapping or intermingling future studies 
will perhaps show. As a result of special attempts to secure 
specimens of C. rufus from Texas, Bailey reported that at the 
time mentioned there were 14 skulls and 4 skins in the U. S. 
Biological Survey collection on the basis of which he outlined 
the range as "the whole of southern Texas north to the mouth 
of the Pecos and the mouth of the Colorado and still farther 
north along the strip of mesquite country east of the plains, 
approximately covering the semiarid part of the Lower Sonoran 
zone." He mentions a specimen from Matamoras, Mexico. 
In this area these wolves are destroyed by the ranchmen, since 
they kill young cattle, goats, and colts. The result is that at 
the present time this, the typical race, may now be extinct 
(Goldman) . 

In Audubon's day "wolves" were common in Kentucky and 
were usually black. They may or may not have represented a 
race of the red wolf, but in his paper of 1937 Goldman definitely 
names the wolf of the lower Mississippi region a race of it, 
Canis rufus gregoryi. He characterizes it as a "large but slender 
form of a small species . . . decidedly larger and grayer, 
less tawny" than C. r. rufus, with a more slender skull and 
lighter dentition than the Florida race. The middle and sides 
of the face are mixed black and gray, changing to black and 
cinnamon -buff on top of head ; upper parts from nape to rump 
light buff, heavily mixed or overlain with black; outer surface 
of legs between cinnamon and cinnamon-buff, paling on the 
feet. Black individuals occasionally occur. 

Of this race, Goldman has examined over 150 individuals 
from the lower part of the Mississippi Valley. He states that 


it is found mainly on the western side of the river in south- 
eastern Missouri, Arkansas, southeastern Oklahoma, eastern 
Texas, and Louisiana. It grades into the typical form in Texas 
and apparently into the eastern race floridanus in northern 
Alabama. Arthur (1931) publishes a photograph of a captive 
Louisiana individual which was black and shows the relatively 
long coyotelike ears. He says that it is found in the prairie 
and marsh sections of Louisiana, preying on rabbits, native 
rats and mice, squirrels, ground-nesting birds, and fawns, as 
well as on domestic calves, sheep, and hogs. The young are 
born in January and February and number from three to a 
dozen to a litter, averaging about half a dozen, so that the rate 
of increase may be fairly rapid. "While the wolf has been 
persistently hunted by man in Louisiana ... it appears 
to be on the increase, and, slowly but surely, extending its 
range in the state ... In La Salle parish . . . the 
wolves have become very obnoxious because of their depreda- 
tions on live stock, and cattle men in West Feliciana parish now 
fear an increase of their number as the cattle and sheep raising 
business in this former cottonraising territory is growing in im- 
portance . . . It is not unlikely, therefore, that in a few 
years stringent and systematic campaigns against wolves in 
this state will have to be planned. " Similar reports of increased 
numbers are credited by Bellinger and Black (1940), who 
write that " wolves are becoming rather common in the Ozarks " 
with records of several, including two black wolves, taken in 
late years in the northwestern and western part of Arkansas. 
Stanley P. Young (1940) finds that in eastern Texas the red 
wolf is abundant; more than 800 were caught during the pre- 
vious year (1939) on account of livestock depredations. He 
believes that on account of its habit of living in thickets it will 
not easily be exterminated. 

The Florida wolf is now regarded as a race of the red rather 
than of the gray wolf, from which it differs presumably in much 
the same characters as does C. rufus gregoryi, but it has ap- 
parently a much stouter skull and dentition. Black was the 
usual color, but no doubt a mixed phase also occurred. At the 
present time this wolf is probably extinct in Florida, as well as 
in southern Georgia, which formerly perhaps marked its north- 
ward range. Dr. Francis Harper (1927) has reviewed its status 
in the region of the Okefenokee Swamp, where the last one actu- 


ally killed was about 1908, some 10 miles north of Fargo. 
Later accounts of what may very likely have been wolves in 
this region point to its survival there perhaps as late as 1918. 
In peninsular Florida, wolves were present in the Everglades, 
according to Cory (1896), up to the middle nineties. Miller in 
his original description of the race makes as the type a specimen 
obtained by the U. S. National Museum on the St. Johns 
River, August 12, 1890. This and a skull taken by Dr. Henry 
Bryant many years before, and now in the Museum of Com- 
parative Zoology, seem to be the only extant specimens from 
the State. In Bradford County they were reported up to 1895 
(V. Bailey, 1907). 

A. H. Ho well, writing in 1921, says that in former times 
wolves probably ranged over the entire State of Alabama, but 
are now on the verge of extinction. In Bartram's time, late in 
the eighteenth century, they were common, roaming the moun- 
tainous areas in small packs and preying considerably on the 
smaller domestic animals, sheep, goats, pigs, and sometimes 
calves. On the Gulf coast, "strangely enough, in the big 
swamp country in Baldwin and Mobile counties, where deer are 
still numerous, wolves were apparently exterminated many 
years ago; the last one of which there is record was reported 
killed near Carlton about 1894." During the second decade 
of the present century wolves were still present in small num- 
bers in northwestern Alabama in the "rough, hilly country 
stretching from Walker County northwestward to Colbert 
County." Here three or four were killed in 1912 near South 
Lowell, while in 1915 wolves were destructive to stock in west- 
ern Cullman County, afterward moving westward into Winston 
and Marion Counties, killing "thousands of dollars' worth of 
sheep and goats" and some calves. In 1917, a wolf was killed 
in Colbert County and is described by Ho well in some detail. 
It is the latest record mentioned by him. Goldman says of 
this specimen that it is somewhat intermediate between typical 
floridanus and gregoryi, but in its heavy dentition is nearer the 

There is some evidence that this small type of wolf formerly 
ranged northward into Tennessee. Kellogg (1939) refers to the 
race floridanus a right mandible excavated from an Indian 
mound near Citico Creek, Hamilton County, and thinks it 
quite likely that this form ranged over southeastern Tennessee 


at the time of the first white traders. Possibly, too, the form 
gregoryi was the wolf formerly found in the extreme western 
part of the State. At all events, he quotes Benjamin C. Miles 
that the small black wolf was exterminated about 1870 in 
Haywood and Lauderdale Counties, while Major Shaw is au- 
thority for the statement that it had been replaced in later 
times by the larger and fiercer gray wolf. 

Family FELIDAE: Cats 


The pumas, or mountain lions, are typically American and 
constitute a single species having a wide latitudinal range from 
the northern United States and southern Canada southward 
in forested country to Patagonia. In South America they are 
spoken of by people of Spanish descent as "leon" in contra- 
distinction to the "tigre," or jaguar. Other names are 
"puma," used in Peru, and "couguar, " said to be a corruption 
of a Tupi word signifying an animal of the same color as a deer. 
For pumas are associated with Seer, on several species of which 
they prey in different parts of their vast range. In disposition, 
these cats are not especially fierce in spite of many tales told by 
frightened persons, and very seldom indeed do they attack 
human beings. In South America they are not feared, the 
jaguar being held in high respect. Slightly smaller than the 
latter, the pumas are characterized by a uniform coloration of 
tawny shades above, whitish lips and eye-stripes, and a black 
spot on each side of the muzzle. With a total length of about 
8 feet for an adult male, and a weight ranging up to some 200 
pounds, there is a wide local variation in size, shade of color, 
and in cranial details, which has led to the distinction by Nelson 
and Goldman (1929), the last to review the matter, of no less 
than 19 subspecies, of which 9 are South American, including 
the typical race of southern Brazil, and 10 are found in North 
and Central America. The tropical races tend to be smaller 
and brighter in color, becoming a distinct reddish in tone. 
The southernmost races are paler, the race pearsoni of Pata- 
gonia being a silvery gray. In the United States, at the more 
northerly part of their range, pumas may show two color 
phases, a redder and a grayish brown, sometimes spoken of as 


red and blue, corresponding to the color phases of deer, though 
not seasonal as in the latter. 

While in South America there seems as yet little to indicate 
a widespread reduction in numbers of any of these races, except 
in the more settled regions, in North America pumas have been 
exterminated over most of their range in the eastern United 
States; in the West, where ranching interests are paramount, 
they are locally gone, and will doubtless ere many years become 
few and restricted to large areas of forest country or to national 
parks, where with proper measures their numbers can be kept 
under control. Nevertheless, in other areas a good many yet 
remain and probably will continue for a long time to come. 
The North American races are here considered in order. 


Felis couguar Kerr, Linnaeus's Anim. Kingdom, p. 151, 1792 ("Pennsylvania"). 
FIG.: Nelson, 1916, p. 413, upper fig. (colored). 

Originally the range of the eastern mountain lion extended 
from central New England westward to the edge of the Plains 
and southward through the forested country to Georgia and 
northern Alabama, where presumably it merged with that of 
the Florida puma. Over this region it is at the present time 
probably extinct, with the possibility, however, that a very few 
may remain in the southern Alleghenies. Its range coincided 
more or less with that of the Virginia deer, which formed its 
main object of prey. 

The eastern mountain lion is described as dark reddish 
brown, darker along the middle of the back, the tail similar, 
becoming blackish at the tip, and the feet dark also. The 
muzzle is whitish, with a black marking just back of the white 
and in advance of the eye ; a small white line in front of the eye ; 
under side paler, yellowish white. A grayer phase probably 
occurred as well. Length of body when full grown, about 6 
feet, tail about 3 feet; weight about 175 pounds for a Vermont 
specimen. Smaller animals, perhaps females, measure less, 
about 7 feet over all, with a weight of 118 pounds (Vermont 
specimen) . 

The catamount, or panther, as these big cats were usually 
called in New England, was not especially common even in 


early days but was nevertheless evidently troublesome to the 
first white settlers on our shores, for bounties were offered and 
many were killed though with little record. At first they were 
thought to be young lions, and one is so reported by John 
Josselyn at Cape Ann in earlier times. As early as 1694 Con- 
necticut offered a bounty of 20 shillings apiece for catamounts 
and as late as 1769 paid for four or five. Massachusetts first 
offered a reward of 40 shillings for killing these animals in 1742. 
In 1753 this was increased to 4. Probably by the time of the 
Revolution panthers were fairly well gone from the three 
southern States of New England. Wood, in his early account 
of New England, tells that "Plymouth men have traded for 
Lyons' skins in former times, " but they seem to have been few 
in eastern Massachusetts and were early driven back. In the 
Boston Gazette of April 20, 1741, is a notice of a strange animal 
exhibited at the Greyhound Tavern in Roxbury that had been 
"caught in the woods about 80 miles to the westward," hence 
in western Massachusetts. "It has a tail like a Lyon, its legs 
are like Bears, its claws like an Eagle, its Eyes like a tyger." 
It was "called a Cattamount." Judd, in his "History of 
Hadley, Massachusetts," 186^?, mentions that one was killed 
by some Northampton hunters in 1764, and others were said 
to have been shot in later years in Hampshire. Emmons in 
1840 regarded the species as then extirpated in Massachusetts, 
yet one was killed in 1847 or 1848 in West Greenwich, R. I., 
was mounted and for many years was preserved in the museum 
of the Providence Franklin Society, and then was secured by 
the Boston Society of Natural History, in whose museum it 
still is. This seems to be the last record for the State. In 
Connecticut it disappeared at about the same time, for Linsley 
(1842), writing of the mammals of Connecticut a century ago, 
had no later record of it than of one he saw some years previ- 
ously that had been killed in the northern part of the State. 
Panthers were occasional in the southern half of Maine during 
the early days of settlement but were practically gone by 1815, 
although what may have been the last one killed in the State 
was shot about 1845 at Sebago (Norton, 1930). From time to 
time there have been rumors of panthers in the wilder parts of 
Maine even down to the present day, but it seems safe to say 
that these reports deserve little credence. In southern New 
Hampshire panthers were occasionally killed, and there are 


several accounts in local histories of such events in the latter 
part of the eighteenth century, but the last one actually cap- 
tured and still preserved seems to be the specimen now in the 
possession of the Woodman Institute, Dover, N. H., which was 
shot on November 1, 1853, in the town of Lee. I have one or 
two later accounts of panthers seen at short range in the 
White Mountains as lately as about 1880, the details of which 
are so circumstantial that they seem quite credible. Vermont, 
however, was the best country for these animals, and in the 
days of settlement previous to the American Revolution they 
were often hunted and killed. At the present time there are 
extant perhaps three specimens, possibly four, that were killed 
in the State. The last one was taken in the town of Barnard 
in 1881 and is in the State Museum at Montpelier. Since then 
there have been a few other reports, some of which seem 
credible, but probably the species has ceased to exist in Ver- 
mont for nearly 50 years 

The wilder parts of New York State were formerly good 
country for panthers. In DeKay's time, a century ago, there 
were still a few in the Catskills, and in his book of the mammals 
of New York he mentions that as a boy he recalled the appear- 
ance of one in Westchester County within 25 miles of the city 
of New York and was also informed of one that had been killed 
in Warren County. Merriam has given an account of the 
species in the Adirondacks in the latter part of the last century. 
Here they were then present in some numbers, and when in 
1871 a bounty was offered on them no less than 46 were re- 
ported killed in the succeeding decade, about half of them in 
St. Lawrence County. Merriam (1882) comments that in 
1882, owing to the incentive of the bounty, they had been 
killed down to nearly the point of extermination, yet Miller 
(1899) continues the bounty tabulation down to 1890, with an 
additional 107 panthers on which $20 each had been paid. 
Since the last date no further bounties were claimed up to 
1897, when a panther, now mounted at Albany, was killed on 
Sumner Stream. The last bounty was paid on one killed 
December 27, 1899. Since that date, however, there were oc- 
casional reports of panthers from the Adirondacks, the last in 
1903, said to have been seen at Big Moose, though it seems 
safe to say that they were virtually extinct in the State by 
about the close of the century. 


To the westward there were formerly a few panthers in 
southern Ontario and to the northward they extended into ex- 
treme southern Quebec. But the Quebec records, according to 
Seton, were long ago: Sherbrooke, about 1840; and near Sorel, 
October 3, 1863; and there is one old record for the Toronto 
region of Ontario. Ohio never seems to have had many of 
these animals, or else they were early killed out. Kirtland, in 
the Ohio Geological Survey for 1838, wrote that they had al- 
ready disappeared but that there were Ohio specimens in the 
museum of a Mr. Dorfeuille at Cincinnati. The bounty of $4 
for a grown panther killed in Ohio was discontinued in 1818. 
The last record for the State is perhaps of one killed in 1805 
near Newark (Brayton, 1882). For Indiana, Lyon (1936) 
writes that they were gone from the southern part of the State 
at the time of the Prince of Wied's sojourn there in 1832-33 
but that they persisted a little longer in the northern part. 
A few final records take the panther to about 1838, with 
additional vague reports up to about 1850, when Hahn be- 
lieved it had become extinct in Indiana. Cory (1912) lists 
several later occurrences in Illinois: the last one killed in Ma- 
coupin County about 1840; one killed in Alexander County 
about 1862; one in Daviess County in 1840. In Wisconsin one 
was shot on the headwaters of Black River in December 1863, 
and one was killed in Vernon County, near Westly, about 1870. 
Possibly a few remained into the next decade. What is perhaps 
the only extant specimen from this State is one recorded by 
Schorger (1938) that was killed at Appleton, Wis., November 
22, 1857, and is preserved at Lawrence College. The last 
record of panther in Minnesota is of one killed at Sunrise, 
Chisago County, in 1875 (Herrick). 

According to Bailey (1926) this species must once have 
ranged over much of the Dakotas, but it is not mentioned by 
Alexander Henry and his trappers in the early years of the last 
century. After detailing a few records for western North 
Dakota up to the late years of the nineteenth century, Bailey 
mentions a report of a pair seen at Sully s Lake in 1907, and 
comments: "It is not improbable that a few may still lurk in 
the very rough Badlands country in the western part of the 
State, but it is more probable that the last record for the State 
has been made." He refers the Dakota panther to "Felis 
hippolestes," the type locality of which is Wyoming in the 


Wind River Mountains, but it seems likely that those formerly 
believed to occur in the eastern parts of the Dakotas were of 
the race couguar. 

Summarizing the former presence of the mountain lion in 
New Jersey and Pennsylvania, Rhoads (1903) says that though 
originally found in every part of both States it was always 
more plentiful in the Allegheny Mountains. As early as 1697, 
on account of its destructiveness to stock, a bounty was placed 
on this animal in New Jersey, amounting to 20 shillings for 
"whatsoever Christian shall kill and bring" in the head, or 
half that sum to a Negro or an Indian. In 1730, this bounty 
was reduced to 15 shillings, and evidently it had been effective, 
for the species seems to have been extirpated by about 1830 
or 1840. In the wilder parts of Pennsylvania, however, the 
mountain lion persisted till much later. In Audubon and 
Bachman's time, about the middle of the last century, it was 
so common "among the mountains of the headwaters of the 
Juniata River . . . that one man has killed for some years 
from 2 to 5, and one very hard winter, 7." The latest date 
when one was killed in Pennsylvania was, according to Rhoads, 
about 1871, although later reports of two panthers killed in 
Treaster Valley take the date down to 1893, and may be 
trustworthy. Three subsequent reports of tracks and even an 
animal seen are as recent as 1913 (Shoemaker, 1914). It is not 
at all impossible that panthers may still survive in the wilder 
parts of the West Virginia and Tennessee Alleghenies. Kellogg 
(1937, 1939) has summarized the available information on this 
matter and finds that although numerous enough at the time 
of settlement to give the pioneers some trouble, their numbers 
had been much reduced by the middle of the last century. He 
cites statistics showing that in West Virginia 11 were killed in 
1853, 14 in 1856, 11 in 1858, and 6 in 1859. The last record 
for Lewis County was in 1855, and there is a skeleton in the 
U. S. National Museum from Hampshire County, taken in 
1850. Although no record of any having since been killed is 
given, Kellogg includes various reports of late date, such as 
"tracks of a panther in the snow on Black Mountain during 
the winter of 1935 and also in 1936"; but all such reports 
should be received with caution, even though made by persons 
who might be capable of correctly identifying the species. 
Shoemaker (1914), however, states that one was killed in No- 


vember, 1913, several miles north of Washington, D. C. The 
panther records assembled for Tennessee relate mostly to 
earlier days, with some evidence that they persisted up till 
about 1875, or slightly after. Merriam, who traveled through 
the Great Smoky Mountains in 1888, concluded as a result of 
his inquiries that it was at that time unknown to the local 
inhabitants. Nevertheless, Kellogg accepts a report of one 
said to have been killed in 1929 in the Holston Mountains, 
Johnson County, and another of one "seen crossing the trail 
on Roan Mountain on September 18, 1937," without adducing 
further evidence. C. S. Brimley (1905) records that the last 
authentic report for North Carolina is of one killed near Rose 
Bay, Hyde County, some years before the Civil War. Prob- 
ably the range extended slightly farther south into the northern 
parts of Georgia and Alabama and then merged with that of 
the lowland Florida race. Ho well (1921) writes that in the 
latter State (Alabama), although it is now "nearly, if not 
quite, exterminated . . . recent reports, although rather 
indefinite, indicate that a very few may still remain in the big 
swamps of the southern counties." If these reports may be 
credited, it is likely that they Vefer to the Florida form, and 
Dr. Francis Harper regards the animal of southern Georgia as 
the same, giving records of its presence up till at least the early 
years of the present century. 

From the foregoing brief survey, it is clear that the eastern 
panther, though fairly common for so large a beast of prey, 
was exterminated from the more settled parts along the 
Atlantic coast in the colonial days, while in the wilder and more 
mountainous regions, as in Vermont and New Hampshire, the 
Adirondacks, and the Alleghenies, it persisted in some numbers 
till about the middle of the last century, after which the last 
surviving scattered individuals were gradually shot or trapped 
until they were exterminated, or practically so, by the last 
years of the nineteenth century. Later reports, though in 
some cases perhaps credible, must nevertheless be received 
with skepticism for, as shown by Cory in his (1912) account of 
the animal, even a trained eye and a person familiar with the 
appearance of the animal may sometimes be deceived. The 
testimony of those who report "tracks" requires careful sub- 
stantiation, since few hunters or trappers in the East have 
ever had opportunity to examine these, much less to recognize 
the animal in the wild. 



Felis coryi Bangs, Proc. Biol. Soc. Washington, vol. 13, p. 15, Jan. 31, 1899 ("Wilder- 
ness back of Sebastian, Brevard County, Florida"). 

SYNONYMS: Felis concolor floridana Cory, Hunting and Fishing in Florida, p. 109, 1896 
(not Felis floridana Desmarest, 1820); Felis arundivaga Hollister, Proc. Biol. Soc. 
Washington, vol. 24, p. 176, June 16, 1911 ("12 miles south of Vidalia, Concordia 
Parish, Louisiana"). 

The puma of the Florida Peninsula and the adjacent parts of 
southern Georgia, Alabama, and Louisiana is a brighter- 
colored animal and smaller than the eastern puma, of a "rich 
ferruginous or intense rusty red" above, with smaller feet. 
Hollister, in describing the Louisiana puma as distinct, sepa- 
rated it partly on the basis of its color, which is a "grayish 
fawn . . . with a decided cast of ecru drab," especially 
on flanks and legs. Since, however, Nelson and Goldman 
(1929) have relegated this name to the synonymy of the 
Florida puma, it seems evident that his description applies to 
the grayer color phase of the same animal. Hollister describes 
the skull as "much larger than the skulls of F. couguar, with 
well developed crest and much larger nasals" and ear bullae. 
The condylobasal length of the skull in an adult male described 
by this author was 193.5 mm.; greatest length of nasals, 63. 

The Florida puma, though fairly common in many parts of 
Florida and the adjacent States at the end of the last century, 
has apparently become much reduced in numbers of late years. 
Cory (1896), in his book "Hunting and Fishing in Florida," 
has much to say of his experiences with this animal. He speaks 
of it as "still not uncommon in the more unsettled portions of 
the State" on both the east and the west coasts. He himself 
had found tracks of seven in one week within 30 miles of Lake 
Worth in southern Florida and mentions that one John Davis 
had killed six in that region in the season of 1895. The Indians 
reported to him that they were also "numerous" in the vicinity 
of the Big Cypress south of Fort Myers on the western side of 
the peninsula. Farther north they were even then less common 
and were "scarce" on the peninsula east of the Indian River, 
"but were common there a few years ago. " He recounts that 
in the eighties a hunter named Quarterman killed several in 
the vicinity of Canaveral, once making a double shot at two 
old males that he discovered fighting. Outram Bangs, in 1898, 


secured several through a hunter who obtained them in the 
wilderness back of Sebastian and in his paper on Florida mam- 
mals writes (1898) that though the exact range could not then 
be given, "the puma is extinct in all the region directly north- 
east of Florida, and I believe in northern Florida as well." 
At the present time there are still a number of pumas in the 
Everglades, where they live largely on deer, but probably they 
have been extirpated from most other parts of the State. 

Dr. Francis Harper (1927) has gathered a mass of notes as 
to the presence of the puma in the region of the Okefenokee 
Swamp in southern Georgia and refers the animal to this form. 
Here he says "it is now very nearly if not entirely extinct. Yet 
it lingered well into the present century, and it is perhaps not 
beyond the bounds of possibility that some solitary survivor 
may yet be taken. " At all events, he records one killed about 
1883 and one seen alive about 1903. Older residents of the 
region supplied other instances of animals killed about 1885. 
Most of the later reports are of tracks seen or other signs, 
with a final record of one said to have been killed about 1925 
on the southern edge of the Swamp. The residents in that 
region universally speak of it*as the "Tiger." Hunting is 
usually done in Florida with dogs. When started, the puma 
usually runs a short distance and then takes to a tree. The 
dogs keep it at bay until the hunters come up and shoot it. 

While probably now quite gone from northern Florida and 
Georgia, this race doubtless still occurs in parts of southern 
Alabama and the canebrakes of Louisiana. Writing of the 
former State, Howell (1921) speaks of it as now "nearly, if not 
quite, exterminated"; recent reports, however, "although 
rather indefinite, indicate that a very few may still remain in 
the big swamps of the southern counties. " Tracks were seen 
near Scale about 1912 by an old trapper, and another was re- 
ported as actually seen about 1905 in Baldwin County, but 
these are rather questionable bits of evidence. In 1911 Hoi- 
lister wrote that pumas "are still fairly common in the wilder 
parts of the cane brake region of eastern Louisiana," and he 
himself while hunting in the Bear Lake Cane in February, 
1904, several times heard panthers "crying." 

Panthers formerly were found in somewhat similar country 
in Arkansas, and True (1889) reports a few killed or seen up to 
the late eighties, but it is not certain that they were of this race. 


At the present time what few may remain are probably to 
be found in the Florida Everglades and in southern Louisiana. 


Felis concolor Stanley ana Goldman, Proc. Biol. Soc. Washington, vol. 51, p. 633, Mar. 

18, 1938 ("Bruni Ranch, near Bruni, southeastern Webb County, Texas"). 
SYNONYM: Felis concolor youngi Goldman, Proc. Biol. Soc. Washington, vol. 49, p. 137, 

Aug. 22, 1936. (Preoccupied by Felis youngi Pei, for a fossil Chinese species.) 

Goldman has lately distinguished the cougar of southern 
and central Texas and northeastern Mexico as larger than 
azteca, with lighter upper parts of a more grayish tint, black on 
tail usually more restricted, the dentition heavier, the nasals 
depressed. It is smaller and paler than hippolestes to the 
north. Total length of the type, 2,134 mm.; greatest length of 
skull, 220 mm. A series of specimens examined by Goldman 
represents 16 localities in Texas, and there were also two from 
Matamoros, Tamaulipas, across the border in northeastern 
Mexico. On the east it presumably intergrades with the race 
coryi of Louisiana and Florida. Its general status may be 
considered together with that of its near ally: 



Felis hippolestes aztecus Merriam, Proc. Washington Acad. Sci., vol. 3, p. 592, Dec. 11, 
1901 ("Colonia Garcia, Chihuahua, Mexico"). 

The characters claimed for this cougar are its large size, 
though slightly smaller than the Montana hippolestes, "general 
color dull fulvous [as in the latter] ... but tail darker, 
browner, with longer black tip and no white underneath"; ears 
almost wholly black. The skull is large and massive, though 
not equalling that of hippolestes, but shorter and the teeth 
somewhat smaller. The total length of an adult male, the 
type, was 2,268 mm.; tail vertebrae, 731; basal length of skull, 
171.5. It is thus a somewhat larger, duller animal than browni. 

The range of this race is believed to extend from the southern 
border of the United States in Arizona and New Mexico, south 
into Mexico for an undetermined distance, but probably over 
the tableland in at least the northern half. Of its present 
status in Mexico and Arizona, little information is at hand. 


Bailey, writing in 1905, says: "In most of eastern Texas 
panthers are reported as formerly common, but now as very 
rare or entirely extinct. Individuals have been killed, however, 
within a few years in the swamps not far from Jefferson in the 
northeastern part and Sour Lake in the southeastern part of 
the State. At Tarkington Prairie Mr. A. W. Carter says there 
were a few panthers when he was a boy in 1860, but he has not 
seen one since. In the Big Thicket of Hardin County a few 
panthers have been killed in past years, and Dan Griffin, who 
lives 7 miles northeast of Sour Lake, says a very large one 
occasionally passes his place. He saw its tracks in the winter 
of 1903-4 ... In the rough and sparsely settled western 
part of the State mountain lions are still fairly common in 
certain sections, where they often lay a heavy tribute on colts, 
calves, and sheep." In the region about Langtry they were 
reported as common a few years previously, but in 1903 were 
become already scarce; and there were a few at that time in the 
Franklin Mountains, where they annually killed numbers of 
colts. "In the Davis Mountains these cougars have been 
hunted with hounds till scarce, but in the Santiago Range, in 
the Chisos Mountains, and along the canyons of the Rio 
Grande and Pecos they" were still common in 1905 (Bailey). 
Even at the present day a few cougars still trouble the cattle 
ranchers in this region, but are constantly being hunted down. 
Of their future prospects, Bailey writes: "The rough desert 
ranges, full of canyons, cliffs, and caves, are the favorite haunts 
of the panthers, and will be their last strongholds, not only 
because of the advantages they offer for foraging but because 
of the protection they afford from hounds and hunters. " In 
extreme southern Texas they were, according to Attwater in 
1896, "not as scarce as the Jaguar in the country west of San 
Antonio, but they are fast becoming killed out" (J. A. Allen, 
1896). He knew of but two records in recent times that he 
considered trustworthy; these, in Bexar and Kerr Counties, 
respectively, were sight records made in 1893 and 1894. 

Mountain lions probably referable to the race azteca "are 
or have been common over practically all of New Mexico, but 
they are rapidly decreasing in numbers and in 1931, writes 
Bailey, "are rare or absent from most of the open plains 
country, but are still found in many of the rough or timbered 
mountain ranges, which afford them cover and game." In a 


report for 1917, J. S. Ligon had killed during the year 84 and 
estimates that there were still about 400 left alive in New 
Mexico. "At the present time they are probably most com- 
mon in the Mogollon Mountain region, and in the Animas, 
San Luis, and Sacramento Mountain Ranges, with a few 
scattered through some of the small desert ranges over the 
southern and western parts of the State." The continual 
hunting down of these animals by ranchers and by government- 
paid hunters can not fail within a few years more to reduce 
their numbers to very small proportions. Bailey (1931) quotes 
the following figures for panthers killed by hunters in govern- 
ment employ in New Mexico: In 1917, 17; in 1918, 14; in 1919, 
41; in 1920, 63; and in 1921, 29, a total of 164 in five years. 
With such a determined campaign waged against them, it 
seems likely that the species will soon be largely wiped out over 
much of the State. When one locality is rid of them, they may 
come in from surrounding territory, for they are great travelers. 
Bailey mentions a case where a hunter followed a track for two 
days and estimated that the animal had covered 30 miles 
without making a kill or stopping for any length of time. 
They may have as many as six young at a litter but usually 
four or even two; the gestation period was 96 days in the case 
of a captive individual (J. A. Allen, 1896). Hence animals 
killed in early or mid winter have probably not yet bred, but 
with females shot late in winter the potential increase is like- 
wise destroyed. 


Felis aztecus browni Merriam, Proc. Biol. Soc. Washington, vol. 16, p. 73, May 29, 
1903 ("Lower Colorado River 12 miles south of Yuma, Arizona"). 

Very little is known concerning this puma. Grinnell, Dixon, 
and Linsdale (1937) describe it as "closely similar to the Cali- 
fornia Mountain Lion but with shorter pelage, paler tone of 
coloration, and smaller skull and teeth." Merriam (loc. cit.) 
in describing the original specimen emphasized similar charac- 
ters in comparison with the more eastern race, azteca, calling 
attention further to the smaller and lower audital bullae, small 
canines and carnassials. Merriam suggested that the slender 
canines indicated that the animal fed upon smaller game than 


do the larger neighboring races. The type specimen was said 
to have measured in the flesh 7 feet 4 inches from tip of nose to 
tip of tail, of which the latter was 28.5 inches. It weighed 170 
pounds (perhaps an estimate). Greatest length of skull, 198 

This small race of the puma was probably confined to the 
desert regions of the lower Colorado River in southeastern 
California and the adjacent parts of Arizona, where, according 
to the authors above quoted, it "is now rare, perhaps even near 
to extinction." They add that in the "spring of 1910, while 
exploring the Colorado River from Needles to Yuma," they 
obtained evidence, both direct and through report, of the pres- 
ence of mountain lions about Riverside Mountain in the ex- 
treme northeastern part of Riverside County, California. The 
animals here "appeared to range chiefly through the densely 
wooded bottom lands, though one was reliably reported to 
have been seen among the rough desert hills which constitute 
'Riverside Mountain'." They saw a fresh track on May 1, 
1910, four miles below Potholes, on the California side of the 
River, and had further reports from Ehrenberg, Cibola, and 
Potholes; and secured two hides and skulls from a rancher 18 
miles north of Picacho who had shot the animals the previ- 
ous autumn. These pumas had appeared in the region where 
none had been seen for ten years, and one by one killed off 
eight of the rancher's hogs. Finally one was hunted down and 
shot. "Meanwhile the hogs had become thoroughly frightened 
and had taken to swimming the river twice daily to forage for 
mesquite beans on the Arizona side, where they appeared to 
feel safer. " A second lion continued the depredations and was 
eventually shot. Inquiry of the rancher 12 years later brought 
out the fact that he had seen no more pumas since that event. 
In 1909, a pair of mountain lions appeared near Calexico and 
were occasionally seen by the ditch tender swimming the main 
Imperial Canal at that point. These lions lived on pigs that 
had gone wild. Since 1910 no information is available from 
California as to the status of this race, and it is doubtful if it 
still exists. 

Outside of the immediate delta region of the Colorado, it is 
likely that this subspecies was found in northern Lower Cali- 
fornia. Nelson (1921) includes it without comment in his lists 
of the fauna of the region. Probably it was closely similar to 


the race improcera of the central and southern parts of the 
peninsula. How far it may have ranged into the adjacent part 
of Sonora is also unknown, but it may have already become 
nearly extirpated. 



Felis improcera Phillips, Proc. Biol. Soc. Washington, vol. 25, p. 85, May 4, 1912 

("Calmalli, Lower California"). 
FIG.: Phillips, 1912, pi. 5 (skull). 

Little seems to be known of this puma. The adult male 
upon which the subspecies was founded is small, with a skull 
of only 150 mm. in basal length, and of a rounder, less elongate 
form than in neighboring races. The color is "dark fawn," 
darker along the back, the general hue less reddish than in 
F. c. azteca. 

This small puma is believed to be characteristic of central 
and southern Lower California. Calmalli, the type locality, 
lies in the Vizcaino Desert district, which occupies the middle 
third of the peninsula and includes the great lava plateau, 
with scattered mountains, low ridges, and groups of granite 
mountains. Two remarkable plants are largely characteristic 
of this region, the "cirio, " which forms polelike forests, and 
the " elephant wood, " with fantastic massive trunks and small 
branches. There is abundance also of giant and other cactuses. 
In this arid region of the Lower Sonoran desert and in the Arid 
Tropical area of southern Lower California, this little puma is 
found. Practically nothing is recorded of it. Dr. Phillips be- 
lieved it "probably a rare animal throughout the entire length 
of the peninsula, " for he adds that he is informed by W. W. 
Brown, Jr., that it occurs "even south to the vicinity of Cape 
St. Lucas. " On account of its restricted habitat and the barren 
country in which it lives, its numbers can not be large and are 
not likely to increase. Unless changes come in this habitat, it 
is likely to remain as it is except as it comes in contact with the 
scattered ranches and interferes with stock raising. At the 
present time there is no evidence at hand. 



Felis hippolestes Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 219, July 15, 1897 
("Wind River Mountains, Wyoming"). 


Felix (sic) oregonensis Rafinesque, Atlantic Journ., vol. 1, p. 62, 1832 (Northwest coast 

of United States). 
SYNONYM: Felis hippolestes olympus Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 

220, July 15, 1897 (Lake Cushman, Mason County, Washington). 
FIGS.: Elliot, 1904, pt. 2, pis. 44, 45 (skull). 


Felis calif ornica May, California Game "marked down," p. 22, 1896 (Upper Kern 

River, Kern County, California). 
FIGS.: Grinnell, Dixon, and Linsdale, 1937, pi. 11, col. (showing red and gray phases), 

figs. 215-239 (skull, photographs of old, young, habitat, etc.). 


Felis concolor Vancouver -crisis Nelson and Goldman, Proc. Biol. Soc. Washington, vol. 
45, p. 105, July 15, 1932 (Campbell Lake, Vancouver Island, British Columbia). 

The present status of these races may be considered briefly 
together, for although none of them seems now in actual danger 
of extermination, nevertheless in the years to come, with the 
constant encroachment of human occupation, with hunting for 
sport or for bounties, and with forest fires and other causes, 
it is likely that their ranks will eventually dwindle, and that 
they can survive in any numbers only in large reserves or in 
country that remains in a more or less primeval condition, 
with shelter of forests and food in the shape of larger game, 
meaning chiefly deer. The fallacy of completely killing out 
these larger predators in deer reserves has been several times 
demonstrated. In the case of the Kaibab Forest, from which 
panthers and wolves were eliminated by purposeful hunting 
and trapping, and from which deer hunters were excluded, the 
mule deer in the course of a few years became so abundant 
that not only did the animals destroy all the available food but 


also permanently injured the sparse forest of the area so that 
hundreds died from starvation. A nearly similar state of affairs 
took place in the Murderers Creek Game Refuge of eastern 
Oregon, which was established in 1929 in the belief that the 
deer there needed protection (Englis, 1939). In the course of 
some years the mule deer so increased that in one section 
12,000 deer were trying to exist "where even 6,000 would have 
gone hungry. " Bitterbush, mountain mahogany, and juniper, 
the chief winter food plants, were so overbrowsed that bitter- 
bush became mere stubs, and the junipers were trimmed as 
high as a deer could reach. Many deer starved to death and 
the others were small and undernourished. In 1935 the area 
was reopened to hunting and the refuge given up. In such 
cases it is clear that a proper proportion (to be determined) of 
panthers would not only keep the deer down to a number 
commensurate with the carrying capacity of the range, but also 
afford a certain return from fur, besides adding to the human 
interest of the situation, in hunting or observation. This view 
of the value of predators has only in recent years been brought 
out forcibly as a result of the experiments in "control" that 
have been more or less intentionally carried out by federal and 
other agencies in our West during the past 60 or 70 years. It 
now becomes clear that "predatory animals are to be con- 
sidered as an integral part of the wild life protected within 
national parks, and no widespread campaigns of destruction 
are to be countenanced, " but native predators may be allowed 
to continue their normal utilization of other park animals, unless 
in special cases where the need for some regulation becomes 
obvious (Cahalane, 1939b). This logical and scientific policy, 
based on an accurate knowledge of the various species con- 
cerned, should become increasingly established over all the 
larger areas of unsettled country. 

Of the four races listed above, Felis c. hippolestes is the largest 
member of the species; it is of a dull fulvous color and has a 
large massive skull with a highly developed sagittal crest. A 
large male measured 8 feet in total length and weighed 227 
pounds (Merriam, 1901, p. 586). The skull measured 196 mm. 
in basal length. Young are born in January and February in 
Colorado. This race ranges from extreme northern New 
Mexico (probably) northward through Colorado and the ad- 
jacent parts to Wyoming, Montana, and British Columbia. 


Bailey (1931) refers to this race a large specimen in the collec- 
tion of the U. S. Biological Survey from the Jemez Mountains 
of northern New Mexico and believes that other dark-brown 
skins from Taos and Pecos River Mountains of that region are 
the same. In the latter district they were "common" in 1903, 
and a few were killed every year in the Taos Mountains. "In 
1910 officials of the Forest Service reported mountain lions as 
fairly abundant on the Carson National Forest and as still very 
common in the Jemez Mountains. In 1914 they reported 3 
killed on the Carson, 3 on the Pecos, and 2 in the Jemez Na- 
tional Forest; in 1915, 4 on the Carson and 4 on the Santa Fe 
National Forests; and in 1916, 4 on the Carson and 7 on the 
Santa Fe National Forests." These large cougars could and 
did kill even elk, as well as deer, but as these and "other native 
game animals become scarce, the mountain lions turn their at- 
tention to domestic stock and seem especially to relish colts, 
but if these are not to be found, they take horse meat of any 
kind. In spite of the bounty usually paid for their destruction 
and the efforts of stockmen and hunters, they have until re- 
cently held their own in the rougher parts of the country, but 
with the present organized effort it will not be long before they 
are sufficiently reduced in numbers to prevent any great 
losses" (Bailey, 1931). 

In Colorado the range of this race, which formerly inhabited 
all the "rough parts of the State, and in early times" was oc- 
casionally seen "even well out on the plains along the more 
heavily brush-fringed streams," has now become much more 
circumscribed. In 1911 Cary wrote that it was already rare 
east of the Continental Divide, though holding its own fairly 
well in the rough canyon and mesa country of the western and 
southwestern parts of the State, where locally the animals 
were still sufficiently numerous to be very destructive to stock, 
especially young colts. Cary (1911) lists eight regions in the 
northern mountains of Colorado in all but one of which they 
were said to be from uncommon to "common"; in the Snake 
River region where formerly they were common, none remain. 
Reports from southern Colorado for ten regions where they are 
found indicate that they are much less numerous, though a few 
still occur. They may be found locally up as high as 10,000 
feet in some of the mountain ranges. At least until recent 
years the cougar was of local occurrence in the mountainous 


parts of Utah and Idaho, but its numbers can not be very 
great, as a result in part of determined efforts of stockmen and 
professional hunters. 

Bailey (1918) writes that they were still common in Glacier 
Park, northwestern Montana, 20 years ago but were confined 
for the most part to the western slopes, where dense forest 
cover and abundance of deer offered excellent living conditions. 
"Apparently they are still almost as numerous as they were in 
1895," when he first went through the region. "In this 
region," he adds, "their food consists mainly of the white-tail 
deer, which abound on the west slope of the park. " Although 
Bailey advocates their destruction, as contributing little to the 
tourist interest of the park, since they are seldom seen, never- 
theless their value as a check on the over-increase of deer might 
well justify the maintenance of a proper number. 

In southern British Columbia the cougar is uncommon but 
extends into the Peace River district. Cowan in a recent report 
(1939) writes of a female killed in March 1937 by Ted Strand 
of Little Prairie, and quotes Seton ("Lives of Game Animals") 
as to a specimen shot in November, 1921, near the junction of 
Cypress Creek and Halfway River. This latter "seems to be 
the northernmost record for North America." 

On Vancouver Island the panther still occurs in some num- 
bers in wilder areas and has lately been distinguished as a sepa- 
rate race, characterized by its darker, more rufescent upper parts 
and the more elevated frontal region of the skull, in comparison 
with the Rocky Mountain and Oregon races. Swarth, in his 
paper of 1912, wrote that for an animal of this type it was 
abundant "throughout the wilder parts of Vancouver Island, 
and frequently seen near many of the smaller towns also." 
They are shy and secretive but are often hunted successfully 
with dogs. One farmer in the Beaver Creek Valley, near 
Alberni, had killed as many as 13 during the previous winter. 
In a letter dated July 8, 1939, Maj. Allan Brooks writes me 
that "over the whole northern third of this island beavers have 
been decimated by cougars of late years, and they will take 
many years to recover if they ever do. " Cougars have also 
been an active agent in the destruction of deer, concentrating 
on the more healthy upland population, on account of the 
dying off of deer in the lowlands from infection by liver flukes. 
At the present time, therefore, the Vancouver cougar seems in 


no danger, unless more intensive operations for its destruction 
are undertaken to preserve the deer. 

Farther south, along the coasts of southern British Columbia, 
Washington, and Oregon, occurs the race oregonensis, which is 
nearly as large as the Rocky Mountain subspecies but darker 
and richer in color, with more black on the tail. A large one 
was said to have weighed 150 pounds. Concerning its status 
in the State of Washington, Taylor and Shaw (1929) write 
that it is "probably more common in the Olympic Mountains 
than elsewhere, but present also in the Cascades and Blue 
Mountains. Predatory-animal control campaigns are markedly 
reducing its numbers." In Oregon cougars were common in 
the forests west of the Cascade Mountains up till recently but 
at the present time have been so reduced by professional 
hunters over the State "that they are no longer a serious 
menace to livestock industries" (Bailey, 1936). Some idea of 
their numbers in recent years may be gained from the figures 
given by Bailey (1936), who states that for the period from 
October 1, 1913, to December 31, 1914, bounties totalling 
$4,035 were paid on 269 mountain lions killed in Oregon. In 
Curry County alone no less th&n 60 were killed, and smaller 
numbers in 19 other counties. In the fiscal year 1930, Jewett 
reported 17 killed by U. S. Biological Survey hunters in Oregon, 
"where they had been reported killing stock or game. While 
the number is insignificant, it shows a marked decrease in these 
big cats during recent years and that their destruction of live- 
stock and game is being well curbed. " It seems clear from these 
figures that mountain lions will ere long become uncommon in 
the State if this policy is continued. In the wilder sections, it 
may be that a certain number of these predators will be valu- 
able in keeping down too great an increase of deer. 

Grinnell, Dixon, and Linsdale (1937) have published an ex- 
cellent summary of our knowledge of the race calif ornica as it 
occurs in California. The race is less dark and richly colored 
than the Oregon cougar. Both red and gray phases occur, 
with an intermediate condition, all three in about equal pro- 
portions. Adult males weigh upwards of 165 pounds, with an 
average of about 140. It occurs in forested and chaparral- 
covered areas mainly in the mountains west of the Great Basin 
and desert divides from Oregon to the Mexican line. For the 
most part it lives at middle altitudes in the mountains between 


2,000 and 5,000 feet. Young seem to be born in almost any 
month of the year, but with a peak in April. The usual number 
seems to be two young to a litter in California. The chief 
prey is deer, on the increase of which it forms a natural check. 
Some valuable data on the abundance and feeding habits of 
cougars, based on much field experience, give perhaps for the 
first time a fairly accurate picture of the economic status of 
mountain lions, at least in California. In good lion and deer 
country there is on an average about one lion to a township 
(36 square miles), and in a few very favored localities about one 
to each 10 square miles. Where deer are most plentiful, as in 
national parks and game reserves, there also the mountain 
lions tend to be most concentrated. A careful study indicates 
that at the time of writing, according to the authors quoted, 
the mountain-lion population of California was about 600 and 
that the annual toll of these animals upon the deer was roughly 
21,600. Yet in spite of this, and of a nearly equal number an- 
nually taken by hunters in the State, deer are actually increas- 
ing. From 1908 to 1921 the annual kill of mountain lions 
averaged 258, and the State even employs a professional hunter 
for their reduction. Yet it seems that the species is about 
holding its numbers in spite of this toll and a natural mortality. 
In regard to stock-killing propensities, it appears that usually 
those lions that have learned by experience that this sort of 
prey is easily secured are the main ones to be feared, and with 
their elimination in grazing areas the trouble is much abated. 
The present prospect for the California mountain lion is that 
although inevitably it must retreat before the spread of settle- 
ment and agriculture, nevertheless in wilder areas it will per- 
sist for many years to come and continue to be a source of 
slight revenue for hunters and perhaps in places a real asset in 
the natural control of deer populations in protected areas. Its 
situation is perhaps more favorable than is that of the other 
three races of the West, the Rocky Mountain, the Vancouver, 
and the Oregon cougars. 


In a recent paper Nelson and Goldman (1933) have re- 
viewed the systematic relationships of the local races of jaguars 
in America. These, the largest of American cats, constitute 
but a single species, which inhabits the tropical and subtropical 


regions from south-central Argentina northward to southern 
United States. With the general appearance of a leopard, 
having a yellowish ground color marked with black spots or 
circles of spots, the jaguar is slightly heavier of build than a 
leopard, with a more massive head and relatively shorter tail. 
The color pattern is variable, but usually at least some of the 
rosettes of black spots have a central spot in the jaguar, which 
is absent in the leopard. The authors mentioned have given 
descriptions of no less than 16 local races of jaguar over the 
wide range covered by the species as a whole. The characters 
on which these are based are usually not striking and depend 
largely on slight peculiarities in the skull. Of the 16 races, 
five occur in North America, the others in South America. 
Concerning most of these, information is rather scanty and in 
general, although jaguars are frequently hunted wherever they 
come into proximity of man, there is little to indicate that they 
have been effectively depleted in numbers except perhaps in 
the regions of thickest settlement. Since the two forms that 
occur along the southwestern border of the United States have 
probably become much reduced in numbers and in the extent 
of the country they occupy, they^may be included together here. 


Felis onca arizonensis Goldman, Proc. Biol. Soc. Washington, vol. 45, p. 144, Sept. 9, 

1932 ("Near Cibecue, Navajo County, Arizona"). 


Felis onca veraecrucis Nelson and Goldman, Journ. Mammalogy, vol. 14, p. 236, Aug., 

1933 ("San Andres, Tuxtla, Vera Cruz, Mexico"). 
FIG.: Nelson, 1916, colored fig. on p. 413. 

These two races of the jaguar are characteristic, respectively, 
of the subtropical areas of the "mountainous parts of eastern 
Arizona north to the Grand Canyon, southern half of western 
New Mexico, and northeastern Sonora, " and the "Gulf slope 
of eastern and southeastern Mexico from the coast region of 
Tabasco north through Vera Cruz and Tamaulipas to central 
Texas." Their ranges are therefore separated by the high 
tableland of north -central Mexico. The race arizonensis is a 


large northern form distinguished by its flatter, more depressed 
nasal bones from other subspecies; it differs additionally from 
the race hernandesii to the south by its more massive skull 
with broader rostrum, wide anterior nares, and narrower 
posterior nares. The race veraecrucis is the largest of the North 
American races and has the nasals more arched. A tanned 
skin of arizonensis measures in total length, 2,145 mm.; tail, 
660; the skull has a condylobasal length of 237 mm. An adult 
male of veraecrucis has a total length of 1,993 mm., the skull a 
condylobasal length of 247 mm. (Nelson and Goldman, 1933). 
The northeastern jaguar just reaches the southern border of 
the United States in Texas but apparently has never been 
known to be common, and by now it is doubtless nearly extir- 
pated in this area. J. A. Allen in 1894 wrote that in Aransas 
County, Texas, on the Gulf of Mexico, it was then already gone 
from the region. He mentions a skin formerly owned by a 
Captain Bailey that was killed in 1858 on Live Oak Peninsula, 
but Attwater who contributed the note had heard of none 
since in that region. The same collector, quoted by J. A. Allen 
(1896), speaks of it as formerly present in Bexar County, 
southern Texas, but adds that it was then (1896) "rare east of 
the Nueces River, but still taken occasionally in the chaparral 
thickets in the counties bordering the Rio Grande. " How far 
to the eastward it may once have extended is uncertain. True, 
in 1885, in his provisional list of the mammals of North and 
Central America and the West India Islands, even stated that 
it ranged from Louisiana to Patagonia, but the basis of the 
Louisiana report is not given. Seton (1920) has called atten- 
tion to an account of what seems to have been the jaguar in 
an old book on "Rocky Mountain Life," by Rufus B. Sage, 
who while encamped on Soublets Creek, headwaters of the 
Platte within 30 or 40 miles of Longs Peak, Colo., mentions a 
"strange looking animal" encountered by one of his party. 
He believed it to have been "of the Leopard family" and adds 
that they (meaning jaguars) "are not infrequently met in some 
parts of the Cumanche country, and their skins furnish to the 
natives a favorite material for arrow-cases. " If this refers to 
the jaguar, as seems likely, it furnishes the most northeasterly 
record. Possibly, however, the ocelot was the animal meant. 
Bailey (1931) mentions a jaguar killed near Center City, Texas, 
as lately as 1903. It was treed by some dogs and after being 


wounded with a revolver shot was finally driven into some 
brush, surrounded, and shot, but not before it had killed a dog 
and a horse. Nelson and Goldman mention a specimen ex- 
amined from Goldthwaite, Mills County, Texas. 

Concerning the Arizona jaguar, the latter authors write that 
at the present time its range marks the most northern point 
where the species is known. Formerly, according to old rec- 
ords, it ranged into extreme southern California, and in recent 
years, "while not very abundant, it appears to be a regular 
resident of southeastern Arizona. " They examined specimens 
from Cibecue, Greaterville, and Nogales in that State. That 
jaguars occurred in southern California in the first half of the 
last century is attested by Merriam (1919), who quotes several 
older accounts: that of Langsdorff, 1814, who mentions it as 
among the species of the Monterey region ; Beechey (Narrative, 
1831), who in 1826 says it was reported to be found in the 
country between San Francisco and Monterey; and Saint- 
Amant, who included it in 1854 as a California species. He 
further calls attention to a circumstantial account of the finding 
of a jaguar family in the Tehachapi Mountains by the famous 
James Capen Adams about the> middle of the last century, as 
detailed in Hittell's (1860) account of the adventures of this 
old hunter. The Indians of a former generation living in 
southern California apparently were well acquainted with this 
jaguar, and Merriam (1919) was told by an old chief of the 
Kammei tribe that in the Cuyamaca Mountain region in San 
Diego County it was there known as "big-spotted lion" in their 
native tongue. Another early writer, Pattie, mentions one 
killed on an island at the mouth of the Colorado River about 
1822. Other Indian records, as gathered by William D. Strong 
(1926) in this region, indicate that the jaguar formerly occurred 
in the mountains bordering the Mohave Desert and that the 
"old people made a practice of following jaguar and mountain 
lion trails in order to uncover and eat the deer remains the 
animals buried. The last jaguar that his informant could re- 
member as having been killed in the region was near Palm 
Springs, Calif., about 1860. In New Mexico the jaguar is now 
rare if not quite extirpated. Bailey (1931) has summarized a 
number of later occurrences of jaguars in the southern part of 
the State: Grafton, in 1900; Ute Creek, San Miguel County, 
1902-3; near Fulton, 1903; Datil Mountains, 1902; Clanton 


Creek ranch, 1903; and Sierra de los Caballos, in about 1904 
or 1905. Since these dates, no later instances are mentioned, 
and probably the species has been reduced to near the vanishing 
point in the United States. It has been found to kill cattle on 
the ranches in these regions and hence is regarded as a menace 
by the ranchers, to be killed wherever found. Probably it still 
occurs in small numbers in northeastern Mexico. According 
to Nelson (1916) the jaguar has little of the truculent disposi- 
tion of the leopard. In parts of Mexico he made careful in- 
quiry without hearing of a single case where one had attacked 
human beings, although the natives everywhere fear it on ac- 
count of its size and strength. He writes: "Jaguars are very 
destructive to the larger game birds and mammals of their 
domain and to horses and cattle on ranches. On many large 
tropical ranches a 'tigrero,' or tiger hunter, is maintained, 
whose duty it is immediately to take up the trail when a 'tigre' 
makes its presence known, usually by killing cattle. The 
hunter steadily continues the pursuit" in which dogs are ordi- 
narily used, until the animal is killed or driven far away. He 
mentions that along the Mexican coast in spring, when sea 
turtles come ashore to lay eggs, fresh tracks of jaguars may be 
found showing where they have traveled along the beach for 
miles in search of the eggs. In the province of Guerrero, 
Mexico, the hunters have a way of imitating the call of the 
jaguar during the mating period and thus enticing it within 

Order ARTIODACTYLA : Even-toed Ungulates 

The cloven-hoofed herbivores form a large and complex 
group. About a hundred genera are usually recognized from 
the Holarctic, Oriental, Ethiopian, and Neotropical Regions. 
The artiodactyls do not occur in Australia, but the deer reach 
Celebes, and the pig family is found on most of the oceanic 
islands, probably carried by man to the islands west of Celebes. 
European and American deer have been introduced into New 
Zealand and some of the other islands, where they are doing 

Since these herbivores are everywhere hunted for food and 
sport, many forms have become greatly reduced in numbers. 
There are three major groups: 

(1) Suina, the hippopotamuses, pigs, and peccaries. 


(2) Tylopoda, the camels and their South American rela- 
tives, the huanaco and vicuna. 

(3) Pecora, the ruminants, to which belong the three families 
treated herein: 

(1) Cervidae, the deer. Six forms of wapiti, 11 forms of the 
American deer, and 13 forms of caribou are considered to come 
within the scope of this work. 

(2) Antilocapridae, the North American pronghorn antelope. 
Four races of the single Recent species are here recognized. 
The familv is the only endemic group of the Artiodactyla in the 
New World. 

(3) Bovidae, the cattle and their relatives. All the American 
representatives of this group, except the Rocky Mountain 
goat (19 forms, 3 genera), are included here as vanishing mam- 
mals. J. E. H. 

Family CERVIDAE: Deer 


These large deer, although almost universally called elk in 
America, should more properly be termed by the American 
Indian name wapiti, since elk in the Old World was originally 
applied to the species resembling our moose, found across 
northern Europe and Asia. In general appearance this animal 
is much like the familiar red deer of Europe but is much larger 
and of a darker color, without the whitish eye ring of the latter. 
The antlers, too, in addition to their larger size, seldom show 
the "cup" of the Old World species, in which the three termi- 
nal tines tend to come off together. Originally found over 
most of temperate North America, this species has become very 
greatly restricted in range within the three centuries since 
active settlement by Europeans. At present some half dozen 
geographic races are recognized, which may be taken up in 
detail, as follows : 


Cervus elaphus canadensis Erxleben, Syst. Regni Animal., vol. 1, p. 305, 1777 (Eastern 

Canada; as later restricted, "near Montreal," Quebec). 
FIGS.: Audubon and Bachman, 1851, vol. 2, pi. 62 (colored figure of animals caught 

when young in western Pennsylvania). 


The general appearance of the eastern wapiti is described by 
Audubon and Bachman (1851), the latter of whom kept on his 
grounds near New York City a pair that were obtained as 
young animals in western Pennsylvania and that later served 
as the originals from which their plate was drawn by Audubon. 
Head dark brown, neck darker, blackish; a white patch on 
each side of the under jaw, with a black stripe between, passing 
down on to the throat. No pale eye-ring. General color dark 
gray all over except for the prominent white rump-patch. 
The male in winter develops a heavy fringe of long hair on the 
throat and back of the neck. Under surface of body, brown. 
The winter pelage is somewhat grayer than that of summer 
which is redder, and the young are spotted with white. The 
large antlers have a rounded beam and extend back, giving off 
a brow tine and two others, the bez and trez, before the longest 
is reached, beyond which is another fork, its tines in the same 
anteroposterior plane. Although there are a number of antlers 
in existence, there is apparently but a single skin of the eastern 
wapiti preserved, namely, one in the possession of the Academy 
of Natural Sciences of Philadelphia. Bailey (1937), on the 
basis of the slender evidence available, believes that the 
eastern animal was not so large and heavy as the Rocky Moun- 
tain race and was brighter brown and more richly colored than 
any of the western forms. 

Three hundred years ago the eastern wapiti ranged from 
southern Quebec to the edge of the Plains and southward 
through extreme western New England and western New 
York, perhaps as far south as North Carolina on the Atlantic 
seaboard, and through the Allegheny Mountains to northern 
Alabama. Such a large animal could not fail to have been 
useful to the early settlers on account of its meat and hide; 
hence it is not surprising that localities the elk haunted or 
where they were killed became commemorated and distin- 
guished from other wilderness spots by appropriate place- 
names. A fairly accurate map of the former range of this 
animal in the eastern United States might be made by plotting 
the cities, counties, creeks, and rivers named after it. Thus in 
western New York we have Elkcreek (town) and Elk Creek 
(stream), Elkdale; in Pennsylvania, Elk City, Elk County, 
Elk Creek, Elk Grove, Elk Hill, Elk Horn, Elk Lake, Elkrun; 
in Maryland, Elk Neck and Elkton; in Virginia, Elk Creek, 


Elk Garden, Elk Hill, Elkton, Elkwood; in West Virginia, Elk 
Garden, Elk Horn; in North Carolina, Elkville; in Tennessee, 
Elk Horn, Elk Mills, Elk River, Elkton; in Kentucky, Elkfork, 
Elk Horn, Elkton; in Alabama, Elk and Elk River. Other 
place names of a similar sort are on the maps of Ohio, Indiana, 
Michigan, and Iowa. But the native elk has long since gone, 
although in a few places as in Corbin Park, N. H., western 
stock has been imported and maintained under fence. 

In the northeastern part of its range the animal was known 
to the French explorers of the St. Lawrence River, but Dr. 
R. M. Anderson (1939a) writes that although found in the prov- 
ince of Quebec south of the St. Lawrence in early days it has 
been gone from there for at least a century. He quotes W. P. 
Lett that about a hundred years prior to 1884 elk were present 
in small numbers in Carleton County, Ontario, and the antlers 
are frequently turned up by the plow in the vicinity of Ottawa. 
Presumably elk were present at that time on the Quebec side 
of the Ottawa River. In New England there is no record of 
the wapiti in historic times, yet it must occasionally have 
reached western Vermont, for antlers have been found in bogs 
in that region. Western New York was a part of the wapiti's 
range at one time. DeKay, in 1842, after correcting a few 
erroneous conceptions concerning this species, writes that they 
were at that time found in "the northwestern counties of 
Pennsylvania, and the adjoining counties of New- York. In 
1834, I am informed by Mr. Philip Church, a stag was killed 
at Bolivar, Allegany county. My informant saw the animal, 
and his description corresponds exactly with this species." 
He also quotes a certain "Mr. Beach, an intelligent hunter," 
that he had shot at one in 1836 on the north branch of the 
Saranac and another hunter, Vaughan, who had actually 
killed one at nearly the same place. Merriam (1884), however, 
is inclined to doubt these last records on the ground that no old 
hunter with whom he had talked in the Adirondack region had 
ever seen or heard of one; nevertheless he admits that they 
must once have been common in the Adirondacks, since a 
number of antlers have been discovered, the best preserved of 
which that he had seen was found in a bog on Third Lake of 
Fulton Chain, in Herkimer County. The largest of several 
discovered at Steels Corners in St. Lawrence County measured 
12 1/2 inches in circumference at the burr and 10 inches immedi- 


ately above it. Thus, in spite of Merriam's doubts, it is clear 
that the eastern wapiti formerly occurred in the northwestern 
part of the State and probably became extinct a century or 
more ago. 

For Pennsylvania the record is much fuller, and Rhoads 
(1903) in particular has gathered what evidence he could find 
of their former presence. Early accounts tell of their abundance 
in that State, especially in the vicinity of salt licks, to which 
they had, with other large mammals, worn broad trails. Peter 
Kalm related how in the winter of 1705 great numbers came off 
the mountains because of the deep snows and were killed. 
Barton in 1806, a century later, implies the reduction of the 
herds: "In the memory of many persons now living, the droves 
of elks which used to frequent the salines near the river Sus- 
quehanna in Pennsylvania were so great that for 5 or 6 miles 
leading to the licks the paths of these animals were as large as 
many of the great public roads of our country. Eighty elks 
have sometimes been seen in one herd upon their march to the 
salines." In the vicinity of these congregating places they 
were apparently much hunted and killed, which may in part 
explain their early extermination. Rhoads says : " In the north- 
eastern Alleghenies of Sullivarr, Luzerne and Wyoming counties 
they seem to have totally disappeared in the second decade of 
the 19th century, although a few remained in a favorite haunt 
called 'Elk Forest' in the Pocono range of Wayne Co. until 
exterminated between 1830 and '40. In Tioga, Lycoming and 
Potter counties they haunted the headwaters of Pine Creek 
and the Black Forest until 1862, when the last was killed. In 
Somerset and Bedford counties, where the mountain glades 
and saline or sulphur springs were sought out by numerous 
bands of wapiti and buffalo in early colonial times, their ex- 
termination must have been of very early date . . . Even 
more obscure is the evidence of their former occurrence in the 
southwestern counties of Pennsylvania, and in the parts of 
New Jersey pertaining to the valley of the upper Delaware. " 

Stragglers, however, driven from the Pennsylvania moun- 
tains are believed occasionally to have been taken in northern 
New Jersey in former days. A writer in 1835 (R. C. Taylor, 
in London's Mag. Nat. Hist., vol. 8, p. 536) says that at that 
time elk were almost extinct in Pennsylvania; nevertheless a 
few continued for a number of years and were shot by hunters 


at the salt licks, or were tracked down with the help of dogs 
and shot. It was not thought unusual for a hunter to follow a 
stag 40 miles before finally bringing it to bay. Shoemaker 
(1915) writes of these final captures: "Taken by sections, the 
last elk in the Blue Mountains was killed about 1800; in the 
Pocono Mountains in 1845; in Lacka wanna County five or ten 
years earlier. Caleb Mitchell killed the last elk of the Seven 
Mountains at the head of Treaster Valley, Mifflin County, in 
1857; James David killed the last elk in Clearfield County in 
May, 1865. It was brought to Lock Haven on a raft from the 
mouth of Medix Run, where it was killed . . . Jim Jacob - 
son, a half-breed Indian, killed an elk in Elk County in 1867," 
which according to Rhoads is the last to have been shot in the 
State. Shoemaker (1915), however, says that this same 
Jacobson killed others "annually until Nov. 19, 1875, when he 
killed his last near Roulette, Potter County," and adds that 
the very last Pennsylvania elk was shot on September 1, 1877. 
Up to the middle of the last century "there was quite a thriving 
business of catching elks alive in northern Pennsylvania. " 

In the adjacent State of West Virginia wapiti were common 
till about the middle of the eighteenth century and thereafter 
dwindled in numbers up to about the time of the Civil War. 
Kellogg (1937), in summing up their brief history, notes that 
"between 1830 and 1835, elk were killed at a deer lick near 
'The Sinks' on Gandy Creek . . . Three elk were killed 
on the Black Fork of Cheat River near Davis, Tucker County, 
in 1843 . . . During 1845, seven elk were seen near 
Durbin, Pocahontas County." McWhorter in a historical 
account of the region states that an elk was killed in 1867 at 
Elk Lick on Middle River, in the same county, and tracks were 
said to have been seen even in 1873. At the present time a few 
elk, escaped from an enclosure in Marlinton, are said to be at 
large on the ridges of the eastern part of the State, but these 
were imported. 

Still farther south, C. S. Brimley (1905) states that this 
animal doubtless occurred in North Carolina a century and a 
half previously, and it seems also to have been present in 
Virginia in the years after early settlement, up to about 1847 
(Audubon and Bachman, 1846-54). For Tennessee, Kellogg 
(1939) has assembled evidence to show that "elk at one 
time were plentiful . . . occurring not only in the high 


passes and narrow valleys of the mountainous sections but 
also in association with the buffalo visited the licks of middle 
Tennessee, browsed along the rivers and creeks in the southern 
counties, and wandered through the canebrakes of the Missis- 
sippi bottomlands." Crockett refers to them repeatedly be- 
tween the years 1820 and 1830 as then present in the bottom- 
lands of Ob ion and Dyer Counties. The last records as given 
by Rhoads (1896) seem to be: About 1849 one killed at Reelfoot 
Lake by David Merri wether and one reported killed in 1865 in 
Obion County. Doubtless a few must have reached the 
mountains of northern Alabama as evidenced by the place- 
names, previously mentioned, and may have wandered into 
the adjacent part of Georgia, but no contemporary record is 
known to me. 

West of the Alleghenies to the edge of the Plains this race of 
elk was formerly common. Historical accounts of Ohio tell of 
its presence in the pioneer days of settlement, but it seems to 
have been much reduced by the end of the eighteenth century. 
Kirtland, however, writing in 1838, says it was "frequently to 
be met with in Ashtabula county until within the last six years. 
I learn from Col. Harper, of that county, that one was killed 
there as recently as October "of the present season." There 
were elk in Kentucky in pioneer days, affording the early 
settlers a desirable source of meat and leather but they were 
early reduced in numbers. Audubon wrote that when he first 
settled in that State (in 1808) there were still some to be found 
and a few in Illinois across the Ohio River, but by 1847 they 
seem to have been pretty well gone. Cory has assembled many 
older accounts for Wisconsin and Illinois. In 1818 they were 
no longer to be found east of the Illinois River, in Illinois, 
though a few seem to have remained in the northern part of 
the State. In southern Illinois they are said to have been com- 
mon up to about 1820; the last records in Indiana are of one 
killed in Knox County in 1829 and another in the following 
year. In Wisconsin they apparently held on till much later. 
Hoy wrote in 1882 that they were found on Hay River in that 
State in 1863, but Bray ton, in 1882, said they were still found 
in the vicinity of Green Bay, Wis. They seem to have been 
"numerous" in eastern Michigan as late as 1860 in Huron and 
Sanilac Counties about the headwaters of Cass River, but so 
relentlessly were they being hunted with rifle and trap pens 


that their early extermination was foreseen by Miles. In 1856 
they were not uncommonly seen at Saginaw Bay. Elk were 
formerly abundant in Minnesota, and according to Herrick 
(1892) they were occasionally killed by the Indians to the north 
of Lake Superior as late as 1885, and he was informed of their 
presence in that year at Red Lake. Although Cory (1912) 
stated that a few individuals were said still to remain in the 
extreme northern part of Minnesota, no evidence of this is 
given. Cahn (1937) says that in Quetico Park, adjoining the 
Superior National Forest in that region, elk have long been 
extinct, but that in 1914 or 1915 they were introduced into the 
latter reserve, and on at least one occasion "some members of 
this herd were seen in the Quetico Park" on the Canadian side 
of the common boundary. At the present time, according to 
a Department of Agriculture press statement dated January 29, 
1939, there are in Minnesota some 45 elk, and in Michigan 5, 
but what part of these, if any, represents the native stock is 
uncertain. From the same source it appears that estimates of 
elk populations show the following figures : In New Hampshire, 
250; in New York, 100; in Virginia, 140. Presumably these are 
all from stock introduced fronrthe West. 

How far to the westward the typical eastern elk ranged may 
perhaps never be demonstrated. From analogy with climatic 
conditions and from what is known concerning other animals 
of wide distribution, it may be presumed that somewhere 
along the eastern edge of the Great Plains the characters 
showed intergradation with those of the Rocky Mountain elk, 
so that available notes from the States of the eastern Plains 
may be grouped under that race. In North Dakota, where 
this transition may have been found, Bailey (1926) writes of 
the former abundance of elk all over the State, where they were 
equally at home in the timber and on the open prairie. Ex- 
plorers and others tell of the numbers found at the beginning 
of the last century, but they rapidly disappeared before on- 
coming settlement. A few remained into the early eighties, 
one of the last reports being that of six killed in 1883 near 
Elkton in Cavalier County. In former times the valleys seemed 
to be their wintering grounds, where they found a dense cover 
of timber and abundant food. The antlers were shed by the 
males mainly during March and April, and quantities of these 
might be picked up. "Next to the buffalo," says Bailey, "the 



elk at the height of their abundance were the easiest to hunt 
and hence the most rapidly killed of the large game, but when 
much hunted they became very wild, and it is probable that 
besides the vast numbers killed in the State, many were driven 
out of its borders." Running down elk on horseback was a 
favorite form of plains sport. The meat was in great demand as 
about the best of game animals, while the hides were much 
used as leather not only by the Indians for moccasins but by 
the whites for various purposes. The upper canine teeth were 
a prized decoration of Indian women, and in later times were in 
great demand as watch charms among the whites, a demand 
that for a time led to much ruthless slaughter. 

The possibility of raising elk commercially under fence has 
been somewhat developed and is the subject of more recent 

Eastern wapiti (Cervus canadensis canadensis} 


experiment. Bailey (1926) says that "in domestication elk 
have proved more hardy and prolific than other stock and al- 
most as easily handled under well-fenced range. If in the future 
the production of elk meat proves as profitable an industry as 
it promises, there will be found ideal conditions for elk pastures 
in many parts of western North Dakota, where rough and 
steep slopes lie close to brushy bottomlands, and winter browse 
and summer grass can be inclosed in single or adjoining areas. 
The severe winter weather which means suffering and loss to 
domestic stock without shelter is a joy to these native born and 
bred deer if a suitable and adequate food supply be available. 
Along many of the stream valleys with Badlands borders, 
which now lie idle or are of little use for stock, elk would find 
an abundance of their favorite food and choice living condi- 
tions. The time seems ripe for adding this industry to the 
many resources of the State." 


Cervus canadensis nelsoni Bailey, Proc. Biol.^Soc. Washington, vol. 48, p. 188, Nov. 15, 

1935 ("Yellowstone National Park," Montana). 
FIGS.: Elliot, 1901, pi. 12 (skull and antlers); Nelson, 1916, p. 454 (col. fig.). 

The wapiti of the Rocky Mountain region, from northern 
New Mexico northward into Canada, and including formerly 
perhaps the Great Plains area, is regarded as a race larger in 
size of body and in antlers than that of the eastern woodlands, 
and in color it is the palest of the races except the dwarf elk 
(C. c. nannodes) . Bailey describes the type as light buffy fawn 
in summer, fading to creamy buff; rump-patch creamy buff or 
whitish; head, neck, legs, and belly dull rusty brown to dark 
umber and blackish; eye-rings buffy. In winter, the body 
color is buffy gray over lavender, with dusky tips that wear 
off leaving a creamy or soiled-whitish appearance; large rump- 
patch, including the tail, whitish or clear white; head and neck 
dull rusty brown with dark-brown manes, darkest on lower 
throat; ears dull light brown, lined with pale buff. The basal 
length of the skull of the type (male) was 430 mm.; length 
of antler over the beam, 1,250-1,260 mm. 

It may now be impossible to ascertain where this race 
intergraded with the eastern form or whether the range of the 


Rocky Mountain wapiti should be regarded as extending east- 
ward to the edge of the Great Plains. At all events, the species 
formerly was found over much of this region. Thus, in 1804-5, 
Lewis and Clark recorded elk along the Missouri River all the 
way through North Dakota, and 40 years later Audubon 
found them abundant along the Missouri and Little Missouri 
Rivers. Bailey (1926) mentions being told of "thousands" 
seen along the Lower Yellowstone in 1864 and they remained 
common in the Dakotas till about 1880, but "as the country 
filled up with settlers they rapidly disappeared," and were 
practically gone from the State by the middle eighties. The 
late J. A. Allen (1870) wrote of their occurrence in Iowa, that 
though formerly numerous, they were by 1879 "extinct in 
most of the region." An old resident with whom he talked in 
1867, at New Jefferson, Greene County, told him that only 
seven years before "the elk were abundant in some parts of 
that county. Prior to this date he used to see herds nearly 
every day, and sometimes several in a day, some of them of 
very large size. During the early settlement of this part of 
Iowa they were of great value to the settlers, furnishing them 
with an abundance of excellent food when there was a scarcity 
of swine and other meat-yielding domestic animals. But, as 
has been the case too often in the history of the noblest game 
animals of this continent, they were frequently most ruthlessly 
and improvidently destroyed. In the severer weather of winter 
they were often driven to seek shelter and food in the vicinity 
of the settlements. At such times the people, not satisfied with 
killing enough for their present need, mercilessly engaged in an 
exterminating butchery. Rendered bold by their extremity, 
the elk were easily dispatched with such implements as axes 
and corn-knives. For years they were so numerous that the 
settlers could kill them whenever they desired to, but several 
severe winters and indiscriminate slaughter soon greatly re- 
duced their numbers, and now only a few linger where formerly 
thousands lived, and these are rapidly disappearing. Their 
home here being chiefly the open country, they much sooner 
fall a prey to the Vest ward march of civilization', through the 
most merciless treatment they receive at the hands of the emi- 
grant, than does the deer." 

On the plains of Kansas the elk was formerly abundant but 
has long been extinct. In these open regions elk were con- 


spicuous because of their large size, while the excellence of their 
meat made them an object of pursuit by the pioneers. Add to 
these points their gregarious habits, the dangers to which 
they were exposed through severe winters, prairie fires, and 
diseases, and it becomes obvious that they would be among 
the first to go in the face of these perils. Shortly after the 
Civil War, with the increase of settlers over the prairie country, 
elk became reduced in numbers on the Plains until they were 
left only in the more rugged country in the foothills of the 
Rocky Mountains. 

This race of elk reached its southern limits in the mountains 
of extreme northern New Mexico. In the San Juan Mountains 
they were reported as abundant by Pike in 1807 and by Cope 
as not uncommon in 1874, but in succeeding years these num- 
bers seem to have quickly dwindled, although they were still 
to be found in 1892. In September, 1909, forest rangers 
reported two elk in these mountains. Bailey (1931), who re- 
cords these facts, was unable to learn of elk in the Jemez 
Mountains when he was there in 1906. Nelson, who in 1883 
was staying near the head of Pecos River, had a few reports 
of elk in the nearby mountains at that time, but the last 
record that Bailey could obtain was of one killed in 1902. 
While probably the native stock was by then wiped out, 
several attempts at restocking have since taken place, by im- 
porting animals from Colorado and Wyoming, as in 1911, 
when small numbers were "planted" in Potato Canyon, 
Cimarron Canyon, Gallinas Canyon, and later in San Miguel 
County. These introductions, in part under fence, were re- 
ported as doing well and increasing slowly. With the setting 
aside of several game reserves, it seems likely that a stock may 
eventually be built up. 

In Colorado, elk were early exterminated in the eastern parts 
and in 1871 were already rare even in Park County in the 
eastern hills. Cary, writing in 1911, says that they were then 
exterminated over much of their former range in Colorado, 
and "the few bands which remain in the wildest parts of the 
western plateaus and mountains are small and widely scat- 
tered . . . Estimates in 1898 placed the number of elk 
in Colorado at 7,000; in 1902 at 3,000. In 1909 their numbers 
were reduced to considerably less, and were divided about 
equally between northern and southern Colorado . . . 


Careful estimates made by the Forest Service officers in the 
spring of 1911 show a total of about 2,100." Since Gary's 
report in 1911, the herds seem to have been built up very 
greatly, for the U. S. Biological Survey in its census published 
in 1939 gave an estimated total of 23,000 elk for Colorado. 

At the present time, owing to adequate protection in recent 
decades, the elk herds of Wyoming, Idaho, and Montana 
have increased to such large proportions that they are no 
longer in danger. The chief centers of abundance are in such 
national parks as the Yellowstone, where formerly hundreds 
were slaughtered by hunters; another famous place is the 
Jackson Hole region, Wyoming, a regular wintering area for 
elk coming down off the surrounding mountains where they 
spend the summers. In Glacier Park, Montana, there are 
smaller numbers, but these are apparently increasing. One 
difficulty in maintaining elk in large preserves is, as Bailey 
points out, the problem of providing upland mountainous 
areas for their summer range and suitable lowland places 
where food is available for wintering them. The 1939 wildlife 
census provides the following figures for elk populations in the 
Rocky Mountain States, from south to north: New Mexico, 
5,000; Colorado, 23,000; Utah, 3,800; Wyoming, 40,700; 
Idaho, 24,400; Montana, 26,700; Nevada, about 200. 

In addition to these there are a good many elk in other 
States that have been introduced from the Rocky Mountain 
area. Some of these are living under fence, and others may be 
free. Thus the report above quoted gives the following: 
Arizona, 4,400; California (in several places in the northern 
part); Michigan, 5; Nebraska, 31; New Hampshire, 250; New 
York, 100; Oklahoma, 230; South Dakota, 3,400; Texas, 350; 
Virginia, 140. The Rocky Mountain elk is obviously in no 
present danger. The feasibility of restoring elk to regions 
from which they have been extirpated is evident from the 
following quotations (gathered by Dr. Francis Harper). Ac- 
cording to Barker (1936, p. 177), in New Mexico, "elk, com- 
pletely exterminated about 1900, have been restored so that 
we now have about 5,000 animals." "Prior to 1912, elk had 
disappeared from the ranges of the State of Utah. Since 1912 
there have been planted within our State, a few at a time, 
193 head in seven different areas. These plantings were made 
during the period from 1912 to 1925, and the elk were imported 


from Jackson Hole and Yellowstone Park . . . We now 
have approximately 3,500 head on the ranges" (Cook, 1936, 
p. 187). In Wyoming, Montana, and Idaho they have been 
completely saved from extinction through adequate protective 
measures. In 1932 there were, in national parks of the United 
States, an estimated total of 15,420 and in national forests 
over 96,800 (Phillips, 1935). 

In a recent survey of the elk situation, it is shown by Skinner 
(1928) that in the Yellowstone region, for example, any ade- 
quate plan for maintaining the elk population should consider 
providing a sufficient food supply for winter as well as summer. 
This seems to consist under natural conditions of "browse" 
as well as grass, so that the hay supplied in recent years for the 
Jackson Hole herd in winter is probably not an adequate 
diet. He gives a partial list of food plants used from month 
to month. Protection against unusually heavy snows and cold 
is another problem, for when elk are forced out from Yellow- 
stone Park into the surrounding low country in winter, thou- 
sands are killed by hunters, without supervision or restraint. 
There is the further problem of competition with bison herds 
of the region for food, the depredations of coyotes, wolves, 
and mountain lions, though these are less important. Finally 
diseases, some of which may be introduced through domestic 
stock, and parasites are as yet insufficiently known. Murie 
(1930) has made a special study of winter losses among the 
elk at Jackson Hole, Wyoming, and attributes a large portion 
of them to necrobacillosis, the symptoms and general etiology 
of which he describes and illustrates. The sharp awns of the 
squirreltail grass, which produce lesions in the skin of mouth, 
tongue, or throat and so afford entrance for the disease organ- 
isms, seem to be a contributory factor in this. The elimination 
of this grass, if that is possible, might aid in preventing the 
trouble. All these problems are matters of importance in 
maintaining the elk herds and increasingly demand careful 
consideration. A beginning in this had already been made by 
the U. S. Biological Survey, in concert with the Forest Service 
and a program of management laid 30 years ago (see Graves 
and Nelson, 1919). The present excellent condition of the 
herds in Yellowstone Park reflects the wisdom of the policies 

A decade ago, during prohibition days, a number of elk 


were imported into eastern Massachusetts and ostensibly 
maintained for breeding and experimental purposes, until it 
developed that the pine grove in their spacious and well- 
guarded paddock concealed a large illicit still! 

North of the United States- Canadian Boundary this race 
of elk extended its range in former times to about latitude 60 
nearly to Lake Athabaska but was driven back in the last 
century. Nevertheless there are still "some numbers in East 
Kootenay district, British Columbia, where they are hunted 
to some extent . . . Some small herds of typical Rocky 
Mountain elk have within the past few years been demon- 
strated to occur on some of the southern tributaries of Nelson 
River in northeastern British Columbia" (Anderson, 1939b). 
They are, however, apparently now gone from the Peace River 
region where they formerly occurred. In a manuscript report 
of Harry Snyder to the Canadian Secretary of the Interior, 
as communicated by Dr. Francis Harper, the former tells of the 
discovery of a herd of 150 to 200 wapiti on the Prairie and 
Henry River watersheds of northern British Columbia, an area 
comprising 750 to 800 [Psquare] miles, about 500 miles north of 
the nearest known area now inhabited by elk. 


C\ervus\ c\anadensis\ manitobensis J. G. Millais, The Gun at Home and Abroad, vol. 4, 
p. 281, 1915 ("Manitoba and eastern Saskatchewan," Canada). 

The elk in the most northerly part of its range in North 
America are "apparently larger and duller colored than typical 
canadensis; not so large or light colored as nelsoni of the Rocky 
Mountains. In summer rich chestnut brown, darker on head 
and neck and belly; in winter somewhat lighter. The name 
manitobensis, applied to this northern animal by Millais in 
1915, is regarded as valid by Bailey (1935) for the race found 
in Manitoba and Saskatchewan and presumably still farther 
to the westward. Its precise area of distribution, however, 
is somewhat uncertain. In former times the wapiti, whether of 
this race or nelsoni, ranged to the Rocky Mountains on the 
west and northwestward, to the plains of Peace River, where 
Alexander Mackenzie found it in abundance near the trading 
post that he established at the mouth of Smoky River in the 


autumn of 1792 (see Preble, 1908). In that day it ranged as 
far north as Fort Nelson River, according to Richardson, but 
the nineteenth century saw its numbers so depleted that in 
1894 Caspar Whitney placed its northern outpost as on the 
Saskatchewan, between Edmonton and Lac la Biche. Loring, 
in 1896, reported that a few were said still to exist near the 
head of Pembina River, where, however, they had been nearly 
exterminated by Indians through "crusting." Preble (1908), 
whose account summarizes these facts, adds that a few were 
seen on the south side of the Athabaska River in 1897. The 
most recent account of the status of this race is that of Dr. 
R. M. Anderson (1939b), who writes that in 1923 there were 
estimated to be 400 or 500 elk at Riding Mountain, Manitoba, 
and other bands on Duck Mountain Forest Reserve, but none 
elsewhere in Manitoba. The Riding Mountain herd in seven 
years under careful protection had by 1930 increased to about 
8,000 head, but in the severe winter of 1934 the heavy mortality 
cut this down to about 6,000, with an estimated 300 in Duck 
Mountain Reserve, 250 in Porcupine Reserve, and 600 in the 
Inter-Lake district. In 1923 the Provincial Game Guardian of 
Saskatchewan estimated that in* the Moose Mountain Reserve 
in that province there were about 100 elk, and in the region 
north and east of Prince Albert about 1,200 more. In 1927 a 
short open season was allowed here in the second half of Novem- 
ber. The Prince Albert National Park, established that same 
year, "now shelters numbers of elk, but the exact figures are 
not known, and there are several bands outside the Park." 
It thus appears that in these two provinces, to which this race 
may be considered restricted at present, there are something 
over 8,000 head, largely in government reserves under careful 
supervision, so that in these areas there is as yet no fear for 
the elk's extinction. 



Cervus merriami Nelson, Bull. Amer. Mus. Nat. Hist., vol. 16, p. 7, Jan. 16, 1902 

("Head of Black River, White Mountains, Arizona"). 
FIGS. : Nelson, 1902, figs. 1, 3, 5 (skull); Bailey, 1931, pi. 3 (antlers). 

Equaling in size the Rocky Mountain elk, this race is de- 
scribed as paler and more reddish in color, with more massive 


skull and more erect antlers; the nasal bones are remarkably 
broad and flattened, with a well-marked constriction in the 

Fifty years ago this elk was common in the Sacramento, 
White, and Guadalupe Mountains east of the Rio Grande 
River and in the Mogollon group of mountains west of it, in 
southern New Mexico, as well as in the White Mountains of 
Arizona. There are old records for the Datil and Gallina 
Mountains of Socorro County, New Mexico, and a doubtful 
record for the Manzano Mountains. At the present time this 
race, with a restricted range in the mountains of southern 
New Mexico and Arizona, is believed to be extinct (Bailey, 

Bailey has summarized the history of this deer. He surmises 
that it was antlers of this race that Montezuma showed to the 
followers of Cortez. That it was once common in southern 
New Mexico is inferred from the statement published by J. A. 
Allen in 1874 that a trapper and guide in the Rocky Mountain 
region had met with droves of 2,000 individuals as far south 
as the Mexican boundary. In the White Mountains and the 
Blue Range of Arizona, "more particularly on the head of 
Black River," according to D. B. Rudd, a forest ranger, they 
were present in large bands in 1876 when his father moved to 
that part of the country. "As late as 1890 elk could be found 
but not so plentifully. Since the year 1895 I cannot find that 
any have been seen. " Nevertheless, another ranger had seen 
"bedding signs" of a small band between Black River and the 
higher plateau of the Blue Range, about 1904, which seems to 
be the latest report of them for that region. In the Mogollon 
Mountains H. W. Henshaw found tracks of these elk in 1873, 
and in 1882 Nelson heard reports of their presence near the 
extreme headwaters of the Gila River. In 1886 he collected 
specimens in the White Mountains of Arizona. Reports of the 
forest rangers indicate that these elk became scarce in the 
Mogollon Mountains about 1888, although up to that time 
they seem not to have shown any alarming decline. Bailey 
adduces a few instances of animals shot in the Mogollon Moun- 
tains up to 1890, when a cow and a bull were killed. The last 
record, however, seems to have been in 1894, when a fine male 
was reported seen on Lily Mountain, north of the main peaks 
of the Mogollons and tracks of three later in the same year. 


In the Mimbres Range Bailey was unable to find any tradition 
of elk, nor in the Datil Mountains could Hollister in 1905 find 
any ranchers who remembered seeing them, although old 
antlers were occasionally discovered. A similar story of traces 
of their former presence was told in the Bear Spring and Indian 
Spring Mountains, which probably once formed the northern 
boundary of this elk's range. In the Sacramento Mountains 
elk were said to have been killed up to 1898, but Bailey him- 
self, in 1902, could get no record of elk killed or even seen 
there later than 1893. 

Concerning these elk in the White Mountain region of 
Arizona, Nelson (1902) wrote that their main range covered an 
area about 30 by 50 miles in extent, at an elevation 8,000 to 
10,000 feet above sea level. At that time, between 1885 and 
1888, "elk were far from numerous" but seemed to be most 
often found "about some beautiful damp meadows in the 
midst of the dense fir forest on the rolling summit of the 
Prieto Plateau, between the Blue and the Black Rivers." 
W. W. Price, who collected in this region in 1894, saw several 
and shot a male at 9,000 feet elevation. The report of tracks 
seen here in 1901 by a local httnter, who followed them un- 
successfully for two days, indicates that a few survived until 
that time, perhaps till about 1904, as previously noted. The 
fact that this survivor was trailed for two days shows that elk 
were still hunted in spite of legal prohibition. 

In the Chiricahua Mountains of southeastern Arizona 
these elk in early times were numerous, but with the extension 
of grazing interests in the latter part of the last century there 
came "a long slow reduction in numbers due chiefly to the 
elks' inability to compete with cattle on the over-grazed range." 
Hunting accomplished their final extermination in the region, 
for the last small band was killed in the vicinity of Fly and 
Chiricahua Peaks about 1906 (Cahalane, 1939a). 



Cervus nannodes Merriam, Proc. Biol. Soc. Washington, vol. 18, p. 23, Feb. 2, 1905 

(" Buttonwillow, Kern County, California"). 
FIG.: Merriam, 1905, p. 23, fig. (from photograph of a male). 

Although called "dwarf elk," this race of southern Cali- 


fornia is after all only slightly smaller than the form of the 
Rocky Mountains. Compared with the latter it is paler, with 
a smaller and narrower rump-patch. The legs are shorter, 
and the skull is slightly smaller but has longer palatal bones 
in proportion. Premolar and molar teeth are of the same length 
in both, hence relatively larger in nannodes. Antlers in general 
similar to those of the Rocky Mountain form but smaller and 
with the posterior terminal prong less developed. "Front of 
legs and feet bright golden fulvous; back and flanks varying 
from buffy gray, slightly washed with fulvous, to grizzled 
buffy whitish"; inner side of the ears "buffy white, the white, 
particularly at posterior base, much more extensive than in the 
other" races. The type specimen, a two-year old male, had a 
total length of 2,030 mm.; tail, 140; hind foot, 620; basilar 
length of skull, 358 mm. as against 388 in a male of same age 
representing the Rocky Mountain form. 

Prior to 1860 this form of elk was common in nearly the 
entire San Joaquin and Sacramento Valleys, California, 
especially in their lower parts. It was found at least as far 
north as Butte Creek, in Butte County, and south to near 
Bakersfield, Kern County, and westward through the southern 
inner coast ranges as far as the plains of the Cuyama Valley, 
San Luis Obispo County, and extreme northern Santa Barbara 
County, and to the south end of San Francisco Bay, Santa 
Clara County (Grinnell, 1933). "The encroachments of civili- 
zation have resulted in the gradual extermination of this elk 
over the greater part of its former range, until" by the end of 
the last century it had become "restricted to a small area 
between Tulare and Buena Vista lakes, where at present 
[i. e. in 1905] the survivors are confined almost exclusively to 
lands included in an extensive cattle ranch (Buttonwillow 
Ranch) owned by Miller and Lux. " About 1903 these owners 
presented the herd to the United States Government and a 
park for its reception was constructed on Kaweah River in 
the Sequoia National Park. In November, 1904, an attempt 
was made to round up and corral this herd and move the 
animals to the new area, but it was not a success, for "the elk 
refused to be driven and escaped to the adjacent foothills of the 
Temploa Mountains. " During the process several were roped, 
with fatal results, but the skins and skulls were preserved for the 
U. S. Biological Survey collection (Merriam, 1905). Since 


then, apparently, the remnant of this herd has continued, wild 
or in part under fence, in "western Kern County between 
Tulare and Buena Vista lake basins and at times in adjacent 
hills to westward. In 1932, about 170 individuals remained" 
(Grinnell, 1933). "Many transplan tings of Dwarf Elk from 
the remaining 'herd' in the Button willow district to other parts 
of the State have been made, first to Sequoia National Park, 
Tulare County, in 1904 and 1905, and more recently to Yosem- 
ite Valley, to Monterey County, etc. None of these has re- 
sulted in establishment under conditions of really wild freedom. 
The Sequoia animals had entirely disappeared by 1926" 
(Grinnell, 1933). With the protection now accorded them, 
unless local requirements change, it seems likely that this 
remnant in the Buttonwillow district may continue for a long 



Cervus roosevelti Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 271, Dec. 17, 1897 
("Mt. Elaine (on ridge between heads of Hoh, Elwah, and Soleduc rivers) near 
Mt. Olympus, Olympic Mts., State of Washington"). 

SYNONYM: ? Cervus occidentalis Hamilton Smith, Griffith's Cuvier, Animal Kingdom, 
vol. 4, p. 101, 1827. [Not certainly applicable to any American deer (Bailey)]. 

FIGS.: Bailey, 1936, pi. 20 (photographs of wild individual and a herd). 

This elk of the Northwest coast is perhaps the largest of the 
races, the males with massive skull and antlers that are often 
"cupped," that is, the three terminal tines tend to come off 
together. The beam of the antler is relatively short and 
straight, with the terminal prong reduced. The body color in 
summer is a rich cinnamon-buff; head, neck, and belly dark 
brown with much black; in winter the body is dark gray with 
a dusky dorsal stripe and with dusky on the face, mane, and 
belly. In the skull, the frontals are broad and much flattened, 
and the preorbital cavity is small, as compared with the Rocky 
Mountain race. Total length of an adult male, 2,490 mm.; 
tail, 80; spread of antlers, 990 (3 feet 3 inches); length of left 
beam (in type), 1,050 (41.25 inches). The length of antler 
is about 500 mm. (20 inches) shorter than in a Rocky Mountain 
elk of comparable development. 

The range of this race is confined to the humid forest belt 
along the Pacific coast from northern Vancouver Island south 
through the coast ranges of Washington, Oregon, and Cali- 


fornia to (formerly) about San Francisco Bay. Eastward in 
California the range included Mount Shasta. Altitudinally 
the area inhabited extended in California to about 7,000 feet 
(Grinnell). On Vancouver Island, its northward limit, it was, 
according to Swarth (1912), formerly abundant everywhere 
in the forests, decreasing in numbers southward to within 30 
miles of Victoria. "They keep well back from the settled 
districts and are quite scarce near the east coast and the ad- 
jacent woods. They are most abundant in the north end of the 
island, particularly in the northwestern section and the vicinity 
of Kuyuquot Sound . . . The wapiti are seldom hunted 
by the Indians, who prefer the more easy task of killing deer on 
the beaches." In the interior of Vancouver Island the wapiti are 
still fairly safe, since the forests are relatively unexplored and 
the animals are little disturbed (Sheldon, 1912). The Govern- 
ment took active steps to stop the killing of these wapiti for 
their canine teeth when, about 30 years ago, they were in 
active demand. Dr. I. M. Cowan, in 1937, told Dr. Francis 
Harper that there were supposed to be only about 700 elk left 
on Vancouver, this decimation being due in part to the ravages 
of wolves and cougars, but possibly also to disease that may 
have come among them, as in the Rocky Mountains. 

This elk is not found on the mainland of British Columbia 
opposite Vancouver but appears again across the Strait of 
San Juan de Fuca in the mountains of the Olympic Peninsula 
and in scattered bands in the coast ranges of Washington and 
southward. Not many years ago its numbers seemed to have 
reached a low level in this State. Taylor and Shaw, in 1929, 
reported that in addition to the small groups found south to 
Pacific County, a small herd was said to be living on the head- 
waters of the Cispus River, south of Mount Rainier. It was 
at one time feared that this remnant might be even further 
reduced, but recent reports indicate that it is at present 
thriving under protection and is in no present danger. It 
should be borne in mind, however, that according to these 
authors Rocky Mountain elk have been introduced into several 
counties of Washington: Stevens, Garfield, and Walla Walla 
in the eastern part and Skagit, Snohomish, King, and Yakima 
in the central part. It is believed that in former times the 
ranges of these two forms did not overlap in the Northwest. 

In Oregon the status of this deer has been summarized by 


Vernon Bailey (1936), who reports that there are still con- 
siderable numbers of them in the coast ranges to which they 
are confined, from the Columbia River to the California border. 
Originally they inhabited the western slope of the Cascade 
Mountains and thence to the coast over all western Oregon. 
In 1841 Wilkes found them plentiful in the Willamette Valley, 
and Peale reported them from the mountains south of the 
Columbia River. Suckley and Gibbs about 1854 met with them 
abundantly in the mountains west of Astoria. "Later as the 
country was settled, elk were reported from almost every 
valley and mountain range of western Oregon, including the 
west slope of the Cascade Mountains, but there seems to be 
no record from the east [drier] slope of the range. In recent 
years, under rigid protection, apparently they have been 
holding their own, while in some localities actually increasing. 
The continual spread of settlement, though, is restricting their 
range, and the greater number of hunters each year makes it 
more difficult to prevent poaching in out-of-the-way places" 
(Bailey, 1936). In 1926 there was an estimated total popu- 
lation of 436 elk on the national forests of western Oregon, a 
figure that, says Bailey, covers the greater part of the Roose- 
velt's elk in the State. This is a slight increase (of about 40) 
over the estimate for 1933, but less than those for 1930 and 
1931. It may be pointed out that the 17,000 elk estimated in 
1939 as inhabiting the State of Oregon (Federal census), are 
doubtless mostly the Rocky Mountain elk, native and im- 
ported, in eastern parts of the State. 

Still farther south this elk formerly occurred in abundance 
all the way to San Francisco Bay in the humid coast belt of 
northwestern California and eastward to the nearer inner 
coast ranges and to Mount Shasta. Of recent years, however, 
its numbers have dwindled, until by 1933, according to Grinnell 
(1933), it was to be found only in Del Norte and northern 
Humboldt Counties, while the total population was believed 
not to exceed 400 head. Its vertical range extended originally 
from sea level up to at least 7,000 feet, as formerly in the Scott 
Mountains, Trinity County. Its usual habitat was in and 
about openings in forest. 

While the population of this elk is small, with a few hundred 
in Vancouver Island, and in each of these three northwestern 
States, nevertheless with the protection now given it there 


seems no reason to be apprehensive for its survival for a long 
time to come. Nevertheless, in order to preserve the race in 
its purity, care should be exercised that introductions of the 
Rocky Mountain elk are avoided within the range of this 
coastal form. 


Cervus hemionus Rafinesque, Amer. Monthly Mag., vol. 1, p. 436, Oct., 1817 (Sioux 

River, South Dakota). 
SYNONYM: Cariacus virgultus Hallock, Forest and Stream, vol. 52, p. 404, May 27, 

1899 (Near Hallock, Kittson County, Minnesota). 
FIG.: Nelson, 1916, col. fig. on p. 455. 

Unlike the white-tailed deer, this is more a species of open 
or rough brushy country, so that where its range meets that 
of the former it is found in a different type of habitat. The 
antlers, while often having a basal prong, are usually charac- 
terized by the dichotomous forking of the main beam. The 
ears are longer than in the white-tailed group; the tail is short 
and either black-tipped or largely black above (black-tailed 
deer of the Pacific coast) . The metatarsal gland is much longer 
and higher up on the foot. Both these types of deer are charac- 
terized by the extension of the vomer backward to form a 
complete longitudinal division of the posterior nasal passage; 
both also belong to the group Telemetacarpalia, in which the 
lateral digits of the forefeet are represented by the outer in- 
stead of the inner ends of the metacarpals. 

As a species the mule deer and its west coast forms, the black- 
tailed deer, are or originally were found from the eastern edge 
of the Plains to the Pacific Ocean. While most of the western 
races are still abundant locally, the eastern form is largely 
extinct and may therefore be mentioned here. Baird describes 
its color as "ashy brown, pointed or varied with gray, and 
without any rusty tinge whatever. There is a distinctly marked 
stripe from the crown of the head to the root of the tail, the 
hairs in which are much darker throughout their extent than 
elsewhere. The under parts generally appear to be ashy 
brown, like the back, but without annulation of the hairs. 
The only whitish portion of the inferior surface is seen beneath 
the head, around the axillae, and in the groin. On each side 
of the tail, on the end of the rump, is a dull white patch, 


crossing above the tail and involving its entire basal half or 
two-thirds. The tail itself is quite slender and cylindrical on 
the basal two-thirds; the hair there being compact and close 
and then expanding into a dense tuft, which is entirely black" 
(Baird, 1857). Total length from tip of nose to tip of tail 
vertebrae, about 6 feet 6 inches. 

Formerly the eastern mule deer was found in the Plains 
country from northern Oklahoma, Kansas, Nebraska, the 
Dakotas, and extreme northwestern Minnesota northward 
into southern Manitoba and westward to the foothills of the 
Rocky Mountains, where it is believed to have intergraded 
with the Rocky Mountain mule deer, 0. h. macrotis, which is 
still common in the region from northern New Mexico north- 
ward into southern British Columbia. In the Dakotas this 
was a common species in the early days, and Bailey (1926) 
has brought together many notes of their presence from the 
accounts of early travelers in the region. In the seventies of 
the last century they were more abundant than the white- 
tailed deer on the upper Missouri, but in the succeeding decade 
the increasing influx of settlers and hunters had greatly reduced 
their numbers. J. S. Weiser, emoted by Bailey, reported them 
so common in the vicinity of Valley City in 1878 "that one 
could not travel 5 miles without seeing them," and JohnHailand 
stated that though common at that time the last one was 
shot in 1885 or 1886. The latter adds: "There was so much 
venison in camp during the first years that visitors' ponies 
were usually loaded down with it before they returned. There 
was no sale for venison nor for skins, they were so plentiful. 
Skins were used for mattresses; they would get damp and 
deteriorate during summer and a new supply was provided 
each fall for the winter's sleeping." In 1896, Seton (1909) in 
a 15-mile ride across the Badlands of the Little Missouri, 
counted only three where ten years before he and his companion 
had counted 160 over the same ground. The numbers seem to 
have declined rapidly in the early years of the present century. 
Bailey states further that in 1912 they were reported rare in 
the Turtle Mountains of North Dakota, but just across the 
line in Manitoba they were more common and a number were 
killed each year. In 1913 Jewett found them "still fairly 
common" in the Badlands along the Little Missouri, but in 
the Killdeer Mountains he found that all had been killed off 


near settlements. He was told that at that time they were 
found only in the rougher parts of the Badlands, where a few 
may still persist, "but, if not already extinct, this finest of all 
native species of the smaller deer will soon have vanished from 
the State" (Bailey, 1926). They are apparently rare also in 
South Dakota. 

There were mule deer in Nebraska in the seventies, but their 
numbers have been much reduced, and from Kansas they are 
quite gone. At the end of the last century a few existed in the 
open northwest corner of Minnesota, from Red Lake to Lake 
of the Woods, but they were exterminated soon after. In 1911 
Gary wrote that "apparently none remain on the plains east 
of the mountains" in Colorado, "where they were common in 
early times." The mule deer of the mountainous areas here 
are perhaps of the race macrotis, although, as Bailey says, so 
few specimens of the typical plains form are in existence that 
we may never know its precise characters. 

In the 1939 estimate of game animals by the U. S. Biological 
Survey, the following figures are given for the mule deer in the 
States east of the Rockies where it formerly was found: 
Oklahoma, none; Kansas, none; [eastern Colorado, gone;] 
Nebraska, 350; South Dakota, 4,900; North Dakota, 100; 
Minnesota, none. Although the basis of the figures for South 
Dakota is not given, it is likely that their center is in the 
Black Hills region, now a national park. Here Dr. Walter 
Granger reported them "numerous" in 1895 although "about 
extinct in the Bad Lands" (see J. A. Allen, 1895). With 
strict protection it seems likely that they might be permanently 
preserved in such areas as parts of the Badlands afford, with 
due consideration of the fact that they are wont to move down 
from the higher hills in autumn to winter in the more sheltered 
places of the lowlands. 



Dama virginiana Zimmermann, Specimen Zoologiae Geogr., p. 351, 1777 ("America," 

assumed to be Virginia). 
FIGS.: Elliot, 1901, pi. 15 (skull), as 0. americanus; Nelson, 1916, p. 458, upper fig. 


The white-tailed deer is found in woodlands of the temperate 


zone quite across the United States, barely entering southern 
Canada; to the southward it extends to Central America and 
northern South America. Over this wide range it develops 
several local races, some of which are in danger of being un- 
duly reduced. A brief survey of these is here given, since this 
animal is one of our most important game species. 

The total length from nose to tail is given by Baird as 59 
inches for a doe from Virginia; tail about 9 inches. Males are 
slightly larger and heavier, weighing up to 200 pounds. In 
winter coat, "pale grayish chestnut, faintly annulated"; in 
summer "bright uniform rufous" the "blue" and the "red" 
coats of hunters. Under parts of body and tail, inside of the 
limbs, the area between the jaws, and lining of ears, white; 
a pale eye-ring. Tips of ears and a spot at the angle of the 
mouth black. Antlers with a basal prong, the beam curving 
up, out, and then slightly inward and forward, usually with 
three erect prongs, but often with additional small points. 

The typical race of Virginia deer is believed to range from 
central or southern Pennsylvania south to the region of Palm 
Beach in eastern Florida; westward it extends at least to the 
Mississippi basin in Ohio and Kentucky, and formerly to the 
edge of the Plains. Over most of this range it still is found in 
woodland regions and survives in spite of persistent hunting, 
even in fairly well-inhabited areas. 

In New Jersey and southern Pennsylvania the white-tailed 
deer formerly abounded, but Rhoads (1903) in the early years 
of this century writes of it as confined to limited areas in 
southern New Jersey, while from the lowlands of the Susque- 
hanna, Allegheny, Monongahela, and Delaware River Valleys 
it was exterminated. In New Jersey it was found then in 
Burlington, Atlantic, Cape May, and Ocean Counties in small 
numbers. At the same time in West Virginia deer were "still 
rather plentiful" though much less abundant than in earlier 
times (F. E. Brooks, 1911), and this is generally true for other 
nearby States. Thus C. S. Brimley (1905) reports it rare or 
absent in the more thickly settled parts of North Carolina 
but "not uncommon" in the eastern section of the State and 
in the mountains. Dr. Francis Harper (1937) notes that in 
the Okefinokee Swamp region of southern Georgia it was 
"abundant in former times" but "had become greatly reduced 
in numbers through excessive hunting in late years, even before 


the cutting of the timber and the great increase of the human 
population" of the area. On account of its ready adaptability 
it can maintain itself in varied cover and persist even in prox- 
imity to settled districts if given a reasonable chance. With 
two young at a birth its numbers readily come up with pro- 
tection, so that even where much hunted it will continue in 
the face of considerable persecution. In the Great Smoky 
Mountains National Park it is said that there were believed 
to be "not more than a score of deer" remaining in this region 
when it was established in 1931, but already they show some 
evidence of coming back (Campbell, 1939). Kellogg (1939) 
summarizes records showing the "incredible number" of deer 
formerly found in parts of Tennessee and the importance of 
their meat and hides to the early settlers. By the latter part 
of the last century they had been so reduced in many areas of 
the State that the "Cumberland Mountain range has been 
almost entirely depleted of its stock of deer. " Action of the 
legislature in 1895 prohibiting their killing for five years in 
five of the counties of Tennessee and the later prohibition of 
their hunting in the Great Smoky park will doubtless save the 
remnant. In 1882, according to Brayton, the deer was then 
"rarely met with in Ohio," and the same story holds elsewhere 
of its great depletion in the eastern States. Even in Alabama, 
Ho well (1921) tells that they "once ranged in large numbers 
over all" parts of the State but are "now exterminated in all 
but the wilder and more inaccessible" districts. 

As a good example of what may be done, however, is the 
result reported by F. B. Chapman (mimeographed report of 
the Ohio Wildlife Research Station, release 105, 1939). Deer, 
almost if not quite extirpated from Ohio by 1904, were re- 
introduced in 1922-30, and under protection they have so 
increased that by 1939 they numbered more than 2,000, dis- 
tributed in 31 counties, in spite of a certain amount of poaching 
and occasional accidents and disease. The sex ratio is found 
to be about four or five does to one buck. As might be ex- 
pected, the abundance of deer is roughly proportionate to the 
amount of forest cover. 

Estimates of the deer population of the States in which this 
race occurs were given by the U. S. Biological Survey in Janu- 
ary, 1939, as follows: New Jersey, 8,000; Indiana, 400; Ohio, 
2,000; Iowa, 450; Virginia, 15,000; Kentucky, about 700; 


Missouri, 700; West Virginia, 17,500; Tennessee, 1,700; 
North Carolina, 51,000; South Carolina, 23,000; Georgia, 
16,400; Alabama, 20,200; Arkansas, 8,400; and Mississippi, 
4,600. A comparison of these figures with the statements 
given above indicates that under recent legal protection the 
numbers have probably increased very greatly over what 
they were a few decades ago, so that a certain amount of hunt- 
ing will become warranted. 

At the present time the white-tailed deer is the largest and 
most valuable of our game animals in the eastern United States. 
As an object of interest to the increasing numbers of persons 
who delight in seeing wildlife, as an important item of food for 
those living in rural or wilderness areas, and as a chief object 
of pursuit by the many who enjoy hunting in the invigorating 
autumnal season, this deer may be reckoned as one of our 
most important species in the regions where it is found. The 
readiness with which it adapts itself to the proximity of man 
and the rapidity with which its ranks may build up if given 
proper protection should make its continuance in reasonable 
abundance a matter easily managed by any efficient game 



Odocoileus americanus borealis Miller, Bull. New York State Mus., vol. 8, p. 83, Nov. 

21, 1900 ("Bucksport, Hancock County, Maine"). 
FIGS.: Stone and Cram, 1902, pi. opposite p. 42; Barbour and Allen, 1922, pi. 4, fig. 2 


The northern Virginia deer is a slightly larger animal than 
the more southern race, weighing when full grown up to 250 
pounds or even more. In winter coat the pelage is slightly 
longer and thicker, and the antlers of the males tend to be 
wider, more spreading, and less bent in at the tips. 

The northern Virginia deer probably did not range farther 
north in early times than the coastal regions of Maine and 
adjacent parts of New Brunswick and Nova Scotia, avoiding 
the central parts of Maine, northern New Hampshire, and 
northern Vermont and southern Canada. The northern and 
central parts of New England and New Brunswick are regions 
of heavy and lasting snowfall, in which the movements of deer 
become much restricted in winter, while in times of crust forma- 


tion, wolves would probably get most of the deer in such 
regions. A hundred years ago, says Dr. R. M. Anderson 
(1939a), "deer were said to be very seldom seen north of the 
Ottawa River." Within the past 50 or 75 years, however, 
considerable changes have taken place. Wolves are gone 
south of the St. Lawrence River, and even to the northward 
are much reduced in Quebec, so that they hardly form a 
deterrent to the spread of deer. Furthermore, with the in- 
tensive lumbering industry in New Brunswick, northern 
Maine, Michigan, and Wisconsin, areas of primeval spruce 
and white-pine forest have been cleared, with a consequent 
springing up of a great sprout growth of deciduous trees, which 
afforded cover and abundant food. These two factors have 
probably contributed in great part to the spread northward of 
this deer over Nova Scotia, New Brunswick, northern Maine, 
and southern Quebec, until "in 1930 a deer was taken in 
Abitibi River district within one hundred miles of James Bay" 
(R. M. Anderson, 1939a). Deer have been introduced and are 
thriving on Anticosti Island. They are common in northern 
Michigan, northern Wisconsin, northern Minnesota and the 
adjacent parts of Canada as in Quetico Park (Cahn, 1937), 
where, though some are killed by wolves during winter, "such 
depredations are not at all serious." 

In his map of the distribution of eastern races of the Vir- 
ginia deer, Cory (1912, p. 66) shows as blank areas from which 
the deer is "now practically extinct," western New York, 
northwestern Pennsylvania, practically all of Ohio, Indiana, 
and Illinois, the southern third of Michigan and Wisconsin, 
all of Iowa, the southern third of Minnesota, and a part of 
northern Missouri. Over much of this area it has in recent 
years come back. According to Dr. M. W. Lyon, Jr. (1936), 
deer at the present time are extinct in Indiana, where in pioneer 
days they were abundant. "The last stand of the deer in the 
state was in the northwest in the Kankakee region and in 
Knox County," where the last wild deer were seen near Red 
Cloud in 1893. Probably the estimate of 400 deer in the 
State given by the Government census in 1939 requires con- 
firmation, but indicates a very recent return. Hollister, writing 
in 1908, says that the last wild deer were exterminated in 
southeastern Wisconsin nearly 60 years before. In Walworth 
County, where they were abundant a century ago, they rapidly 


decreased after 1842, and the last one was seen near Delavan 
in 1852. "Probably the most southern limit of their range in 
Wisconsin at the present time is Sauk County" (Cory, 1912). 
Although formerly abundant in Illinois, deer seem to have 
been greatly reduced by the last decade or so of the last century, 
and by 1900 were regarded as practically exterminated within 
the State, although even as late as 1910 Cory had reliable 
reports of a very few in Alexander County in the southern 
portion of the State. Possibly this remnant has increased in 
the succeeding quarter century enough to justify the figure 
given in the 1939 Federal estimate of 250 animals. 

The way in which this deer will "come back" if depleted 
numbers are allowed to breed up for a period of years is well 
illustrated by the experience of the three southern States of 
New England. Before the settlement of the country deer were 
a staple source of food for the Indians. From deer skins they 
prepared clothing, and from the straight metapodial bones 
they skilfully cut out portions for use as implements. The 
first settlers also depended in part on these animals for sus- 
tenance and buckskin, and there was a large trade in the hides. 
As early as 1646 Rhode Island had a closed season on deer 
from May 1 till November 1. In 1698 Connecticut ordered 
that deer should not be killed between January 15 and July 15, 
and this period was soon after extended to include the re- 
mainder of these two months. Massachusetts soon followed 
suit, and deer reeves were appointed to enforce the law. For 
nearly a century the numbers continued without notable de- 
pletion, but by the close of the eighteenth century deer had 
become scarce or nearly exterminated over much of southern 
New England, until in 1842 Linsley, in his account of the mam- 
mals of Connecticut, makes mention only of one killed the 
previous year in Waterbury. In 1869 J. A. Allen wrote that 
in Massachusetts they were gone except for a few in Plymouth, 
Barnstable, and Berkshire Counties, where they were "strin- 
gently protected by law. " Much of southern New Hampshire 
and parts of Massachusetts were in the middle of the nineteenth 
century cleared and used as grazing country, supplying the 
nearby markets until the opening of the more extensive areas 
of the Middle West, after which these regions gradually re- 
verted to woodlands. Toward the end of the century the 
remnant of deer that had hung on in the Berkshires of the 


western part of Massachusetts and in the Plymouth region of 
the eastern section gradually built up a small population, 
which overflowed into the surrounding regions and, helped by 
a few introductions, began at length to attract notice. Under 
protective laws the numbers have increased so greatly that by 
1910 Massachusetts declared an open season of six days late 
in November. Since then there has been an annual open sea- 
son with few exceptions, and of about the same length. The 
number killed each year has varied from as low as 832 to over 
2,000 (in 1924). There seems no reason to believe that with 
proper regulation the deer may not continue to be a game 
animal not only in Massachusetts but also in the other New 
England States where a somewhat similar increase also took 
place, in Connecticut, Rhode Island, and Vermont (in the 
last-named as a result in part of introduction). A similar wise 
administration has brought about a like result in Pennsylvania, 
where in early days it abounded, but by the end of the last 
century, Rhoads (1903) said, it had become "sparsely scattered 
or locally exterminated" though most often found in the 
Pocono and South Mountain regions. At the present time, 
the estimate published (in 1939) by the U. S. Biological Survey, 
places the white-tailed deei population of Pennsylvania at 
793,000 ! The success of these eastern States in the replenish- 
ment of their forest covers with deer, and the asset which this 
game may be to the commonwealths, should prove an example 
to other States, in which the deer have been greatly depleted, 
to encourage an attempt at restocking. 


Cariacus osceola Bangs, Proc. Biol. Soc. Washington, vol. 10, p. 26, Feb. 25, 1896 

(" Citronelle, Citrus County, Florida"). 
SYNONYM: Odocoileus virginianus mcilhennyi F. W. Miller, Journ. Mamm., vol. 9, p. 

57, Feb. 9, 1928 ("Near Avery Island, Louisiana"). 
FIGS.: Barbour and Allen, 1922, pi. 5, figs. 5-7 (antlered skulls). 

In the southern part of the Florida Peninsula, and extending 
up on the western side and along the Gulf coast to Louisiana, 
the representative of the Virginia deer is a small animal, with 
notably smaller antlers than the typical race. Although the 
type locality is in the intergrading area of the two races, the 
name may for the present be retained. Of these small Florida 


deer, Cory (1896) states that he has killed full-grown bucks 
that did not weigh over 110 pounds, although they average 
larger. Apparently the recently described small deer from the 
coast marshes of Louisiana may be considered the same and 
is here included as a synonym. 

The exact status of the small Florida deer is not well ascer- 
tained. It is apparently common in southern Florida and was 
considered to be "of very general distribution" over much of 
the peninsula in 1898 by Bangs, although "in the more thickly 
settled and accessible parts of the State it has been much re- 
duced in numbers. " This reduction has of late been acceler- 
ated, because of the killing of deer on account of their being 
hosts to ticks that spread disease. On the Gulf coast they are 
said to be no longer found in western Florida, but in extreme 
southwestern Alabama what may possibly be this same race 
still exists. In this State, according to A. H. Howell (1921), 
deer are now practically exterminated except in a few areas in 
the northern part (presumably typical mrginianus) and in 
"the big wooded swamps of the lower Tensaw and Mobile 
Rivers " (presumably osceola) . Here a number are killed during 
the open season every fall. Ttey are hunted with dogs that 
drive the deer past the waiting hunter at some favorable 
"stand." "The deer take readily to the water and swim 
easily from one island to another in this great swamp; in this 
way they are able to keep ahead of the dogs, but are often 
shot while swimming a creek or river or when crossing an 
opening in the timber. Deer are still found in moderate num- 
bers in the sandhills and swamps of southern Baldwin County 
. . . Twenty years ago or more they were common in the 
sandhills and small swamps of Mobile County, but now ap- 
parently all have been exterminated from that region." 
Both these counties border the Gulf of Mexico in extreme 
southwestern Alabama. No data are at hand concerning the 
deer in coastal Mississippi, but it seems probable that the 
small deer with large toothrows of south-central Louisiana 
described as a race, mcilhennyi, are the same and are of local 
distribution in the coastal region. 

Probably some restriction on the killing of deer in these 
areas is desirable. 




Odocoileus virginianus clavium Barbour and Allen, Journ. Mammalogy, vol. 3, p. 73, 

May, 1922 ("Big Pine Key, Florida"). 
FIG.: Barbour and Allen, 1922, pi. 5, fig. 4 (an tiered skull). 

This is the smallest of the eastern races of this deer, charac- 
terized by its pale colors, very small antlers, and small teeth 
(upper cheek teeth 67 mm. in combined length). It is short- 
coated at all seasons and somewhat buffier in color than the 
mainland form. 

This small deer is confined to a few of the outer keys off the 
southeast coast of the Florida Peninsula, at the present time 
being found only on the chain of islands from Big Pine Key 
southwestward to Boca Chica, a small island 7 or 8 miles from 
Key West; on the latter there were deer in the early nineties. 
"Deer were killed on Stock Island, a small key adjoining Key 
West," about 1910, but none has since been known there. 
There were deer on Boca Chica until about the same time, 
when they disappeared and were not again noticed there until 
late in 1920, when two were seen. "Deer have always been 
found from time to time on Ramrod Key, all three of the Torch 
Keys, and probably Newfound Harbor. They swim readily 
from key to key and if hunted on the smaller islands they leave 
and go back to Big Pine Key," the largest of the "lower 
keys. " On this island and some of the others the dense thorny 
thickets of pricklypear and other growth offer them a safe 
refuge, and in 1922 they were said to be more plentiful than 
they had been a few years previously. 

On account of its restricted range and the assiduity with 
which it is hunted, even by parties coming over from Cuba, 
this small deer may be considered to maintain a rather pre- 
carious existence, but unless its stronghold of thorny growth 
is burned or destroyed in some other way it is likely to hold 
out in small numbers for a long time. 



Dorcelaphus texanus Mearns, Proc. Biol. Soc. Washington, vol. 12, p. 23, Jan. 27, 1898 
(Fort Clark, Kinney County, Texas). 


This is the race of the semiarid part of southern and middle 
Texas, but the exact limits of its range remain to be defined. 
It is said to be of small size, with relatively short legs, small 
ears, and with small and strongly incurved antlers. The molar 
and premolar teeth are relatively very large, the colors pale. 
Total length, 1,585 mm. Weight of bucks up to 100 pounds^ 
females 75 pounds (Mearns, 1907). 

The range of this race extends into Mexico from southern 
Texas at least as far as San Luis Potosi. Mearns (1907) 
wrote of it that in the early years of this century it was abun- 
dant in the bottomlands and low mountain ranges of southern 
Texas, notwithstanding the fact that the Seminoles kill num- 
bers of them every year. He had seen them in great bands in 
Devils River Valley. Bailey (1905) says that "on many of the 
large ranches between Corpus Christi and Brownsville, where 
the oak and mesquite thickets are interspersed with prairie and 
grassy openings, deer find ideal conditions, with abundant food 
and cover. The nature of the ground is such as to protect them 
from wolves and other natural enemies; but it is well suited to 
either hunting on horseback or still hunting, which, if freely 
allowed, would soon exterminate them. With protection, 
however, they increase rapidly, and in many places are abun- 
dant . . . On certain large ranches they are still numerous, 
while on others they have become extremely scarce and would 
be entirely exterminated but for the recruits from surrounding 
and better protected ranches ... In spite of the pro- 
tection of State laws and ranch owners there are still remote 
sections of rough, uncontrolled range where every year hunters 
kill wagonloads of deer for the market, or worse, kill the deer 
for the hides only, leaving the carcasses to rot. I was told 
that in the winter of 1901-2 hundreds of deer skins were 
brought out of the country west of Kerrville." Bailey at- 
tributes much of the present abundance of this deer to the en- 
lightened attitude of the ranch owners who give it reasonable 



Odocoelus [sic] virginianus louisianae G. M. Allen, Amer. Nat., vol. 35, p. 449, June, 

1901 (Mer Rouge, Morehouse Parish, Louisiana). 
FIG.: Allen, 1901, fig. on p. 453 (antlered skull). 


The status of this form is imperfectly known. It is a large, 
rather pale animal, not very different apparently from mr- 
ginianus. Its range is believed to be the northeastern part of 
Louisiana, northeastward into the adjacent region. No 
statistics of its numbers or notes on its status are at hand. 
Probably, however, the 2,000 deer credited to Louisiana by 
the 1939 Federal census of big game are largely of this race. 



Corvus (i. e. Cervus) macrourus Rafinesque, Amer. Monthly Mag., vol. 1, p. 436, 1817 

(Plains of Kansas River, eastern Kansas) . 
FIG.: Bailey, 1931, pi. 2, fig. D (antlered skull). 

This race of the mountains and plains of the western United 
States is described by Bailey as large for a white-tailed deer, 
color in winter pale gray, yellowish red in summer; tail long 
and bushy, ears small. He describes a doe from Texas as 
lacking the black tips on the ears, and in general brighter 
colored than the race texanus. It is larger and paler than 
typical virginianus. 

Originally well distributed in wooded and brush-grown 
areas in valley bottoms and along streams from the Dakotas 
west to the Rocky Mountains and south to eastern New 
Mexico, it is naturally restricted in these regions to the local 
habitats it favors, avoiding open country, but seeking the pro- 
tection of thickets along the streams. In North Dakota, 
Bailey (1926) writes that though much reduced in numbers 
locally, this deer, because of its secretive habits has hung on 
and is even in places common in spite of settlement. It may 
even be farmed with profit, like domestic stock, so readily 
does it respond to careful handling. Elsewhere its numbers 
vary. Thus in Kansas, where it was originally described, it is 
now extinct (Hibbard, 1933). In early days, wrote Gary 
(1911), it was found generally over the Plains region of Colo- 
rado, but it is at present uncommon in the State and largely 
restricted to the foothills and eastern slopes of the front 
ranges, where it occurs sparingly across the entire width of the 
State. In recent years deer are reported as locally common in 
the mountains of eastern New Mexico, although much hunted. 
They especially frequent the "willow thickets along stream bot- 


toms. Bailey (1931) writes that "this white-tailed deer will 
soon become exterminated from the open country in New 
Mexico, and the only possible hope of keeping it from entirely 
disappearing from the State will be to give it a permanently 
protected breeding ground where conditions are suitable for 
food and shelter. Within its present range these conditions 
could be obtained in great perfection on the eastern slope of 
the Sacramento Mountains and along the southern and eastern 
slopes of the Sangre de Cristo Range. " 

In recent years, according to Seton, this deer has extended 
its range into parts of Utah. 



Odocoileus virginianus ochrourus Bailey, Proc. Biol. Soc. Washington, vol. 45, p. 43, 

Apr. 2, 1932 ("Coolin, south end of Priest Lake, Idaho"). 
FIGS.: Bailey, V., 1918, pi. 7, fig. 2 (photograph of doe), fig. 3 (head). 

This race is found from Idaho and Montana to the eastern 
slopes of the Cascade Mountains, Oregon, and a short distance 
into northern Nevada and the extreme northeastern corner of 
California, and extends northward into the southeastern corner 
of British Columbia (Cowan). 

Equaling the Plains white-tailed deer in size, it differs in 
being darker with less black and more ochraceous on the upper 
side of the tail. In winter coat it is dark buffy gray above, 
becoming bright ochraceous on top of tail, on the legs, and on 
the edges of the belly. Forehead and top of head dark brown, 
brisket dusky ; sides of nose and eye-ring light gray. In summer 
the upper parts are bright tawny or light bay, legs but little 
paler, not yellowish as in the race macrourus. Antlers large 
and heavy when well developed, and frequently having the 
brow tine and the proximal long tine slightly forked. 

Concerning this race Bailey (1936) writes: "There is very 
little on record of the habits of this deer in Oregon, except 
that they are found mainly in thickets and willow bottoms 
along the streams and valleys. In recent years they are some- 
times found back in the hills, crowded back probably by 
settlements. Like most of their group they are secretive and 
would rather hide than run. In the more extensive thickets 


and forests of Idaho and Montana they are still abundant in 
favorable locations but over much of their ranges they are 
doomed to be crowded out by settlements, or killed by preda- 
tory animals as they gather on winter ranges and are easily 
pulled down by coyotes and dogs in the deep snows." Bailey 
(1918) has given a further account of this deer in Glacier 
National Park (under 0. v. macrourus) where it is abundant 
and protected. In extreme northeastern California, it was 
evidently at one time common in Modoc County but apparently 
now is no longer so. Grinnell (1933) gives as "a late definite 
record" one taken in extreme eastern Lassen County in 
January, 1922. On the eastern slopes of the Cascade Moun- 
tains in Oregon and Washington this race seems still to be 

Estimates of white-tailed deer populations for the States in 
which this race occurs are given by the U. S. Biological Survey 
for 1939 as follows: Idaho, 10,000; Montana, 24,000; Oregon, 
not given; Washington, 7,500. 

By Executive Order in 1939 a Federal refuge for this deer 
and other forms of wildlife was established in Stevens and 
Pend Oreille Counties, Washington. It will contain eventu- 
ally about 65,000 acres and is well adapted to the management 
of these deer, since they spend the spring and summer in the 
mountains and the autumn and winter in the lowlands of the 
area. The need for such a refuge has for some time been urged 
because the deer have become locally much limited through 
the operations of farming and logging. 



Cervus leucurus Douglas, Zool. Journ., vol. 4, p. 330, 1829 (Lower Columbia River, 
Oregon; description based on examples from Falls of the Willamette and mouth 
of the Columbia). 

FIGS.: Bailey, V., 1936, pi. 23, fig. A (antlered skull); Scheffer, 1940, p. 275 (photo- 
graphs of a doe). 

This is the only race of white-tailed deer to reach the Pacific 
coast of the United States. 

Small in size, standing about 3 feet 5 inches at the shoulder, 
and characterized by its small delicate antlers, this is likewise 
a darker race than ochrourus, its nearest neighbor, an appear- 


ance due to the narrowness of the pale subterminal rings of 
the hairs. There is a poorly defined median dark dorsal line 
on neck and shoulders ; upper side of tail light ochraceous-buff 
at base, more dusky toward tip; brow patch, with prominent 
suffusion of chestnut; white of under parts not reaching the 
axillae. Summer coat redder. 

Formerly this deer was found "at least in the Cowlitz and 
lower Columbia River Valleys of western Washington, possibly 
also eastward for some distance along the Columbia, there 
being one report from Pasco" but at present is much restricted 
and has even been supposed to be extinct in that State (Taylor 
and Shaw, 1929). South of the Columbia River it formerly 
ranged along the coast west of the Cascades nearly to the Cali- 
fornia line. It was typically an animal of the river bottoms. 
In 1826 Douglas reported this as the commonest deer in the 
districts adjoining the Columbia River and in the fertile 
prairies of the Cowlitz and Willamette Rivers. He found it 
also near the head of tidewater on the Umpqua River. There 
are no positive records of its occurrence farther south in the 
Coast Range valleys but northward these deer extend to the 
Puget Sound country of Washington. In 1872-75 they were 
still of regular occurrence in the Willamette Valley, though 
less common than 20 years before, but after that time they 
rapidly disappeared. According to the report of a reliable 
hunter in that valley, the last whitetail he knew of was killed 
about 1898 near Sweet Home, Linn County (Bailey, 1936). 
These deer were said to be common in the foothills about 
Beaverton, Washington County, Oreg., from 1860 to 1875. 

Until very recently the only remaining deer of this race were 
supposedly to be found in Douglas County, Oregon, where, 
according to Vernon Bailey (1936), Stanley G. Jewett in 1915 
reported a few still inhabiting the Long Tom Swamp west of 
Eugene and a few on the oak-covered hills along the Umpqua 
River, some miles northeast of Roseburg. In this latter region, 
according to Scheffer (1940), the "herd" is now believed to 
number 200 to 300 animals, upon which the State Game 
Commission enforces a strict closed season. Curiously, how- 
ever, Scheffer reports that this deer is still to be found in some 
numbers on both sides of the mouth of the Columbia River, 
a fact that had previously quite escaped notice. According to 
his detailed account (Scheffer, 1940), the present range of these 


deer includes chiefly low brushy islands and tidelands, below 
the 100-foot contour; above this the hills rise sharply from the 
river valley and the whitetails give place to the black-tailed 
deer. In the State of Washington the area inhabited by the 
former includes chiefly Skamokawa, Elokomin, and Cathlamet 
Islands, Puget Island, Tenasillahe Island, and Price Island, a 
total area of over 12,000 acres, with an estimated whitetail- 
deer population of between 400 and 500. On the Oregon side 
of the river's mouth the area inhabited is slightly greater 
(nearly 14,800 acres), but the deer population is less than half 
that in Washington, since the bulk of it is confined to the less 
cleared Webb and Westland districts. "As far as the main- 
land is concerned, the four corners of the range are fixed by 
steep bluffs that rise from the water's edge on both the Oregon 
and Washington shores." With practically no competition 
from blacktail deer, and with nearly complete freedom from 
large predatory mammals, the danger to this group seems small. 
Nevertheless in some of the islands as on Puget Island, the 
deer "have increased to the point where their depredations 
were distinctly annoying to the farmers." A few are annually 
killed in spite of legal protection, but with the gradual clearing 
and settlement of their restricted range, this remnant, Scheffer 
believes, will be in danger of extermination. "The obvious 
solution," he writes, "is to move one or more breeding stocks 
of the deer to nonagricultural areas," if suitable ones can be 
found, or to make a sanctuary of some tideland area already 
occupied by them and as yet only slightly devoted to farming, 
as on Skamokawa Island. This deer is well known to the 
farmers and fishermen of this region as "tideland deer" or 
"cottontail deer." 

ODOCOILEUS COTJESI (Coues and Yarrow) 

Cariacus virginianus var. couesi Coues and Yarrow, Rept. Geogr. and Geol. Explor. 

and Surv. west of 100th Meridian (Wheeler), vol. 5, p. 72, 1875 (Camp Crittenden, 

Pima County, Arizona). 
FIG.: Nelson, 1916, p. 458, lower fig. (colored). 

Though possibly to be considered a race of the Virginia deer, 
Bailey (1931) believes the Sonora deer are best regarded as a 
distinct though closely allied species. About half the size 
and weight of a Virginia deer, they are extremely pale in color, 


a light gray, somewhat more rusty in summer, but in pattern 
much the same as their larger relative. Old bucks reach a 
maximum weight of about 100 pounds, does 75 pounds. Total 
length of a large buck, 1,530 mm.; tail, 270; ear, 203. 

This small deer is still common on many of the wooded 
mountains "of middle and southern Arizona, southern New 
Mexico, western Texas, and in the Sierra Madre of Chihuahua 
and Sonora, Mexico," which constitute its range, but "with 
the growing occupation of their territory by cattle and sheep 
and the increase in the number of hunters, these once abundant 
deer are rapidly diminishing" (Nelson, 1916). In winter, if 
snows became deep on the upper levels of their mountain 
haunts, these little deer would move down into the valleys, 
and it was not uncommon to see bands of 20 to 100 in the 
White Mountains of Arizona in the nineties of the last century 
(Nelson). Concerning their status in New Mexico, Bailey 
(1931) writes that they occupy the mountain ranges "west of 
the Rio Grande as far north as the Datils and possibly to the 
Zuni Mountains ... In 1908 Goldman reported them as 
formerly abundant in the Burro Mountains, but at that time 
apparently all gone. In 1909 he 5 was told by residents that there 
were a very few of them in the Zuni Mountains, but in previous 
years" others had failed to find them. In the Mimbres Range, 
Bailey reports that Goldman in 1909 found them in limited 
numbers and that same year the Forest Service reported them 
in the Datil, Gallinas, Magdalena, and San Mateo Mountains. 
In the Animas Mountains they were found common by Bailey 
in 1906, but he believes that their numbers are a result of the 
protective natural features of their haunts, and that "a few 
persistent hunters could easily exterminate them." This deer 
probably reaches its eastern limit in the Chisos Mountains of 
western Texas, where according to Bailey (1905) it ranges from 
5,000 to 9,000 feet in the oak, juniper, and nut-pine cover. 
Where undisturbed they may often be seen feeding by day. 

As to the future outlook for this deer, Bailey writes of it in 
New Mexico: "In September, 1915, J. S. Ligon reported Sonora 
deer still common in parts of the Mogollon Mountains, but 
much less so than formerly, owing to unrestricted hunting in 
season and out. On December 31, he wrote: 'The number of 
deer killed in New Mexico during the season just closed far 
outnumbers the increase for the year, I am quite sure.' These 


mountains now form the principal range of the species in New 
Mexico, but as the country fills up with settlers the deer will 
entirely disappear unless given better protection than they 
have received in the past . . . The many game refuges 
in these mountains will now doubtless insure the perpetuity of 
this interesting little deer." In the Chiricahua Mountains of 
southeastern Arizona, Cahalane (1939a) has lately reported 
them as numerous "from the tops of the highest peaks to the 
lower limit of the upper Sonoran Zone." Owing to their 
aversion to leaving the shelter of forest cover, they are prac- 
tically "marooned" on these isolated ranges and in 1933 were 
found to be increasing so rapidly that already they "were too 
abundant for the carrying capacity of the range . . . The 
depletion of food was becoming a serious problem at this 
time" and reduction of their numbers by hunting was insuffi- 
cient on account of the rugged nature of the country they 
inhabit. "A check on this undesirable increase would be 
effected by dropping control of mountain lions," the reduction 
of which during the previous decade by the "predatory -animal 
control" "has been followed by a too rapid growth of the deer 
population. " 


Two types of caribou are found in the northern parts of the 
New World, the barren-ground and the woodland. Both are 
divided into several geographic races, not very sharply defined. 
The barren-ground caribou, as its name implies, inhabits the 
Arctic tundra. It is shorter of limb but often longer in its 
antlers than the woodland caribou, which is found slightly to 
the southward of the barrens, in regions where open bogs 
alternate with evergreen forest. The term "caribou," almost 
universally applied to these animals in America, is from a 
Micmac word meaning "a shoveler," in allusion to the way in 
which these animals paw away the snow in winter to get at the 
reindeer moss on which they feed. In contrast to other mem- 
bers of the deer family, the females as well as the males may 
have antlers, although less often in the woodland than in the 
barren-ground species ; when present, however, they are smaller 
in the females and in the woodland species are generally spike- 
like. The hoofs are broad and spreading as an adaptation to 
living on boggy or snow-covered ground, and the body is stocky 


for a deer. Because of their gregarious habits, their manner of 
living in more or less open country, and a sort of stupid curi- 
osity they may often be closely approached or attracted and 
so in former days were easily brought within gunshot and 
several killed before the herd could escape. In many parts of 
their range they have become greatly reduced in numbers and 
merit protection wherever found. As a source of food and 
clothing they are a mainstay of Eskimo and Indians in the 
parts of their range where these primitive people live. 


Cervus tarandus var. arctica Richardson, Fauna Boreali-Americana, vol. 1, p. 241, 1829 

(Fort Enterprise, Mackenzie, Canada). 
FIGS.: Grant, 1902, pis. [18, 19] (antlers and skull). 

As Murie (1935) points out, the characters distinguishing 
the barren-ground type from the woodland type of caribou 
are sometimes "confusing." In general, however, the former 
is shorter of limb and lighter in color, the antlers tend to have 
the main beam more nearly cylindrical rather than flattened, 
and the antlers themselves are less compact, with long, backward 
then forward curved beam, branching dichotomously at the 
summit. The two brow tines tend to show a wide triangular 
expansion or palm. So great is the variation in these structures, 
however, that scarcely any two are alike, and the opposite 
antlers of the same individual may be considerably different. 
Hollister (1912a) states that in the barren-ground type the 
lower incisors (which in both are small and weak) decrease in 
size from the middle to the outer pair by conspicuous steps, 
with the outer pairs very small, whereas in the woodland type 
the gradation from the middle to the outer pair is more uniform. 

General color in summer "clove-brown, mingled with reddish 
and yellowish brown, under-parts white; in winter entire coat 
dirty white " (Lydekker, 1915) . In size this is one of the smaller 
American races, standing about 46 inches in height at the 
shoulder. The basal length of skull averages about 365 mm., 
the maxillary tooth row about 89, but there is a slight over- 
lapping with the larger Alaskan race (Murie, 1935). The 
antlers of the male are "very long, slender, and rounded, with 
few points on the expanded portion of the beam, which is 


separated by a long interval from the third tine, . . . back 
tine usually, if not always, wanting; female antlers much 
smaller, simpler, and scarcely curved at all" (Lydekker). In 
the few specimens available the antlers seem to be less spreading 
than in other races, but, as Murie has shown, there is great 
individual variation in their form. 

The range of this subspecies is believed to be continental 
North America from the west side of Hudson Bay to the Mac- 
kenzie River and northeastward to include Boothia, South- 
ampton Island, and Baffin Island. The migratory habit so 
characteristic of caribou leads to more or less irregular wander- 
ing, apparently in search of better feeding areas, and the ani- 
mals often congregate and pass through certain sections in 
great numbers, while in other seasons they may hardly be 
found at all. This uncertainty leads at times to much hardship 
on the part of Indians and Eskimos who depend upon them as 
a source of food. Preble (1902) states that the southern limit 
of the barren-ground caribou on the west coast of Hudson Bay 
is Churchill River. "Even in former years these caribou were 
seldom known to cross that river, and they are still killed within 
a few miles of Fort Churchill. Farther inland they reach the 
south end of Reindeer Lake." In his later paper on the 
Athabaska-Mackenzie region, Preble (1908) has given a de- 
tailed account of the presence of this animal as recorded by 
various travelers along the western part of the barren grounds, 
the shores of the Arctic Ocean, and the outlying large islands 
as Banks Land and Victoria Land. "As winter approaches, 
the caribou which have summered on the Barren Grounds 
move southward in herds, many of which enter the wooded 
country. Their movements are more or less irregular. " Few 
persons traverse the inland barrens so that this race probably 
remains, as Dr. R. M. Anderson says (1939b), "the most 
numerous of all the caribou, although now very rare or missing 
from much of its fairly recent former range along the Arctic 
coast." Nevertheless, he adds, it "has perhaps not been 
greatly reduced in total numbers, as where it has disappeared 
many of the natives have not followed the caribou. The 
caribou have also retreated farther from the shores of Hudson 
Bay, and there have been many reports of large concentrations 
of caribou in the interior east of Great Slave Lake and south of 
Bathurst Inlet. In some cases shifting of range is due to human 


crowding and in others the destruction of winter forage of 
lichens by fire may have been the prime cause of extended 
movement . . . Within the past few years more caribou 
than usual have come into parts of northern Manitoba and 
Saskatchewan, in winter, and quite recently barren ground 
caribou have crossed the Slave River into the Wood Buffalo 
Park and still farther south of the Park in northeastern Alberta 
where the species was never known before. This southward 
trend of the barren ground caribou of course brings them out 
of the comparative protection of the large native hunting 
reserves and restrictive regulation of white trappers in the 
Northwest Territories into the northern parts of the Prairie 
Provinces where white trappers abound. Reduced killing of 
caribou in parts of the north has perhaps been counterbalanced 
by increased slaughter in parts of the winter range. Recent 
expert investigators inform the writer that there has not been 
much new evidence to change his estimate [of 1930] of about 
three million caribou in Canada." 

On Southampton Island in northern Hudson Bay there is a 
local herd of which Sutton and Hamilton, in 1932, saw a few 
animals. They write that this caribou was "once a very abun- 
dant animal all over Southampton Island. At the present time 
it appears to have disappeared almost altogether from the 
southern part and to be restricted principally to the region of 
the high country between East Bay and Duke of York Bay, 
and to the more or less unknown country inland from the 
coast and north of Cape Kendall." The Eskimos evidently 
hunt them only occasionally, and there is no evidence that they 
are greatly depleted. In Baffin Land both Hantzsch and Soper 
report the barren-ground caribou as local in occurrence and 
irregular in its appearance about the misson settlements of 
Cumberland Sound. Almost nothing is known of their num- 
bers or their migrations, but it seems likely that they keep 
mainly to their archipelago, which they must have originally 
reached by way of Boothia. 



Rangifer arcticus caboti G. M. Allen, Proc. New England Zool. Club, vol. 4, p. 104, 
Mar. 24, 1914 (30 miles north of Nachvak, eastern Labrador, Canada). 

SYNONYM: Tarandus rangifer labradorensis Millais, The Gun at Home and Abroad, 
vol. 4, p. 259, 1915 ("Nain, Davis Inlet, and Fort Chimo"). 

FIGS.: Grant, 1902, pis. [8, 9] (skull and antlers). 

This is the race of the Labrador Peninsula north of its 
forested base. It is believed to be characterized by the wide 
antlers, with very sweeping backward and forward curve, and 
the great palmation of the brow and bez tines. If other dif- 
ferences occur, as in color, these have not been made out. 

Dr. R. M. Anderson (1934b) writes that this caribou "is 
found in more or less scattered bands over the treeless Arctic 
Zone area and through parts of the scantily timbered Hud- 
sonian Zone of the peninsula. In most of the districts near the 
coast they have been greatly reduced in numbers, but con- 
siderable numbers are still found in the more isolated inland 
districts." These are regions seldom penetrated by white 
hunters, and accurate information as to the status of this 
animal is difficult to obtain. Low, at the beginning of this 
century, was told by the Nascopie Indians that there were 
three main herds or stocks: One near the coast on the high- 
lands between Nachvak and Nain; one in the hinterland of 
Ungava Bay that crosses the lower Koksoak and passes the 
summer on the tundra along Ungava Bay and Hudson Strait; 
and one on the east coast of Hudson Bay, south to Clearwater 
Lake. Before 1886 the last herd was said to have fallen off 
greatly. Of the second, W. B. Cabot (1912) and others have 
witnessed something of the migrations and mention the hunting 
of the migrating animals by the Indians who rely partly on the 
caribou for sustenance. Prichard (1910) states that these 
animals often work out to the coast by November, and at 
that time a good many may be taken, but their appearance is 
not to be counted on, and they may come out at different 
points in different seasons. There is some evidence that ex- 
tensive fires have destroyed the available food in parts of their 
range, thus excluding them from return to such districts. 

There is every reason to believe that this caribou never 
crosses the rough water and ice of Hudson Strait to Baffin 
Land but is isolated on the Labrador Peninsula. To the south- 


ward its range is limited by the forest country of the base of 
the peninsula, where the woodland caribou is found. 


Rangifer stonei J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 14, p. 143, May 28, 1901 
("Kenai Peninsula, Alaska"). 

SYNONYMS: Rangifer excelsifrons Hollister, Smithsonian Misc. Coll., vol. 56, no. 35, p. 
5, Feb. 7, 1912 ("Meade River, near Point Barrow, Alaska"); Tarandus rangifer 
ogilvyensis Millais, The Gun at Home and Abroad, vol. 4, p. 263, 1915 (Ogilvie 
Mountains, north of Dawson, Yukon, Canada); Rangifer mcguirei Figgins, 
Proc. Colorado Mus. Nat. Hist., vol. 3, no. 1, p. 1, Dec. 28, 1919 (Kletson Creek, 
a tributary of White River, Yukon, Canada). 

FIGS.: Allen, J. A. 1901, figs. 1-4 (head, skull, antlers). 

In his recent review of the Alaska- Yukon barren-ground 
caribou, Murie (1935) has shown that only one race is recog- 
nizable over most of central and northern Alaska (exclusive 
of the Alaska Peninsula and Unimak Island), ranging to the 
western part of Yukon, Canada. This is recognizable by its 
large size, with an upper maxillary tooth row averaging 94 mm. 
The large cheek teeth, large size, maximum for the arcticus 
type, dark color, with well-developed white fringe on the 
throat, and large, heavy, and "rangy" antlers are given as 
diagnostic characters. 

Murie states that this caribou has now disappeared from the 
Kenai Peninsula, whence it was first described, but the form 
of the interior of Alaska is the same, and it is probable that the 
Kenai animals simply represented those remaining from a 
periodic overflow of the herds of the interior. 

Stone's caribou, named in honor of Andrew J. Stone, who 
collected the type, is at present represented by four herds, 
which Murie (1935) briefly characterizes as follows: 

(1) The Bering seacoast group, consisting of scattered bands 
distributed over a wide area of lowland bordering Bering Sea 
from Bristol Bay to Bering Strait, but which even by the 
early eighties had begun to decline. Residents agree that at 
the beginning of this century caribou were much commoner 
in the country north of the lower Yukon and inland from 
Norton Sound and may still be found in the hills of that region 
particularly at the head of Unalakleet, but at the present time 
are "entirely absent or occur only as stragglers on the Bering 


Sea coast, in most of the Kuskokwim region, along the lower 
Yukon, and in the vicinity of Norton Sound, and much of this 
area is now occupied by domestic reindeer" (Murie, 1935). 

(2) The Alaska Range herds, consisting of scattered bands 
along the entire length of the Alaska Range. Murie believes 
that an approximate estimate of this group would be between 
25,000 and 30,000 animals, of which the principal aggregation 
is included in Mount McKinley National Park. 

(3) Northern herds, including the scattered remnants on 
the Arctic slope, comprise the animals "ranging along the 
Endicott Mountains, through the upper Koyukuk and Chanda- 
lar watershed, across the Porcupine into Yukon Territory." 
An estimate of 60,000 for the animals of this area is "probably 
conservative." In early times caribou were plentiful along 
the Arctic coast, but "in later years they largely disappeared 
in this area, owing no doubt partly to the activities of whalers, 
fur traders, and natives." 

(4) The Yukon-Tanana herds, which comprise the largest 
numbers of all, occupy the country west of the Mackenzie 
River, on the uplands between the Yukon and Tanana Rivers. 
Migrations of these herds, often numbering many thousands 
of animals, are described by Murie from accounts gathered. 
Except locally, their numbers show no depletion of importance. 


Rangifer granti J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 16, p. 122, Mar. 31, 

1902 ("Western end of Alaska peninsula, opposite Popoff Island"). 
FIGS.: Allen, J. A., 1902a, figs. 1-6 (skulls and antlers). 

The caribou of the Alaska Peninsula and Unimak Island is 
distinguished by "averaging somewhat smaller and paler than 
R. a. stonei, many antlers diverging widely and with sharply 
recurving beams, although these antler characters may not 
hold uniformly in a large series." The average length of the 
maxillary tooth row is 94.5 mm., or about as in stonei. 

"In their present depleted numbers" the caribou of the 
peninsula are "islolated from the interior herds" but appear 
to have been "sufficiently isolated to form a local race" 
(Murie, 1935). The antlers show a "tendency toward wide 
divergence of the beams," which "average lighter in weight 


than those of stonei. " At one time these caribou were abun- 
dant throughout the length of the Alaska Peninsula from a 
point nearly opposite Kodiak Island west to the island of 
Unimak and thence all around the Alaskan shore of Bering 
Sea (Nelson, 1887), as well as on Nunivak Island. At present, 
writes Murie (1935), "the numbers on the peninsula have 
dwindled greatly, but a few herds are still holding their own. 
One herd, numbering possibly 2,500, ranges from Morzhovoi 
Bay to Herendeen Bay, with the center of abundance in the 
vicinity of Pavlof Valley. The other herd, apparently a little 
larger, ranges from Moller Bay to the vicinity of Black Lake. 
The latter herd appears to be less molested by hunters than the 
others, but as a whole the caribou of the peninsula have been 
and are rapidly dwindling . . . Caribou are scarce to the 
eastward of Port Heiden, but a few have been recorded as far 
east as Becharof Lake in comparatively recent years. Several 
islands adjacent to the peninsula were inhabited at one time. 
Caribou occurred in considerable numbers on Unga Island, 
according to A. J. Stone, and Deer Island is also said to have 
had caribou, but today none are found on either. In July 1925 
a caribou skeleton was found on Amak Island, 12 or 14 miles 
off the Bering Sea shore . . . This island cannot be much 
over 2 miles square. Unimak Island, roughly 30 by 75 miles 
in extent, represents the westernmost point of distribution for 
caribou and harbors a group that has been estimated at from 
7,000 to 10,000 animals; a visit to the island in 1925 inclines 
the writer to favor the lower figure. " According to reports of 
Donald H. Stevenson, quoted by Murie, the caribou here were 
decreasing rapidly in the eighties and early nineties, when it 
was thought many caribou were killed for food by men hunting 
sea otter. With the cessation of these activities the caribou 
commenced to increase until "by 1905 the island held all that 
the range could carry. About 1908 they again began to decline 
in numbers, but soon were able to hold their own and later 
increased once more. " At intervals these island animals have 
been known to cross to the mainland of the peninsula, and 
there is some evidence of migratory movements lengthwise up 
and down the country. 

From the evidence brought together by Murie it seems that 
in spite of a depletion that has restricted these caribou to 
parts of the peninsula and islands, and cut them off from the 


herds of interior Alaska, they are at present in no danger of 
extinction and with proper regulation of hunting should hold 
their numbers for a long period. 


Rangifer osbarni J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 16, p. 149, Apr. 16, 
1902 ("Cassiar Mountains (60 miles southeast of Dease Lake), British 

FIGS.: Grant, 1902, pis. 18-21 (unnumbered) (head with antlers). 

This race of the barren-ground caribou is darker than R. a. 
stonei with the dusky tipping of the white throat fringe more 
pronounced and the maxillary tooth row longer, averaging 
about 100.5 mm., according to Murie. The differences are, 
however, actually slight, and it is hard to find any very tangible 
distinctions between the two, beyond these average ones of 
size and color. The antlers sometimes show the flattened form 
of beam characteristic of the woodland caribou. 

The geographic range of R. a. osborni includes southern 
Yukon and northern British Columbia, where according to 
late information it is still -"reasonably common" (R. M. 
Anderson, 1939b). This race is believed to be much less mi- 
gratory than the others. 


Rangifer pearyi J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 16, p. 409, Oct. 31, 1902 

("Ellesmere Land, N. Lat. 79"). 
FIGS.: Allen, J. A., 1908, figs. 1-6 (antlered skulls); Nelson, 1916, p. 421 (col. fig.). 

The caribou of Ellesmere Land, Grinnell Land, and Grant 
Land is smaller than the Greenland caribou or the typical 
R. a. arcticus, its nearest neighbors, and is nearly pure white 
in color, with, however, a slight admixture of gray to gray- 
brown in the middle area of the back. The maxillary tooth 
row ranges from 83 to 95 mm. The antlers "have a much 
greater upward curvature than in arcticus, in proportion to 
their length," and the main beam has an average length of 
about 1,019 mm. 

From Peary's notes on these caribou, which range the Arctic 
islands mentioned, it appears that they live in small scattered 


groups of up to a dozen individuals of both sexes, and penetrate 
as far as latitude 83 N. on the northern coasts of these areas. 
They are not known to show any distinct mass migrations but 
remain the year round thriving on the lichen, moss, and other 
dwarf and scanty Arctic vegetation. Their chief enemies are 
the Arctic wolf and, of course, man, particularly the native 
Eskimo, who hunts them for food and clothing. "In these 
northern wilds, amid the most intense cold, the caribou passes 
from three to five months of continuous night, its wanderings 
lighted only by the moon, stars, and the marvelous displays of 
waving northern lights" (Nelson, 1916). According to Ander- 
son these caribou are not very numerous, although Sverdrup 
found them in abundance on the west coast of Ellesmere 
Land. Since there are rather few human inhabitants it is 
likely that the caribou will persist for a long time to come. 


[Cervus tarandus] groenlandicus Gmelin, Linnaeus's Systema Naturae, ed. 13, vol. 1, 

pt. 1, p. 177, 1788 (Greenland). 
FIGS.: Grant, 1902, two unnumbered plates (4, 5) (skull with antlers). 

Lydekker (1915) describes the Greenland caribou as "closely 
allied to R. t. arcticus, with a broad sharply defined white ring 
round each eye, and distinct broad white bands above the 
hoofs; skull with an elevated frontal region." Whether or 
not these color characters are really distinctive, J. A. Allen 
(1908) states that it is darker in color than typical arcticus 
and much darker than pearyi, resembling greatly the coloration 
of the Woodland caribou in its dark-brown body, with neck and 
ventral area much lighter. Three male skulls average in 
condylobasal length, 368 mm., which is greater than in arcticus 
and considerably more than in the little pearyi. The upper 
maxillary tooth row is also greater, 81-99 mm. Antlers of 
adult males are large and spreading; the brow tines are not 
greatly palmate, the main beam is bent sharply forward at 
near a right angle at the most posterior point, at which there 
is a small but well-developed backward tine. Although the 
Greenland caribou has by some naturalists been associated with 
the Old World reindeer as a subspecies, and by others is re- 
garded as a derivative of the New World barren-ground cari- 


bou, it seems quite as likely that Jacobi (1931) is correct in 
calling the forms of both merely races of a single circumboreal 
species, Rangifer tarandus. 

The Greenland caribou, or reindeer, is believed to be derived 
from North America by way of the islands lying to the west- 
ward, and in that way to have reached the Greenland coast. 
Since, however, Peary's caribou now occupies the northern 
Arctic Archipelago, one must assume either that both have 
since differentiated from a common stock or that the Greenland 
animal arrived in some other way. 

Formerly the Greenland caribou was found over the whole 
outer land fringing the ice cap of this country, except, ap- 
parently, the northern part of the coast. During the last 
century and a half constant hunting on the part of both 
Eskimo and white population has greatly reduced their num- 
bers, so that at the present time they are found only locally in 
the areas most suitable to their needs on the southwest and 
west coasts but are gone from the eastern coast. Jensen 
(1928) has given some interesting data on the past and present 
distribution. These caribou were frequent on the southern- 
most part of the west coast even as late as the end of the 
eighteenth century, but they disappeared from this southern 
tip of the peninsula during the early part of the nineteenth 
century. Nevertheless, a good many still remain not far to 
the northward, near Narssalik (south of Frederikshaab) . 
The best hunting district is a little farther to the northwest, 
Sukkertoppen, at latitude 65 N., where "even now" about a 
thousand animals are killed annually. The Godthaab district, 
slightly to the south, is also a favored habitat, where Jensen 
says about 600 are taken yearly. The present distribution of 
these caribou extends still farther northwestward to the 
southern part of the Upernavik district, in about latitude 73, 
so that the range of the animal now is mainly limited to the 
stretch of coast covering about 800 miles from the southern 
Frederikshaab district to the Upernavik area. They no longer 
occur, however, on Disko Island about in the center of this 
stretch. Jensen states that in former times these animals 
extended their range still farther north to the Thule district, 
"at least as far north as Rensselaer Bay (lat. 78 40' N.)," 
where now they have "practically disappeared." Neverthe- 
less, a remnant still persists here and apparently forms the 


dependence of the Eskimo of the Etah and Smith Sound area. 
Macmillan (1918), in his account of experiences here, tells of 
the Eskimo setting forth on their annual hunt for the caribou, 
about 50 miles north of Etah, on September 10, 1914, and 
returning on the 23rd with "forty -two warm skins, invaluable 
for bed-robes, coats, and sleeping-bags for the extreme tem- 
peratures to come." At another time the Eskimos reported 
killing 19 in two weeks. In February, 1916, Macmillan 
noticed the wandering of the caribou herd southward to the 
vicinity of Etah, because of deep snows covering their feeding 
grounds between there and Humboldt Glacier, which forms a 
natural barrier to the northward extension of their range. Of 
six killed near Etah, the heaviest weighed but 120 pounds. 
It is believed that this northern herd is much preyed upon by 
wolves, for in 1916, Macmillan writes, the Eskimos returning 
late in October from the annual caribou hunt throughout the 
region extending from Etah to Humboldt Glacier, reported "no 
young caribou whatever and tracks of wolves everywhere." 
It was his belief that the wolves had crossed over Smith Sound 
on the ice from Ellesmere Land and had accounted for the 
young caribou. The Eskimo ^stated that they had seen a 
number of caribou sleeping on the ice in the center of lakes, 
probably for security against attacks by wolves. 

Returning to the southern tip of Greenland, Jensen notes 
that these caribou were in former times found at Angmags- 
salik but are exterminated. During recent decades they have 
also disappeared from the coast between Cape Dalton and 
Cape Brewster. In the northern part of East Greenland the 
Ryder expedition in 1891-92 "encountered reindeer with com- 
parative frequency in the region round Scoresby Sound; in 
1899 Nathorst only saw a few herds." The "Danmark Ex- 
pedition" in 1906-8 found "innumerable traces (cast-off 
antlers and excrements) of the former occurrence of reindeer 
in great numbers, as far north as Holm Land (lat. 80 20' N.), 
but now they have entirely disappeared"; this disappearance 
from the northeast coast is corroborated by later reports of 
explorers and hunters. Coincident with this disappearance 
may have been the abandonment of the region by the Eskimo, 
whose deserted camps have been investigated in the same 
region. Nathorst, who in 1899 saw only a few herds in the 
Scoresby Sound region, attributed this disappearance to wolves, 


which are said to have appeared in East Greenland "rather 
suddenly" at about the time of the disappearance of the cari- 
bou. This, however, may be only part of the reason. Quite 
possibly overhunting had much to do with it, or there may 
have been other contributory causes. 

From an economic standpoint, the Greenland caribou is an 
important asset to the Eskimo and white inhabitants of that 
country and one that should be exploited with care. For a 
great many years the export of caribou hides to Denmark has 
been going on, to an extent that was evidently unwarranted 
by the size of the herds. Jacobi (1931) quotes figures showing 
a decline in the numbers during part of last century. It is 
said that 37,000 caribou were killed in Greenland in 1839; 
from 1841 to 1850 an average of 13,900 hides were exported; 
from 1851 to 1860, an average of 5,667 yearly; from 1860 to 
62, the average yearly number was below 1,000; and since 
1862, below 100. In 1891, none was shipped. Nevertheless, 
there seems to be some evidence of an increase in numbers 
locally at the close of the last century. 

"Among land mammals the reindeer is the most important 
object of hunting to the Eskimos. The meat is eaten, and the 
fat is used, for instance, as cfeam for coffee, and the contents 
of the paunch are considered a special delicacy; the skins are 
used as underlayers on sleeping platforms and for sleeping 
bags and garments, the antlers for hunting implements, the 
sinews for thread" (Jensen, 1928). Apparently caribou hunt- 
ing is a summer occupation in South Greenland, when the 
Eskimo and their families seek the great plateaux in the neigh- 
borhood of the inland ice, and having selected their camp sites 
the men carry on the hunting till about the first of September, 
when they return to their dwellings on the outer coast. In the 
Etah region the hunting season is apparently in October. 
Since 1924 this pursuit has been somewhat regulated by the 
government in South Greenland. It has forbidden the hunting 
of caribou or molesting of the calves between May 20 and July 
20, and during the open season no more animals may be killed 
than can be utilized on the spot or taken care of for future use 
by the hunters. Obviously these animals are of great impor- 
tance in the economy of the Eskimo, to whose use it would 
seem they should mainly be relegated. Macmillan writes 
(1918, p. 74) that caribou meat "is lamentably lacking in 


strength and stamina-producing properties, " yet it is neverthe- 
less more palatable than walrus or polar bear and forms a 
welcome change from seal meat. 


Rangifer dawsoni Seton, Ottawa Nat., vol. 13, p. 260, Feb., 1900 (Graham Island, 

Queen Charlotte Islands, British Columbia). 
FIGS.: Seton, 1900, pis. 4, 5 (type specimen, antler and skull); Sheldon, 1912, pi. 

opposite p. 234. 

This is supposed to be a small island form, darker in color 
than the barren-ground caribou of the mainland. The single 
antler of the type is peculiar in that the brow tine does not 
project parallel to the forehead but turns upward and is well 
separated from it. The terminal forks of the main beam turn 
backward instead of extending forward. Length of type 
antler following outer curve to highest point, 28% inches. 

Very little is known of this caribou. Indeed, its very exist- 
ence in the Queen Charlotte Islands was doubted by Osgood 
(1901) and it was not until years after its description that 
satisfactory evidence of its presence on Graham Island, the 
most northerly island of the group, was obtained. Dr. G. M. 
Dawson, while making a geological survey of the islands in 
1878, was the first to learn of a caribou there, but from the 
meager description given him by the natives he at first was 
inclined to suppose the animals were wapiti. Nevertheless he 
noted the use of pieces of caribou antler in the implements of 
the native Haidas. After some effort the portion of cranium 
with an antler that afterward served as the type was secured 
through the Indians with the aid of a local trader, on Virago 
Sound. In 1904, 1905, and 1906, the Rev. Charles Harrison, 
a missionary stationed there, found evidence in the shape of 
tracks, dung, and hair. In 1906 the late Charles Sheldon 
undertook an intensive search for caribou in the country west 
and southwest of Virago Sound on Graham Island but found 
nothing more than old tracks and dung. In 1908 a bull and two 
cows were killed, and a calf was seen by two half-breeds near 
Virago Sound, and the specimens were acquired by the Pro- 
vincial Museum in Victoria, B. C. (Keen, 1909). In October, 
1910, Francis Kermode found two old tracks in the same dis- 
trict, and expressed the opinion that the form was nearly 


extinct. Wharton Huber informed Dr. Francis Harper that 
in 1930 two prospectors again found tracks on Graham Island 
but saw no animals. This was on grass-covered mountains 
about 2,000 feet high in the northwestern part of the island. 
I. M. Cowan further informed Dr. Harper that in November, 
1935, "Ed White reported fresh caribou tracks on Massett 
Island." It seems likely, then, that a few still exist in the 
interior mountainous parts of these islands. Since 1910 the 
British Columbia authorities have prohibited the killing of 
these animals, and because the local Haidas are more a fishing 
than a hunting race and do not care for caribou meat, it seems 
probable that the law will be observed. 

There has been much speculation as to the origin of this 
caribou group and how it should have survived in such seem- 
ingly small numbers with no known natural enemies. The 
nearest point of land is about 30 miles away to the coastal 
archipelago, and the nearest relatives of the caribou live 150 
miles distant at the present time. Dawson has suggested that 
the ancestral stock reached the archipelago during the Ice Age, 
when the islands were more easily accessible. If the race 
proves to be one of the barren-ground group, as seems likely, 
the fact will be all the more interesting. 


[Cervus tarandus] caribou Gmelin, Linnaeus's Systema Naturae, ed. 13, vol. 1, pt. 1, 

p. 177, 1788 (eastern Canada). 
FIGS.: Grant, 1902, 2 pis. opposite p. 18 (photographs); Nelson, 1916, p. 459 (colored 


The woodland caribou, as its name implies, is a more south- 
ern animal than the barren-ground species, inhabiting the 
boreal evergreen forests and their bogs quite across North 
America from Newfoundland to British Columbia. It is larger 
of body but with less elongate antlers than the latter type; the 
antlers usually tend to have a more flattened beam, and the 
lower incisor teeth are more uniformly graded in size from the 
middle to the outer ones. The females occasionally have small 
antlers, but these are ordinarily much less developed than in 
females of the barren-ground caribou. About five closely allied 
forms are at present recognized. 

The eastern woodland caribou is usually darker than the 


barren-ground species, with the head in front of the ears, the 
body from shoulders to tail, and the legs to just above the 
hoofs a dark brown; the lips and neck with its fringe of long 
hair below, and the legs just above the hoofs white or hoary. 
There is a narrow white area bordering the buttocks, but the 
upper surface of the short tail is dark. The antlers of the male 
are shorter than in the barren-ground type, less sweeping, and 
extend outward and forward, with a short tine at the bend, 
palmate brow tine on one or both sides and a varying number 
of tines on the terminal part of the main beam. Large antlers 
may measure in length of beam 42 inches on the outside curve. 
The range of this eastern race originally extended from Nova 
Scotia westward across New Brunswick, northern Maine, ex- 
treme northern New Hampshire, and Vermont to southern 
Ontario, and on the north side of the Gulf of St. Lawrence to 
the wooded parts of southern Labrador west to James Bay. 
Westward of the latter region it grades into the race sylvestris. 
Its southward limit was the north shore of Lake Superior. It 
apparently did at one time reach the Adirondack region of 
New York. Over this range the woodland caribou is now 
largely gone, chiefly as a resulfe of hunting and partly perhaps 
as a result of lack of reproduction on account of reduced living 
area and the harrying of civilization. The caribou south of 
the St. Lawrence have probably long been cut off from their 
western and northern neighbors since the settlement of southern 
Canada and may be thought of as a separate herd having a 
tendency to wander irregularly over the area. Thus during 
the nineteenth century at intervals of about 15 years, the 
records show a great influx of their numbers into northern 
Maine, so that they came down as far as tidewater in the 
eastern part of the State and to the large bogs near Bangor in 
the interior. For a few years there would be caribou in some 
numbers in the Maine woods, then they would drift away again 
eastward into New Brunswick and Nova Scotia and might be 
nearly if not quite absent from Maine for another period. 
New Brunswick with its greater extent of barrens and southern 
Quebec south of the St. Lawrence were probably more attrac- 
tive feeding grounds. Caribou are easily killed on account of 
their gregarious habits and a curious inquisitiveness that leads 
them to stop their flight and approach an unfamiliar object. 
In former times a hunter on being discovered might lie on his 


back and kick his legs in the air or wave a red cloth, when 
presently the whole herd would turn and approach to investi- 
gate. The hunter, waiting his chance, would leap up and get 
in several shots before the caribou could dash off. This south- 
ern herd must have become gradually reduced by hunting and 
the effects of settlement without much cognizance being taken 
of it. The last great infiltration in Maine was in the early 
nineties, when they were reported for a time more numerous 
than deer. In 1895 and again in 1896 the Bangor and Aroostook 
Railroad alone shipped out about 130 caribou each year that 
had been killed by visiting sportsmen, but after that the num- 
ber in the State began to decline, owing in part probably to the 
eastward movement of the animals once more and in part to 
summer and winter killing in both Maine and the adjacent 
parts of Canada. There were a few caribou in the bogs of 
northern New Hampshire until at least 1885. In 1899 a closed 
time on caribou in Maine for six years was established and on 
its expiration was renewed for another six years. Meanwhile, 
however, the animals had practically gone from the State. 
The last caribou in the Mount Katahdin region, formerly their 
favorite haunt, was seen in 1905 and a small herd remained on 
the northwestern border of the State near the St. Johns River 
until 1916, but since then there seem to be no certain records. 
The history of this herd is much the same in New Brunswick 
and Nova Scotia a gradual diminution and retreat. Dr. 
R. M. Anderson (1939b) writes that in Nova Scotia caribou 
were so numerous in the sixties that a certain English sportsman 
killed 120 in a single day in Lunenberg County. "The last 
caribou on the mainland of Nova Scotia was killed in Guys- 
borough County about 1912," and they were "almost certainly 
gone" from the once famous caribou grounds of Victoria and 
Inverness Counties in northern Cape Breton Island by 1925, 
although in only the previous year he found recent "traces" 
of them. "In New Brunswick a closed season was estab- 
lished for caribou in 1919," but it was too late to help them, 
for in a few years they seem to have altogether gone. "Fairly 
authenticated records in the upper Tobique River in 1924" 
are mentioned by Dr. Anderson, and the New Brunswick 
Game Department is quoted as saying that "reports from 
wardens and woodsmen show that caribou probably remained 
in the Province in small quantities until about 1926, and one 


warden actually reported seeing five caribou east of Bartibogue 
Station during the spring of 1927." Since then, however, the 
herd south of the St. Lawrence seems to have vanished, except 
for a small remnant still surviving in the Gaspe region. This 
remnant is chiefly confined to the Shickshock Mountains in 
the northern part of the country and though small is never- 
theless of sufficient size to allow a few to be shot each year. 
There is a considerable barren area on the summits of some of 
these mountains that is attractive to caribou, and the sur- 
rounding wilderness is at present little disturbed. As the last 
remaining woodland caribou south of the St. Lawrence River, 
this little group should be given careful protection until it 
appears that it is numerous enough to survive. One may 
believe, however, that it will not persist for a great many years, 
when annually hunted. In southern Labrador "a few" wood- 
land caribou are found "here and there in the interior north of 
the St. Lawrence River and a small number scattered through 
the back districts of northwestern Quebec . . . The situ- 
ation in Ontario is even less hopeful. On a map prepared at 
the Royal Ontario Museum of Zoology in 1935, caribou are 
considered to have disappeared Entirely from the east of a line 
drawn from east end of Lake Superior to James Bay, the dates 
of extirpation being dotted on the map here and there 1894, 
1908, 1912, 1919, 1926, etc. The most southern band at present 
is said to occupy Shakespeare Island in Lake Superior, with a 
few about Lake Nipigon, Lake of the Woods, and the Rainy 
Lake area. Local bands exist north of the Canadian National 
Railway lines, but the most recent reports are to the effect 
that the numbers of caribou are not large even in the more 
northern parts of Ontario." The range in southern Labrador 
probably extends to Hamilton Inlet and Sandwich Bay on the 
east, and on the Hudson Bay side to Great Whale River or 
thereabouts. According to Eidmann (1935) it has been killed 
out over most of this range, but small herds are still occa- 
sionally met with, especially in the southeastern part of Labra- 
dor, where a small herd appeared in the Matamek area in 1930 
and several were killed. Its extermination has been especially 
hard for the native Indians, who relied on it in part for food 
and clothing. 

Minnesota is the only one of the eastern United States in 
which caribou still exist in a wild state. C. L. Herrick (1892) 


found them 50 years ago "only about the headwaters of the 
White-face River and along the St. Louis River near Knife 
Fall. There it was in 1884 not rare, though so shy as to be 
secured with difficulty. Along the North Shore of Lake 
Superior it is less shy and the animals may be seen feeding 
quietly in groups along the upland meadows." G. S. Miller 
(1897b) in 1896 found these caribou "very abundant on the 
north shore of Lake Superior" and saw "heads, antlers, and 
jaws of caribou at White River, Peninsula Harbor, Schreiber, 
and Nepigon. A wet pasture among the hills a mile or more 
northeast of Peninsula Harbor is a favorite feeding ground of 
these animals. " At the west end of Lake Superior, in what is 
now Quetico Provincial Park of Ontario, caribou are extinct, 
"although this region was once the approximate southern 
limit of its winter migration. There are no records in recent 
years. Nash reported it as very shy but not rare in the White- 
face and St. Louis River country of northern Minnesota as 
late as 1894 . . . There is an abundance of its favorite 
food, the caribou moss, Cladonia rangiferina, throughout the 
region" (Cahn, 1937). A century ago there were caribou 
in Chippewa County, northern Michigan (Schorger, 1940). A 
recent report by BreckenridgeXl935) shows that a last remnant 
of the woodland caribou still holds out "in the muskeg country 
lying between Upper Red Lake and Lake of the Woods in 
northwestern Minnesota. " In company with game officers he 
spent from February 28 to March 3, 1935, investigating the 
status of these animals. Of three adults seen at close range on 
March 1, one retained its antlers. Evidence of at least six ani- 
mals in this region was obtained, a number that is reported in the 
August, 1939, Biological Survey census to have been increased 
to 12, but the basis of this estimate is not given. The district 
where these caribou live is now included in the Red Lake 
Game Refuge, where they may be considered fairly safe except 
from wolves which still are found in small numbers. The 
refuge includes over 400,000 acres of land, "which is prac- 
tically worthless, not only to the agriculturalist, but to the 
forester" (Swanson, 1936). 

Finally, in the hope of building up the stock more rapidly, 
W. T. Cox (1941), in charge of the refuge, arranged to introduce 
new stock from Saskatchewan. With the aid of Indians, seven 
calves and an adult bull were captured alive and after pre- 


liminary conditioning were brought in 1938 to the refuge. 
Nearly a year later the bull was seen ranging with the cows. 
The calves, however, were kept in a corral and small pasture 
and later were transferred to a larger fenced area, to be liberated 
later. At last accounts the little group was thriving encourag- 
ingly, and every effort is being made to ensure the success of 
the attempt at colonization. The danger is that the animals 
may wander outside the reserve and fall prey to settlers or the 
Indians on a nearby reservation. 

To sum up, the woodland caribou, once so common in the 
muskeg areas from Nova Scotia to western Minnesota and 
north to southern Labrador and James Bay, is now everywhere 
so greatly reduced that it is in actual danger of extermination. 
South of the St. Lawrence a single small group still exists in 
the mountains of northern Gaspe; it is gone from Nova Scotia, 
New Brunswick, southern Quebec and northern New England. 
A few scattered groups still remain in southern Labrador, but 
to the westward it is all but gone from Ontario, and a very 
small group still holds out in northern Minnesota. The main 
factor in its extermination has no doubt been overshooting; 
other factors are doubtless the animal's innate aversion to the 
presence of settlements, thus tending to a restriction of its 
range and lowered prolificness, its gregarious habits and curi- 
osity often making easier the killing of several individuals at 
a time by hunters; finally forest fires may affect the food 
supply, and wolves, though fewer now than formerly, may be 
a factor. Of possible diseases nothing is known, but that deer 
of this group are subject to serious attack by bot flies is well 
ascertained, although what effect these may have in causing 
lowered vitality is not clear. 


Rangifer terraenovae Bangs, Preliminary Description of the Newfoundland Caribou, 
p. 2, Nov. 11, 1896 (privately printed) ("Codroy, Newfoundland"). 

FIGS.: Grant, 1902, two pis. unnumbered (14, 15) (heads); Prichard, 1910, col. pi. 
opposite p. 80; Dugmore, 1913, many photographs. 

The caribou of Newfoundland is not very different in general 
from the woodland caribou of the neighboring mainland, even 
though usually accorded specific status. It is said to be lighter 
in color, the back, sides, and legs drab, somewhat mixed with 


yellowish- white hairs paling on flanks to white. Face and inner 
surface of ears rich bister brown, nose and chin white; tail 
short, drab above, white beneath (Bangs). The antlers are 
rather distinctive, being "low, widely spread, much forked and 
with the points pointing forward and inward" (Bangs). They 
are, in a well-developed head, rather shorter of beam, with 
large palmated brow and bez tines. The skull is said also to 
be somewhat larger than in the typical woodland caribou. 
An adult male stands from 46 to 49 inches at the shoulder. 
Does are in about half of the cases provided with small branch- 
ing antlers. Weight of a full-grown male up to about 300 

Both Prichard (1910) and Dugmore (1913) have given 
excellent accounts with illustrations of this caribou. According 
to the former, the population of this species in the first decade 
of the present century was said to consist of three groups: 
One north of the railway that crosses the island from east to 
west, especially frequenting the Humber River valleys and 
Birchy Lake region; the main herd, largely south of the rail- 
road, inhabiting the central and southern parts; and a small 
"herd" consisting of a few animals that still survive in the 
Avalon Peninsula, south of St. John's. The last group is 
believed to be nonmigratory, but the other groups perform 
annual north and south migrations, with more or less regularity, 
although they may not always be found at the same places in 
different seasons. The first of these groups is said to be the 
best known and most accessible and crosses the railway in its 
southward wanderings, especially in the vicinity of Howley. 
To this region resort the greater number of sportsmen in 
search of trophies and of settlers who to some extent depend 
on caribou meat. 

In 1913 Dugmore believed that a conservative estimate of 
the caribou population of the island was not less than 150,000 
animals. Some estimates were much larger. It is difficult to 
judge of the general status, since those visiting the hunting 
grounds annually for short seasons may not find the larger 
groups each time or the caribou may be for various reasons 
less in evidence in different years. The interior barrens are 
vast and travel is difficult. Nevertheless the consensus seems 
to be that the numbers have greatly declined in the last two 
decades, owing in large part to overshooting. Jacobi (1931) 


writes that in February, 1899, an observer saw 550 carcasses 
unloaded in the harbor of St. John's and two weeks later about 
the same number, so that the price of meat fell to 3 cents per 
kilogram. However, as late as 1913 they were still common 
and could be seen in large bands, especially during the migra- 
tions, but large heads were more difficult to secure. 

In a letter to Dr. Francis Harper, dated October 4, 1934, 
Greville Haslam writes that he had spent nine of the previous 
fourteen summers in the island, and found in 1919, 1920, and 
1921 "a few caribou" each year in the region north of the 
railroad near the Humber River and Birchy Lake. "There 
was no question in anybody's mind that the caribou were dis- 
appearing, but they all claimed that this was because the 
animals no longer migrated, and that they would be found in 
large numbers in some other section, especially the country 
lying at the head of the Gander River and extending around 
Mount Cormack and west around Meelpaig Lakes. In 1929 
we spent three weeks in this country but saw not a single cari- 
bou and only a few tracks made some months before. In the 
summer of 1934 we spent three weeks on the Serpentine River 
but saw no tracks." It seems \hat the numbers have appar- 
ently fallen off considerably in late years, but exact statistics 
are unavailable. At all events there was a general impression 
in 1934 that caribou were again increasing and in that year 
the Government permitted an open season, charging $50 for 
a license to shoot a caribou. Brooke Dolan reported that 
according to a Newfoundland official, 60 licenses (including 
two to aliens) were issued in 1936 and that some caribou were 
killed. One herd of about 700 was reported seen north of the 
railroad, but shooting was permitted to the southward of the 
railroad only. In 1938 John K. Howard found small numbers 
a little distance north of the head of White Bay. Thus it 
seems that there are still a good many caribou left in the 
country, but in order that the numbers may be maintained at 
an economic level some care in administration is needed. 

Dugmore (1913) presents some interesting notes on habits. 
Summer in Newfoundland begins late in June. During this 
month the young caribou are born, and at this time the does 
seek the thick forests of spruce and fir. Usually there is a single 
young at a birth but twins are not rare. With the passing of 
wolves on the island, the lynx is about the only potential 


enemy. On the other hand, the swarms of black flies and 
mosquitoes must be a great source of annoyance to the animals 
as they are to human beings at this season. "During the 
warmer months the caribou are more or less solitary in habit, 
going about singly or in pairs and only rarely in small herds of 
half-a-dozen or more. In the day-time they keep very largely 
to the woods, coming out to feed at the approach of evening 
. . by September their habits have completely changed 
and they become almost entirely diurnal," and bands gather. 
An attempt is being made to introduce this caribou from 
Newfoundland into Nova Scotia, in the hope that it may, if the 
plan is successful, in time replace to a certain extent the 
eastern woodland caribou, which has for a number of years 
been extinct in this province. On April 10, 1939, nine females, 
of which five were "with calf," arrived in Halifax from New- 
foundland. The five pregnant females were at once taken to the 
Liscombe Game Sanctuary in Guysboro County, while the 
four others were being held temporarily at Halifax pending the 
arrival of the three males, which were to be imported in order 
to start this new herd (R. W. Tufts, 1939). The result of this 
experiment may be awaited with interest. 


Genus tarandus var. (3 sylvestris Richardson, Fauna Boreali-Americana, vol. 1, p. 250, 

1829 (southwestern shores of Hudson Bay). 
FIG.: Hollister, 1912b, pi. 2, fig. 2 (skull). 

The western woodland caribou is much the same as the 
eastern animal, but according to Hollister (1912b) it may be 
distinguished by its longer and slenderer skull, with a narrower 
rostrum and larger teeth. The tooth rows, especially the lower, 
are longer. The neck and head are darker, the ears, back, and 
sides of the neck are much darker, the hairs brown to their 
roots. Total length of a male skull, 417 mm.; upper tooth 
row, 107 mm. These characters, while not very sharply 
marked, are supposed to distinguish the animals west of Hud- 
son Bay; presumably those of the Lake Superior region and 
James Bay are intermediate but may be referable to this 
rather than to the typical race. 

The precise range of this race has not been well defined but 
is believed to have originally extended from the southwestern 


shores of Hudson Bay westward across northern Manitoba to 
the Anderson River, overlapping the winter range of the 
barren-ground caribou to some extent and possibly reaching 
the Mackenzie River region. 

Dr. R. M. Anderson (1939b) writes that it is widely dis- 
tributed in the timbered portions of the Northwest Territories, 
but never common, ranging as far north as Great Bear Lake. 
It is fairly certain that human settlement is the controlling 
factor in the present range of the species. "Small numbers of 
woodland caribou are found in the wooded parts of northern 
Saskatchewan and Alberta, but many reports indicate that 
the large number of trappers and prospectors who have 
strung trap lines all along the northern parts of the Prairie 
Provinces are rapidly causing the woodland caribou to disap- 
pear. Bush fires destroy their winter food of lichens which are 
slow to recuperate, and the woodland caribou are shy of settle- 
ments as well as being easy to kill on the 'barren' openings in 
the forest in winter." Writing of this caribou in 1902, Preble 
says that he found it throughout the region he traversed be- 
tween Norway House and Hudson Bay. Between York 
Factory and Fort Churchill 3, few small bands are found 
throughout the year on the "barrens." Dr. Milne, who had 
resided 14 years at York Factory, told Preble that Cape 
Churchill was considered a good place to hunt them at any 
time of year and that he believed the small bands occurring 
here formed the northern fringe of those that migrate to the 
coast in spring, the great majority of which cross to the south 
of Nelson River. Their return movement occurs from about 
the middle of October to the last of November. "During 
these semiannual movements the animals are much pursued, 
especially in the fall, when the weather is usually cold enough 
to preserve the meat for winter use." A resident at Oxford 
House said that the species was much less common than 
formerly. In his report on mammals of the Athabaska- 
Mackenzie region, Preble (1908) wrote that it is found in the 
"country between Lake Winnipeg and Athabaska Lake, and 
though nowhere in large numbers is more abundant on the 
southern than on the northern shores of this lake. Between 
Athabaska and Great Slave lakes . . . the animal is 
met with chiefly on the west side of Slave River, and through 
all the country lying between Peace River and Great Slave 


Lake." Caribou still exist in small numbers in the Peace 
River district, as in the vicinity of Peavine Lake and on 
Mount Bickford and Tuscoola Mountain (Cowan, 1939), and 
are presumably of this race. According to Anderson, there is 
some evidence that in areas visited by game officers in the 
winter of 1937-38, the caribou are increasing. With proper 
protection, they should hold their own in most of their range 
for the present. 


Rangifer montanus Seton, Ottawa Nat., vol. 13, p. 129, Aug., 1899 (Selkirk Range, 

near Revelstoke, British Columbia). 
FIGS.: Allen, J. A., 1902b, pp. 154, 155, 156, 157, figs. 3-6 (antlered skulls). 


Rangifer fortidens Hollister, Smithsonian Misc. Coll., vol. 56, no. 35, p. 3, Feb. 7, 1912 
("Head of Moose Pass branch of the Smoky River, Alberta (northeast of 
Mount Robson) "). 

FIG.: Hollister, 1912a, pi. 10 (antlered skull of type). 

Hollister, in 1912, described the large caribou of the woodland 
type from Mount Robson region as a species distinct from R. 
montanus of the Selkirk and Gold Ranges in southeastern 
British Columbia. The two must be closely related (the type 
localities are little over a hundred miles apart) and may even 
prove to be much the same when further study can be made 
with adequate specimens. Hence it is convenient to treat both 
together here. 

Both agree in being darker in color than the more eastern 
forms. Dr. J. A. Allen (1902b) describes R. montanus in 
September coat as in general terms "a black caribou, with the 
neck and shoulders, especially in the males, much lighter than 
the body and limbs." The "whole body and legs blackish 
brown, varying (in different specimens) to glossy black over 
the middle of the dorsal area from shoulders to rump; lighter, 
more brownish black, on the flanks and ventral surface; 
inguinal region, sides and under surface of tail, a narrow band 
bordering the hoofs, and ventral median line of neck, grayish 
white; nose and edge of lips grayish white; sides of neck 
grayish brown varied with blackish and, in the males, tinged 
more or less with rusty. The females are much darker than 
the males, especially on the neck and shoulders, but have the 



grayish white areas of the males replaced by nearly pure 
white." The distinction between the two "species" seems to 
be that in R. montanus the size is somewhat less in skull and 
teeth, with shorter beams to the antlers and a more exuberant 
development of tines ; whereas fortidens is said to be largest of 
the caribou with less branching and less shortened antlers. 
No doubt both should eventually be regarded as races of R. 

The range of the more southern montanus was the Selkirk 
and Gold (or Columbia) Ranges and doubtless the adjacent 
country, southward to the northern border of Washington. 

Newfoundland caribou (Rangifer terraenovae) 


Anthony states that it formerly reached the mountains of 
Montana and Idaho, but certainly it has not been known in 
these States for a long time. In the State of Washington, 
Taylor and Shaw (1929) wrote that it was formerly "of 
occasional occurrence along the Canadian boundary in the 
northeastern part of the State, south to Usk, and west to 
Okanogan County" but that it was exterminated a decade 
ago (about 1920). Nevertheless the latest wildlife census by 
the IT. S. Biological Survey credits the national forests of the 
State with an estimated four individuals in 1938. In a letter 
to Dr. Francis Harper in April, 1937, I. M. Cowan wrote that 
in British Columbia there are still a number of montanus but 
none now south of the main line of the Canadian Pacific Rail- 
way. In general, caribou are "definitely on the wane through 
overhunting," he says, "mainly by Indians." 

Cowan's statement probably applies in part to the Mount 
Robson form, fortidens, as well, for he says they are "almost 
gone" in the Chilcotin region. Hollister in 1912 said they were 
not common along the line of the railroad east of the boundary 
line of Jasper Park but were formerly common down to the 
eastern foothills. Dr. R. M. Anderson wrote in 1938 that 
R. montanus "is still fairly common in the Rocky Mountains 
of western Alberta, but the animals are extremely scarce on 
most of their former range in central and southern British 
Columbia." In 1929 he had "fairly conclusive evidence of the 
presence of a small band ranging along Summit Creek on both 
sides of the British Columbia-Idaho border, and if these still 
exist they are probably the most southern representatives. 
At that time a few were still supposed to be in the mountains 
southwest of Nelson, British Columbia." Carl Rungius (in 
Ely et al., 1939) has lately published an account of hunting R. 
fortidens in the Mount Robson region, but he gives no dates. 
At the time, caribou were fairly numerous there. 

Family ANTILOCAPRIDAE : Pronghorns 

The pronghorns are the last surviving members of a family, 
the Antilocapridae, that has developed exclusively in North 
America. Though related to the true antelopes of the Old 


World, they differ in that the horn sheaths are forked instead 
of simple and are shed and renewed annually instead of being 
permanently retained. In many of the fossil forms, the bony 
core of the horns was forked to correspond to the forked sheath, 
but in the living species this front branch of the horn core is 
represented by a slight bulge only. In some of the extinct 
species the horns showed a twisted core recalling that of the 
Old World bushbucks. Only a single species is known to have 
lived down to the Recent period, but smaller forms, such as 
Capromeryx, with a forked horn core, must have persisted to no 
very distant period, and have left their remains in the tar pits 
of Rancho La Brea, California, and elsewhere in the Southwest. 
Four races of the pronghorn have been described, but they 
differ in such slight characters that they may be best con- 
sidered together. 


Antilope americana Ord, Guthrie's Geography, 2d Amer. ed., vol. 2, pp. 292, 308, 1815 

("Plains and highlands of the Missour^"). 
FIGS.: Stone and Cram, 1902, pi. facing p. 64 (photographs); Nelson, 1916, p. 451, 

upper fig. (col.); 1925, pis. 1-6 (photographs and habitat). 


Antilocapra americana mexicana Merriam, Proc. Biol. Soc. Washington, vol. 14, p. 31, 
Apr. 5, 1901 ("Sierra en Media, Chihuahua, Mexico"). 


Antilocapra americana oregona V. Bailey, Proc. Biol. Soc. Washington, vol. 45, p. 45, 
Apr. 2, 1932 ("Hart Mountain (Warner Mts.), Oregon"). 


Antilocapra americana peninsularis Nelson, Proc. Biol. Soc. Washington, vol. 25, p. 
107, 1912 ("Forty-five miles south of Calmalli, Lower California"). 

Light of body and limb, the pronghorn stands about 34 to 
36 inches at the shoulder, with a total length of about 54 inches 
in adult males, females slightly less (Anthony). The forehead, 


nape, and upper part of the back and the outside of the limbs 
are rich reddish brown, with a mixture of blackish on the 
muzzle and in the short mane. On the throat the reddish 
brown of the upper parts extends as two collars, separated by 
a white space, around the throat, although the lower one is 
often incomplete in the middle line below. Under parts and 
flanks and a prominent rump-patch, white. Tail and a central 
dark line dividing this patch are colored like the back or 
darker. Horns erect and diverging, the posterior and longer 
point curving backward, the anterior point short and blunt, 
projecting forward. 

The Mexican race is similar but paler, with a tinge of cinna- 
mon. That of the peninsula of Lower California has the ears 
darker, with the facial markings dark and strongly contrasting 
with the pale areas. The Oregon race is slightly larger than 
the typical race of the plains, with relatively larger feet, 
longer horns, and slightly paler color. The distinctions between 
these races are slight and the distributional areas need more 
careful mapping. 

Pronghorns are characteristic of open plains country of a 
semi-desert nature and seldom enter tree-grown areas except 
in winter to find shelter from storms. Their flashing of the 
erectile white hair of the rump-patch makes a remarkably 
striking semaphore visible at a great distance in sunlight and 
affords an automatic danger signal to others of their kind. 
They go in bands of varying size according to the circumstances 
and when once alarmed can reach a great speed, seeming 
fairly to fly over the ground. A curious habit seen in this and 
in some other animals, as gazelle in the Gobi or seabirds in open 
ocean, is a seeming desire to match speed with pursuers, and, 
attaining a sufficient lead, to cross in front. Another char- 
acteristic trait is curiosity, leading animals often to approach 
unfamiliar appearing objects. Nelson tells of enticing shy 
animals up within gunshot by donning a white sheet and ap- 
proaching them on all fours. 

In an important paper on the status of the pronghorned 
antelope, Dr. E. W. Nelson (1925) presents a careful summary 
of its former and recent distribution. It originally ranged over 
an enormous area, from the present provinces of Manitoba, 
Saskatchewan, and Alberta in the north to the southern part 
of Texas and the Mexican tableland in the south, and from the 


eastern edge of the plains in Minnesota, Iowa, Kansas, and 
Oklahoma westward to eastern Washington, Oregon, and the 
valleys and coast of southern California and most of the penin- 
sula of Lower California. "In Mexico it occupied the open 
plains country of the tableland south almost to 20 of latitude, 
nearly to the valley of Mexico " and the western part of Sonora. 
Originally it was abundant and well distributed over this 
territory and exceeded the bison in numbers where both 
occurred together. It is estimated by Nelson that at the time 
of the settlement of this continent by Europeans the prong- 
horn population was "not less than thirty to forty millions, 
possibly more." A recent census of those still existing in the 
United States, Canada and Mexico (1922-24), indicates that 
of these vast numbers about 30,000 then remained, but a later 
figure (1939) makes this over 180,000. 

The pronghorn just reached the prairie country of south- 
western Minnesota in its northeastward distribution, but 
Herrick, writing of it in 1892, speaks of it as having "long since" 
gone. That it may even have penetrated farther east is 
possibly indicated by the recent discovery of a horn buried in 
a few inches of earth at Moline, ill., although it may have been 
brought there by Indians in earlier years (Fryxell, 1926). 
Formerly abundant all over the Dakotas, it began to decline 
in the late seventies, and with the encroachment of settlement 
gradually disappeared. By 1924, Nelson wrote that but five 
small herds aggregating 225 animals remained, and these in 
the southwestern corner; in South Dakota about three times 
that number were found in the western half of the State. 
"Of the countless thousands of antelope which once roamed 
the plains of Nebraska but 10 small bands remain, containing 
a total of about 187 animals." "At one time Kansas was 
inhabited by myriads of pronghorns, and for years after the 
construction of the transcontinental railroads they were a 
familiar sight to passengers on the trains. In 1923, however, 
they had become almost exterminated throughout the State," 
and the few remaining were to be found only in the extreme 
southwest corner where they wandered at times into the 
adjacent parts of Oklahoma. Otherwise the species once so 
common in this State was gone, but in 1910 an attempt was 
made to establish a herd in the Wichita National Game Pre- 
serve in Comanche County. After several unsuccessful im- 


portations of antelope from Yellowstone National Park, the 
difficulties were finally overcome, and in 1925 Nelson reported 
that the birth of three pairs of young had brought the total 
number of this small band up to 17. 

Except for Oregon the Rocky Mountain States seem now to 
hold the largest antelope populations. Nelson's account shows 
Montana with some 44 herds and a total population of over 
3,000; Wyoming leads with 27 areas in which a total of nearly 
7,000 is carried; Colorado has 28 localities mostly in the eastern 
half of the State where over 1,200 pronghorns are found; in 
New Mexico the numbers were decreasing, but a total of 31 
localities and nearly 1,700 animals was recorded in 1924. 
In Arizona the estimates were 18 bands totaling about 650 
animals; they appeared to be increasing on cattle ranges but 
decreasing on sheep ranges in the State. In Utah the number 
of bands was 10, with about 670 animals; in Idaho 14 bands 
with about 1,500 animals. In Nevada, where once they were 
plentiful, they were found in only 11 limited areas, but since 
some of these are areas sparsely occupied by man the numbers 
of antelope are large, aggregating over 4,200. Southeastern 
Oregon, where the form oregonus is found, "forms part of a 
rough, rocky desert covering also northern Nevada and south- 
western Idaho, on which natural conditions have been ex- 
ceedingly favorable for antelope. This region constitutes one 
of the few areas in the United States where large herds of these 
animals numbering hundreds still continue to congregate during 
the winter season." Here they have increased in recent years 
and were in 1924 estimated at over 2,000 animals. In Texas, 
antelope formerly abounded on the plains in the western part, 
"but with the occupation of the country they have decreased 
until it has been possible to obtain definite information of only 
42 existing bands, numbering about 2,400 animals, for the 
entire State." While most of these are in the western third, a 
few hundred are also found in the southwestern tip. A few 
antelope remain in California, where once they were abundant; 
in 1923 there were six widely separated areas in the State 
where small bands were found totaling about 1,057. 

In comparison with these estimates of 1924, the following 
issued by the U. S. Biological Survey are significant: South 
Dakota, 4,508 (large increase) ; Nebraska, 750 (large increase) ; 
Kansas, none (decrease); Oklahoma, 36 (increase); Montana, 


6,740 (doubled); Wyoming, 24,071 (more than tripled); Colo- 
rado, 1,770 (slight increase); New Mexico, 26,564 (great in- 
crease); Arizona, 9,410 (great increase); Utah, 35,350 (great 
increase); Idaho, 12,328 (great increase); Nevada, 12,700 
(about tripled) ; Oregon, 28,550 (great increase) ; Texas, 9,075 
(great increase); California, 14,212 (great increase). The 
estimate published by Nelson in 1925 gave a total of 26,604 
pronghorns in the United States; the 1939 estimate by the 
U. S. Biological Survey was 186,114, or nearly a sevenfold 
increase for the same States. It is clear that the animal is no 
longer in danger; in fact, in some areas it is common enough to 
cause resentment among the ranchers for fear that the herds 
will make the grazing less available for their cattle. In several 
of the States antelope refuges have been established, which, 
especially in Nevada and Oregon, are well populated by the 

Obviously, with the continued increase of these herds and 
their tameness as a result of protection, there will come points 
where the numbers are too great, and their competition for 
food with domestic stock must result in measures being taken 
to reduce the numbers to a reasonable proportion. It remains 
to be seen whether this is best done by having brief open 
seasons such as have been declared in Wyoming and Nevada 
or in some other way. The natural enemies of the pronghorn 
are chiefly coyotes, which kill the fawns, and bobcats, which in 
winter may kill a few. Wolves formerly were doubtless a 
principal enemy. With present measures against these ani- 
mals, the menace seems slight. These antelope are said not to 
do well under fence, for it has been found that "within such 
areas" they seem "to lose their freedom of movement and be- 
come extraordinarily helpless. This is particularly the case 
during heavy snowstorms, when they remain within more or 
less definite areas, in which predatory animals capture them 
with surprising ease." 

In Canada the pronghorn formerly ranged eastward to 
southwestern Manitoba but is now restricted to southwestern 
Saskatchewan and southern Alberta. Nelson's (1925) estimate 
for 1924 gave a total of 1,327 antelope for Canada, but Ander- 
son (1939b) shows a figure nearly double this, about 2,400 in 
1932, as a result of protection, and adds that in 1938 the Fish 
and Game Commissioner of Alberta estimated the antelope 


population of that province at 15,000, and there have been two 
short open seasons. In certain districts they have even ex- 
tended their range north of the South Red Deer River, as well 
as into the adjacent parts of Montana and Saskatchewan. 
In the latter province there was an open season in 1936 during 
which 267 antelope were taken. 

In Mexico the situation is less clear, but Nelson's (1925) 
report gives an estimate of about 2,400 animals, of which 500 
are in Lower California, the rest being in the States of Sonora 
adjoining and Chihuahua, Durango, and Coahuila to the east- 
ward. In Lower California the pronghorn was formerly 
found over much of this desert country nearly to the tip of the 
peninsula, which is mountainous and unsuitable for them. 
At present the range extends, according to Nelson, only to the 
basal half, on the plains east of the central mountain range, 
and on the Desert of Vizcaino west of it. With the establish- 
ment of a close season at the time of Nelson's report, he be- 
lieved that their prospect of surviving in these sparsely settled 
desert lands was very good. 

From the human viewpoint pronghorn antelope may be a 
source of food in some areas, and they have an esthetic and 
recreational value as well for those who enjoy the sight of wild 
game or take pleasure in hunting it with gun or camera. 
Large herds such as gather in winter may, on the other hand, 
cause some competition with grazing stock of which ranchmen 
are jealous. On the whole, however, it seems that the preserva- 
tion of the species at least in certain large areas of our West is 
well assured at present and that its numbers can be readily 
controlled where the necessity exists. Some pertinent sugges- 
tions for the management of this antelope in the arid South- 
west have been put forth by Dr. W. P. Taylor (1936), who 
points out that in these regions herds of pronghorns in severe 
winter weather are wont to come down off the mountains to 
lower levels where they come into competition with cattle on 
the ranches for winter food. Ordinarily they do not like sheep 
ranges but prefer those suitable for larger stock. These factors 
are often of critical value in the selection of reserves for their 
preservation. The ranchmen have often done much to en- 
courage these animals and give them protection. 


Family BOVIDAE: Sheep, Goats, Cattle 

Much has been written on the relationships of the muskox to 
other ruminants. Although many fossil skulls have been found 
of this and related animals, there seems to be little evidence 
that the type represented by the living species was derived 
from the Old World. It is therefore believed that it must have 
originated in the New World and in late geological times spread 
into northern parts of the Old. While some zoologists have 
thought it related to the sheep or the goat-antelopes, the latest 
suggestions favor its close affinity to the bison. The single 
living species is of high Arctic distribution from (until recently) 
Alaska to Hudson Bay and the Arctic Archipelago, and thence 
across northern Greenland to the eastern coast of that country. 
Over this area it becomes differentiated locally into three 
generally recognized races, for a full account of which the 
reader is referred to the extensive papers of J. A. Allen (1913) 
and Elisabeth Hone (1934). 


Bos moschatus Zimmermann, Geographische Geschichte, vol. 2, p. 86, 1780 (based on 
Pennant's description; type locality "therefore the region adjoining Hudson Bay 
between the Seal and Churchill rivers (about latitude 59) ", Keewatin (J. A. 
Allen, 1913)). 

SYNONYM: Ovibos moschatus mackenzianus Kowarzik, Fauna Arctica, vol. 5, pt. 1, pp. 
97, 116-122, 1910. 

FIGS.: Allen, J. A., 1913, figs. 28, 30, 32, 39 (exterior and skull). 

The typical race of the muskox is "very dark, nearly black 
on the head, neck, sides, and underparts, with the feet and 
nose white; the back is lighter (browner black), with a still 
lighter "saddle" behind the shoulders. There is no white 
area on the head in adult males, although individual white 
hairs may often be found on the face; in young animals and 
females there is sometimes quite a trace of white on the front 
of the head" (J. A. Allen, 1913). The long hair of the body 
forms a protecting fringe on the sides. The horns are charac- 
teristically dark brown and are very broad at the base in pro- 
portion to their length. They sweep downward and slightly 
forward, then turn upward to form a hook; the record length 


of horn on the outside curve is 29 inches. A large bull killed 
at Aylmer Lake was measured by Seton as follows: Total 
length, 96 inches; tail, 4 inches; height at shoulder, 59 inches. 
The basal length of the skull in adult males is about 466 mm., 
with a maxillary toothrow of 132 mm. 

The continental range of the muskox formerly was more or 
less coextensive with the barren grounds west of Hudson Bay, 
extending to the Arctic coast and the adjacent islands, which 
must have been reached by crossing on the ice in winter. On 
account of certain characteristic habits this species became 
peculiarly liable to extermination when hunted by men with 
modern weapons. Lacking the speed to escape such enemies 
as wolves, and having the habit of living in small groups, these 
animals when attacked would form a close circle, the adults 
with heads out to the enemy, the young huddled at the inner 
side against their mothers' quarters. When closely approached 
by wolves, one of the company would suddenly dash out from 
the ring in an endeavor to rip one of the assailants with its 
horns, then as quickly dart back into its place. This method 
of defense, while probably efficient against wolves, its only 
natural enemy, was often its undoing when attacked by men 
with modern guns, for the entire group might then be killed 
at close quarters. Against Eskimo and Indians, who in former 
days often depended in part upon muskox for winter food, this 
defense was probably more effective before the use of firearms. 
During the summer season the almost impenetrable muskegs 
of the north form a fairly safe retreat, and the animals' long 
silky coat seems sufficient protection against cold and bliz- 
zards. But against Eskimo and Indians with modern arms, 
they can make little stand, while the thoughtlessness of human 
pursuers in wantonly killing entire groups has in places brought 
the species to the verge of extinction. 

There is evidence that at no very distant time the range of 
the muskox extended westward at least to Point Barrow, 
Alaska, where according to Frank Russell, in 1898, the oldest 
natives "say that their fathers killed muskox, which were then 
abundant." The fact that skulls of muskoxen have been found 
on the neighboring tundra confirms this statement. J. A. 
Allen (1913), in reviewing this evidence, believes that muskoxen 
existed on the tundra of northern Alaska till about the middle 
of the nineteenth century. A. J. Stone, after extensive inquiry 


among the Indians and Eskimo west of the Mackenzie, came 
to the conclusion that the species had not inhabited that region 
for a very long period. Their western limit, at the close of the 
nineteenth century, he writes (1900); is "far to the east of 
Anderson River and Liverpool Bay." In the region about 
Artillery Lake, where the muskox was common shortly before 
1901, it was practically exterminated a few years later. Con- 
tinual hunting by the Eskimo from the coasts and by the 
Indians from the southern borders of the barren grounds had 
reduced their numbers still further in the early years of this 
century, so that Dr. R. M. Anderson, in 1917, believed that 
they were extinct west of the Tree River and much reduced 
elsewhere, especially near the accessible regions along the 
Arctic coasts, 

In 1913 Dr. J. A. Allen well summed up the situation at that 
time as follows: "With all its natural fitness to survive, the 
muskox is doomed wherever it can be utilized by man as a 
commercial asset. The history of its restriction in range and 
decline in numbers over the western part of its former range 
during the last quarter of a century . . . indicates clearly 
its fate wherever it can be reafched by the white man, either 
directly or through his Eskimo or Indian allies . . . But 
wherever its range is shared by the Eskimos, as many portions 
are, the muskox's fate is sealed, as shown by its extermination 
over the greater part of the large expanse of land known as 
Victoria Island. It will also rapidly decline over the more 
accessible parts of Ellesmere Land and Grinnell Land, through 
intrusions of ambitious sportsmen. It is doomed throughout 
the mainland of northern Canada unless the Canadian Govern- 
ment takes the utmost care in restricting the killing . 
Much could be done to preserve a considerable remnant of 
these unsuspicious animals if the Danish, Canadian, and other 
governments would declare muskox peltries contraband and 
suppress all traffic in them, while the Canadian and Danish 
governments might set aside reservations within which neither 
Eskimos and Indians nor white hunters should be allowed to 
kill them." Fortunately the Canadian Government in 1927 
established a sanctuary known as the Thelon Game Sanctuary, 
with an area of 15,000 square miles, northeast of Great Slave 
Lake, which so far as then known included the last important 
herd of muskoxen remaining on the mainland of Canada. No 


person may hunt or trap within its borders, and entry is pro- 
hibited except under permit from the Minister of the Interior. 
In 1929 it was estimated that there were 250 muskoxen in this 
sanctuary. In 1935 a representative of the department made 
an airplane survey and counted 180 muskoxen north of the 
junction of the Thelon and Hanbury Rivers (Cameron, 1936), 
and Dr. R. M. Anderson (1937) adds that there are a few 
scattered bands and individuals still farther north, although 
few if any animals are now known to occur very near the 
Arctic coast. With this protection it seems likely that the 
remnant of the continental muskox is safe enough at present. 



Ovibos moschatus niphoecus Elliot, Proc. Biol. Soc. Washington, vol. 18, p. 135, Apr. 

18, 1905 (type locality, head of Wager Inlet, Hudson Bay, not "600 miles north 

of Hudson Bay" as originally given). 
FIG.: Kowarzik, 1910, vol. 5, fig. 8. 

In his key, J. A. Allen (1913) characterizes this race as having 
"usually no coronal nor facial white areas in adult males, but 
traces of them (often well developed) in young males and 
females; horns more slender and longer in proportion to their 
basal breadth, and generally light-colored; toothrow relatively 
longer (max[illary] series, cf, 130 [mm.]); basal length of 
skull in old males, 442 mm." The males are usually more 
intensely black than the typical race, and the horns are 
lighter colored. While males generally lack the white on face 
and crown, the females and young are much like the Green- 
land race. In body size it is much as in the typical race. 

The range of this race, and indeed its characters, do not 
seem to be very well defined. The region whence came the 
original specimens was at the head of Wager Inlet, in the 
northwestern corner of Hudson Bay. According to Captain 
Comer, who collected them, the muskox does not now range 
south of Chesterfield Inlet, but it is believed that the distribu- 
tion of this race extends northward from the latter point to the 
Arctic coast of the mainland and inland for an undetermined 
distance. They are not found on Melville Peninsula or in 
Baffin Land, nor do the natives there have any tradition of 
their former occurrence. North of Baker Lake the natives say 
that the muskoxen are larger, perhaps representing the typical 


race. Captain Comer states that formerly the native Eskimo 
hunted these animals but seldom on account of the risks in- 
volved in making the inland trips, but are encouraged to under- 
take their pursuit by traders who are eager for the pelts. Up 
till recent years at least, the remoteness of the region where 
these muskoxen occurred was itself a factor of safety, for few 
hunters would enter it unless well provisioned (J. A. Allen, 

Though explorers agree that no muskoxen are now known to 
occur on Melville Peninsula or on Baffin Island, there is some 
evidence that they may formerly have been found there but 
have been exterminated by the Eskimos. "The Canadian 
commission report (1922, p. 13) notes a tradition of a Muskox 
once killed on Baffin island . . . ," while on Melville 
Peninsula Freuchen states that it has so recently been exter- 
minated there that the Eskimos "still knew the names of men 
who have hunted it" (Hone, 1934). Apparently there is 
nothing to indicate that muskoxen ever occurred naturally on 
Southampton, although, as noted by Miss Hone, Freuchen 
"says that some teeth were found in the settlement at Kuk on 
the west side of York Bay, and* according to the Eskimos there 
was a skull in a house ruin on the south shore." 


Ovibos moschatus wardi Lydekker, Nature (London), vol. 63, p. 157, Dec. 13, 1900 

(Clavering Island, off East Greenland). 
SYNONYM: Ovibos moschatus melmllensis Kowarzik, Fauna Arctica, vol. 5, pp. 113-116, 

FIGS.: Allen, J. A. ( 1913, figs. 29, 31. 33, 34-38, 40-44, pis. 11-17 (exterior, live animals, 

skull, teeth). 

This northernmost race of the muskox has "conspicuous 
areas of white between and behind the horns, and face and 
sides of the head sometimes suffused with white to a greater or 
less extent in old males, in which much of the original white 
area is obliterated by the development of the horn bases; 
horns long and slender in proportion to their basal breadth, 
very light creamy white; tooth row relatively longer than in 
moschatus (max[illary] series in males, 140 mm.); basal length 
of skull in old males, 442 mm." "In general coloration wardi 
is not so dark as either moschatus or niphoecus; the saddle 


area and especially the horns are much lighter in color" (J. A. 
Allen, 1913). Tables of cranial measurements for this and 
other races of the muskox are given by the latter author. 

The range of the white-faced muskox occupies "a narrow 
coast belt of Greenland from about latitude 70 on the east 
side north as far as land extends, and thence southward along 
the west coast to about 81, and within historic times as far 
south as Westenholme Sound (latitude 78), where its further 
progress south appears to have been checked by impassable 
glaciers. It formerly occupied practically the whole Arctic 
Archipelago from Grant Land and Ellesmere Land westward 
to Prince Patrick Island and south to Lancaster Sound and 
Coronation Gulf. Thus it must have nearly met the range of 
niphoecus on the mainland west of the Gulf of Boothia, and 
the range of moschatus thence westward to Coronation Gulf 
and Dolphin and Union Strait . . . They have been ex- 
terminated from the greater part of Victoria Island, including 
Victoria Land, Wollaston Land, and part of Prince Albert 
Land. Hundreds have been killed on Melville Island, and 
thousands in northern Ellesmere Land, Grinnell Land, and 
Grant Land, mainly by explorers for the support of their dogs 
and men. They are found in* winter as well as in summer on 
the most northern known land, being in no sense migratory" 
(J. A. Allen, 1913). 

On Banks Island muskoxen were formerly numerous, for 
there was much good pasturage there. They were extermi- 
nated, however, by the Eskimos, who killed entire bands, 
utilizing little of the meat. The latest record given by Miss 
Hone (1934) is of a band killed by Eskimos in the spring of 
1911. At the present time the muskox is said to be extinct on 
this island. On Victoria Island there were formerly "plenty" 
in the little- visited northern part, and there are reports of them 
having been killed as recently as 1924 (Hone, 1934) ; the stock 
is, however, apparently small. Prince of Wales Island, at 
least within a few years, held a population of muskoxen that 
Anderson estimated at 1,500. They were present on Somerset 
Island in the middle of the last century but are now gone. 
Melville Island was the first of the Arctic islands where musk- 
oxen were found; Parry in 1819-20 reported them "in con- 
siderable numbers." Storkersen in 1916 estimated that there 
were about four thousand on this island. They occur also on 



Byam Martin Island and on Bathurst Island, where Anderson 
estimates there are about 1,500. The numbers on Devon 
Island seem fewer according to recent reports. On Axel 
Heiberg they are plentiful, although, according to Anderson, 
they are found only on the east side and may number in all 
about 1,000. On Ellesmere Island they were common along 
the east coast up till the eighties, but since then they have 
seldom been seen there, although numbers occur in the western 
areas and may in these less-visited parts be still numerous. 

On the short stretch of coast opposite Ellesmere Island in 
west Greenland muskoxen formerly occurred, as far south as 
Cape Alexander. South of this point the obstructing glaciers 
prevent further extension. A few animals were to be found 
northward of these points till about 1850, when the last living 
muskoxen in the region were reported seen by Eskimo near Cape 
George Russell. Macmillan (1918) mentions numerous skulls 
to be found from Etah north to Humboldt Glacier, but the ani- 
mal no longer occurs there and is believed to have become ex- 
tinct about 1860. Various expeditions have reported muskoxen 
in some numbers at various localities on the north coast of 
Greenland and Miss Hone (1$34) has lately assembled many 

White-faced inuskox (Ovibos moschatus wardi) 


notes on their occurrence in bands large or small along the east 
coast as far as the region of Scoresby Sound. Anderson, a 
decade ago, believed there were approximately 1,500 head in 
East Greenland, but others believe this figure too low. Jennov 
places the numbers between 6,000 and 10,000. There is some 
evidence that they are more abundant than they were when 
this region was inhabited by Eskimo. Their distribution may 
be sporadic, depending on the depth of winter snows and on 
whether the summer has been warm enough to produce pasture. 

There has been some controversy as to whether muskoxen 
are in need of protection in Greenland. While the inhospitable 
nature of their habitat prevents much hunting by visitors, it 
is obvious that near the Eskimo settlements, as in northwestern 
Greenland, they will continue to dwindle, unless perhaps, as 
has been hmted, these Eskimo themselves may be decreasing. 
After four years in East Greenland, Pedersen is of the opinion 
that protection is more than ever needed, partly on account of 
the excessive use made of the animals as food by trappers from 
Norway, and partly on account of the trade in young animals 
for zoological gardens, for in order to secure the calves, the 
herd of adults must first be shot. Although regulations have 
been proposed for limiting the number of animals that may be 
killed as well as prohibiting hunting for sport or for the capture 
of calves for commercial purposes, the latest reports at hand 
do not indicate great progress in this direction. 

An interesting experiment was undertaken a few years ago 
in introducing this muskox into Alaska, in order if possible to 
reestablish the animal over areas formerly occupied by it. 
In 1930, on the suggestion of the legislature of Alaska, the 
United States Congress appropriated $40,000 for obtaining the 
necessary stock. Thirty-four animals, both calves and adults, 
were captured in East Greenland, transported to Norway, 
thence to New York, and by rail to Seattle, whence they were 
finally brought by boat to Seward, Alaska (Bell, 1931). This 
herd has prospered, so that in 1936 Dr. Bell wrote to Dr. 
Harper in response to inquiry that the animals were doing 
well, had reached breeding age, and already had produced two 
crops of calves, five the first year and six in the second year. 
In 1935 two pairs of animals were transferred to Nunivak 
Island as an experiment and have thrived there, since food is 
plentiful. The herd was later placed on Nunivak National 


Wildlife Refuge. At last accounts the herd was in excellent 
condition and had increased to 90 animals (Dufresne, 1942). 


Although evidence is accumulating that at least one large 
species of extinct bison was contemporaneous with early man 
in North America, only one species was found on the continent 
at the time of its discovery by Europeans (see Meserve and 
Barbour, 1932). This species is closely related to the Old 
World bison, Bison bonasus, but differs in numerous details, 
as in the larger chest, smaller pelvus and shorter tail. In 
early days the "buffalo," as it is almost universally called, was 
found in great numbers over a vast range in this continent, 
but with the westward expansion of settlements it became an 
object of exploitation on a tremendous scale, so that literally 
millions were killed, and it was wiped out in the East and later 
over much of its western range. The story of this decimation 
has been many times told, but more especially by J. A. Allen 
(1876a, 1876b, 1877), Hornaday (1889), and recently by 
Garretson (1938). Although .attempts have been made to 
distinguish several local races, the characters of these are for 
the most part imperfectly known, and the respective ranges 
undefined. The northern form, or wood bison, however, 
constitutes a fairly well marked race. 


[Bos] bison Linnaeus, Systema Naturae, ed. 10, vol. 1, p. 72, 1758 (Mexico). 

Bos americanus Gmelin, Linnaeus's Systema Naturae, ed. 13, vol. 1, pt. 1, p. 294, 

Bison bison hanningtoni FIGGINS, Proc. Colorado Mus. Nat. Hist., vol. 12, no. 4, p. 
30, pis. 8, 9, Dec. 5, 1933 ("Head of Rock Creek, northeast South Park, Park 
County, Colorado"). Doubtfully distinct. 

Bison bison septentrionalis Figgins, op. cit., p. 28, pi. 7, Dec. 5, 1933 ("Six miles north- 
east of Palmer, Nebraska"). Doubtfully distinct. 

FIGS.: Allen, J. A., 1876a, pis. 5, 6, 9, 10, 12, fig. 1-6 (skulls and teeth); Garretson, 
1938, pis. facing title page and p. 5. 



Bison americanus pennsylvanicus H. W. Shoemaker, A Pennsylvania Bison Hunt 
(Middleburg, Pa.), p. 9, 1915 ("Pennsylvania"). 


Since the first accounts of the American bison were those 
brought back by the Spanish explorers of northeastern Mexico, 
this is taken as the type locality of the so-called Plains bison, 
the range of which is believed to have covered much of interior 
North America from the tableland of Mexico and the grass- 
lands of the West to the eastern Alleghenies, perhaps even 
reaching the coast in the southeastern States. In a recent 
history of the bison in Pennsylvania, Shoemaker has named the 
animal ranging "between the east and west slopes of the 
Alleghenies, migrating between the Great Lakes and the valleys 
of Southern Pennsylvania, Maryland and Virginia, to Georgia, " 
as a distinct eastern race, pennsylvanicus, but unfortunately 
the description is not based on a comparison of specimens, but 
upon local tradition that the bison of this region was larger 
than the Plains animal, and "very dark, many of the old bulls 
being coal black, with grizzly white hairs around the nose and 
eyes"; the hump "was notable by its absence" (which seems 
strange), while the legs were "long" without the contrast 
between the height of the fore and hind quarters seen in more 
western animals. It is difficult to know what value to give 
such an account, but the probability that these eastern bison 
were somewhat different from those of the Mexican tableland 
warrants the tentative recognition of the name. The charac- 
ters claimed for the bison of Colorado and of Nebraska, named 
B. b. hanningtoni and B. b. septentrionalis, respectively, seem 
more likely to be merely individual variations in tooth struc- 
ture, so that I have for the present considered these names as 
synonyms of B. bison bison. 

An adult male Plains bison stands about 5% to 6 feet at the 
highest point of the shoulder, but only about 4% feet at the 
hip, so that the hind quarters are proportionately small and 
the back is sloping. The females are somewhat smaller than 
males. J. A. Allen (1876a) gives the following measurements: 
Muzzle to insertion of tail, male, about 9 feet (2.75 m.) ; female, 
about 6.5 feet (2 m.). The horns are short, thick at the base, 
curving outward and upward, then somewhat inward. In the 
female they are slenderer than in the male. "In winter the 
head, neck, legs, tail, and whole under parts, are blackish- 
brown; the upper surface of the body lighter. The color above 
becomes gradually lighter towards spring; the new short hair 
in autumn is soft dark umber or liver-brown. In very old 


individuals the long wooly hair over the shoulders bleaches to 
a light yellowish-brown . . . The chin and throat are also 
covered with long hair, which under the chin forms an immense 
beard, eight or ten inches to a foot or more in length. Thick 
masses of long hair also arise from the inner and posterior 
surfaces of the fore legs, where the hair often attains a length 
of six or eight inches. A strip of long hair also extends along 
the crest of the back nearly to the tail. The tail is covered 
with only short soft hair till near the tip, from which arises a 
tuft of coarse long hair twelve to eighteen inches in length" 
(J. A. Allen, 1876a). Rarely, black or melanistic individuals 
occur, and still more rarely an albino. Many cranial measure- 
ments are tabulated in the monograph of J. A. Allen (1876a, 

Much has been written on the history, distribution, decima- 
tion, and reestablishment of this species. In the eastern part 
of North America the bison (" pennsylvanicus") occurred as 
far east as the western parts of New York State, but in inter- 
glacial times it probably extended to New England, as proved 
by the discovery of a piece of the maxilla with characteristic 
milk premolars found in glacia*! till on Cape Cod (G. M. Allen, 
1920). From western New York southward small herds were 
found in former times in the mountains of Pennsylvania, 
West Virginia, and Tennessee, into the upper parts of North 
and South Carolina, following the valleys of the New, Holston, 
and French Broad Rivers. Probably the extreme southeastern 
limit was in Georgia, where in the southeastern part is a creek 
still known as Buffalo Creek. There seems to be no certainty, 
however, that it was found in the present limits of the State of 
Florida, although its occasional presence there in former 
times is not unlikely. It is believed that the eastward extension 
of the bison's range was taking place at the time of the dis- 
covery, aided in part by the clearing of forests by the Indians 
and in part by the attraction of salt licks as in the mountains 
of Pennsylvania and West Virginia. To the northward the 
animals reached the shores of Lake Erie in their annual mi- 
grations and thence ranged westward to Lake Winnipeg and 
in increasing numbers over the Great Plains, to the edge of 
the Great Basin, and north of that to the extreme northeastern 
part of California and southeastern Oregon. The bison of the 
latter State has recently been described as a distinct race 
(see below). 


In its southward extent, the bison seems to have reached 
northern Alabama, central Mississippi, and Louisiana but at- 
tained the Gulf coast only in extreme southern Texas and 
northeastern Mexico (see map in J. A. Allen, 1876a). 

There is but little contemporary record of buffalo in early 
days east of the Alleghenies, but what evidence remains indi- 
cates that they were locally common in western Pennsylvania, 
West Virginia, and the Carolinas, but were constantly perse- 
cuted by settlers, explorers, and to some extent by the Indians, 
for meat and hides, and even wantonly destroyed. In their 
spring migration northward and during the autumnal migra- 
tion southward over a considerable extent of country, they wore 
deep trails following the easiest gradients and natural passes, 
going to better feeding grounds. Many of these paths came to 
be the trails and routes used later by the settlers pushing west- 
ward from the coast. This emigration received impetus after 
the Revolution when "thousands of officers and men who had 
served in the war received their pay in land script" (Garretson, 
1938). Even before this, the settlers had found that numbers 
of bison were incompatible with their own safety, for on oc- 
casion the herds would eat and trample down their slender 
crops and on one occasion an e&rly settler on Toby and Licking 
Creeks (now Oil and Clarion Creeks), Pennsylvania, had his 
cabin demolished by a herd of these animals that persisted in 
rubbing their backs and sides against its timbers. In his first 
two seasons this man and his companions killed 600 or 700 
bison, the skins of which brought but 2 shillings apiece. Such 
continued slaughter together with the destruction of the natural 
food of the bison, through firing of the grass and canebrakes by 
the settlers, gradually reduced their numbers so that by the 
close of the eighteenth century the "buffalo" in Pennsylvania 
"had been reduced to one herd, numbering between three and 
four hundred animals which had sought refuge in the wilds of 
the Seven Mountains, where, surrounded on all sides by settle- 
ments, they survived for a short time by hiding on the most 
inaccessible parts of the mountains" (Garretson, 1938). Ac- 
cording to Garretson, the last buffalo migrations from the Ohio 
country into Pennsylvania had ceased prior to 1783, and the 
year 1795 marked the disappearance of the last herds in the 
northwestern parts of the State. In the very severe winter of 
1799-1800, what was probably the last herd in Pennsylvania 


was slaughtered when, huddled together in the deep snow in 
a great hollow known as the "Sink" in the White Mountains 
of Union County, they were rendered nearly helpless. In the 
following year a bull, a cow, and a calf were seen in the same 
county, and the bull was killed the following year; it was be- 
lieved to be the last wild buffalo to be shot in the State. The 
cow and calf were hunted but eventually disappeared, and 
with that the bison became extinct in Pennsylvania. The 
story of the buffalo in West Virginia is very similar, but they 
persisted a little longer. According to Garretson (1938), "the 
last buffalo killed in Kanawha County, West Virginia, was in 
1815, on the waters of the Little Sandy Creek of Elk River, 
about twelve miles from Charleston. It is also recorded that 
as late as 1 825 a buffalo cow and her calf were killed at Valley 
Head, near the source of Tygart's River, and these are be- 
lieved to be the last buffalo killed in the East. " Thus by 1825, 
with the rapid opening up of the Middle West, and the 
slaughter of these animals by the settlers, the buffalo had be- 
come practically extinct east of the Mississippi, although a few 
stragglers were killed in Wisconsin as late as 1833 (Cory, 1912). 

With the transcontinental Purveys followed by the trans- 
continental railways in the two or three decades after 1830, 
added to the expansion of the fur trade in the West, the slaugh- 
ter of the vast herds beyond the Mississippi began in earnest. 
"As early as 1840 the American Fur Company sent 67,000 
robes to St. Louis and in 1848, 110,000 robes were received, 
also 25,000 tongues." The skins of cows only were used for 
robes, for those of bulls were too heavy. Hayden, of the U. S. 
Geological Survey, who visited "the upper Missouri country 
in 1850-1860, estimated the number of buffalo killed every 
year to be about 250,000 of which 100,000 were for robes" 
(Garretson). The railroad companies, advancing their lines 
across the western country, employed hunters to keep their 
camps supplied with buffalo meat, and the hunters likewise 
shipped back incredible quantities of tongues and hides for 
sale in the East. 

After the Civil War the army posts on the Plains increased 
in number, and hunters on contract supplied the camps with 
meat. In the seventies bison were recklessly slaughtered by 
the hundreds of thousands, and for every one utilized Hornaday 
(1889) believes two were wasted. Only the best hides and the 


choicest parts of the meat were saved. South of the main 
transcontinental railroad lines, what came to be known as the 
"southern herd" centered in Kansas, Oklahoma, and Texas. 
In December 1877 and January 1878 the "last great slaughter" 
of this group took place when more than 100,000 hides were 
taken by an army of hunters. By the end of the latter year, 
this herd had been practically wiped out. A few remained in 
the southwestern corner of Kansas until 1879, when the last 
one in that State was killed west of Dodge City. In Texas, 
scattered herds survived later, but it is believed that a little 
group of four killed near Buffalo Springs were the last survivors 
of the southern herd. It was said that at this period, when 
war was still being carried on against some of the tribes of 
Plains Indians, the extermination of the buffalo on which they 
chiefly depended for food was a factor also in the destruction of 
the Indians. This period of the seventies marked the turn of 
the tide for the buffalo, and their numbers rapidly decreased 
both in the southern and the northern part of their range. 
Garretson (1938) has published some interesting pictures of the 
vast quantities of bones bleaching on the sites of slaughter, 
and for years after persons made a living by collecting these 
bones in cartloads and shipping them east to be made into 

In the late eighties naturalists and others interested in wild 
life began to realize that the bison was approaching extermi- 
nation. Apart from a few privately owned herds and a herd 
closely protected by the Government in Yellowstone National 
Park very few remained in a wild condition in the United 
States. It was at this time that Dr. Hornaday (1889) made 
his stirring protest against the extermination of the species. 
By 1900 there were but two herds of bison remaining in a wild 
state in North America: the small one in Yellowstone Park 
and the wood bison in Athabaska. Notwithstanding supposed 
protection of the former, there was for a time considerable 
poaching, and it was not until May 1894 that an effective 
law for the preservation of the bison was passed by Congress. 
In 1902 Congress appropriated $15,000 for the purchase of 
buffalo from privately owned herds to build up the small stock 
then remaining in Yellowstone Park. Soon after, through the 
efforts of the New York Zoological Society, the Wichita herd 
was established, following the plan that numerous small herds 


in different areas would be the wisest way to build up a stock 
of these animals. In 1905 the American Bison Society was 
founded, starting with a group of 16 public-spirited citizens 
who saw the need for steps being taken if the bison was not to 
become endangered. The story of its work in establishing at 
various suitable places small herds as nuclei, and thus building 
up a sufficient population so that the species would no longer 
be in danger, has been graphically told by Garretson (1938). 
The effectiveness of these efforts is evident from the table 
published by the latter, showing that in 1889 the total popu- 
lation of bison in existence was placed at 1,091 but by 1933 
had been built up to well over 21,000, of which the greater part 
(17,043) were in Canada. There are now some 121 small herds 
in 41 States totaling 4,404 animals. Thus the species seems 
no longer in danger of vanishing from the face of the earth, 
but there is good assurance that in suitable places and under 
supervision sufficient numbers may be maintained under fence 
to make certain of its preservation. 

From a practical point of view, bison were formerly the main 
source of meat for many of the Plains Indians, as well as for 
the early settlers in certain regions. Their later exploitation 
by hunters was no doubt largely unnecessary, yet one should 
not lay too great a blame upon the shoulders of those who, in 
the presence of seemingly unbounded stores of animals, made 
use of them for their own gain. For as yet the needs of "con- 
servation" had not been made obvious. 

Experiments have from time to time been made in crossing 
the bison with domestic cattle, but although the resulting 
"cattaloes" possess certain desirable qualities, it does not 
appear that the hybrids have proved popular with agricul- 

One of the largest of the protected herds has been that 
maintained at Buffalo National Park at Wainwright, Alberta. 
Its numbers were lately estimated at about 3,000 head, and in 
addition on the reserve were nearly half as many wapiti, as 
well as numbers of deer and moose, together with some im- 
ported yak from Tibet. Now, 1940, as a late development of 
the war in Europe, comes word that this national park must 
be utilized for other purposes, which according to rumor, 
include the training of aviators, and the animals must be 
cleared from the area. According to a quotation in Science 


(vol. 91, pp. 12-13, Jan. 5, 1940), the bison are to be killed and 
the meat and hides sold. 



Bison bison oregonus Bailey, Proc. Biol. Soc. Washington, vol. 45, p. 48, Apr. 2, 1932 
("Dry bed of Malheur Lake, Oregon"). 

In the extreme western extension of its range, the local bison 
was slightly different from the typical Plains bison of south- 
western Texas, being "slightly larger, with relatively longer 
and straighter and less abruptly tapering horn cores, indicating 
wider and straighter horns of a somewhat larger animal. The 
rostrum or arch formed by the upper premaxillary bones is 
slightly longer and relatively narrower than in southern 
specimens; interpterygoid fossa wider and larger; auditory in- 
flations smaller than in typical Texas skulls; molars larger." 
The external characters are unknown. 

This race of the bison is now extinct, but a century or more 
ago it was found over southern Idaho and extreme northern 
Nevada to southeastern Oregon and northeastern California, 
areas that it reached from the more eastern plains through 
broad flat valleys such as those of the Quinn River and Alvord 
Valley. Vernon Bailey (1936) has gathered together what is 
known of its history. From the lips of older Indians of the 
region, he heard that these animals formerly abounded about 
the Cow Creek Lakes country in the early part of the last 
century, and Townsend in 1834 found them across southern 
Idaho to the Malad River. They seem to have vanished from 
Oregon before the arrival of the white man, and disappeared 
from Idaho soon after the coming of the early explorers. This 
disappearance Bailey attributes largely to the fact that by the 
beginning of the last century the Indians of this region were 
then well supplied with horses and thus were able to make 
more serious inroads into the ranks of the bison, finally ex- 
terminating them. Merriam (1926) was able to obtain definite 
accounts of its former presence in northeastern California 
where old Indians of several tribes said that their fathers had 
killed these animals, as in Pine Creek Valley west of Eagle 
Lake. The Indians believed that they came in small bands 
from still farther north, indicating that here as elsewhere the 


bison made seasonal migrations in search of better feeding 
grounds. Bailey learned from an old chief of the Piute Indians 
that in the Malheur Lake region of Oregon bison were found 
all over the district at a time probably about the middle of the 
last century. They went into the mountains in summer and 
came down into the valleys in winter. During dry years, 
Bailey writes, the waters of Malheur Lake became very low, 
and numerous bison skeletons were laid bare, so that a series 
of specimens was collected on which the new race was described. 
These remains were evidently of animals that "had bogged 
down in search of water at some dry period long ago when the 
water had receded; or else, in attempting to cross the lake on 
the ice in winter, or to get out to open water, they had broken 
through and drowned." There is therefore "no question that 
only a few generations back buffalo covered in considerable 
numbers many of the large valleys of southeastern Oregon, 
and that they disappeared after the introduction of horses 
among the Indians and before many firearms were obtained. " 
According to an old chief, Yakima Jim, said in 1916 to be 110 
years old, the "last of the buffalo were killed during a hard 
winter when the snow was so 1 deep that they could not get 
grass and a good many tumbled over the high bluffs on the 
Owyhee River" (Bailey, 1936). Grinnell (1933) comments 
that although the bison evidently was sporadically distributed 
in the northeastern part of California in Modoc and Lassen 
Counties a century ago, there seems to be no good evidence 
that it ever reached the Sierra Nevadas. It may be that 
future comparisons will show, if material becomes available, 
where this race intergraded with the typical Plains bison. 


Bison bison athabascae Rhoads, Proc. Acad. Nat. Sci. Philadelphia, for 1897, p. 498, 
Jan. 18, 1898 ("Within 50 miles southwest of Fort Resolution, Mackenzie, 

FIGS.: Garretson, 1938, pi. opposite p. 12 (head and refuge map). 

The wood bison is a distinct northern race characterized by 
darker color, more dense and silky coat, somewhat larger size, 
and more particularly by its longer and slenderer horns and 
horn cores, as compared with the Plains bison to the south. The 
longer and more incurving horns give the animals a distinctly 


different look from their shorter- and stouter-horned relatives, 
which is obvious even in photographs of the living animals. 

The range of this race is (or was) north of the United States, 
in northwestern Canada, east of the Rocky Mountains to 
about the 95th meridian, and between latitudes 55 and 63 
N., approximately. According to Dr. R. M. Anderson (1937) 
there "is evidence that the wood bison formerly was found 
some little distance northwest of Great Slave Lake as far as 
Horn Mountains and Liard River, and for an indeterminate 
distance up the Peace River valley, and southward, but it is 
now restricted to the Wood Buffalo Park area, on both sides of 
the 60th parallel." Preble (1908) has assembled a large 
amount of data gleaned from the accounts of older travelers 
and explorers concerning the bison of this northern region. 
It appears that the animals "formerly ranged over immense 
areas north to Great Slave Lake and Liard River," where in 
1772, Samuel Hearne found them "very plentiful." At the 
beginning of the last century Mackenzie recorded "numerous 
herds" on the plains near Vermilion Falls and in the Peace 
River region, which was probably close to its southern limit. 
There is some evidence that by 1828 it was already diminishing 
in numbers here, and according to Cowan (1939) the last 
record of its presence in the district is furnished by Dawson, 
who, in his report of his expedition down the Peace River in 
1879 and 1880, mentions the many scattered bones and the 
saucershaped wallows of the buffalo, adding that "the Beaver 
Indians report having seen in the summer of 1879, six woodland 
buffaloes of which they killed one in the vicinity of Pouce 
Coupe." Probably they did not long persist in this part of 
British Columbia after that time. Their destruction in this 
region may not have been wholly due to man, for in 1877 
J. A. Allen published a letter giving observations of two young 
men who had reached the Yukon through British America in 
which they state that in making a portage from Peace River to 
Hay River, they saw "thousands of buffalo skulls and old 
trails in some instances two or three feet deep, leading east and 
west. They wintered on Hay River, near its entrance into 
Great Slave Lake, and there found the buffalo still common, 
occupying a restricted territory along the southern border of 
the lake. This was in 1871. They made inquiry concerning 
the large number of skulls seen by them on the portage, and 


learned that about fifty years before snow fell to the estimated 
depth of fourteen feet, and so enveloped the animals that they 
perished by thousands." Such wholesale decimation must 
have been rather rare, yet nevertheless shows what an unusual 
season might so. Another danger to which these animals 
sometimes must have succumbed is illustrated by an account 
given to Preble (1908) of a herd of about 50 bison that were all 
drowned in attempting to cross a small lake too early in the 
season, before the ice would bear their weight. This was near 
where the Petite Riviere Bouffante enters the Athabaska. 

Macfarlane, who resided at Fort Chipewyan during the 
years 1870-85, said that the fort hunters seldom failed to kill 
a few bison each winter, mainly on the north side of lower 
Peace River (Preble, 1908). Writing in 1888, William Ogilvie 
(quoted by Preble, 1908) reported that the wood bison was 
"nearly a thing of the past ... In the winter of 1887-88, 
on the headwaters of Hay River, which flows into Great 
Slave Lake, and west of Battle River, a tributary of the Peace, 
the Indians saw three bands containing 17, 10, and 4, respec- 
tively; they killed 5 ... The same winter three bands 
were seen between Salt River and Peace Point, on Peace 
River, numbering 50, 25, and about 25, respectively . . . 
During the winter of 1886-87, between the north end of Birch 
and the south end of Thickwood Mountains, distant about 
one day or 30 miles from Fort McMurray, on Athabasca River, 
one band of about 13 was seen. Since then 5 of this band have 
been killed. Below Red River, a tributary of the Athabasca, 
and between Birch Mountains and Athabasca River, and 
ranging down to Poplar Point, on the Athabasca, another band 
said to contain about 20 was seen. Altogether we have only 
about 180 head of wood buffalo in this vast extent of territory. " 
In 1891 the same explorer estimated the wood-buffalo popula- 
tion as not exceeding 300 in all. He speaks of the Indian 
method of killing them by stampeding the animals into a bog 
where they soon become mired and are easily slaughtered. 

In February, 1894, Caspar Whitney "estimated the total 
number then living as about 150" (Preble, 1908). That the 
numbers were indeed low is indicated by the fact that Preble 
on his journey through their region in 1903 and 1904 was able 
to obtain but few reports of their presence. In the winter of 
1902-3 he found two small bands, aggregating 24 individuals, 


in the thickly forested region about 125 miles southwest of 
Fort Smith. There were apparently no young animals with 
this herd, and it was believed that wolves had accounted for 
any there may have been. In 1907 Inspector A. M. Jarvis and 
Ernest Thompson Seton in much the same region found two 
herds of 13 and 20. 

Fortunately, in December, 1922, the Canadian Government 
set aside as Wood Buffalo Park an area of 10,500 square miles, 
which included the entire habitat of the known herds, as a 
sanctuary. This was subsequently enlarged to 17,300 square 
miles and placed under the charge of a dozen skilled rangers. 
By 1929 the number of animals had increased to about 1,500, 
or nearly three times the number estimated to inhabit the 
region in 1914 (Harper, 1932). This satisfactory increase has 
continued, but an undesirable element has been that since 1925 
there have been transported to Wood Buffalo Park the increase 
of the Wainwright herd of Plains buffalo from central Alberta 
in at least four annual shipments totaling 6,673 animals. On 
account of the close relationship of the two races, it seems in- 
evitable that they must interbreed and that the remnant of the 
wood buffalo will eventually be represented by a mongrel 
stock. From a zoological point of view this result will be 
highly undesirable, but on the other hand may, through the 


Wood bison (Bison bison athabascae) 


increase of the herd, tend to add to the living resources of the 
human population in the region. The combined herds have so 
increased that in 1934 the total number of buffalo in the park 
was estimated at about 8,500 (Anderson, 1937, p. 103). 

Outside and to the northwest of the park there is said to be 
a small band of 20 to 30 buffalo in a place known locally as the 
Buffalo Mountains on the South Nahanni River, and it is not 
impossible that these may help to perpetuate the woodland 
stock, since its requirements are slightly different from those 
of the Plains bison (notes of Dr. Harper from manuscript 
report of H. M. Snyder) . Thus, while the wood bison may be 
considered as no longer in danger of extermination by man, it 
remains to be seen whether its type will disappear through 
interbreeding with the imported Plains stock. 


Of all American game animals, probably the bighorn offers 
the greatest thrill to the sportsman. A dweller in open moun- 
tainous country, keenly alert to the least danger, with excep- 
tional eyesight and hearing, it is the most difficult to approach 
of all American mammals, while its magnificent horns make a 
fine trophy for the successful hunter. The American members 
of this genus appear to have come in from Asia at a time when 
it was possible to cross from the northeastern extremity of 
that continent to Alaska, probably during the Pleistocene 
period. Thence they have spread southward through the 
western mountain chains to northern Mexico and eastward to 
the Dakotas. At the present time zoologists recognize two 
American species, the white, or DalPs, sheep in the northwest 
and the mountain sheep, or bighorn, from British Columbia 
south. Several races of each have been named at various 
times, but only recently has a comprehensive study of the 
group been made (Cowan, 1940), the results of which have 
been followed here. 



Ovis montana dalli Nelson, Proc. U. S. Nat. Mus., vol. 7, p. 13, June 3, 1884 (mountains 
west of Fort Reliance, Alaska, on the divide between Tanana and Yukon Rivers). 

FIGS.: Hornaday, 1901, figs, on pp. 83, 86, and pis. facing pp. 86, 92, 94; Nelson, 1916, 
p. 450, lower fig. (col.); Sheldon, 1930, pis. facing pp. 78, 79 (photographs). 


This is slightly smaller than the other North American sheep 
and with its races is considered to represent a species different 
from the more southern 0. canadensis and its races. In general 
appearance it is similar, but the horns are slenderer and the 
coat is white at all seasons, with sometimes a few scattered 
black hairs, especially near the base of the tail, where at times 
they may form a distinct black line. As a species distinct from 
0. canadensis, Ovis dalli, the white sheep, is smaller, with horns 
of greater length but of less basal circumference, "with the 
anterior surfaces strongly rugose, the orbital angle prominent, 
and often developed into a pronounced ridge overhanging the 
orbital surface of the horn" (Cowan, 1940). Height at shoul- 
ders, 39 inches; total length, 58.5 inches; tail, 4 inches. Record 
length of horn, 47.5 inches. Weight of rams about 200 pounds 
(Seton) ; of ewes 50 or more pounds less. 

Originally white sheep were found in the mountain ranges 
of Alaska "from those bordering the Arctic coast south 
through the interior to the cliffs on Kenai Peninsula, but are 
now scarce or gone from some mountains. " " Coming within a 
few miles of the Arctic coast south of Herschel Island," they 
follow the Mackenzie Mountains as far south as Nahanni 
River, north of the Liard River. Probably at one time their 
range was practically continuous over this area. Hornaday 
(1901) mentions the presence of this sheep in the mountains 
east of Nome, Alaska; it was not uncommon in the region about 
Cook Inlet, and extended as far eastward as about fifty miles 
from the Mackenzie delta. Southward it extends along the 
Rocky Mountains to about latitude 60 N. At the beginning 
of this century it was regarded by A. J. Stone as common 
throughout most of its range, but "not nearly so abundant as 
formerly. Where hundreds roamed eight years previously, we 
saw but sixty -four. " Sheldon (1930) found them still abun- 
dant in the Mount McKinley region, and Dr. R. M. Anderson 
(1939b) writes that it is "holding its range in most parts of 
Yukon Territory and limited numbers are found along the 
eastern slopes of the Mackenzie Mountains in Northwest Ter- 
ritories from the Arctic coast south nearly to Liard River." 

Thus, although on the borders of its distributional area it 
has undoubtedly been reduced in numbers, it is still common 
on the main mountain ranges of Alaska and Yukon. Various 
excellent accounts are in print of its habits and hunting, no- 


tably by A. J. Stone (1900), and in the Mount McKinley region 
by Charles Sheldon. The natural enemies are chiefly the wolf, 
less often the lynx (Sheldon), and Lee emphasizes particularly 
the depredations of grizzly bears, which ambush them and suc- 
ceed often in killing them. During winter, when snowstorms 
cover their upland pasturage, they may lie up for days among 
sheltering rocks without venturing forth to feed. Their range 
is usually above timberline on the open slopes where they can 
readily detect the approach of danger from below. Two 
young are usually produced at a birth, some time between 
May 1 and August 1. "The males and females are hardly 
ever found together during the summer months. The males 
generally inhabit the roughest and highest peaks, while the 
ewes and lambs keep along the high plateaus" (Lee, in Horna- 
day, 1901). 

Except for food and trophies these sheep are little hunted, 
for the coat even in winter is so brittle that it is not in demand. 
Also since their area of distribution is mainly outside tracts 
used for grazing domestic sheep, the bighorns of this type are 
relatively safe from infection with scabies, which has proved 
so destructive to their more southern relatives in the United 
States. Nevertheless, there is always danger of introducing 
new diseases where domestic animals are brought in. Thus 
Maj. Allan Brooks (1923) writes that "lumpy -jaw was very 
prevalent in the range of Ovis dalli and its subspecies (north 
side of Stikine River) up to 1908, but is now apparently 
stamped out. " 



Ovis dalli kenaiensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 16, p. 145, 1902 
("Head of Sheep Creek, Kenai Peninsula,' 5 Alaska). 

This race, confined to the Kenai Peninsula of Alaska, is 
similar externally in its white pelage to typical 0. d. dalli. It 
is distinguished, however, on the basis of cranial characters: 
basilar length less; molar series significantly shorter (70-72 
mm.); basioccipital narrower; angle between basioccipital axis 
and palatal axis apparently greater (Cowan, 1940). 

The sheep on the Kenai Peninsula constitute a population 
now quite isolated from their mainland relatives and charac- 
terized by slight though significant differences in certain cranial 


proportions, so that Cowan believes they deserve recognition 
as a distinct geographical race. This fact, he comments, 
"probably indicates a very long period of residence there and 
an effective degree of isolation by the lower ground between 
the peninsula and the mountains of the contiguous mainland. " 
Very little seems to be recorded as to the present status of the 
sheep on the Kenai Peninsula. W. T. Hornaday (1901) repro- 
duces an account of hunting them in the Kenai Mountains by 
Harry Lee, who mentions their great reduction in numbers at 
the hands of prospectors and miners in the region forty years 
ago. Previous records for this race from the base of the Alaska 
Peninsula are regarded by Cowan (1940) as referring to typical 

Undoubtedly on account of its limited range it should be 
protected if it is to be saved from eventual extermination. 

Ovis DALLI STONEI J. A. Allen 

Ovis stonei J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 9, p. Ill, Apr. 8, 1897 (head- 
waters of the Stikine River, British Columbia, altitude 6,500 feet). 

SYNONYMS: Ovis liardensis Lydekker, Wild Oxen, Sheep, and Goats, p. 215, fig. 41, 
1898 (Liard River, Canada); Ovisfannini Hornaday, Fifth Ann. Rept. New York 
Zool. Soc., p. 78, June 1, 1901 (Dawson City, Northwest Territories); Ovis cowani 
Rothschild, Proc. Zool. Soc. London, 1907, p. 238 (mountain chain near Mount 
Logan, British Columbia); Ovis canadensis niger Millais, The Gun at Home and 
Abroad, vol. 4, p. 324, 1915 (Skeena River, mountains at the head of, British 

FIGS.: Hornaday, 1901, col. pi. opposite p. 78 (0. 'fannini'}, pis. opposite pp. 92, 98, 
100; Nelson, 1916, p. 450, upper fig. (col.). 

South of the range of the typical form of the white sheep 
occurs a dark-colored race, with intermediate conditions shown 
by specimens from the mountains west of the Yukon between 
Selkirk and Forty-mile River (the 0. "fannini") while vari- 
ations from the mountain ranges in northern British Columbia, 
though given distinct names, are now regarded as all referable 
to Stone's sheep. 

Of the same general type as Ball's sheep, with slender, out- 
curving horns of triangular cross section, Stone's sheep de- 
velops a partly pigmented coat, varying from one with a gray 
saddle to a more gray or dark-brown condition, with a black 
dorsal stripe from the back of the head to the tip of the tail. 


Face and mid ventral area white. Height at shoulders in males 
said to be as in dalli, in an adult female 32 inches. Record 
length of horn on front curve, 51% inches, with a tip-to-tip 
spread of 31 inches (Muskwa River, British Columbia) (Ely 
et al., 1939). 

The range of Stone's sheep approaches that of the Rocky 
Mountain bighorn south of the upper Peace River, British 
Columbia. It is still common about the "upper Stikine River 
and its tributaries; thence it extends easterly to Laurier Pass 
in the Rocky Mountains, north of Peace River, and south 
perhaps to Babine Lake. Unfortunately it seems to have be- 
come extinct in the southern border of its range" (Nelson, 
1916). Cowan (1939), however, mentions the occurrence of a 
single young ram on the shore of Charlie Lake during the sum- 
mer of 1937, in the Peace River district. It is said to be espe- 
cially common east of Dease Lake. The intermediate form, the 
so-called Fannin's sheep, or saddleback sheep, occurs in the 
same groups with white animals, and there is a condition in 
which the white sheep show in some individuals a decidedly 
black tail, especially in the Nahanni Mountains. All such, 
however, are best regarded as connecting steps in the passage 
from the two extremes of coloration, rather than as distinct 

Nelson (1916) writes that this sheep occurs in "one of the 
most notable big-game fields of the continent. Its home above 
timberline is shared with the mountain goat and in the lower 
open slopes with the caribou, while within the adjacent forests 
wander the moose and two or more species of bear. Owing to 
its frequenting remote and sparsely inhabited country it con- 
tinues to exist in large numbers; but if its range becomes more 
accessible, only the most stringent protection can save this 
splendid animal from the extermination already accomplished 
on the southern border of its range. " 



Ovis canadensis Shaw, Naturalist's Miscellany, vol. 15, text to pi. 610, 1804 (mountains 

on Bow River, near Calgary, Alberta, Canada). 
SYNONYMS: Ovis cervina Desmarest, Nouv. Diet. Hist. Nat., vol. 24, pp. 5, C, 

1804 (Alberta); Ovis montana G. Cuvier, Regne Animal, vol. 1, p. 267, 1817 ( = 

Dec., 1816) (Alberta). 


FIGS.: Hornaday, 1901, figs, on pis. opposite pp. 102, 106, 110; Nelson, 1916, col. fig. 
on p. 447; Chapman, W. and L., 1937, pis. following p. 86. 

The bighorn group has the facial part of the skull an inch 
or two longer than in the white sheep and its race, Stone's 
sheep; the body is larger and the horns stouter, less spreading. 
The pelage is coarser and in winter somewhat shorter, crinkled. 
In color the Rocky Mountain bighorn is a uniform gray -brown, 
varying much in individuals, even in the same group, and on 
the front of the hind legs becoming darker. The end of the 
muzzle, a conspicuous patch on the rump, and the backs of the 
limbs, white to creamy. "This dark-colored abdomen seems . . . 
to render this mountain animal less conspicuous from below" 
(Hornaday, 1901). The massive horns of the male usually 
make one complete turn, sometimes more, in a rather close 
spiral. A record head with about one and one-half turns is 
figured by Hornaday (1901, opp. p. 106). In females the horns 
are shorter, more nearly erect. There is no groove under 
either edge such as is seen in horns of male white sheep. 
Height at shoulder, 40 inches (1,018 mm.); length from nose to 
base of tail, 58 inches (1,476 mm.); tail, 3 inches (77 mm.). 
In the record head, the greatest length of horn was 52.5 inches 
on outer curve; the longest listed by Ely et al. (1939) is 49.5 
inches. Cowan (1940) states that the short ear is additionally 
diagnostic of the typical race. 

The range of this sheep is from the mountains of western 
Alberta and southeastern British Columbia to about 120 miles 
south of Peace River, thence southward in the Rocky Moun- 
tains to Colorado, Utah, and northern New Mexico. Over 
parts of this range, owing to persecution, and to some extent 
to other causes, it has become scarce or even extirpated. 
"Their wariness, their strength and agility in climbing, and 
the rugged mountains which they inhabit combine to render 
them difficult to find and difficult to kill," while the impressive 
horns of an adult male form a spectacular trophy, so that 
more perhaps than any other game mammal of North America 
they have a strong appeal to sportsmen. According to Dr. 
Hornaday, the young, usually one or more rarely two, are 
born late in spring, between May 15 and June 15. The young 
lambs are sometimes the prey of eagles, and the adults are 
occasionally killed by wolves in winter. The same author 
states that although they have been exhibited in zoological 


gardens in eastern Unites States, they do not long survive 
change in altitude, humidity, and food. 

In the Rocky Mountains of western Alberta, Dr. R. M. 
Anderson writes (1939b) that this sheep is "still fairly common 
. . . and many good specimens are taken annually." 
Under present conditions and with adequate legal protection, 
the prospect for their survival seems excellent. In Canada 
ewes are protected at all times, and the rams during about 
two-thirds of the year, while in a national park created by the 
Canadian Government on the boundary adjacent to Glacier 
National Park in northwestern Montana, there is of course 
year-round protection. In the Glacier Park, Montana, Bailey 
(1918) reported that these sheep are abundant "on practically 
all the high, rugged ranges . . . especially on the rocky 
slopes above Two Medicine Lakes and around Chief and 
Gable Mountains. In summer they scatter out over the high 
and more inaccessible ridges above timberline . . . but 
during the winter they come down on the lower slopes and, 
especially in spring and early summer, are much in evidence." 

Bailey estimated the total sheep population in the park as 
about 2,000. Their chief ene*my here is the large mountain 
coyote, but outside the Park "the animals are not easily 
protected from poachers" who hunt for meat. In the adjacent 
State of Idaho, sheep were formerly plentiful and widely dis- 
tributed. In 1884 "thousands of sheep were in the Lost River 
area; Hornaday (1901) reports that in 1887 trappers encoun- 
tered 2,000 to 2,500 head on the Middle Fork of the Salmon 
River" (Davis, 1939); Merriam a few years later found them 
common in the Lemhi and Pahsimeroi Mountains and in 
smaller numbers in the Sawtooth Mountains. A 1939 estimate 
of the bighorn population in Idaho was but 2,450, of which 
the greater part were in national forests. The same source 
(U. S. Dept. Interior, Wildlife Leaflet BS-142) placed the 
bighorn population of Wyoming at 5,079, or nearly as great as 
the number in Montana and Idaho combined. These animals 
were said to be about equally divided between grazing areas 
and national forests, and hence largely in the western part of 
the State. The estimate given for Utah was 252, probably 
mostly in the Ashley National Forest. When about 20 years 
ago I accompanied Dr. Theodore Lyman to the Uinta Moun- 
tains in search of sheep, we could gather nothing but vague 


reports of a few animals said to have been seen within recent 
years. However, Cowan (1940) reports 150 head there at 
present. Most of the upper parts of the range are grazed by 
domestic sheep in summer. In Colorado, Merritt Cary in 1911 
reported that bighorns were to be found in small numbers on 
nearly all the high mountain ranges, especially in the northern 
parts of the State. Since 1885 they had been protected by law, 
but this apparently had not always been well enforced. With 
better protection and a developing public sentiment against 
killing them, an encouraging increase in numbers took place 
in the early years of this century. In 1902, Dall De Weese 
estimated that there were probably 200 in the State. By 1907, 
"sheep were reported on most of the mountain ranges of south- 
ern Colorado, and seemed to be on the increase." The 1939 
census credited the State with some 2,285 sheep, of which all 
but about 150 were in national forests. To the southward the 
typical Rocky Mountain bighorn extended formerly into the 
northern parts of New Mexico, but at the present day there 
are few if any left there. Bailey gives as its former limits of 
distribution the Sangre de Cristo Mountains as far south as 
the Truchas Peaks, Pecos Baldy, and Santa Fe Baldy on the 
east of the Rio Grande Valley, and probably through the San 
Juan Mountains to the Jemez on the west of the valley. Sheep 
disappeared from the lower and more accessible San Juan 
Mountains some time toward the close of the last century for 
in 1904 Bailey could obtain no recent record of them from 
local ranchmen, while from the Jemez Mountains still farther 
to the south they must have gone soon after 1880. Sheep were 
common in the Santa Fe region in 1873, and probably a few 
lingered among the mountains at the head of the Pecos till 
the early years of the present century. Bailey (1931) believes 
that given adequate protection bighorns will in course of time 
increase in Colorado and the overflow will repopulate northern 
New Mexico, following the mountain chains. The future of 
the species seems well assured, for summing the censuses of 
1939 for the five Rocky Mountain States gave an estimated 
total of nearly 11,000 animals, of which about half are in the 
State of Wyoming. 

Cary (1911) as well as Warren (1910) mentions the coyote 
as an occasional enemy, especially in winter when, in deep 
snows, it is sometimes impossible for the sheep to escape, 


whereas with a light crust the coyotes are able to give pursuit 
effectively. Another danger that has developed is the intro- 
duction of scab by the grazing of domestic sheep on the upper 
ranges in summer, in the States where bighorn occur. Warren 
was told of an instance in which 75 bighorns were found dead 
from this cause in the West Elk Mountains. More recently 
the bighorn in Glacier Park have in some instances shown the 
development of a fatal pneumonia, to the extent in one case 
that 15 out of a band of a hundred had died. The primary 
trouble seems due to a lungworm, which opens the way for 
secondary bacterial infection. Most of this seemed to occur 
among bighorns that in a certain locality were fed with hay, 
where local pasturage had been overgrazed. Other cases of 
young lambs dying of pneumonia due to infection by Pasteu- 
rella, without primary infection by the lungworms, were also 
found. No suggestions for eradicating this danger have so far 
been made. The lungworms probably pass one stage in a 
snail and then, if the small snails are eaten by the sheep while 
grazing, the parasite continues its development in the new 
host, and invades the lung tissue (see Marsh, 1938). * The late 
George Bird Grinnell (1928) wrote that "many years ago Col. 
Edward L. Munson expressed the belief that an epidemic of 
anthrax communicated to them from domestic sheep feeding 
on the plains below had exterminated the wild sheep of the 
Bear Paw Mountains in Montana." They were formerly 
common there, but in later years none has been found. Thus 
it seems likely that bighorn may be susceptible to various 
introduced diseases that are brought in by domestic sheep and 
against which they may need in some way to be guarded. 

As a preliminary to more intelligent management and pro- 
tection of this species, H. B. Mills (1937) has made an intensive 
study of the bighorn population of the Yellowstone National 
Park in the northwestern corner of Wyoming and the adjacent 
edge of Idaho. Here, notwithstanding Vernon Bailey's earlier 
estimate that the area might easily support ten times the 
present population, the number of bighorns has remained 
about 200 for the past 20 years or more and even dropped to 
about half that number as from 1927-33. Thus under protec- 
tion from both man and natural predators, the animals show 
a decrease. A careful census of the bighorns in the park in 
1934-35 gave a total of 240. They keep more or less in groups, 


each having its range, migrating in autumn to wintering 
areas and in spring to the extensive summer ranges. A proper 
winter range thus becomes essential to their well-being, but 
it appears that at the time when the report was made (1937), 
the winter range had "been so depleted by the large bands of 
wapiti that the bighorn were feeding on very short grass all 
winter . . . cropping grass so closely that plants were up- 
rooted and the roots and adhering soil were swallowed." In 
addition to this competition for food, "two diseases, scabies 
and lung worm infection, have caused reduction of the bighorn 
population in this region. " There is also a certain amount of 
infection from nematodes, which may be more or less normal. 
Mills believes that the conditions may be alleviated by re- 
storing the winter range to a better condition (involving some 
reduction in the wapiti herd as well as a possible acquisition of 
additional park territory) and maintaining a proper proportion 
of large predators that will eliminate unhealthy individuals. 
The study points the way to a better understanding of the 
requirements of the bighorn and of methods for maintaining a 
normal population. 


Ovis canadensis auduboni Merriam, Proc. Biol. Soc. Washington, vol. 14, p. 31, Apr. 5, 

1901 ("Upper Missouri," believed to be the Badlands of South Dakota). 
FIG.: Audubon and Bachman, 1846-54, vol. 2, pi. 73 (col). 

This is the most eastern race of the bighorn, but it is now un- 
happily extinct. It seems to have been but very slightly 
different from the typical Rocky Mountain bighorn, but the 
upper tooth row averaged longer, and the tooth rows were less 
nearly parallel. Audubon and Bachman (1846-54) described 
the color as "light grayish brown," the rump and under parts 
grayish white; weight of an adult male 344 pounds, and of a 
female 240 pounds. 

A century ago this sheep was abundant in the broken country 
or Badlands of the upper Missouri and Little Missouri in 
western Dakota. Bailey (1926) has summed up its history in 
North Dakota; its former range in that State "included all of 
the very rough Badlands country along and west of the Mis- 
souri River. " It was found in western South Dakota, western 


Nebraska, and to an undetermined distance westward probably 
into eastern Montana and Wyoming, where it must have inter- 
graded with the typical form of the Rocky Mountains. At 
the present time it is believed to be extinct over all this region. 
Audubon in ascending the Missouri River in 1843 first met with 
the bighorn in western North Dakota above the mouth of the 
Little Knife River, probably, as Bailey points out, at about 
the same place where Maximilian ten years before had found 
them. Probably this was very near the northeastern limit of 
distribution, for Bailey quotes a resident long acquainted with 
that region as stating that there "never were any mountain 
sheep near the Missouri at Cannon Ball, but that formerly 
the Indians went farther west to hunt them." Dr. George 
Bird Grinnell (1928) pointed out that in those days sheep were 
far less shy than later, when the white hunter with his rifle 
proved a deadly enemy; it was then common to find the 
bighorn grazing on open prairie near the high buttes to which 
they could escape if threatened with danger. The Platte 
River in southern Nebraska marked about the southeastward 
extent of the range. Grinnell recalls that in earlier days, 
about the middle of the last century, "on that stream certain 
isolated buttes and pinnacles were their favorite resorts, but 
when the country about these high points for example, 
Scott sbluff began to settle up, the sheep were cut off from the 
mountains and could neither escape to other refuges nor could 
their numbers be added to by others of their kind from the 

"In the early eighties Theodore Roosevelt . . . hunted 
mountain sheep in the Badlands along the Little Missouri," 
but their numbers were apparently not large. In 1888 three 
were killed near the present town of Oakdale, in the Killdeer 
Mountains, North Dakota. Three others were killed from a 
little band of five in the Badlands of the Little Missouri in 
1898. The last positive record for the bighorn in North 
Dakota is said by Bailey to be an old ram killed about 1905 on 
Magpie Creek, in the Killdeer Mountains. There are later 
reports, however, for the Badlands of South Dakota. Ap- 
parently the sheep held out longest in the Black Hills region, 
where in 1895 Dr. Walter Granger was told of the presence of 
a small herd in the vicinity of Harney Peak. "In the Bad 
Lands," he wrote (in J. A. Allen, 1895), "they are quite com- 



mon. Several were seen by our party, and their tracks could 
be seen at any time. They live mostly in the high flat-topped 
buttes, where there is good grass." Just when the last ones 
disappeared from this State and from Nebraska, where they 
were earlier exterminated, is difficult to say, but they must 
have held out in the Black Hills region till rather recent dates. 

Badlands bighorn (Ovis canadensi? auduboni] 




Ovis calif ornianus Douglas, Zool. Journ., vol. 4, p. 332, Jan., 1829 ("Near Mt. Adams, 

Yakima County, Washington"). 
SYNONYMS: Ovis canadensis samilkameenensis Millais, The Gun at Home and Abroad, 

vol. 4, p. 324, 1915 (Similkameen Mountains, British Columbia); Ovis cervina 

sierrae Grinnell, Univ. California Publ. Zool., vol. 10, p. 144, 1912 ("East slope of 

Mount Baker, Sierra Nevada, Inyo County, California"). 
FIGS.: Bailey, V., 1936, pi. 17 (horns). 

This race was believed to have been characterized by its 
slightly darker color than the Rocky Mountain bighorn, with 
heavier jaws and teeth, and especially by its horns, which 
were slightly more spreading and less closely coiled, as well 
shown in Bailey's (1936) figure. Cowan (1940), however, be- 
lieves the color hardly different from that of canadensis. 

Originally described on the basis of a specimen from Mount 
Adams in western Yakima County, Washington, sheep referred 
to this race extended northward into the mountains of south- 
central British Columbia, and southward through the lava- 
beds region of extreme northeastern California and perhaps 
the adjoining part of western Nevada to Tulare and Inyo 
Counties, California. The race at the present time is believed 
to be nearly extinct in California, where probably it occurred 
west to "include neighborhood of Mount Shasta, and to Sheep 
Rock . . , east side of Scott Valley, and to Siskiyou 
Mountains, in Siskiyou County . . . ; also south as far 
at least as Observation Peak, near Nevada line in eastern 
Lassen County" (J. Grinnell, 1933). In their account of the 
vertebrate natural history of the Lassen Peak region in ex- 
treme northeastern California, Grinnell, Dixon, and Linsdale 
(1930) adduce a few last records of sheep in that corner of the 
State. A band of about 40 "that had lived on Observation 
Peak were thought all to have perished in the severe winter of 
1922," and many skeletons were found there the following 
summer. In 1927 a small band thought to consist of four 
females and two males was located in the extreme southeastern 
corner of the region, close to the Nevada line. In Lassen 
County the last sheep of which the authors could secure in- 
formation was seen in 1872, but on Lassen Peak itself there 
seems to be no evidence that the animal ever was found. 


According to Dr. Joseph Grinnell (1933), it ranged in the 
high Sierra Nevada of California, from the vicinity of Sonora 
Peak, in Alpine County, southward to southeastern Tulare 
County; probably also this was the race formerly present on 
the upper parts of the White Mountains in Mono County, 
eastern California. At the date of writing this account, Dr. 
Grinnell said that it was still to be found from the vicinity 
of Mammoth Pass, Mono County, south to the vicinity of 
Olancha Peak and the Kaweah Peaks, Tulare County. Its 
altitudinal range extended from 5,000 feet (in winter) in Tulare 
County up to 13,000 feet on the ridge east of Whitney Pass. 
There was some evidence of an autumnal movement to lower 
altitudes in fall and winter, on the eastern slopes of the Sierra. 

In a consideration of the present status of this sheep in 
California, Dr. J. Dixon (1936) believes it is now in a very 
precarious position. In various ways this is a result of human 
influence, the chief factors being deer hunting, in the course of 
which bighorns may at times be unlawfully killed though ac- 
corded full legal protection, and grazing by domestic sheep on 
the lower winter ranges, which results in the eating of food that 
should be reserved for the bighorns. Intensive human intru- 
sion in the form of summer camps is an added disturbing 
factor. In September, 1935, Dr. Dixon knew of but a single 
band still in existence, probably numbering seven animals, 
near Mount Baxter. Testimony of deer hunters and others in 
the region is nearly unanimous that some bighorns are shot 
every year as camp meat and not for trophies. The critical 
time is only about a month in the hunting season, so that by 
bringing in a "roving ranger" at that period and by obtaining 
the active cooperation of the Forest Service, which is with- 
drawing domestic sheep from all parts of the bighorn's range 
where they might compete with the latter for food, there is 
still a chance for its survival. Those who have eaten it agree 
that no other meat that the game animals of this continent 
yield is equal to that of mountain sheep; hence arises the temp- 
tation to hunters to kill an occasional animal for the pot, even 
at considerable risk of detection. 

In Oregon this was a sheep of relatively low country, oc- 
curring over most of the State east of the Cascade Mountains. 
Bailey (1936) has gathered much detailed information on their 
former presence, from which a few notes are here extracted. 


"Originally mountain sheep inhabited every canyon, cliff, and 
lava butte as well as many of the rough lava beds of Oregon 
east of the Cascade Mountains. They were common until 
recent years in the Steens and Warner Mountains and are 
still found in the Wallowa Mountains and along the canyon 
walls of the Imhaha River . . . There are also records of 
sheep seen within the memory of many now living over most 
of the extensive lava beds and buttes of eastern Oregon." 
They were formerly abundant along the Deschutes Canyon; 
old settlers who came to the Bridge Creek region on the John 
Day River in 1873 found these sheep in bands of as many as 
50 or more, but they have long been extinct. In the early 
nineties they were numerous "on all the rimrock of the sur- 
rounding country from Burns to Bend, on the rough rim of 
Dry Basin, on Glass Mountain, Rams Rock, Juniper Moun- 
tain, in the Warner and Abert Mountains, around Christmas 
lake, and even out on the sagebrush plains where they some- 
times joined the herds of domestic sheep and fought the rams. 
They seem to have been last seen in the Mount Warner area 
about 1912. Captain Louis, an old Indian, told Bailey that in 
former days they frequently crossed the sagebrush valleys and 
then were hunted on horseback witji bows and arrows. His 
people reported them in the Wagontire and Juniper Mountains 
as lately as the autumn of 1915. In the Steens Mountains the 
last one was killed in 1911; at least a thorough search of these 
mountains in 1916 revealed no trace of living sheep. To the 
east of these mountains, they had gone at about the same time. 
In south-central Oregon the bighorn sheep disappeared shortly 
after the settlement of the country by white men. "In 1905 
James H. Gaut was told by people living west of lower Klamath 
Lake that mountain sheep had been numerous on the lava 
ridges near there up to 1885 and that the last was killed in 
1890." In Lake County, where formerly sheep were numerous, 
they had nearly disappeared by 1897. In the 1939 census of big 
game by the Fish and Wildlife Service, Oregon is credited with 
50 Rocky Mountain bighorns, but though these may include a 
few referred to this race that reach the extreme northeastern 
part of the State (see Bailey, 1936, p. 65), the figure probably 
also includes the remaining few Lava Beds bighorns mentioned 
as still found in the Wallowa Mountains. To the southeast 
there are probably still small numbers in western Nevada, but 
their precise subspecific status is unknown. 


In the State of Washington this sheep is "now nearly extir- 
pated, though a few are reported as of irregular occurrence in 
the Mount Chopaka and Mount Bonaparte region. " Formerly 
they occurred easterly in the Cascade Mountains from the 
Canadian boundary south to the Columbia River (Taylor and 
Shaw, 1929). In the 1939 census this State is credited with 
only ten bighorns. A few still exist also, according to Dr. 
R. M. Anderson (1939b), "in the southern interior of British 
Columbia (Okanagan, Similkameen, Lillooet, and Chilcotin 
districts), but apparently no sheep ever ranged in the Selkirk 
Mountains. " 

With this race now near extinction from its former range, 
one asks again why it should so suddenly have begun to dis- 
appear in the late eighties and nineties, and the answer is no 
doubt in part that its habit of frequenting lower levels, where 
it came in contact with the domestic sheep then on the increase, 
resulted in many areas in its contracting scabies and perhaps 
other diseases, for there is much testimony to the effect that 
hunting by white settlers alone did not account for the rapid 
decline. For example, Bailey (1936) quotes W. F. Schnabel 
that in the Mahogany Mountains of Oregon, where mountain 
sheep were plentiful, the^y practically disappeared in 1885, 
during the winter of that year and the one before. "They did 
not starve but were killed by some disease. I found their 
carcasses everywhere and grass and feed were plentiful in those 
days." Nowadays, with the greater care taken by ranchers 
to free their domestic stock from the ravages of the scabies 
parasite, the danger of infecting the feeding ranges is probably 
lessening, but in many instances this precaution may have 
come too late to save the bighorns. 

According to Maj. Allan Brooks (1923), the "mountain 
sheep of the greater part of the dry interior [of British Colum- 
bia] were wiped out forty or fifty years ago by the introduction 
of rifles to the Indians and the introduction of domestic sheep 
to their range. Scab decimated the sheep of the region east of 
the Fraser River about 1870. A virulent disease that affects 
the heart and liver is now being introduced in [the Chilcoten 
and Similkameen River districts] ... by domestic sheep 
that are brought over to graze from the state of Washington 
by sheep-herders. The government veterinary at Osoyoos 
is unable to determine this disease Ticks were 


very bad in 1897 and 1898. The ears of rams killed were 
packed to the drum with larval ticks, pale blue with sulphur- 
yellow legs. None were found in the ears of rams killed 
in 1902 and 1905." Brooks believes that coyotes and golden 
eagles annually kill "at least seventy -five per cent of all 
the lambs on two of the ranges on which" he has an op- 
portunity to observe, which "prohibits all possibility of any 
increase." As corrective measures he recommends the total 
prohibition of grazing permits for domestic sheep on any range 
inhabited by mountain sheep and the appointment of wardens 
whose duty it should be to enforce the game laws and to reduce 
the predatory eagles and carnivores. 

According to Cowan (1940) the largest remnant of the Cali- 
fornia bighorn is now to be found in British Columbia; a few 
are still living in the Ashnola and Similkameen district where 
they were formerly abundant. Small bands are present in the 
mountains near Vasseaux Lake and the northern end of 
Okanagan Lake. 



Ovis mexicanus Merriam, Proc. Biol. Soc. Washington, vol. 14, p. 30, Apr. 5, 1901 
("Lake Santa Maria, Chihuahua, Mexico"). 

SYNONYMS: Ovis canadensis texianus V. Bailey, Proc. Biol. Soc. Washington, vol. 25, 
p. 109, June 29, 1912 ("Guadalupe Mountains, Texas"); Ovis canadensis gaillardi 
Mearns, Mamm. Mexican Boundary, U. S. Nat. Mus. Bull. 56, p. 240, 1907 ("Gila 
Mountains, between Tinajas Altas and the Mexican Boundary Line in Yuma 
County, Arizona"); Ovis sheldoni Merriam, Proc. Biol. Soc. Washington, vol. 29, 
p. 130, Sept. 6, 1916 ("El Rosario, northern Sonora, Mexico"). 

FIGS. : Elliot, 1904, pt. 1, pis. 24, 25 (skull of type) ; Mearns, 1907, figs. 35-38 (feet, 
skull, horns). 

This is a pale desert race of the bighorn, with long ears, a 
short broad skull, and relatively short, moderately spreading 
horns. Height of ear from notch, 106 (95-120) mm. (Cowan, 
1940). The latter author describes the color in September 
topotypes as paler than in any of the other races in comparable 
pelage with the possible exception of cremnobates; above, pale 
vinaceous-fawn and vinaceous-buff, tail wood brown, lower 
legs between avellaneous and wood brown; much white on 
abdomen, extending forward narrowly on to chest, and as a 
narrow white line down fore and hind legs. Horns pale, taper- 
ing more rapidly than in nelsoni. 


In his recent review, Cowan (1940) regards the Texas and 
the Arizona bighorn as insufficiently distinguished, while 0. 
sheldoni of Sonora is based on a 'runt' or dwarfed individual, 
as pointed out by Dr. H. H. T. Jackson (in Ely and others, 

The range is from extreme southwestern Texas, southern 
New Mexico, and Arizona south across northwestern Chi- 
huahua and in Sonora to Seriland opposite Tiburon Island. 
Northward it intergrades with neighboring races. 

This race of mountain sheep was formerly found in the most 
arid desert ranges of western Texas in the San Andreas and 
Guadalupe Mountains and on the western slopes of the latter 
range in extreme eastern New Mexico as well as in eastern 
Chihuahua. Its numbers and distribution are now greatly 
reduced. Bailey (1905, 1931) has given a good summary of 
notes he obtained as to the status. In 1912-14 the number of 
these sheep in eastern New Mexico was estimated at 200 in 
the Guadalupe Mountains, but a more thorough census in 
1916 resulted in a count of about 100 or less. "All the reports 
from the Sacramento, Capitan, and Jicarilla Mountains assert 
that no sheep have been known in these ranges in modern 
times." To the northwest oJ the Guadalupe Range, in the 
San Andreas Mountains of New Mexico, there were a very few 
left in 1902, according to Gaut who made a special trip in 
search of them and as late as 1914 located about 30. At the 
same time he was "informed that sheep formerly were found 
along the crest of the Organ Mountains, but that none had 
been seen there in recent years." On account of the accessi- 
bility of their range, they were much hunted over this region, 
and Bailey in 1931 concluded that within the limits of New 
Mexico this race is now to be found only on the Guadalupe and 
San Andreas Ranges. He believes that "with adequate pro- 
tection for a term of years the mountain sheep could doubtless 
be brought back to its original range and abundance. The 
difficulty of enforcing game laws, however, in these uninhabited 
mountains is almost insurmountable, and wholly so without 
the full cooperation of the resident population. The fact that 
these sheep occupy land that will always remain practically 
worthless for stock raising or other agricultural purposes makes 
it doubly important that their numbers should be increased. " 
Since now the "whole summit and eastern slope of the Guada- 


lupe Mountains from Guadalupe Peak in Texas north to Dog 
Canyon in New Mexico is "a permanent game refuge that 
could easily support a thousand bighorns, the desirability of 
reestablishing the local race is apparent. "Under wise control 
and a definite plan for use of the surplus game, either for 
hunting or stocking other ranges, such an area could be made 
not only self-supporting but a valuable piece of property" 
(Bailey, 1931). 

In western Texas, in addition to the Guadalupe Mountains, 
a few sheep formerly were found in the Eagle and Corozones 
Mountains and on the northwest side of the Chisos Mountains. 
"They come into the Grand Canyon of the Rio Grande 
mainly from the Mexican side . . . The sheep are by no 
means confined to isolated mountain ranges. In several 
valleys I saw tracks," writes Bailey (1905) "where they had 
crossed from one range to another through open Lower Sonoran 
country. In this way they easily wander from range to range 
over a wide expanse of country in western Texas, and might be 
considered to have an almost or quite continuous distribution 
between the Guadalupe Mountains and the desert ranges of 
Chihuahua." Here they have more or less held their own for 
many years, and in the wildlife census of 1939 were estimated 
to number 280 individuals. . 

No data are at hand as to the status of this race in eastern 
Chihuahua, Mexico, but since the time of Bailey's investiga- 
tions a study of the bighorn sheep in Texas was carried out in 
1938 by W. B. Davis and W. P. Taylor (1939), which affords 
an excellent summary of present numbers and living condi- 
tions. This sheep is now confined in Texas to the extreme 
western wing of the State west of the Pecos River, an arid 
region having a scanty rainfall averaging between 9 and 17 
inches annually, depending somewhat on elevation. About 1 
percent of this area is under cultivation, and the rest is used 
for grazing. "The present range of the Texas bighorn in Texas 
is considerably more restricted than it was when Vernon Bailey 
worked in trans-Pecos Texas at the turn of the century." 
None occur now in the Chisos and Corozones Mountains or in 
the Grand Canyon of the Rio Grande, and they seem to have 
been absent in these areas for the past ten or fifteen years. 
"The heaviest concentration is north of the Texas and Pacific 
Railway in the Beach, Baylor, Carrizo, and Diablo mountains, 


where the bighorns doubtless cross freely from one mountain 
range to another." The authors quoted state that "probably 
not more than 25 mountain sheep occur in all the Guadalupes, 
none of them on the top in the heavily wooded sections." 
The numbers here seem to be decreasing but were probably 
never very high. In other ranges single rams or rarely small 
bands appear sporadically from time to time, but in other of 
the 16 mountain masses for which data were gathered sheep 
have been absent for years. Perhaps not more than 30 occur 
south of the Texas and Pacific Railway. Summarizing the 
figures given, there are approximately 410 bighorns in western 
Texas, of which some 300 are found in the four mountain 
ranges Baylor, Beach, Carrizo, and Sierra Diablo, while the 
remainder are chiefly in the Apache (40), Guadalupe (25), 
Delaware (17), Eagle (8), and Glass (13) Mountains. The 
requirements of these sheep include not only craggy regions 
but also a particular type of vegetation, of low, shrubby, 
xerophytic sorts, low enough to afford an unobstructed view. 
In the Baylor Mountains the evidence seems to show that 
bighorns move into them seasonally and leave "when condi- 
tions are adverse." Young are born in March and April, 
usually a single one at a time, and they follow their mothers 
throughout at least the first summer and autumn. Davis 
found that in the Yellowstone National Park grass constituted 
nearly 60 percent of the food for Rocky Mountain bighorns, 
but for the Texas bighorns investigation shows that grass con- 
tributes about 3 percent only, and that the animals prefer 
mountain-mahogany, Mexican tea, yellow trumpetflower, 
mock-orange, and wild onion. Davis and Taylor consider 
carefully the reasons for the decrease in numbers of this big- 
horn and conclude that it is not due to overhunting, for the 
season has been closed since 1903; nor is it a result of depre- 
dations by mountain lions, for these predators have been re- 
duced to very small numbers. Golden eagles may take toll of 
a few lambs, but these birds are not a very significant factor, 
and besides predators have always lived in the region. They 
conclude that "the incursion of domestic sheep possibly has 
been the most serious factor affecting bighorn numbers" and 
suggest that this is in part because of the introduction of 
diseases to which domestic sheep are liable, in part to direct 
competition for food and water, and in part perhaps to an 


intangible factor of psychic incompatibility, for bighorns are 
commonest where sheep are fewest. On account of the rather 
unfavorable conditions of the area, they doubt if it will ever be 
possible to increase the bighorn population enough to warrant 
an open season in western Texas. 

There is little recent information at hand as to the present 
status of the bighorn in northern Chihuahua or the adjacent 
parts of New Mexico, beyond the fact of its evident deple- 
tion within the past 50 years. In extreme southwestern New 
Mexico Mearns in 1892 found them in some abundance in 
the Dog, Big Hatchet, and San Luis Mountains. In 1900 
sheep were brought in and sold for meat at Deming from the 
mountains near the international boundary; but by 1908 there 
were only a few remaining in the Big Hatchet Mountains, 
"and probably none in the other ranges of southwestern New 
Mexico. In the country north of Deming there seem to be no 
recent records," nor could Goldman and Bailey at that time 
procure any records for the Burro and Carlisle Mountains or 
the Mogollon Mountains. In 1905 old horns were reported to 
Hollister as found from time to time in the little ranges from 
the Magdalenas to the Zunis, But no one could tell when the 
animals had last lived there. 

In northern Chihuahua and the adjacent sierras of Sonora 
these sheep were found up to a few years ago on all the ranges, 
but now, owing to relentless hunting, and in spite of the fact 
that these sheep are not to be killed legally, they are in danger 
of becoming extinct. A reservation for this animal is contem- 
plated in the Altar district of Sonora (Zinser, 1936, p. 9). 

Bighorn sheep, presumed to be of this race, formerly in- 
habited the Chiricahua Mountains in the extreme southeastern 
corner of Arizona but are now extinct there. In former years, 
according to information supplied to Cahalane (1939a, p. 439), 
a life-long resident of the mountains recalled that they were 
"fairly numerous" in all the lava hills of the vicinity but were 
gradually shot out, though he believed that the last remnant 
(probably in the Rincon Mountains) succumbed to the drought 
of 1903-5. 

During the early nineties mountain sheep were common along 
the international boundary and the adjacent parts of Arizona. 
Mearns (1907) writes that he saw many horns in the Papago 
Indian settlement, and the Indians reported them common in 


the higher ranges visible from Nariz, where they killed many, 
in consequence of which the animals were much scarcer than 
they had formerly been. The surveyors of the party saw sheep 
"in the rugged Tule Mountains in 1892. When my party was 
there, in February, 1894, no sheep were seen, but many tracks 
and heaps of horns were noted, as also in the neighboring 
Granite Mountains. During our stay at Tinajas Altas, at the 
foot of the Gila Mountains, from February 14 to 23, 1894, 
sheep were seen on four occasions, in flocks of 6, 3, 3, and per- 
haps as many as 20. They were feeding largely upon a Cy- 
lindropuntia cactus, in valleys at the base of the mountain, 
but tracks and beds were seen at all altitudes." 

Dr. W. P. Taylor (1936, p. 653) speaks of this region as a 
desert of rare beauty, and one of the most arid in the United 
States. "There is but one small cow outfit in the entire 
region, at least on the American side. A few prospectors 
work the territory. Aside from a handful of Indians and an 
occasional smuggler or United States custom agent, there are 
no other inhabitants of the area, and no permanent resi- 
dents at all. Both antelope and bighorns are at present subject 
to poaching on both sides of the international boundary 
through lack of adequate warden service. " 

In 1939, by Executive order, two game ranges were estab- 
lished in Arizona, with a view especially to protecting the 
sheep, mule deer, antelope, and peccaries of this region. One 
in central Yuma County, the Kofa Range, comprises over 
660,000 acres and is at one point within 15 miles of the border 
of southeastern California. The other, the Cabeza Prieta 
Range, lies 35 miles to the south on the international border, 
in Yuma and Pima Counties, and includes 866,880 acres. 
Negotiations are still pending with the Mexican Government 
to set aside a corresponding adjacent area on the Mexican side 
of the border. It is believed that these game ranges will prove 
of great value in preserving the larger wild mammals of the 
State. "A few years ago this section abounded in game, but 
indiscriminate hunting and poaching came dangerously near to 
eliminating some of the most valuable species. " 

As to the present status of this sheep in Arizona, Cowan 
(1940) quotes A. A. Nichol, who has made it the subject of 
special investigation lately. He gives an approximate census 
for 1937 as follows: Granite Mountains, 10; Aquila Moun- 


tains, 4; Gila Mountains, not including the Tinajas Altas, 15; 
Kofas, 25; Trigos, 10-15; Chocolate Mountains, 6 or 7; Buck- 
skin Mountains, 10-12; west end of Harcuvar Mountains, 7 or 
8; Black Mountains, 60-75; Superstition Mountains, 47; while 
C. T. Vorhies adds that there were known to be at least 6 in 
the Tucson Mountains and about 71 in the Santa Catalina 
Mountains altogether approximately 275 animals. 



Ovis nelsoni Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 218, July 15, 1897 
("Grapevine Mountains, on boundary between California and Nevada"). 

This is a "comparatively small Bighorn, with rather slender 
horns somewhat triangular in cross-section near base; color 
very pale; white rump patch divided lengthwise to base of tail 
with dark line" (H. H. T. Jackson, in Ely and others, 1939). 
Molar teeth heavy, horns slender. Total length, 1,280 mm.; 
tail, 100; height at shoulder, 830. In its pale salmon-gray 
pelage, this subspecies is strikingly different from the Rocky 
Mountain bighorn ; and its ears' are noticeably longer. 

This pale, small race is found in the desert mountain ranges 
of southeastern California east of the Sierras and in the ad- 
jacent parts of southern Nevada. Although the limits of dis- 
tribution have not been carefully worked out, Dr. J. Grinnell 
(1933) regarded it as the form now or lately in the mountain 
ranges of Inyo and Mohave Desert regions, northeast to Owens 
Valley, Mono County, south to the Chocolate Mountains in 
Imperial County, and west to San Bernardino, San Gabriel, 
and (formerly) the Tejon Mountains and the southern end of 
San Joaquin Valley, California. 

Recent observations on the status of this desert race are few. 
According to Burt (1934, p. 424) they are still common in the 
Sheep Mountains of southern Nevada, and he was informed of 
a herd of 24 at Corn Creek which watered at a cattle tank. 
They were formerly common in the Charleston Mountains, 
but few if any were left by 1934, although Burt was told of a 
small band seen shortly before, along the southwestern border 
of the mountains. In California they are somewhat more re- 
stricted than formerly but still remain in some numbers in the 
desert regions of the southeastern parts of the State. The 


1939 estimate of their numbers by the Biological Survey 
credited California with 1,770 desert bighorns, and Nevada 
with 1,485. If these figures are approximately correct, the race 
is no longer in immediate danger. In both States they are 
protected by law throughout the year. Cowan (1940) has 
given a good summary of its recent history with a list of speci- 
mens examined, including one from as far west as Caliente 
Peak, San Luis Obispo, Calif. 



Ovis cervina cremnobates Elliot, Field Columbian Mus. Publ. 87, zool. ser., vol. 3, p. 

239, Dec., 1903 (Matomi, San Pedro Martir Mountains, Lower California, 

FIGS.: Elliot, 1904, pt. 1, figs. 26, 27 (drawings of adult ram, immature ram, and ewe). 

This race of bighorn is described as resembling 0. c. nelsoni 
"but of a much lighter color, the head of a three-year-old ram 
being nearly white, with a very small caudal patch not divided 
from color of upper parts by any perceptible line; fore part of 
legs almost black . . . ; head very broad between orbits, 
from 20 to 25 mm. broader in old rams than the head of 0. c. 
nelsoni; horns of adult rams very large. " Length along outer 
curve of horns in an old ram, 850 mm. (Elliot, 1904) . "Perhaps 
the palest" of the races. 

The isolated desert mountains of the northern half of Lower 
California, Mexico, northward to extreme southern California, 
on the eastern and northeastern faces of the mountains along 
the west side of the Colorado Desert, and to the lower northern 
slopes of the San Jacinto Mountains in San Gorgonio Pass, 
form the range of this palest of the desert bighorns. In south- 
ern California it is not known ever to have invaded the Pacific 
slopes but reached the eastern edges of San Diego and River- 
side Counties. 

This was the first of the races of the bighorn known to Euro- 
peans, for it was briefly described as long ago as 1702, in a 
memoir by the Jesuit priest Francis Maria Piccolo, on the 
discovery of a passage by land to Lower California. The 
animal was called the "Taye" and was figured in 1758 by 
Venegas, as detailed by J. A. Allen (1912) in whose historical 
paper the figure is reproduced. He writes: "Sheep still exist 
where the first Spanish missionaries found them in 1697. Dr. 


Charles H. Townsend on his recent 'Albatross' expedition to 
Lower California . . . obtained some imperfect skulls of 
mountain sheep from the natives at Conception Bay, and saw 
a living specimen in the low mountains at the head of that bay. 
He also informs me that mountain sheep are said still to in- 
habit the low mountains near the Gulf coast as far south as 
Saltillo del Rey, or to within about one hundred miles of La 
Paz, and that they range thence northward in all the high hills 
and mountains, especially on the Gulf side, nearly to the 
United States boundary." Mearns (1907), during the survey 
of the international boundary, was informed by Indians that 
sheep were then (1894) abundant in most of the rocky ranges 
of northern Lower California. In more recent years, however, 
the needs of miners and prospectors have jeopardized the con- 
tinued existence of bighorns in the region. Dr. E. W. Nelson 
(1921) wrote that they were then "still widely distributed on 
the main mountains of the eastern half of the peninsula," 
but "the difficulties of transportation and the scarcity of live 
stock through most parts of the peninsula render the securing 
of food supplies so difficult that great numbers of game ani- 
mals, especially mountain sheep^ have been killed and the meat 
dried, or jerked, to supply mining camps and other communi- 
ties. This deplorable and wasteful slaughter still continues 
and unless checked will ultimately result in the extermination 
of the sheep . . . Mountain sheep are peculiarly en- 
dangered by this slaughter owing to their habit of going in 
bands and drinking at certain watering places, where hunters 
lie in wait behind blinds built of loose stones and kill indi- 
viduals of all ages and sexes, often in a few minutes destroying 
an entire band. I have been informed of one party having 
killed more than 100 sheep in this manner to make dried meat 
during a single season." 

Although in 1917 the Mexican Government prohibited the 
hunting or exploitation of mountain sheep in Lower California, 
"no effective enforcement of this regulation appears to have 
been undertaken" (Nelson, 1921, p. 132). 

Cowan (1940) regards specimens from slightly north of 
Calmalli, Lower California, as intermediate between this and 
the race weemsi of the lower part of the peninsula, while others 
from the Chuckawalla Mountains, California, are intergrades 
between cremnobates and nelsoni. 




Ovis canadensis weemsi Goldman, Proc. Biol. Soc. Washington, vol. 50, p. 30, Apr. 2, 
1937 (Cajon de Tecomaja, Sierra de la Giganta, about 30 miles south of Cerro 
de la Giganta, southern Lower California, Mexico). 

This recently named race occupies the mountainous country 
at the tip of the peninsula of Lower California and is remarka- 
bly dark for a desert-frequenting animal. In its "usually 
darker" color it seems chiefly to differ from the race cremno- 
bates of the upper parts of the peninsula, "varying from very 
dark brown more or less mixed with black, " rump patch white, 
almost divided by the tail stripe. Horns in the adult female 
are said to be remarkably long and gradually tapering; in males 
the lateral surface is flatter and the external ridge more promi- 
nent than in other races. 

Very little information is at hand as to the status of this 
sheep. Major Goldman describes the country in which it 
lives as supporting a richer vegetation than that to the north 
where the race cremnobates is found, indicating a greater rain- 
fall. The Sierra de la Giganta is the highest range of moun- 
tains in the southern part of the Cape, about 60 or 70 miles 
long and isolated by a low gap on the north and the south, 
close to the Gulf of Mexico. Its eastern face is much steeper 
and more broken than the western. The flora includes a num- 
ber of subtropical species not found to the north. Here this 
southernmost race of mountain sheep still occurs in small 
numbers. Goldman mentions four specimens secured by his 
party at the type locality in 1936. There is a mounted head 
in the Museum of Comparative Zoology obtained in 1925 near 
La Paz, and it is of the dark-brown color characteristic of the 
race. Although doubtless the puma occasionally attacks this 
sheep, as Goldman intimates, yet this animal is probably not 
now common enough in Lower California to be nearly so im- 
portant an enemy as man. The Mexican Government nomi- 
nally gives the sheep of this area full legal protection, but, as 
earlier mentioned by Dr. Nelson, the law is difficult to enforce. 
Nevertheless it is encouraging to learn that this small remnant 
of mountain sheep still holds out in the isolated ranges of the 
tip of Lower California. Cowan (1940) regards specimens 
from Sierra San Borjas, 20 miles north of Calmalli, as inter- 
mediate between this race and 0. c. cremnobates. 


Order EDENTATA: Edentates 

THE NEW WORLD edentates are discussed elsewhere in 
the present volume, p. 34. The only edentate considered 
in this section is the Patagonian giant ground sloth, which be- 
came extinct after man reached southern South America. 

Family MEGATHERIIDAE : Giant Ground Sloths 



Neomylodon listai Ameghino, Premiere Notice sur le Neomylodon listai, un Representant 

Vivant des Anciens Edentes Gravigrades Fossiles de 1'Argentina, La Plata, Aug. 

1898; transl. in Natural Science, vol. 13, pp. 324-326, Nov., 1898 (Consuelo Cove, 

Last Hope Inlet, Patagonia). 
SYNONYM: Grypotherium domesticum Hauthal, Roth, and Lehmann-Nitsche, Revista 

Mus. La Plata, vol. 9, pp. 409-474, 1899. 
FIGS.: Hauthal, Roth, and Lehmann-Nitsche, 1899, pis. 1-5; Woodward, 1900, pis. 

5-9 (piece of skin). 

Much interest was aroused by the announcement in 1898 of 
the discovery of a large piece of skin of a ground sloth in south- 
ern Patagonia. In a preliminary notice, F. Ameghino briefly 
described this and, believing it to represent a new genus re- 
lated to the extinct giant Mylodon, named it Neomylodon listai, 
after the ill-fated explorer Lista, who described what he be- 
lieved to have been a living example in the region of the eastern 
base of the Andes. Although it is probable that he was mis- 
taken in identifying the animal he briefly saw, nevertheless the 
finding soon afterward of the portion of skin with the hair in- 
tact, and with the small bony nodules imbedded in it, as 
characteristic of the Pleistocene Mylodons, raised at the time 
high hope that a living example might yet be found. With 
this object in view, H. Hesketh Prichard (1902a) undertook a 
journey to the region, an account of which he gives in his book 
"Through the Heart of Patagonia"; no evidence did he find, 
however, that this mysterious animal was still extant. 



The piece of skin on which the accounts of the animal are 
based was so fresh that the hair and serum on it, though 
dried, were still intact. The hair was 4 or 5 cm. long, reddish 
tinged with gray, while in a large portion of the fragment were 
numerous small oblong or rounded dermal ossicles. The skin 
was discovered in a large cavern, where at a shortly subsequent 
date further investigations were made by Hauthal, geologist 
of the La Plata Museum. Here he found not only "another 
piece of skin, but also various broken bones of more than one 
individual of a large species of ground-sloth in a remarkably 
fresh state of preservation. Moreover, he discovered teeth of 
an extinct horse and portions of limb bones of a large feline 
carnivore, in association with these remains; he likewise met 
with traces of fire, which clearly occurred in the same deposits 
as the so-called Neomylodon. All these remains were found 
beneath the dry earth on the floor of an enormous chamber, 
which seemed to have been artificially enclosed by rude walls. 
In one spot they were scattered through a thick deposit of 
excrement of some gigantic herbivore, evidently the ground- 
sloth itself; in another spot they were associated with an ex- 
tensive accumulation of cut hay. Dr. Hauthal and his col- 
leagues, indeed, concluded that the cavern was an old corral in 
which the ground-sloths had been kept and fed by man" 
(Woodward, 1900). 

Remains of at least three individuals were found in this cave. 
So fresh were these that bits of dried tissue still adhered to 
some of the bones, and one of the skulls showed evidence of 
having been cut away at the occiput by an instrument. In the 
lower jaw there were four cheek teeth without any evidence of 
a caniniform tooth in front of them. The upper jaw likewise 
had four cheek teeth, thus fixing the identity as Grypotherium 
instead of Mylodon, in which there are five, as also in 
other large ground sloths. Of particular interest are the 
masses of excrement, which have been determined to consist of 
grasses largely. A few pieces of stems appear sharply cut and 
suggest that, like the quantity of cut hay found in association 
with the remains, these had been fed to the animals by their 
human captors. Complete measurements are not available, 
but these ground sloths were doubtless as large as a cow, 
though differently and heavily proportioned. 


Order RODENTIA: Gnawing Mammals 

The rodents, as an order, are discussed hereinbefore (p. 41). 
Representatives of four family groups in South America are 
considered vanishing forms: 

(1) Cricetidae. The rats and mice of the New World are 
separated from most of the Old World types by important 
dental characters, although frequently similar in appearance. 
Six species of two genera of rice rats, peculiar to the Galapagos 
Islands, are in danger of extinction. 

(2) Myocastoridae, hystricoid water rats, or coypus. Three 
races are in need of protection because of persecution for their 

(3) Dinomyidae, hystricoid giant rats. A single species is 
considered here. 

(4) Chinchillidae. Three races of the rare chinchilla are in 
need of protection. J. E. H. 

Family CRICETIDAE: Hamsterlike Rodents 

The Galapagos group of islands, lying on the Equator nearly 
600 miles due west of the western coast of Ecuador, is remark- 
able for the various peculiar endemic birds found there, in- 
cluding the flightless cormorant, the dwarf penguin, and such 
land birds as Geospiza, while among reptiles there are sundry 
species of giant land tortoises, large iguanas of a genus peculiar 
to the islands, and others. The land mammals, however, until 
a few years ago were believed to comprise but two species 
related to the continental rice rats of the genus Oryzomys. 
Later explorations have now raised the number of indigenous 
rodents to six, representing two related genera, Oryzomys and 
Nesoryzomys. Commenting on these discoveries, Osgood 
(1929) writes that the native rodents now "take on consider- 
ably more importance than formerly and will doubtless need 
serious consideration in speculation regarding the derivation 
of the insular fauna. Until thorough study of mainland forms 
is made, however, no satisfactory conclusions are to be ex- 
pected. While the oryzomyine rodents are widely distributed 
and greatly varied in South America, they are also highly 
developed in Central America, and no competent study of the 


whole group has yet been undertaken. When such a study is 
made, it may be possible to determine something more than 
the general affinity of the insular and continental forms. At 
present, it can not be affirmed even that the nominal genus 
Nesoryzomys is confined to the Galapagos, for at least in some 
of its characters it is closely paralleled by certain forms of the 
mainland. However, the number and diversity of the island 
rodents may perhaps be taken as indicating that their existence 
on the islands is not an accidental matter and the view is 
somewhat substantiated that the present land area has been 
reduced from former larger proportions. Furthermore, it 
seems quite certain that before the introduction of house rats 
the native rodents were more generally distributed throughout 
the different islands of the group than at present." Since the 
various species have much general similarity in appearance, they 
may have been frequently confused with house rats by casual 
visitors, and so neglected with the result that very little is as 
yet known of them. The six species hitherto described are the 



Mus galapagoensis Waterhouse, Zool. Voyage of the Beagle, Mammalia, pt. 2, p. 65, 

1839 ("Chatham Island," Galapagos group). 
FIGS.: Waterhouse, 1839, pi. 24 (animal in color), pi. 33, fig. 8, a-c (skull and teeth), 

pi. 34, fig. 14, a (lower jaw). 

Slightly smaller and more delicately built than a roof rat, 
this species has rather large thin ears, a tail about as long as the 
body, and in color is brownish above, with a mixture of black 
hairs and others having a subterminal yellowish band; the sides 
are yellowish, and the under surface of the body is whitish, with 
a faint yellowish wash and with dark gray bases to the hairs. 
Feet whitish; tail distinctly bicolor, dark above, white below. 
Head and body about 6 inches long (150 mm.) ; tail, 4.75 inches 
(120 mm.); hind foot, iVe inches (about 30 mm.); length of 
skull, 31 mm. 

The colors and general appearance of many of the small 
rodents of South America are rather nondistinctive, so that 
cranial characters give the best clues to the genera. Oryzomys 
is characterized by the double series of enamel tubercles on 
the anterior molars, with narrow secondary enamel folds 
between them. 


When Darwin visited Chatham Island in 1835 he found this 
rice rat abundant. "It frequents the bushes," he wrote, 
"which sparingly cover the rugged streams of basaltic lava, 
near the coast, where there is no fresh water, and where the 
land is extremely sterile." He seems to have made search for 
it or other native rats on other islands, for he remarks that he 
was unable to find it on other parts of the archipelago; nor has 
any one since taken specimens. Indeed, Heller (1904), who 
visited the island in 1898-99 and searched for it unsuccessfully, 
believes "it is now probably extinct or else restricted to the 
barren eastern part of the island where Darwin secured his 
specimens." If it has really been extirpated, one may suppose 
that the introduced brown and black rats have been too 
active competitors. 



Oryzomys bauri J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 4, p. 48, May, 1892 
("Barrington Island," Galapagos group). 

The rice rat of Barrington Island, in the Galapagos Archi- 
pelago, is believed to be very Closely allied to 0. galapagoensis 
of Chatham Island but "differs mainly in having a somewhat 
shorter tail and less yellow on the upper parts" (Heller, 1904). 
The general color is said by its describer to resemble very much 
that of the cotton rat, Sigmodon hispidus. A series of average 
measurements given by Heller: Total length, 273 mm.; tail, 
136; hind foot, 30-31; greatest length of skull, 31.5. 

This rodent is known only from Barrington Island, about 30 
miles to the east of Chatham Island. Here Dr. George Baur 
obtained the three specimens that served for the description of 
the species a number of years ago. As Heller points out, the 
lack of actual specimens of 0. galapagoensis makes close com- 
parison out of the question, but there is every reason to believe 
that it is distinct from the latter. Dr. Baur found it rather 
common on Barrington, "between the bushes near the shore, 
and also high up between grass and the lava rock." When 
Heller visited the island ten years later, in 1898-99, he found 
it "very abundant," inhabiting crevices among the loose lava 
rocks and burrows and runways beneath bushes and brush 
piles. "In habits it appears to be somewhat diurnal and was 
as often seen at midday as at other times." Nothing further 


seems to be known of the species, but presumably it still is to 
be found on its island home. 


Oryzomys indefessus Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 4, p. 280, 1899 ("Inde- 
fatigable Island," Galapagos group). 
FIGS.: Orr, 1938, pi. 25, figs. 3, 3a (skull). 

This and the three other remaining rodents of these islands 
are placed in a somewhat aberrant genus, Nesoryzomys, which, 
though thus far considered endemic, may nevertheless eventu- 
ally prove to be represented on the mainland of South America, 
as Osgood (1929) has remarked. Compared with typical 
Oryzomys, the frontal bones are much narrower, with rounded 
orbital edges; the snout is more elongate, the nasals narrower 
and less convex in profile, and the tooth rows are shorter. 
Externally the fur and the tail are shorter. The present species 
is dull mouse gray above, heavily lined with black, and with a 
slight tinge of fawn on the posterior back; below, the hairs are 
slaty at the base, tipped with white. The tail is about as long 
as head and body, well haired, black above and white below. 
An average specimen measures: Total length, 269 mm.; tail, 
108; hind foot, 29; ear, 22. Skull length, 35.3. 

This species was described from a specimen taken on Inde- 
fatigable Island by the Rothschild Expedition. Later, in 
1898-99, Heller found it abundant on the same island, as well 
as on South Seymour Island, and secured a series of specimens. 
He did not find it on the adjacent North Seymour Island. 
He writes (1904) that it "seems to be more nocturnal in habits 
than Oryzomys. It inhabits burrows or rock crevices beneath 
bushes." Nothing further is known of its habits. Although 
at present believed to be still common, it is probable that if 
Old World rats and mice or house cats (the latter recently 
introduced) should become established on these islands, as 
they have on some others of the group, this native rat would 
be in jeopardy. 


Nesoryzomys narboroughi Heller, Proc. California Acad. Sci., ser. 3, zool., vol. 3, p. 242, 

Aug. 31, 1904 ("Narborough Island," Galapagos group). 
FIGS.: Orr, 1938, pi. 25, figs. 2, 2a (skull). 


The rice rat of Narborough Island is described as similar to 
N. indefessus of Indefatigable but larger, the feet and ears 
especially larger, the former 31 mm. or greater in length; 
coloration much darker above, chiefly blackish mixed with 
some rusty brownish; below, darker gray (Heller, 1904). The 
skull is wider, with longer nasals and shorter palatal foramina 
than in the latter. Total length, 303 mm. ; tail, 131 ; hind foot, 
33; ear, 23. An adult male skull measured 41.5 in length. 

According to Heller's account, "this species was found in- 
habiting the cracks and fissures in barren black lava fields near 
the coast of Mangrove Point, Narborough Island. Individuals 
were rather scarce at this locality, perhaps owing to the paucity 
of the vegetation. The contents of several stomachs were 
examined and found to contain a reddish material resembling 
pulverized Crustacea." At the time of Heller's visit the Euro- 
pean rats and house mouse apparently had not reached Nar- 
borough Island. Should they later become established there, 
it is likely that this native species will suffer. 


Nesoryzomys swarthi Orr, Proc. California Acad. Sci., ser. 4, vol. 23, p. 304, Sept. 1, 
1938 ("From the vicinity of Sulivan [ = Sullivan] Bay, James Island, Galapagos 

FIGS.: Orr, 1938, pi. 25, figs. 1, la (skull). 

This island race is evidently closely related to N. indefessus 
of Indefatigable Island and is indistinguishable from it in 
color, but the skull shows broader nasals, a larger rostrum, 
greater diastema and much larger molariform teeth than any 
of the other species of the genus yet known. The average 
measurements of three males are given as: Total length, 312.3 
mm.; tail, 133.7; hind foot, 35.7; length of skull, 40.6. 

The four specimens from James Island, from which no 
mammals had previously been known, were collected in 1906 
by J. S. Hunter for the California Academy of Sciences and 
may be regarded as the representative form on that island, of 
the indefessus type, which is again present on Narborough 
Island in the much darker N. narboroughi. In the 30 years and 
more since the species was discovered, no other examples have 
been collected by more recent expeditions, so that, as Orr 
writes, "the possibility exists that this form may now be 


extinct as a result of the introduction of non-native old world 
rats. Examples of Rattus rattus alexandrinus were taken dur- 
ing the same visit to James Island on which the specimens of 
N. swarthi were secured." Heller (1904) remarks that "leav- 
ing out of consideration Albemarle, James, Charles, and 
Duncan, the islands which are inhabited by introduced species, 
it seems remarkable that large islands like Abingdon, Bindloe, 
and Hood should lack indigenous species of Muridae [Criceti- 
dae]. A careful search, however, failed to discover any mam- 
mals on these islands. It is probable that James, Duncan, and 
Albemarle islands, intervening between the ranges of the two 
species of Nesoryzomys, were until recently inhabited by 
indigenous species of this genus which became extinct upon 
the introduction of" the house rats. This suggested distribu- 
tion is now partly substantiated through the subsequent dis- 
covery of Nesoryzomys of a related type on James Island. 
Moreover, the fact that Heller failed to find it in 1898-99, 
while Hunter secured four specimens in 1906, merely shows 
how elusive small mammals may be, even when sought for by 
so skilled a collector as Heller. 


Nesoryzomys danvini Osgood, Field Mus. Nat. Hist., zool. ser., vol. 17, p. 23, July 12, 

1929 (Academy Bay, Indefatigable Island, Galapagos group). 
FIGS.: Orr, 1938, pi. 25, figs. 4, 4a (skull). 

From its nearest island relatives, N. indefessus and nar- 
boroughi, this species differs in its decidedly smaller size and in 
its bright fulvous coloration, which extends to the entire under 
parts. The skull is slender and without sharp ridges or angles. 
Total length, 222 mm.; tail, 89; hind foot, 27; length of skull, 

This small, bright-colored species is remarkable in that it 
furnishes the only instance known of two species of the genus 
occurring side by side on the same island, "indicating a dis- 
tinction of long standing." Three specimens were taken at 
Academy Bay and a fourth at Conway Bay, Indefatigable 
Island, while on the same dates 12 examples of N. indefessus 
were taken at the former and 13 at the latter locality, indicating 
that it is probably much the less numerous. Orr (1938) records 


the capture of two additional specimens at Academy Bay by 
the California Academy's Expedition of 1905-6, making six 
known examples. Though this species at present may be in no 
danger, it is likely to disappear first if Old World murids are 
introduced or other unfavorable circumstances intervene. 

Family MYOCASTORIDAE : Coypus 


Mus coypus Molina, Saggio Stor. Natur. Chile, p. 287, 1782 (Chile). 

SYNONYM: Guillinomys chilensis Lesson, Nouv. Tabl. Regne Anim., p. 126, 1842. 


Myopotamus bonariensis E. Geoffroy, Ann. Mus. Hist. Nat. Paris, vol. 6, p. 81, 1805 
(Parana River, Paraguay). 


Myocastor coypus santacruzae Hollister, Proc. Biol. Soc. Washington, vol. 27, p. 57, 
Mar. 20, 1914 ("Rio Salado, near Los Palmares, Santa Cruz, Argentina"). 

FIGS.: Waterhouse, 1848, pi. 15, fig. 1; pi 16, fig. 1 (skull) ; Hudson, 1892, p. 12 (female 
and young) 

The coypu, or nutria, is so important as a fur animal and has 
in some places become so depleted that it may be briefly con- 
sidered here. With somewhat the appearance of a large musk- 
rat, this is a dark yellowish-brown or reddish-brown animal, 
with longer and coarser guard hairs than a muskrat, which 
when plucked disclose a soft velvety coat of under fur of a dark 
slaty color. The tail is about as long as head and body and is 
covered with coarse scales, but unlike the tail of the muskrat 
it is round instead of compressed from side to side. The large 
hind feet are webbed for swimming. Tip of muzzle and chin 
white. Length of head and body, about 25 inches; tail, 17.5 
inches; hind foot, 5.5 inches. The large incisor teeth are of a 
deep orange-red; the cheek teeth are four in each row, with 
three outer and one inner enamel fold in each of the upper and 
the reverse in each of the lower series set at an angle directed 
forward. Length of skull, 4.8 inches. 

As a species this large aquatic rodent is found in the tern- 


perate parts of southern South America, from the valleys of 
central Chile in about latitude 33 S., southward to the Straits 
of Magellan and eastward across Argentina to extreme southern 
Brazil, Paraguay, and Uruguay. Over this wide area there is 
some geographic variation, so that at present three living 
races are recognized: the typical form of Chile at lower alti- 
tudes; the race bonariensis of northern Argentina, Uruguay, 
Paraguay, and southern Brazil, in which the notch at the 
posterior border of the palate is broadly arched and concave 
instead of strongly V-shaped as in the typical coypus; and 
lastly the Patagonian race M. c. santacruzae, of southeastern 
Argentina, in which the skull is larger with the posterior border 
of the palate V-shaped as in the Chilean animal, but the upper 
cheek teeth increase conspicuously in size from before back- 
ward, the last with nearly twice the crown area of the first, 
instead of being of uniform size. All three races are evidently 
closely related and to be distinguished only upon comparison 
of skins and skulls. 

The coypu, like the North American muskrat, is an animal 
of lakes, streams, and swamps, but on the south coast of Chile 
it is said to frequent the coastal tide waters as well, living in 
the bays and channels among the small islands. Although 
primarily a vegetarian, it is said also to eat shellfish. According 
to Bennett the breeding season is in September and October, 
but Gay (1847) states that there are two litters, or even three, 
in a year, and young may be up to five or seven to a litter. 
Hudson (1892), however, says that the number may be as 
many as eight or nine, and he has published a sketch of an 
adult female, with her brood, swimming, some of the young 
being ferried along on her back, the others paddling behind. 
He mentions the peculiar moaning cries the animals make, 
but these may be given mostly at mating seasons, as with the 

The soft, plushlike underfur of this animal early attracted 
the notice of Europeans. At the beginning of the last century, 
Azara in his account of the species in Paraguay (where it was 
known by the native name of "quouiya") says that the fur 
was then beginning to be used for felt hats. Since his day this 
use has greatly increased, while the pelts also, after the long 
coarse guard hairs have been plucked, are much used in fur. 
They dye easily and give a soft plushlike effect. So great has 


become the demand for this fur that in recent decades the 
animals have in places become greatly depleted. 

Hudson, writing in 1892, said that it was much more abun- 
dant in the La Plata region 50 years previously than at that 
time. Its skin was largely exported to Europe. "About that 
time the Dictator Rosas issued a decree prohibiting the hunting 
of the coypu. The result was that the animals increased and 
multiplied exceedingly, and abandoning their aquatic habits, 
they became terrestrial and migratory, and swarmed every- 
where in search of food. Suddenly a mysterious malady fell 
on them, from which they quickly perished, and became 
almost extinct." This account of a sudden and decimating 
epizootic among these rodents recalls the cyclic fluctuations 
known among such northern rodents as meadow mice and 
snowshoe hares. No other evidence is known to me of such an 
epizootic elsewhere among the coypus. 

In his account of the mammals of Uruguay, Sanborn (1929) 
writes: "Coypus have been so hunted for their fur that they 
are now quite scarce in settled districts. I heard of many at 
the Laguna Negra in Rocha but did not have a chance to 
visit there. At most places tBe people said there were a few 
left where many had been." The Argentine Government, in 
1931, issued a bulletin on the nutria, remarking on the great 
persecution to which it had been subjected on account of the 
value of its fur and calling attention to the fact that in conse- 
quence it had now disappeared from many parts of Argentina, 
while recent droughts had augmented these losses. The export 
of the fur is now under governmental control, and already 
there are farms for breeding it. The report adds that the 
hunters are now their worst enemy. They hunt not only for 
the fur but also for the meat, which is palatable and is said 
even to have been served by a hotel in Goya. 

Numerous breeding farms have of recent years been estab- 
lished in Uruguay and have given excellent results. Devin- 
cenzi (1935) states, however, that "pitiless persecution by the 
professional hunters, without regard to season, sex, or size 
has so decimated the numbers in many sections of Uruguay, 
that where once it was abundant, it is now rare." Frank G. 
Ashbrook, summing up the situation, writes to Dr. Harper 
(1935): "The nutria is perhaps not in as great danger of ex- 
termination as the chinchilla, but strict conservation laws 


applied to this animal are necessary." There is the added 
circumstance that the nutria is much more prolific than the 
chinchilla, so that with reasonable opportunity of breeding it 
should maintain its numbers better. 

Live animals have also been exported to various European 
countries for breeding purposes and appear to have flourished. 
In France, where it is known as the "ragondin," the nutria 
was bred in captivity, at least as early as 1882, by Pays- 
Mellier at Champigny on Veude, and in 1888 by Edgard 
Roger in the park of the Chateau de Naudy (Seine et Marne) . 
In the two decades preceding the World War of 1914-18, 
there were apparently a number of persons breeding these 
animals in France, but the war seems to have put a stop to 
such enterprises until about 1925, when the raising of this 
animal was again taken up, especially in southern and south- 
western France. The nutria seems to adapt itself readily to 
conditions under captivity and reproduces normally. Eco- 
nomic conditions during the past ten years in France appear 
to have been unfavorable, however, for the hopes entertained 
for financial profit from this source and most of these breeding 
colonies have been given up. Many animals have escaped and 
established the species as an -element of the wild fauna, which 
may in time prove a valuable asset or the reverse. At present 
the capture of such wild individuals is subject to the hunting 
restrictions applied to other game (Bourdelle, 1939). 

Family DINOMYIDAE: Giant Rats 


Dinomys branickii Peters, Monatsb. Akad. Wiss. Berlin, 1873, p. 551 (Colonia Amable 
Maria, Montana de Vitoc, Andes of central Peru). 

SYNONYMS: Dinomys pacarana Ribeiro, Arch. Escola Sup. Agric. Med. Veterin., vol. 
2, pp. 13-15, 1919 (" Amazonas, Brazil "); Dinomys branickii occidentalis Lonnberg, 
Arkiv Zool., vol. 14, pp. 49-53, 1921 ("Gualea, Ecuador"); Dinomys gigas 
Anthony, Amer. Mus. Nov., no. 19, pp. 6-7, 1921 ("La Candela, Huila, Colombia"). 

FIGS.: Peters, 1873, pis. 1-4; Sanborn, 1931, pi. 5 (skull); Mohr, 1937b, figs. 1-9 

It is remarkable that this large rodent should continue to be 
so rare in collections that, although first described in 1873 on 
the basis of a specimen captured in the Andes of central Peru, 


it remained practically unknown thereafter until 1904, when 
Goeldi (1904) published an account of two living individuals 
sent to him at Para from the upper Rio Purus, Brazil. These 
were the first he had known of in 20 years' acquaintance with 
the fauna of the country. Another, perhaps from the same 
region, was described by Ribeiro in 1919 as Dinomys pacarana 
on the basis of its brown instead of black color; and two years 
later Lonnberg named as a new race, occidentalis, a specimen 
from near Gualea, Ecuador. The animal had meanwhile been 
taken in Colombia, near La Candela, Huila (J. A. Allen, 1916b), 
and this specimen was in 1921 made the type of D. gigas 
Anthony. In 1922-23, Edmund Heller secured a series of 23 
specimens from the natives about Buena Vista and Vista 
Alegre on the Rio Chinchao, and at Pozuzo, Peru, and pur- 
chased another at Manaos, Brazil. On the basis of the speci- 
mens available in North American museums, Sanborn (1931) 
reviewed the history of the species and showed that all the 
names given doubtless applied to but a single form. In 1937 
Dr. Erna Mohr (1937b) published an account, with photo- 
graphic illustrations, of a live one in the zoological gardens at 
Hamburg, with remarks on jireviously known specimens in 

From Sanborn's account it appears that this large and 
heavily built rodent is found in the Andean region from 
"central Colombia through Ecuador to central Peru, and east 
to the Rio Purus region of Brazil." Attaining a length of head 
and body of about 730-790 mm., with a tail of about 190 mm., 
it is of a black or brown color, with on each side of the midline 
two more or less continuous, broad white stripes, and on the 
sides two shorter rows of white spots. Apparently older 
animals have the stripes broader and more conspicuously 
white. The ears are short and rounded, the tail stout and 
cylindrical; the skull measures in length some 153 mm. (about 
6 inches) in the adult male, but the females are smaller, with a 
skull length of about 141 mm., and the bodily proportions 
correspondingly less. The incisors are disproportionally 
large, the cheek teeth small relatively, each showing three 
transverse enamel folds, with in the three last teeth a small 
additional posterior fold. 

The Tupi name, pacarana, signifies "false paca," since the 
size and color pattern recall those of the common agouti-like 


paca (Cuniculus). The present animal, however, is extraor- 
dinarily different in its stout, heavy appearance and thick 
tail of about half the body length. Goeldi describes the gait 
as "waddling" on account of the plantigrade position of the 
feet and the shortness of the legs, while the whole appearance 
"reminds one of an immense rat well advanced in develop- 
ment towards a bear." Of his captive animals Goeldi (1904) 
writes that they are "of a peaceful, phlegmatic disposition," 
exhibiting as a predominant trait "a combination of leisurely 
movements and supreme good nature . . . It is not easily 
irritated, and permits one to stroke and to scratch its head 
and back, and only occasionally manifests its displeasure by a 
low guttural growl. I have never yet observed a manifest 
intention to bite. When let out of the cage it makes no attempt 
to escape, and limits its excursions to an exploration of the 
immediate neighborhood in search of something to eat . . . 
This phlegmatic disposition seems to me to be a very precarious 
endowment for the struggle for life ; and considering the evident 
advantages which result to the smaller domestic rodents . . . 
from their nervously active constitution, it would not be 
strange if the species should tend to disappear. The apparent 
rarity of Dinomys may possibly find its explanation in the con- 
sequences of such a psychological endowment in a more 
nervous environment; but it is also possible that this rarity 
is because of the circumstance that the real habitat of the 
species has not yet been ascertained." Goeldi's captive 
female shortly gave birth to a young one and died in unsuccess- 
ful parturition of the second fetus, so that two young at a 
birth is probably normal, indicating a slow rate of increase 
(see also Tate, 1931). Dr. Mohr corroborates Goeldi's account 
of the slow and gentle behavior of the animal. 

Heller, who secured the series in the Field Museum, is the 
first to have gleaned much information as to the habitat of the 
species, and owes the skulls he secured to the habit of the 
natives of the upper Huallaga River of preserving the skulls 
of the game they kill, hanging these in their huts for good luck 
in hunting. He states (in Sanborn, 1931) : "It is not a fighter 
but merely fights as a last resort to save its life. It is slow in 
motion and can not turn about quickly, therefore it has no 
rear protection from alert foes like ocelots, tayras, coatis, etc. 
It therefore lives in rocky cliffs, or holes in the ground by 


preference, where it can back up and secure rear protection." 
In several months' stay in the region where it occurs he never 
met with one of these animals, though he was taken to their 
haunts by the natives, who use dogs to trail it. 

Because of its large size the pacarana is sought out by 
natives for food, while its lethargic manners and inoffensiveness 
make it easily vulnerable to its enemies. In habits it is likely 
to prove largely nocturnal, but captives seem to be active by 
day as well. Unless in some way given protection it is likely 
to be exterminated in the limited area where it occurs. A 
similar fate at the hands of the natives very likely in former 
days overtook its West Indian relatives, Elasmodontomys and 

Family CHINCHILLIDAE : Chinchillas 



Chinchilla lanigera Bennett, Gardens and Menagerie Zool. Soc. London, vol. 1, p. 1, 
Oct., 1829 (Coquimbo, Chile). 

SYNONYM: Eriomys chinchilla Lichtenstein, Darstellungen neue oder wenig bekannt. 
Saugeth., pi. 28 and text, 1830 (Chile implied) ; Chinchilla velligera Prell, Zool. 
Anz., vol. 108, p. 100, Nov. 1, 1934 (based on Bennett's description; hence Co- 
quimbo, Chile). 

FIGS.: Lichtenstein, 1830, pi. 28 (colored fig. of animal); Bennett, 1833, vol. 1, pi. 1, 


Chinchilla brevicaudata Waterhouse, Nat. Hist. Mammalia, vol. 2, Rodentia, p. 241, 
1848 ("Peru"). 


Chinchilla boliviana Brass, Aus dem Reiche der Pelze, vol. 2, p. 613, 1911 (Bolivia). 

The nomenclature of the chinchilla has been much confused 
owing to the fact that the naturalists of over a century ago 
believed that Molina's description of his Mus laniger applied 
to it. In 1830 Lichtenstein pointed out that Molina's animal 


was too small to be regarded as identical with the chinchilla, 
and himself proposed a new genus Eriomys and named the 
animal Eriomys chinchilla. However, in the year previous 
Bennett had already described the animal anew from specimens 
and given it the name Chinchilla lanigera, thus creating for it 
a special genus. Molina's name, Mus laniger, is now regarded 
as pertaining to a small, long-haired mouse, Abrocoma, so that 
Bennett's new name is not invalidated and becomes the correct 
term for the chinchilla. His account was based, he says, on 
two animals brought back to England by Surgeon Collie, 
who accompanied Captain Beechey on his exploring voyage 
around Cape Horn to the northwest coast of North America in 
1825-28. On the voyage out their vessel stopped at Concep- 
cion and Valparaiso, Chile, but on the return put in for a few 
days at Coquimbo, a short distance to the north of these ports. 
Here evidently were obtained two chinchillas, one of which 
survived the voyage and reached England alive, while the 
other died and was preserved as a skin and skull to serve later 
as the basis of Bennett's description. In the later account by 
Waterhouse (1848) this specimen is mentioned as in the British 
Museum. The living specimen seems to have been presented 
by Surgeon Collie to Lady Kriighton, who in turn gave it to the 
London Zoological Gardens, where it lived "for some time" 
and was "said to be from Coquimbo." It was perhaps the 
skeleton and internal anatomy of this individual that were 
described by Yarrell and Bennett. It seems clear that what- 
ever Molina's M us laniger was it could not have been a true 
chinchilla and that the first tenable name for the latter was 
applied to the animals brought back by Collie from Coquimbo. 

This is a stocky, short-limbed rodent, with a head and body 
about 9 inches long, and a well-haired, somewhat tufted tail 
about 5 inches long to the end of the vertebrae, beyond which 
the tuft projects about two inches. The ears are broad, about 
two-thirds the length of the head, prominent and oval. The 
fur, about an inch in length and extremely soft, is a beautiful 
gray strongly mottled with dusky or black above, passing into 
an "impure yello w- white " below. Feet dull white. Tail 
along the middle line above and below, as well as its tuft, 
black, with the sides dull white. Length of cranium about 2.5 

From time immemorial the thick fur of a delicate buffy-gray 


tint has been used in wearing apparel, first by the Incas and 
other native peoples of the southern Andes and later by Euro- 
peans. The former likewise utilized the long hair for weaving 
into cloth and the flesh was highly regarded as food (Ashbrook) . 
When the fur was introduced by dealers to the European 
trade, it became highly prized and much sought after. The 
price in recent decades has been so high and the animals 
themselves have become so reduced that naturalists have 
found it difficult to assemble sufficient series to determine the 
limits of geographic variation, so that the definition of races 
has been to this day unsatisfactory. A recent writer (Prell, 
1934) has attempted, however, to distinguish and allot names 
to three races, the Chilean, the Bolivian, and the Peruvian, 
but it can not be said that the characters of these are as yet 
sufficiently defined or their respective ranges traced. The 
name Eriomys chinchilla was given by Lichtenstein to a skin of 
a chinchilla obtained through fur traders and is said to be still 
in the Berlin Museum. A careful perusal of that author's 
account reveals nothing of the origin of the specimen beyond 
the fact that it was one of others traded through Carthagena 
and La Guayra, Venezuela. t)n the other hand, he implies 
that Chile is the home of the species, rather than Peru, as 
sometimes given, for in mentioning Molina's chinchilla or 
Mus laniger from Chile, he adds, "Die Uebereinstimmung des 
Namens, sowie der Fundort, machen es sehr wahrscheinlich, 
das damit unser Thier gemeint sei" (the correspondence of the 
name as well as of the locality makes it very probable that our 
animal is the same) . Moreover, the tail length of the specimen 
figured of half size, would be 6 inches, of the hind foot about 2.5. 
It was the largest of various skins at a fur dealer's. It thus 
seems most likely that Lichtenstein's name is a synonym of 
C. laniger a, published by Bennett very shortly before. This 
leaves the latter's Chinchilla brevicaudata as the first name 
applied unequivocally to the Peruvian chinchilla. Prell (1934) 
believes that the Bolivian chinchilla is also a distinct race and 
adopts for it the name boliviana given by Brass in a German 
treatise on fur. The ranges and characters of these races may 
be further outlined, as follows : 

The Chilean chinchilla is said to occur from about the Rio 
Chupa in latitude 32 S. northward along the western base of 
the Andes to the region of Copiapo in northern Chile. It is 


said by Gay to prefer the warmer areas along the coastal 
foothills and the inner valleys. It is thus somewhat of a low- 
land animal. In the fur trade it passes under the names of 
small, or coast, chinchilla, Chilean chinchilla, and bastard 
chinchilla, and its fur is the least valuable. 

The Peruvian chinchilla is described as somewhat shorter- 
tailed and larger in body (length of head and body, 14 inches ; 
tail vertebrae 3 inches). The general tint is more silvery than 
in the Chilean form, the back silvery gray with a clouding 
of blackish. Its range is said by Prell to be at higher altitudes 
on the western slopes of the coastal Cordillera of Peru at alti- 
tudes of 8,000 to 10,000 feet. In the trade this is known as the 
big chinchilla or royal chinchilla and is the most prized as fur. 

The Bolivian chinchilla has more rounded ears than the 
two others and is short-tailed like the Peruvian race, but in 
color it is with difficulty distinguished from the coast chinchilla, 
even by the fur dealers among whom it passes under the names 
Bolivian, La Plata, or Argentine chinchilla, after the ports 
whence it is shipped out. The range of this race is the eastern 
Andes and upper plateaus of Bolivia and northern Argentina, 
especially the provinces of Jujuy, Salta, Catamarca, and La 

According to Ashbrook (1928) the chinchillas are very 
swift in movement and in the early morning and late forenoon, 
when they are abroad, are shy, dashing at once to the shelter 
of their holes at the least alarm. Their curiosity, however, 
often prompts them to reappear shortly after, to see the cause 
of their fright. They generally feed actively early in the 
evening, sitting on their haunches and holding the food in their 
fore paws. They are fond of grains, seeds, fruits of shrubs, dry 
and green herbs, mosses and lichens. "Of all fruits, they seem 
to prefer the Algarrobilla, the seed of which is sweet and nutty, 
although the pod is astringent. The pods are found stored in 
their dens." 

Because of the extremely soft quality of the fur and its 
delicate tints of gray, the chinchilla is the most valuable and 
most sought after of all South American furs. Formerly so 
numerous that travelers in their haunts could see "thousands 
of them daily," they have now become so rare that in parts of 
the range, especially in Peru, whence the royal chinchilla 
comes, they are practically exterminated. Ashbrook (1928) 


states that till recently at least they were still common in 
Villenar Province, Chile, but are gone from the provinces of 
Antofagasta and Tacna. The countries in which these animals 
occur have of recent years realized the value of chinchillas as 
fur bearers, and not only have passed laws to safeguard them 
from extermination if possible but also have made some at- 
tempts to breed them in confinement for fur. Dr. Francis 
Harper through correspondence has assembled some notes on 
this subject, from which some of the following details are 
extracted: In the winter of 1923, M. F. Chapman succeeded 
in securing five male and seven female coast chinchillas and 
bringing them to California for breeding purposes. At the end 
of the third breeding season (1926) his stock had increased to 
65 animals. He found them easy to handle, for they quickly 
become tame, but they do not thrive in a damp atmosphere. 
In Chile captive animals have been found to produce two 
litters a year and are polygamous. The gestation period is 
about 111 days. Females will mate and commence breeding at 
four months of age, but it is inadvisable to breed them so 
young. Those brought to the United States have, after four 
years in captivity, raised even \hree litters a year. The num- 
ber of young at a birth averages two but varies from one to 
four (Ashbrook, 1928). Following Chapman's success with 
these animals, the stock was divided in order to establish 
other "ranches." In 1935 M. L. Weaver, who has had a share 
in raising this stock, wrote that "we have ranches at Logan, 
Utah; Idaho Falls, Idaho; Afton, Wyoming; and Madison, 
Wisconsin ; all doing well." He knew of several later shipments 
of live animals from South America, but none had been success- 
ful. In Chile, the home of this chinchilla, local attempts are 
being made to raise the animals and to develop an improved 
cross-breed, but efforts to hybridize the coast chinchilla with 
other forms have been unsuccessful. The former, though 
prolific, is less valuable than the Peruvian chinchilla, has 
coarser and less grayish fur, and is worth about half as much in 
price. Hitherto most of the efforts of breeders in the United 
States have been directed to building up a stock, some of 
which has been sold for fabulous prices to prospective breeders 
as royal chinchillas, which strictly they are not. There are 
several chinchilla farms in Chile, where the coastal form is 
still locally common. 


Concerning the Bolivian race, the so-called "Indian" or 
"Cordilleran" chinchilla, Carlos Garcia-Mata, of the Argentine 
Embassy, wrote Dr. Harper that in April, 1933, the Argentine 
Government started a chinchilla farm in Abra Pampa, Jujuy, 
at 12,548 feet altitude in the Andes, and has been very success- 
ful with it. From a start of 9 animals, in three years the number 
had increased by breeding to 61, an average per pair of 3.8 a 
year. It was planned to start selling breeding stock to local 
breeders by 1937. A decade before, the Argentine Government 
had made a similar attempt in cooperation with local hunters 
at the same place, but most of the stock was lost through 
ignorance of proper methods of care. They prove very sensi- 
tive to humidity and quickly succumb if kept in a damp place. 
At the present time there are stringent laws establishing close 
seasons and regulating the hunting of these animals. In 1926 
the hunting, exportation, transportation, and sale of chin- 
chilla skins were prohibited by law. In Bolivia, likewise, laws 
have been passed for its protection, and the exportation of the 
fur was prohibited under laws of 1920 and 1922. The hunting 
of chinchillas in Bolivia is said to be in the hands of a monopoly, 
but because of the scarcity of the animals very few are taken. 
In 1931, according to the American vice-consul at La Paz, 
there were no chinchilla farms in Bolivia. 

The royal chinchilla ("chinchilla real") of the Andes of 
western Peru is the most valuable of the three forms because of 
its longer and silkier fur and its pale "bluish" tint. The most 
valuable skins formerly came to market at Oruro or Tacna and 
Arica. This form has now been brought nearly to the verge of 
extinction. Not only were they persistently trapped and 
hunted by the natives for the fur trade, but it is said by Water- 
house (1848) on the authority of Bridges, who traveled in Peru 
a century ago, that they even trained a species of grison (of the 
weasel family) to hunt them by entering their burrows and 
capturing them as they endeavored to escape. Dogs also were 
used in hunting. In a letter to Dr. Harper in 1935, Carlos 
Garcia-Mata, of the Argentine Embassy, wrote that a few 
still remain in the steep and inaccessible rocks of the lower 
Andes, but that attempts to capture a pair for breeding stock 
have resulted in failure. Where 30 or more years ago they were 
common, and the pelts brought only $6 or $7 a dozen, by 1930 
they had become so scarce that pelts brought as much as $200 



apiece. However, as early as 1920, hunting and the sale of 
pelts as well as exportation were prohibited by the Peruvian 
Government, except under license. There continued to be 
nevertheless a small amount of illicit trade, but this is now 
apparently reduced to a minimum, for the animals are too 
scarce to make the trade remunerative. 

In 1931, William C. Burdett wrote to Dr. Harper, in response 
to inquiries, that "it is almost impossible to obtain accurate 
information in regard to the Peruvian chinchilla, since it is 
practically extinct. No one in this office (American consul- 
general's) has ever heard of a chinchilla ranch. As far as is 
known no live chinchillas have been seen in Lima for over 20 
years. The Peruvian center for the traffic in wild animals and 
their pelts is Sicuani, a mountain town near Cuzco in southern 
Peru, and the Indian dealers there report that they have not 
seen any specimens of chinchilla for many years." 

Formerly "most of the Chilean pelts were purchased by 
buyers in Coquimbo," and some statistics from the customs 
office of that city presented by Bidlingmaier (1937) are inter- 
esting. It seems that in 1905 the quantity sold amounted to 
18,153 dozen valued at from $100 to $110 (? per dozen). In 
the following year the number sold was about half that figure, 
and in 1907 it was again reduced to half that of 1906. In 1909 
it had dropped to 2,328 dozen, and the price had risen to $400 
to $500. European markets instruct their agents "to purchase 

Peruvian chinchilla (Chinchilla lanigera brevicaudata) 


skins at any price" thus increasing the greed of the "chinchil- 
lero," or professional native hunter, who redoubles his efforts. 
The relentless and systematic methods of destruction by the 
latter are recounted and the difficulties of exercising any effec- 
tive control by governmental agencies. Bidlingmaier suggests 
that perhaps the most feasible way would be to enact "stringent 
laws especially aimed at the traffic in contraband skins." 
Some attempts have been made to breed these animals in 
Chile, as in the vicinity of Vallemar and Copiapo, where their 
natural food plants are available, especially the "algarrobilla," 
but so far "nothing of great value has been revealed . . . 
Today there remain only five licensed criaderos (breeding 
farms), two of which are now presumed to be working in 
conjunction with a syndicate from the United States." 

Order CARNIVORA: Dogs, Cats, and Their Relatives 

The families of carnivores are discussed on page 134. Four 
groups occur in South America, the cats, the dogs, the raccoons 
and their relatives, and the bears. The bears occur only in the 
Andean Mountains, while the raccoon family developed in this 
continent. The bear and the Falkland fox, or wolf, are the 
only carnivores considered in this section. J. E. H. 

Family URSIDAE: Bears 



[Ursus] ornatus F. Cuvier, in E. Geoffrey and F. Cuvier, Hist. Nat. Mammiferes, 

vol. 3, pt. 50, p. 2 and pi, June, 1825 (Cordillera of Chile). 
SYNONYM: Ursus frugilegus Tschudi, Fauna Peruana, pp. 11, 90, 1844 (Peru, probably 

near Lima). 


Tremardos ornatus majori Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 9, p. 216, 1902 
("Southern Ecuador, probably the province of Azuay"). 

SYNONYMS: Ursus ornatus thomasi Hornaday, Bull. New York Zool. Soc., no. 45, p. 
748, 1911 ("Andes of southern Colombia"); Tremarctos lasallei Maria, Bol. Soc. 
Colombiana Cien. Nat., vol. 13, no. 76, p. 115, Aug. 1924 ("Arauca, Boyaca 
Province, Colombia"). 

FIGS.: Geoffroy and Cuvier, 1825, pi. 50; Hornaday, 1911, pp. 747-748, 4 figs, (photo- 


This is the only member of the bear family living in South 
America, where as a species it occurs from the Andes near the 
Venezuela-Colombia boundary, across Colombia and Ecuador, 
to Peru, northern Chile, and Bolivia. Although several names 
have been given to specimens of the species, the supposed 
differences appear to rest mainly on individual variations. 
Thomas (1902) found that skulls from Ecuador differed from 
the typical southern skulls in being larger, longer in proportion, 
and more slender, and the teeth "rather larger throughout." 
He therefore named these latter specimens as a northern race, 
majori, but even yet it is uncertain what are the limits of 
variation in the more northern bears, or how the ranges of the 
two races should be drawn. As long ago as 1844, Tschudi dis- 
tinguished the Peruvian animal as Ursus frugilegus on account 
of the supposed shorter head, shorter soles of the feet, and 
much slenderer body, but the differences appear again to be 
of an individual nature, and the Peruvian animal is currently 
believed to be inseparable from that of Chile. In 1911 Horna- 
day casually bestowed the name Ursus ornatus thomasi on a 
bear of this species from the Andes of southern Colombia 
living in the New York Zoological Gardens, but, again, the 
supposed diagnostic character, the lack of white markings, is 
now known to be individually variable ; and Maria's Tremarctos 
lasallei from Arauca, Colombia, based on a mounted skin, 
having longer muzzle and claws than typical majori, can 
hardly be other than a synonym of the latter. 

In coloring and pattern both northern and southern races 
are apparently alike. The entire body is uniformly black or 
blackish brown, except that usually there is a narrow line of 
white beginning on each side about halfway between the ear 
and the eye, continuing forward nearly to the bridge of the 
muzzle in front of the eye, then turning downward across the 
cheek to the midline of the throat, and continuing parallel 
with the corresponding mark of the other side, to the chest. 
This marking, which from the semicircle about the eyes, gives 
the bear its English name, shows, however, considerable varia- 
tion. Its upper part may be washed with pale yellowish; 
again, the muzzle may be light tan, or more extensively 
whitish, to include the forehead, nose, cheeks, and throat. 
As an opposite extreme, the white markings may be lacking 
altogether in animals from the same region. Cuvier mentions 


the total length as about 3 feet in the original specimen, height 
at the shoulder about 20 inches. Thomas's measurements for 
the type specimen of the northern race are: Head and body, 
1,625 mm.; hind foot with claws, 210; ear, 75; greatest length 
of skull, 263 mm. (in one of the typical race, 231, a male in 
each case); zygomatic width, 169 mm. (in typical race, 163). 

Various writers reiterate that very little is known of this 
bear beyond the fact that it seems to be rare throughout its 
range and is seldom seen or hunted. This fact and its restricted 
range may warrant its inclusion among species in a precarious 
situation. The specimen on which Frederic Cuvier founded his 
description and of which he published a colored figure, was said 
to have been brought from the Cordillera of Chile, and it 
doubtless did come from that region, probably in northern 
Chile. It is also known in Bolivia, where as well as in Peru, it 
is called "hucamari" in the Quichua tongue. Krieg (1931) 
mentions that it is reported from the Bolivian (?) Chaco, and 
he obtained a skin from near Caraparisito. There it was said 
to be exceedingly rare and very shy of men, though occasionally 
accused of killing cattle. The individual exhibited in the New 
York Zoological Gardens in 1911 was from near Quito, Ecuador, 
and it seems likely that the center of abundance now lies in 
southern Colombia, Ecuador, and parts of Peru. 

Osgood (1912, 1914), who made special efforts to learn 
something of the habits of this bear, writes of securing an adult 
female and wounding a male accompanying it, in the moun- 
tains about 10 miles northwest of Menocucho, east of Truxillo, 
northern Peru. The region inhabited by this bear is here 
extremely arid and mountainous, with a scanty vegetation 
consisting mainly of cacti and small thorny bushes. The 
mountains range from 1,000 to 5,000 feet in height and are not 
greatly different in character from the desert plain stretching 
westward to the sea. The principal food of the bears seemed to 
be a pear-shaped fruit with a hard outer shell enclosing numer- 
ous seeds, a species of Capparis locally known as "chapote." 
"The region is for long periods almost waterless and animal 
life is very limited . . . From reports received from local 
sources, it is evident that bears are fairly common in numerous 
localities in the arid region." The two seen by Osgood were 
feeding at midday in the full glare of the tropical sun, but the 
natives advised that the early morning was the best time for 
hunting them and they use dogs in the pursuit. 


Quite different is the type of country inhabited by these 
bears in southwestern Ecuador, where Tate (1931) found them, 
and farther northeast in the Paramo de Tama near the Colom- 
bia-Venezuela boundary, where, writes Osgood (1912, p. 58), 
they are very seldom seen and are decidedly rare. In four 
weeks' hunting he found only a fragment of dung and no tracks; 
"natives say the bears live almost exclusively in the forest and 
it is only on the very rare occasions when they wander out into 
the cultivated clearings that they have been killed." Tate, in 
company with a native hunter, found traces of an adult pair in 
the rain forest on a large plateau in southwestern Ecuador, on 
the Andean slope at about 4,000 feet altitude. He followed 
the tracks a long distance up the slopes, and came upon six 
places where the bears had "bedded down." He saw also 
where they had broken down many trees 2 inches in diameter 
and "were feeding on the seeds of a palm called "pambili" 
trees from 80 to 100 feet high. They evidently climbed the 
trees and brought down the whole fruit-stalk, which looks 
somewhat like that of the royal palm. Numbers of the trees 
had been climbed, some of them several times, since they had 
both old and fresh claw marksX Other food was secured by 
breaking down young palms, tearing open the green stalk and 
eating the unopened leaves in the interior. According to 
Olalla [a skilled collector], in the region about Quijos in eastern 
Ecuador bears appear at a certain season of the year upon the 
mountain sides to feed upon the ripe fruit of a certain tree," 
but the statement that they make large nests of sticks in the 
tree tops for sleeping purposes, as Olalla told him, seems open 
to question. 

Nearly a century ago Tschudi (1844) gathered together many 
notes on this bear in Peru, from which the following are culled. 
He transcribes a few interesting points from the accounts of 
the early Spanish explorers. Thus Ulloa said that the bear was 
common in the provinces of Guijos, Macas, and Jaen de Braca- 
moros. It was sometimes lassoed from horseback. It was 
also found in the forests east of La Paz, Bolivia. According 
to the account of Garcilasso de la Vega, it was rare in Peru, a 
fact that he attributed to the method of hunting. Great 
annual drives were held in which the Incas used as many as 
25,000-30,000 Indians, whose lines would cover 20 to 25 
leagues, converging toward a funnel into which the game of the 


enclosed area was driven. As the circle became smaller, the 
lines of Indians would become deepened by the addition of 
their number on the outer edge of the line, to prevent animals 
from breaking through. When the enclosed animals were 
finally penned, the carnivores were all killed, but of the deer, 
vicunas, or other such game, only a certain number of males 
and old females were slain, the others freed. Tschudi had seen 
similar drives in his time, but on a smaller scale, and agrees 
that the number of bears thus captured or killed stands in very 
small proportion to the number of other large carnivores, so 
that their apparent scarcity is probably an actual one. He 
doubts if the bear that served for Cuvier's description and 
figure really came from the Chilean Cordillera but suggests 
that the only likely harbor on the west coast of South America 
from which such an animal might be shipped was Truxillo. 
In distinguishing his supposed new species, "frugilegus," from 
ornatus, Tschudi stresses the reports that while the latter 
preys upon young deer, vicunas, and huanacos, the former is 
chiefly a vegetarian and often does much damage in plundering 
the maize fields of the natives. The available evidence does not 
indicate that this bear is much of a predator but finds abun- 
dant food in the way of fruits', leaves, or roots. 

Family CANIDAE: Wolves, Dogs, Foxes 



Canis vulpes australis Kerr, Linnaeus's Animal Kingdom, p. 144, 1792 (West Falkland 

Island; see Osgood, Journ. Mamm., vol. 1, p. 35, 1919). 
SYNONYMS: Canis antarcticus Bechstein, Uebers. Vierfiiss. Thiere Pennant, vol. 1, p. 

271, footnote, 1799; Dusicyon antarcticus Thomas, Ann. Mag. Nat. Hist., ser. 8, 

vol. 13, p. 353, 1914. 
FIGS.: Mivart, 1890, pi. 8 (col.); Pocock, 1913, figs. 70B, 71 A, 73, 74B, D (skull and 


The Falkland Island fox, although often called a wolf, is not 
a wolf at all and is not even closely related to the North 
American coyotes, as Huxley formerly supposed. Instead it 
is a near relative of the group of South American foxes, which 
are now regarded as distinguishable under the generic title 
Dusicyon. How this animal reached the Falkland Islands 
will doubtless ever remain a matter for speculation. 


Foxlike in appearance, this animal stands 15 inches at the 
shoulder. Its coat is thick and soft, lacking coarse long hair 
and having a bushy tail. Pocock (1913) describes a specimen 
as having the prevailing color of the body brown "relieved by 
fine speckling due to the narrow pale band on the individual 
hairs." Below, the color is brownish, except that the posterior 
part of the belly and upper end of the throat are white, the 
chin and lower jaw white with a fuscous tint. There is a 
marked fuscous patch above the hock of the hind leg. The tail 
on its basal two-fifths is like the back, the middle part black 
and the tip white. The ears, according to Pocock, are unusu- 
ally small. The skull is characterized among other things by 
the lyrate sagittal area and truncated instead of pointed occip- 
ital crest. In these respects it agrees with some other of the 
South American "foxes." Mivart (1890) gives the following 
measurements : Length of head and body, 970 mm. ; tail, 285 ; 
hind foot, 180; ear, 65; skull length, 110. It is said that the 
animal formerly occurring on the East Falkland Island was 
smaller and redder than the one on West Falkland. 

The history of the Falkland fox has been several times writ- 
ten, most recently by Renshawtl931). He points out that the 
animal was first discovered in January, 1690, on South Falk- 
land by Strong's party, who captured one alive and kept it for 
several months on their ship. Unfortunately the vessel 
finally had occasion to discharge its guns, which so startled the 
fox that it leaped overboard and perished. In his voyage of 
1763-64, the French navigator Pernetty again found the 
species, and the discovery of it had been credited to him by 
writers, until Strong's earlier account was brought to notice. 
In 1765 Commodore Byron took possession of the islands for 
Great Britain, and his landing party records that several of 
these foxes came to meet the men, wading out toward them in 
curiosity at the strange apparition. The sailors, however, 
believing the "wolves" were actuated by ferocity, at first 
retreated, but later found that the animals were quite without 
fear. The animals were so numerous that the men set fire to 
the long grass and presently saw great numbers of them run- 
ning to escape the flames. Byron brought a live one back to 
England, and this individual was later described by Pennant 
and thus became the basis of Kerr's name. 

When Darwin during the voyage of the Beagle stopped at 



the Falklands in 1833, the animal was still common there, but 
its absurdly tame habits, he foresaw, would lead to its early 
extermination. Indeed, he mentions that the Gauchos would 
capture them by holding out a bit of meat to a fox in one hand 
and stab the animal with a knife held in the other, when the fox 
came within reach. 

Such an abundant and soft-furred animal, thus easily caught, 
attracted the notice of John Jacob Astor, then active in the 
fur trade, who in 1839 sent men to the Falklands to collect 
pelts, and great numbers of them were taken. Hamilton 
Smith mentions seeing quantities of them in Astor's warehouse 
at New York. Renshaw even says that the extermination of 
the species from East Falkland may date from this exploita- 
tion; at all events by 1863 they were already extinct in the 
eastern part of East Falkland. When, later, the Scotch set- 
tlers started raising sheep on the Falklands, the foxes seem to 
have developed a taste for mutton and would kill sheep by 
attacking one or two or three together. As a result a poisoning 
campaign was undertaken and many were destroyed. In 1870 
Byng wrote to the Zoological Society that they were almost 
exterminated, and the last one is said to have been killed in 
1876 at Shallow Bay, West Falkland. 

Antarctic wolf (Dusicyon australis) 


The Antarctic "wolves" were said to feed on various native 
birds, harrying the penguin colonies and driving the upland 
geese to nest on small islands off the coasts. Seals were eaten 
too. Their extreme tameness may have been a result of long 
isolation and lack of contact with man, but their failure to 
develop any wholesome fear of him may have been in part a 
result of the use of such silent weapons as bolos and knives 
and probably traps, rather than firearms. 

Captain Fitzroy of the Beagle and Darwin in 1836 brought 
back four of these animals, two of which are still preserved as 
specimens in the British Museum, which has a skeleton in 
addition. The Royal College of Surgeons had two skulls 
which may now be in the British Museum, for according to 
Pocock (1913) there are five crania in that institution. Pocock 
adds that the other known material representing this animal 
is in Paris, but he does not tell of what it consists. Renshaw, 
however, states that the Leiden Museum has three specimens. 
It was first exhibited by the Zoological Society of London in 
1845. Twenty years later, in 1868, a pair was again sent to 
the Society, but one only survived the journey. Again in 
1870, a pair was sent by Byng, of which the male died on the 

Order PERISSODACTYLA: Odd-toed Ungulates 

This order is represented by three Recent families: 

(1) Equidae, the horses, asses, and zebras. The wild horses 
are Eurasian, but all except a few individuals of the Przewalski 
horse of Mongolia have become extinct. 

(2) Rhinocerotidae, the rhinoceroses of southern Asia and 

(3) Tapiridae, the tapirs, with nose and upper lip produced 
into a short proboscis, are found in Central and South America, 
the Malay Peninsula, and part of the Malay Archipelago. 

The odd-toed ungulates are represented in South America 
only by the tapirs. The mountain tapir is poorly known and 
is thought to be endangered by the agricultural development 
of the northern Andes. J. E. H. 


Family TAPIRIDAE: Tapirs 


Tapirus roulinii Fischer, Synopsis Mammalium, Addenda, p. 602, 1830 ("Summos 

monies Andes Americae australis")- 
SYNONYMS: Tapirus villosus Wagler, Syst. des Amphib., p. 17, 1830; Tapirus pinchacus 

Blainville, Osteograph., Unguligrades, Genus Tapirus, pi. 3, fig., 1845; Tapirus 

leucogenys and enigmaticus Gray, Proc. Zool. Soc. London, 1872, pp. 488, 490 

(Assuay and Sufiac, Ecuador). 
FIGS.: Gray, 1872a, pi. 21, fig. 1; pi. 22, fig.l (col. fig. of young and subadult) ; Sclater, 

1878, pi. 39 (col., adult); Hatcher, 1896, pi. 4, figs. 2, 2a; pi. 5, fig. 2 (skull). 

Very little seems to be known of the habits and present 
status of this tapir of the northern Andes. It was first brought 
to the notice of naturalists by M. Roulin, who communicated 
to Baron Cuvier an account of the animal, which was published 
by Cuvier in the Annales des Sciences Naturelles in 1829 and 
further elaborated by Roulin in the following year in the same 
journal. Roulin gave the tapir a French name, "Tapir pinch- 
aque" from the name used by the native Indians to denote a 
fictitious monster, but the Latin form was not used. In 1830, 
both Fischer in the supplement to his Synopsis Mammalium 
and Wagler in his System des Amphibiens, etc., proposed new 
Latin names based on Roulin's description, but since it is not 
evident which had precedence, it seems better to follow Thomas 
(1880) in adopting Fischer's name, which commemorates the 
discoverer of the species. 

This tapir does not apparently differ in size from the lowland 
species of the Amazon Valley, which is about as large as a pony. 
The hair, contrary to the usual belief, is said by Sclater (1878) 
to be rather short, the individual hairs about an inch long, and 
the color nearly uniform black, shading to brownish; outer 
edge of the ears and a spot at the corner of the mouth white. 
Iris light bluish hazel, rather than brown as in the lowland 
tapir. The form of the nasals is very different from that of 
the latter, being long, and tapering, with concave outer borders. 
The brain case is shorter and the sagittal crest less high. 
Measurements given by Roulin are : Tip of snout to tip of tail, 
5.5 feet; height at shoulder 2 feet 9 inches, but these are per- 
haps not of a fully adult animal. Greatest length of skull (M. 
C. Z.), 380 mm.; median length of nasals, 95. 

The home of the mountain tapir seems to include the high- 


level forests from central Colombia to Ecuador and possibly 
northern Peru. Most of the specimens seem to come from the 
Andes of Ecuador. Its discoverer, Dr. Roulin, obtained his 
specimens from the Paramos of Quindiu and Suma-Paz, during 
his residence at Bogota, and it was met with by Goudot on the 
peak of Tolima between 1,400 and 4,400 meters, in south- 
eastern Colombia. P. L. Sclater (1870) quotes a letter from 
Robert B. White dated from Popayan, Colombia, June 8, 1869, 
in which he speaks of finding this tapir on the central Cordillera, 
in the region of the volcano of Purace, adding that "they are 
very shy, and I have not been able to get near them, but have 
seen them at a distance of half a mile, with a telescope, bathing 
themselves in a small lake. I have also seen the skins occa- 
sionally brought in by the Indians . . . It is never found 
at a lower elevation than 3,500 metres above sea-level 
and it exists up to 4,200 metres. These animals are rarely 
killed, because the skin only sells for" 3 shillings. 

In the Santa Marta region of northern Colombia, there 
are tapirs in the somewhat isolated Santa Marta Range, but 
judged from a skull from Dibulla, in that region, the species 
there is T. terrestris. Farther s\mth, however, in the Depart- 
ment of Santander, M. A. Carriker, Jr., informs us that tapirs 
are rather common in the southern part of the area at altitudes 
from 8,000 to 10,000 feet, and are much hunted by the natives. 
Owing to clearing of the forests for cultivation, the tapirs are 
likely in time to become driven out or much depleted in 

Thomas (1880) mentions that a Mr. Buckley secured a series 
of 15 specimens of this tapir at Sarayacu, Ecuador, but "un- 
fortunately before the skins were prepared, a troop of revo- 
lutionary soldiers put in an appearance and cut off the hoofs of 
every specimen to make into amulets," which so vexed Mr. 
Buckley that he abandoned the entire lot! 

Mountain tapirs were first exhibited in the London Zoologi- 
cal Gardens in 1878, according to Sclater (1878), who probably 
did not then regard as the same the two youngish ones from 
Ecuador that Gray in 1872 named enigmaticus and leucogenys. 
There is a mounted skin and a skeleton from Ecuador in the 
Museum of Comparative Zoology, and there are two speci- 
mens in the Academy of Natural Sciences of Philadelphia 
taken in the Uanganatis Mountains on the headwaters of the 
Curaray River, Ecuador, at 14,000 feet, in 1935-36. 


Whether the range of this tapir extends to Peru, or formerly 
did, is uncertain. Tschudi (1844), however, wrote that the 
presence of this species seems almost certainly to be indicated 
on the testimony of the natives that a tapir is to be found in 
the eastern slope of the inner Andes at 7,000-8,000 feet, 
especially in the Ceja region. No one seems to have verified 
this supposition, nor is there enough available information on 
the species at hand to give any idea of its present abundance. 

Order ARTIODACTYLA: Even-toed Ungulates 

This order is represented in South America by two families, 
the camel group and the deer; these are discussed on page 256. 
The wild South American camel-like mammals, the guanaco 
and the two races of the vicuna, are in need of protection from 
hunters, chiefly natives. Four species of deer, representing 
three endemic genera, are rare and appear to be decreasing in 

Family CAMELIDAE: Camels, Llamas 


Camelus huanacus Molina, Saggio Storia Nat. Chile, vol. 1, p. 317, 1782 (probably the 

Chilean Andes). 

SYNONYM: Auchenia llama Water-house, Zool. Voyage Beagle, Mamm., p. 26, 1839. 
FIGS.: Prichard, 1902a, pi. opp. p. 160; Cutright, 1940, pi. opp. p. 119 (photograph). 

The wild huanaco, or guanaco, is believed to be the original 
source of the domesticated llamas and alpacas of the Peruvian 
aborigines, and with the smaller vicuna it is the only living 
member of the camel family in America. The family seems to 
have originated in North America, becoming much diversified 
in middle and late Tertiary times and spreading to the Old 
World and to South America. It has since completely died out 
in North America. 

The guanaco has somewhat the slender build and long neck 
of the camel, but the ears are proportionately long, the tail is 
short and bushy, there is no "hump," and the coat is soft and 
woolly. In color the wild species is a dark fawn-brown above, 
with a blackish face and white under surfaces. It stands about 
3 feet 7 inches high at the shoulder, and the skull is about 11.5 


inches in basicranial length. The upper incisor and canine are 
lancet-shaped and capable of inflicting a bad wound, but the 
lower incisors are procumbent. There are callosities on the 
inner side of the fore limbs. 

The range is from the Andes of south-central Peru south- 
ward to Patagonia, where it comes to the lower altitudes, and 
thence to Tierra del Fuego. Very likely, when sufficient 
series of specimens are available, more than the single geo- 
graphic form may be distinguished. Indeed, Lonnberg (1913) 
has described as Lama huanachus cacsilensis, an animal of 
small size from Cacsile, Nunoa, Peru, which seems to be more 
like the vicuna and according to Osgood (1916) is "not closely 
related" to the common guanaco. Its status may await 
further study. 

Osgood (1916) states that the herds he saw on the Pampa de 
Arrieros, between Arequipa and Puno, at the northern part of 
the animal's range, are "almost if not quite the northernmost 
now existing." Inhabiting as they do the alpine zone between 
14,000 and 18,000 feet above the sea, they are perhaps also 
the most lofty-living of the species. Barren though these 
heights appear, the animals obtain pasturage sufficient to sus- 
tain existence. Though little hunted by white man, they are 
persecuted by the Indians, who shoot them from blinds 
erected at waterholes. The skins are used in making beautiful 
rugs or for saddle cloths, while the meat is of excellent quality. 
B. T. Colley, writing to Dr. Francis Harper from Oroya, Peru, 
in April, 1934, states that to the south of the Puno region, and 
beyond the Atacama Desert, the guanaco is rather plentiful, 
and he has seen herds of over a hundred animals on the con- 
tinental divide above Santiago, while farther to the south 
there are many more. Here, as winter sets in, they move down 
from the higher regions to lower altitudes, retiring chiefly to 
the Argentine side because of the abruptness of the Chilean 
Cordillera. They have a characteristic habit of making zigzag 
trails in which the angles tend to be equal and regular. 

In Chile, Jose M. B. Toro reports (in litt., 1934), the guanaco 
has become so much reduced in numbers in the past 20 years 
or more because of persecution that in 1929 a decree was 
passed prohibiting the hunting of it for three years, after 
which the close time was extended for two years more to 
December 1, 1935. At the present time there are still a few 


scattered bands to be found in the steeper parts of the Cordil- 
lera and along the Argentine border, but any close estimate of 
their numbers is difficult. 

On the pampas of southern Argentina and over most of 
Patagonia the guanaco seems to be still common from the Rio 
Colorado in about latitude 40 S., and is even said to have 
increased within recent years. 

Darwin has given a brief account of the abundance and 
habits of the guanaco as he saw it a century ago in the Santa 
Cruz region of Patagonia. They go in small herds of from half 
a dozen to thirty, but he mentions one herd of "at least five 
hundred." He speaks of their taking readily to water and 
swimming to islands near the mainland. They have a certain 
curiosity when alone but in herds easily become bewildered and 
stampeded, a fact of which the Indians take advantage in 
killing them. Both Darwin and Prichard (1902a) mention 
coming upon places where great numbers seem to have perished 
and left bones bleaching on the ground. Prichard was told 
that in the winter previous enormous numbers of guanacos had 
sought Lake Argentine and perished there of starvation. "In 
the severities of winter they seek drinking-places where there 
are large masses of water likely to be unfrozen. The last few 
winters " had been so severe that great havoc had been wrought 
among the animals. Prichard crossed Patagonia from the Rio 
Chubut southwesterly to the Andean foothills, and thence 
proceeded southward, coming out at Punta Arenas on the 
Straits of Magellan. Over most of this country guanacos 
were common, and in regions where they were not much hunted 
were not very shy. 

Apparently the main danger to which this species may be 
exposed, apart from the decimating effect of severe winters, is 
the extension of grazing by the cattle and sheep of the settlers, 
and the exploitation of the guanacos for their hides and meat. 
As to the former, a writer in an Argentine journal (La Chacra, 
1936) states that in 1913 ranchers in Santa Cruz pleaded for 
the destruction of the guanaco on the ground that it was a 
detriment to sheep raising and a national plague. Wire fences 
used to enclose ranges prove the ruin of many guanacos, 
victims of cold and hunger. On the other hand, the animal 
does not interfere with sheep raising in parts farther south that 
are unsuitable for sheep. It is much persecuted for its hide 


and meat. In the report of the world's fur production for 1928 
(Journ. Soc. Preserv. Fauna Empire, pt. 12, p. 64, 1930) the 
number of guanaco skins brought to market is given as 300,000. 
While at present it can hardly be said that the species is in 
danger of extermination, nevertheless the demand for hides 
to be used as robes, and the encroachment of grazing and 
hunting, will doubtless much restrict its area and its numbers 
in years to come. 


Camelus vicugna Molina, Saggio Storia Nat. Chile, vol. 1, p. 313, 1782 ("Probably 
Peru" Lydekker; but Molina says in the Cordillera of the provinces Coquimbo 
and Copiapo). 


Lama vicugna mensalis Thomas, Smithsonian Misc. Coll., vol. 68, no. 4, p. 3, Apr. 10, 

1917 ("Incapirra, Junin, Peru"). 
FIGS.: Tschudi, 1844, pi. 17 (col.); Royal Nat. Hist., vol. 2, p. 412, fig., 1894. 

The vicuna resembles the guanaco in general form but is 
about a fourth smaller, somewhat slenderer, and paler in color, 
a pale fawn, without black on the face. The fore limbs have 
no callosities such as are present on the inner side of the 
"knees" of the guanaco. Although formerly placed in the 
same genus with the latter, Miller (1924c) regards it worthy of 
generic distinction, since the lower incisors are peculiar in 
being long, slender, and ever-growing from persistent pulps, 
like those of rodents, but with the enamel on the inner side. 
This condition is unique among living artiodactyls but recalls 
that of the dwarf wild goat (Myotragus) , the remains of which 
are found in Pleistocene cavern deposits of the Balearic Islands. 
Osgood (1916) gives the following measurements of a Peruvian 
specimen: Length from between ears to root of tail, 1,250 mm.; 
base of ear to point of shoulder, 670. The height at the shoul- 
der, according to Lydekker, is about 2 feet 9 inches; Tschudi 
(1844) gives 2 feet 6 inches. Length of skull, about 220 mm. 

Although Lydekker says that the "typical locality is prob- 
ably Peru," Molina's account, on which the name is based, 
states that they are found in the parts of the Cordillera belong- 
ing to the provinces Coquimbo and Copiapo, but keep to the 
steep summits of the mountains, minding neither snow nor ice. 


Whether or not the vicuna did occur in Molina's day, 1782, as 
far south as the Cordillera of Coquimbo, in Chile, it apparently 
no longer does so, but one may regard the type locality as the 
latter region on Molina's authority. Northward the range 
extends to northern and central Bolivia, Peru at high altitudes, 
and southern Ecuador. Thomas has named, as a distinct race, 
mensalis, the northern animal, basing his description on speci- 
mens from Incapirra, Junin, Peru. The characters claimed are 
the more strongly fulvous color and slightly smaller size and 
smaller teeth, as compared with true V. vicugna, the type 
locality and size of which are not indicated. The range of this 
race is given as "Peru and Bolivia" and is assumed to include 
also southern Ecuador at high altitudes. This would leave 
typical V. vicugna as the form of Chile, where it is possibly now 
extinct. The two races may be considered together. An adult 
male measured from "between ears to root of tail," 1,250 mm.; 
base of ear to point of shoulder, 670, or more than in the guana- 
co. The skull of the type of mensalis, a male, had a greatest 
length of 240 mm.; length of molars, 45. 

The vicuna is apparently at the present time gone from 
Chile, but the British Museum has a mounted skin from Cata- 
marca, northwestern Argentina. In Bolivia it is found hi a 
restricted area in the north-central part. In Peru, its chief 
centers of abundance are said by Maccagno (1932) in his 
recent monograph to be: Junin, Huanta, Ayacucho, Puno, 
Cuzco, Apurimac, Huancavelica, and Arequipa. 

These animals live on the vast plains at altitudes between 
11,500 and 18,500 feet in Peru, going in small droves of 10 to 
15 females and an adult male. In general they are said to be 
easily tamed, and often one or two may be seen about ranch 
houses, but in confinement they do not breed readily. On the 
other hand, if allowed practical liberty under fence with 
sufficient area, they are said to breed freely and may be caught 
and sheared for their wool, which is much sought for its fine 

In Peru the vicuna was accorded legal protection as long 
ago as 1825, but the law apparently remained a dead letter 
until the animals had become much reduced through hunting. 
Finally, on October 8, 1920, a decree was passed prohibiting 
the making of goods from vicuna wool and forbidding the sale 
of skins. A heavy fine was provided for infractions. As a 



result it is said on good authority that in Peru the vicuna is 
fast recuperating and is now locally common as in the Junin 
area (William C. Burdett, in litt.). Nevertheless it is difficult 
to enforce these laws and many of the animals are killed. 
In Bolivia the exportation of vicuna wool and hides is pro- 
hibited under laws of 1920 and 1922. Vicuna rugs are often 
manufactured in Peru, but their export is forbidden except that 
a person leaving the country may take one out by securing an 
official permit. The robes are made of small strips of the hide 
perfectly matched in color and texture. Several grades are 
made by using the skin from different parts of the body, the 
back and sides for one type, the neck for another, and the legs 
and belly for a third. The most valuable are those made from 
the back and sides, and some of the best may sell for as high as 
$100 (Carriker). It is believed that it may eventually be 


Vicuna (Vicugna vicugna) 


feasible to raise vicunas on ranches and shear them for the 
wool, which brings a good price, as much as $5 a pound. 
This would require special sanction of the government con- 
cerned. Like the llama and alpaca, this animal is subject to 
such parasitic infections as lungworm, scab, and flukes, particu- 
larly at lower altitudes (B. T. Colley, in Hit.). 

In earlier days the native Indians made much use of this 
species. Tschudi (1844) describes how they carried out ex- 
tensive drives, gradually working the animals into a narrow 
funnel-shaped place among the rock walls, where many were 
captured, their wool sheared, and the captives then freed. 

Family CERVIDAE: Deer 


Pudua mephistopheles de Winton, Proc. Zool. Soc. London, 1896, p. 508. 
FIGS.: De Winton, 1896, pi. 19; Lydekker, 1898, pi. 24, fig. 1. 

This small deer stands about 14 inches high at the shoulder 
and is peculiar in its short metapodials, very short spikelike 
antlers in the male, short ears, and practical absence of an 
external tail. Very little is known of it, but since its habitat is 
restricted and is likely to be further limited by various develop- 
ments in the future, it may deserve mention here. 

The general color is a rich brown, due to a blackish-brown 
ground color, sprinkled with bright rufous. Ears relatively 
short, with long white hairs lining them. Face and legs nearly 

Originally made known from a specimen taken on the paramo 
of Papallacta, Ecuador, very few examples of this deer have 
since found their way into collections. The Swedish consul 
Soderstrom sent a specimen to the British Museum and one to 
the Royal Museum at Stockholm, the latter with antlers about 
78 mm. long. All these are from an altitude of about 12,000 
feet at Papallacta, outside of which the species is unknown. 
It is believed that even here it is uncommon. 




Cervus antisiensis d'Orbigny, Ann. Mus. Hist. Nat. Paris, vol. 3, p. 91, 1834 (Andes, 

near La Paz, Bolivia). 
SYNONYMS: Anomalocera huamel Gray, Scientific Opinion, p. 384, 1869; Xenelaphus 

chilensis Gray, Ann. Mag. Nat. Hist., ser. 4, vol. 12, p. 61, 1873. 
FIG.: Lydekker, 1898, pi. 23 (col.). 

This small deer is confined to a rather restricted alpine 
habitat in the Andes and may be included here as a game 
animal that may need more protection. Somewhat smaller 
than a Virginia deer, having a coarse brittle coat and lacking 
metatarsal glands, this species is of a nearly uniform speckled 
brown and buffy, with a darker line on the forehead. Tail dark 
brown on base and most of the upper surface, but its tip and 
lower side are white. The antlers are present in males only, 
and consist of a short fork commencing close to the burr. 
Height at shoulder about 34 inches (Lydekker). 

In the Andes of Ecuador, Peru, Bolivia, and northern Chile 
this species is found at high altitudes, mainly between 14,000 
and 16,000 feet but at times lower. Lydekker in 1898 wrote 
that it was abundant in Ecuador on Chimborazo, Pichincha, 
and Cotapaxi, but Richardson in 1912-13, collecting for the 
American Museum of Natural History, failed to obtain speci- 
mens when he was in that region. In Peru, Tschudi gives some 
account of its habits and speaks of it as frequenting rocky 
areas, often hiding in caves by day, and coming out in the 
evening to feed on mosses and other vegetation, and lichens. 
Osgood (1914) in his journey to northern Peru in 1912 did not 
find even a track of one and writes that "so far as learned from 
inquiry, it never has been common in the region and it was only 
at rare intervals that we met a man who ever had seen one. 
A few doubtless remain in the higher parts of both the western 
and the eastern cordillera but at the points we were able to 
touch " none was found. The Peruvian Indians call it "taruga," 
and Tschudi mentions that he was to have named it Cervus 
taruga but discovered that d'Orbigny had already described it. 
It seems likely that the drives formerly held on a large scale by 
the Peruvian Indians must often have captured these small 
deer. On his later visit to the Arequipa region, Osgood (1916) 
secured a single specimen at Pampa de Arrieros, Peru. It was 
found up to an altitude of 13,000 feet, ranging somewhat lower 


than the guanaco and vicuna. In Bolivia, Neveu-Lemaire 
and Grandidier (1907) have recorded it from the Yuru district, 
at 5,000 meters. On account of the limited distribution and 
the hunting to which this deer is subject, it is evidently becom- 
ing uncommon and requires adequate protection. 



Cervus bezoarticus Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 175, 1758 (Brazil). 
SYNONYMS: Cervus campestris F. Cuvier, Diet. Sci. Nat., vol. 7, p. 484, 1817; Cervus 

azarae Wiegmann, Isis, col. 954, 1833 (Paraguay). 
FIGS.: Lydekker, 1898, pi. 22 (col); 1915, p. 190. 

The pampas deer is much smaller than its congener the 
marsh deer, about the size of the European roebuck, but more 
delicately built. The color is a light reddish brown, the face 
darker, and occasionally a black patch on the crown; "tarsal 
tuft, a patch at base of backs of ears, a ring round pedicles of 
antlers, another round each eye, lips, throat, chest, under- 
parts, fronts and inner sides of thighs, and inner sides of but- 
tocks and upper part of fore-legs whitish; . . . tail dark 
blackish brown above and white below." The antlers of the 
male are small, with a large brow tine and a posterior beam 
that forks usually once; length of beam up to 14.5 inches. 
Males usually have the upper canine present. 

The general range of this small deer includes the open 
campos of Brazil, Paraguay, and Uruguay to the pampas of 
Argentina and northern Patagonia (Lydekker, 1898). Possi- 
bly the animals of the more southern part of the range may be 
separable as a distinct subspecies for which, as pointed out by 
Lydekker, the name azarae is available. It is said to inhabit 
dry open plains, avoiding forests and thickets. "Formerly, 
when the tussocks of tall pampas-grass were dotted more or 
less thickly over all the plains, it had plenty of covert; but in 
the more settled districts it now has to live almost completely 
in the open, and has consequently become wary in the extreme" 
(Lydekker, 1898). It may be found in small groups or pairs, 
but the adult males are often solitary for most of the year. 
In the evening they are active but during the daytime lie up in 
concealment. They are said to have a strong and characteristic 
odor. When started, they bound off at a considerable speed, 


so that a good horse is necessary to overtake one. The natives 
sometimes capture them with the bolas. In parts of Brazil the 
animal goes by the native name of "guazuti," and in Uruguay 
it is known as "gama." 

According to Lydekker (1901) it has completely disappeared 
from many districts of Argentina and Uruguay. As long ago 
as 1894, Aplin (1894) wrote that in the neighborhood of Santa 
Elena, Uruguay, it had been exterminated except for a small 
herd of about a dozen does and seven bucks preserved in a 
certain district. He found it rare on the Rio Negro, but in 
some other areas it was still common. Sanborn (1929) in 
1926-27 found it plentiful in one locality in Rocha, Uruguay, 
but in most other places rare, and this is corroborated by 
Devincenzi (1935), who speaks of Rocha as the district where 
it is now to be found, although 30 years before it was abundant 
in the whole country. 

Dr. Roberto Dabbene, in a letter to Dr. Francis Harper in 
1937, stated that though formerly common in the northern 
and central regions of Argentina as far as northern Patagonia, 
it is now very scarce as far as the Chaco, and if not protected 
adequately is certain to disappear from the Argentine fauna. 
He deplores the introduction of exotic deer into the national 
parks, rather than the encouragement of the native species. 

While further and more precise information on the present 
and past status of this species is much needed, it is evident 
that it is much reduced in many districts and requires careful 
protection. In Argentina hunting this species was prohibited 
by presidential decree several years ago. 


Cervus dichotomies Illiger, Abh. Akad. Wiss. Berlin, for 1811, pp. 108, 117, 1815 

SYNONYMS: Cervus paludosus Desmarest, Mammalogie, pt. 2, p. 443, 1822; Cervus 

palustris Desmoulins, Diet. Classique Hist. Nat., vol. 3, p. 379, 1823. 
FIGS.: Lydekker, 1898, pp. 284, 285 (antlers, animals). 

This is the largest South American deer, attaining about the 
size of the British red deer, but more slenderly built, standing 
about 46 inches at the shoulder (Azara). "General colour in 
summer bright rufous chestnut, in winter brownish red, becom- 
ing lighter on flanks, neck, and chest; legs black from knees 


and hocks downwards, and tarsal tuft also black; abdomen, 
inside of thighs, chin, and insides and bases of backs of ears 
white ... a whitish line above, or a ring round eyes, 
most marked in females . . . tail yellowish rusty red 
above and black beneath . . . Fine antlers attain a length 
of from 21 to 24^ inches" (Lydekker, 1915); they are doubly 
forked, each of the two branches having a simple fork. 

The range is extensive in the tropics and subtropics of South 
America, from probably Guiana southward through Brazil to 
Paraguay and Uruguay and the Chaco or wooded districts of 
northern Argentina. Unlike its smaller relative, the pampas 
deer, it inhabits dense jungle on the borders of streams or 
swamps and it is said to go in small parties of three to five 
individuals. The skin is much used for leather by the natives, 
but the meat is apparently not held in high esteem. Lydekker 
remarks on the similarity in color between this deer and the 
maned wolf, which is found in the same areas in parts of the 

Although little information is at hand as to the details of 
distribution and relative abundance of this large deer, it is 
evident that in certain of the more settled regions in the south- 
ern part of the range it is becoming much reduced in numbers. 
Devincenzi (1935) writes that in Uruguay the Departments of 
Rocha and Treinta y Tres have been "considered the last 
refuge of the species/' while Sanborn comments that it is now 
very rare in Uruguay but was reported to be found in small 
numbers in Rocha. No effort has been made to preserve them. 
Dr. Roberto Dabbene writes, in response to inquiry by Dr. 
Harper, that in Argentina it was formerly common and oc- 
curred as far as the islands of the Delta of La Plata but is now 
(1937) confined to the Territory of Formosa, where it is not 
common. By presidential decree the hunting of this deer is 
prohibited in Argentina. 

Though no data of importance are available for Brazil, it is 
apparently to be found in fair numbers. 


Order CARNIVORA: Dogs, Cats, and Their Relatives 

E carnivores, considered as an order on p. 134, are repre- 
- sen ted by the sea otter (family Mustelidae) of the northern 
Pacific Ocean. Two races of this species are recognized; both 
were brought to the verge of extinction because of their 
valuable fur, but now they are recovering in numbers and 
may be an important resource in the future. 

Family MUSTELIDAE: Weasels, Martens, Otters 


[Mustela] lutris Linnaeus, Systema Naturae,, ed. 10, vol. 1, p. 45, 1758 (Kamchatka). 
SYNONYM: Lutra marina Schreber, Saugthiere, vol. 3, p. 465, pi. 128, 1778 (ex Steller) ; 
Nov. Comment. Petropol., vol. 2, p. 367, pi. 26, 1751. 


Latax lutris nereis Merriam, Proc. Biol. Soc. Washington, vol. 17, p. 159, Oct. 6, 1904 

("San Miguel Island, Santa Barbara Islands, California"), 
FIGS.: Royal Nat. Hist., vol. 2, p. 98, fig., 1894; zur Strassen, 1914, pp. 12*. 13*; 

Nelson, 1916, col. fig. p. 434. 

The distinction between the northern sea otter and the 
southern race rests apparently upon slight cranial differences; 
hence it is not possible to distinguish the two without recourse 
to the skulls. Accordingly they may be treated together as 
representing but a single species, while the limits of geographi- 
cal and individual variation still remain to be more precisely 
defined. It is believed that the southern race is the one 
occurring north at least to the coast of Oregon. 

The sea otter is of rather heavy, robust form, about 4 feet in 
length, of which the tail is about a foot. The fore feet are small, 
with naked palms, but the hind feet are long and broad, 
webbed and with furry soles, recalling the flippers of a seal. 



The color is dark brown, becoming hoary on the head. The 
sparse whiskers are stout and short. The posterior teeth are 
remarkably enlarged, broadened, and their cusps blunted to 
form low rounded knobs suitable for crushing the shellfish that 
form their diet. Dr. Merriam states that the skin of the type 
of the southern race was 6 feet long, but it may have been 
stretched, or perhaps a very large individual. Barabash- 
Nikiforov (1935) gives a maximum length of 1,635 mm., of 
which the tail was 330, and a maximum weight of 35 kilograms. 

Sea otters were abundant formerly from the coasts of south- 
ern Kamchatka to the Kurile Islands in the western North 
Pacific and in the waters about the islands of the Bering Sea 
and Alaskan coasts southward following the cooler currents 
even to the coasts of southern California. On the Asiatic side 
they ranged at one time as far south as Yezo. On account of 
the richness of its fur the sea otter was ruthlessly pursued 
since its discovery by the Russians in about the middle of the 
eighteenth century, down to more modern times, until by the 
first decade of the present century it was very nearly exter- 
minated. The few skins that then came on the market sold for 
as much as $1,000 apiece. Finally the United States Govern- 
ment in 1910 passed a law forbidding its capture in American 
waters and negotiated treaties with other interested nations 
for giving similar protection. Now, after a lapse of little over 
a quarter of a century, there is encouraging evidence that the 
species is recovering and it has lately appeared in some num- 
bers off the California coasts. 

The history of the pursuit and exploitation of the sea otter 
has at various times been written. From some of these ac- 
counts the following pertinent facts are gleaned. Active 
trade in sea-otter skins seems to have begun in 1742, when 
Bering, after being wrecked in the sea bearing his name, 
returned to Petropaulovsk with about 900 skins stowed in the 
small boat built from the wreck of his vessel, the St. Peter. 
These at first were chiefly traded with the Chinese, by whom 
they were highly valued. Shelikof, a Russian trader, at once 
saw great possibilities in further trade in this fur. He founded 
a colony on Kodiak Island and made plans for collecting sea- 
otter skins on a large scale. He died before his object was 
accomplished, but his son-in-law, Nicholas P. Rezanof, carried 
on the work and in 1799 obtained from Emperor Paul the 


charter of the Russian-American Co. Shortly after, under the 
management of Shelikof and Baranof, the work was organized, 
and a fleet of bidarkas, manned by native Aleuts, started an 
intensive campaign. H. W. Elliott (1875) writes: "During 
the first few years the numbers of these animals taken all along 
the Aleutian chain, and down the whole northwest coast as 
far as Oregon, were very great ; for instance, when 

the Prybilov Islands were first discovered, two sailors, Lukan- 
non and Kaiekov, killed at St. Paul's Island, in the first year 
of occupation, five thousand, the next year they got less than a 
thousand, and in six years after not a single sea-otter appeared, 
and none have appeared since. When Shellikov's party first 
visited Cook's Inlet, they secured three thousand; during the 
second year, two thousand; in the third, only eight hundred; 
the season following they obtained six hundred; and finally, in 
1812, less than a hundred, and since then not a tenth of that 
number. The first visit made by the Russians to the Gulf of 
Yahkutat, in 1794, two thousand sea-otters were taken, but 
they diminished so rapidly that in 1799 less than three hundred 
were taken. In 1798 a large party of Russians and Aleuts 
captured in Sitka Sound and "^neighborhood twelve hundred 
skins, besides those for which they traded with the natives 
there, fully as many more; and in the spring of 1800 a few 
American and English vessels came into Sitka Sound, and 
anchored off the small Russian settlement there, and traded 
with the natives for over two thousand skins, getting the 
trade of the Indians by giving fire-arms and powder, ball, &c., 
which the Russians did not dare to do, living then, as they were, 
in the country. In one of the early years of the Russian- 
American Company, 1804, Baranov went to the Okotsk from 
Alaska with fifteen thousand sea-otter skins, that were worth 
as much then as they are now [1875], viz., fully $1,000,000." 
These tremendous inroads very quickly had an alarming 
effect. A Russian report, quoted by Elliott, tells that in 1826 
the total number secured in the entire Aleutian chain was only 
15, where previously more than 1,000 had been regularly taken 
each year. In 1835 the number from this district was 70 to 
150 annually. The natives employed in collecting these furs 
were frequently subjected to great dangers, not only from the 
elements but from other native tribes who were unfriendly. 
When Alaska was purchased by the United States, writes H. 


W. Elliott (1875), "the Russians were taking between four 
and five hundred sea-otters from the Aleutian Islands and 
south of the peninsula of Alaska, with perhaps a hundred and 
fifty more from Kenai, Yahkutat, and the Sitkan district; the 
Hudson's Bay Company and other traders getting about two 
hundred more from the coast of Queen Charlotte's and Van- 
couver's Islands, and off Gray's Harbor, Washington Territory. 
Now, during the last season, 1873, instead of less than seven 
hundred skins, as obtained by the Russians, our traders secured 
not much less than four thousand skins. This immense differ- 
ence is not due to the fact of there being a proportionate in- 
crease of sea-otters, but to the organization of hunting parties 
in the same spirit and fashion, as in the early days . . . 
The keen competition of our traders will ruin the business in 
a comparatively short time if some action is not taken by the 

"Over two-thirds of all the sea-otters taken in Alaska are 
secured in two small areas of water, little rocky islets and reefs 
around the island of Saanach and the Chernobours, which 
proves that these animals, in spite of the incessant hunting 
all the year round on this ground, seem to have some particular 
preference for it to the practical exclusion of nearly all the rest 
of the coast in the Territory. This may be due to its better 
adaptation as a breeding ground. It is also noteworthy that 
all the sea-otters taken below the Straits of Fuca are shot by 
the Indians and white hunters off the beach in the surf at 
Gray's Harbor, a stretch of less than twenty miles; here some 
fifty to a hundred are taken every year, while not half that 
number can be obtained from all the rest of the Washington 
and Oregon coast-line; there is nothing in the external appear- 
ance of this reach to cause its selection by the sea-otters, except 
perhaps that it may be a little less rocky. 

"As matters are now conducted by the hunting-parties, the 
sea-otters at Saanach and Chernobours do not have a day's 
rest during the whole year. Parties relieve each other in 
succession, and a continuous warfare is maintained. . . So 
the bad work goes on rapidly, though a majority of the natives 
and traders deprecate it." The optimum region mentioned by 
Elliott is described as a chain of small islets, most of them bare 
at low tide but with numerous reefs and rocky shoals with beds 
of kelp. "As the natives have never caught the mothers 


bringing forth their offspring on the rocks, they are disposed to 
believe that the birth takes place on kelp-beds, in pleasant or 
not over-rough weather. The female has a single pup [rarely 
two], born about fifteen inches in length. . . The sea-otter 
mother sleeps in the water on her back, with her young clasped 
between her fore-paws." 

The methods employed in taking sea otters Elliott describes 
as four: Shooting them in the surf at long range with a rifle, 
and waiting till they drift ashore if the surf is too rough to 
permit of launching a boat; surrounding an otter by a party 
of spearers in their boats and awaiting its return to the surface 
after it becomes exhausted in several dives; clubbing them in 
winter when they may at times be stealthily approached among 
rocks and kelp; and using nets 16 to 18 feet long and 6 to 10 
feet wide, of coarse meshes, spread out on the kelp beds. 
Frequently several at a time are thus taken, for when enmeshed 
they seem to make little or no attempt to get free. It is said 
that this method is preferred by the Japanese, since it permits 
the release of inferior animals or breeding females. The young, 
according to Scammon (1874), are met with at all times of the 
year, so that there appears to 1be no definite breeding season ; 
the period of gestation is believed to be eight or nine months. 

Writing in 1874, Captain Scammon speaks of the Lower 
California coasts as the haunt of sea otters and adds that 
Cerros, San Geronimo, Guadalupe, San Nicolas, and San 
Miguel Islands were "regarded as choice places to pursue 
them." Earlier, when California was part of Mexico, the 
pursuit of sea otters was prohibited by that Government under 
severe penalty. In recent years bones of the sea otter have 
been found in Indian shell heaps on Santa Cruz Island and 
near Monterey (E. M. Fisher, 1930). 

Previous to the purchase of Alaska from Russia, Wrangell 
had already instituted somewhat more far-sighted methods of 
making the annual catch, allowing no part of the hunting 
grounds to be used for two consecutive years, and thus some- 
what restricting the number killed. From 1842 to 1862 the 
average catch, including that of the Kurile Islands, was about 
1,249, and the total for the 20 years was 25,899. The result 
was to restore the sea-otter population to a slightly better con- 
dition. But these careful methods were abandoned when 
Alaska became part of the United States, and the pursuit was 


followed with the greatest intensity, so that from 1881 to 1890 
the take was 47,842, or nearly 5,000 a year (C. L. Andrews, 
1937). The immediate reduction of the breeding stock by 
thus overexploiting these animals caused the Alaska Commer- 
cial Co., which had practically secured control of the Alaskan 
fur production, to close four or five of its posts by 1897. In 
1900 the company was able to secure but 127 skins, and ten 
years later the entire catch of its fleet of 16 schooners was but 
31 skins. Even at the price of $1,000 a skin there was no 
profit in this undertaking. Fortunately, in 1910, the taking of 
sea otters by citizens of the United States was prohibited, and 
by treaties of similar import with other interested nations the 
species was given respite just in time. 

The gradual recuperation of the sea otter since 1910 from 
the verge of extinction to appreciable numbers during the last 
30 years since protective laws were established has been most 
encouraging, although relatively meager d