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Full text of "North American fauna"

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U. S. DEPARTMENT OF AGRICULTURE 

BUREAU OF BIOLOGICAL SURVEY 

K. \V. NELSON, Chief 



NORTH AMERICAN FAUNA 

No. 41 

[Actual date of publication, February 9, 1918] 




REVIEW OF THE GRIZZLY AND BIG BROWN 
BEARS OF NORTH AMERICA 

(GENUS URSUS) 

WITH DESCRIPTION OF A iNEW GENUS, VETULARCTOS 



C. HART MERRIAM 

CONSULTING BIOLOGIST, BIOLOGICAL SURVEY 
RESEARCH ASSOCIATE, SMITHSONIAN INSTITUTION 




• / 



WASHINGTON 

GOVERNMENT URIXTINi; OFFICE 

1918 






NORTH AMERICAN FAUNAS. 

Copies of North American Faunas not out of print are for sale, at the prices 
named, by the Superintendent of Documents, Government Printing Office, Wash- 
ington, D. C. 

No. 1. Revision of the North American Pocket Mice. By C. Hart Merriam. Pp. 
36, pis. 4. 1S89 Price, 10 cents. 

No. 2. Descriptions of Fourteen New Species and One New Genus of North 
American Mammals. By C. Hart Merriam. Pp. 52, pis. 8, figs. 7. 1889. 

Price, 10 cents. 

No. 3. Results of a Biological Survey of the San Francisco Mountain Region 
and Desert of the Little Colorado, Arizona. By C. Hart Merriam and 
Leonhard Stejneger. Pp. 136, pis. 14, maps 5 (colored), figs. 2. 1890. 

Price, 25 cents. 

No. 4. Description of Twenty-six New Species of North American Mammals. 
By C. Hart Merriam. Pp. 60, pis. 3, figs. 3. 1890 Price, 10 cents. 

No. 5. Results of a Biological Reconnaissance of South-Central Idaho. By 
C. Hart Merriam and Leonhard Stejneger. Descriptions of a New Genus and 
Two New Species of North American Mammals. By C. Hart Merriam. Pp. 
132, pis. 4 (1 colored), figs. 4. 1891 Price, 15 cents. 

No. 6. Not issued. 

No. 7. The Death Valley Expedition : A Biological Survey of Parts of Cali- 
fornia, Nevada, Arizona, and Utah. Part II. — 1. Birds, by A. K. Fisher. 
2. Reptiles and Batrachians, by Leonhard Stejneger. 3. Fishes, by Charles 
H. Gilbert. 4. Insects, by C. V. Riley. 5. Mollusks, by R. E. C. Stearns. 
6. Desert Trees and Shrubs, by C. Hart Merriam. 7. Desert Cactuses and 
Yuccas, by C. Hart Merriam. 8. List of Localities, by T. S. Palmer. Pp. 402, 
pis. 15, maps 5, figs. 2. 1893 [Out of print.] 

No. 8. Monographic Revision of the Pocket Gophers, Family Geomyidse (exclu- 
sive of the species of Thomomys) . By C. Hart Merriam. Pp. 258, pis. 20, 
figs. 71, maps 4 (colored). 1895 Price, 35 cents. 

No. 9. Not issued. ) 

No. 10. Revision of the Shrews of the American Genera Biarina and Notiosorex. 
By C. Hart Merriam. The Long-tailed Shrews of the Eastern United States. 
By Gerrit S. Miller, jr. Synopsis of the American Shrews of the Genus 
Sorex. By C. Hart Merriam. Pp. 124, pis. 12, figs. 3. 1895 Price, 15 cents. 

No. 11. Synopsis of the Weasels of North America. By C. Hart Merriam. Pp. 
44, pis. 6, tigs. 16. 1S96 *_ Price, 10 cents. 

No. 12. The Genera and Subgenera of Voles and Lemmings. By Gerrit S. 
Miller, jr. Pp. 84, pis. 3, figs. 40. 1896 Price, 10 cents. 

No. 13. Revision of the North American Bats of the Family Vespertilionidae. 
By Gerrit S. Miller, jr. Pp. 140, pis. 3, figs. 40. 1897 Price, 10 cents. 

No. 14. Natural History of the Tres Marias Islands, Mexico. Prepared under 
the direction of C. Hart Merriam. General Account of the Islands, with Re- 
ports on Mammals and Birds, by E. W. Nelson. Reptiles, by Leonhard 
Stejneger. Notes on Crustacea, by Mary J. Rathbun. Plants, by J. N. Rose. 
Bibliography, by E. W. Nelson. Pp. 97, pi. (map), figs. 2. 1899. 

Price, 10 cents. 

No. 15. Revision of the Jumping Mice of the Genus Zapus. By Edward A. 
Preble. Pp. 42, pi., figs. 4. 1899 Price, 5 cents. 

No. 16. Results of a Biological Survey of Mount Shasta, California. By C. Hart 
Merriam. Pp. 179, pis. 5, figs. 46. 1899 [Out of print.} 

No. 17. Revision of American Voles of the Genus Microtus. By Vernon Bailey. 
Pp. 88, pis. 5- figs. 17. 1900 [Out of print. I 

No. 18. Revision of the Pocket Mice of the Genus Perognathus. By Wilfred H. 

Osgood. Pp. 72. pis. 4 (inch 2 maps), figs. 15. 1900 Price, 10 cents. 

(Continued on page 3. of cover.) 



North American Fauna No. 41. U. S. Dept. Agr. Biological Survey. 



Plate I. 




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U. S. DEPARTMENT OF AGRICULTURE 

BUREAU OF BIOLOGICAL SURVEY 

E. W. NELSON, Chief 



NORTH AMERICAN FAUNA 

No. 41 

[Actual date of publication, February 9, 1918] 




REVIEW OF THE GRIZZLY AND BIG BROWN 
BEARS OF NORTH AMERICA 

(GENUS URSUS) 

WITH DESCRIPTION OF A NEW GENUS, VETULARCTOS 



C. HART MERRIAM 

CONSULTING BIOLOGIST, BIOLOGICAL SURVEY 
RESEARCH ASSOCIATE, SMITHSONIAN INSTITUTION 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



/ t/6 




LETTER OF TRANSMITTAL. 



United States Department or Agriculture, 

Bureau of Biological Survey, 
/ Washington, D. C, July 18, 1917. 
Sir : I have the honor to transmit for publication as North Ameri- 
can Fauna No. 41 a review of the grizzly and big brown bears of 
North America, by Dr. C. Hart Merriam, consulting biologist and 
former chief of the Biological Survey and research associate of the 
Smithsonian Institution. This review was prepared and originally 
submitted in September, 1916, but before composition was begun 
it was recalled from the printer in order that additional informa- 
tion, developed by the discovery of new material, might be included. 
The work is based largely upon material in the collection of the 
Biological Survey. Up to 20 years ago only 8 species of grizzly 
and big brown bears were known, but since then, largely through 
the investigations of Dr. Merriam, the number of recognizable forms 
has increased to 86. Additional study and material may solve cer- 
tain points now in doubt, but it is not deemed advisable to delay 
further the publication of our present state of knowledge of this 
group of America's historic big game animals, now vanished from 
great stretches of their former domain. This review will be of 
material assistance to students and others interested in our native 
wild life, past and present. 
Respectfully, 

E. W. Nelson, 
Chief, Biological Survey. 
Hon. Davto F. Hpu?ton^ , r o « : .«. : . 

Secretary &f,Agnb&1iWe,: I ..; 
2 



CONTENTS. 



Introduction •. 7 

Geographic distribution 9 

Sexual differences 10 

Age differences 10 

Material examined 10 

Technical terms 11 

Classification of grizzly and big brown bears 12 

Relative values of cranial and dental characters 13 

List of species and subspecies, with type localities 14 

Descriptions of species and subspecies 17 

Horribilis group 17 

Planiceps group 34 

Arizonx group 53 

Hylodromus group 77 

Horrixus group 84 

Stikeenensis group 88 

Alascensis group 94 

Richardsoni group 99 

Kidderi group 106 

Innuitus group 110 

Townsendi group 115 

Dalli group 116 

Gyas group 124 

Kenaiensis group 127 

Vetularctos, a new genus related to Ursut 131 

Index 135 

3 



ILLUSTRATIONS. 



PLATES. 

[Plate I, frontispiece; Plates II-XVI following page 134.] 

Plate I. Wild grizzly in northwestern Wyoming. From photograph by Frederick 
K. Vreeland. 
II. Skull of Ursus gyas, old S , from Bear Bay, Alaska Peninsula. No. 91690. 

III. Skull of Ursus middendorffi, old S, from Chiniak Point, Kodiak Island, 

Alaska. No. 96509. 

IV. Skull of Ursus kenaiensis, old d 1 , from Kenai Peninsula, Alaska. No. 

210291. 
V. Skull of Ursus sheldoni, adult c?, from Montague Island, Prince William 
Sound, Alaska. No. 137318 (type). 
VI. Skull of Ursus shirasi, old <?, from Admiralty Island, Alaska. No. 203030 

(type). 
VII. Skull of Ursus innuitus, old <?, from Golofnin Bay, Alaska. No. 179780 

(type). 
VIII. Skull of Ursus kidderi kidderi, old <?, from Belkofski, Alaska Peninsula. 
No. 91698. 
IX. Skull of Ursus stikeenensis, adult <? , from Tatletuey Lake, near head of 

Skeena River, British Columbia. No. 202794 (type). 
X. Skull of Ursus ophrus, old cf , from eastern British Columbia. No. 210252 

(type). 
XI. Skull of Ursus rungiusi rungiusi, old d\ from Indian Point Creek, near 

Barkerville, British Columbia. No. 209899. 
XII. Skull of Ursus alascensi-s, old d, from Unalaklik River, Alaska. No. 76466 
(type). 

XIII. Skull of Ursus nortoni, old 9 , from Yakutat, Alaska. No. 178763 (type). 

XIV. Skull of Ursus horribilis horribilis, old c? , from Missouri Breaks, eastern 

Montana. No. 202739. 
XV. Skull of Ursus horriseus, old <$ , from Coppermines, New Mexico No. 990 

(type). 
XVI. Skull of Ursus arizonse, adult <? , from Escudilla Mountains, Apache County, 
Arizona. No. 177332 (type). 

[The plates of skulls, owing to restriction of space, are limited to the side view, and 
consequently in some cases are misleading. Skulls having similar profiles often differ sur- 
prisingly when viewed from above or below, as would be seen at a glance were it prac- 
ticable to give two views. J 

& 



No. 41. NORTH AMERICAN FAUNA. Feb., 1918. 

REVIEW OF THE GRIZZLY AND BIG BROWN 
BEARS OF NORTH AMERICA 

(GENUS URSUS) 
WITH DESCRIPTION OF A NEW GENUS, VETULARCTOS. 



By C. Hart Merbiam. 



INTRODUCTION. 

When Audubon and Bachman published their great work on the 
Mammals of North America (1846-1854), and in fact up to the year 
1857, it was commonly believed by naturalists as well as by hunters 
and the public generally that there was only a single species of 
grizzly bear — the one described by Lewis and Clark in 1804-5, and 
named Ursus horribilis by Ord in 1815. Baird, in 1857, described 
another species, from Coppermines, New Mexico, which he named 
Ursus horriceus. 

Nearly 40 years later, in my " Preliminary Synopsis of the Ameri- 
can Bears," 1 eight grizzlies and big brown bears were recognized, 
of which five were described as new. It was not then suspected 
that the number remaining to be discovered was anything like so 
great as has since proved to be the case. The steady influx of speci- 
mens resulting from the labors of the Biological Survey, supple- 
mented by the personal efforts of a number of hunter-naturalists, 
brought to light many surprises, most of which have been published ; 
and beginning in the spring of 1910, a fund placed at my disposal 
made it possible to offer hunters and trappers sufficient inducement 
to tempt them to exert themselves in securing needed specimens. As 
a result, the national collection of bears has steadily grown until, in 
number of species represented, in completeness of series, and in num- 
ber of type specimens, it now far excels all other collections in the 
world together. Nevertheless there are many gaps in the series. 

1 Proc. Biol. Soc. Washington, X, pp. 65-S3, AprU 13, 1896. 



8 NORTH AMERICAN FAUNA. [No. 41. 

Knowledge of the big bears is by no means complete and many years 
must pass before the last word on the subject will be written. Many 
bears now roaming the wilds will have to be killed and their skulls 
and skins sent to museums before their characters and variations will 
be fully understood and before it will be possible to construct accu- 
rate maps of their ranges. Persons having the means and ambition 
to hunt big game may be assured that bears are still common in many 
parts of British Columbia, Yukon Territory, and Alaska, and that 
much additional material is absolutely required to settle questions 
still in doubt. 

Among the localities from which specimens are greatly needed 
may be mentioned Lynn Canal and Lituya Bay, Alaska, and in fact 
the entire coast strip between Cross Sound and Yakutat Bay; the 
Cook Inlet and Susitna regions; the mountains between the Yukon 
and Tanana; the Endicott Range and other mountains between the 
Yukon and the Arctic coast all the way from Seward Peninsula 
to the Alaska- Yukon boundary; the Rocky Mountains of Canada, 
from British Columbia northward, including the sources of the 
Pelly, Macmillan, Stewart, and Porcupine Rivers; the Mackenzie 
River and Great Bear Lake region; the southwest corner of Yukon 
Territory; the western part of Alberta; and the interior of British 
Columbia. In the United States, skulls of adult males are much 
needed from all localities inhabited by grizzly bears, particularly in 
Colorado, Idaho, Montana, and Wyoming — including the Glacier and 
other National Park regions. 

What is most needed to-day in this line is a series of adults col- 
lected by absolutely trustworthy persons and labeled on the spot 
for locality and sex. Many specimens in museums are not labeled 
for sex ; others have the sex wrongly marked ; and many either lack 
localities or the localities given are open to serious doubt. A speci- 
men is of little value unless one can pin his faith on the label. 

Some writers have advanced the view that the various species of 
bears freely interbreed. Let those so minded ask themselves the 
question, If promiscuous interbreeding were to take place, what 
would become of the species? From the nature of the case, the sta- 
bility of species depends on the rarity of crossings with other species, 
for if interbreeding were to take place frequently the species so inter- 
breeding would of course cease to exist, having merged into a com- 
mon hybrid. Hybrids now and then occur, particularly in zoological 
gardens, but among wild animals in their native haunts they are ex- 
ceedingly rare. 

The number of species here given will appear to many as preposter- 
ous. To all such I extend a cordial invitation to visit the National 



1918.] INTKODUCTION. 9 

Museum and see for themselves what the bear skulls show. Recog- 
nition of species is a matter of interpretation. If the material is 
adequate there can be little room for difference of opinion; if in- 
adequate, many important points must remain in doubt. It is not 
the business of the naturalist either to create or to suppress species, 
but to endeavor to ascertain how many Nature has established, and 
having discovered this, to point out their characters and learn as 
much as possible about them. 

One of the unlooked-for results of the critical study of the Ameri- 
can bears is the discovery that the big bears, like mice and other small 
mammals, split up into a large number of forms whose ranges in some 
cases overlap so that three or more species may be found in the same 
region. 

Another surprising result is the discovery that Admiralty Island in 
Southeastern Alaska appears to be inhabited by no less than five 
distinct species, each of which is obviously related to and representa- 
tive of an adjacent mainland species. The recognition of this very 
remarkable state of affairs makes it possible to understand what 
before had seemed a most anomalous condition, namely, the extraor- 
dinary diversity or variability of the skulls and teeth of the island 
bears. It was not until material essential for the determination of 
the mainland forms had been collected that it was possible to recog- 
nize and define the island forms. 

The varying degrees of divergence of the island bears furnish an 
interesting index to the relative time when each obtained a foothold 
on the island. In this connection it is well to remember that the 
breadth of the strait separating Admiralty Island from the main- 
land at its narrowest point does not exceed 5 miles. 

The mainland big bears with their representatives on Admiralty 
Island here provisionally recognized are : 

Mainland Species. Admiralty Island Species. 

Vrsu8 dalli Ursus shirasi. 

Ursus stikeenensis Ursus mirabilis. 

Vrsus tahltanicus Ursus insularis. 

Ursus ktoakiutl Ursus neglectus. 

Ursus caurinus Ursus eulophus. 

GEOGRAPHIC DISTRIBUTION. 

In early days grizzly bears were common in most parts of west- 
ern North America, their range being nearly continuous from north- 
ern Mexico northward through the Western States and western 
Canada to northern Alaska; but now within the United States sev- 



10 NORTH AMERICAN FAUNA. [No. 41. 

eral of the species are extinct, and those still living are confined in 
the main to remote or inaccessible mountain ranges, where two or 
more species not infrequently occur together. In many cases the 
original distribution areas overlapped, as they do to-day in parts of 
British Columbia, Yukon, and Alaska; in other cases, owing to the 
settlement of the country, species inhabiting easily accessible areas 
were either exterminated or forced into mountains where they now 
occupy the same ground with other species, so that it is impossible to 
ascertain what the original distribution was. As a rule, in cases where 
two or more species inhabit the same area, the species occurring to- 
gether belong to different superspecific groups. Thus in the Yellow- 
stone Park and Stikine River regions representatives of the kor- 
ribilis, absarokus, tahltanicus, and chelan groups occur, so far as 
known, in the same localities. 

SEXUAL DIFFERENCES. 

In most species of bears the males are much larger than the fe- 
males. In some the disparity in size is very remarkable, as in mid- 
dendorffi of Kodiak Island and magister of southern California. 
In a few cases the difference is slight, as in kidderi of Alaska 
Peninsula. 

AGE DIFFERENCES. 

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. In species having the frontal shield highly ele- 
vated, as in middendorfji, kluane, stikeenensis, and mi?'abHis, the f ron- 
tals reach their maximum of arching or bulging in early adult life 
(about the sixth year), after which they gradually become flatter. 

MATERIAL EXAMINED. 

It is a pleasure to express appreciation of the invaluable assistance 
rendered by the loan or presentation of skins and skulls of grizzly 
and brown bears utilized in the preparation of the accompanying 
descriptions. To Charles Sheldon, G. Frederick Norton, and the 
late Charles R. Cross, jr., and to J. H. Kidder, Robert P. Blake, 
Waldo Emerson Forbes, George Mixter, Samuel Mixter, Dr. William 
Jason Mixter, Charles S. King, and Homer E. Sargent, special thanks 
are due for their generosity in placing at my disposal the valuable 
specimens and notes obtained on their private hunting trips. Most 
of these specimens have been presented to the national collections. 
And to Miss Annie M. Alexander, of Oakland, California, special 
acknowledgment should be made for the loan of her collection of 



1918.] INTRODUCTION. 11 

Alaska bears, now in the Museum of Vertebrate Zoology of the Uni- 
versity of California, and second only to that of the Biological 
Survey and National Museum. 1 

In conclusion it is only proper to state that the material on which 
the present publication is based could not have been brought together 
during my lifetime, nor the results prepared for publication, but for 
the generous assistance of Mrs. E. H. Harriman in establishing 
under the Smithsonian Institution a special research fund for my 
scientific work. 

TECHNICAL TERMS. 

In describing the skulls of bears a few terms are used in a special 
sense which it is desirable to understand. 

The frontal shield is that part of the top of the skull extending 
from the base of the rostrum backward to the meeting point of the 
temporal impressions. It is elevated above the surrounding parts 
and is sharply defined. Its posterior point, confined between the tem- 
poral impressions, is longer in female than in male skulls, and up 
to a certain limit becomes shorter with age. 

The postorbital processes stand out from the sides of the frontal 
shield, limiting the orbits posteriorly. 

The term sulcate is applied to skulls having a longitudinal median 
depression or groove in the frontal shield, usually shallow and rather 
broad and without definite lateral limits. 

1 Others who have helped hy the presentation or loan of material are : C. E. Aiken. 
Dr. J. A. Allen, Dr. R. M. Anderson, Edward F. Ball, Dr. Arthur H. Bannon, Br. William 
Bebb, H. C. Beggs, Mrs. C. C. Beggs, John P. Bird, W. C. Bradbury, J. Stanley-Brown, 
Fred K. Burnham, Mr. and Mrs. E. S. Cameron, Dr. Frank M. Chapman, R. H. Chapman, 
Elton Clark, James L. Clark, Charles B. Cory, Prof. Charles R. Cross, Heyward Cutting, 
Frank S. Daggett, E. W. Deming. Howard Eaton, Charles Farwell Edson, Lincoln Ells- 
worth, Lieut. G. T. Emmons, J. D. Figgins, J. Stanley Foster, Charles A. Gianini, Dr. 
J. B. Girard, Dr. Joseph Grinnell, Samuel Henshaw, Charles J. Hittell, Dr. W. J. Holland, 
Dr. R. Houston, James T. Jardine, Remington Kellogg, Francis Kermode, Charles S. King. 
Paul Kleineidam, Prof. S. H. Knight, Frederick Lambart, Edward H. Litchfield, Col. J. A. 
McGuire, John Murgatroyd, Prof. C. C. Nutting, Wilfred H. Osgood, John M. Phillips, 
the late W. Hallett Phillips, the late Warburton Pike, Wilson Potter, George D. Pratt, 
Dr. E. P. Richardson, Powhatan Robinson, Archibald Rogers, Carl Rungius, Homer E. 
Sargent, Prof. W. B. Scott, George Shiras 3d, George Shiras 4th, Dr. H. A. Sifton, Henry 
A. Stewart, Dr. Walter T. Swingle, P. A. Taverner, Prof. S. D. Thacher, Dr. Charles H. 
Townsend, Frederick K. Vreeland, E. R. Warren, A. Bryan Williams, and W. W. Wood. 

Fullest acknowledgments are due the following institutions for their courtesy in loaning 
specimens : American Museum of Natural History, New York ; Carnegie Museum, Pitts- 
burgh ; Colorado Museum of Natural History, Denver ; Field Museum of Natural History, 
Chicago ; Museum of Comparative Zoology, Cambridge ; Museum of History, Science, and 
Art, Los Angeles ; Museum of Vertebrate Zoology, University of California ; Peabody 
Museum of Salem ; Peabody Museum of Yale University ; Provincial Museum, Victoria, 
British Columbia ; U. S. National Museum, Washington ; Victoria Memorial Museum, 
Ottawa ; Zoological Society of Philadelphia ; and the Museums of the Universities of Iowa, 
Kansas, Nebraska, and Wyoming. 

For the frontispiece, showing a wild grizzly in a pine forest near Yellowstone Park, 
western Wyoming, thanks are due Frederick K. Vreeland, who was so fortunate as to 
take the photograph. 



12 NOBTH AMERICAN FAUNA. [No. 41. 

The term dished means that the nasal or fronto-nasal region is 
depressed, producing a change of angle from the plane of the anterior 
part of the nasals to the plane (or slope) of the frontal shield. 
Some skulls are strongly dished, some are flat, while some have the 
fronto-nasal region elevated and compressed, giving a very different 
outline from that of the normally dished skull. 

The term braincase is loosely applied to the whole upper rounded 
part of the skull between the occiput and postorbital processes, but 
not including the wedge-shape posterior part of the frontal shield. 
Used in this way, it covers the parietal bones and posterior part of 
the frontals on each side of the temporal ridges, including not only 
the actual bony case inclosing the brain, but also its anterior con- 
tinuation (the sinus case). 

The term sinus case is applied more definitely to the smoothly 
rounded part of the frontals below the shield and in direct continu- 
ation of the braincase — the outer shell covering the large sinuses or 
air cells lying between the nasal chamber and the brain. It is not 
always discriminated from the braincase. 

The term bellied is applied to the posterior part of the inferior 
border of the ramus of the under jaw to indicate a swelling or thick- 
ening common in many species. 

The term subangvZar border is applied to the posterior part of the 
inferior border of the under jaw, immediately anterior to the angular 
process, and usually set off from the rest of the ramus by a step or 
small tubercle. 

The term keeled is applied to a not uncommon condition of the 
upper part of the sinus case, in which the top or arch is compressed, 
rising rather narrowly into the anterior part of the sagittal crest 
and posterior part of the frontal shield. The condition is marked 
in Ursus eulophus and occurs in several other species. In most 
species, however, this part of the braincase is rather broadly rounded, 
the sagittal crest rising abruptly from the median line. 

Measurements are always in millimeters unless otherwise specified. 

The museum number of the skull, unless otherwise stated, is under- 
stood to be that of the United States National Museum. 

CLASSIFICATION OF GRIZZLY AND BIG BROWN BEARS. 

The differences formerly supposed to exist between the grizzlies 
and the big brown bears appear, in the light of the material now 
available, to distinguish certain groups of species from certain other 
groups, rather than the grizzlies collectively from the big brown 
bears collectively. In other words, the differences between the griz- 
zlies on the one hand and the big brown bears on the other are 



1018.] INTRODUCTION. 13 

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 posi- 
tively referred to either group. In fact, it seems at least possible 
that certain species which appear to belong with the grizzlies are 
closely related to certain other species which clearly belong with 
the big brown bears. 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 sulcate heel of the true grizzlies. But these are average differ- 
ences, not one of which holds true throughout the group. Most of 
the specimens in museums consist of skulls only, unaccompanied by 
skins or claws, leaving a doubt as to the external characters; and 
in old bears the important fourth lower premolar is likely to be so 
worn that its original form can not be made out. And, worst of all, 
some of the grizzlies lack the distinctive type of premolar, leaving 
only the skull as a guide to their affinities. The present classification, 
therefore, must be regarded as tentative and subject to revision. 

RELATIVE VALUES OF CRANIAL AND DENTAL CHARACTERS. 

In my judgment cranial characters among the bears of the genus 
Ursus are more permanent and of more significance from the stand- 
point of classification than minor tooth characters. The teeth are 
strongly modified by food and consequently in some cases present 
marked variations in the same group. Thus the skull of adult male 
chelidonias from the coast of southern British Columbia is almost in- 
distinguishable from that of imperator from the Yellowstone Park, a 
member of the horribilis group ; but imperator has very large molars, 
nearly as big as those of horribilis and bairdi, while chelidonias, be- 
ing a fish eater, has such small molars that were it not for the skull 
no one would think of placing it in the horribilis group. 

Cranial and dental characters among the big bears are very subtle. 
As a rule comparison 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 re- 
semblances far outnumber the differences. Hence the greatest caution 
is necessary to avoid misleading conclusions. 

The present paper is merely a review of the existing state of knowl- 
edge of the grizzlies and big brown bears of America and does not 
include either the polar or the black bears. It is not intended as a 



14 NORTH AMERICAN FAUNA. [No. 41. 

monographic revision, but aims to supply a list of the species, to- 
gether with descriptions and comparisons of adult skulls, chiefly 
males. Little is said of external characters, for the reason that little 
is known, only a few skins with claws being available for study. 

List of Species and Subspecies of Grizzly and Big Brown Bears, with Type 

Localities. 1 

(Classification provisional.) 
Horribilis group (pp. 17-34): 

Ursus horribilis horribilis Ord Missouri River, northeastern Montana. 

horribilis bairdi Merriam Blue River, Summit County, Colorado. 

horribilis impera tor Merriam Yellowstone National Park, Wyoming. 

chelidonias nobis Jervis Inlet, British Columbia. 

atnarJco nobis / . .Atnarko River, British Columbia. 

kwdkiutl Merriam Jervis Inlet, British Columbia. 

nortoni Merriam Southeastern side Yakutat Bay, Alaska. 

warburtoni Merriam Atnarko River, British Columbia. 

neglectus Merriam Near Hawk Inlet, Admiralty Island, 

Southeastern Alaska. 

californicus Merriam Monterey, California. 

tularensis Merriam Fort Tejon, California. 

colusus Merriam Sacramento Valley, California. 

dusorgus nobis 2 .Jack Pine River, lberta-British Columbia 

boundary. 
Planiceps group (pp. 34-53): 

Ursus nelsoni Merriam Colonia Garcia, Chihuahua, Mexico. 

texensis texensis Merriam Davis Mountains, Texas. 

texensis navaho Merriam Navajo country near Fort Defiance, Ari- 
zona. (Probably Chuska Mountains.) 

planiceps nobis Colorado (exact locality uncertain). 

macrodon nobis Twin Lakes, Colorado. 

mirus nobis Yellowstone National Park , Wyoming. 

eltonclarhi Merriam Near Freshwater Bay, Chichagof Island, 

Alaska. 

tahltanicus Merriam Klappan Creek (=Third South Fork Stikine 

River), British Columbia. 

insularis Merriam Admiralty Island, Alaska. 

orgilos Merriam Bartlett Bay, east side Glacier Bay, South- 
eastern Alaska. 

orgiloides nobis Italio River, Alaska. 

pallasi Merriam Donjek River, southwestern Yukon. 

rungiusi rungiusi nobis Rocky Mountains, headwaters Athabaska 

River, Alberta. 

rungiusi sagittalis iiobis Champagne Landing, southwestern Yukon. 

macfarlani nobis Anderson River, 50 miles below Fort 

Anderson, Mackenzie. 

canadensis Merriam 2 Moose Pass, near Mount Robson, British 

Columbia. 

1 Nearly 130 years ago Prof. Zauschner proposed the name Ursus saribur for an animal 
'• from the region of Canada " (Bestlmmung der Hundsart Krokute, und der Barenart 
Saribur, p. 8, 1788), but the species appears to be impossible of identification. 

* Reference to group provisional. 



1918.] LIST OF SPECIES AND SUBSPECIES. 15 

Arizonse group (pp. 53-76): 

Ursus arizonse Merriam Escudilla Mountains, Apache County, 

Arizona. 

idahoensis nobis North Fork Teton River, eastern Idaho. 

pulchellus pulchellus nobis Ross River, Yukon. 

pulchellus ereunetts nobis Beaverfoot Range, Kootenay District, 

British Columbia. 

oribasus nobis Upper Liard River, Yukon. 

chelan Merriam East slope Cascade Mountains, Chelan 

County, Washington. 

shoshone Merriam Estes Park, Colorado. 

kennerlyi Merriam Mountains of northeastern Sonora, near Los 

Nogales, Mexico. 

utahensis Merriam Salina Creek, near Mayfield, Utah. 

perturbans nobis Mount Taylor, northern New Mexico. 

rogersi rogersi nobis Upper Greybull River, Absaroka Moun- 
tains, Wyoming. 

rogersi bisonophagus nobis Black Hills (Bear Lodge), northeastern 

Wyoming. 

pervagor Merriam Pemberton Lake (now Lillooet Lake), Brit- 
ish Columbia. 

caurmus Merriam Berners Bay, east side Lynn Canal, South- 
eastern Alaska. 

eulophus Merriam Admiralty Island, Southeastern Alaska. 

klamathensis Merriam * Beswick, near mouth Shovel Creek, Kla- 
math River, northern California. 

mendocinensis Merriam ' Long Valley, Mendocino County, Califor- 
nia. 

magister Merriam l Los Biacitos, Santa Ana Mountains, South- 
ern California. 
Hylodromus group (pp. 77-84): 

Ursus hylodromus Elliot Rocky Mountains, western Alberta. 

kluane kluane Merriam McConnell River, Yukon. 

kluane impiger nobis Columbia Valley, British Columbia. 

pellyensis nobis Ketza Divide, Pelly Mountains, Yukon. 

andersoni nobis ■ Dease River, near Great Bear Lake, Mac- 
kenzie. 
Eorrixus group (pp. 84-88): 

Ursus apache Merriam Whorton Creek, south slope White Moun- 
tains, eastern Arizona (a few miles west 
of Blue). 

horriseus Baird Coppermines, southwestern New Mexico. 

henshawi Merriam Southern Sierra Nevada, near Havilah, 

Kern County, California. 
Stikeenensis group (pp. 88-94): 

Ursus stikeenensis Merriam Tatletuey Lake, tributary to Finlay River, 

near head Skeena River, British Colum- 
bia. 

crassodon nobis Klappan Creek (=Third South Fork Stilrine 

River), British Columbia. 

crassus nobis ' Upper Macmillan River, Yukon. 

mirabilis Merriam 1 Admiralty Island, Alaska. 

absarokus Merriam * Little Bighorn River, northern Bighorn 

Mountains, Montana. 

'Reference to group provisional. 
64854°— 18 2 



16 • NORTH AMERICAN FAUNA. [No. 41. 

Alascensis group (pp. 94-99): 

Ursus alascensis Merriam Unalaklik River, Alaska. 

t klat Merriam Head of Toklat River, north base Alaska 

Range, near Mount McKinley, Alaska. 

latifrons Merriam Jasper House, Alberta. 

Richardsoni group (pp. 99-106): 

Ursus richardsoni Swainson Shore of Arctic Ocean, west side Bathurst 

Inlet, near mouth of Hood River. 

russelli Merriam 1 West side Mackenzie River delta, Canada. 

phxonyx Merriam Glacier Mountain, Tanana Mountains, 

Alaska (about 2 miles below source of 
Comet Creek, near Fortymile Creek, be- 
tween Yukon and Tanana Rivers). 

internationalis Merriam Alaska- Yukon boundary, about 50 miles 

south of Arctic coast. 

ophrus Merriam Eastern British Columbia (exact locality 

unknown). 

•washake Merriam North Fork Shoshone River, Absaroka 

Mountains, western Wyoming. 
Kidderi group (pp. 106-110): 

Ursus kidderi kidderi Merriam Chinitna Bay, Cook Inlet, Alaska. 

kidderi tundrensis Merriam Shaktolik River, Norton Sound, Alaska. 

eximius Merriam Head of Knik Arm, Cook Inlet, Alaska. 

Innuitus group (pp. 110-115): 

Ursus innuitus Merriam Golofnin Bay, south side Seward Penin- 
sula, northwestern Alaska. 

cressonus Merriam Lakina River, south slope Wrangell Range, 

Alaska. 

alexandrse Merriam * Kusilof Lake, Kenai Peninsula, Alaska. 

Townsendi group (pp. 115-116): 

Ursus townsendi Merriam Mainland of Southeastern Alaska (exact 

locality uncertain). 
Dalli group (pp. 116-124): 

Ursus dalli Merriam Yakutat Bay (northwest side), Alaska. 

hoots Merriam Clearwater Creek, a north branch of Stikine 

River, British Columbia. 

sitkensis Merriam Sitka Islands, Alaska. 

shirasi Merriam Pybus Bay, Admiralty Island, Alaska. 

nuchek Merriam 1 Head of Nuchek Bay, Hinchinbrook 

Island. Prince William Sound, Alaska. 
Gyas group (pp. 124-127): 

Ursus gyas Merriam Pavlof Bay, Alaska Peninsula. 

middendorffi Merriam Kodiak Island, Alaska. 

Kenaiensis group (pp. 127-131): 

Ursus kenaiensis Merriam Cape Elizabeth, extreme west end Kenai 

Peninsula, Alaska. 

sheldoni Merriam Montague Island, Prince William Sound, 

Alaska. 



Vetularctos genus nobis fpp. 131-133): 

Vetularctos inopinatus nobis Rendezvous Lake, northeast of Fort Ander- 
son, Mackenzie. 
1 Reference to group provisional. 



1918.J HORRIBILIS GROUP. 17 

DESCRIPTIONS OF SPECIES AND SUBSPECIES. 
Horribilis Group. 
URSUS HORRIBILIS HORRIBILIS Obd. 

Big Plains Gbizzly. 
(Pi. XIV.) 

Ursus horribilis Ord, Guthrie's Geography, 2d Amer. Ed., pp. 291, 300, 1815 
(Rhoads' reprint, 1894). Based on the white bear of Lewis and Clark, 
particularly the one from eastern Montana killed May 5, 1805, on the 
Missouri River, near the mouth of Poplar River (called by them Porcupine 
River, but not the same as the Porcupine of to-day, which is about 50 miles 
farther west). 

Ursus ferox Rafinesque, Amer. Monthly Mag., I, p. 437, Oct., 1817 (nomen 
nudum). 

" Ursus ferox Lewis & Clarck," Desmarest, Mammalogie, p. 164, Paris, 1820 
(under U. cinereus). 

Ursus cinereus Desmarest, Mammalogie, p. 164, Paris, 1820. 

Ursus griseus Choris, Voyage Pittoresque autour du Monde, folio colored plate 
and text (unpaged), Paris, 1822. 

Ursus candescens Ham. Smith, in Griffith's Ouvier, II, p. 229 and facing plate, 
1827 ; ibid. V, p. 112, 1827. 

Type locality. — Missouri Kiver, a little above mouth of Poplar 
River, northeastern Montana. 

Characters. — Size huge; skull long and massive; claws long, mod- 
erately or slightly curved, and smoothly polished; usually streaked 
lengthwise with whitish or yellowish, which increases with age until 
in some old individuals the claws are almost wholly white or whitish. 
Color variable, usually light. 

Cranial characters. — Old male (topotype, No. 202739) from Breaks 
of Missouri River, about 100 miles north of Fort Miles, eastern Mon- 
tana, April 4, 1890; killed and presented by E. S. Cameron and wife: 
Skull huge (total length, 400 mm.=15| inches) ; vault of cranium 
moderately arched; zygomata rather squarely spreading posteriorly, 
moderately outbowed, the squamosal part rather broadly expanded 
vertically ; frontal shield rather broad, flattish, sloping gradually up- 
ward to apex, the posterior part broader than usual (not cut away on 
sides by incurving temporal ridges) ; sagittal crest long, straight on 
top, high posteriorly, reaching forward over posterior fourth of fron- 
tals; postorbital processes large, flat, and horizontally outstanding; 
rostrum high, ascending posteriorly ; f ronto-nasal region rather high 
but distinctly dished and somewhat sulcate; palate long, concave in 
cross section ; postpalatal shelf broad and flat, well rounded on sides ; 
underjaw large and rather massive; coronoid blade high, its apex not 
strongly produced posteriorly; ramus rather flat; diastema long. 



18 NORTH AMERICAN FAUNA. [No. 41. 

Teeth large, especially lower canine and last upper and middle lower 
molar. Skull of female (No. 13245, from head of Big Porcupine 
Creek [not Porcupine River] between Musselshell and Yellow- 
stone Rivers, eastern Montana) : Of generalized grizzly type; 
large and rather massive; vault of cranium rather flat; braincase 
rather broad; frontal shield of moderate breadth, slightly convex, 
slightly depressed or sulcate between orbits, elongate-lyrate pos- 
teriorly; postorbital processes weak and slightly decurved; sagittal 
crest not yet reaching fronto-parietal suture; muzzle moderate or 
rather short; zygomata moderately spreading and rather angular; 
palate rather narrow. Teeth large ; M- 2 - large and subrectangular. 

Cranial comparisons. — Adult male (No. 202739, topotype) com- 
pared with two adult males of bairdi, one from near Cheyenne, south- 
eastern Wyoming (Yountz), the other from Blue River, Summit 
County, Colorado (No. 203805): Size slightly larger; fronto-nasal 
region (interorbitally) dished instead of elevated, not compressed in 
front of orbits; frontal shield slightly broader, rising from plane 
of nasals (in bairdi continuing plane of nasals) ; palate longer and 
broader; postpalatal shelf much broader and more rounded on sides; 
mastoids short, appressed, vertical (in bairdi longer and divergent) ; 
meatus tube compressed between mastoid and glenoid (in bairdi not 
compressed); underjaw longer; ramus longer and flatter (much 
less swollen on outer side) ; coronoid blade less falcate (notch shal- 
lower) ; lower canine and last upper molar larger. 

Male adult compared with male adult absarokus (type) : Size de- 
cidedly larger; vault of cranium less arched; frontal shield rising 
more gradually; rostrum longer; palate broader and much longer; 
ramus much longer; meatus tube compressed between mastoid and 
glenoid (in absarokus not compressed). Teeth much larger, espe- 
cially lower canine, last upper molar, and middle lower molar. 

Remarks. — Until recently the absence of authentic specimens from 
the neighborhood of the type locality, in connection with the pres- 
ence of several species of grizzly in Montana, caused an embarrassing 
uncertainty as to which species was entitled to the name horribilis. 
But the slow accumulation of material during the long period in 
which I have been engaged in a study of the group made it possible 
to map the ranges of some of the species with some degree of con- 
fidence; and finally, through the generosity of Mr. and Mrs. E. S. 
Cameron, of Marsh, Montana, I have been presented with a splendid 
skull of an old male horribilis from the Breaks of the Missouri, 
about 100 miles north of Fort Miles, Montana (practically the type 
locality). This skull proves that the huge buffalo-killing grizzly 
of the Great Plains bordering the Missouri in eastern Montana and 
the Dakotas — the "White Bear" of Lewis and Clark — is really the 
species to which Ord in 1815 gave the name Ursus horribilis. 



1918.] HOREIBILIS GKOUP. 19 

Skull measurements. — Old male (No. 202739, from Missouri Breaks, 
eastern Montana): Basal length, 351; occipito-nasal length, 350; 
palatal length, 192 ; zygomatic breadth, 247 ; interorbital breadth, 86. 

URSUS HORRIBILIS BAIRDI Mebriam. 1 

Baied Grizzly. 

Ursus bairdi Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 192-193, August 
13, 1914. 

Type locality. — Blue River, Summit County, Colorado. 

Type specimen. — No. 203805, S old, U. S. National Museum, Bio- 
logical Survey collection (=3147, Warren collection). 

Range. — Southern Rocky Mountain region from San Juan Moun- 
tains, southwestern Colorado, northward through Wyoming to Mon- 
tana, and perhaps to southeastern British Columbia. Probably a 
mountain animal, while its neighbor horribilis was a plains species. 

Characters. — Size large — in the Rocky Mountain region exceeded 
only, if at all, by horribilis; skull long, with narrow elevated fronto- 
nasal region ; claws of moderate length, smooth. 

Cranial characters. — Old male (type) from Blue River, Colorado, 
and a closely similar old male from Sabille Hole, Laramie County, 
southeastern Wyoming, a little north of Cheyenne : Size large ; fronto- 
nasal region high and rather narrow; rostrum rather long, narrow, 
and strongly compressed in front of orbits; face long sloping; frontal 
shield flat, short pointed, faintly depressed medially ; shield and nasals 
in essentially same plane except that anterior third of nasals is 
slightly upturned; postorbital processes large, outstanding, flat, and 
blunt; orbital rims prominent; sagittal crest moderately high pos- 
teriorly; temporal impressions short, incurved, beaded; zygomata 
strongly outbowed, squarely spreading posteriorly; lachrymal duct 
notching orbital rim ; squamosal shelf short, arched over meatus, the 
free edge thickened ; palate and postpalatal shelf of moderate breadth ; 
mastoids rather long, divergent. Molars large, especially M- 2 -. 

Immature males, up to at least the fifth or sixth year, have rather 
narrow convex frontal shields with weak decurved postorbital proc- 
esses, and may be recognized at once by the form of the f ronto-nasal 
region, which is high, narrow, and strongly pinched in immediately 
in front of the orbits. 

Cranial comparisons. — Old male compared with old male horribilis 
(No. 202739, from Missouri Breaks, eastern Montana) : Size essentially 
the same; frontal shield slightly lower posteriorly, higher anteriorly, 
the point lyrate and more slender ; f ronto-nasal region elevated (never 
dished or sulcate between orbits) ; base of rostrum much more com- 

1 Named for S. F. Baird, former Secretary of the Smithsonian Institution and founder 
of the U. S. National Museum. 



20 NORTH AMERICAN FAUNA. [No. 41. 

pressed laterally in front of orbits ; palate narrower ; mastoids longer 
and more divergent; meatus tube broadly rounded and free (not 
compressed between mastoid and glenoid) ; under jaw shorter; ramus 
more swollen on outer side ; inferior border of ramus shorter and less 
upturned; coronoid blade more recurved, the apex narrower and 
reaching farther backward. Teeth similar but slightly smaller; 
M 1 - and M> smaller and less massive. 

Skull measurements. — Old male (type) : Basal length, 348; occipito- 
nasal length, 325; palatal length, 157; zygomatic breadth, 235; inter- 
orbital breadth, 81. 

URSUS HORRIBILIS IMPERATOR Mereiam. 

/ 
Yellowstone Pabk Big Grizzly. 

Ursus imperator Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 180-181, 
August 13, 1914. 

Type locality. — Lake Hotel, Yellowstone National Park, Wyoming. 

Type specimen. — No. 176297, $ old, U. S. National Museum. Died 
in National Zoological Park. 

Range. — Yellowstone National Park, Wyoming; limits unknown. 

Characters. — Size large; skull massive, with large horizontally out- 
standing postorbitals ; closely similar to horribilis. 

Cranial characters. — Old male (type) and a much finer male (No. 
216205) : Size large (only slightly smaller than horribilis) ; skull 
massive; vault of cranium rather high; fronto-nasal region moder- 
ately dished; sagittal crest long, high, and straight; frontal 
shield exceedingly short, flattish, shallowly sulcate medially, 
slightly swollen over orbits, ending in short obtuse point about 30 
millimeters behind plane of postorbitals; postorbitals long, peglike, 
horizontally outstanding; lachrymal duct opening on orbital rim; 
zygomata moderately outstanding and outbowed, the squamosal base 
broadly expanded; squamosal shelves broad, arched over meatus; 
palate moderate; postpalatal shelf rather long; occipito-sphenoid 91 
mm.; mastoids short; underjaw large, long, and massive, with high 
vertical coronoid blade. Teeth large : M. 2 - with long heel. 

Cranial comparisons. — Two old males (Nos. 176297 and 216205) 
from near northern boundary of Yellowstone National Park 1 com- 
pared with an old male topotype of horriMUs (No. 202739) from 
Missouri Breaks, eastern Montana : Size only slightly less, general 
appearance essentially same; frontal shield slightly broader inter- 
orbitally and decidedly broader across postorbital processes; postor- 

1 While both are old males, the type is the older of the two and the recessive changes 
incidental to senility have already begun. It differs from the other in having the frontal 
shield shorter pointed posteriorly (with consequent lengthening of sagittal crest), the 
rostrum narrower, the fronto-nasal region more strongly dished. 



1918.] HORRIBILIS GROUP. 21 

bital processes more widely outstanding horizontally; palate and 
postpalatal shelf narrower; mastoids less appressed (apex farther 
from glenoid surface, leaving broader space for tube of auditory 
meatus). Last upper molar somewhat smaller, heel equally long but 
narrower posteriorly ; middle lower molar narrower. 

Old males (type and topotype) compared with old male bairdi: 
Size essentially same; frontal shield broader throughout; distance 
between tips of postorbital processes much greater; fronto-nasal 
region more dished (in bairdi elevated) and less compressed; inion 
more strongly developed. 

Old male (type) compared with adult male absarokus (type) : 
Size slightly larger; frontal shield broader and flatter; postorbital 
processes longer and more widely outstanding; rostrum higher and 
more nearly horizontal (in absarokus lower anteriorly and more 
sloping) ; sagittal crest longer; squamosal root of zygoma very much* 
more expanded vertically; palate much longer. 

Skull measurements. — Old male (type) : Basal length, 336 ; occipito- 
nasal length, 317; palatal length, 191; zygomatic breadth, 232; inter- 
orbital breadth, 90. Another and more perfect old male from Yel- 
lowstone Park (No. 216205): Basal length, 340 j 1 occipito-nasal 
length, 332 ; palatal length, 186 ; zygomatic breadth, 230 ; interorbital 
breadth, 90. 

URSUS CHELIDONIAS sp. NOV. 
Jeevis Inlet Grizzly. 

Type No. 223133, $ old, U. S. National Museum, Biological Sur- 
vey collection. Collected on river at head of Jervis Inlet, British 
Columbia, in 1916, by Forrest and Fred Johnstone. 

Characters. — Size very large; external characters unknown; skull 
of male strikingly like that of imperator but teeth, especially upper 
molars, very much smaller, resembling those of kwakiutl. 

Cranial characters. — Adult male (type) : Skull very large, massive, 
flat on top, with horizontally outstanding or slightly uplifted post- 
orbital processes ; shield moderately broad, flat, the point long, reach- 
ing back to fronto-parietal suture; rostrum elevated in same plane 
with shield; sagittal crest short, confined to parietals; zygomata 
moderately outstanding and outbowed ; nares small ; postpalatal shelf 
rather broad; occipito-sphenoid unusually long (102) ; basicranial 
axis flat ; mastoids long and spreading ; meatus tubes large and free ; 
underjaw large and massive ; coronoid blade high and nearly vertical ; 
ramus rather broad posteriorly, its inferior border concave under 
anterior molar, convex behind plane of last molar; subangular 

1 In part restored. 



22 NORTH AMERICAN FAUNA. [No. 41. 

border short ; canines moderate, rather small for size of skull ; upper 
and lower molars small; heel of M> short and rather broadly 
rounded ; last lower premolar of grizzly type. 

Cranial comparisons. — Ursus chelidonias requires comparison with 
only two species, imperator and warburtoni. Adult male (type) 
compared with equally old male imperator from Hell Roaring Creek 
near northern boundary of Yellowstone National Park: Size 
slightly greater; general form and proportions almost identical; 
point of shield somewhat longer ; rostrum more completely in frontal 
plane; zygomata more outbowed anteriorly; basicranial axis flatter 
(in imperator somewhat arched) ; mastoids longer and more diver- 
gent; meatus tube longer; postpalatal shelf broader; coronoid blade 
higher; canines about same size; molars (both upper and lower) 
smaller, the difference most marked in M 1 , M 1 , and M T . 

Compared with two adult males of warburtoni, one (No. 223946) 
from Iskut River near its junction with the Stikine; the other (No. 
210142) from Chilkat River valley, Southeastern Alaska: Skull as a 
whole slightly larger ; frontal shield more nearly horizontal, somewhat 
broader, and completely flat (in warburtoni slightly convex) ; zygo- 
mata more outbowed anteriorly; postpalatal shelf broader; basi- 
cranial axis flatter; occipito-sphenoid longer (102 mm., contrasted 
with 95); palate longer; postpalatal shelf broader; under jaw more 
massive; coronoid blade higher (posterior part of jaw more uplifted, 
raising condyle and coronoid) ; upper canines conspicuously more 
slender; lower canines much smaller; upper and lower molars 
smaller ; middle lower molar not only shorter but differing markedly 
in proportions, the posterior moiety small and narrow (14.5) in com- 
parison with the anterior (17) ; PM^more distinctly of grizzly type; 
M T with saddle open (cusplet on inner side nearly obsolete). 

Skull measurements. — Old male (type) : Basal length, 346 ; occipito- 
nasal length, 337 ; palatal length, 191 ; zygomatic breadth, 240 ; inter- 
orbital breadth, 91. 

URSUS ATNARKO sp. NOV. 

Atnabko Gbizzly. 

Type No. 211452, 6* ad., U. S. National Museum, Biological Sur- 
vey collection. From Lonesome Lake, 1 Atnarko River, one of 
the upper forks of the Bella Coola, British Columbia. Collected in 
September, 1915, by E. H. Edwards. 

1 Lonesome Lake la about 30 miles from the Junction of the Whitewater, or Talchawko, 
and the Atnarko, which two rivers unite to form the Bella Coola. The Bella Coola Is 
about 45 miles long. Lonesome Lake is nearly on the fifty-second parallel, and by the 
river about 75 miles from the head of Burke Channel. 



1918.] HOREIBILIS GROUP. 23 

Characters. — Size large; external characters unknown; skull of 
male long and narrow, similar in general to that of kwakiutl but 
much narrower; skull of female surprisingly different from that of 
kwakiutl, being long, low, and very narrow, while kwakiutl is ex- 
ceptionally high and broad. 

Cranial characters. — Adult male (type) : Skull long, low, and 
narrow, highest in posterior frontal region; braincase anteriorly 
keeling into sagittal crest; shield narrow and rather sharp pointed 
posteriorly ; postorbitals moderate, horizontally outstanding ; rostrum 
long and slender, rising gradually into plane of shield ; zygomata not 
widely outstanding ; sagittal crest long ; palate and postpalatal shelf 
long and narrow; lower jaw long; coronoid blade moderate; sub- 
angular border short. Dentition weak: canines small for size of 
skull (as in kwakiutl) ; molars even smaller than in kwakiutl. 
Adult female (based on 3 adult and old females, Nos. 223177, 223182, 
and 223183) from Atnarko River: Skull long, low, slender, nearly 
flat on top; shield long and narrow, not rising above plane of ros- 
trum; postorbitals weak, outstanding; rostrum long and slender, 
passing insensibly into frontal plane; zygomata appressed; palate 
long and narrow ; coronoid blade moderate ; subangular border short. 
Dentition weak. 

Cranial comparisons. — Compared with kwakiutl the skull as a whole 
is longer and narrower; frontal shield much narrower and more 
evenly sloping (rising less abruptly from rostrum) ; postorbitals 
much less widely outstanding; posterior frontal region higher, keel- 
ing into sagittal crest; palate and postpalatal shelf notably longer 
and narrower. 

Adult and old female compared with old female kwakiutl of 
slightly greater basal length: Breadth very much less (across post- 
orbitals 105 or less, contrasted with 130) ; shield low, narrow, and 
flat instead of broad, high, and rising abruptly from rostrum ; post- 
orbitals much smaller and weaker; zygomata appressed instead of 
outstanding; palate, postpalatal shelf, and palatal notch very much 
narrower ; sagittal crest much weaker. 

Remarks.— The males of kwakiutl and atnarko are so much alike 
that their differences would naturally be regarded as of only sub- 
specific weight; but the females are so strikingly different that it 
seems necessary to give them independent specific rank. Their spe- 
cific distinctness is the more certain by reason of the geographic 
contiguity of the localities where they were killed, the type localities 
being on the same watercourse and not more than 75 miles apart. 

Skull measurements. — Adult male (type) : Basal length, 345 ; * 
occipito-nasal length, 325; palatal length, 199; zygomatic breadth, 
214; interorbital breadth, 75. 

1 Partly restored. 



24 NORTH AMERICAN FAUNA. [No. 41. 

URSUS KWAKIUTL Merriam. 

Kwakitjtl Grizzly. 

Ursus kwaJciutl Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 143-144, Sep- 
tember 6, 1916. 

Type locality. — Jervis Inlet, coast of southern British Columbia. 

Type specimen. — No. 211748, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected May 17, 1916, by Fred Mansell. 

Range. — Coast region of British Columbia from southwestern cor- 
ner (Burrard Inlet, Howe Sound, Jervis Inlet) northwesterly to or 
beyond the lower Bella Coola. 

Characters. — Size large; color dark; ears densely furred; claws 
unknown ; skull long, but little arched. 

Color. — Skin of head of adult male (type) : Nose brown ; head 
and face from front of eyes posteriorly very dark brown, darkest 
on ears, slightly grizzled on occiput by golden-tipped hairs. 

Cranial characters. — Adult male (type): Size large; skull long, 
rather low and narrow, with long high rostrum, gradually ascend- 
ing frontal shield, rather low fronto-parietal region, and strongly 
outstanding postorbitals. Frontal shield of moderate breadth, shal- 
lowly sulcate medially, swollen over orbits, short pointed posteriorly ; 
rostrum long, high, and rather narrow; nasals flattened, nearly 
horizontal; fronto-nasal region sloping gently in plane of shield; 
braincase long, its anterior part keeling into elongate sagittal crest; 
zygomata moderately spreading (becoming, of course, more strongly 
outbowed in old age) ; palate long, concave, slightly arched antero- 
posteriorly ; postpalatal shelf large and broad, nearly square ; denti- 
tion rather light for size of skull; underjaw long, its inferior border 
slightly sinuous (slightly bellied under last molars) ; coronoid blade 
broad at base, not very high, the apex slightly produced posteriorly 
but falling far short of plane of condyle. 

Old female (No. 215432) : The skull of an old female from Kwatna, 
on the lower Bella Coola, is obviously of this species. It is large 
and rugged for a female, with strongly dished fronto-nasal region 
and broad massive frontal shield rising abruptly from a very small 
rostrum; shield sulcate, swollen between sulcus and orbits, lyrate 
pointed, ending at fronto-parietal suture ; postorbitals large and out- 
standing; zygomata outstanding; sagittal crest strongly developed 
for a female; palate and postpalatal shelf very broad; teeth small; 
PM T badly worn but apparently subcorneal, as in a male (No. 215433) 
from the same locality. 

Cranial comparisons. — The only species requiring comparison with 
kwakiutl are the much larger nortoni and the very different pervagor. 
Adult male (type) compared with adult male nortoni (No. 213705) 
from southeast side Yakutat Bay: Similar in general but much 



1918.] HORRIBILIS GROUP. 25 

shorter; rostrum slightly longer; hraincase materially shorter; 
shield much less elevated; fronto-nasal region much less dished; 
postpalatal length much less. Compared with male pervagor: Skull 
longer and less highly arched; braincase and rostrum materially 
longer ; frontal shield shorter pointed posteriorly ; rostrum more ele- 
vated anteriorly; nasals more nearly horizontal, shorter posteriorly, 
longer anteriorly; postpalatal notch longer; under jaw decidedly 
longer and more massive, but inferior border of ramus from symphy- 
sis to tubercle of essentially same length; subangular border much 
longer. Dentition heavier (both upper and lower canines, incisors, 
and molars larger). 

Skull measurements. — Adult male (type) : Basal length, 340; 
occipito-nasal length, 330; palatal length, 184; zygomatic breadth, 
212 ; interorbital breadth, 85. 

URSUS NORTONI 1 Mebriam. 

Yakutat Grizzly. 

(PI. XIII.) 

Ursus nortoni Merrtam, Proc. Biol. Soc. Washington, XXVII, pp. 179-180, April 
13, 1914. 

Type locality. — Near Yakutat village, southeastern side Yakutat 
Bay, Alaska. 

Type specimen. — No. 178763, $ ad., U. S. National Museum, 
Biological Survey collection. (Mother of $ cub of second year, 
No. 178764.) Collected May 15, 1910, by G. Frederick Norton. 

Range. — Limited apparently to coastal plain on southeastern side 
of Yakutat Bay. 

Characters. — A true grizzly, apparently of the califomicus- 
kwakiutl group; of large size, with smoothly polished horny claws. 
Fourth lower premolar large and of ultra-grizzly type (long heel 
with continuous sulcus and complete lateral ridges ending in up- 
turned posterior cusplets) ; molars, especially M- 2 -, small for size of 
skull. Coloration normal for a grizzly; skull large, massive, and 
very broad. 

Color. — Of pale grizzly type; head grizzled yellowish or golden 
brown; muzzle pale brown; neck and shoulders to middle of back 
pale buffy from the long whitish buff-tipped hairs, giving the skin 
viewed from behind a decidedly whitish appearance; hinder back 
and rump dark, well washed with pale brown tips; lower part of 
legs and feet dark brown; back of fore feet browner and not so 
dark ; underchin and throat pale soiled buffy whitish. 

1 Named for G. Frederick Norton, who collected and presented the type specimen. 



26 NOBTH AMERICAN PATJNA. [No. 41. 

Cranial characters. — Adult male (No. 213705, from southeast side 
of Yakutat Bay, northeast of Yakutat village) : Skull large, 
long, massive, and strongly dished; braincase exceptionally long; 
facial part of skull relatively short; frontal shield rather broad, 
moderately convex transversely and sulcate medially (swollen 
between sulcus and postorbitals), rising strongly and abruptly from 
rostrum; postorbital processes well developed, slightly arched; ros- 
trum rather short, horizontal; zygomata moderately spreading, 
outbowed; sagittal crest highly developed; squamosal shelves broad 
and long; palate and postpalatal shelf rather long and of moderate 
breadth; palate troughed between molar series; mastoids moderate; 
underjaw massive; coronoid blade high, broad, and not recurved; 
ramus long and straight. Teeth small for size of skull; M- 2 - rather 
short, the heel evenly emarginate and broadly rounded posteriorly. 

Adult female (type) : Rather massive, moderately dished, broad, 
with large broadly rounded braincase, bowed (but not widely out- 
bowed and not angular), moderately spreading zygomata; short 
weak sagittal crest confined to posterior half of parietals; rather 
broad lambdoid crest defining a rather broadly open groove or sulcus 
continuous with that of the deeply sulcate and broadly expanded hori- 
zontal shelf of the squamosal ; squamosal base of zygoma moderately 
expanded; frontal shield broad and long, its lyrate point reaching 
back to middle of parietals, broadly sulcate between orbits; fronto- 
nasal region moderately dished; postorbital processes large and 
broadly outstanding, infraorbital process of jugal and infra jugal 
process of maxillary well developed (probably not constant) ; ros- 
trum broad, short, and rather depressed; lachrymal opening within 
orbit; palate and postpalatal shelf broad, the palate rather deeply 
concave from incisive foramina to plane of hind molars ; postpalatal 
notch broad and rather squarely truncate; occipito-sphenoid length 
90 (about equal to distance from posterior rim of alveolus of outer 
incisor to front of last upper molar) ; basioccipital considerably 
longer than basisphenoid ; occiput broad, rather low, truncate ; coro- 
noid blade rather high. Basicranial axis nearly straight and hori- 
zontal, parallel to palatal axis. Dentition light in both sexes ; canines 
and molars rather small for size of skull ; PM^ of ultra grizzly type, 
with slender cone, well-developed completely sulcate heel ending pos- 
teriorly in pair of upturned cusplets ; PM> large and broad ; heel of 
M^ rather short, not strongly narrowed posteriorly, third cusp 
small; incisors rather large. 

Cranial comparisons. — Male adult (type) compared with male 
adult kwakiutl (type) : Similar in general hut very much longer both 
basally and on top; rostrum slightly shorter; braincase materially 
longer; frontal shield rising higher and more abruptly from ros- 



1918.] 



HOBRTBILIS GROUP. 27 



tram; fronto-nasal region more conspicuously dished; postpalatal 
length very much greater; under jaw longer and more massive; 
ramus longer; subangular border shorter; coronoid blade much 
larger and higher ; teeth closely similar, but upper canines and upper 
molars slightly larger. 

Specimens examined. — About a dozen specimens have been ex- 
amined, mainly females and young, all from the coastal plain south- 
east of Yakutat Bay. Three (including type) were collected by G. 
Frederick Norton, and by him presented to the Biological Survey; 
others were collected by Miss Annie M. Alexander, and are in the 
Museum of Vertebrate Zoology, University of California ; still others 
(including a splendid adult male) were obtained from Yakutat In- 
dians by E. M. Axelson and purchased by me. The localities are: 
Peninsula between Yakutat and Disenchantment Bays (northeast of 
Yakutat village) ; near Yakutat village; front of Yakutat Glacier; 
Ankow River ; Setuk River ; and Anklin River. 

Skull measurements. — Adult male (No. 213705) : Basal length, 
353; occipito-nasal length, 346; palatal length, 185; zygomatic 
breadth, 241; interorbital breadth, 91. Adult female (type) : Basal 
length, 306; occipito-nasal length, 284; palatal length, 165; zygo- 
matic breadth, 210 ; interorbital breadth, 80.5. 

URSUS WARBURTONI * Mekriam. 

Warbtjrton Pike Geizzly. 

Ursus Jcwakiutl toarburtoni Merriam, Proc. Biol. Soc. Washington, XXIX, p. 145, 
September 6, 1916. 

Type locality. — Atnarko River, British Columbia. 

Type specimen. — No. 210576, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected July 15, 1915, by E. H. 
Edwards. 

Range. — Coast region (but perhaps not the immediate coast strip) 
of Southeastern Alaska and adjacent parts of British Columbia from 
Chilkat River southeasterly to Atnarko River, one of the upper 
forks of the Bella Coola (skulls of adult males examined from At- 
narko River, Stikine River, Iskut River near junction with Stikine, 
and Chilkat River valley). 

Cranial characters. — Adult males : Skull large and massive, rather 
long and flattish on top, not arched. Similar to male Jcwakiutl but with 
much broader (less peglike) and flatter postorbitals, flatter frontal 
shield (not deeply concave in old age), much shorter sagittal crest, 
somewhat heavier dentition, especially broader and more massive 

1 Named in honor of the late Warburton Pike, author of The Barren Grounds of North- 
ern Canada, and The Subarctic Forest, who obtained a fine large typical skull (No. 
223946) on the Iskut a few miles from its junction with the Stikine. 



28 NORTH AMERICAN FAUNA. [No. 41. 

M^. Easily distinguished from stikeenensis of the same region by 
the much greater length of skull and under jaw and lesser elevation of 
frontal region. 

Skull measurements. — Old male (type) : Basal length, 340; oc- 
cipito-nasal length, 324 ; palatal length, 185 ; zygomatic breadth, 233 ; 
interorbital breadth, 85. Old male (No. 223946) from Iskut Eiver, 
a branch of the Stikine: Basal length, 326; occipito-nasal length, 
340; palatal length, 176; zygomatic breadth, 230; interorbital 
breadth, 86.5. 

URSUS NEGLECTUS Merriam. 

Admiralty Island Grizzly. 

Ursus kwakiutl neglecims Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 
144-145, September 6, 1916. 

Type locality. — Near Hawk Inlet, Admiralty Island, Southeastern 
Alaska. 

Type specimen. — No. 209889, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected April, 1914, by W. H. Spaulding. 

Cranial characters. — Skull rather large and massive, flat topped, 
with rather broad outstanding postorbitals. Frontal shield moder- 
ate, flattish, shallowly sulcate, rather short pointed; fronto-nasal 
region elevated in plane of shield and slightly compressed ; rostrum 
moderate, high; zygomata moderately outstanding and somewhat 
bowed; postpalatal shelf short and broad; notch rather broad. 
Underj aw rather short, ramus rather flat and broad vertically, 
especially posteriorly, strongly bellied posteriorly; coronoid blade 
high and narrow, its apex not reaching plane of condyle. Teeth 
moderate. 

Cranial comparisons. — Ursus neglectus requires comparison with 
both kwakiutl and warburtoni. Old male (type) compared with 
adult male kwakiutl (type from Jervis Inlet) : Size similar; shield 
flatter, more nearly horizontal and less sloping anteriorly ; postorbit- 
als broader and flatter on top ; fronto-nasal region elevated instead of 
depressed; rostrum larger and more elevated; zygomata less out- 
bowed; nasals more produced anteriorly, projecting broadly over 
nares (about 7 mm. beyond premaxillse at point of contact) ; nares 
more truncate: underjaw and subangular border much shorter; 
coronoid blade narrower. Upper canines and crown of last upper 
molar longer. 

Old male (type) compared with three old males of warburtoni 
(from Atnarko and Iskut Rivers and Chilkat Valley) : Size slightly 
smaller but occipito-sphenoid length same ; top of skull more nearly 
horizontal (shield anteriorly and rostrum more elevated) ; zygomata 



1918.] HOERIBILIS GROUP. 29 

less outbowed; nasals more projecting anteriorly; palate shorter; 
postpalatal shelf broader, flatter, and much shorter; mastoids 
shorter; under jaw slightly smaller; subangular border much shorter; 
coronoid blade narrower and more nearly vertical. Canines (espe- 
cially upper) more slender; M^ narrower and less massive but 
difference not great. 

Skull measurements. — Old male (type) : Basal length, 322 ; oc- 
cipito-nasal length, 325 ; palatal length, 177 ; zygomatic breadth, 229 ; 
interorbital breadth, 83. 

URSUS CALIFORNICUS Merbiam. 
California Coast Grizzly. 

[Ursus horriceus] subspecies califonvicus Merriam, Proc. Biol. Soc. Washing- 
ton, X, pp. 76-77, April 13, 1896. 

Ursus californicus Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 186, 188, 
August 13, 1914. 

Type locality. — Monterey, California. 

Type specimen. — Skull No. 3630, $ old, U. S. National Museum. 

Range. — Humid coast region of California from San Francisco 
Bay south about to San Luis Obispo (apparently passing into 
tularensis in the dryer interior) . 

Characters. — Size large; claws long and smooth; pelage variable. 
Dentition heavy ; last upper molar large, its heel long and broad. 

Cranial characters. — Adult male: Skull long and narrow; vault of 
cranium flat — not arched; frontal shield flat (or concave medially), 
short pointed posteriorly, gradually sloping almost in same plane 
with rostrum (dishing slight) ; temporal impressions beaded; postor- 
bitals massive and moderately spreading ; zygomata slightly bowed ; 
rostrum long, slightly compressed in front of orbits; nasals nearly 
horizontal, slightly rising posteriorly in plane of shield; frontal 
shield moderately broad, slightly swollen over orbits, the point lost 
in sagittal crest 25 to 50 mm. anterior to fronto-parietal suture; 
postorbital processes strongly developed, subtriangular, rather mas- 
sive, outstanding, and slightly decurved over orbits; sagittal crest 
strongly developed and nearly straight; squamosal shelf broad; 
squamosal base of zygoma vertically expanded in aged skulls ; frontal 
part of braincase in young-adult skulls somewhat elevated and tend- 
ing to "keel" into crest; mastoid processes long; interpterygoid 
fossa long and usually narrow; under jaw long and massive, ramus 
broad vertically. Teeth large and heavy ; M- 3 - subrectangular, heel 
long, broad posteriorly. Skull of female: Similar to that of male, 
but smaller. 

Cranial comparisons. — Adult and old male compared with adult 
male klamathensis (type) : Skull longer ; zygomatic breadth essen- 



30 NORTH AMERICAN FAUNA. [No. 41. 

tially the same; face and rostrum longer; rostrum more compressed 
in front of orbits; fronto-nasal region more dished; frontal shield 
more concave medially and more swollen at orbital rims ; postorbital 
processes much more massive, slightly arched, the tips slightly de- 
curved; temporal beads more strongly incurved; sagittal crest 
shorter posteriorly; lambdoid crest more outstanding laterally; oc- 
cipital overhang much less; squamosal shelf behind zygoma much 
shorter; occipito-sphenoid and palate longer; mastoids much longer 
and more strongly outstanding. Under jaw much longer. Dentition 
heavier, the last upper and middle lower molar in particular much 
larger ; heel of M 1 very broad and broadly rounded posteriorly. 

Old male from Monterey, compared with old male colusus (type, 
from Sacramento River) : Frdnto-nasal region slightly higher and 
less depressed; rostrum higher, less depressed, and less horizontal; 
postorbitals much larger and more swollen; orbital rims more 
swollen ; orbits more nearly vertical, squamosal base of zygoma more 
expanded; palate narrower anteriorly; ramus much broader verti- 
cally, its inferior border less convex in middle part and more bellied 
posteriorly; apex of coronoid more produced posteriorly; angular 
process larger and longer; mastoids longer and directed less ante- 
riorly, not reaching so near glenoid process. Dentition stronger and 
strikingly different : canines larger ; upper incisors and both upper and 
lower molars very much larger; heel of M^ large and not normally 
emarginate. 

/Skull measurements. — Old male (type): Basal length, 361; oc- 
cipito-nasal length, 346; palatal length, 196; zygomatic breadth, 
224 ; interorbital breadth, 82. 

URSUS TULARENSIS Mebbiam. 

Tejon Gbizzlt. 

Ursus californicus tularensis Merriam, Proc. Biol. Soc Washington, XXVII, 
p. 188, August 13, 1914. 

Type locality. — Fort Tejon, Canada de las Uvas, Tehachapi 
Mountains, California. 

Type specimen. — No. 3536, £ old, U. S. National Museum. Col- 
lected by John Xantus. 

Range. — Dry chaparral hills of interior coast ranges between the 
San Joaquin Valley and Los Angeles plain, comprising the Teha- 
chapi, Tejon, Sierra Madre, and San Gabriel Ranges, and probably 
San Bernardino Mountains also, and ranging northward an un- 
known distance, doubtless covering the San Rafael and Gabilan 
Ranges, and southern part of the Diablo Range ; limits unknown. 

Characters. — Size large, but smaller than californicus; claws of 
grizzly type, but those of a female dark, thick, and broad for a fe- 



1918.] HORRIBILIS GROUP. 31 

male, rather straight (tips worn off), straighter and broader than in 
female magister. 

Color. — Nearly full-grown male killed by Walter Richardson, near 
head of Tejunga Canyon, San Gabriel Mountains, Southern Cali- 
fornia, in 1897: General color very dark brown, almost dusky; 
grizzled on upperparts by admixture of pale-tipped hairs; muzzle 
reddish brown. 

Cranial characters. — Old male: Skull large, rather broad and flat 
frontally, moderately dished, moderately high, with large outstand- 
ing postorbitals and beaded temporal ridges. Frontal shield rather 
broad, swollen, and somewhat elevated on orbital rims and base of 
postorbitals, depressed interorbitally, sloping gradually into rostrum ; 
rostrum large and rather high ; f ronto-nasal region not depressed ; 
sagittal crest rather short, elevated and produced posteriorly ; occipi- 
tal overhang marked; zygomata moderately spreading, subtriangu- 
lar; palate and postpalatal shelf broad and flat, mastoids rather long 
and inclined strongly forward. Under jaw large and rather mas- 
sive; ramus broad vertically, strongly bellied posteriorly; coronoid 
rather high, its apex not strongly recurved. Teeth large; M> sub- 
rectangular, the long heel only slightly emarginate on outer side. 
Adult and old females : Skulls more easily distinguished than in males 
from their nearest relative, calif ornicus. The skull of the female is 
much more like the normal female grizzly type, not resembling the 
males as does the female of calif ornicus. 

Cranial comparisons. — Adult and old females compared with fe- 
male calif ornicus: Skull decidedly smaller; frontal shield behind 
plane of postorbital processes more elevated and convex — not de- 
pressed and concave as in calif ornicus; point of shield longer and 
broader ; orbital rims more swollen ; sagittal crest shorter and lower ; 
palate at least 10 mm. shorter; occipito-sphenoid about 10 mm. 
shorter; underjaw smaller and lighter; last upper molar decidedly 
smaller. Normal M^ subrectangular as in calif ornicus (in henshawi 
subtriangular and small). 

Adult and old males compared with male calif ornicus: Similar but 
smaller; base of cranium shorter; palate slightly shorter; occipito- 
nasal length decidedly less; braincase decidedly shorter; zygomatic 
breadth same or slightly greater — the skull as a whole relatively 
broader than in calif ornicus; postpalatal shelf broader; underjaw 
materially shorter; M 1 decidedly smaller (both shorter and nar- 
rower) ; lower series of teeth smaller, M T and M^ particularly 
smaller, much narrower and less massive; heel of last upper molar 
less broad than in calif ornicus. 

Adult male compared with henshawi (type) : Skull larger and 
more massive; fronto-nasal region much higher and much less 
dished; rostrum larger, higher, and not depressed; zygomata more 
64854°— 18 3 



32 NORTH AMERICAN FAUNA. [No. 41. 

broadly spreading; postpalatal shelf broader; coronoid larger and 
less falcate ; ramus broader vertically ; last upper molar much larger, 
the heel long and broad posteriorly, contrasted with the short sub- 
triangular heel of henshawi. 

Skull measurements. — Average of two old males from Fort Tejon, 
California: Basal length, 329; occipito-nasal length, 320.5; palatal 
length, 179.5 ; zygomatic breadth, 228 ; interorbital breadth, 78. Old 
female from Fort Tejon: Basal length, 296; occipito-nasal length, 
287; palatal length, 162; zygomatic breadth, 187; interorbital 
breadth, 75. 

URSUS COLUSUS Merriam. 

Sacramento Valley Grizzly. 

Ursus colusu.s Merriara, Proc. Biol. Soc. Washington, XXVII, pp. 187-188, 
August 13, 1914. 

Type locality. — Sacramento River valley, California (probably 
between Colusa and Sacramento). 

Type specimen. — No. 3837, $ old, U. S. National Museum. Col- 
lected by the Wilkes U. S. Exploring Expedition and marked " C. P. 
Ex. Ex. 6.16 " (the numerals uncertain, there being indication of a 
figure in front of the first 6). The words "Grizzly Bear, Sacra- 
mento " are written on the right parietal in pencil. 

Range. — Sacramento (and perhaps also San Joaquin) Valley and 
adjacent foothills; westerly in the hot inner coast mountains to 
Dobbins Creek canyon on the boundary between southeastern Hum- 
boldt and southwestern Trinity Counties. 

Characters. — Size large, external characters unknown. Skull 
large and long, resembling that of californicus, but teeth smaller 
and last upper molar very different. 

Cranial characters. — Old male: Skull large, long, and low, the 
frontal shield flat, postorbital processes moderate or small, fronto- 
nasal region moderately dished, palate long, sagittal crest high, 
temporal ridges beaded, and occipital overhang pronounced. In gen- 
eral resembling largest skulls of calif omicus but having very much 
smaller teeth and differing in numerous minor cranial characters. 
Young-adult female: One from San Jose Mission (No. 1143, Yale 
Museum) appears to be a not quite grown female colusus. The last 
upper molar is short and subtriangular, the heel emarginate on 
outer side. 

Cranial comparisons. — Old male (type) compared with old male 
calif ornicus (from Monterey) : Fronto-nasal region slightly lower and 
more depressed ; rostrum lower, more depressed and more nearly hori- 
zontal ; postorbitals much smaller and less swollen ; orbital rims less 
swollen; orbits less nearly vertical; squamosal base of zygoma less ex- 



1918.] HORRIBILIS GROUP. 33 

panded ; palate broader anteriorly ; ramus much less broad vertically, 
its inferior border more convex in middle part and less bellied pos- 
teriorly ; apex of coronoid less produced posteriorly ; angular process 
smaller and shorter; mastoids shorter and directed more anteriorly, 
reaching nearer to glenoid process. Dentition weaker and strikingly 
different : canines too badly broken to admit of satisfactory compari- 
son, but obviously smaller ; upper incisors and both upper and lower 
molars very much smaller ; heel of M^ small and strongly emarginate 
on outer side, of same size and approximately same form as in kla- 
mathensis but even more strongly constricted on outer side immedi- 
ately behind second cusp ; lower molars of same size as in klamathen- 
sis though the jaw is much longer. 

Old male (type) compared with klamathensis : Postorbitals, 
length of sagittal crest, form of zygomata, and dentition essentially 
the same, but skull longer; vault of cranium much lower; frontal 
shield narrower and much shorter; temporal beads much more 
strongly incurved; fronto-nasal region dished instead of elevated; 
rostrum much lower, more depressed, and more nearly horizontal; 
ramus longer, less broad vertically ; diastema much longer. 

Skull measurements. — Old male (type): Basal length, 352; oc- 
cipito-nasal length, 337 ; palatal length, 185 ; zygomatic breadth, 228 ; 
interorbital breadth, 82.5. 

URSUS DUSORGUS sp. nov. 1 
Rindsfoos Grizzly. 

Type No. 217426, $ old, U. S. National Museum. From head of 
Jack Pine River near Mount Bess, Alberta (close to British Columbia 
boundary). Collected September 4, 1916, and presented to the Na- 
tional Museum by William Rindsfoos. 

Cranial characters. — Skull large, broad, and massive, with flat 
gradually sloping frontal shield and high sagittal crest; shield 
broad, short pointed, slightly thickened at orbits, with horizontally 
outstanding slightly elevated postorbitals; fronto-nasal region 
faintly dished, the broad slightly depressed rostrum rising imper- 
ceptibly into shield; sinus case keeling into anterior part of sagittal 
crest, the crest rising above plane of top of skull and point of f rontals 
and reaching forward nearly half way from fronto-parietal suture 
to postorbitals; temporal impressions strongly beaded; zygomata 
broadly outstanding; palate and postpalatal shelf rather broad; 
underjaw massive; ramus strongly bellied posteriorly; coronoid 
rather high. Teeth moderate; last upper molar long. 



1 Tentatively included in horribilis group. (See Introduction, pp. 12-13.) 



34 NORTH AMERICAN FAUNA. [No. 41. 

Cranial comparisons. — So far as known dusorgus requires com- 
parison with only a single species — imperator of western Montana 
and the Yellowstone Park region. Old male (type) compared with 
old male imperator (No. 216205) : Similar in general but size some- 
what less; vault of cranium, frontal region, and rostrum lower; 
shield flatter, more strongly sloping, and shorter pointed; rostrum 
more depressed; sagittal crest much higher anteriorly, rising well 
above point of shield; palate shorter; meatus tube longer; ramus of 
jaw shorter. Teeth similar but M- 1 smaller. 

Skull measurements. — Old male (type) : Basal length, 324; oc- 
cipito-nasal length, 322.5 ; palatal length, 180 ; zygomatic breadth, 
227; interorbital breadth, 86. / 

Planiceps Group. 

URSUS NELSONI * Merriam. 

Nelson Grizzly. 

Vrsus nelsoni Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 190-191, 
August 13, 1914. 

Type locality. — Colonia Garcia, Chihuahua, Mexico. 

Type specimen. — No. 99657, 9 ad., U. S. National Museum 
Biological Survey collection. Collected November 13, 1899, by H. A. 
Cluff. 

Range. — Sierra Madre of Mexico from northwestern Chihuahua 
and northeastern Sonora south to southern Durango. 

Characters. — Smallest of the grizzly bears. General color, pale 
buffy yellowish, varying to grayish white, grizzled from darker color 
of underfur. Specimens in worn pelage vary to yellowish brown and 
even rusty. Hairs of throat and flanks longer than elsewhere ; belly 
sparsely haired, lacking the thick underfur of upperparts. Claws 
long, smooth, and moderately curved; brownish horn color streaked 
with yellowish. Longest claw 56 mm. from upper base to tip (tip 
worn off). 

Color. — Type specimen in fresh fall pelage: Muzzle pale brown, 
much darker around eyes; top of head yellowish buff; back grayish 
brown, heavily overlaid with pale buffy gray tips (color more buffy 
across shoulders, more whitish gray on back) ; hump dark brown, 
small; rump grizzled grayish and brown, the light-tipped hairs 
failing posteriorly and on the sides, the dark brown ground color 
passing into blackish brown on thighs, legs, feet, and tail; forelegs 
also blackish brown; lips and point of chin yellowish buff, followed 
by area of dark brown; underneck and underparts generally long 
haired and grizzled, the prevailing color yellowish buff. 

* Named for E. W. Nelson, who collected the original series in the type region. 



1918.] PLANICEPS GROUP. 36 

Cranial characters. — Adult male: Skull small and wolflike. Simi- 
lar in general to texensis, but smaller and more wolflike; rostrum 
strikingly narrow; frontal shield flat, narrow, and only faintly 
sulcate medially; temporal impressions not beaded; occipito-sphe- 
noid length 82 mm. (in $ texensis, 86) ; palate very short (149 mm. 
contrasted with 111 in texensis) ; posterior root of zygoma slender; 
postorbital process slender, peglike, and outstanding horizontally. 
Teeth small. 

Skull measurements.— Old male (No. 16025, Field Mus. Nat. Hist., 
Chicago, from Casas Grandes, Chihuahua): Basal length, 284; 
occipito-nasal length, 264; palatal length, 149; zygomatic breadth, 
199; interorbital breadth, 65. Average of three adult females from 
Colonia Garcia, Chihuahua : Basal length, 260 ; occipito-nasal length, 
243; palatal length, 138; zygomatic breadth, 168; interorbital 
breadth, 61. 

URSUS TEXENSIS TEXENSIS Merbiam. 

Texas Gbizzly. 

Ursus horriceus texensis Merriam, Proc. Biol. Soc. Washington, XXVII, p. 191, 
August 13, 1914. 

Type locality. — Davis Mountains, Texas. 

Type specimen. — No. 203198, $ old, U. S. National Museum, 
Biological Survey collection. Killed November 2, 1890, by C. O. 
Finley and John Z. Means. 

Range. — Restricted, so far as known, to Davis Mountains, Texas, 
and mountains of southern Colorado. 

Characters. — Size small; external characters unknown. Affinities 
with shoshone rather than with horriceus. 

Cranial characters. — Old male (type) : Size small; frontal shield 
low, narrow, flat (slightly depressed medially), short pointed pos- 
teriorly, gently sloping in same plane with rostrum; postorbital 
processes small, peglike, horizontally outstanding; fronto-nasal re- 
gion elevated in fronto-facial plane; rostrum narrow; zygomata 
rather broadly outbowed, moderately expanded vertically, lachrymal 
duct wholly anterior to orbit ; sagittal crest long and nearly straight ; 
occipital overhang marked; palate broad, deeply excavated between 
molars (may be abnormal) ; postpalatal shelf broad and flat; post- 
palatal notch broad and short; meatus tube slightly recurved and 
markedly upturned. Under jaw light; inferior border of ramus 
long, slightly bellied posteriorly; subangular border rather short; 
coronoid blade high, its recurved apex falling short of plane of 
condyle. Teeth rather small; M- 2 - small, with small heel obliquely 
narrowed on outer side. 

Old female (No. 213002, from Navajo Range, near Cromo, Colo- 
rado) : Skull short, dished, the braincase broad; frontal shield nar- 



36 NORTH AMERICAN FAUNA. [No. 41. 

row ; zygomata broadly spreading, slightly bowed ; occiput truncate ; 
and postorbitals slender, widely outstanding. Rostrum small, slen- 
der, tapering, rising posteriorly into flat lyrate shield; postorbital 
processes long, slender, and directed forward as well as outward; 
braincase rather broad and depressed; palate rather short; post- 
palatal shelf broad and flat. Teeth moderate; canines small and 
slender; molars nearly as large as in male, and M> of same form. 

Cranial comparisons. — Adult male (type) compared with adult 
male planiceps (type) : Size smaller throughout; shield much nar- 
rower, less flat, and less nearly horizontal ; f ronto-nasal region dished 
(in planiceps not dished); rostrum much smaller and narrower; 
palate deeply concave between posterior molars (in planiceps flat) ; 
ramus of under jaw more tapering anteriorly; angular processes not 
bellied (in planiceps bellied) ; teeth smaller. 

Old male (type) compared with male shoshone: Size smaller; 
vault of cranium lower and more nearly horizontal; frontal shield 
less sloping; anterior part of braincase broadly depressed (not 
compressed or keeling into crest) ; palate broader. Teeth smaller. 

Old male (type) compared with old male horriceus (type) : Skull 
similar in basal length, but materially smaller, lower, flatter, nar- 
rower, more smoothly rounded, broader across squamosals, much 
less massive, and with wholly different postorbitals. Postorbitals 
small, slender, peglike, and horizontally outstanding instead of 
large, broad, massive, and decurved; frontal shield narrower and 
flatter; orbital rims less swollen; rostrum smaller (shorter and more 
slender) ; fronto-nasal region even less dished ; palate somewhat 
broader between molars and deeply excavated between last molars 
(slightly depressed in horriceus) ; interpterygoid canal shorter; zygo- 
mata more widely outstanding posteriorly and bowed (in horriceus 
more angular and more outstanding anteriorly) ; inferior border of 
jaw (symphysis to subangular tubercle) much longer. Canines and 
molars smaller. 

Old female (from Navajo Mountains) compared with old female 
horriceus (from mountains north of Silver City, N. Mex.) : Skull as 
a whole and rostrum shorter; occiput doubtless more truncate (that 
of horriceus sawed off) frontal shield shorter and more dished; 
postorbitals longer, more slender, and directed anteriorly as well as 
outward; zygomata much more broadly spreading, more swollen at 
anterior base, and distinctly bowed instead of angular; palate 
broader, flat instead of concave; lower jaw thicker and heavier. 

Skull measurements.— Old male (type) : Basal length. 308 ; occi- 
pito-nasal length, 301; palatal length, 171; zygomatic breadth, 218; 
interorbital breadth, 71. 



1918.] PLANICEPS GROUP. 37 

URSUS TEXENSIS NAVAHO Merriam. 

Navaho Grizzly. 

Ursus navaho Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 191-192, 
August 13, 1914. 

Type locality. — Navajo country near Fort Defiance, Arizona (Moll- 
hausen) ; type probably killed in 1856 in Chuska Mountains, on boun- 
dary between northeastern Arizona and northwestern New Mexico. 

Type specimen. — No. 3500, $ old, U. S. National Museum. 

Range. — Probably restricted to the isolated Chuska Mountains 
(including the so-called Lukachukki and Tunitcha elevations, and 
perhaps also the neighboring Carriso Mountains on the north). 

Characters. — Size small; external characters unknown; skull 
short, broad and slightly dished. 

Cranial characters. — Old male (type; badly damaged, only the 
front part with zygomata and underjaw remaining) : Skull short; 
zygomata broadly spreading and outbowed, the anterior root much 
swollen, posterior root not expanded; frontal shield flat, short 
pointed; postorbital processes peglike and outstanding; temporal 
impressions finely beaded; rostrum short and small; nares rather 
small and nearly vertical; lachrymal duct opening on orbital rim, 
but more in than out; palate broad for so small a skull; postpalatal 
shelf broad ; coronoid blade high. Teeth small : M 1 very small ; M- 2 - 
short, with broadly rounded heel (tooth nearly as broad posteriorly 
as in middle and anteriorly). 

Cranial comparisons. — Old male (type) compared with old male 
texensis (type) : Size somewhat larger; frontal shield and rostrum 
broader; jugal longer posteriorly. Palate and postpalatal shelf 
much shorter (157 mm. contrasted with 173), and less deeply concave 
between posterior molars ; postorbital processes slightly larger ; ante- 
rior root of zygoma much more swollen (in texensis hardly swollen) ; 
nares much more truncate; underjaw straighter (less upcurved 
posteriorly) ; canines about same size; molar series slightly shorter; 
M. 2 - of same length but heel broadly rounded posteriorly instead of 
obliquely truncate, the sides of tooth nearly parallel; M 1 , M T , and 
PM A smaller. 

Skull measurements. — Old male (type) : Palatal length, 157.5 ; 
interorbital breadth, 81. Skull too badly broken to furnish other 
measurements. 

URSUS PLANICEPS sp. nov. 
Flat-Headed Grizzly. 

Type No. 13289, $ ad. (rather old), U. S. National Museum, from 
Colorado (exact locality unknown). Collected by Dr. F. V. Hayden. 1 

1 Dr. Hayden worked in Colorado in 1869, mainly in the mountains and foothills of the 
east-central part of the State. Inasmuch as two other grizzlies, hairdi and macrodon, 
inhabit the higher mountains of Colorado it seems highly probable that the home of 
planiceps was in the foothills or along the western edge of the plains. 



38 NORTH AMERICAN FAUNA. [No. 41. 

Cranial characters. — Adult male (type) : Skull rather large, low, 
and flat ; shield broad, flat, and nearly horizontal ; postorbitals hori- 
zontally outstanding ; rostrum broad, somewhat depressed ; zygomata 
broadly outstanding and outbowed; palate, postpalatal shelf, and 
postpalatal notch broad; occipito-sphenoid long (95 mm.) Under- 
jaw rather long; ramus flat and rather thin; condyle high, vertical; 
subangular border short; angular process bellied on underside. 
Canines moderate; molars rather large but much smaller than in 
macrodon. 

Cranial comparisons. — Ursus planiceps requires comparison with 
its two neighbors, macrodon and texensis, of which its affinities are 
closest with macrodon. Old male (type) compared with old male 
macrodon: Size and vault of cranium about same; shield slightly 
broader and flatter; rostrum broader; zygomata very much more 
broadly outstanding and outbowed; palate, postpalatal shelf, and 
palatal notch much broader; posterior part of underjaw more up- 
curved, lifting condyle and coronoid ; upper canines about same size ; 
lower canines and upper and lower molars decidedly smaller. 

Old male (type) compared with old male texensis (type) : Size 
larger throughout; shield much broader, flatter, and more nearly 
horizontal ; f ronto-nasal region more elevated in plane of shield (not 
dished as in teaensis) ; rostrum much larger, broader, and more mas- 
sive; zygomata about the same but squamosal arm longer; palate 
flat (in texensis deeply concave between posterior molars) ; occipito- 
sphenoid longer; underjaw much more massive; ramus less tapering 
anteriorly; angular processes strongly bellied (in texensis not bel- 
lied) ; teeth larger. 

Skull measurements. — Old male (type) : Basal length, 319.5; occi- 
pito-nasal length, 305 ; palatal length, 173 ; zygomatic breadth, 215 ; 
interorbital breadth, 75. 

URSUS MACRODON sp. nov. 
Twin Lakes Grizzly. 

Type skull No. 15707 (skin No. 12678), $ old, U. S. National 
Museum. From Twin Lakes, Colorado, July 28, 1876. Collected by 
C. W. Derry. 

Characters. — Old male (type) : Size large; hump evident; claws 
(worn short) large, broad, mainly yellowish on top, horn color on 
sides. Skull rather large and flat ; last upper molar very large. 

Color. — Type specimen: Peculiar for a grizzly, resembling some 
of the Alaska brown bears; general body color rich brown, almost 
reddish brown, darker on hump and lightly washed with pale tipped 
hairs on upper part of back; head rather uniform brown. Adult 
female (No. 203178) from South Branch Williams Kiver, Colorado, 



1918.] PLANICEPS GROUP. 39 

killed September 13, 1895 : General color very dark, almost black. 
Muzzle reddish brown with a golden tinge on top, becoming very 
dark brown on cheeks, chin, and around eyes ; top of head dusky, over- 
laid by deep rich glossy hazel or between hazel and chestnut; top of 
neck and upperparts generally blackish, moderately washed with 
golden-tipped hairs; hump marked and nearly black; legs and feet 
nearly black ; the long hairs below ears and on sides of neck washed 
with golden. Claws long, slender, and smoothly polished; dark 
horn color, paler toward tips and on sides. Longest claw from upper 
base 61 mm. (tip worn off; greatest breadth of claws 8 mm.). 

Cranial characters. — Old male (type) : Skull long, low, flat, and 
rather narrow; shield narrow, flat, nearly horizontal, sloping grad- 
ually into rostrum without noticeable dishing ; zygomata not broadly 
outstanding; squamosal root long but not vertically expanded; palate 
long and narrow; postpalatal notch narrow; underjaw long; ramus 
straight, swollen on outer side below premolars ; coronoid blade high ; 
subangular border rather short ; angular process bellied ; canines and 
molars large ; M- 2 - 40 mm. in length ; heel long, slightly emarginate. 

Female from South Branch Williams River, Colorado (No. 203178, 
September 13, 1895, collected and presented by J. P. Bird) : Skull 
similar to that of male but much smaller, with correspondingly nar- 
rower shield and rostrum, and slightly more dished fronto-nasal re- 
gion; shield low, narrow, flat, and gently sloping, the point slender 
and remarkably short .for a female, ending anterior to fronto-parietal 
suture; sagittal crest proportionately long but low anteriorly; post- 
orbital processes small, slightly elevated, and directed slightly back- 
ward; orbital rims somewhat thickened; rostrum small; nasals an- 
teriorly horizontal, posteriorly rising slightly and passing into frontal 
shield in same plane ; zygomata moderately spreading, angular ; palate 
and postpalatal shelf short. Canines very small; molars large; M> 
38 mm., which is materially larger than in males of texensis and 
planiceps. 

Cranial comparisons. — Skull of old male (type) similar in general 
to old male texensis (type) but slightly larger, with longer, flatter, and 
more nearly horizontal frontal shield, and somewhat higher fronto- 
nasal region and rostrum; shield somewhat longer pointed; fronto- 
nasal region elevated in plane of shield (not dished as in texensis) ; 
postorbitals more broadly outstanding; rostrum broader and some- 
what longer ; opening of lachrymal duct within orbital rim (in texen- 
sis anterior to rim); zygomata less widely outstanding; squamosal 
root longer; postpalatal shelf narrower; palate flatter; underjaw more 
massive; ramus less tapering anteriorly; inferior border of angular 
process strongly bellied or keeled (may be individual) ; canines and 
molars very much larger throughout (especially M>). 



40 NORTH AMERICAN FAUNA. [No. 41. 

Remarks. — In size and general appearance macrodon resembles 
planiceps, but the frontal shield is more nearly horizontal ; postorbital 
processes less widely outstanding; zygomata much less broadly out- 
standing and not howed; squamosal arm of zygoma longer; canines 
and molars, particularly M- 2 - , decidedly larger. It may prove to inter- 
grade with planiceps. 

Skull measurements. — Old male (type) : Basal length, 312 ; occipito- 
nasal length, 302 ; palatal length, 171 ; zygomatic breadth, 202 ; inter- 
orbital breadth, 72. 

URSUS MIRUS sp. nov. 

Yellowstone Pabk Grizzly. 

/ 

Type No. 206595, $ ad. (rather old), U. S. National Museum, Bio- 
logical Survey collection. From Slough Creek, Yellowstone Na- 
tional Park, March 27, 1915. Collected by Henry Anderson. 

Cranial characters. — Old male (type) : Size medium; skull long, 
rather narrow and low arched ; shield rather narrow and flat, nearly 
horizontal postorbitally, anteriorly sloping gently into rostrum; 
postorbitals long, slender, and horizontally outstanding; rostrum 
high, subterete above ; nares high ; zygomata widely outbowed ; squa- 
mosal root arched and broadly expanded; palate and postpalatal shelf 
narrow ; mastoids vertical ; underjaw moderate, swollen on lower part 
of outer side under diastema; subangular border rather long and 
sloping upward posteriorly. Teeth of medium size. 

Cranial comparisons. — Old male (type) compared with old male 
tahltanicus (type) : Basal length and vault of cranium essentially 
same ; occipito-nasal length greater ; shield and rostrum flatter ; zygo- 
mata more outbowed; squamosal arm of zygoma longer and more 
broadly expanded; underjaw slightly longer; coronoid blade nar- 
rower in middle part ; inferior border of ramus of same length ; sub- 
angular border longer. 

Old male (type) compared with old male planiceps (type) : Size 
essentially same except that the rostrum is decidedly narrower and 
more elevated anteriorly; point of shield slightly more elevated; 
shield more sloping; postorbitals longer; rostrum narrower, higher, 
and subterete instead of flattened above ; palate and postpalatal shelf 
narrower ; zygomata even more strongly outbowed ; occipito-sphenoid 
shorter; subangular border of lower jaw longer; angular processes not 
bellied; teeth about same size. 

Old male (type) compared with old male texensis (type) : Similar 
in general characters but somewhat larger; vault of cranium slightly 
higher ; rostrum much higher anteriorly and more nearly horizontal ; 
nares much higher ; postorbitals more slender and much more widely 
outstanding (121 contrasted with 104) ; posterior frontal region less 



1918.] PLANICEPS GEOUP. 41 

elevated ; zygomata more outbowed (less triangular) ; squamosal root 
of zygoma much more broadly expanded ; postpalatal shelf narrower; 
underjaw larger and longer; ramus less tapering anteriorly; sub- 
angular border somewhat longer. 

Old male (type) compared with old male shoshone (type), with 
which it agrees in basal length: Vault of cranium lower; shield 
lower posteriorly and much less strongly sloping; rostrum higher 
anteriorly and more nearly horizontal; nares higher; sinus case not 
definitely keeling into sagittal crest; zygomata much more broadly 
outstanding and outbowed; squamosal root much more broadly ex- 
panded ; postpalatal length less ; canines * and molars closely similar. 

Compared with old male idaJwensis (type) : Skull, palate, and 
rostrum longer ; basicranium and vault less arched ; squamosal arm of 
zygomata longer and more broadly expanded. 

Remarks. — The cranial characters indicate that mints of the 
Yellowstone Park region is rather closely related to tahltanicus of 
the Stikine region, and also, though apparently less closely, to 
planiceps of Colorado and southern Wyoming. It is so much smaller 
and has teeth so much smaller than horribilis and imperator that 
comparison in detail is unnecessary; and compared with absarokus 
the skull is so much lower, flatter, and narrower and the teeth so 
much smaller that the two can not be confused. 

Skull measurements. — Old male (type): Basal length, 315; oc- 
cipito-nasal length, 307 ; palatal length, 174 ; zygomatic breadth, 220 ; 
interorbital breadth, 77. 

URSUS ELTONCLARKI ■ Merbiam. 

Sitka Grizzly. 

Ursus eltonclarJci Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 175-176, 
August 13, 1914. 

Type locality. — Near Freshwater Bay, Chichagof Island, the more 
northern of the Sitka Islands, Alaska. 

Type specimen. — No. 179066, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected May 19, 1912, by Elton Clark 
and by him presented to the Biological Survey. 

Range. — The Sitka Islands, Baranof and Chichagof. 

Characters. — A grizzly of medium or rather small size; skull 
small, long, narrow, and rather low, with flat frontal shield. Claws 
of true grizzly type, smoothly polished, strongly curved and rather 
short: longest claw (in type specimen) from upper base 70 mm.; 
dark bluish or plumbeous horn color streaked with whitish or 
yellowish. 



1 In the type specimen of shoshone the canines are absent, but they are present in a 
young male, No. 113410, from Marvine, Colorado, affording the desired comparisons. 
* Named in honor of Elton Clark, of Boston, who killed and presented the type specimen. 



42 NORTH AMERICAN FAUNA. [No. 41. 

Color. — Color of type very dark and rich. Nose pale brown, dark- 
ening just in front of eyes; face, head, and throat rich dark choco- 
late brown, with golden-brown wash in front of ears ; ears and patch 
under each ear dusky; occiput and neck grizzled golden-brown; 
back pale, overlaid by buffy tips ; legs and feet varying from blackish 
brown to brownish black. 

Cranial characters. — Adidt male (type) : Skull elongate, narrow ; 
zygomata moderately spreading, outbowed, rounded posteriorly, 
squamosal part not vertically expanded ; frontal shield in same plane 
with rostrum, narrow, low, flat, or slightly concave, acutely rather 
short pointed posteriorly, the point entering sagittal crest about one- 
third the distance from fronto-parietal suture to postorbital 
process; sagittal crest moderate, reaching more than halfway from 
occiput to postorbital processes; postorbital processes rather thick, 
outstanding; fronto-nasal region elevated (not dished) ; rostrum 
high and sloping gently upward in plane of frontals; lachrymal 
opening within orbit; palate long and narrow; postpalatal shelf 
long; postpalatal notch long and narrow; occipito-sphenoid short, 
about 80 mm. ; basisphenoid deeply concave, without trace of median 
ridge. Under jaw long. Teeth moderate; canines rather long; PM ¥ 
with moderately sloping heel slightly upturned at tip, sulcus very 
shallow; M T rather short, with cusplet on inner side of saddle 
posteriorly; My with anterior moiety decidedly longer than pos- 
terior and twin cusps of entoconid small and not deeply notched; 
PM A large and rather broad ; M 1 broad and rather short ; M- 2 - broad 
in anterior half, then narrowing strongly, the heel obliquely truncate 
on outer side ; cusps rather weak. 

Adult female: Similar, but, much smaller, distinctly dished, point 
of shield lyrate, zygomata more angular. 

Cranial comparisons. — Adult male (type) compared with adult 
orgilos (type) : Size essentially same; vault of cranium higher; ros- 
trum and fronto-nasal region longer and more elevated; postorbital 
processes heavier and shorter; occipito-sphenoid much shorter (80 
mm. contrasted with 90) ; mandible more massive. Canines larger 
and longer; My shorter; M 1 and MA shorter and broader (M* 
broader in middle.) 

Remarks. — Ursus eltonclarki falls in the tahltanicus group. Oddly 
enough, it resembles mirus, a geographically remote member of the 
group from the Yellowstone Park country, much more closely than it 
does tahltanicus. It agrees with mirus essentially in narrowness of 
skull as a whole, elevation and narrowness of rostrum and narrow- 
ness of palate, and even exceeds mirus in narrowness of shield and 
postpalatal notch. Even the under jaw agrees surprisingly with that 
of mirus, and the molar series are of approximately the same length, 
but the lower canines are materially larger. 



1918.J 



PLANICEPS GROUP. 43 



Skull measurements. — Adult male (type) : Basal length, 316; 
occipito-nasal length, 322; palatal length, 173; zygomatic breadth, 
215; interorbital breadth, 69. 

URSUS TAHLTANICUS Meeriam. 

Tahltan Grizzly. 

Ursus tahltamcus Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 181-182, 
August 13, 1914. 

Type locality.— Klappan Creek (=Third South Fork Stikine 
River), British Columbia. 

Type specimen. — No. 179928, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected in September, 1906, and pre- 
sented by G. Frederick Norton. 

Range. — Middle and upper Stikine-Skeena region, limits uncer- 
tain. 

Characters. — Size medium; skull of male slightly smaller than 
male stikeenensis and canadensis; skull of female about same size as 
female stikeenensis, decidedly larger than female canadensis. Color 
of type very dark. 

Color. — Type specimen: Black (head absent, but entire body and 
legs almost coal black, lightly grizzled on shoulders and anterior 
part of back by tips of golden brownish). 

Cranial characters. — Adult and old males: Rostrum rather narrow 
(in old age compressed in front of orbits) ; vault of cranium low, 
flattish; frontal shield of moderate breadth, flat or depressed (some- 
times sulcate medially), rather short pointed, sloping gradually to 
plane of muzzle, and only slightly dished in fronto-nasal region; 
postorbital processes peglike, horizontally far outstanding; sagittal 
crest low, reaching forward a little more than halfway from inion 
to postorbitals ; zygomata broadly spreading and bowed; palate of 
medium breadth; postpalatal shelf broad and flat; underjaw short — 
much shorter than in stikeenensis, shoshone, and canadensis — its 
ramus bellied and upcurved posteriorly, not flattened. Teeth rather 
small ; canines and molars much smaller than in stikeenensis. 

Female: Skull relatively large (about same length as female 
stikeenensis but much narrower) ; much larger than female sho- 
shone and canadensis; vault of cranium moderately elevated, with 
tendency to a fronto- parietal hump; facial part of skull large and 
long; frontal shield long, rather narrow, lyre shaped posteriorly, 
meeting sagittal crest some distance posterior to fronto-parietal su- 
ture; fronto-nasal region well dished; palate and postpalatal shelf 
long; zygomata moderately spreading and conspicuously outbowed; 
underjaw long. Young females are best told by the teeth, especially 
the lower canines. In tahltanicus the lower canines are shorter and 



44 NORTH AMERICAN FAUNA. [No. 41. 

more slender than those of stikeenensis. In the latter species they 
are longer and thicker, especially thick in middle. 

Cranial comparisons. — Old male (type) compared with adult male 
orgilos (type) : Size essentially same (basal length slightly less) ; 
top of skull higher; frontal shield broader and less flat; postorbital 
processes less slender ; palate and postpalatal shelf shorter ; zygomata 
less bowed but more widely outstanding; underjaw shorter, much 
more massive; inferior border of ramus shorter and more bellied 
posteriorly; coronoid blade more recurved. 

Adult male compared with adult male stikeenensis (both inhabit- 
ing the same region) : Basal length, zygomatic breadth, and frontal 
breadth essentially the same, but/frontal region much lower, frontal 
shield not rising abruptly at orbits, but sloping gently in plane of 
rostrum; rostrum narrower, higher, and less nearly horizontal (ap- 
pearing longer) ; postorbitals much smaller and more horizontally 
outstanding; palate shorter; underjaw shorter. Dentition lighter. 

Skull measurements. — Old male (type) : Basal length, 305 ; x occi- 
pito-nasal length, 304 ; palatal length, 163 ; zygomatic breadth, 220 ; 
interorbital breadth, 84. 

URSUS INSULARIS Merriam. 
Island Gbizzly. 

Ursus eltonclarhi insularis Merriam, Proc. Biol. Soc. Washington, XXIX, p. 
141, September 6, 1916. 

Type locality. — Admiralty Island, Alaska. 

Type specimen. — No. 205186, $ old, U. S. National Museum, Bio- 
logical Survey collection, 1914. Purchased from W. H. Case, of 
Juneau. 

Characters. — An island grizzly, apparently related to tahltanicus 
and orgilos of the mainland, and to eltonclarki of Baranof and Chich- 
agof Islands; in some respects a miniature of hoots. PM^ indis- 
tinctly of grizzly type. External characters unknown. 

Cranial characters. — Adult male (type) : Size medium; frontal 
shield broad and flattish, with exceptionally large, broad, and flat 
postorbitals; zygomata broadly outstanding; ramus of underjaw 
strongly upturned posteriorly, elevating coronoid and condyle. Teeth 
of medium size. 

Cranial comparisons. — Adult male (type) compared with old male 
tahltanicus (type) : Basal length, zygomatic breadth, breadth and 
form of frontal shield, breadth across postorbitals, truncation of 
nasals, and length of jaw essentially same, but postorbitals very 
much larger, especially broader and more decurved; fronto-nasal 

Restored. 



1918.] PLANICEPS GROUP. 45 

region more dished; rostrum slightly longer; palate, postpalatal 
shelf, and postpalatal notch narrower; underjaw more massive and 
more upturned posteriorly; ramus longer; coronoid much higher; 
subangular border shorter. Molar series of approximately same 
length but more massive ; M 1 longer, M 2 - shorter. 

Adult male (type) compared with adult male eltonclarki (type) : 
Similar in general, agreeing essentially in basilar length, zygomatic 
breadth, length and narrowness of palate and postpalatal shelf, and 
narrowness of postpalatal notch, but differing widely in other charac- 
ters. Vault of cranium less highly arched ; frontal shield and post- 
orbitals very much broader and more massive (shield interorbitally 
82 mm. contrasted with 69), less flat, faintly sulcate medially; post- 
orbital processes very much larger, broader, more widely outstanding 
(from tip to tip 120 mm. contrasted with 101) and more decurved; 
rostrum lower, broader, and shorter; nasals shorter (89 contrasted 
with 105 — probably not constant). Underjaw more massive; inferior 
border of ramus longer, more swollen and more upcurved posteriorly ; 
outer side of ramus not depressed or excavated below anterior base 
of coronoid; coronoid blade narrower and higher. Canines (both 
upper and lower) somewhat shorter; molars, especially M- 1 , My, and 
Mj decidedly larger. 

Compared with orgilos and tahltanicus of the mainland: Easily 
distinguished by great size of postorbitals and upturning of posterior 
part of ramus. 

Compared with kwakiutl: Quickly told by general shortness of 
skull, including palate and underjaw, and by large size and breadth 
of postorbitals. 

SkuU measurements. — Old male (type) : Basal length, 311; occip- 
ito-nasal length, 310; palatal length, 171; zygomatic breadth, 216; 
interorbital breadth, 82. 

URSUS ORGILOS Merriam. 

Glacier Bay Grizzly. 

Ursus orgilos Merriam, Proc. Biol. Soc. Washington, XXVII, p. 176. August 13, 
1914. 

Type locality. — Bartlett Bay, east side of Glacier Bay, Southeast- 
ern Alaska. 

Type specimen. — No. 180280, probably $ , rather old, U. S. Na- 
tional Museum, Biological Survey collection. Collected August 22, 
1912, by A. Hasselborg. 

Cranial characters. — Size medium ; skull long, rather narrow, low, 
flat on top, slightly dished. Frontal shield rather narrow, flat, concave 
between orbits, acute pointed posteriorly ; postorbital processes long, 
slender, outstanding, slightly decurved and recurved (posteriorly) ; 



46 NORTH AMERICAN FAUNA. [No. 41. 

rostrum normal or rather small ; nasals nearly horizontal, slightly up- 
lifted and decurved anteriorly; braincase long and low; squamosal 
shelves long ; zygomata moderately spreading and strongly outbowed, 
the broadest part more anterior than usual; sagittal crest straight 
and nearly horizontal, reaching only to fronto-parietal suture; palate 
long and rather narrow ; postpalatal shelf long, flat, and rather broad 
for size of skull ; jugal broad anteriorly, rising well above lachrymal 
duct; lachrymal duct opening within orbit; occipito-sphenoid long 
(90 mm.) ; underjaw long; coronoid blade broad at base, moderately 
high, the apex moderately recurved. Teeth of medium or rather 
small size. 

Cranial comparisons. — Ursus orgilos needs to be distinguished 
from its neighbors, orgiloides, iahltanicus, and pallasi. Compared 
with orgiloides, with which it agrees essentially in size of skull and 
length of braincase : Shield narrower, gently sloping instead of rising 
strongly from rostrum; rostrum longer and narrower; palate nar- 
rower. Compared with tahltanicus: Skull and shield narrower; 
postorbitals more slender and delicate; zygomata much less out- 
standing; underjaw more slender and delicate. Compared with pal- 
lasi: Skull as a whole, braincase, and posterior part of frontal shield 
much longer; shield less elevated above rostrum; sagittal crest 
shorter. 

Skidl measurements — Probably old male (type) : Basal length, 
316; occipito-nasal length, 304; palatal length, 177; zygomatic 
breadth, 207 ; interorbital breadth, 75. 

URSUS ORGILOIDES sp. nov. 
Alsek Gbizzly. 

Type No. 223275, probably $ , U. S. National Museum, Biological 
Survey collection. From Italio River, Alaska. Collected November, 
1916. Purchased from E. M. Axelson, of Yakutat. 

Range. — Coast strip southeast of Yakutat Bay. Specimens have 
been received from near Yakutat village and from Ankow and Ank- 
lin Rivers and mouths of Alsek and Italio Rivers. 

Cranial characters. — Skull of medium size, long, low, and smoothly 
rounded, rather narrow, with long braincase, long-pointed shield, 
rather short rostrum, and large broadly rounded lambdoid crest; 
shield medium, rising strongly from horizontal rostrum, sulcate inter- 
orbitally and swollen between sulcus and orbits; point ending at or 
near fronto-parietal suture; postorbital processes moderate, somewhat 
decurved; sagittal crest rather short; occipital overhang and inion 
well developed; rostrum short, broad for size of skull; nares trun- 
cate; zygomata not widely outstanding and not bowed: palate long, 



1918.] PLANICEPS GROUP. 47 

of moderate breadth; mastoids appressed; meatus tube large and 
free; underjaw long; coronoid blade moderate; apex only slightly 
recurved. Teeth rather small. 

Cranial comparisons. — Adult male (type) compared with the type 
of orgilos: Size, length of braincase, and length of postorbital part 
of frontal shield essentially same; shield broader, rising strongly 
from rostrum instead of sloping gradually into rostrum, sulcate 
anteriorly and swollen between sulcus and orbits; rostrum shorter 
and broader; lachrymal duct opening on orbital rim instead of 
within rim ; zygomatic arches more subtriangular (less bowed) ; 
palate broader; meatus tube much larger; lambdoid crest more 
highly developed and more broadly rounded. Teeth about same 
size; PM ¥ distinctly of grizzly type (in orgilos subcorneal). 

Compared with the type of pallasi, which it strongly resembles: 
Length about an inch greater (mainly in posterior part of skull) ; 
braincase and point of shield much longer; sagittal crest much 
shorter; occipito-sphenoid and palate longer; mastoids appressed 
instead of divergent; underjaw much longer, but inferior border 
of ramus of essentialty same length; subangular border slightly 
longer. 

Remarks. — Unfortunately there is possible doubt as to the sex of 
the type specimens of orgilos and orgiloides, though both are believed 
to be males. Both skulls have the appearance of males, except that 
in orgiloides the point of shield is longer than usual in males and 
the canines are small for males of corresponding size. 

Skull measurements. — Adult male (type): Basal length, 316.5; 
occipito-nasal length, 300; palatal length, 169; zygomatic breadth, 
218 ; interorbital breadth, 81.5. 

URSUS PALLASI Mebriam. 

Pallas Gbizzly. 

Ursus paZlasi Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 149-150, Sep- 
tember 6, 1916. 

Type locality. — Donjek River, southwestern Yukon. 

Type specimen. — No. 205160, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected August, 1913, by T. A. Dixon. 

Range. — Southwest corner of Yukon Territory, east of the St. 
Elias Range (Kluane Lake, Donjek River, St. Clair River) and 
adjacent eastern border of Alaska; easterly to McConnell River 
and Teslin Lake and south into northern British Columbia. 

Cranial characters. — Old male (type) : Size small, one of the 
smallest of the grizzlies; skull moderately elevated, flattish on top, 
with relatively broad frontal shield rising strongly at orbits. Frontal 
64854°— 18 4 



48 NORTH AMERICAN FAUNA. [No. 41. 

shield flattish, exceedingly short pointed posteriorly, faintly de- 
pressed medially between orbits, slightly swollen on sides of median 
depression, strongly sloping to rostrum; postorbital processes small, 
peglike, horizontally outstanding; fronto-nasal region strongly 
dished; rostrum short, somewhat depressed and pugged; nasals ris- 
ing anteriorly; nares small and subtruncate; sagittal crest long, 
reaching to halfway between fronto-parietal suture and plane of post- 
orbitals; zygomatic arches moderately outstanding, narrow, and 
slender, not expanded vertically; palate and postpalatal shelf short 
and broad; mastoids long and spreading. Underjaw long for size of 
skull; coronoid narrow above, the apex not reaching plane of con- 
dyle ; teeth rather large for size of skull. 

Old female (No. 205162) from St. Clair River, Yukon; collected 
September 6, 1914, by A. Hoyt: Size very small; frontal shield 
remarkably broad for so small a skull, convex and medially sulcate 
interorbitally ; fronto-nasal region strongly dished; postorbitals 
small, outstanding; rostrum short and depressed; palate and post- 
palatal shelf short and broad ; zygomata rather strongly outstanding, 
subtriangular. Underjaw very small and light; coronoid moderate, 
apex strongly recurved; teeth very small, nearly as small as in 
nelsoni. 

Cranial comparisons. — Ursus palTasi, owing to the number of 
species occurring in or adjacent to its range, should be known from 
tahltanicus, orgilos, orgiloides, pulchellus, and kluane. It may be 
related to tahltanicus but is easily distinguished by its low broad 
rostrum, strongly dished fronto-nasal region, very short shield, less 
elevated midfrontal region, long and rather high sagittal crest, and 
larger teeth; while tahltanicus has a higher and narrower rostrum, 
sloping fronto-nasal region, much longer shield, higher midfrontal 
region, shorter and lower sagittal crest, and smaller teeth. From 
orgilos and orgiloid.es it may be told by the shortness of the skull as 
a whole, shortness of braincase and point of frontal shield, and 
greater length of sagittal crest. From pulchellus it differs in lower 
vault of cranium, much broader shield and rostrum, shorter rostrum, 
broader palate and postpalatal shelf, longer underjaw, and smaller 
molars. From kluane it differs strikingly in much smaller size, 
much lower arch of cranium, conspicuously shorter braincase and 
sagittal crest; very much smaller, narrower, and less strongly de- 
curved postorbitals ; very much less elevated and more dished fronto- 
nasal region; much lower rostrum; much shorter palate and much 
shorter underjaw. 

Adult female compared with adult female kluane: Size of skull 
essentially same or slightly smaller; canines smaller; molars very 
much smaller. 



1918.] PLAN7CEPS GROUP. 49 

Skull measurements. — Old male (type) : Basal length, 302.5; occi- 
pito-nasal length, 279; palatal length, 159; zygomatic breadth, 209; 
interorbital breadth, 72.5. 

URSUS RUNGIUSI RUNGIUSI * sp. nov. 
Rungius Geizzlt. 

Type No. 179893, $ young-adult, U. S. National Museum, Biolog- 
ical Survey collection. Collected September, 1910, in Rocky Moun- 
tains on headwaters of Athabaska River, Alberta, by Carl Rungius, 
and by him presented to the Biological Survey. 

Characters. — Young -adult male (type) : Size small; skull low and 
flat, with low depressed braincase, very broad depressed sinus case, 
medium or narrow and exceptionally short frontal shield, long sagittal 
crest, and long, peglike outstanding'postorbital processes. Fully adult 
males differ somewhat. An old male from Fortress Lake, head of 
Athabaska River (No. 40091, Amer. Mus. Nat. Hist.), killed in May, 
1916, by Malcolm S. Mackay; another old male (No. 1919, Ottawa 
Museum) from Kootenay Pass, Alberta; and a fully adult male (No. 
209899) from Indian Point Creek, near Barkerville, B. C, present 
the following characters: Skull small, low, and nearly flat; shield 
low, of medium breadth, broadly flat-concave between orbits, sloping 
gradually into rostrum, very short pointed posteriorly; postorbitals 
peglike, outstanding, and elevated; orbital rims slightly thickened 
and everted; rostrum small, narrow, and rather high for so small 
a skull; zygomata widely outbowed; palate and postpalatal shelf 
narrow; base of cranium rather narrow; mastoids appressed (not 
outstanding); underjaw rather light; apex of coronoid recurved. 
Dentition moderate. PM ¥ distinctly of grizzly type. Canines and 
upper molars rather large for so small a skull. 

Cranial comparisons. — Adult male (No. 209899), from Indian 
Point Creek, B. C, compared with adult male macfarlani (type) : 
Size slightly less; skull as a whole much lighter; rostrum much 
smaller and less elevated; fronto-nasal region slightly dished; 
zygomata much less widely outbowed; palate and postpalatal shelf 
much narrower; sagittal crest shorter; base of skull narrower; 
underjaw much less massive; apex of coronoid more recurved. 
Dentition lighter (except M- 2 -, which is of about same size in both) ; 
M T less swollen; PM ¥ distinctly of grizzly type (not conical as in 
macfarlani) . 

Adult male (No. 209899, from Indian Point Creek, B. C.) compared 
with adult male ophrus (type) : Size smaller; vault of cranium very 

1 Named for the artist, Carl Rungius, of New York, who collected and presented the 
type specimen. 



50 NORTH AMERICAN FAUNA. [No. 41. 

much lower and flatter; frontal shield broadly concave, not deeply 
sulcate medially ; postorbital processes more slender and horizontal ; 
orbital rims slightly or not swollen ; f ronto-nasal region not notably 
dished ; rostrum lower ; nares smaller ; sagittal crest less highly 
developed and straighter (less convex) ; zygomata very much less 
outbowed and only slightly arched (in ophrus very highly arched) 
mastoids much shorter; under jaw much shorter; coronoid about the 
same size ; upper canines and upper and lower molars about the same 
size; lower canines smaller. 

Adult and old males compared with old male hylodrormis (No. 
205170), from Selkirk Mountains, B. C: Skull shorter; shield flat- 
concave, rising at orbits and postorbitals (in hylodromus decurved 
laterally) ; posterior part of shield very much shorter; vault of 
cranium much lower; braincase more depressed; rostrum smaller, 
narrower, more nearly horizontal; zygomata more outbowed (less 
triangular) ; sagittal crest longer; postpalatal shelf narrower. 

Skull measurements. — Young-adult male (type) : Basal length, 
293; occipito-nasal length, 282; palatal length, 162; zygomatic 
breadth, 190; interorbital breadth, 73. Adult male (No. 209899) 
from Indian Point Creek, B. C. : Basal length, 294 ; x occipito-nasal 
length, 278.5; palatal length, 161; zygomatic breadth, 198; inter- 
orbital breadth, 75. Old male (No. 40091 Amer. Mus. Nat. Hist.) 
from head Athabaska River, B. C. : Basal length, 295 ; occipito-nasal 
length, 276; palatal length, 166; zygomatic breadth, 214; inter- 
orbital breadth, 75. 

URSUS RUNGIUSI SAGITTALIS subsp. nov. 
Crested Gbizzly. 

Type No. 210705, $ ad. (rather old), U. S. National Museum, 
Biological Survey collection. From Champagne Landing, south- 
western Yukon. Collected in the fall of 1915. Purchased from 
Mackay & Dippie. 

Cranial characters. — Adult male (type) : Size small; skull low and 
narrow; shield flat or flat-concave, narrow, short pointed, sloping 
gradually into rostrum; postorbitals slender, outstanding; rostrum 
slender ; nares large and oblique ; zygomata moderately outstanding, 
subtriangular ; sagittal crest long, high, and arcuate ; palate narrow ; 
underjaw long and slender; coronoid small and falcate; subangular 
border short. Dentition moderate. 

Cranial comparisons. — Adult male (type) compared with adult 
male rungiusi (No. 209899) : Skull similar in general but shield nar- 
rower and flatter; postorbitals more slender; orbital rims not thick- 
ened or everted; nares larger and more oblique; sagittal crest very 

1 Restored. 



1918.] PLANICEPS GROUP. 51 

much higher, rising well above point of shield; squamoso-j ugal 
suture much shorter; unclerjaw longer; coronoid smaller and more 
falcate; subangular border shorter. Canines slightly larger; M 2 
somewhat smaller. 

Adult male (type) compared with orgilos (type) : Skull about an 
inch shorter; braincase very much shorter: point of shield much 
shorter; sagittal crest longer, much higher, and arcuate instead of 
straight; rostrum much smaller; squamosal arm of zygoma much 
shorter; underjaw and ramus much shorter; subangular border nearly 
the same ; coronoid blade narrower. 

Skull measurements. — Adult male, old : Basal length, 295 ; * oc- 
cipito-nasal length, 281 ; palatal length, 160 ; zygomatic breadth, 200 ; 
interorbital breadth, 71.5. 

URSUS MACFARLANI * sp. nov. 
MacFabiane Bear. 

Type No. 6551, $ ad., U. S. National Museum. Collected on 
Anderson River, 50 miles below Fort Anderson, Mackenzie, May 
8, 1863, by R. MacFarlane. (Original No. 551.) 

Characters. — External characters unknown. Relationship ap- 
parently with rungiusi. 

Cranial characters. — Size medium; skull of adult male (type) 
massive, low, broad, flat ; tip of nose to point of shield in same plane 
without trace of dishing. Frontal shield low, flat-concave, rather 
broad, exceptionally short posteriorly, shallowly concave between 
orbits, not rising above plane of rostrum; postorbitals large, hori- 
zontally outstanding and slightly elevated; rostrum broad and high, 
large for size of skull; zygomata widely outbowed; sagittal crest 
long and low; palate and postpalatal shelf broad; underjaw massive, 
coronoid blade high, narrow, nearly vertical, obtusely rounded above ; 
subangular border exceptionally short. Dentition heavy: Canines 
and molars large for size of skull; PM T subcorneal; M T swollen; 
M- 2 - moderate, the heel slightly emarginate and rather broadly 
rounded posteriorly. In immature and young-adult males the frontal 
shield is less flat, the sides (between sulcus and orbits) strongly 
swollen (as shown by No. 7146, from Franklin Bay; and 2773, 
Ottawa Museum, from Stapylton Bay). 

Cranial comparisons. — Adult male (type) compared with adult 
male rungiusi (No. 209899) from Indian Point Creek near Barker- 
ville, B. C. : Skull slightly larger and much more massive, with higher 

1 Partly restored. 

2 Named in honor of Roderick MacFarlane, who collected the specimen and presented it 
to the Smithsonian Institution. 



52 NORTH AMERICAN FAUNA. [No. 41. 

and much broader rostrum, and much more widely outbowed zygo- 
mata; sagittal crest longer; palate, postpalatal shelf, and base of 
skull much broader; underjaw much more massive; coronoid blade 
more nearly vertical and less recurved at apex. Dentition heavier: 
Canines and molars, upper and lower, larger; M T more swollen; PM ? 
subconical (not at all of grizzly type). 

Skull measurements. — Adult male (type): Basal length, 303; 
occipito-nasal length, 283; palatal length, 164; zygomatic breadth, 
218 ; interorbital breadth, 79. 

URSUS CANADENSIS Meebiam. 1 
Canada Gkizzly. 

Vrsus shoshone canadensis Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 
184-185, August 13, 1914. 

Type locality. — Moose Pass, near Mount Robson, British Columbia. 

Type specimen. — No. 174511, $ ad., U. S. National Museum. Col- 
lected by N. Hollister and Charles D. Walcott, jr., July 23, 1911. 
(Original No. 3792, Hollister catalogue.) 

Range. — Eastern British Columbia; limits unknown (type from 
near Mount Robson ; and an adult female from Kootenay Lake ) . 

Characters. — Size medium; color brown, grizzled with buff; claws 
short for a grizzly, rather thick, moderately curved, pale yellowish 
on upper surface and tips, brownish horn color on sides. 

Color. — Muzzle very pale drab brown, changing to darker brown 
on head, face, and chin, darkest around ears; top of head, cheeks 
posteriorly, 2 ears, back, and thighs washed with buffy whitish from 
abundance of buffy-tipped hairs; foreleg and lower part of hind leg 
and feet very dark (almost blackish brown) ; long hairs of throat 
and axillary region pale yellowish, of rest of underparts dark 
brown. 

Cranial characters. — Adult male (type) : Skull of medium size, 
rather long, low, and narrow, flat in frontal region, with long, high 
sagittal crest rising above general level of top of cranium. Frontal 
shield small, flat, narrow, faintly depressed medially, short pointed 
posteriorly, gently sloping ; postorbitals slender, peglike, horizontally 
outstanding; rostrum relatively high, tapering anteriorly; sagittal 
crest remarkably long, arcuate, and high, reaching nearly to midway 
between fronto-parietal suture and plane of postorbitals; zygomata 
moderate, slightly outbowed, and only slightly expanded vertically; 
palate excavated between molar series; postpalatal shelf rather long 
and broad; notch medium or narrow. Underjaw rather massive; 

1 Tentatively included in planiceps group. (See Introduction, pp. 12—13.) 

2 The old whitish-tipped hairs of the old coat have fallen out on the cheeks and anterior 
part of head nearly to ears. 



1918.] ARIZONA GROUP. 53 

ramus moderately bellied posteriorly; coronoid blade broad and 
rather short, the apex cutting plane of condyle. Dentition rather 
heavy; canines thick and short; molars broad. 

Adult female (No. 209902 from Kootenay Lake, British Colum- 
bia) : Skull rather small, long, narrow, low, and slightly dished. 
Frontal shield narrow, flattish, slightly depressed medially between 
orbits, long pointed, the point reaching to midparietal region; post- 
orbitals moderate; rostrum long and slender, tapering; zygomata 
moderately spreading. Under jaw similar to that of male but much 
smaller; coronoid blade relatively narrower and higher. Dentition 
heavy; teeth similar to those of male and only slightly smaller; 
canines large and swollen; molars broad. 

Cranial comparisons. — Adult male (type) compared with adult 
male macrodon (type) : Size essentially same; shield more strongly 
sloping ; postorbitals more slender ; rostrum lower and more tapering ; 
sagittal crest higher ; inion much more strongly produced ; subangular 
border of ramus shorter ; angular process not bellied ; canines smaller ; 
M^ and M- decidedly smaller. 

Skull measurements. — Adult male (type) : Basal length, 313; oc- 
cipito-nasal length, 312 ; palatal length, 171 ; zygomatic breadth, 208 ; 
interorbital breadth, 74. 

Arizonae Group. 

URSUS ARIZONA Meebiam. 

Aeizona Geizzly. 

(PI. XVI.) 

Vrsus arizonw Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 135-136, 
September 6, 1916. 

Type locality. — Escudilla Mountains, Apache County, Arizona. 

Type specimen. — No. 177332, $ ad., U. S. National Museum, 
Biological Survey collection. Collected September 3, 1911, by C. H. 
Shinn. 

Cranial characters. — Size rather large; skull as a whole rather 
long and narrow, with broad rostrum; vault of cranium moderately 
elevated but not arched, highest about two-thirds distance from plane 
of postorbitals to fronto-parietal suture; frontal shield rather nar- 
row, nearly flat, gently sloping in plane of rostrum, the posterior 
point in type specimen reaching to about 25 mm. in front of parietals 
(in older specimens shorter) ; postorbitals broad and broadly 
rounded, nearly horizontal, but not widely projecting; fronto-nasal 
region and rostrum elevated and swollen, continuing plane of frontal 
shield without trace of dishing, tapering anteriorly; zygomata not 



54 NORTH AMERICAN FAUNA. [No. 41. 

widely outstanding, bowed, anterior roots swollen ; palate rather short 
and broad; postpalatal shelf broad; meatus tube long; coronoid 
blade rather broad above, its recurved apex cutting plane of con- 
dyles. Teeth rather small for size of skull; canines of good size; 
molars rather small for size of skull, especially last upper molar. 

Cranial comparisons. — Adult male (type) compared with old male 
apache (type) : Basilar length, occipito-nasal length, length of palate, 
interorbital breadth, and occipito-sphenoid length essentially the 
same; zygomata very much less outstanding and subtriangular in- 
stead of bowed ; frontal shield flatter, of essentially the same breadth 
interorbitally, but very much narrower across postorbital processes; 
postorbital processes much less broadly outstanding; orbital rims 
less swollen; fronto-nasal regu^/much more elevated and swollen; 
rostrum much larger, broader, more swollen, and tapering instead 
of depressed basally, narrow, and horizontal ; palate and postpalatal 
shelf much broader. Under jaw weaker; ramus less broad vertically; 
coronoid blade less high; molars slightly larger; heel of M- 2 - longer, 
more distinctly emarginate on outer side (less tapering). 

Skull measurements. — Adult male (type) : Basal length, 326; oc- 
cipito-nasal length, 323; palatal length, 175; zygomatic breadth, 208; 
interorbital breadth, 82. 

URSUS IDAHOENSIS sp. nov. 
Idaho Grizzly. 

Type No. 93, $ old, Merriam collection (=187888 U. S. National 
Museum), from North Fork Teton River, eastern Idaho, September 
23, 1874. Killed by the late Richard Leigh (better known as " Beaver 
Dick"). 

Characters. — Size rather large, about equaling arizonw (much 
smaller than horribilis and subspecies, and shorter than rogersi, 
bisonophagus, and perturbans) ; frontal shield convex both longi- 
tudinally and transversely; fronto-nasal region elevated and some- 
what compressed (much as in rogersi, but rostrum much shorter) ; 
postorbital processes rather weak and decurved as in rogersi and 
arizonce; sagittal crest rather short and not strongly developed; 
zygomata moderately outbowed ; coronoid blade rather high, but less 
high than in bisonophagus and rogersi; ramus flattish, broad verti- 
cally, but much less broad and massive then in rogersi; dentition 
rather heavy; M 1 large, its heel elongate, emarginate, but not nar- 
rowed posteriorly, slightly everted ; canines rather small, about as in 
rogersi and arizonce. 

An old female (No. 160153) from Wallowa Mountains, Oregon, is 
assumed to be typical of idahoensis. It has a long, low, slender, 
smoothly rounded skull with narrow zygomatic arches, narrow 
palate, rather broad postpalatal shelf, and very small teeth. 



1918.] ARIZONA GROUP. 55 

Skull measurements. — Old male (type) : Basal length, 317; oc- 
cipito-nasal length, 318; palatal length, 177; zygomatic breadth, 
206; interorbital breadth, 81. 

URSUS PULCHELLUS PULCHELLUS sp. NOV. 
Upper Yukon Gkizzly. 

Type No. 221599, $ ad., U. S. National Museum, Biological Sur- 
vey collection, from Ross River, Yukon Territory, Canada. Col- 
lected July 20, 1916, by Fred E. Enevoldsen. 

Characters. — Size small; frontal shield and rostrum narrow; 
vault of cranium well arched; base of cranium moderately arched; 
shield rising rather strongly from plane of rostrum, convex trans- 
versely but shallowly sulcate medially, short pointed posteriorly 
(point ending about halfway between plane of postorbitals and 
fronto-parietal suture); rostrum slender and high; fronto-nasal 
region dished; nasals nearly horizontal; sagittal crest only slightly 
developed, not high posteriorly, somewhat decurved; occiput rather 
low and shortly truncate; palate and postpalatal shelf rather nar- 
row; zygomata subtriangular ; squamosal root expanded vertically; 
occipito- sphenoid short (80 mm.) ; under jaw short, its inferior bor- 
der straight; coronoid rather low, broad basally, apex strongly re- 
curved; subangular border short. Teeth, particularly molars, large 
for so small a skull; last upper molar broadly quadrate anteriorly, 
the heel abruptly and strongly emarginate, narrowly rounded pos- 
teriorly ; M 1 large, broad, and massive. 

Cranial comparisons. — Ursus pulchellus requires comparison with 
its near relative ereunetes, and also with pallasi and kluane. Adult 
male (type) compared with adult male ereunetes (type) : Size 
slightly smaller; fronto-nasal region more dished; shield less flat; 
zygomata broad (in ereunetes slender); occipito-sphenoid shorter; 
M 1 larger and more massive. 

Compared with old male pallasi (type) : Basal length slightly less; 
occipito-nasal length same; zygomatic breadth less; cranium higher 
and more arched; frontal shield and rostrum conspicuously nar- 
rower; palate and postpalatal shelf narrower; rostrum longer, 
higher, and more slender; squamosal root of zygoma more broadly 
expanded vertically; mastoids less elongate; underjaw shorter and 
less massive. Upper molars larger and more massive. 

Compared with adult male kluane : Similar in general appearance 
but skull as a whole, braincase, palate, and underjaw very much 
shorter,' frontal shield very rmich narrower; inferior border of ramus 
more abruptly upcurved; subangular border more nearly horizontal 
and much more sharply defined. 



56 NORTH AMERICAN FAUNA. [No. 41. 

Remarks. — The type of pulchellus came from Ross River, a north- 
ern tributary of the Pelly. Skulls of males from the southwestern 
corner of Yukon (Donjek River and Champagne Landing) differ in 
greater occipito-nasal length, more highly arched cranium, more ele- 
vated rostrum, and less deeply emarginate heel of 1VP-. 

I refer to pulchellus an adult female, No. 204187, from McConnell 
River, Yukon, and a still older female, No. 215113, from Ross Moun- 
tains. 

Skull measurements. — Adult male (type) : Basal length, 292; oc- 
cipito-nasal length, 281 ; palatal length, 160 ; zygomatic breadth, 196 ; 
interorbital breadth, 66. 

URSUS PULCHELLUS / EREUNBTES subsp. nov. 

KOOTENAY GRIZZLT. 

Type No. 222323, $ ad., from Beaverfoot Range, Kootenay Dis- 
trict, British Columbia. Collected October 1, 1916, by George Hill, 
of Field, British Columbia. 

Characters. — Size rather small; occiput shortly truncate; vault of 
cranium and basicranial axis well arched; fronto-nasal region ele- 
vated, sloping gradually upward, not dished; rostrum small, rather 
narrow, rising gradually into shield; point of shield ending about 
two-thirds distance from plane of postorbitals to fronto-parietal 
suture; zygomata slender, rather broadly outbowed for size of skull. 
Last upper molar broad anteriorly, the anterior part of cingulum 
on inner side produced, the heel emarginate, narrowing posteriorly, 
subtriangular ; middle lower molar large and massive. 

Skull in general similar to that of pulchellus but slightly larger, 
with more elevated fronto-nasal region, flatter shield, much more 
slender zygomata, and much longer occipito-sphenoid ; teeth as in 
pulchellus except that MMs smaller, and M^ is less quadrangular 
anteriorly. 

Skull measurements. — Adult male (type) : Basal length, 297; 
occipito-nasal length, 278; palatal length, 165; zygomatic breadth, 
203 ; interorbital breadth, 72. 

URSUS ORIBASUS sp. NOV. 
Liasd River Grizzly. 

Type No. 223991, $ ad. (rather old), U. S. National Museum, 
Biological Survey collection. From Upper Liard River, Yukon, 
near British Columbia boundary. Killed by J. Thompson in the 
spring of 1916. (Purchased from William Drury, of Whitehorse.) 

Characters. — Adult male (type) : Size large; hump absent or in- 



1918.] ARIZONA GROUP. 57 

conspicuous; color dark; claws long (longest 90 mm. 1 ) and unusually 
straight; top convex in section, dark horn color, paler at tips and 
along upper surface; skull long, narrow, and arched, with elevated 
straight-sloping fronto-nasal region (much like that of ereunetes 
but much longer posteriorly). Frontal shield, rostrum, and molar 
teeth narrower than in any other member of the shoshone group. 

Color. — General ground color dark brown to dusky ; muzzle dull 
golden brown, becoming much darker between eyes; a dark ring 
around each eye; cheeks chestnut brown; top of head, nape, and 
shoulders strongly washed with yellowish buffy; back washed with 
soiled buffy; rump dark brownish dusky; legs and feet dusky 
blackish. 

Cranial characters. — Adult male (type) : Skull rather large, long, 
narrow, rather strongly ;: relied both above and below, with high 
straight-sloping (not dished) fronto-nasal region. Frontal shield 
narrow, flat, gently sloping, the point reaching two-thirds distance 
from postorbitals to parietals ; postorbitals rather small, horizontally 
outstanding; rostrum narrow and high, in same plane with frontal 
shield; braincase and sagittal crest long; inion and occipital over- 
hang marked; zygomata well outstanding, strongly subtriangular, 
squamosal base broadly expanded ; basicranium and palate arched ; 
palate and postpalatal shelf narrow ; occipito-sphenoid 92 mm. ; mas- 
toids large, divergent. Under jaw long; ramus long and flat; sub- 
angular border short and broad; coronoid blade narrow. Canines 
rather long ; molars narrow and rather small ; MA small, narrowly 
triangular, the heel small, thin, and pointed; PM^ imperfectly of 
grizzly type. 

Cranial comparisons. — Old male (type) compared with old male 
chelan and idahoensis (both types) : Length, height, and arching 
essentially same; shield narrower, flatter, and more nearly hori- 
zontal ; sinus case and rostrum narrower ; top of rostrum higher and 
more completely in fronto-nasal plane ; zygomatic arches shorter and 
more angular (less outbowed), the inclosed space (temporal fossa) 
much smaller. Under jaw shorter and weaker; subangular border 
shorter; coronoid blade narrower. 

Old male (type) compared with adult male ereunetes (type) : 
Similar in general appearance but length much greater; shield flat- 
ter; rostrum more elevated; braincase much longer; sagittal crest 
much longer and higher; inion more strongly produced; squamosal 
base of zygoma broadly expanded (in ereunetes not expanded). Ca- 
nines longer; molars smaller and very much narrower. 

1 Claw of second or index finger longest, but second, third, and fourth practically sub- 
equal ; claw of thumb very long. Claw measurements of right hand (those of left hand 
more worn at tips) from upper exposed base to tip : First, 82 mm. ; second, 90 ; third, 88 ; 
fourth, 89 ; fifth, 68. 



58 NORTH AMERICAN FAUNA. [No. 41. 

Remarks. — Ursus oribasus appears to be closely related to pulchel- 
lus and ereunetes, both of which have decidedly smaller skulls with 
larger teeth. It is related also to idahoensis and chelan, which are 
about the same size, but have much broader skulls and differ other- 
wise as already pointed out. 

Skull measurements. — Adult male (type) : Basal length, 310; oc- 
cipito-nasal length, 304 ; palatal length, 172 ; zygomatic breadth, 215 ; 
interorbital breadth, 75. 

URSUS CHELAN Mereiam. 

Chelan Grizzly. 

Ursus chelan Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 136-137, Sep- 
tember 6, 1916. / 

Type locality. — East slope Cascade Mountains, northern Chelan 
County, Washington. 

Type specimen. — No. 205185, $ old, U. S. National Museum, 
Biological Survey collection. Killed in Township 30 N, Range 16 
East, Willamette Meridian, Wenatchee National Forest. Collected 
September 1, 1913, by D. S. Rice. 

Range. — Cascade and Cassiar Mountains from northern Washing- 
ton to upper Stikine River and Dease Lake, British Columbia. 

Cranial characters. — (External characters unknown). Skull of 
medium or rather large size; facial axis strongly deflected from 
basicranial axis; vault of cranium well arched, highest over poste- 
rior frontal region; sagittal crest long, high, arcuate, rising ante- 
riorly above general level of top of cranium. Affinities apparently 
with hylodromus on the one hand, and with shoshone and pervagor on 
the other. Frontal shield narrow, flattened, short pointed poste- 
riorly, ending about midway between fronto-parietal suture and 
plane of postorbitals, slightly sulcate medially ; postorbital processes 
rather broad, flat, outstanding horizontally (not depressed or de- 
curved) ; fronto-nasal region including posterior two-thirds of nasals 
sloping strongly, forming part of long fronto- facial plane; rostrum 
small, short, somewhat depressed, sloping anteriorly to nares, grad- 
ually rising posteriorly into frontal plane; braincase long, arched, 
frontal part keeled into sagittal crest; palate arched antero-poste- 
riorly, slightly concave; postpalatal shelf broad; zygomata broadly 
spreading, rounded and outbowed posteriorly, vertically expanded 
and strongly arched ; mastoids of medium length, spreading ; under- 
jaw massive; ramus swollen on outer side over roots of M-j and 
M T , bellied under last molars; coronoid blade high, its anterior bor- 
der rather strongly recurved, the apex overarching high coronoid 
notch but barely reaching plane of front of condyle; teeth rather 
small for size of skull (so badly worn in type specimen that propor- 
tions of canines can not be determined). 



1918.] ARIZONA GROUP. 59 

Cranial comparisons. — Old male chelan differs from all its rela- 
tives in the degree of deflection of the facial part of the skull, and ex- 
ceeds all except Kylodromus in the arching of the palate. 

Old male (type) compared with pervagor (type) : Basal length 
slightly less; zygomatic breadth greater; frontal shield shorter 
pointed and flatter; postorbitals flatter and apparently broader; 
sagittal crest longer, higher anteriorly and more convex; rostrum 
shorter and more strongly sloping anteriorly; nares more truncate; 
zygomata more strongly outbowed, more arched, more expanded 
vertically, squamosal arm longer (squamoso-jugal suture much 
longer) ; palate shorter and more strongly arched ; under jaw 
shorter, its inferior ramus much shorter; coronoid blade slightly 
higher and more falcate. Some of these differences may be due to 
age, the skull of the type of chelan being very old, while the type of 
pervagor is only adult. However, it differs rather conspicuously 
from equally old male pervagor from Bridge River (No. 4, Provin- 
cial Museum, Victoria, B. C.) as will be seen from the following: 
Basal length 20 mm. less; occipito-nasal length slightly greater; 
zygomatic breadth less; facial part strongly deflected (in pervagor 
not deflected) ; palate arched (in pervagor not arched) ; frontal 
shield shallowly sulcate medially (in pervagor broadly concave) ; 
postorbitals broad, flat, horizontally outstanding (in pervagor long, 
peglike, uplifted, and arched) ; braincase and sagittal crest arched 
(in pervagor straight and nearly horizontal) ; occipital overhang 
much greater; zygomata less widely outbowed. 

Old male (type) compared with old male Kylodromus (No. 205170) 
from Selkirk Mountains, British Columbia : Size larger (basal 
length only slightly greater but occipito-sphenoid and occipito- 
nasal lengths much greater, and skull as a whole distinctly 
larger) ; vault of cranium decidedly more highly arched ; facial 
angle more strongly deflected from basicranial axis; zygomata 
much more widely spreading and outbowed and much more 
arched; frontal shield rising less abruptly from rostrum, more 
evenly sloping, rising higher posteriorly, and much shorter pointed ; 
braincase and sagittal crest much longer, the crest higher and con- 
vex or arcuate anteriorly; occipital overhang greater; palate more 
strongly arched (antero-posteriorly) ; mastoids longer and strongly 
spreading. Underjaw longer and more massive, more swollen on 
outer side below middle and posterior molars; its inferior border 
more bellied posteriorly ; coronoid blade very much higher. 

Old male (type) compared with old male washake (type) : Size 
slightly larger (basal length essentially the same, but upper part 
of skull much longer) ; vault of cranium more highly arched; frontal 
shield continuing to rise posteriorly (instead of flattened) and much 
shorter pointed ; rostrum decidedly broader ; postorbital processes 



60 NORTH AMERICAN FAUNA. [No. 41. 

not elevated; lachrymal duct within orbit (not cutting rim as seen 
from front); braincase compressed and keeled anteriorly (in 
washake depressed) ; sagittal crest much longer and convex instead 
of straight; squamosal arm of zygoma longer and more broadly ex- 
panded vertically; palate concave and arched antero-posteriorly, in- 
stead of flat; postpalatal shelf longer and less broadly flattened ; oc- 
cipito-sphenoid longer ; mastoids longer and more spreading. Molars 
smaller. 

Skull measurements. — Old male (type): Basal length, 314; oc- 
cipito-nasal length, 323 ; palatal length, 170 ; zygomatic breadth, 225 ; 
interorbital breadth, 86. 

URSUS SHOSHONE Merriam. 

Shoshone Grizzly. 

Ursus shoshone Merriam, Proc. Biol. Soc. Washington, XXVII, p. 184, August 
13, 1914. 

Type locality. — Estes Park, Rocky Mountains of northern Colo- 
rado. 

Type specimen. — No. 203185, $ old, U. S. National Museum, 
Biological Survey collection. 

Range. — Mountains of Colorado and Wyoming. 

Characters. — Size medium or rather large, but much smaller than 
horribilis and bairdi — skull about same size as absarokus, but nar- 
rower and widely different. External characters unknown. 

Cranial characters. — Adult nude (type) : Skull rather long and 
high, with flattish, short-pointed, long-sloping frontal shield con- 
tinuing plane of rostrum to highest point, about midway between 
postorbitals and fronto-parietal suture; zygomata moderately 
spreading, outbowed; anterior (frontal) part of braincase keeling 
into sagittal crest ; sagittal crest long and high ; lambdoid crest high ; 
postorbital processes peglike, outstanding, rather slender; nasal 
region slightly dished and sulcate in middle third (nasals dipping 
toward one another — may be individual) ; rostrum of moderate 
breadth, strongly ascending in plane of frontal shield ; palate slightly 
dished between posterior molars ; postpalatal shelf broad ; postpalatal 
notch long and narrow; lachrymal duct cutting orbital rim but 
mainly on inner side. Under jaw rather long; ramus broad vertically, 
flattish, highest posteriorly; coronoid blade high, rather strongly 
sloping, the apex cutting plane of condyle. Canines (absent in type 
but present in other males) slender; molars medium; M- 3 - large with 
long heel. 

Adult female (No. 203761, from Fort Fred Steele, Wyoming) : 
Skull long, low, and rather slender; frontal shield narrow, long, 
lyre pointed posteriorly, flat interorbitally ; postorbitals slender and 



1918.] ARIZONA GROUP. 61 

outstanding; rostrum slender; sagittal crest short, reaching only 
halfway from inion to fronto-parietal suture; lambdoid crest mod- 
erate; palate dished between posterior molars; postpalatal shelf 
broad ; notch moderate ; meatus tube short ; ramus flat and light ; apex 
of coronoid produced posteriorly, overhanging deep coronoid notch. 
Teeth rather small ; upper molars relatively large, M> with long heel, 
cut-turned posteriorly ; lower molars narrow. 

Cranial comparisons. — Adult male compared with adult male hor- 
riceus: Rostrum, nasals, and frontal shield more elevated and much 
more strongly ascending posteriorly (less flattened and less nearly 
horizontal) ; lambdoid much more strongly developed. 

Adult male compared with adult male absarokus: Frontal shield 
flat, rising gradually in long continuous slope to highest point, about 
25 mm. anterior to fronto-parietal suture (in absarokus arched, 
strongly convex, and rising suddenly to highest point, immediately 
behind orbits) ; postorbital processes more slender; fronto-nasal re- 
gion only slightly dished ; rostrum more slender ; braincase narrower ; 
breadth across squamosal shelves less. 

Remarks. — Skulls from the Wind River and Absaroka Mountains 
have the last upper molar smaller, the heel less strongly developed. 

Skull measurements. — Old male (type): Basal length, 320; occi- 
pito-nasal length, 317; palatal length, 166; zygomatic breadth, 208; 
interorbital breadth, 78. 

URSUS KENNERLYI 1 Mebriam. 

Sonoea Grizzly. 

Ursus kennerlyi Merriam, Proc. Biol. Soc. Washington, XXVII, p. 194, August 
13, 1914. 

Type locality. — Mountains of northeastern Sonora, near Los 
Nogales, Mexico. 

Type specimen.— Skull No. 2086, $ old ; skin No. 1047, U. S. Na- 
tional Museum. Collected in June, 1855, by Dr. C. B. Kennerly. 

Range. — Nothing is known of the range of kennerlyi except that 
the type specimen came from mountains near Nogales, Sonora. Its 
affinities with utahensis suggest that formerly it may have had a dis- 
connected distribution northward in the mountains of central 
Arizona. 

Characters. — Size rather small. Ursus kennerlyi is a strongly 
marked member of the arizonce-utahensis group, most nearly related 
to utahensis but very much smaller, although the teeth are about same 
size. The skull, though that of an old male, agrees in size (length) 
with that of female utahensis. 

1 Named In honor of Dr. C. B. Kennerly, who collected the type specimen. 



62 NORTH AMERICAN FAUNA. [No. 41. 

Color. T- The prevailing color is dull pale brownish yellow with 
amber tinge. The tips only are of this color, the basal and larger 
portion being of a dark chestnut-brown, passing into blackish, which 
extends nearly to yellowish tips, the blackish predominating along 
median line of back and posteriorly; legs blackish brown slightly 
tinged with chestnut (Baird). 

Cranial characters. — Size rather small; skull long, narrow, and 
high, but not much arched ; rostrum narrow and high, in same plane 
with shield; fronto-nasal region strongly elevated, making a con- 
vexity slightly above otherwise continuous plane of rostrum and 
frontal shield ; rostrum and fronto-nasal region subterete, constricted 
(but not strongly pinched in) in front of orbits ; nares much higher 
than broad; frontal shield flaf, rather short pointed posteriorly, 
passing into sagittal crest about one-third distance from fronto- 
parietal suture to postorbitals ; postorbital processes long, rather 
slender, outstanding, and slightly decurved; frontal part of brain- 
case elevated ; zygomata moderately spreading and outbowed ; palate 
long, somewhat concave ; squamosal shelves broad ; coronoid broadly 
falcate; ramus flat and broad vertically. Teeth of medium size 
(badly worn). 

Cranial comparisons. — Old male (type) compared with old male 
horriceus (type) : Size essentially the same; vault of cranium higher 
over posterior frontal region; frontal shield slightly convex (not 
sulcate or depressed between orbits or elsewhere), longer pointed 
posteriorly; fronto-nasal region markedly elevated instead of de- 
pressed; rostrum much higher posteriorly, rising in same plane with 
frontal shield and strongly compressed; postorbital processes more 
slender and less decurved; sagittal crest much shorter; angle of jaw 
shorter ; inferior border of ramus decidedly longer ; apex of coronoid 
more slender. Teeth so badly worn that detailed characters are lost ; 
in size, however, they agree essentially with those of horriceus except 
that the large lower premolar is decidedly smaller. 

Skull measurements. — Old male (type): Basal length, 314; * 
occipito-nasal length, 306; palatal length, 165; zygomatic breadth, 
205 ; interorbital breadth, 75. 

URSUS UTAHENSIS Merriam. 

Utah Grizzly. 

Ursus utahensis Merriam, Proc. Biol. Soc. Washington, XXVIT, pp. 193-194, 
August 13, 1914. 

Type locality.— North Fork Salina Creek, 10 or 12 miles southeast 
of Mayfield, Utah. 

1 Restored. 



1918.] ARIZONA GROUP. 63 

Type specimen— No. 180193, $ old, U. S. National Museum, 
Biological Survey collection. Collected May 22, 1911, by Mart 
Martenson. 

Range. — Southern Wasatch and Pine Valley Mountains ; limits un- 
known. 

Characters. — Size large; coloration apparently normal. Skull 
long, narrow, and high, but not arched; fronto-nasal region high 
and very narrow — strongly pinched in. 

Color. — Skin of head of male killed on Pine Valley Mountain, 
southwest Utah (obtained from forest ranger, September 24, 1907, by 
Clarence Birdseye; original No. 989) : Muzzle pale brown; face and 
throat, except pale lip edgings and long hairs of median line of 
throat, dark brown, becoming grizzled posteriorly ; top of head very 
dark; grizzled posteriorly by brown-tipped hairs. 

Cranial characters. — Adult male (type, and equally old male from 
northeast corner Sevier National Forest) : Size large; skull very long, 
high, and exceedingly narrow; zygomata moderately spreading and 
outbowed; frontal shield narrow, flattened posteriorly, falling away 
laterally immediately in front of orbits, leaving a high fronto-nasal 
ridge ; short pointed posteriorly ; sagittal crest long and high, reach- 
ing anteriorly nearly to midway between fronto-parietal suture and 
plane of postorbital processes; postorbital processes very long, 
slender, peglike, and horizontally extended; rostrum long, high, 
rather narrow, and strongly compressed below nasals; palate and 
postpalatal shelf exceedingly long; postpalatal shelf and notch nar- 
row; interpterygoid fossa exceptionally deep; basisphenoid strongly 
concave. Under jaw very long, ramus flat and exceedingly broad 
vertically; coronoid blade high and moderately recurved. Dentition 
light for so large a skull; canines rather small; upper and lower 
molariform series medium or rather small; middle lower molar de- 
cidedly narrow ; M- 2 - small, the heel narrowed on outer side. 

An imperfect skull of an old male (No. 167390) from Pine Valley 
Mountain, southwest Utah, differs from the type in having still 
smaller teeth both above and below, the molars, fourth premolar, and 
canines being but little larger than those of the female from the type 
locality. 

Adult female (No. 180207, from type locality) : Similar in general 
to male, but much smaller and somewhat less extreme. Skull long 
and narrow ; f rontals and fronto-nasal region essentially the same but 
sagittal crest shorter; zygomata relatively as well as actually much 
narrower (much less spreading) and not outbowed; rostrum narrow- 
est anteriorly ; molars smaller ; canines much smaller. 

Cranial comparisons. — Old male (type) compared with adult and old 
males of bairdi (the only neighboring species of approximately same 
size) : Eostrum longer and decidedly narrower; base of rostrum in 
64854°— 18 5 



64 NORTH AMERICAN FAUNA. [No. 41. 

front of orbits more compressed; postorbital processes longer and 
more slender; palate longer; postpalatal shelf narrower; interptery- 
goid canal much deeper; ramus of jaw longer, decidedly broader 
(vertically), flatter, and much thinner under M-y and M ? ; upper 
molariform teeth, middle lower molar, and lower canines much 
smaller. 

Skull measurements. — Old male (type) : Basal length, 348 ; occipito- 
nasal length, 337; palatal length, 194; zygomatic breadth, 226; inter- 
orbital breadth, 79. 

URSUS PERTURBANS sp. NOV. 
Mount Taylob Gbizzlt. 

Type No. 222102, $ old, U. S. National Museum, Biological Sur- 
vey collection. Collected near Mount Taylor, northern New Mexico, 
July 9, 1916, by Ed. Anderson. 

Characters. — Size very large; skull long and narrow, with nar- 
rowly spreading zygomata and exceedingly high sagittal crest ; affini- 
ties with utahensis, idahoensis, and more remotely with arizonoe. 
Claws moderate, slightly curved, mainly ivory whitish on top, 
darker on sides. 

Colon/'. — Adult male (type) : General ground color dusky; face and 
head dark brown, becoming dusky around eyes; body dusky, back 
grizzled with dark golden tips; legs and feet black. 

Cranial characters. — Adult male (type) : Skull conspicuously long 
and narrow, the narrowness marked in braincase, frontal shield, 
rostrum, palate, postpalatal notch, and basicranial axis; postorbital 
processes moderately outstanding horizontally, frontal shield broadly 
and shallowly concave, becoming flat in old age, very short poste- 
riorly, the point entering sagittal crest in midfrontal region (less 
than halfway from postorbital processes to fronto-parietal suture) ; 
f ronto-nasal region very slightly dished ; rostrum narrow and rather 
high in type specimen (somewhat broader in very old skull from 
Datil Mountains, No. 140086) ; frontal part of braincase keeling into 
crest; underjaw long; coronoid blade high; ramus straight, its in- 
ferior border slightly concave under M T , only slightly upcurved 
posteriorly, and not broadly expanded vertically; diastema long. 

Teeth of medium size, about as in idahoensis (much smaller than 
in horribUis and bairdi) ; heel of M- 2 - long, flat, emarginate, and 
slightly everted, resembling that of idahoensis; PM ¥ strikingly 
small — much smaller than in any other known grizzly and no larger 
than in some of the black bears (Euarctos), its crown falling below 
plane of molar crowns. Upper canines rather small, as in idahoensis 
and arizonw; lower canines more slender than in these species. 



1918.] ARIZONJE GROUP. 65 

The skull of a still older male (No. 140086) killed some years ago 
at Kid Springs, Datil Mountains, New Mexico, 10 miles northeast of 
Datil, and secured for the Biological Survey by N. Hollister, in 
October, 1905, resembles the type in essential characters, but is even 
longer and owing to greater age has the frontal shield flatter, the 
fronto-nasal region less dished, the sagittal crest even more highly 
developed, the interpterygoid fossa even longer and narrower. 

Cranial comparisons. — Old male (type) compared with old male 
utahensis (type) : Size and general appearance similar, but underjaw 
widely dissimilar ; nasals and fronto-nasal region less elevated ; ros- 
trum smaller; sagittal crest more highly developed; zygomata less 
spreading; palate and postpalatal shelf shorter; postpalatal shelf 
less narrowed; underjaw very much smaller, shorter, and lighter, 
the ramus much less broadly expanded vertically, its inferior border 
shorter and less upcurved posteriorly; coronoid blade much smaller 
and lower. Canines much smaller; molars decidedly larger, espe- 
cially M^; heel of M^ much larger and broader posteriorly. 

Remarks. — Ursus perturbans appears to have affinities in several 
directions. In length and slenderness of skull it resembles utahensis 
more closely than any other species, but the underjaw differs amaz- 
ingly from that of utahensis, being relatively small and light, while 
that of utahensis is large and remarkably broad vertically. In 
dental characters, especially the form of M- 2 -, it resembles idahoensis. 
In the great development of the sagittal crest it exceeds all known 
grizzlies, not excepting horribilis. 

Skull measurements. — Old male (type) : Basal length, 338; oc- 
cipito-nasal length, 339; palatal length, 182.5; zygomatic breadth, 
210; interorbital breadth, 83. 

URSUS ROGERSI ROGERSI " sp. nov. 
Rogers Grizzly. 

Type No. 222983, $ ad., U. S. National Museum, Biological Survey 
collection. Collected high up on Greybull River, Absaroka Moun- 
tains, Wyoming, in the fall of 1890, by Archibald Rogers, and by him 
presented to the Biological Survey. 

Characters. — Skull very large and long, the length of the adult 
male equalling or slightly exceeding that of horribilis, bairdi, utahen- 
sis, and perturbans ; fronto-nasal region elevated and compressed, 
forming part of long fronto-nasal plane, as in bairdi and utahensis; 
frontal shield rather narrow, faintly convex transversely; post- 
orbitals rather weak and somewhat decurved ; sinus case keeling into 

1 Named for Archibald Rogers, of New York, who collected and presented the type 
specimen. 



66 NORTH AMERICAN FAUNA. tNo. 41. 

sagittal crest; rostrum long and rather slender, high posteriorly, 
strongly sloping; palate narrow and very long; interpterygoid fossa 
narrow and very long ; zygomata moderate, rather low, as in utahensis 
and idahoensis — not arched as in horribilis, bairdi, and imperator; 
underjaw long and massive with ramus rather broadly expanded 
posteriorly, but very much less so than in utahensis. Canines small; 
molars moderate. Relationships with arizonaz, bisonophagus, and 
idahoensis, but size much larger; and also with utahensis, with which 
it agrees essentially in size. 

Cranial comparisons. — Adult male (type, perhaps not quite fully 
adult) compared with males of the three related forms, arizonce 
(type, adult), bisonophagus (Jype, young-adult) and idahoensis 
(type, old) : General appearance similar, but skull as a whole, palate, 
and interpterygoid fossa much longer; underjaw very much longer, 
larger, and more massive ; coronoid blade broader and higher ; canines 
of approximately same length but lower canines more massive 
basally ; molars in general similar, but last upper molar longer, with 
heel rather broadly rounded posteriorly, instead of emarginate or 
sub triangular ; M T smaller; crown of M^much longer. More detailed 
comparisons seem unnecessary, though it may be remarked that from 
idahoensis, its nearest neighbor on the west, it differs markedly not 
only in greater size, but also in very much longer, larger, and more 
massive underjaw, and in the following dental characters: M> 
smaller ; heel of M ^ broader and not emarginate ; M-g- much longer. 

Adult male (type) compared with old male utahensis (type) : Size 
essentially the same though the basal length is greater; frontal 
shield somewhat broader; postorbitals less strongly developed, 
shorter, depressed instead of horizontally outstanding ; rostrum larger 
and less compressed; interpterygoid fossa longer and less deep, 
palate and postpalatal shelf broader; underjaw of essentially same 
length, but ramus less broadly flattened. Canines apparently less 
elongate (in utahensis broken) ; last upper molar larger, the heel 
broader and more broadly rounded posteriorly ; crowns of middle and 
last lower molars longer. 

Skull measurements. — Adult male (type) : Basal length, 353 ; oc- 
cipito-nasal length, 345 ; palatal length, 193 ; zygomatic breadth, 211 ; 
interorbital breadth, 86. 

URSUS ROGERSI BISONOPHAGUS subsp. nov. 

Black Hells Gbizzly. 

Type No. 181089, $ young-adult, U. S. National Museum, Biologi- 
cal Survey collection. From Bear Lodge, Sundance National Forest, 
Black Hills, northeastern Wyoming. Collected in February, 1887, 
by Paul Kleineidam. 



1918.] ABIZONiE GROUP. 67 

Range. — Black Hills of South Dakota and adjacent northeast cor- 
ner of Wyoming. 

Characters. — Size large; skull long, slender, and rather low, 
smoothly rounded on sides, with weak decurved postorbitals, and 
elevated fronto-nasal region. Affinities with arizonm and rogersi. 
Claws of moderate length, strongly curved, smoothly polished, dark 
horn color, marked toward tips with pale yellowish, and most of them 
with whitish (superficially) on upper side of basal half. 

Color. — Type: Muzzle pale brown (apparently old pelage) ; head 
and face blackish, becoming slightly grizzled posteriorly and on 
lower part of cheeks by wash of yellowish-brown-tipped hairs ; entire 
body, legs, and feet very dark brown overlaid on back by wash of 
light tips. 

Cranial characters. — Young-adult male (type) : Similar in general 
to rogersi. Viewed from above: Closely similar except for smaller 
size and differences in the development of certain parts attributable 
in the main to lesser age (shield more convex transversely; post- 
orbitals slightly less outstanding; fronto-nasal region slightly 
higher, almost forming a hump). Viewed from below: Palate and 
postpalatal shelf very much shorter/ postpalatal notch less narrow; 
underj aw smaller and lighter; canines longer; M 1 slightly larger; 
M- 2 - with heel subtriangular, strongly narrowed on outer side as in 
arizonce (in rogersi not narrowed but rather broadly rounded) ; M T 
slightly larger; M T much smaller. Fully adult skulls would doubt- 
less show other differences. 

Cranial comparisons. — Young-adult male (type) compared with 
adult male arizonw (type) : Skull and teeth similar but skull longer 
and narrower anteriorly; vault of cranium slightly less arched; 
rostrum narrower ; fronto-nasal region slightly more compressed and 
more elevated, continuing frontal plane; frontal shield slightly nar- 
rower, somewhat more convex transversely, its sides more smoothly 
rounded (doubtless because slightly younger) ; postorbital processes 
less developed; lambdoid crest more strongly developed; postpalatal 
shelf smaller. Underj aw slightly longer; inferior border of ramus 
longer and more upcurved posteriorly; ramus more broadly ex- 
panded vertically; coronoid blade higher and flatter (fossa less 
deeply excavated), its anterior border more nearly vertical; upper 
molars somewhat larger ; middle lower molars smaller ; upper canines 
essentially the same; lower canines somewhat longer and more 
slender. 

Remarks. — The range of bisonophagus appears to be completely 
isolated from that of its nearest relative rogersi of the mountains be- 
tween Yellowstone Park and Bighorn Basin. On the other hand, its 
range appears to overlap parts of those of absarokus, horribilis, and 
bairdi. From bairdi. which it resembles in form of skull, it is easily 



68 NORTH AMERICAN FAUNA. [No. 41. 

distinguished by decidedly smaller size and very much smaller 
canines and molars. 

Skull measurements. — Young-adult male (type) : Basal length, 
331 ; x occipito-nasal length, 323 ; palatal length, 173 ; zygomatic 
breadth, 200 ; interorbital breadth, 80. 

URSUS PERVAGOR Mebeiam. 

Lillooet Grizzly. 

Ursus pervagor Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 186-187, 
August 13, 1914 

Type locality. — Pemberton Lake (now Lillooet Lake), British 
Columbia. 

Type specimen. — No. 187887, $ ad., U. S. National Museum 
(=No. 6510, Merriam collection). Collected in May, 1883, by John 
Fannin. 

Range. — Interior of southwestern British Columbia; known only 
from Lillooet Lake and Bridge River. 

Characters. — Size rather large. External characters unknown. 

Cranial characters. — Adult male (type) : Size large; skull long, 
rather narrow, high, moderately dished; zygomata moderately 
spreading and outbowed ; frontal shield of moderate breadth, rather 
flat, strongly sloping, shallowly sulcate medially, swollen on each 
side just behind plane of postorbitals ; postorbital processes outstand- 
ing, thick, peglike; postpalatal shelf rather broad and flat; frontal 
part of braincase elevated and compressed, supporting posterior part 
of frontal shield ; palate long ; squamosal shelf long ; mastoids long ; 
interpterygoid fossa short and rather broad; under jaw long; coro- 
noid broad and rather vertical; ramus long, swoll n on outer side. 
Teeth small, particularly the canines and lower molars. 

Cranial comparisons. — Adult male (type) compared with adult 
male canadensis (type) : Considerably larger ; vault of cranium much 
higher; frontal shield much broader, higher, more strongly sloping, 
and less flat; frontal part of braincase elevated and compressed, 
rising strongly to temporal impressions (in canadensis not com- 
pressed except at sagittal crest) ; postorbital processes much larger; 
rostrum more strongly ascending; occipito-sphenoid longer (98 mm. 
against 90) ; underjaw much longer; coronoid blade higher, its apex- 
less recurved. Canines about same size ; large upper premolar, upper 
molars, and M T and M T decidedly smaller. 

Adult male (type) compared with adult male caurinus: Frontal 
shield broader, less elevated posteriorly; fronto-nasal region less 
strongly dished; coronoid blade less nearly vertical; upper canines 
slightly shorter; lower canines conspicuously smaller and shorter. 

1 Restored. 



1918.] ARIZONA GROUP. 69 

Adult male (type) compared with adult male eulophus (type and 
older skulls) : Similar in general, both having the fronto-facial 
region long sloping and flattish, but differing in many characters. 
The skull of pervagor differs from that of eulophus in being slightly 
shorter, less highly arched, frontal shield less elevated posteriorly; 
postorbitals larger, more horizontally outstanding; sagittal crest 
shorter and straighter, not arched; braincase, rostrum, and palate 
shorter; squamosal shelf shorter; postpalatal shelf broader and 
shorter; interpterygoid notch broader; under jaw smaller and less 
massive ; inferior border of ramus shorter and less broadly expanded ; 
coronoid lower, the apex less narrowed and less recurved, with less 
development of inferior ridge of fossa; teeth (canines and molars) 
slightly smaller. 

Skull measurements. — Adult male (type) : Basal length, 330; 
occipito-nasal length, 322; palatal length, 178; zygomatic breadth, 
224 ; interorbital breadth, 81. 

URSUS CAURINUS Merbiam. 

Lynn Canal Gbizzly. 

Vrsus caurinus Merriam, Proc. Biol. Soc. Washington, XXVII, p. 187, August 18, 
1914. 

Type locality. — Berners Bay, east side of Lynn Canal, Southeast- 
ern Alaska. 

Type specimen. — No. 176591, 2 ad., U. S. National Museum, Bio- 
logical Survey collection. Collected June 8, 1911 by A. Hasselborg. 

Range. — Coast of mainland of Southeastern Alaska from Cnilkat 
River valley and Lynn Canal south an unknown distance. 

Characters. — Very closely related to eulophus of Admiralty Island. 
Size rather large; skull long and rather narrow; canines long, the 
lower ones massive; claws smoothly polished. 

Color. — Upperparts yellowish buff; face and most of head pale 
brown or drab; ears, hump, and underparts conspicuously darker; 
legs and feet dark brown or brownish black. 

Cranial, characters. — Male and female : Both skulls long and nar- 
row, strongly arched posteriorly, moderately dished; frontal shield 
of medium breadth, strongly ascending; postorbital processes weak 
and decurved except in old age,; frontal part of braincase elevated, 
forming an uplifted base for posterior part of frontal shield, behind 
which it keels into sagittal crest as in eulophus; palate long; post- 
palatal shelf rather narrow; squamosal shelf long; lachrymal duct 
opening within orbital rim; ramus of jaw broad and flattened, 
notably higher posteriorly than anteriorly. Teeth rather small for 
size of skull; last upper molar long and narrow, tapering posteriorly. 



70 NORTH AMERICAN FAUNA. [No. 41. 

Unfortunately, no fully adult male caurmus has been obtained, 
but I have secured a skull of a young-adult male (No. 205169) from 
Berners Bay, Lynn Canal, and another (No. 210140) from Chilkat 
River valley. These present the following characters: Skull long, 
high, and narrow, rather highly arched and dished; frontals rising 
rather strongly from rostrum; frontal shield rather narrow, long 
sloping, convex transversely in these youngish skulls (doubtless flat- 
tish in adults), slightly sulcate medially; long pointed, the point 
nearly reaching parietals; postorbital processes rather broad and 
slightly decurved (doubtless more outstanding with age) ; fronto- 
nasal region dished; rostrum moderate; nasals nearly horizontal 
anteriorly, rising posteriorly inj:o frontal shield; frontal part of 
braincase compressed and elevated, keeling into temporal impressions 
and anterior part of sagittal crest ; squamosal shelves long. Under- 
jaw rather long; ramus moderate, its inferior border bellied pos- 
teriorly; coronoid blade moderate and rather vertical; apex not 
strongly recurved ; palate, postpalatal shelf and notch medium ; teeth 
rather small for size of skull ; molars only slightly larger than those 
of female ; M- 2 - rather narrow, with narrow slightly everted heel ; 
canines, especially lower canines, decidedly thicker than those of 
female; M T rather swollen in both male and female; M-j- small and 
narrowest posteriorly; PM T with main cusp rather small, conical, 
anterior, with gradually sloping incompletely sulcate heel, rarely 
with traces of posterior cusplets, sometimes with anterior cusplet 
on inner side of cingulum. 

The Chilkat Valley male is a year older than the Berners Bay 
skull, and broader across the frontals; postorbital processes more 
outstanding, ramus of underjaw more bellied posteriorly; coronoid 
blade broader above; M T smaller and thinner. 

Cranial comparisons. — Vrsus caurinus appears to be rather closely 
related to pervagor of the Lillooet region in the interior of British 
Columbia, and to eulophus of Admiralty Island, Southeastern Alaska, 
but unhappily no skulls of fully adult males are available for com- 
parison. 

Young-adult male (No. 210140, from Chilkat River valley) com- 
pared with adult male eulophus (type) and with a young adult 
eulophus (No. 203284), both from Admiralty Island: Frontal shield 
narrower anteriorly and much narrower posteriorly (narrowed be- 
hind postorbitals by usual incurving temporal ridges, while in 
eulophus the ridges are nearly straight and the posterior part of 
shield correspondingly broader) ; fronto-nasal region more strongly 
dished; underjaw longer; ramus thicker (more swollen on outer 
side) ; coronoid blade broader above and much less recurved. Canines 
essentially same; M T smaller; M- 2 - broader in middle and posteriorly. 

Adult female (type) compared with young-adult female eulophus 
(No. 137470) : Basal, occipito-nasal, and occipito-sphenoid lengths 



1918.] ARIZONA GROUP. 71 

essentially same; vault of cranium much less arched; frontal shield 
flatter, narrower interorbitally and postorbitally ; postorbitals much 
smaller and outstanding instead of strongly decurved; fronto-nasal 
region scarcely dished ; rostrum lower ; nasals smaller ; palate shorter ; 
under jaw more massive; coronoid broader and less recurved; M T 
thicker; M^ narrower throughout, the heel narrowed on outer side. 

Young-adult male compared with adult male pervagor: Frontal 
shield narrower, more elevated posteriorly ; fronto-nasal region more 
strongly dished ; coronoid blade more nearly vertical. M- 2 - narrower, 
with narrower heel. Upper canines slightly longer; lower canines 
conspicuously longer and larger. 

Skull measurements. — Adult female (type): Basal length, 295; 
occipito-nasal length, 285; palatal length, 161; zygomatic breadth, 
196 ; interorbital breadth, 66. 

URSUS EULOPHUS Mebriam. 

Admiralty Island Crested Bear. 

Ursus eulophus Merriam, Proc. Biol. Soc. Washington, XVII, p. 153, October 6, 
1904. 

Type locality. — Admiralty Island, Southeastern Alaska. 

Type specimen. — No. 81102, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected in 1896 by Lieut. G. T. Emmons. 

Range. — Admiralty Island. 

Characters. — Size large; color rich dark brown; claws blue-black, 
of moderate length ; skull long, rather narrow, and high, with weak 
decurved postorbital processes. 

Color. 1 — General color of head and body in fresh pelage, rich dark 
brown or seal brown; muzzle paler; legs, feet, and belly dusky or 
blackish; neck and shoulders sometimes grizzled by admixture of 
yellowish-tipped hairs. 

Cranial characters. — Adult males: Skull large, long, high, and 
rather narrow; frontal shield long and rather narrow, shallowly 
grooved medially, gradually sloping (not abruptly elevated) ; tem- 
poral impressions long and only slightly incurved, meeting at fronto- 
parietal suture; postorbital processes weak and decurved; fronto- 
nasal region elevated in plane of shield; rostrum long and high; 
anterior third of nasals horizontal, posterior two-thirds rising in 
fronto-nasal plane; zygomata moderately spreading, subangular; 
squamosal root only slightly expanded; palate and postpalatal shelf 
long and narrow ; ascending arms of maxillae long, passing far 
beyond nasals and reaching back over anterior two-thirds of orbit; 
nares rather small; braincase long anteriorly, rather narrow, com- 

1 Color from skins in Mus. Vert. Zool., Univ. California, obtained on Admiralty Island 
by Miss Annie M. Alexander. 



72 NORTH AMERICAN FAUNA. [No. 41. 

pressed, and keeling into sagittal crest; sagittal crest high, curved, 
and relatively short; ramus of jaw high vertically, its inferior border 
strongly bellied posteriorly; coronoid blade large and moderately 
high. Canines long ; molars rather small for so large a skull. 

Cranial comparisons. — Adult male (type) compared with adult 
male pervagor (type) : Skull longer; arch of cranium higher; frontal 
shield longer sloping, rising higher posteriorly; rostrum longer; 
braincase longer; sagittal crest more convex; palate and postpalatal 
shelf longer; shelf and notch narrower; under jaw larger, longer, 
more massive ; inferior border of ramus longer, more strongly bellied 
posteriorly and broader vertically ; coronoid fossa deeper, its inferior 
border much more sharply defined by strongly developed ridge for 
muscular attachment. Canines and molars larger. 

Skull measurements. — Adult male (type) : Basal length, 346; oc- 
cipito-nasal length, 343; palatal length, 190; zygomatic breadth, 
221 ; interorbital breadth, 81. 

URSUS KLAMATHENSIS Meeeiam. 1 

Klamath Gbizzly. • 

Vrsus klamathensis Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 185- 
186, August 13, 1914. 

Type locality. — Beswick, near mouth of Shovel Creek, Klamath 
Eiver, northern California. 

Type specimen. — No. 178735, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected and presented by Charles Far- 
well Edson. 

Range. — Siskiyou Mountains of northern California and southern 
Oregon, ranging north in recent times to Fort Klamath region and 
Rogue River valley ; in early days to lower Willamette Valley (pre- 
sumably same species) ; south in Sierra Nevada an unknown distance. 
(Skull from lower McCloud River referred to this species.) 

Characters. — Size of male large ; skull in general of the idahoensis 
type, but larger and with heavier canines. Claws moderate, rather 
strongly curved, horn color, washed with yellowish basally and with 
pale yellowish markings at tips, marked longitudinally with fine 
parallel striae. Skin characters unknown. 

Cranial characters. — Skull large and high, highest about 40 
mm. in front of fronto-parietal suture; rostrum long, high, and 
ascending in plane of frontal shield; fronto-nasal region elevated, 
scarcely if at all dished ; frontal shield broad, flat, sloping, and rather 
short pointed; postorbital processes moderate, peglike, horizontally 
outstanding ; sagittal crest long and well developed ; lachrymal duct 

» Tentatively included in arizonce group. (See Introduction, pp. 12-13.) 



1918.] ARIZONA GROUP. 73 

opening on orbital rim (rather posteriorly than anteriorly) ; zygo- 
mata moderately spreading, only moderately expanded vertically; 
anterior (frontal) part of bra incase keeling into sagittal crest; 
occiput produced posteriorly (overhang much greater than in 
calif ornicus) ; squamosal shelves long and broad; palate rather flat, 
slightly arched lengthwise; postpalatal shelf broad and flat; occipito- 
sphenoid short for so large a skull (length 89 mm.) ; basioccipital 
very broad anteriorly; mastoids vertical and short. Under jaw long; 
ramus exceptionally broad and flat vertically; coronoid blade large 
and high, broad basally. Canines very large ; molars moderate ; last 
upper molar relatively small, the heel emarginate or obliquely trun- 
cate on outer side; middle lower molar with anterior moiety much 
larger than posterior. Large upper premolar absent and no trace 
of alveolus. 

Cranial comparisons. — Curiously enough klamathensis does not re- 
quire close comparison with any of the other species inhabiting Cali- 
fornia, its only near relatives being members of the shoshone- 
idahoensis group of the Rocky Mountains, and pervagor of interior 
British Columbia. 

Adult male (type) compared with old male idahoensis (No. 187888, 
=No. 93, Merriam collection, from North Fork Teton River, 
eastern Idaho) : Similar in general form and proportions but larger 
( condylobasilar length 350 mm. contrasted with 335) ; vault of cra- 
nium somewhat higher; frontal shield broader and flatter; palatal 
length about the same; postpalatal length much greater (150 con- 
trasted with 135 ) ; opening of lachrymal duct slightly more posterior ; 
occipital overhang greater; basioccipital anteriorly very much 
broader; ramus of under jaw longer and much broader vertically; 
coronoid blade higher. Canines larger and longer ; last upper molar 
shorter. 

Adult male (type) compared with adult male pervagor (type) : 
Similar in size and general characters; vault of cranium slightly 
higher ; frontal shield flatter and somewhat broader ; postorbitals not 
quite so large; fronto-nasal region more elevated (in pervagor 
slightly dished); rostrum broader anteriorly; occipital overhang 
greater; ramus of under jaw much more broadly expanded vertically 
and flatter; canines and molars very much larger. 

Adult male (type) compared with old male henshawi (type) : Ursus 
klamathensis and U. henshawi belong to widely different groups and 
do not require detailed comparison. U. klamathensis may be distin- 
guished at a glance by its much larger size, much higher vault of 
cranium, highly elevated and continuously sloping fronto-nasal 
region and rostrum, and peglike postorbitals — in striking contrast 
to the much smaller, lower, and strongly dished skull of henshawi, 



74 NORTH AMERICAN FAUNA. [No. 41. 

with its low depressed rostrum and large broadly rounded post- 
orbitals. 

Adult male (type) compared with adult male californicus (from 
coast region south of San Francisco Bay) : The differences are 
marked in the skull and striking in the teeth. In klamathensis the 
vault of the cranium is lower posteriorly and higher anteriorly ; the 
frontal shield flatter laterally; the rostrum shorter; the base of the 
cranium (occipito-sphenoid) decidedly shorter. The last upper and 
middle lower molars are widely different, the heel of M> in cali- 
fornicus large, long, and broad posteriorly, while in klamathensis it 
is small and emarginate on outer side; the anterior part of M 7 in 
californicus is normal, while in klamathensis it is disproportionately 
large. 

Skull measurements. — Adult male (type) : Basal length, 331 ; oc- 
cipito-nasal length, 337; palatal length, 175; zygomatic breadth, 223; 
interorbital breadth, 85.5. 

URSUS MENDOCINENSIS Meekiam. 1 

Mendocino Grizzly. 

Ursus mendocinensis Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 145-146, 
September 6, 1916. 

Type locality. — Long Valley, Mendocino County, California. 

Type specimen.— No. 206625, $ old, U. S. National Museum, Bio- 
logical Survey collection. Obtained through Charles J. and Frank 
H. Hittell. 

Characters. — Size rather large, about equaling klamathensis but 
apparently smaller than californicus and colusus; external characters 
unknown. Affinities with klamathensis, with which it may inter- 
grade at the north. 

Cranial characters. — Skull short, broad, highly arched, and 
strongly dished, with widely outstanding zygomata and truncate occi- 
put. Frontal shield of moderate breadth, short pointed posteriorly, 
slightly convex between orbits, strongly sloping to rostrum ; rostrum 
short, broad, and strongly depressed ; postorbital processes moderate, 
sub-peglike, horizontally outstanding; sagittal crest high, thick, 
humped anteriorly, short posteriorly ; occiput obliquely truncate ; oc- 
cipital overhang slight compared with that of californicus and colu- 
sus; palate short and rather broad; postpalatal shelf of moderate 
breadth, flat; postpalatal notch moderate; mastoids rather short, 
directed anteriorly. Under jaw absent. Teeth gone except left hind 
molar, which is short, with small heel, obliquely truncate on outer 
side (as in klamathensis). 

1 Tentatively included in arizonw group. (See Introduction, pp. 12-13.) 



1918.] ARIZONA GROUP. 75 

Cranial comparisons. — Similar in general to klamathensis but 
frontonasal region strongly dished, rostrum shorter, broader, flatter 
on top, and depressed instead of elevated; zygomata more widely 
outstanding; palate broader; occipital overhang less. 

Skull measurements. — Old male (type) : Basal length, 327; occi- 
pito-nasal length, 323; palatal length, 183; interorbital breadth, 84.5. 

URSUS MAGISTER Merriam. 1 

Southeen California Grizzly. 

Ursus magister Merriam, Proc. Biol. Soc. Washington, XXVII, p. 189, August 
13, 1914. 

Type locality. — Los Biacitos, head of San Onofre Canyon, Santa 
Ana Mountains, Southern California. 

Type specimen. — No. 160155, $ old, U. S. National Museum, Bio- 
logical Survey collection. Killed in August, 1900 or 1901, by Henry 
A. Stewart and by him presented to the Biological Survey. 

Range. — Santa Ana or Trabuco Mountains, Cuyamaca and Santa 
Rosa Mountains, and probably San Jacinto Mountains. Believed to 
be extinct. 

Characters. — Size of male huge (estimated weight over 1,400 
pounds), largest of known grizzlies, considerably larger than cali- 
fomicus of the Monterey region, and even than horribilis, the 
great buffalo-killing grizzly of the Plains (only equaled by the 
largest alexandrce of Kenai Peninsula) ; sexual disparity great; skull 
of female hardly half the bulk of male; skull of male of a rather 
generalized type; not dished. Claws of old female from head of 
Trabuco Canyon, Santa Ana Mountains, exceedingly long, strongly 
curved, mainly yellowish above. 

Color. — (Old female from head of Trabuco Canyon) : General 
color dusky or sooty all over except head and grizzling of back. 
Muzzle gray or mouse brown, palest above; top of head and neck 
very dark brown, sparsely grizzled with pale-tipped hairs; back 
dusky grizzled with grayish; legs and underparts wholly blackish. 

Cranial characters. — Adult male (type) : Skull exceedingly large, 
long; vault of cranium arched, but not abruptly; rostrum long and 
high ; f ronto-nasal region elevated, in same plane with frontal shield 
and rostrum; frontal shield flattish-convex, faintly sulcate medially 
and slightly swollen on each side between postorbital processes, the 
point decurved and reaching fronto-parietal suture; zygomata only 
moderately spreading, angular, the posterior root expanded and 
rising abruptly from plane of squamosal shelf; sagittal crest rather 
short; palate scooped out anteriorly, forming a basinlike depression 
surrounding the anterior palatine foramina; occipito-sphenoid 

1 Tentatively included in arizona group. (See Introduction, pp. 12-13.) 



76 NORTH AMERICAN FAUNA. [No. 41. 

length 103.5 mm. Underjaw long; ramus broad and flat vertically, 
its inferior border moderately bellied and incurved posteriorly; 
coronoid blade large, its apex strongly recurved, cutting plane of 
middle of condyle. Teeth large and broad but by no means dis- 
proportionate to large size of skull ; M- 1 broad ; last upper molar 
absent, but from its alveolus and its form in female, obviously broad, 
short, strongly triangular, the heel small, narrowed posteriorly, ob- 
liquely emarginate on outer side; PM ¥ broad, with rather short 
slightly sloping heel, narrow imperfect sulcus without posterior 
cusplets; M T apparently normal (much worn in type specimen) ; 
middle lower molar absent in type specimen but apparently normal 
(judging from the female, in which, however, it is badly worn). 

Female of extreme age (No. 156594, from Trabuco Canyon, killed 
January 5, 1908, by Andrew Joplin and Edward Adkinson) : Size 
small; rostrum short and depressed; fronto-nasal region strongly 
dished; frontal shield flattish, slightly sulcate interorbitally, short 
pointed, beaded posteriorly by elevated temporal impressions, rising 
rather abruptly at orbits; sagittal crest long and nearly horizontal; 
palate and postpalatal shelf broad, flat posteriorly, concave ante- 
riorly. 

Cranial comparisons. — Ursus magister does not require close com- 
parison with any other species. While the largest skulls of old male 
calif ornicus equal it in basal length, they are so much lower, nar- 
rower, and smaller in every way that detailed comparisons are un- 
necessary. The species which it most nearly resembles is bairdi from 
the mountains of Colorado, but the resemblance is not close. It 
differs from bairdi in somewhat larger size, much more highly 
arched vault of cranium, much broader and more strongly sloping 
frontal shield, more posterior mastoids, longer underjaw with much 
more broadly flattened and less massive ramus, and in important 
tooth characters. 

Between the two geographically is utahensis, which, like magister, 
has the ramus of the underjaw very broadly flattened vertically, but 
in form of cranium utahensis goes to the opposite extreme, the frontal 
shield, rostrum, and braincase being exceptionally narrow, and the 
fronto-nasal region compressed and elevated. 

Flesh measurements. — Old male (type) : Height at shoulder from 
flat of foot 4 ft. (=1,220 mm.) ; total length, snout to tail, 9£ ft. 1 
(=2,900 mm.); sole of largest foot without claws: length 12 in. 
(=305 mm.) ; breadth 8 in. (=204 mm.). Length of old female from 
Trabuco Canyon, measured in the flesh by Andrew Joplin, 6 ft. 3 in. 

Skull measurements. — Old male (type): Basal length, 365; oc- 
cipito-nasal length, 366; palatal length, 197; zygomatic breadth, 236; 
interorbital breadth, 97. 

* Apparently an error ; possibly intended for snout to claws of extended hind foot. 






1918.] HYLODROMUS GROUP. 77 

Hylodromus Group. 

URSUS HYLODROMUS Elliot. 

Forest Grizzly. 

(Plate XI.) 

Ursus hylodromus Elliot, Field Columb. Mus. Pub. 87, Zool. Ser. Ill, pp 257- 
258, December, 1913. (Described as a black bear!) Purchased from 
Mackay & Dippie, taxidermists, Calgary. 

Ursus selkirki Merriam, Proc. Biol. Soc. Washington, XXIX, p. 150, September 
6, 1916. (From Selkirk Mountains, Upper Columbia River, British Co- 
lumbia.) 

Type locality. — Eocky Mountains of western Alberta (precise lo- 
cality unknown). 

Type specimen. — No. 19065, 5 young-adult, Field Museum of Nat- 
ural History. 

Range. — Rocky Mountain region of western Alberta and eastern 
British Columbia, including Selkirk Range. 

Characters. — Size of male large, of female small; external charac- 
ters unknown. 

Cranial characters. — Young -adult female (type *) : Size small, skull 
short posteriorly (occiput less extended than in most species) ; mod- 
erately arched and dished; zygomata not outstanding and only 
slightly bowed; frontal shield of moderate breadth, rising rather 
strongly from rostrum, its apex sublyrate ; postorbital processes large, 
outstanding; rostrum rather high, strongly ascending posteriorly; 
sagittal crest short, reaching only two-thirds distance from occiput 
to fronto-parietal suture; squamosal shelf narrow and nearly hori- 
zontal; postpalatal shelf rather broad. 

Old male (No. 205170) from Selkirk Mountains, upper Columbia 
River, British Columbia (assumed to be typical) : Size medium; skull 
long, low arched, highest immediately in front of fronto-parietal 
suture, and of medium breadth; braincase and palate arched; shield 
flattish, long pointed, sloping gradually from point to rostrum, 
faintly sulcate medially; postorbitals broadly subtriangular, flat on 
top and slightly decurved, convex posteriorly, concave anteriorly; 
rostrum moderate, nearly horizontal ; f ronto-nasal region rising very 
gradually into shield ; sagittal crest short ; palate of medium breadth, 
slightly troughed and arched ; postpalatal shelf broad and flat ; zygo- 
mata subtriangular, not widely outstanding, strongly arched antero- 
posterior^; squamosal root long but not broadly expanded; mas- 

1 Through the courtesy of the officials of the Field Museum I have had the privilege of 
examining the type skull Of hylodromus and comparing it with skulls in the National 
Museum collection. 



78 NORTH AMERICAN FAUNA. [No. 41. 

toids short, not divergent; meatus tube large and free; ramus of 
underjaw straight; coronoid blade low, broad in middle part. Teeth 
of medium size (too badly worn to admit of description). 

Cranial comparisons. — Old male (No. 205170, from Selkirk Moun- 
tains) compared with old male kluane (type) : Size smaller; vault of 
cranium materially lower ; top of skull lower and flatter throughout ; 
shield narrower, much flatter, and longer pointed ; sagittal crest much 
shorter and lower ; occipital overhang and inion much less developed ; 
zygomata more triangular (less bowed) ; braincase, palate, and 
underjaw much shorter; coronoid blade lower; subangular border 
shorter and more strongly defined. Teeth very badly worn but ca- 
nines and molars evidently mucl> smaller. 

Compared with idahoensis and chelan (both types) : Similar in 
general but smaller; vault of cranium much lower; frontal shield 
somewhat narrower, longer pointed posteriorly ; postorbital processes 
broader basally but less widely outstanding; sagittal crest much 
shorter, its anterior part less distinctly keeled from sinus case; 
zygomata shorter and more sharply triangular; underjaw shorter; 
coronoid blade lower; subangular notch and border similar. 

Compared with latifrons, whose range it approaches on the north 
but with which it does not appear to be related, it is easily dis- 
tinguished by smaller size, flatter and very much narrower frontal 
shield, more elevated and evenly sloping fronto-nasal region, less 
outbowed and more triangular zygomata, much shorter underjaw and 
ramus, and much lower coronoid blade. 

Remarks. — The type specimen of hylodromus is a skull of a young- 
adult female from western Alberta, exact locality unknown. Until 
recently so few males have been available from this region that I was 
long in doubt as to which was its proper mate. In the light of pres- 
ent material, however, it has been possible to match up males and 
females of most of the species of eastern British Columbia and west- 
ern Alberta with some confidence, and I now feel reasonably certain 
that the type specimen of the species described by me as Ursus sel- 
hirhi from the Selkirk Mountains on the upper Columbia River is 
in reality an old male hylodromus, the name selkirki thus falling as a 
synonym. 

Skull measurements. — Old male (No. 205170) from Selkirk Moun- 
tains: Basal length, 305; occipito-nasal length, 306; palatal length, 
169; zygomatic breadth, 206; interorbital breadth, 74. Female 
young-adult (type): Basal length, 275; occipito-nasal length, 257; 
palatal length, 154 ; zygomatic breadth, 173 ; interorbital breadth, 69. 



1918.] HYLODROMUS GROUP. 79 

URSUS KLUANE KLUANE Meebiam. 

Kxuane Grizzly. 

Ursus kluane Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 141-143, Sep- 
tember 6, 1916. 

Type locality. — McConnell River, Yukon. 

Type specimen. — No. 204188, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected by Smith and Geddis July 15, 
1914. 

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 
(north-northeast of Teslin Lake) and probably south into northwest 
corner of British Columbia. 

Cranial characters. — Adult male: Skull medium, rather long, nar- 
row, somewhat arched and dished, with long braincase, long convex 
sagittal crest, and unusually broad decurved postorbitals. Frontal 
shield of medium width, strongly convex both transversely and 
antero-posteriorly, rising rather strongly from rostrum, slightly sul- 
cate medially and moderately swollen over orbits ; very short pointed, 
the point ending about midway between parietals and plane of post- 
orbitals; postorbitals broad, decurved, strongly convex anteriorly, 
concave posteriorly ; f ronto-nasal region somewhat depressed ; ros- 
trum high and narrow, rounded above (subterete) ; nares truncate; 
sagittal crest very long and arcuate; occipital overhang and inion 
well developed; zygomata not widely outstanding, somewhat bowed, 
rounded posteriorly; palate moderate; postpalatal shelf large and 
broad; notch rather broad and short; mastoids long and divergent; 
underjaw rather long; coronoid blade high and narrow, the apex 
rather strongly recurved; teeth too badly worn to admit of descrip- 
tion (apparently large for size of skull). 

Adult female: Size small, nearly as small as female pallasi; 
f ronto-nasal region moderately dished and usually sulcate ; braincase 
moderately arched, highest just in front of fronto-parietal suture; 
temporal impressions meeting over anterior part of parietals (prob- 
ably somewhat more anteriorly in old skulls) ; zygomata moderately 
outbowed, subtriangular ; frontal shield of medium breadth, lyrate 
pointed posteriorly; postorbital processes rather broad for so small 
a skull, moderately decurved; underjaw short; coronoid blade broad 
basally and rather short. Teeth (canines, incisors, and molars) 
rather large for size of skull, decidedly larger than in pallasi; molars, 
both upper and lower, very much larger. 
64854°— 18 6 



80 NORTH AMERICAN FAUNA. [No. 41. 

Cranial comparisons. — The species requiring comparison with 
hluane are toklat, pulchellus, and pallasi. Old male (type) com- 
pared with old male toklat, from Alaska Range, near north base of 
Mount McKinley: Size slightly larger; occipito-nasal length, length 
of braincase, and length of sagittal crest very much greater ; frontal 
shield more convex transversely; postorbital processes much larger 
and broader; rostrum higher, more rounded on top; nares more 
squarely truncate; underjaw longer; inferior border of ramus more 
convex posteriorly ; coronoid blade decidedly higher, narrower above, 
the apex more strongly recurved; teeth badly worn in both, but 
canines decidedly longer in kluane; molars apparently somewhat 
larger. / 

Compared with adult male pulchellus : Similar in general, but skull 
as a whole, braincase, palate, and underjaw very much longer; 
frontal shield very much broader; inferior border of ramus less 
abruptly upcurved; subangular border less nearly horizontal and 
much less sharply defined. 

Adult female compared with adult female toklat (comparison 
hardly necessary because of the great difference in size) : Basal length 
at least 20 mm. less ; vault of cranium and frontal shield lower ; brain- 
case less constricted anteriorly ; posterior part of shield much longer 
and broader, reaching or passing the f ronto-parietal suture ; sagittal 
crest much shorter; postpalatal shelf less broad; underjaw and 
inferior border of ramus shorter ; coronoid blade about same height ; 
canines about same size; molariform series (upper and lower) about 
same length, but proportions of individual teeth different: M^-much 
larger ; M> with shorter heel ; M T larger. 

Old male (type) compared with old male pallasi (type) : Size de- 
cidedly greater; skull about an inch longer and much more highly 
arched, with conspicuously longer braincase and longer sagittal crest; 
crest strongly arched instead of nearly straight; postorbitals very 
much larger, broader, and more strongly decurved; fronto-nasal 
region much more elevated and less dished; rostrum much higher, 
rounded above instead of depressed ; palate much longer, more arched 
and more concave; underjaw much longer; coronoid higher. 

Adult female compared with adult female pallasi: Skulls very 
much alike in size and appearance (that of kluane slightly larger), 
but teeth strikingly different. In kluane, canines larger; molars very 
much larger. 

Skull measurements. — Old male (type): Basal length, 317; oc- 
cipito-nasal length, 324; palatal length, 177; zygomatic breadth, 210; 
interorbital breadth, 85. 



1918.] HYLODROMUS GROUP. 81 

URSUS KLUANE IMPIGER subsp. nov. 

Industrious Geizzlt. 

Type No. 210708, $ , not quite fully adult. From Columbia Valley, 
British Columbia. Collected in April, 1914, by Mackay & Dippie. 

Cranial characters. — Adult male: Similar in general to kluane (No. 
221620, $ ad., Kluane River), but shield narrower and less sulcate, 
less swollen over orbits ; nasals less elevated ; palate much shorter and 
narrower; postpalatal shelf narrower; under jaw shorter; subangular 
border shorter; subangular notch and space much more strongly de- 
fined. Teeth smaller throughout, especially M> and My, but My 
swollen as in kluane. 

Female young-adult from Brisco, 1 Columbia Valley (No. 210707) : 
Vault well arched; shield posteriorly broadly lyrate, elevated by 
rising sinus case; rostrum long and slender; fronto-nasal region 
dished; palate long, troughed; underjaw long and straight. Teeth 
large. 

Cranial comparisons. — Male young-adult (type) compared with 
old male hylodromus from Selkirk Mountains (No. 205170) : Vault 
of cranium much more highly arched and narrower, the narrow 
frontal shield rising higher and more strongly, but not abruptly, 
from rostrum; shield strongly convex (in old hylodromus nearly 
flat) ; postorbitals much more slender, outstanding, depressed, and 
somewhat decurved as in kluane'; fronto-nasal region more dished; 
palate narrower; inferior border of underjaw much longer; sub- 
angular border shorter; coronoid blade higher. 

Female young-adult (No. 210707) compared with female kluane: 
Skull as a whole, rostrum, occipito-sphenoid, palate, and underjaw 
much longer ; ramus straighter ; subangular border same ; teeth essen- 
tially same except that M T is smaller. Compared with female hylo- 
dromus (type) : Skull as a whole, rostrum, occipito-sphenoid, palate, 
and underjaw longer (but not so much longer as in comparison with 
female kluane) ; ramus straighter; subangular border shorter; teeth 
similar. 

Remarks. — Ursus hylodromus, U. impiger, and U. kluane form a 
rather closely related group ranging from western Alberta and south- 
eastern British Columbia northwesterly to southwestern Yukon. 
From the material now in hand impiger appears to be most nearly 

1 Skulls of females identified as impiger have been examined from Brisco, Columbia 
Valley, British Columbia (No. 210707) ; Morley, Alberta (No. 210706) ; Jasper, Alberta 
(No. 222745) ; and headwaters North Fork Blackfoot River, western Montana (No. 
203188). 



82 NORTH AMERICAN FAUNA. [No. 41. 

related to kluane, but additional skulls of adults may show that its 
affinities with hylodromus are equally close. 

/Skull measurements. — Adult male (type): Basal length, 309; oc- 
cipito-nasal length, 293; palatal length, 162; zygomatic breadth, 197; 
interorbital breadth, 72. 

URSUS PELLYENSIS sp. nov. 
Pelly Grizzly. 

Type No. 215477, $ young-adult. U. S. National Museum, Bio- 
logical Survey collection. From the Ketza Divide, Pelly Moun- 
tains, Yukon. Collected September 30, 1915, by Fred E. Enevoldsen. 

Characters. — Skull of medium size, rather long and narrow, ap- 
parently related to hylodromus : Frontal shield narrow, long pointed 
(point reaching fronto-parietal suture but in older specimens ending 
much more anteriorly), rising strongly from rostrum; rostrum 
rather narrow and high; fronto-nasal region dished; postorbitals 
broad, strongly decurved (in old skulls doubtless more outstanding) ; 
sinus case keeling into posterior part of shield; palate moderate 
(rather broad for so narrow a skull), slightly arched and slightly 
troughed; under jaw long for size of skull; inferior border of ramus 
long, not upturned; subangular border short; coronoid rather high; 
canines long, the lower ones large for size of skull ; molars long and 
narrow, especially M T and M T ; anterior part of M ¥ exceptionally 
long and posterior part relatively narrower than in most species; 
PM ? subconical. 

Female skull (based on four rather old specimens from Pelly and 
Ross Mountains, Nos. 215710, 215711, 215713, and 221600) : Size small; 
frontal shield of moderate breadth, rather short pointed posteriorly, 
entering sagittal crest anterior to fronto-parietal suture; postorbital 
processes well developed, outstanding ; rostrum rather small ; frontal 
shield rising moderately from rostrum, shallowly sulcate medially; 
zygomata subtriangular. Dentition moderate; heel of last upper 
molar slightly emarginate, rounded posteriorly; canines small and 
slender. 

Cranial comparisons. — Young-adult male (type) compared with 
young-adult male hylodromus (No. 210708 from Columbia Valley, 
British Columbia) : Size smaller (basal length about 10 mm. less) ; 
vault of cranium less highly arched; sinus case more definitely keeled 
to support posterior part of shield; postorbitals broader, more de- 
curved, less outstanding; palate broader; molar series about same; 
upper canines essentially same; lower canines larger; upper molar 
series essentially same length; lower molar series longer and nar- 
rower; M T much longer and more slender. 



1918.] HYLODROMUS GROUP. 83 

Young-adult male (type) compared with adult male pulchellus 
from the neighboring Eoss River (No. .221599) : Size slightly 
greater; shield and rostrum broader (in pulchellus exceptionally 
narrow); postorbitals broader and more decurved; palate longer 
and broader; postpalatal notch much shorter; coronoid blade larger; 
molars, upper and lower, strikingly narrower and less massive. 

Female pellyensis compared with female toklat: Size smaller; vault 
much lower and depressed — sinus case not keeled into front of sagit- 
tal crest; fronto-nasal region less strongly depressed and dished; 
rostrum and nasals flatter; palate about same length; postpalatal 
length much less; under jaw much shorter. Canines smaller; molar 
series shorter; heel of M- 2 - less elongate. 

Skull measurements. — Young-adult male (type) : Basal length, 
299; occipito-nasal length, 294; palatal length, 168; zygomatic 
breadth, 186 ; interorbital breadth, 73. 

URSUS ANDERSONI 1 sp. Nov.* 
Anderson Bear. 

Type No. 34402, $ ad. (rather old), American Museum of Nat- 
ural History. Collected on east branch Dease River 3 near Great 
Bear Lake, Mackenzie, May 12, 1911, by Dr. R. M. Anderson. 

Characters. — Size medium or rather large; frontal region includ- 
ing postorbitals rather flat and narrow as in pellyensis (in strong 
contrast with the broader swollen frontals and elevated outstanding 
postorbitals of richardsoni) ; vault moderately arched. 

Cranial characters. — Frontal shield rather narrow, strongly slop- 
ing, flattened, short pointed posteriorly ; postorbitals moderate (short 
in contrast with those of richardsoni), horizontal (not decurved); 
sagittal crest long and strongly developed, humped anteriorly, pro- 
jecting posteriorly in prominent inion; rostrum narrow, rather 
high, compressed below middle of nasals; orbits strongly sloping 
(retreating) ; occipital overhang marked; squamosal shelf long; pal- 
ate long and narrow; postpalatal shelf moderate; zygomata well out- 
standing and somewhat bowed. Underjaw moderate ; coronoid blade 
strongly recurved, apex sharp pointed. Molars broad and rather 
large: M A and M„ together 62 mm. (in type skull) ; lower molars 74 
mm. Canines large (too much broken to afford measurements). 

Cranial comparisons. — The adult male resembles, though not very 
closely, both pellyensis and tahltanicus. Compared with the type 
specimen of pellyensis: Size essentially same; shield shorter and 
flatter with more horizontally outstanding postorbitals (differences 

1 Named for Dr. R. M. Anderson, who collected the type specimen. 

2 Tentatively included in hylodromus group. (See Introduction, pp. 12-13.) 

8 Not to be confused with the better-known Dease River of northern British Columbia. 



84 NORTH AMERICAN FAUNA. [No. 41. 

attributable in large part at least to difference in age) ; zygomata 
much less broadly outstanding (also attributable to age) ; postpalatal 
notch decidedly longer; coronoid blade lower. Canines apparently 
larger (badly broken) ; molars broader and more massive. Com- 
pared with an old male (No. 134486) from Macmillan River, pre- 
sumably old petty ensis, it is much smaller and narrower; shield 
narrower, flatter, and shorter posteriorly; postorbitals flatter and 
more nearly horizontal on top; rostrum narrower; sagittal crest 
longer and rising higher anteriorly; palate, postpalatal shelf, di- 
astema, and occipito-sphenoid much shorter; under jaw much shorter; 
coronoid lower. Dentition heavier ; canines and molars larger. 

Compared with old male tahltdnicus (type), with which it agrees 
essentially in size, height of vault of cranium, slope and flatness of 
shield, form of rostrum, spread and form of zygomata, and size and 
form of underjaw, it differs as follows: Shield narrower both inter- 
orbitally and across postorbitals; point of shield much shorter; 
fronto-nasal region slightly dished (in tahltanicus not dished) ; 
sagittal crest much longer, higher anteriorly, and keeled from sinus 
case. Dentition heavier ; canines and molars much larger. 

Skull measurements. — Adult male (type) : Basal length, 300; oc- 
cipito-nasal length, 285 ; palatal length, 165 ; zygomatic breadth, 211 ; 
interorbital breadth, 76. 

Horriaeus Group. 

URSUS APACHE Merriam:. 

Apache Grizzly. 

Ursus apache Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 134-135, Sep- 
tember 6, 1916. 

Type locality. — Whorton Creek, south slope of White Mountains, 
eastern Arizona (a few miles west of Blue). 

Type specimen. — No. 212436, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected April 3, 1913, by B. V. Lilly. 

Cranial characters. — Skull short, broad, and low, rather massive, 
moderately dished, with broad frontal shield and exceedingly broad 
outstanding postorbitals. Frontal shield broad, shallowly sulcate 
medially between orbits ; very slightly and rather flatly swollen over 
orbits; long pointed posteriorly, meeting short sagittal crest at 
fronto-parietal suture; rostrum short, high, and rather narrow; 
zygomata strongly outbowed and outstanding anteriorly as well as 
posteriorly ; ramus of jaw rather short, bellied under last molars; 
coronoid blade high, sloping strongly outward, the apex overarching 
shallow coronoid notch, but not cutting plane of condyle; dentition 
moderate. 

Cranial comparisons. — Old male (type) compared with old male 
horriceus (type) : Size somewhat greater ; vault of cranium materially 



1918.] HOKBLEUS GROUP. 85 

higher; shield much broader both interorbitally and across postor- 
bital processes; the posterior part much longer and horizontal or 
sloping posteriorly (in horriceus short and sloping strongly forward 
from apex of posterior point); sagittal crest very much shorter; 
palate broader and flat (in horriceus concave and arched) ; zygomata 
broadly outbowed (in horriceus narrowly triangular) ; meatus tube 
strongly compressed between mastoid and glenoid (in horriceus large 
and free) ; underjaw much more massive; ramus longer; broadly ex- 
panded vertically and upcurved posteriorly; coronoid blade higher. 
Teeth apparently similar (in horriceus badly worn). 

Compared with adult male absarokus, to which it appears to be re- 
lated, the skull of apache differs as follows : Vault of cranium lower, 
less arched; frontal shield broader and flatter; postorbitals much 
broader and flatter, standing out more horizontally ; f ronto-nasal re- 
gion more depressed; rostrum shorter; orbits notably smaller (lower 
vertically); squamosal trough shorter antero-posteriorly ; zygomata 
very much more strongly outbowed and conspicuously more out- 
standing anteriorly; underjaw and inferior border of ramus shorter; 
coronoid blade of equal height; teeth slightly smaller (difference 
slight). 

Old male (type) compared with adult male arizonce (type) : Basal 
length, occipito-nasal length, length of palate, interorbital breadth, 
and occipito-sphenoid length essentially the same; zygomata very 
much more outstanding and bowed instead of subtriangular ; frontal 
shield less flat, of essentially the same breadth interorbitally as in 
arizonce, but very much wider across postorbital processes, rising 
strongly from plane of rostrum; postorbital processes much more 
broadly outstanding; orbital rims more swollen; fronto-nasal region 
much less elevated; rostrum much smaller, narrower, less swollen, 
depressed basally, and horizontal instead of tapering; palate and 
postpalatal shelf much narrower. Underjaw stronger; ramus 
broader vertically; coronoid blade higher; molars slightly smaller; 
heel of M> shorter, less distinctly emarginate on outer side (more 
tapering) . 

Skull measurements. — Adult male (type) : Basal length, 325 ; 
occipito-nasal length, 315; palatal length, 171.5; zygomatic breadth, 
234 ; interorbital breadth, 89. 

URSUS HORRI^US Baibd. 

New Mexico Grizzly. 

(PI. XV.) 

Ursus horribilis var. horriceus Baird, Mammals North Amer., Pacific RR. 

Reports, VIII, pp. 224, 225, 1857 (name, type locality, and measurements). 
Ursus horriMUs var. horriceus Baird, Mammals Mexican Boundary Survey, pp. 

24-29, 1859. (Full description, including a specimen from Nogales, since 

made the type of another species — kennerlyi.) 



86 NORTH AMERICAN FAUNA. [No. 41. 

Ursus {Darin) horriaceus Gray, Catalog. Cam., Pach., and Edent, British 
Museum, p. 229, 1869. 

Type locality. — Coppermines, southwestern New Mexico. 

Type specimen. — No. 990, $ old, U. S. National Museum. Col- 
lected in 1855, by J. H. Clark. 

Range. — Parts of New Mexico, south to Casas Grandes, Chihuahua, 
Mexico; probably extending into eastern Arizona. 

Characters. — Size medium; external characters unknown; skull 
low and flat with broad outstanding postorbital processes. 

Cranial characters. — Old male (type) : Skull long, low, narrow, and 
flat; frontal shield short pointed posteriorly, temporal impressions 
conspicuously beaded, curving strongly inward, and meeting half- 
way between plane of postorbital processes and fronto-parietal 
suture ; frontal shield flatfish, shallowly concave, swollen over orbits ; 
postorbital processes large, broad, widely outstanding and moderately 
decurved; fronto-nasal region slightly dished; rostrum narrow; sag- 
ittal crest long and rather high ; occipital overhang and inion marked ; 
zygomatic arches angular, rather squarely but not widely spreading 
posteriorly; squamosal root of zygoma moderately expanded ver- 
tically; lachrymal duct cutting orbital rim anterior to orbit; palate 
and interpterygoid fossa rather narrow. Teeth of medium size 
(badly worn in type). 

An old female (No. 67405, from mountains north of Silver City, 
New Mexico, near type locality), collected in 1893 by Dr. A. K. 
Fisher, is assumed to be horriceus. Unfortunately the occipital region 
is absent so that measurements of length can not be taken. Skull low, 
with moderately spreading angular zygomata, flat long-pointed 
frontal shield, elevated flat fronto-nasal region (in plane of shield), 
and narrow rostrum, without trace of dishing except slight change of 
angle at middle of nasals; highest point of cranium about 25 mm. 
anterior to fronto-parietal suture; postorbital processes moderate or 
weak, subtriangular, slightly decurved; lachyrmal duct cutting 
orbital rim anteriorly; rostrum somewhat compressed a little below 
nasals; palate short (145), concave between last molars; postpalatal 
shelf moderately broad, short, and flat. Teeth small. 

Cranial comparisons. — Old male (type) compared with old male 
apache (type) : Size somewhat smaller ; vault of cranium lower ; shield 
much narrower, the point much shorter; palate concave and arched 
instead of flat ; zygomata narrowly triangular instead of broadly out- 
bowed; ramus of jaw shorter. 

Old male (type) compared with adult male shoshone (type) : Size 
essentially the same (basal length slightly less but occipito-nasal 
length same); vault of cranium much lower and flatter; frontal 
shield much lower and more nearly horizontal, less strongly sloping, 



1918.] HORRIiEUS GROUP. 87 

slightly broader, more swollen over orbits, and broadly depressed 
interorbitally instead of flat or slightly convex ; postorbital processes 
very much larger, broader, and more widely outstanding; frontal 
part of braincase broader and depressed — not keeling into sagittal 
crest; zygomata angular — not outbowed; occipito-sphenoid shorter; 
lachrymal duct more anterior; inferior border of ramus shorter; 
coronoid blade less high. 

Old female compared with old females of nelsoni: Size larger 
(basal length about 10 mm. longer) ; vault of cranium higher over 
posterior frontal region; frontal shield broader, flatter anteriorly; 
zygomatic arches slightly more spreading. Teeth, especially canines 
and molars, materially larger. 

Skull measurements. — Old male (type): Basal length, 312; oc- 
cipito-nasal length, 310; palatal length, 169; zygomatic breadth, 207; 
interorbital breadth, 79. 

URSUS HENSHAWI * Merriam. 

Henshaw Grizzly. 

Ursus henshawi Merriam, Proc. Biol. Soc. Washington, XXVII, p. 190, August 
13, 1914. 

Type locality. — Southern Sierra Nevada, near Havilah, Kern 
County, California. 

Type specimen. — No. 15671, $ old, U. S. National Museum. Col- 
lected in 1875 by Dr. J. T. Rothrock and Henry W. Henshaw. 

Range. — Lower slopes of southern part of Sierra Nevada; limits 
unknown. 

Characters. — Size rather small — by far the smallest of the Cali- 
fornia grizzlies; size and general cranial characters as in horriceus, but 
fronto-nasal region strongly dished and rostrum strongly depressed. 
Last upper molar short and broad, the heel short and subtriangular. 
Skin characters unknown. 

Cranial characters. — Old male (type) : Skull long, narrow, and 
rather low; zygomata subtriangular, narrowly spreading; frontal 
shield gently sloping, flat-concave ; postorbital processes massive, and 
somewhat arched ; orbital rims swollen ; fronto-nasal region strongly 
depressed and dished ; rostrum low, depressed. 

Cranial comparisons. — Compared with an equally old male hor- 
riceus (type, from Coppermines, New Mexico) : Surprisingly similar 
in general, with similar broad outstanding postorbitals, but rostrum 
strongly depressed ; nasals flattened and horizontal anteriorly ; fronto- 
nasal region concave and strongly dished (in horriceus rather high 
and not dished) ; frontal shield strongly and broadly concave between 
orbits and between postorbital processes (only faintly depressed medi- 

* Named for Henry W. Henshaw, formerly chief of the Biological Survey. 



88 NORTH AMEBIOAN FAUNA. [No. 41. 

ally in horriceus) ; more strongly sloping anteriorly; postorbital pro- 
cesses shorter and blunter; orbital rims more prominent, relatively 
thin, somewhat everted, continuing to lachrymal notch — their promi- 
nence anteriorly due in part to presence of a broad sulcus in ascending 
arm of maxillary immediately in front of orbit ; lachrymal duct open- 
ing in orbit posterior to orbital rim (in horriceus on or anterior to 
rim) ; anterior nares broader than high (contrary true in horriceus) ; 
lambdoid crest higher ; sagittal crest and inion much shorter ; occipi- 
tal overhang much less; palate decidedly broader; interpterygoid 
canal shorter; mastoid processes much longer and more divergent; 
anterior part of pterygoids more broadly expanded vertically and 
articulating with a like expansjxm of posterior arms of palatines 
(probably not constant). Upper molars decidedly broader (canines 
broken off). 

Compared with two old males of tularensis (type, No. 3536, and 
No. 3537, from Fort Tejon, California) : Size smaller; occipito- 
sphenoid length and frontal breadth essentially the same; vault of 
cranium and rostrum very much lower; fronto-nasal region more 
deeply concave, more strongly sloping, and strikingly more dished; 
rostrum smaller, lower, and strongly depressed instead of elevated; 
occipital overhang and development of inion conspicuously less; 
posterior part of sagittal crest not elevated or produced. Under- 
jaw decidedly shorter, ramus much less broadly flattened vertically 
and more strongly bellied posteriorly. Last upper molar much 
smaller, the heel conspicuously shorter and narrowed on outer side. 

Skull measurements. — Old male (type): Basal length, 318; oc- 
cipito-nasal length, 305 ; palatal length, 173 ; zygomatic breadth, 204 ; 
interorbital breadth, 76. 

Stikeenensis Group. • 

URSUS STIKEENENSIS Merbiam. 

Stikine Gbizzlt. 

(PI. IX.) 

Ursus stikeenensis Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 178-179, 
August 13, 1914. 

Type locality. — Tatletuey Lake, tributary to Finlay River, near 
head of Skeena River, northern British Columbia. 

Type specimen. — No. 202794, & ad., U. S. National Museum, 
Biological Survey collection. Collected September 23, 1913, by 
Charles R. Cross, jr., and Edward A. Preble ; original No. 5772. 

Range. — Region about head of Finlay River, and Dease Lake 
region, northern British Columbia, and northerly in Yukon. 



1918.] STTKEENENSIS GROUP. 89 

Characters. — Adult male (type) : Size medium ; skull short, broad, 
and highly arched; face strongly pugged from abrupt rising of 
frontal region; claws short and strongly curved for a grizzly (longest 
60 mm. — tips worn by digging), dark, marked with yellowish on 
tips and sides. Upper molars large. Total length before skinning 
1,830 mm. ; hind foot 267 mm. ; estimated height at shoulder 990 mm. 
(39 inches). 

Color. — Type specimen: General ground color dark brown, griz- 
zled and washed with pale-tipped hairs. Muzzle pale brown, becom- 
ing darker between eyes and on sides of face; top of head in front 
of ears washed with yellowish brown, almost forming a golden 
brown band between the darker ungrizzled frontal region and the 
more whitish color of the neck; general ground color of body very 
dark, bountifully overlaid on neck and back by pale buffy-tipped 
hairs which give a whitish cast to the neck ; ears dark ; legs and feet 
blackish ; underparts dark brown ; hairs on sides of throat long and 
grizzly. 

Cranial characters. — Adult male (type): Size medium or large; 
skull short posteriorly, broad, highly arched, strongly dished, with 
abruptly ascending frontals and large molar teeth. Rostrum short, 
broad, broadening and strongly ascending posteriorly; nasals 
strongly upturned posteriorly; frontal shield rather broad, rising 
abruptly and swollen in front of and above orbits, sulcate medially, 
short pointed, the point ending in sagittal crest about 30 mm. ante- 
rior to fronto-parietal suture; postorbital processes large, broad, 
subtriangular, and decurved (more broadly rounded and more out- 
standing in older skull) ; braincase long for size of skull; zygomata 
rather broadly spreading, subtriangular, not much expanded verti- 
cally; palate and postpalatal shelf broad. Underjaw and inferior 
border of ramus very short. Molars large and broad. 

Cranial comparisons. — Adult male compared with adult male 
absarokus : Size nearly the same though absarokus appears the larger ; 
basal length and frontal breadth approximately the same, but oc- 
cipito-nasal length much greater in absarokus. In stikeenensis ros- 
trum lower, flatter, and more nearly horizontal ; frontal shield and pos- 
terior part of nasals rising much more abruptly ; frontals much more 
swollen in front of upper part of orbits ; point of shield much shorter 
(ending midway between plane of postorbitals and fronto-parietal 
suture, while in absarokus it reaches posteriorly to suture) ; inion 
short; palate broader; underjaw shorter; coronoid lower. 

Adult male compared with adult male tahltanicus (both inhabit- 
ing the same region) : Basal length, zygomatic breadth, and frontal 
breadth essentially the same, but frontal region much higher and 
rising abruptly at orbits instead of sloping gently in plane of ros- 
trum; rostrum broader, flatter, and more nearly horizontal (appear- 



90 NORTH AMERICAN FAUNA. [No. 41. 

ing shorter) ; postorbitals much larger and less horizontally outstand- 
ing; palate longer; under jaw longer. Dentition heavier. 

Skull measurements. — Adult male (type) : Basal length, 321; occi- 
pito-nasal length, 305; palatal length, 171; zygomatic breadth, 217; 
interorbital breadth, 84. 

URSUS CRASSODON sp. kov. 
Big-Tooth Grizzly. 

Type No. 171049, $ old, U. S. National Museum, Biological Sur- 
vey collection. Collected on Klappan Creek (=Third South Fork 
Stikine Eiver), September, 1907, by Dr. E. P. Richardson, of Boston. 

Cranial characters. — Old maZe/( type) : Frontal shield broad, lyrate 
posteriorly, the point reaching or slightly passing fronto-parietal 
suture; shield rising abruptly from rostrum, sulcate anteriorly but 
nearly flat between postorbitals; postorbitals broadly rounded and 
horizontally outstanding; rostrum of medium breadth, nearly hori- 
zontal ; braincase short ; sinus case rising to support postorbital part 
of shield; zygomata moderately outstanding, subtriangular, the squa- 
mosal root not broadly expanded; squamosal shelf narrow; palate 
and postpalatal shelf moderate; ramus short; coronoid blade rather 
small, moderately recurved at apex; canines rather large; molars 
enormous for size of skull ; MA and M 2 - very broad ; heel of M-2- sub- 
triangular, narrowing posteriorly. 

I refer to this species two young males from Yukon, one (No. 
209896) from White River, the other (No. 1839, Ottawa Museum) 
from Wolf Lake near Teslin Lake, both in the third year; and an 
old female (No. 202792) from Tatletuey Lake on the upper Finlay. 
Besides these, two very old male skulls from southern Yukon (No. 
223760 from Glenlyon Mountains and No. 223767 from Quiet Lake 
at head of Big Salmon River) are provisionally referred to the 
species, but not without considerable hesitation. Both are high, 
short, and broad as in crassodon, but the teeth are too much worn to 
admit of reliable comparison. One (No. 223760) is so extremely 
aged that the roots of the canines have worked down and become 
absorbed basally, leaving the middle part of the rostrum narrower 
than normal. 

Skull measurements. — Old male (type) : Basal length, 320 ; * oc- 
cipito-nasal length, 306 ; 1 palatal length, 175 ; zygomatic breadth, 
222 ; interorbital breadth, 84. 

URSUS CRASSUS sp. nov. 1 
Thickset Grizzly. 
Type No. 225473, $ ad., U. S. National Museum, Biological Sur- 
vey collection. From upper Macmillan River, Yukon. Collected in 
September, 1916, by William Drury. 

1 Partly restored. 

* Tentatively included in stikeenensis group. (See Introduction, pp. 12-13.) 



1918.] STTKEENENSIS GROUP. 91 

Characters. — Size large; hump apparent but not conspicuous; 
general color dark; claws of medium length, narrow, moderately 
curved, smooth, whitish above, with dark sides. 

Color. — Muzzle pale brownish; top of head and neck strongly 
washed with yellowish or golden buffy; shoulders and back lightly 
tipped with same on dark background ; legs and feet dusky. 

Cranial characters. — Adult male (type) : Skull rather large, short, 
broad, rather high, but not highly arched, and unusually massive. 
Shield broad, convex, faintly sulcate, sloping gradually into rostrum, 
the point reaching fronto-parietal suture; postorbitals rather broad, 
decurved in convexity of shield; rostrum large and high, rising in 
plane of shield; sagittal crest short; zygomata moderate; palate nar- 
row and strongly troughed; postpalatal shelf short; underjaw rather 
short, massive ; coronoid blade high and nearly vertical ; subangular 
border short. Teeth large; canines massive; M a very long, its heel 
exceptionally long and flat, slightly emarginate and everted. 

Cranial comparisons. — The skull of the adult male (type) viewed 
from above bears a striking likeness to that of adult male hoots, but 
when turned over the likeness ceases. It agrees with hoots essentially 
in size, massiveness, arching of vault of cranium, and frontal breadth ; 
but frontal shield is more convex; postorbitals more decurved; ros- 
trum somewhat more ascending (higher posteriorly) ; orbital rims 
less swollen ; palate decidedly narrower and troughed instead of flat ; 
coronoid more nearly vertical; subangular border shorter; teeth 
strikingly larger throughout — incisors, canines, and molars. The 
teeth of hoots are small for size of skull, those of crassus exception- 
ally large. 

Male adult (type) compared with male adult crassodon (type) : 
Size slightly greater; frontal shield much broader, more gently slop- 
ing, and more convex transversely, with decurved postorbitals (in 
crassodon shield flattish, nearly horizontal, with horizontally out- 
standing postorbitals and descending abruptly to rostrum) ; rostrum 
very much larger and higher and sloping gradually into shield; 
ramus much longer ; subangular border shorter ; coronoid larger and 
higher. Upper canines about same; lower canines much larger; 
upper and lower molar series of about same length but upper molars 
much narrower and less massive. 

Remarks. — The skull of an immature male (No. 6552) collected by 
R. MacFarlane, May 1, 1863, on Anderson River, 50 miles southeast 
of old Fort Anderson, has very large teeth, especially M- 2 -, thus dif- 
fering widely from any adult Barren Ground bear thus far examined. 
Two still younger skulls collected on the Barren Grounds in 1911, 
by Dr. R. M. Anderson (No. 34411 Amer. Mus. Nat. Hist, from Hor- 
ton River, and 34413 from Langton Bay), also have the crown of M* 
very long. These three skulls I provisionally refer to crassus. 



92 NORTH AMERICAN FAUNA. [No. 41. 

It is a singular fact, in view of the wide dissimilarity of the skulls, 
that the teeth of male crassus and male kluane are very much alike. 

Skull measurements. — Adult male (type): Basal length, 325; 
occipito-nasal length, 322; palatal length, 171; zygomatic breadth, 
224; interorbital breadth, 94. 

URSUS MIRABILIS Merriam.* 
Strange Grizzly. 

Ursus mirabilis Merriam, Proc. Biol. Soc. Washington, XXIX, p. 146, September 
6, 1916. 

Type locality. — Admiralty Island, Alaska. 

Type specimen. — No. 137471, ^ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected June 26, 1905, by Cyrus Catt. 

Characters. — A true grizzly, of medium size, related to stikeenensis 
of the mainland, and having the same high bulging forehead; exter- 
nal characters unknown. 

Cranial characters. — Adult male (type) : Skull of medium size, 
short, with rather broadly spreading zygomata and highly arched 
(almost domed) frontal region. Frontal shield of moderate breadth, 
short pointed posteriorly, rising abruptly from rostrum, convex both 
antero-posteriorly and transversely, slightly sulcate medially, mod- 
erately swollen on each side of sulcus ; postorbital processes moderate, 
broader than peglike, slightly decurved (continuing convexity of 
frontals) ; fronto-nasal region strongly dished; rostrum high, nar- 
row, strongly sloping ; zygomata rather broadly outstanding, slightly 
bowed; palate and postpalatal shelf rather broad; under jaw short; 
ramus bellied posteriorly; coronoid blade high and rather vertical, 
the apex not reaching plane of condyle. Upper canines rather long; 
molars rather broad and short. 

Cranial comparisons. — Ursus mirabilis requires comparison with 
only a single species — stikeenensis of the neighboring mainland : Size 
smaller ; frontal shield narrower and more bulging anteriorly, rising 
more abruptly from rostrum; rostrum narrower, materially higher, 
and more sloping; zygomata more widely outstanding; occipital 
overhang more pronounced; palate and postpalatal shelf similar; 
underjaw less massive; coronoid blade narrower and higher; teeth 
similar, but heel of last upper molar much shorter. 

Skull measurements. — Adult male (type): Basal length, 308; 
occipito-nasal length, 310; palatal length, 168; zygomatic breadth, 
230; interorbital breadth, 81. 

* Tentatively included in stikeenensis group. (See Introduction, pp. 12-13.) 



1918.] STTKEENENSIS GROUP. 93 

URSUS ABSAROKUS Merbiam. 1 

Absaroka Grizzly. 

Ursus absarokus Merriam, Proc. Biol. Soc. Washington, XXVII, p 181, August 
13, 1914. 

Type locality. — Near head of Little Bighorn River, northern part 
of Bighorn Mountains, Montana. 

Type specimen. — No. 67391, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected in May, 1893; purchased for 
Biological Survey by J. Alden Loring. 

Range. — Laramie and Bighorn Mountains, eastern Wyoming, 
Black Hills region, South Dakota, and northward along Little Mis- 
souri to Missouri and Yellowstone Rivers. 

Characters. — Size large, but much less than horribilis and with 
much smaller molars — especially M^. 

Color. — Head of young-adult (No. 203524) killed by Howard 
Eaton on the Little Missouri at mouth of Bear Creek, near Middle or 
" Bullion " Butte, October 27, 1880 : Muzzle pale brown, changing 
to grizzled dark brown on head and face; a large patch of dark 
brown free from grizzling on side of face extending from eye to 
angle of jaw ; chin and gular region dark brown (except anterior part 
of chin, which has not yet molted the pale old coat) ; top and sides 
of neck and doubtless body also, strongly grizzled. 

Cranial characters. — Adult male (type specimen, and other males 
from Bighorn and Laramie Mountains) : Vault of cranium rather 
highly arched; zygomata moderately outstanding and slightly 
bowed; frontal shield rather broad, rising strongly from rostrum, 
convex both antero-posteriorly and transversely ; slightly swollen on 
each side of median line just behind plane of orbits, faintly de- 
pressed medially between orbits; postorbital processes large, out- 
standing, and slightly decurved, the tips bluntly rounded; sagittal 
crest nearly straight, reaching anteriorly to fronto-parietal suture; 
rostrum large and high, rising strongly into frontal shield; post- 
palatal shelf broad and flat; nasals large and long; top of coronoid 
high and broadly rounded, its apex short, not reaching posteriorly to 
plane of condyle. Teeth rather large, especially M- 2 - which is long 
and broad, the heel emarginate on outer side. 

Cranial comparisons. — Adult male (type) compared with adult 
male shoshone (type) : Basal length essentially the same but skull 
somewhat larger and more massive, broader, highest point more 

1 Tentatively included in stiheenensis group. (See Introduction, pp. 12-13.) 



94 NORTH AMERICAN FAUNA. [No. 41. 

anterior; frontal shield broader, domed (strongly convex both an- 
tero-posteriorly and transversely) and rising rather abruptly from 
plane of rostrum (in shoshone flat) ; point of shield much longer pos- 
teriorly, reaching to or nearly to fronto-parietal suture. Molars 
larger — especially MA 

Skull measurements. — Adult male (type): Basal length, 322; 
occipito-nasal length, 322; palatal length, 172; zygomatic breadth, 
218; interorbital breadth, 88. 

Alascensis Group. 

URSUS ALASCENSIS Merriam. 

/ 
Alaska Grizzly. 

(PI. XII.) 

Ursus horribilis alascensis Merriam, Proc. Biol. Soc. Washington, X, pp. 74-75, 
April 13, 1896. 

Type locality. — Unalaklik River, Alaska. 

Type specimen. 1 — No. 76466, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected in 1895, by the late Rudolf 
Neumann, of Iliuliuk, Alaska. 

Range. — Norton Sound region, Alaska (Unalaklik and Shaktolik 
Hills) southerly over the Nushagak and Kuskokwim Rivers to 
Chinitna on Cook Inlet. Limits unknown. 

Cranial characters. — Old male (type) : Size small; braincase broad 
anteriorly; frontal shield rather broad, flattish, very short pointed 
posteriorly with correspondingly elongate sagittal crest, moderately 
sulcate interorbitally, otherwise convex in cross section, rising rather 
strongly from facial plane; postorbitals rather small, moderately 
decurved; fronto-nasal region moderately dished; zygomata sub- 
triangular, not broadly outstanding; palate rather broad and con- 
cave; postpalatal shelf broad and short; notch rather broad; jaw 
rather long; inferior border of ramus long, moderately convex 
posteriorly ; coronoid blade moderate, rather high, the apex recurved. 
Teeth moderate; last upper molar broad and short, with short 
obliquely truncate heel ; first lower molar sinuous, a strong concavity 
on outer side. 

Adult female: 2 Long and high; vault of cranium well arched, 
highest about midway of frontals; frontal shield rather narrow, not 
flattened but arching high above facial plane; swollen between me- 

1 In describing this bear 20 years ago I neglected to designate a type, and the original 
material included skulls of more than one species. I take this opportunity therefore to 
fix the type of Ursus alascensis on one of the original specimens (No. 76466, U. S. Na- 
tional Museum, Biological Survey collection) and to redefine the species. 

2 In the absence of skulls of adult females from the type region it is assumed that 
females from the Nushagak, Kuskokwim, and Chinitna Rivers are fairly typical of 
alascensis. 



1918.] ALASCENSIS GROUP. 95 

dian sulcus and orbits; postorbital processes decurved; palate and 
postpalatal shelf long. Last upper molar with short heel. 

Cranial comparisons. — Old male (type) compared with old male 
tohlat (No. 158811, collected by Charles Sheldon at head of Toklat 
River near northern base of Mount McKinley) : Size and general 
appearance essentially the same, but vault of cranium less elevated ; 
frontal shield narrower and shorter (the point reaching about half- 
way from plane of postorbitals to parietals) ; sagittal crest longer ; 
zygomata much less broadly outstanding, less arched; underjaw de- 
cidedly longer; inferior border of ramus longer; coronoid decidedly 
higher ; heel of last upper molar very much shorter. 

Adult female compared with adult female toklat: Skull decidedly 
larger; vault of cranium much higher and more arched (in tohlat 
low and flat), highest about middle of frontals instead of at hinder 
end; rostrum larger (longer, broader, and higher) ; face more slop- 
ing (nasals less nearly horizontal) ; frontals much more swollen be- 
tween sulcus and orbits ; postorbital processes more decurved ; palate, 
postpalatal shelf, and occipito-sphenoid notably longer; last upper 
molar very much shorter. 

Adult male compared with adult male tundrensis of same region : 
Size smaller (basilar and occipito -nasal lengths fully an inch less) ; 
skull less massive; frontals narrower, rising more abruptly from 
facial plane and more swollen over orbits; rostrum more slender; 
palate less flattened ; ramus of jaw less swollen posteriorly ; coronoid 
less falcate; canines smaller, decidedly shorter, and more curved; 
molars decidedly smaller. 

Skull measurements — Old male (type) : Basal length, 310; oc- 
cipito-nasal length, 298.5; palatal length, 166; zygomatic breadth, 
206; interorbital breadth, 79. 

URSUS TOKLAT Meebiam. 

Toklat Gbizzly. 

Ursus toklat Merriam, Proc Biol. Soc. Washington, XXVII, pp. 182-183, 
August 18, 1914. 

Type locality. — Head of Toklat River, north base of Alaska Range, 
near Mount McKinley, Alaska. 

Type specimen. — No. 158813, 9 ad., U. S. National Museum, 
Biological Survey collection. Collected May 24, 1908, by Charles 
Sheldon, and by him presented to the Biological Survey; original 
No. 324. Mother of cub No. 158814 (original No. 325). 

Range. — So far as known, restricted to Alaska Range. 

Characters. — Size medium; skulls of both male and female rather 
highly arched and dished, that of female much smaller and lower 
than male. Last upper molar large; heel very long. Claws horny 
and smooth. 

64854°— 18 7 



96 NORTH AMERICAN FAUNA. [No. 41. 

Color. — Color variable, upperparts ranging from ordinary 
" grizzly bear color " to creamy white ; claws usually dark through- 
out, but in many cases more or less marked with white. 

Cranial characters. — Adult males: Size medium; rostrum high, 
subterete above, nearly horizontal; frontal shield abruptly elevated, 
convex or domed, swollen over orbits, sulcate medially, rather long 
pointed (reaching fronto-parietal suture) ; postorbital processes 
rather small and strongly decurved; squamosal shelves short; palate 
arched and excavated longitudinally; postpalatal shelf rather broad; 
zygomata very broadly spreading and angular (zygomatic breadth 
in proportion to basal length much greater than in any other true 
grizzly, about equaling that of sheldoni) ; sagittal crest short. Last 
upper molar large, its heel normally very long. Adult female: Skull 
rather long and narrow, with broadly spreading zygomata and 
strongly dished f ronto-nasal region ; frontal shield rather flat, sulcate 
between orbits, varying from lyre pointed to short pointed; post- 
orbital processes outstanding horizontally; rostrum rather slender, 
nearly horizontal. In most specimens the highest point of cranium 
culminates in a rather abrupt change of angle at or near the fronto- 
parietal suture, forming a sort of " hump," a condition usual also in 
skulls of female grizzlies from the upper Yukon and northern 
British Columbia. 

Cranial comparisons. — Old male compared with old male alascen- 
sis: Frontal region more elevated; sagittal crest shorter; zygomatic 
breadth much greater ; heel of last upper molar much longer. 

Adult female compared with female alascensis: Size smaller; 
vault of cranium decidedly lower, highest over posterior part of 
frontals instead of over middle of frontals; frontal shield narrow, 
flattened, sloping (not arched and not materially swollen over or- 
bits) ; postorbital processes outstanding horizontally; palate, post- 
palatal shelf, and occipito-sphenoid shorter; M> with heel much 
longer. 

Adult female compared with adult female phmonyx: Size de- 
cidedly greater; teeth much larger, especially the canines and last 
upper molars. 

Adult female compared with adult female kluane: Length much 
greater; facial part of skull about the same; braincase and post- 
palatal parts much longer; vault of cranium higher; sagittal crest 
longer and higher, reaching forward over posterior frontals; point 
of shield much shorter; underjaw and inferior border of ramus much 
longer ; teeth about the same size but M 1 usually smaller ; heel of M- 2 - 
longer. 

Skull measurements. — Old male (No. 158811, from head Toklat 
River, Alaska) : Basal length, 304; occipito-nasal length, 298; palatal 



1918.] AXASCENSIS GBOUP. 97 

length, 169.5 ; zygomatic breadth, 222 ; interorbital breadth, 80. Old 
female (type) : Basal length, 283; occipito-nasal length, 267; palatal 
length, 157; zygomatic breadth, 187; interorbital breadth, 73. 

URSUS LATIPRONS Mebeiam. 

Bboad-Fbonted Gbizzlt. 

Ursus phcBonyx latifrons Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 
183-184, August 13, 1914. 

Type locality. — Jasper House, Alberta. 

Type specimen. — No. 75612, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected September 15, 1895, by J. Alden 
Loring. Original No. 3270. 

Range. — Rocky Mountains of western Alberta and eastern British 
Columbia from Jasper House northwesterly to region between head- 
waters of Parsnip and Great Bend of Fraser River and thence to 
extreme headwaters of Stikine River; limits of range unknown. 

Characters. — Size medium or rather large. External characters 
unknown. Affinities with the geographically remote toklat. 

Cranial characters. — Adult male (type) : Size medium; base elon- 
gate posteriorly; vault of cranium rather low, only slightly arched, 
top flattish; frontal shield very broad (interorbitally 93 mm., be- 
tween tips of postorbitals 130), rising from rostrum rather abruptly, 
broadly but shallowly depressed medially, swollen over orbits, the 
point ending in sagittal crest some distance (apparently more than 
an inch) anterior to fronto-parietal suture; postorbital processes 
broad, outstanding, and only slightly depressed ; f ronto-nasal region 
slightly dished; rostrum rather high, nearly horizontal, rounded 
above; postpalatal shelf rather broad; notch rather narrow; zygo- 
mata strongly outbowed but not widely spreading; under jaw long; 
coronoid blade broad basally, rather low, its anterior border sloping 
strongly backward. Teeth too badly worn to admit of description. 
Adult female (No. 209378) collected by F. K. Vreeland, September 
17, 1915, near head of Big Salmon or North Fork Fraser River be- 
tween Big Bend of the Fraser and headwaters of the Parsnip : Simi- 
lar in general to that of male with the usual sexual difference in 
frontal shield, which is lyrate pointed posteriorly; frontal shield 
broad — broadly depressed medially between orbits, swollen over or- 
bits; postorbital processes strongly developed, outstanding, slightly 
decurved ; sagittal crest short, high posteriorly, covering about three- 
quarters of suture between parietals; postpalatal shelf rather broad 
and flat; mastoids short and appressed; postpalatal notch short, of 
moderate breadth; zygomata moderately outstanding, less bowed 
than in the male; coronoid blade broad basally, rather low, recurved 



98 NORTH AMERICAN FAUNA. [No. 41 

at apex; canines of medium size; molars large, especially last upper 
molar. 

Cranial comparisons. — Old male (type) compared with old male 
toklat (from type locality) : Frontal shield much broader and less 
elevated; braincase less arched; palate flatter (in toklat distinctly 
arched antero-posteriorly) ; zygomata more rounded basally and more 
outbowed (less angular) ; underjaw slightly longer; inferior border 
of ramus much longer ; coronoid blade broader and higher. The basal 
part of the skull is longer in latifrons than in toklat. This is apparent 
whether the skull is viewed from above or below. In latifrons, seen 
from above, the distance from occipital crest to postorbital process is 
materially greater ; while seen from bejow, the distance from occipital 
condyle to postpalatal notch is likewise greater. Another difference 
appears rather strikingly when the skull is viewed from below: in 
latifrons the squamosal base of the zygoma slopes gradually forward, 
while in toklat the curve is much shorter and more abrupt, so that 
the bases of the zygomata stand out much more squarely, practically 
at right angles to cranial axis. 

While skulls of adult male latifrons differ strikingly from those 
of male toklat, skulls of the adult females are surprisingly alike, 
agreeing in general appearance, basal length, breadth of palate, and 
large size of last upper molar, though in latifrons this tooth is ac- 
tually broader than in most specimens of toklat. The two agree 
essentially also in lower molars and canines. The underjaw in lati- 
frons, however, is shorter, and coronoid blade lower than in toklat. 

Old male compared with old male kluane: Frontal region much 
broader and flatter ; postorbital processes much less decurved ; sagit- 
tal crest shorter and straight instead of arched; rostrum less ele- 
vated and less narrowly rounded above; nasals flatter; zygomata 
more widely spreading and more strongly outbowed posteriorly; 
coronoid blade broader at base. 

Adult female compared with adult female kluane: Skull much 
larger, longer, and more massive; frontal shield irmch broader; 
vault of cranium flatter (less arched over posterior frontals) ; ros- 
trum higher; zygomata more widely outstanding; palate much 
broader; last upper molar much larger: lower molars and canines 
approximately same size. While the skull of female latifrons is 
much larger than that of female kluane, the underjaw is only slightly 
larger. 

Adult female compared with adult female phwonyx: Size much 
larger; frontal region much broader; rostrum broader; nasals more 
nearly horizontal; arch of cranium more depressed ; underjaw longer; 
coronoid blade broader; molars much larger. (Comparison of males 
is unnecessary, the male of phmonyx having a large, broad, massive 
skull resembling that of daMi.) 



1918.] RICHARDSONI GROUP. 99 

Skull measurements. — Old male (type) : Basal length, 324; oceip- 
ito-nasal length, 312; palatal length, 173; zygomatic breadth, 214.5; 
interorbital breadth, 94. 

Richardsoni Group. 

URSUS RICHARDSONI Swainson. 

Babren Ground Beab. 

Vrsus Richardsoni Swainson, Animals in Menageries, pp. 54-56, 1838. 

Type locality. — Shore of Arctic Ocean, on west side of Bathurst 
Inlet near mouth of Hood River. 1 

Characters. — Size medium; color variable, from yellowish to 
grizzly brown ; f oreclaws of medium length, smooth ; skull medium 
or small, with broadly spreading zygomata. 

Cranial characters. — Adult males: Two adult or rather old male 
skulls collected by Dr. R. M. Anderson in 1915, and loaned me by the 
Museum of the Geological Survey of Canada, at Ottawa (one No. 
2774 from Dolphin and Union Straits, the other No. 2776 from 
near mouth of Coppermine River), are assumed to be typical: Size 
medium ; cranium high in relation to size, but not much arched ; basi- 
cranium flat; shield rising abruptly from rostrum, of medium 
breadth, broadly and strongly sulcate medially, swollen over orbits, 
short pointed; postorbital processes massive, outstanding, arched 
over orbits and slightly decurved; orbits nearly vertical; rostrum 
moderate, rather high and narrow, sloping into sulcus of shield; 
orbits prominent, rising well above fronto-nasal plane; zygomata 
moderate, somewhat outbowed; nares subrectangular, broader than 
high ; palate rather short ; postpalatal shelf moderate or rather broad ; 
squamosal shelf weak posteriorly; underjaw rather short; infra- 
angular border of ramus short; apex of coronoid only slightly re- 
curved, falling short of plane of condyle; angular process projecting 
beyond condyle. Teeth medium ; canines rather large ; M- 2 - with mod- 
erate heel, narrowing posteriorly ; cusps of posterior molars weak, those 
of inner side of M- 2 - nearly obsolete (in striking contrast with the 
highly developed cusps of both upper and lower molars of pellyensis) . 

An exceedingly old male in the National Museum (No. 6255), 
collected on Anderson River by R. MacFarlane, has the shield less 
swollen - over and in front of orbits, and even shorter pointed poste- 

1 Swainson's description was based on and largely quoted from Richardson's account of 
" an old and lean male, killed on the shores of the Arctic Sea on the 1st of August, 1821.'' 
(Article on the Barren Ground Bear, in Fauna Boreali-Americana, pp. 21-24, 1829.) And 
in Franklin's " Narrative of a Journey to the Shores of the Polar Sea," p. 373, 1823, 
under date of August 1, 1821, the party being at the mouth of Hood River on Bathurst 
Inlet, the killing of a lean male brown bear is chronicled in some detail. It appears 
therefore that the type locality of Vrsus richardsoni is Hood River, Bathurst Inlet. 



100 NORTH AMERICAN FAUNA. [No. 41. 

riorly; rostrum more depressed; sagittal crest longer and somewhat 
higher (but still not high) and somewhat humped over posterior 
frontals; zygomata more broadly outstanding. 

SkiiM measurements. — Old male (No. 6255) from Anderson River: 
Basal length, 311; occipito-nasal length, 286; palatal length, 163; 
zygomatic breadth, 227; interorbital breadth, 72.5. Old male (No. 
2774, Ottawa Museum) from Dolphin and Union Strait: Basal 
length, 316; occipito-nasal length, 290; palatal length, 170; zygo- 
matic breadth, 211; interorbital breadth, 79. Adult male (No. 2776 
Ottawa Museum) from near mouth Coppermine River: Basal length, 
299; occipito-nasal length, 288; palatal length, 163; zygomatic 

breadth, 217 ; interorbital breadth, 82.5. 

/ 

URSUS RUSSELLI * Mebbiam.* 
Mackenzie Delta Gbizzly. 

Ursus russelli Merriam, Proc. Biol. Soc. Washington, XXVII, p. 178, August 13, 
1914. 

Type locality. — West side Mackenzie River delta, Canada. 

Type specimen. — No. 21301, $ old, University of Iowa Museum. 
Collected June 28, 1894, by Frank Russell. Mounted skin with 
skull separate. 

Range. — Lower Mackenzie region ; limits unknown. 

Characters. — Size rather small. Color a curious pale drab-brown, 
somewhat darker on legs and feet; ears conspicuously hairy. Claws 
smooth, moderately to strongly curved; brownish horn color with 
paler (almost amber) tips. Teeth large; last upper molar of great 
size and peculiar glassy texture in type skull. Affinities uncertain. 

Cranial characters. — Adult male (type) : Skull of medium size, 
about equaling old male of alascensis; rather short; fairly broad 
across zygomata ; frontal shield sloping strongly upward, moderately 
sulcate, swollen over orbits, but orbits not everted; posterior point 
of shield rather short, ending about one-third the distance from 
fronto-parietal suture to postorbital processes; postorbital processes 
peglike, standing out nearly horizontally — not depressed as in 
alascensis; muzzle rather narrow and high; zygomata slender, the 
posterior roots not expanded vertically ; palate flat, not excavated or 
arched as in several species; underjaw massive, heavier under Mj 
and My than in alascensis; coronoid blade falcate but not nar- 
rowly so. 

Cranial comparisons. — From richardsoni, its neighbor on the east, 
with which it agrees in size and in certain dental characters, it differs 
in much more highly vaulted cranium; more highly sloping (less 

1 Named for Frank Russell, who collected the type specimen. 

• Tentatively included In riohardaoni group. (See Introduction, pp. 12-18.) 



1918.] EICHAEDSONI GROUP. 101 

nearly horizontal) braincase; much more elevated frontal region, and 
very much narrower rostrum. The frontal shield is much longer than 
in richardsoni, the temporal impressions curving backward to meet 
one another about one-third the distance between postorbital processes 
and f ronto-parietal suture, instead of turning abruptly inward ; post- 
orbital processes more slender than in richardsoni/ sagittal crest 
shorter and less nearly horizontal; upper molars very much larger, 
particularly broader. Zygomatic arches bowed outward in both spe- 
cies — not sharply angular as in many large bears. 

Skull measurements. — Old male (type): Basal length, 310; oc- 
cipito-nasal length, 300 ; palatal length, 163 ; zygomatic breadth, 220 ; 
interorbital breadth, 79.5. 

URSUS PHJEONYX Mebeiam. 

Tanana Geizzly. 

Ursus Jwrribilis pkceonyx Merriam, Proc. Biol. Soc. Washington, XVII, p. 154. 
October 6, 1904. 

Type locality. — Glacier Mountain, Tanana Mountains, Alaska 
(about 2 miles below source of Comet Creek, near Fortymile Creek, 
between Yukon and Tanana Rivers). 

Type specimen. — No. 133231, 2 ad., U. S. National Museum, Bio- 
logical Survey collection. Collected July 12, 1903, by W. H. Osgood. 

Range. — Tanana Mountains between Tanana and Yukon Rivers. 

Characters. — Size of male large; of female small (sexual dis- 
parity great, much greater than in datti). 

Color. — Upperparts varying from creamy or buffy to dark 
" grizzly color " ; underparts and muzzle pale brown ; legs very dark 
brown, varying to blackish brown; claws horny and smooth, usually 
dark but sometimes marked with whitish. Last upper molar of me- 
dium size or rather small. 

Cranial characters. — Old male (No. 201586, from Ketchumstock, 
assumed to be typical of phwonyx) : Size medium or rather large; 
skull broad and short, moderately arched and dished, with broadly 
outstanding zygomata and rather deeply sulcate strongly sloping 
frontal shield. Frontal shield of moderate breadth, strongly swollen 
over orbits and bases of postorbitals, short pointed posteriorly, the 
point ending midway between fronto-parietal suture and plane of 
postorbitals; postorbitals triangular, broad basally, convex poste- 
riorly, straight or concave anteriorly; rostrum moderate, nearly 
horizontal, sulcate- depressed on top; fronto-nasal region distinctly 
dished ; squamosal shelves broad and rather short, the outer margin 
arched and upturned; middle part of zygomata moderately ex- 
panded vertically; palate broad and short; postpalatal shelf broad; 
notch relatively narrow. Under jaw long for size of skull, massive; 



102 NORTH AMERICAN FAUNA. [No. 41. 

coronoid blade high, rather narrow, and rather vertical, the recurved 
apex not reaching plane of condyle. Teeth medium. 

Adult female (type): Size small; vault of cranium well arched 
posteriorly, highest point 25 to 30 mm. in front of fronto-parietal 
suture; braincase rather broad, not much constricted anteriorly, 
not compressed, and with no tendency to keel into anterior part of 
sagittal crest; frontal shield moderately flattened, shallowly sulcate 
medially and slightly swollen on each side of median depression, 
gently sloping into rostrum posteriorly, the lyrate point reaching 
nearly to parietals ; rostrum rather small ; zygomata moderately out- 
standing; palate medium; postpalatal shelf rather broad and short. 
Under jaw long; inferior border of ramus long and straight; coro- 
noids rather small, the apex strongly recurved, cutting plane of 
condyle. 

Cranial comparisons. — Old female (type) compared with adult 
female kluane: Size somewhat larger; braincase broader anteriorly; 
frontal shield broader; palate and postpalatal shelf broader; oc- 
cipito-sphenoid and postpalatal lengths much greater. Under jaw 
much longer ; inferior border of ramus longer and straighter. Teeth 
too badly worn to admit of description but apparently about same 
size as in kluane. 

Old female (type) compared with adult female toklat from Alaska 
Range, near Mount McKinley: Basal length somewhat less; brain- 
case less contracted and much broader anteriorly (not compressed 
and with no tendency to keel into anterior part of sagittal crest) ; 
frontal shield less deeply sulcate anteriorly and much longer pointed 
posteriorly; fronto-nasal region less depressed; under jaw shorter; 
ramus straighter and lighter; coronoid much smaller, narrower, and 
lower; upper canines apparently about the same; last upper molar 
smaller ; lower canines more slender ; lower molars apparently about 
the same, but so completely worn off in type specimen that compari- 
son is impossible. 

Remarks. — The skull of the adult male phmonyx (if the sexes are 
correctly mated) is large, broad, and rather massive, requiring com- 
parison with only a single known species, Ursus dalli. The female 
on the other hand is small and resembles in a general way the 
females of the still smaller kluane, pulchellus, and pallasi. 

Skull measurements. — Old male (No. 205186, from Ketchumstock, 
Alaska) : Basal length, 327; occipito-nasal length, 309 ; palatal length, 
176; zygomatic breadth, 242; interorbital breadth, 85. Old female 
(type) : Basal length, 280; occipito-nasal length, 267; palatal length, 
148; zygomatic breadth, 189; interorbital breadth, 68. 



1918.] RICHARDSONI GROUP. 103 

URSUS INTERNATIONALIS Meeeiam. 

AXASKA BOUNDAEY GbIZZLT. 

Urstts internationalis Merriam, Proc. Biol. Soc. Washington, XXVII, pp. 177- 
178, August 13, 1914. 

Type locality. — Alaska- Yukon Boundary, about 50 miles south of 
Arctic coast (lat. 69° 00' 30"). 

Type specimen. — No. 1763, $ ad., Ottawa Museum. Killed July 3, 
1912, by Frederick Lambart, of Canadian Boundary Survey. 

Range. — Eegion bordering Arctic coast along international bound- 
ary, and doubtless adjacent mountains, between the coast and the 
Yukon-Porcupine; limits unknown. 

Characters. — Size medium or rather large; affinities doubtful. 
Color a peculiar pale yellowish brown. Head strongly arched ; muz- 
zle and frontal region broad. Large lower premolar strictly conical, 
without heel, as in the brown bears. 

Cranial characters. — Skull of medium size, massive, strongly 
arched and dished, highest over anterior part of braincase; frontal 
shield broad, very short pointed posteriorly, sulcate medially and 
swollen over orbits ; postorbitals bluntly rounded, strongly decurved, 
not widely projecting; fronto-nasal region strongly dished; rostrum 
large and broad; sagittal crest long but feebly developed; zygomata 
subtriangular, not widely outstanding, and not much expanded ver- 
tically; palate and postpalatal shelf rather broad; notch moderate. 
Teeth rather small for size of skull; heel of last upper molar small 
and obliquely truncate on outer side; large lower premolar strictly 
of brown-bear type — a single cone without heel, sulcus, or posterior 
cusplets; first lower molar broad and somewhat sinuous; middle 
lower molar narrow and short posteriorly. 

Skull measurements. — Adult male (type) : Basal length, 309; oc- 
cipito-nasal length, 293; palatal length, 169; zygomatic breadth, 
203.5; interorbital breadth, 82. 

URSUS OPHRUS 1 Meebiam. 

High-Bbow Grizzly. 
(Pl.X.) 

Ur8U8 ophrus Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 148-149, Sep- 
tember 6, 1916. 

Type locality. — Eastern British Columbia (exact locality un- 
known). 

Type specimen. — No. 210252, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected in 1915, by E. W. Darbey. 

1 Ophrus, with reference to the unusual brows. 



104 NORTH AMERICAN FAUNA. [Mo. 41. 

Cranial characters. — Skull short, strongly dished, remarkably high, 
the deeply sulcate frontal shield rising abruptly high over orbits, 
with thickened brows and large outstanding arched postorbital proc- 
esses. Frontal shield of moderate breadth ; deeply and broadly con- 
cave between orbits, swollen over orbits and passing out into strongly 
outstanding postorbitals, short pointed posteriorly; fronto-nasal 
region deeply sulcate; middle part of nasals flat; sagittal crest high 
and reaching anteriorly nearly midway from fronto-parietal suture 
to plane of postorbitals; rostrum rather small and narrow; palate 
rather narrow; postpalatal shelf rather broad; zygomata broadly 
and strongly outbowed; mastoids rather long. Under jaw long, its 
inferior margin rather long and nearly straight ; subangular tubercle 
considerably posterior to inferior dental foramen; coronoid blade 
broad, its apex only moderately recurved, ending anterior to plane 
of condyle; canines of medium size, the lower ones rather massive; 
molars of medium size, the upper rather small for size of skull. 

Skull measurements. — Old male (type) : Basal length, 323 ; l oc- 
cipito-nasal length, 304; palatal length, 175; zygomatic breadth, 229; 
interorbital breadth, 85. 

URSUS WASHAKE Merbiam. 

Washakie Grizzly. 

Ursus ivashake Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 152-154, 
September 6, 1916. 

Type locality. — North Fork Shoshone River, Absaroka Mountains, 
western Wyoming (between Bighorn Basin and Yellowstone National 
Park). 

Type specimen. — No. 213005, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Killed September, 1913, by Col. J. A. 
McGuire. 

Cranial characters. — Old male (type) : Size medium, about equal- 
ing male shoshone and male horriwits; skull rather short and high, 
moderately arched, with broad, elevated postorbitals and rather 
broadly outbowed zygomata. Frontal shield rather narrow, sloping 
strongly upward anteriorly, highest at postorbital processes, hori- 
zontal posteriorly, broadly concave between postorbital > processes ; 
postorbital processes large, broad, subtriangular as viewed from 
above, outstanding, elevated and slightly arched, rising well above 
frontal plane and passing anteriorly into thickened orbital rims; 
fronto-nasal region dished (change of angle about middle of nasals) ; 
rostrum rather small, strongly compressed horizontally between 
nasals and roots of canines, making nasals appear elevated ; anterior 
nares small; zygomata rather slender, broadly spreading, rounded 

1 Restored. 



1918.] RICHARDSONI GROUP. 105 

and strongly outbowed posteriorly, only slightly expanded verti- 
cally; sagittal crest low; postpalatal shelf broad, flat, and rather 
short; occipito-sphenoid 87 mm. (= distance from front of canine 
to or slightly beyond middle of M 1 ). Underjaw moderate; ramus 
bellied posteriorly; coronoid blade high and rather falcate, the apex 
cutting plane of condyle (line from apex to tip of angular process 
passing well behind condyle). Teeth moderate or rather large; 
M^ large. 

Cranial comparisons. — Old male (type) compared with adult male 
ophrus (type) : Size about the same, but appearing smaller; frontal 
shield less elevated and less deeply concave; fronto-nasal region 
elevated instead of sulcate-dished ; zygomata less widely outbowed; 
postorbitals much broader and less elevated; orbital rims less 
swollen; postpalatal shelf shorter and broader; mastoids shorter; 
nares smaller and lower. Underjaw more massive; inferior border 
of ramus more swollen and much more bellied posteriorly ; coronoid 
blade higher and more falcate, the apex reaching much farther pos- 
teriorly (cutting plane of hinder part of condyle) . 

Compared with adult male canadensis (type), to which it is not 
related but with which it agrees essentially in basal and occipito- 
sphenoid length : Frontal shield less flat, more elevated laterally, 
highest at postorbitals instead of at posterior point; fronto-nasal 
region more dished ; rostrum smaller, narrower basally, more strongly 
compressed below nasals; postorbital processes very much larger, 
broader, and more massive, elevated, arched, and subtriangular, 
instead of slender and narrowly peglike; zygomata more outbowed 
and arched; sagittal crest low and straight instead of high and 
arched; inion less developed; braincase anteriorly broader and more 
depressed — not tending to " keel " into sagittal crest as in canadensis; 
occipito-nasal length less, although basal length of skull is essentially 
the same in both. Underjaw longer; inferior border of ramus 
shorter and more strongly bellied; coronoid blade higher and more 
falcate, its apex reaching farther posteriorly; distance from angle 
to subangular process much greater; diastema in both jaws much 
longer. Last upper and middle lower molars not quite so broad. 

Compared with adult male absarokus (type) : Size smaller; vault 
of cranium decidedly lower ; braincase anteriorly broadly depressed ; 
frontal shield narrower, lower, and flatter, concave instead of convex 
between postorbital processes; postorbital processes (viewed from 
above) broadly triangular, uplifted, and somewhat arched instead 
of pegshape; orbital rims more thickened and elevated; rostrum 
smaller, lower, more slender, and much more compressed horizontally 
between nasals and roots of canines, making the nasals appear 
elevated; sagittal crest lower; occipito-sphenoid shorter; occiput 
lower; anterior nares smaller. Underjaw shorter; inferior border of 



106 NORTH AMERICAN FAUNA. [No. 41 . 

ramus much shorter; coronoid blade more falcate, its apex reaching 
farther posteriorly; angular process more slender and more pro- 
duced posteriorly (line connecting apex of coronoid with angle pass-)! 
ing well behind condyle — in dbsarohus cutting condyle near middle).! 
Molars smaller. 

Skull measurements. — Adult male (type): Basal length, 310; oc- 
cipito-nasal length, 305; palatal length, 170; zygomatic breadth, 217; 
interorbital breadth, 76. 

Xidderi Group. 
URSUS KIDDERI KIDDERI 1 Merbiam. 

Kidder Beab. 

(PI. VIII.) 

Ursus kidderi Merriam, Proc. Biol. Soc. Washington, XV, p. 78, March 22, 1902. 

Type locality. — Chinitna Bay, Cook Inlet, Alaska. 

Type specimen. — No. 1L6562, $ young (not fully grown), U. S. 
National Museum, Biological Survey collection. Collected June 9, 
1901, by James H. Kidder. 

Range. — Alaska Peninsula for its entire length. 

Characters. — Size medium — small compared with gyas of the 
same , region ; sexual disparity small, female nearly as large as 
male; skull of adult male hardly half the bulk of male gyas; 
skull of adult female nearly the same size as that of female gyas; 
color yellowish brown ; claws rather short, blue-black. 

Color. — June specimens (in left-over winter pelage) : General 
color yellowish brown, darkest on belly and legs, legs much darker 
than body. Most of the Kidder bears in the National Zoological 
Park are pale buffy yellowish, or yellowish cream color. 

Cranial characters. — Adult moJe: Skull long, rather low, narrow, 
and massive; frontal shield narrow, moderately sloping; swollen 
over orbits and rather deeply sulcate or troughed medially; fronto- 
nasal region slightly dished; rostrum and nasals high and rather 
long; postorbital processes peglike, outstanding; braincase elon- 
gate; zygomata angular, only moderately spreading; sagittal crest 
long; palate long; mastoids long; underjaw long, with long ramus 
and moderate or low coronoid blade. Adult female: Skull in general 
like that of male but slightly smaller and with more slender rostrum 
and slightly smaller teeth. The sagittal crest extends much farther 
forward than in the females of most species, in this respect also 
resembling the male. The sexual difference in size of teeth appears 
to be covered by individual variation. 

1 Named for James H. Kidder, who collected and presented the type specimen. 



1918.1 KIDDERI GEOUP. 107 

Cranial comparisons. — Adult male and female compared with adult 
female gyas : Adult males require no comparison, owing to the great dif- 
ference in size of skull and teeth, but with the females the case is very 
different, the size being essentially the same. In fully adult females 
\kidderi is easily distinguished by the lowness of the vault of the 
! cranium and greater length of the sagittal crest — female gyas being 
rather highly domed and having the short sagittal crest of most 
female bears. 

Males of Mdderi are sometimes hard to tell from females of gyas, 
but in the case of fully adult skulls they may be distinguished as 
follows: Male kidderi averages longer, both in basal and occipito- 
nasal length, is much less highly arched, and is more obliquely trun- 
cate posteriorly so that the occiput overhangs, giving the effect of a 
longer braincase. The rostrum also is somewhat longer. The 
frontal shield is quite different, being very much shorter posteriorly, 
ending about an inch in front of the parietals, whereas in female 
gyas the posterior point of the shield extends posteriorly to about the 
same distance behind the fronto-parietal suture. Thus in adult male 
kidderi the frontal shield is horizontal or slopes forward from its 
most posterior point, while in adult female gyas it slopes backward 
for some distance — from a point at least an inch in front of the 
fronto-parietal suture — the shield thus overreaching the highest 
point of the arch of the skull and sloping downward in both direc- 
tions. The occipito-sphenoid length is slightly greater in female 
gyas than in male kidderi. 

Comparison of cubs of the second, third, and fourth years: Cubs of 
kidderi after the molars are in position and the permanent canines 
partly exposed, are easily told from those of male gyas of corre- 
sponding age by the relatively small size of the teeth, especially the 
canines, molars, and outer upper incisors. It is not so easy, however, 
to tell them from female gyas, and in some cases it may be impossible. 
The most constant character appears to be the length of the canines, 
and this can not be determined in cubs less than three or three and a 
half years of age. The canines are slightly longer in male kidderi 
than in female gyas, and their diameter also is usually, but not 
always, greater. As a rule, also, the crowns of the molars are larger, 
especially longer, in male kidderi the combined length of the first 
and second lower molars averaging about 4 mm. greater than in 
female gyas. As the teeth become worn with use the differences 
become less and less obvious, so that after the third year the distinc- 
tions are not easy of recognition. 

Skull measurements. — Average of 3 males from Belkofski, Alaska 
Peninsula: Basal length, 330; occipito-nasal length, 312; palatal 
length, 177; zygomatic breadth, 207; interorbital breadth, 74. 



108 NORTH AMERICAN FAUNA. [No. 41. 

Average of 4 females from Pavlof Bay, Alaska Peninsula: Basal 
length, 330; occipito-nasal length, 312; palatal length, 181.5; zygo- 
matic breadth, 218 ; interorbital breadth, 80.5. 

URSUS KIDDERI TUNDRENSIS Mebbiam. 

TtJNDBA BEAB. 

Ursus kidderi tundrensis Merriam, Proc. Biol. Soc. Washington, XXVII, p. 196, 
August 13, 1914. 

Type locality. — Shaktolik Eiver, Norton Sound, Alaska. 

Type specimen. — No. 76470, $ ad., U. S. National Museum, Bio- 
logical Survey collection. Collected by natives, September, 1894, 
and secured through the late Rudolf Neumann, then of Iliuliuk, 
Unalaska. 

Range. — Tundra region of northwestern Alaska from Shaktolik 
River on Norton Sound, southerly across the lower Yukon, Kus- 
kokwim, and Nushagak Rivers to Bristol Bay and north side of 
base of Alaska Peninsula. 

Characters. — Size medium (small in contrast with gyas), about 
equaling kidderi. External characters unknown, but doubtless little 
different from kidderi. Known to the natives as " Red Bear." 

Cranial characters. — Size medium, about as in kidderi; skull rather 
long and heavy; frontals broad and flat, broadly but not deeply 
sulcate medially; postorbital processes small; coronoid blade falcate 
and rather high. 

Cranial comparisons. — Similar to kidderi in essential cranial and 
dental characters, but differing in having the frontal shield and post- 
orbital processes very much broader and flatter, and the frontal sulcus 
less marked. As in kidderi there is little difference in the sexes 
except that the females have narrower muzzles and narrower frontal 
shields. A young male (No. 16375) from Andreafski on the lower 
Yukon, collected by E. W. Nelson, has somewhat shorter canines 
than the others. 

From alascensis, the grizzly of the same general region, skulls of 
tundrensis (adult males in both cases) may be distinguished by the 
foll®wing characters: Size larger (basilar and occipito-nasal lengths 
fully an inch greater) ; skull as a whole much more massive ; frontals 
broader, rising less abruptly from facial plane, less swollen over 
orbits; rostrum less slender; palate more flat; ramus of jaw thicker 
under M T and M T ; coronoid blade more falcate; canines larger, 
decidedly longer, and somewhat less curved ; molars decidedly larger. 

Skull measurements. — Adult male (type) : Basal length, 333 ; 
occipito-nasal length, 317; palatal length, 178; zygomatic breadth, 
228; interorbital breadth, 93. 



1918.] KIDDERI GROUP. 109 

URSUS EXIMIUS Merkiam. 

Knik Beab. 

Ursus eximius Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 139-140, Sep- 
tember 6, 1916. 

Type locality. — Head of Knik Arm, Cook Inlet, Alaska. 

Type specimen. — No. 122495, $ ad., U. S. National Museum. Col- 
lected by G. W. Palmer. 

Characters. — Size rather large ; color uniform rich dark brown sug- 
gesting seal brown; muzzle brown, paler than rest of head; back of 
head and neck lightly sprinkled with pale-tipped hairs; claws of 
medium thickness, only slightly curved, decidedly short, probably 
from wear, smooth, very dark horn color, becoming paler on sides 
toward tip. Skull long and narrow, with narrow highly arched 
frontals. Related to kidderi. 

Cranial characters. — Adult male (type) : Skull long, extremely 
narrow in fronto-nasal region, rather highly arched and strongly 
dished. Frontal shield exceedingly narrow, convex, shallowly sulcate 
medially, strongly arched anteriorly, horizontal posteriorly, long- 
pointed, the point nearly reaching f ronto-parietal suture ; postorbital 
processes slender, peglike, moderately outstanding; fronto-nasal 
region strongly dished; rostrum long, narrow, high, compressed be- 
tween nasals and canine roots; zygomata moderately outstanding, 
subtriangular ; palate long and narrow; postpalatal shelf relatively 
broad ; notch rather broad ; mastoids long, strongly divergent ; under- 
jaw long, moderately massive, the ramus broad vertically; coronoid 
blade rather broad, the apex not strongly recurved ; teeth of medium 
size ; M> with rather long heel, not much narrowed posteriorly ; M> 
relatively large and broad; PM4 a single cone without distinct heel 
but sulcate posteriorly. 

Adult female (No. 205176, from type locality) : Skull long and 
narrow ; vault of cranium moderately arched, the highest part form- 
ing a hump at f ronto-parietal suture ; frontal shield narrow, flattish, 
sulcate medially, the point reaching f ronto-parietal suture; postor- 
bitals weak, subtriangular, not decurved ; fronto-nasal region moder- 
ately dished ; rostrum narrow, compressed between nasals and canine 
roots; zygomata moderately spreading, subtriangular; postpalatal 
shelf relatively broad; notch moderate; inferior border of ramus 
convex from plane of front molar posteriorly ; coronoid blade broad 
and low. 

Cranial comparisons. — Ursus eximius appears to be related to only 
a single species, kidderi of Alaska Peninsula. Adult male (type) 
compared with a series of kidderi from various points on Alaska 
Peninsula: Size about the same; vault of cranium more highly 
arched; frontal shield narrower, more strongly convex in cross sec- 



110 NORTH AMERICAN FAUNA. INo. 41. 

tion, less deeply silicate; postpalatal processes more slender, peglike, 
and outstanding; fronto-nasal region more dished; rostrum more 
slender; nasals longer posteriorly, more completely wedge-shaped, 
reaching posteriorly to plane of postpalatal processes; mastoids 
longer and more divergent. 

Adult female (No. 205176, from head of Knik Arm) compared 
with adult female Mdderi: Size materially smaller; frontal shield 
and rostrum much narrower; vault of cranium notably higher over 
f ronto-parietal suture ; braincase narrower ; nasals longer posteriorly ; 
under jaw smaller and lighter. 

Adult male (type) compared with adult male alascensis (No. 
76466, from Unalaklik River, Norton Sound, Alaska) : Skull much 
longer, more highly arched, and narrower throughout. Frontal 
shield much more elevated, narrower, and longer posteriorly ; fronto- 
nasal region more strongly dished; rostrum narrower and higher; 
lambdoid crest more strongly developed ; palate and postpalatal shelf 
much longer; occipito-sphenoid much longer; mastoids much longer 
and strongly divergent; under jaw longer; coronoid blade much 
higher; teeth larger; heel of M^ much longer. 

Adult female compared with female alascensis: Length essentially 
the same; skull narrower throughout; frontal shield lower, much 
narrower and flatter, rising less abruptly from rostrum ; fronto-nasal 
region sulcate but less strongly dished; rostrum slightly more 
slender; postpalatal shelf narrower. Underjaw about same length; 
inferior border of ramus more evenly convex (less abruptly bellied) ; 
coronoid blade broader; canines about same size; molars somewhat 
larger. 

Skull measurements. — Adult male (type): Basal length, 331; 
occipito-nasal length, 319; palatal length, 185; zygomatic breadth, 
215; interorbital breadth, 71. 

Innuitus Group. 

URSUS INNUITUS Mebbiam. 

Innuit Beab. 

(PI. VII.) 

Ursus innuitus Merriam, Proc. Biol. Soc. Washington, XXVII, p. 177, August 13, 
1914. 

Type locality. — Golofnin Bay, south side of Seward Peninsula, 
northwestern Alaska. 

Type specimen. — No. 179780, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected in 1886, by Edward F. Ball. 

Range. — Coastal region of Norton Sound, Alaska, from Unalaklik 
northward and westward ; limits unknown. 



1918.] INNUITUS GROUP. Ill 

Characters. — Size large; external characters unknown. Molars 
large and massive, especially M- 2 -. Large lower premolar subcorneal, 
apparently of the brown bear type. But in the north the grizzly 
type of premolar often fails in true grizzlies ; hence not having seen 
the claws, it is at present impossible to say whether innuitus is a 
brown bear or a grizzly. 

Cranial characters. — Old male (type): Skull large; basal length 
essentially the same as in horrihilis and alexandraz but occipito-nasal 
length much less, owing to shortness of occiput ; f ronto-nasal region 
strikingly dished: rostrum short, exceedingly broad (of same breadth 
as in alexandrce, very much broader than in horrihilis), strongly de- 
pressed; frontal shield exceedingly broad interorbitally, rising high 
and abruptly from rostrum, nearly horizontal behind plane of post- 
orbital processes, rather deeply sulcate medially and strongly 
swollen over orbits; postorbital processes large, subtriangular, out- 
standing and decurved; nasals nearly horizontal; palate and post- 
palatal shelf broad; postpalatal notch of medium width; zygomata 
broadly spreading and somewhat outbowed posteriorly, acute ante- 
riorly ; nares broader than high ; sagittal crest short, extending only 
about 25 mm. beyond fronto-parietal suture, straight (not arched), 
high posteriorly ; lambdoid crest large and full ; coronoid blade nar- 
row and high; ramus long and flat. Canines badly broken, appar- 
ently long; last lower premolar broad, broader posteriorly than ante- 
riorly, the cusp small and sloping posteriorly without heel or mar- 
ginal cusplets, but with pit and indication of narrow sulcus; molars 
exceptionally large and broad, the last upper one with heel strikingly 
long and broad, agreeing almost exactly with that of true horrihilis 
from eastern Montana. 

Two youngish skulls from Unalaklik (No. 82024, third year and 
No. 210554, fourth year) are believed to be females of this species. 
They are not old enough to show adult cranial characters except that 
the postpalatal shelf is broad and flat and the notch broad, but the 
teeth are perfect, full grown, and unworn. Canines of medium size 
(in the type badly broken and hence not available for comparison) ; 
molars large but smaller and less massive than those of type; M 4 
large, cusps on inner side nearly obsolete (presenting little more 
than an undulating line), heel long and broad with large flat gran- 
ular grinding surface. 

Cranial comparisons. — The only species needing comparison with 
innuitus are alexandrw and cressonus. The old male skull resembles 
certain old skulls of alexandrai from Kenai Peninsula, but differs 
rather strikingly in truncation of occiput (the occipital overhang 
being very much less) and corresponding shortening of braincase 
and sagittal crest. The crest moreover is straight instead of arcuate 
64854°— 18 8 



112 NORTH AMERICAN FAUNA. [No. 41. 

or arched, and the molars, especially M 1 and M A are very much 
larger. 

Compared with old male cressonus (type) : Basal length and 
zygomatic breadth essentially same ; occipito-nasal length much less ; 
frontal shield broader, much less elevated over orbits ; rostrum much 
broader and lower; occipital overhang much less; braincase and 
sagittal crest much shorter; postpalatal shelf broader and flatter. 

Skull measurements. — Old male (type) : Basal length, 353 ; occipito- 
nasal length, 331; palatal length, 194; zygomatic breadth, 251; 
interorbital breadth, 104. 

URSUS CRESSONUS Mebbiam. 

Chitina Beak. 

Ursus cressonus Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 137-139, 
September 6, 1916. 

Type locality. — Lakina River, south slope of Wrangell Range, 
Alaska. 

Type specimen. — No. 206529, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected by Captain J. P. Hubrick, of 
McCarthy, Alaska, 1914. 

Range. — Chitina River valley and adjacent slopes of Skolai and 
Wrangell Mountains, westerly doubtless through Chugach Moun- 
tains to the west side of Cook Inlet; occurs as far south as the 
Iliamna region. 

Cranial characters. — Old male (type) : Skull peculiar and distinc- 
tive; size large (basal length 357 mm.); skull long, narrow, high, 
and strongly dished ; frontal shield highly elevated, rising abruptly 
from rostrum, rather broad, deeply sulcate throughout medially, 
swollen over orbits, short pointed posteriorly ; orbits nearly vertical ; 
postorbital processes small and strongly decurved; fronto-nasal re- 
gion sulcate and strongly dished; rostrum rather short and narrow; 
nasals horizontal except posteriorly, where they rise strongly ; brain- 
case exceedingly long; sagittal crest high posteriorly, straight, and 
long, reaching anteriorly to halfway between parietals and plane of 
postorbitals ; zygomatic arches moderately spreading, subtriangular 
(not outbowed), expanded vertically; palate and postpalatal shelf 
relatively long and narrow for so large a skull; postpalatal notch 
rather narrow and short; occipito-sphenoid long (about 103 mm.); 
mastoids outstanding; anterior nares rather small, subtruncate, and 
broader than high in type skull, higher and less truncate in the 
Iliamna skulls. Under jaw absent in type specimen, but in an old 
male from Iliamna on north side of Cook Inlet (No. 209885) which 



1918.] INITUITUS GROUP. 113 

in most respects closely matches the type, the ramus is broadly flat- 
tened vertically, much broader posteriorly than anteriorly, and the 
coronoid blade is high and rather vertical. In younger skulls from 
Iliamna the coronoid is broader basally and less high. Canines large 
and massive; molars moderate. The last upper molar is large in 
the type, smaller and more cut away on outer side of heel in the 
Iliamna specimens. No. 209885 from Iliamna agrees with the type, 
except that the nares is higher and less truncate, and the last upper 
molar smaller, with heel more cut away on outer edge. 

Old female (No. 209881, from head of Chitina Eiver, 80 miles from 
McCarthy, Alaska; collected by Capt. J. P. Hubrick) : Size medium; 
cranium moderately arched; frontal shield broad, deeply sulcate an- 
teriorly, strongly swollen over and posterior to orbits, the point 
lyrate and reaching parietals; postorbitals rather large, blunt, and 
somewhat decurved; fronto-nasal region strongly dished and de- 
pressed medially ; rostrum rather large and high, nearly horizontal ; 
palate and postpalatal shelf broad; postpalatal notch moderate and 
rather broad. Underjaw long; coronoid blade high and rather 
narrow, its apex only slightly recurved. Dentition light; canines 
small and short; molars rather narrow, apparently normal (too 
badly worn to admit of description, except that the heel of M> is 
moderately long and rather broadly rounded posteriorly). 

Cranial comparisons. — Old male (type) compared with old male 
dalli: Size about the same; vault of cranium and frontal shield much 
more elevated, less flat, less nearly horizontal, and much more swollen 
over orbits; shield more deeply sulcate; postorbitals weak and de- 
curved (in dalli larger and more horizontally outstanding) ; fronto- 
nasal region more strongly dished; rostrum narrower and longer; 
zygomata much less widely outstanding and much less bowed ; palate 
longer ; molars very much larger. 

Old male (type) compared with adult male nuchek (type) : Size, 
elevation of vault of cranium, and zygomatic breadth about the same ; 
frontal shield somewhat broader, much more highly arched, much 
more swollen over orbits, much more deeply sulcate medially, shorter 
and more acutely pointed posteriorly ; postorbitals smaller and more 
decurved; fronto-nasal region strongly dished; rostrum more de- 
pressed; nares more truncate; last upper molar of normal form, 
large, and with long posteriorly rounded heel, differing widely from 
the short, broad-in-the-middle, obliquely truncate tooth of nuchek. 

Skull measurements. — Old male (type): Basal length, 357; oc- 
cipito-nasal length, 354 ; palatal length, 199 ; zygomatic breadth, 244 ; 
interorbital breadth, 97. 



114 NORTH AMERICAN FAUNA. I No. 41 

URSUS ALEXANDRA 1 Merbiam.* 
Alexander Grizzly. 

Ursus alexandrw Merrlam, Proc. Biol. Soc. Washington, XXVII, pp. 174-175, 
August 13, 1914. 

Type locality. — Kusilof Lake, Kenai Peninsula, Alaska. 

Type specimen. — Skull No. 4752, $ old, Museum of Vertebrate 
Zoology, University of California ; original No. 218. Collected Sep- 
tember, 1906. (Skull, skin, and skeleton complete.) 

Range. — Kenai Peninsula, Alaska. 

Characters. — Size very large ; skull long and narrow ; rostrum ex- 
ceptionally broad for a grizzly; pelage very uniform in color, 
scarcely or not grizzled; clav^s enormous (second foreclaw of type 
specimen measuring: length from upper base, 91 mm; height at 
base, 25; breadth, 11.5). The longest claw in a specimen collected 
by Wilson Potter measures 120 mm.; in a male killed by Dall De- 
Weese, 110 mm. 

Color. — Type, very old male, in fresh short fall pelage: General 
color pale, almost grayish brown, becoming yellowish brown between 
ears, contrasting with pale brown of muzzle; legs and feet only 
slightly darker than back; entire animal remarkably unicolor; under- 
fur plumbeous, crinkled, and wooly. Another male, killed by Wilson 
Potter, of Philadelphia, in May, 1912 (belonging to skull No. 181102, 
presented by Wilson Potter), is pale buffy inclining to light reddish 
brown throughout, without grizzly appearance; legs only slightly 
darker. One killed by Dall DeWeese, of Canyon City, Colorado, 
September 7, 1897, is described by him as "grayish-yellow," with 
legs and sides chocolate-brown. 

Cranial characters. — Skull large, long, rather narrow, with mod- 
erately spreading zygomata, short flattish frontal shield, outstanding 
postorbitals (with age) ; very broad rostrum for a grizzly, and long 
and high sagittal crest. Frontal shield not markedly elevated above 
plane of rostrum; posterior root of zygoma not expanded. Canines 
large and long. 

Cranial comparisons. — Adult male compared with adult male 
kenaicnds: Basal length, palate, and occipito-sphenoid length essen- 
tially the same ; skull as a whole much narrower, frontal shield inter- 
orbitally and across postorbital processes much narrower, flatter, more 
nearly horizontal, not materially elevated above plane of rostrum; 
zygomata much less widely spreading, squamosal part much narrower 
(not expanded) ; sagittal crest much longer, reaching anteriorly over 
posterior third of frontals (in kenaiensis ending on or near fronto- 
parietal suture) ; posterior third of frontals compressed, rising in 

1 Named for Annie M. Alexander, founder of the Museum of Vertebrate Zoology, Uni- 
versity of California, Berkeley. 

* Tentatively included in innuitus group. (See Introduction, pp. 12-13.) 



1918.] TOWNSENDI GROUP. 115 

a keel to sagittal crest. Canine teeth, both upper and lower, but 
especially the lower, much larger and longer. 

Remarks. — The skull of alexandrce is of a generalized type, lacking 
the special distinctive features that characterize several of its neigh- 
bors — as kenaiensis, sheld&ni, and others — none of which are true 
grizzlies. Among the grizzlies it stands alone in the great breadth of 
the rostrum, which in bears of its size is only exceeded by the widely 
different kenaiensis. Ursus alexandrce attains the largest size known 
among the grizzly bears, the biggest skulls equaling those of the 
huge magister of Southern California. 

Skull measurements. — Old male (type) : Basal length, 355 ; occi- 
pito-nasal length, 358; palatal length, 191; zygomatic breadth, 252; 
interorbital breadth, 87. 

Townsendi Group. 

URSUS TOWNSENDI 1 Mebbiam. 

Town send Beab. 

Ursus townsendi Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 151-152, 
September 6, 1916. 

Type locality. — Mainland of Southeastern Alaska (exact locality 
uncertain). 

Type specimen. — No. 216643, $ old, U. S. National Museum. 
Purchased at Sitka, in 1889, by Dr. Charles H. Townsend. 

Cranial characters. — Skull large, long, massive, rather low, and 
flat topped, dished, with extremely small teeth. Shield broad, flat, 
slightly depressed medially, the point ending anterior to parietals, 
sides reaching out broadly into very broad postorbitals, strongly 
sloping to rostrum; rostrum moderate, flat or depressed on top; 
nares truncate; zygomata moderately outstanding and moderately 
bowed; squamosal base broadly and abruptly expanded vertically; 
palate and postpalatal shelf moderate; notch rather narrow; mas- 
toids long; occipito-sphenoid 95 mm.; basisphenoid rather deeply 
concave. Underjaw long; ramus broad and flat vertically; coronoid 
of moderate height, narrowing above, sloping strongly backward, 
apex cutting plane of posterior part of condyle ; upper two-thirds of 
anterior border strongly inflected. 

Cranial comparisons. — Old male (type) compared, with male 
caurinus : Skull much larger, broader, more massive, and less arched ; 
teeth smaller. Frontal shield very much broader interorbitally and 
postorbitally (interorbitally 91 mm. contrasted with 81 or less; 

* Named for Dr. Charles H. Townsend, formerly naturalist of the Fish Commission 
steamer Albatross, now director of the New York Aquarium, who collected the specimen 
and presented it to the U. S. National Museum. 



116 NORTH AMERICAN FAUNA. |No.41. 

across postorbitals 130 contrasted with 116) ; postorbitals very much 
broader and flatter; rostrum more nearly horizontal; nares truncate 
instead of sloping; zygomata more widely outstanding and more 
broadly expanded vertically. 

Skull measurements. — Old male (type): Basal length, 348; occi- 
pito-nasal length, 353; palatal length, 183; zygomatic breadth, 245; 
interorbital breadth, 91.5. 

Dalli Group. 

URSUS DALLI 1 Merriam. 

Dall Brown Bear. 

Urgus dalli Merriam, Proc. Biol. Soc. Washington, X, pp. 71-73, April 13, 1896. 

Type locality. — Yakutat Bay (northwest side), Alaska. 

Type specimen. — No. 75048, $ old, U. S. National Museum. 

Range. — Malaspina Glacier and region northwest of Yakutat Bay, 
Alaska. 

Characters. — Size very large ; skull without very pronounced char- 
acters, although differing sufficiently from its neighbors. General 
color dark brown, strongly grizzled. 

Color. — Adult male (from Malaspina Glacier, belonging to skull 
No. 210293, killed by G. Frederick Norton) : Muzzle pale brown, be- 
coming much darker on head and sides of face; general body color 
dark brown, moderately grizzled with pale-tipped hairs, the grizzling 
beginning on top of head a little in front of plane of ears and ex- 
tending posteriorly to middle part of back. 

Cranial characters. — Adult male: Skull large, vault of cranium 
fairly high but not arched ; frontal shield broad, rather flat, broadly 
sulcate or concave medially, rather short pointed posteriorly, with 
large, rounded, broadly outstanding postorbital processes; fronto- 
nasal region moderately dished; rostrum broad and short, in some 
skulls slightly depressed; sagittal crest of medium length, rather 
high and nearly straight; zygomata rather broadly outstanding and 
strongly outbowed; squamosal root expanded in old age; squamosal 
shelf broad ; braincase narrow anteriorly, tending to keel into sagittal 
crest; nares small. 

Note. — The type skull of dalli (No. 75048, $ old) is abnormal: 
abnormally large, abnormally high (vault of cranium abnormally 
arched) ; fronto-nasal region abnormally elevated — not dished as 
usual; and underjaw abnormally long. No. 75047 (old $ ) and No. 
210293 (adult $ ) are far more typical. 

1 Named for William H. Dall, of the Smithsonian Institution. 



1918.] DALLI GROUP. 117 

Adult females (No. 140085, from Copper River delta, July, 1905, 
A. G. Maddren; and old female, No. 210308, from Bering Lake, 
1915, J. L. Hill) : Size, medium; skull broad and short for a female; 
moderately arched, and moderately dished, with rather broad frontal 
shield and broad rostrum. Frontal shield rather broad, strongly 
sulcate medially, swollen over orbits; postorbital processes rather 
weak and strongly decurved ; point of shield ending at f ronto-parietal 
suture; fronto-nasal region strongly sloping, rostrum large, broad, 
and rather short; sagittal crest confined to parietals, high for a 
female; postpalatal shelf medium or broad, strongly rounded on 
sides; zygomata moderately spreading, subtriangular, the posterior 
base somewhat bowed and vertically expanded; under jaw rather 
short ; coronoid high, falcate, the apex cutting or overreaching plane 
of condyle; teeth medium; heel of M> rather long, slightly emar- 
ginate on outer side, the extreme tip with tendency to turn outward. 

Cranial comparisons. — Adult and old males (Nos. 75047 and 
210293) compared with adult male nuchek (No. 146459, type) : Size 
about the same (basal length essentially same, but occipito-nasal 
length decidedly less) ; frontal shield broader and more acutely 
pointed ; vault of cranium less high and more nearly horizontal ; post- 
orbitals more outstanding; rostrum slightly shorter and more de- 
pressed (appearing broader) ; zygomata more outbowed (less tri- 
angular) ; palate somewhat shorter; mastoids more appressed, closer 
to glenoid processes, constricting meatus tube (in nuchek more out- 
standing, leaving wide postglenoid space with correspondingly large 
open meatus) ; coronoid blade broader above (less falcate). Canines 
almost the same; molars decidedly smaller and less massive and in 
details quite different (as stated under nuchek). 

Adult female (No. 140085, from Copper River delta) compared 
with female nuchek (No. 44049, from near Mount St. Elias) : Skulls 
so different as not to require close comparison, that of dalli being 
massive, broadly arched or domed, and with massive underjaw, while 
that of nuchek is light, slender, and narrow, with low narrow flat- 
tened frontal region, long slender rostrum, and light underjaw. The 
teeth also differ strikingly. 

Adult and old male compared with old male cressonus (type) : 
Size about the same; vault of cranium and frontal shield much less 
elevated, flatter, more nearly horizontal, and much less swollen over 
orbits; shield less deeply sulcate; postorbitals larger and more hori- 
zontally outstanding (in cressonus weak and decurved) ; fronto-nasal 
region less strongly dished; rostrum broader and shorter; zygomata 
much more widely outstanding and much more bowed; palate 
shorter; molars very much smaller. 



118 NORTH AMERICAN FAUNA. [No. 41. 

Adult and old female compared with old female cressonus: Skull 
larger, broader, and more highly arched ; frontal shield broader, less 
deeply sulcate and less swollen over orbits ; f ronto-nasal region more 
strongly dished ; rostrum smaller and lower ; zygomata less outstand- 
ing, less arched, much less expanded vertically and more sharply 
angular; underjaw much shorter and lighter; coronoid smaller and 
lower. Teeth smaller throughout. 

Adult and old male compared with adult and old male kenaiensis: 
Size the same or somewhat smaller ; skull much less massive ; braincase 
conspicuously narrower ; frontal shield interorbitally narrower, shorter 
pointed posteriorly; f ronto-nasal region normally more strongly 
dished ; postorbital processes less broadly rounded ; rostrum less mas- 
sive and less elevated ; sagittal crest much longer ; occipito-sphenoid 
shorter ; mastoids usually less outstanding. 

Skull measurements. — Old male (No. 75047, from Yakutat Bay, 
Alaska) : Basal length, 345 ; x occipito-nasal length, 342 ; palatal length, 
190; zygomatic breadth, 263; interorbital breadth, 91.5. Old male 
(No. 210293, from Malaspina Glacier) : Basal length, 345; occipito- 
nasal length. 338; palatal length, 188; z} T gomatic breadth, 248; inter- 
orbital breadth, 96. 

URSUS HOOTS' Merbiam. 
Stikine Bbown Beab. 

Vrsus hoots Merrlam, Proc. Biol. Soc. Washington, XXIX, pp. 140-141, Sep- 
tember 6, 1916. 

Type locality. — Clearwater Creek, a north branch of Stikine River, 
British Columbia. 

Type specimen. — No. 206136, $ ad., U. S. National Museum, 
Biological Survey collection. Collected by John Hyland; presented 
by Lincoln Ellsworth. 

Cranial characters. — Size medium or large; skull massive; slightly 
dished, rather short, flattish on top, very broad across frontals and 
rostrum. Frontal shield broad, nearly flat, long pointed; broadly 
and shallowly sulcate medially as far back as posterior plane of post- 
orbitals; postorbitals large, broad, and horizontally outstanding; 
f ronto-nasal region sloping ; rostrum broad and rather high ; palate 
and postpalatal shelf broad; postpalatal notch moderate; sagittal 
crest short, ending at fronto-parietal suture; zygomata moderately 
outbowed, not broadly spreading; underjaw rather massive; ramus 
broad vertically, its inferior border upcurved posteriorly; coronoid 
blade broad at base, rather high and subfalcate, the apex curving 
strongly backward, cutting plane of condyle; dentition remarkably 

1 Restored. 

* Hoots, the native Indian name for the big brown and grizzly bears of tbe coast region. 



1918.] DALLI GROUP. 119 

light for so large a skull ; canines and molars (both upper and lower) 
surprisingly small. 

Remarks. — Ursus hoots does not appear to be related to any of 
the other mainland species except the newly discovered crassus from 
the mountains on the upper Macmillan River, Yukon, from which 
it may be distinguished at a glance by the small size of its molar 
teeth. It may be related also to sitkensis of Baranof and Chi- 
chagof Islands, but differs in somewhat smaller size; less elevated 
posterior frontal region; broader postorbital processes; less broadly 
spreading zygomata ; shorter and less spreading mastoids ; less nearly 
vertical and more strongly recurved coronoid blade, the apex over- 
arching a well-defined coronoid notch; smaller molars (both upper 
and lower) ; and smaller upper incisors. The large lower premolar 
has the upturned heel of the Sitka bear, but lacks the posterior sulcus 
and pair of cusplets of the grizzlies. Two additional old male skulls 
of hoots have been recently received from the Stikine River region. 
One of these (No. 224841) from Clearwater branch of Stikine — the 
type locality — agrees closely with the type specimen in size and form, 
but has slightly larger and especially broader molars (difference 
most marked in M ). The skull is older than that of the type and 
the mastoids are more strongly developed and divergent. The other 
(No. 224839) is still older and was killed low down the Stikine. It 
is of approximately the same size as the type, but considerably older ; 
the frontal shield is more broadly concave interorbitally and shorter 
pointed posteriorly; sagittal crest more strongly developed; squa- 
mosal base of zygoma much more broadly expanded; mastoids much 
longer and strongly divergent; postpalatal shelf flatter; coronoid 
blade broader at and above middle ; molars slightly larger. An adult 
female from the type locality (No. 180883, from Clearwater branch 
of Stikine) resembles the male type very closely except for the 
smaller size of skull and teeth, and therefore needs no special descrip- 
tion. A feature deserving mention is that in both sexes the point 
of the frontal shield ends at the fronto-parietal suture. 

Skull measurements. — Adult male (type) : Basal length, 333; oc- 
cipito-nasal length, 325; palatal length, 179; zygomatic breadth, 228; 
interorbital breadth, 96. 

URSUS SITKENSIS Merbiam. 

Sitka Bbown Bear. 

Ursus sitkensis Merriam, Proc. Biol. Soc. Washington, X, p. 73, April 13, 1896. 

Type locality. — Sitka Islands, Alaska. 

Type specimen. — No. 187891, $ ad., U. S. National Museum (No. 
6543, Merriam collection). Collected by an Indian; purchased at 
Sitka, Alaska, and presented to C. Hart Merriam by J. Stanley- 
Brown. 



120 NORTH AMERICAN FAUNA. [No. 41. 

Range. — Sitka Islands (Baranof and Chichagof), Alaska. 

Characters. — Size large; coloration very dark; claws of moderate 
length, curved, dark blue-black, scurfy ; skull broad and massive. 

Color. — Dark; muzzle dark brown, sometimes chocolate brown or 
even sooty, paler in faded summer pelage ; head and body very dark 
brown or even dusky, varying to dull brown in summer, washed on 
back of head, neck, and shoulders with yellowish or golden. 

Cranial characters. — Adult male: Skull large, massive, dished, 
vault moderately elevated, frontal shield (normally) broad, strongly 
sloping, sulcate medially ; postorbitals outstanding, broadly rounded ; 
rostrum normally rather short and somewhat depressed; sagittal 
crest massive, straight; zygomata broadly outstanding and out- 
bowed ; palate moderate ; postpalatal shelf normally rather long and 
of medium breadth ; notch moderate or rather narrow ; basioccipital 
broad ; mastoids long and spreading. 

Cranial comparisons. — Adult and old males compared with adult 
and old male dalli: Frontal shield higher posteriorly (in dalli highest 
at or immediately behind postorbitals), flatter and more sloping at 
base of postorbitals, thus tilting postorbital plane forward (in dalli 
looking more directly upward) ; a distinct thickening or hump pres- 
ent on each side of median sulcus behind plane of postorbitals (lack- 
ing in dalli) ; basioccipital usually broader; inion less strongly de- 
veloped; mastoids longer, usually more outspreading and more dis- 
tant from glenoid processes, leaving broader space for audital canal ; 
auditory meatus larger; rostrum somewhat broader basally; dis- 
tance from last lower molar to middle of condyle usually less; last 
upper molar somewhat longer and more nearly rectangular, the outer 
side of heel less oblique ; M T normally with open saddle between pos- 
terior and anterior parts (in dalli a cusplet occupies the inner side of 
the saddle) ; PM ¥ in sitkensis normally tricuspidate as seen in profile, 
there being both anterior and posterior cusplets on the cingulum (in 
dalli the anterior and posterior cusplets are absent and the main cusp 
is larger and higher and slopes posteriorly without horizontal heel). 

Skull measurements. — Average of 2 adult males from Chichagof 
Island: Basal length, 358.5; occipito-nasal length, 354.5; palatal 
length, 189; zygomatic breadth, 260; interorbital breadth, 93.5. 

URSUS SHIRASI 1 Merbiam. 

Shieas Brown Beab. 

(PL VI.) 

Uraus shirati Merriam, Proa BloL Soc. Washington, XXVII, p. 195, August 
13, 1914. 

Type locality. — Pybus Bay, Admiralty Island, Alaska. 

1 Named for George Shiras, 4th, who collected and presented the specimen. 



1918.] DALLI GROUP. 121 

Type specimen. — No. 203030, $ old, U. S. National Museum, Bio- 
logical Survey collection. Killed September 16, 1913, by George 
Shiras, 4th, and presented to the Biological Survey. 

Range. — Restricted to Admiralty Island. 

Characters. — A huge brown bear larger than the largest sit- 
kensis; head highly arched; color black, except muzzle, which is 
dull brown ; claws dark blue- black, dull, slightly scurfy (not smoothly 
polished as in the grizzlies), rather strongly curved and of mod- 
erate length (middle claw over curve, 92 mm.; from top of base to 
apex, 75), fourth and fifth rounded off on outer side. 

Color. — Entire animal, except muzzle, coal black, showing when 
examined closely a brownish wash along middle of back ; muzzle from 
nose pad to between eyes dull brown. 

Cranial characters. — Old male (type) : Skull large, broad, massive, 
strongly dished, and highly arched; zygomata large, broadly out- 
bowed and rounded anteriorly as well as posteriorly; frontal shield 
remarkably short and broad (nearly twice as broad as long), deeply 
and broadly concave in cross section, with huge uplifted 1 broadly 
outstanding postorbital processes which arch over the orbits and are 
strongly decurved apically, completely roofing the orbits; temporal 
ridges beaded, short, meeting far forward (at least 25 mm. in front 
of fronto-parietal suture) ; sagittal crest long and high, humped 
over fronto-parietal suture; fronto-nasal region strongly concave; 
rostrum broad and short, rising strongly to meet frontal shield; 
palate broad; basioccipital and basisphenoid subequal; ramus 
strongly bellied under posterior molars; coronoid blade broad and 
high. 

Dental characters. — Dentition heavy; canines large, the upper 47 
mm. high above enamel line of outer side; molars large and rather 
broad; M 1 with large and broad heel; M T with strongly developed 
cusplet on inner side of saddle. 

Remarks. — Ursus shirasi is a very large member of the brown 
bear group. Whether it is always black, like the type specimen, is 
not known. But of all the American bears its skull is the most 
striking and distinctive. The short broad frontal shield rising on 
each side into huge postorbital processes, which arch broadly over 
the orbits, serve to distinguish it at a glance from all other species, 
rendering close comparisons unnecessary. In this connection it is 
interesting to observe that shirasi and its neighbor eulophus, an in- 
habitant of the same island, present opposite extremes of departure 
from the normal ursine type — eulophus having a long narrow skull 
with slender elongate rostrum, long and narrow frontal shield, and 

1 Additional skulls of adult males recently received have the postorbitals equally large 
but less uplifted, not rising above frontal plane. 



122 NORTH AMERICAN FAUNA. [No. 41. 

insignificant postorbital processes, while shirasi has an exceptionally 
broad skull with broad short rostrum, excessively broad and short 
frontal shield, and huge massive postorbital processes. 

Skull measurements. — Adult male (type) : Basal length, 355; oc- 
cipito-nasal length, 348; palatal length, 191; zygomatic breadth, 
259; interorbital breadth, 104.5. 

URSUS NUCHEK Mereiam. 1 

Nuchek Brown Bear. 

Ursus nuchek Merriam, Proc. Biol. Soc. Washington, XXIX, pp. 146-148, Sep- 
tember 6, 1916. 

Type locality. — Head of Nuchek Bay, Hinchinbrook Island, Prince 
William Sound, Alaska. 

Type specitnen. — No. 146459, $ old, U. S. National Museum, Bio- 
logical Survey collection. Collected September 15, 1905, by C. Swan- 
son. 

Range. — Prince William Sound easterly to Mount St. Elias ; limits 
unknown. 

Characters. — Size large; external characters unknown; skull long, 
narrow, and moderately high ; molars peculiar. 

Cranial characters. — Adult male (type) : Large, elongate; frontal 
shield relatively narrow, flattish, moderately depressed between 
orbits; orbital rims thickened; postorbital processes broad and flat- 
tish, moderately outstanding; posterior part of shield broad, ending 
about two-thirds distance from plane of postorbitals to fronto- 
parietal suture ; sagittal crest rather long, straight, high posteriorly ; 
rostrum long, high, rather narrow ; f ronto-nasal region sloping in 
facial plane; nasals slightly elevated anteriorly; zygomata moder- 
ately spreading, subtriangular, not much expanded vertically; post- 
palatal shelf moderate, its sides rounded; notch long and narrow; 
anterior nares small; meatus tube short and large. Underjaw mas- 
sive; coronoid blade narrow and falcate. Teeth of medium size; 
molars broad (more massive than in dalli) ; last upper molar excep- 
tionally short, broadest in middle, heel short and obliquely truncate 
on outer side ; M 1 large, much broader posteriorly than anteriorly ; 
middle lower molar peculiar: twin cusps of entoconid very small, 
low, and close together; main cusp of inner side large and high, re- 
ducing the posterior moiety of the tooth to about a third the length 
of the crown instead of about half as usual. 

Young -adult female (No. 44049, from Chaix Hills near Mount St. 
Elias, Alaska; killed July 4, 1891, by the late Prof. I. C. Russell) : 
Skull long, narrow, rather low, with narrow frontals, narrow ros- 

* Tentatively included in dalli group. (See Introduction, pp. 12-13.) 



1918.] DALLI GROUP. 123 

trum, and moderately outstanding subtriangular zygomata. Frontal 
shield flattish, medially depressed interorbitally, sloping gradually 
into rostrum, rather short pointed posteriorly (ending about 15 mm. 
in front of parietals; in fully adult and old females it would be 
still shorter) ; postorbital processes moderate, horizontally outstand- 
ing, the tips rounded (not fully grown) ; palate concave, postpalatal 
shelf rather long and broad; notch rather narrow; basisphenoid 
strongly concave antero-posteriorly and transversely; underjaw long 
and slender. Canines long and slender ; molars and large premolars 
with rather high cusps ; last upper molar short, much broader in mid- 
dle than anteriorly, heel short and obliquely truncate on outer side. 

Cranial comparisons. — Ursus nuchek evidently overlaps the range 
of dalli and may come in contact with rressonus, necessitating com- 
parisons with both. 

Adult male (type) compared with adult and old male dalli (Nos. 
75047 and 210293) : Size about the same; basal length essentially the 
same, but occipito-nasal length decidedly greater; skull appearing 
longer and narrower; more elevated behind orbits and much more 
strongly sloping posteriorly; frontal shield narrower, the point 
broader posteriorly; vault of cranium higher and less nearly hori- 
zontal ; postorbitals less outstanding ; f ronto-nasal region less dished ; 
rostrum longer and not depressed (appearing narrower) ; zygomata 
less outbowed (more triangular) ; palate somewhat longer; post- 
palatal notch longer and narrower; mastoids less appressed, leaving 
wide postglenoid space with correspondingly large open meatus (in 
dalli closer to glenoid process, pressing on and contracting meatus 
tubes); coronoid blade narrower above (more falcate). Canines 
about the same ; molars, both upper and lower, decidedly larger and 
more massive and in details quite different : M^ exceptionally short 
and much broader in middle than elsewhere, the heel short and 
obliquely truncate on outer side; M 1 large, much broader posteriorly 
than anteriorly; middle lower molar peculiar, the twin cusps of 
entoconid very small, low, and close together; metaconid exception- 
ally large and high, reducing the posterior moiety of the crown to 
about one-third its length, instead of about half as in dalli and most 
species. 

Young-adult female (No. 44049, from near Mount St. Elias) com- 
pared with adult female dalli (No. 140085, from Copper Eiver 
delta) : Skulls so strikingly different as not to require close compari- 
son, that of nuchek being light, slender, narrow, with low narrow 
flattened frontal region, long slender rostrum, and light underjaw, 
while that of female dalli is massive, broadly arched or domed, and 
with massive underjaw. The teeth also differ strikingly. 



124 NORTH AMERICAN FAUNA. INo. 41. 

Young-adult female (No. 44049) contrasted with old female cres- 
sonus (No. 209881) : Size slightly smaller (when fully adult probably 
the same) ; frontal shield much narrower and flatter, much less 
deeply sulcate, much less swollen over orbits, and much shorter pos- 
teriorly; fronto-nasal region in same plane (in 2 cressonus strongly 
dished); sagittal crest longer; nares smaller; canines (upper and 
lower) much longer; molars more massive; M^ extremely short, 
much the broadest in middle, with short obliquely truncate heel (in 
cressonus normal). 

Female (No. 44049) compared with female kenaiensis (No. 
133244) : Basal length essentially the same; cranium narrower, with 
narrower braincase, narrower shield, and narrower rostrum; zygo- 
mata less broadly spreading (would be more broadly spreading with 
age) ; occipito-sphenoid shorter; palate essentially same length but 
narrower; postpalatal shelf narrower; ramus more slender (con- 
spicuously thinner below M-y and M 7 ) ; its inferior border straighter, 
less upcurved posteriorly; coronoid lower and less narrowed above; 
cusps of larger premolars above and below much more highly de- 
veloped ; main cusp of upper premolars very much higher relative to 
posterior cusp ; molar cusps also more strongly developed ; last upper 
molar shorter and of peculiar form, as in the male. 

Skull measurements. — Old male (type) : Basal length, 360 ;* occi- 
pito-nasal length, 358; palatal length, 191; zygomatic breadth, 248; 
interorbital breadth, 88. 

Gyas Group. 

URSUS GYAS Mebbiam. 

Peninsula Giant Beab. 

(Pi. II.) 

Ursus dalli gyas Merriam, Proc. Biol. Soc. Washington, XV, p. 78, March 22, 

1902. 
Ursus merriami Allen, Bull. Amer. Mus. Nat. Hist., XVI, p. 141, April 12, 1902. 

Type locality. — Pavlof Bay, Alaska Peninsula. 

Type specimen. — No. 91669, $ ad., U. S. National Museum, Bio- 
logical Survey collection. 

Range. — Entire length of Alaska Peninsula from Cook Inlet to 
Isanotski Strait and adjacent Unimak Island. 

Characters. — Size huge, either largest living bear or second only to 
the great Kadiak bear {middendorffi) . Claws rather long and 
smooth, dark when young, pale when old. Color variable, from 
grizzled brown to pale yellowish. Skull of male large, long, and 
massive, but not highly arched. Sexual disparity great. 

1 Restored. 



1918.] GYAS GROUP. 125 

Cranial characters. — Adult males : Skull large, long, and massive ; 
frontal region moderately elevated, sloping gradually into rostrum, 
strongly depressed or troughed medially; slightly swollen over or- 
bits and bases of postorbitals ; postorbitals rather large, subtriangu- 
lar, moderately outstanding; zygomata moderately outstanding and 
bowed; palate long and relatively narrow. Underjaw long and 
massive; coronoid blade very broad basally. Molars light for so 
large a skull. Viewed from behind, the posterior frontal region, with 
its depressed median trough and massive, outstanding, and elevated 
postorbital processes, suggests the spread wings of a bat or a 
butterfly. 

Skulls of adult males which it seems necessary to call gyas present 
a surprisingly wide range in size and form. Among them are three 
quite different types which if isolated would undoubtedly develop 
into very distinct species. 

1. Typical form, with large elongate skull. — Skull and teeth large and mas- 
sive ; zygomata widely spreading ; sagittal crest high ; f rontals moderately ele- 
vated, broadly sulcate medially, swollen laterally ; postorbital processes thick, 
massive, subtriangular, and decurved ; nasal opening and rostrum proportionate 
to size of skull (but anterior nares very much smaller than in middendorffl). 
Teeth large; lower canines averaging about 19 mm. in diameter at base of 
enamel. This large form ranges over the entire length of Alaska Peninsula 
from Cook Inlet (where it was obtained at Chinitna by Kidder and Blake) to 
Morzhovoi Bay and Unimak Island. Departures from normal: The series of 
skulls at hand shows two prominent departures from typical gyas — one larger 
and more massive, the other smaller and lighter. 

2. Giant form, with exceptionally broad rostrum. — Huge skulls with broad 
massive rostrum and exceedingly heavy jaws. This type is represented in the 
Biological Survey collection by two specimens, No. 91694 from Cold Bay, and 
No. 91704 from Belkofski Bay. The entire skull is larger and more massive, but 
the differences are most pronounced in the face and jaws. The frontal shield 
and postorbital processes are not broader than in some skulls of typical gyas, 
but the postorbital processes are abruptly de flexed at the tips, forming a thick 
massive hook over the orbit, much as in middendorffi. The rostrum is extraor- 
dinarily broad and massive, giving the skull, viewed from the front, a most 
peculiar aspect. The occipital flange (lambdoid crest) also is largely de- 
veloped. The peculiarities of the underjaw are as striking as those of the 
rostrum. The inferior part of the ramus is greatly thickened anteriorly, and 
the posterior half is strongly everted, forming a broad lip, unlike anything 
seen in typical gyas. By reason of this peculiarity the jaws flare strongly out- 
ward under the last molars, and the flaring is so great that it is conspicuous 
even when looked at from above. 

3. Small form, with narrow rostrum. — Smaller skulls, with narrower ros- 
trum, more slender, horizontally outstanding postorbital processes and much 
smaller canines (exemplified by Nos. 82003 and 82004 from Pavlof Bay, and 
No. 91699 from Belkofski Bay). Compared with typical gyas, the skull as a 
whole is considerably smaller, shorter, and lighter ; vault of cranium rather 
more flattened; rostrum decidedly narrower; canine teeth above and below 
decidedly more slender ; last upper molar narrower. But the most conspicuous 
difference is in the postorbital processes, which instead of being broadly tri- 



126 NORTH AMERICAN FAUNA. lNo.41. 

angular, massive, and decurved, are elongate, peglike, and stand out hori- 
zontally. 

The above description applies to fully adult males of the form described by 
Allen under the name Ursus merriami, the type of which was an immature 
male. Unfortunately for merriami, the typical form seems to be connected 
with gyas by a series of intergrades. Thus, skull No. 862, Field Mus. Nat. 
Hist., has the small teeth and narrow muzzle of merriami, but the postorbital 
processes are broader posteriorly and slightly decurved; and Nos. 91691, 
147630, 91675, U. S. Nat. Mus., Biological Survey collection, and No. 4585, Mus. 
Vert. Zool., Univ. California, complete the chain of intergrades, so that it is 
difficult to tell just where to draw a line between them. 

Skulls of adult males of the small form have been examined from various 
localities from Cold Bay and Ugashik Lake and River westerly to Pavlof, 
Belkofski, Bear Bay, and Tonki Point. 

Adult females: Skull of moderate size, conspicuously smaller than 
male; frontal region normally elevated, domed, and rounded off, the 
postorbital processes somewhat decurved, the frontal shield sulcate 
medially. Skulls of adult females differ among themselves in degree 
of elevation and doming of frontal shield, depth of median groove, 
relative massiveness, and other characters, but in the present imper- 
fect state of knowledge it is impossible to assign positively any par- 
ticular female to either of the above-described types of males. 

Cranial comparisons. — Old male compared with old male mid- 
dendorffi of essentially same size : Vault of cranium much less highly 
arched and never domed; postorbital processes more strongly de- 
veloped and less decurved ; zygomata less widely outstanding and far 
less bowed (ratio of zygomatic breadth to basal length much less) ; 
anterior nares normal, not flaring; coronoid blade less high. Last 
upper molar shorter and of different form, the heel obliquely trun- 
cate on outer side, narrowing posteriorly ; lower molars broader and 
heavier. 

Skull measurements. — Old male (type): Basal length, 380; oc- 
cipito-nasal length, 394 ; palatal length, 206 ; zygomatic breadth, 286 ; 
interorbital breadth, 96. 

URSUS MIDDENDORFFI Mebbiam. 

Kadiak Beab. 

(Pi. III.) 

Ursus middendorffl Merrlam, Proc. Biol. Soc. Washington, X, pp. 67-69, April 

13, 1896. 
Ursus kadiaki Kleinschmidt, Outdoor Life, XXVII, p. 3, January, 1911. 

Type locality. — Kodiak Island, Alaska. 

Type specimen. — No. 54793, $ young-adult, U. S. National Mu- 
seum, Biological Survey collection. Collected July 3, 1893, by B. J. 
Bretherton (original No. 176). 

Range.— Kodiak and adjacent islands, Afognak and Shuyak; not 
known from mainland. 



1918.] KENAIENSIS GROUP. 127 

Cranial characters. — Size huge; skull of male exceedingly broad, 
high, and relatively short; frontal shield domed, sulcate medially 
and swollen over orbits (obliquely flattened in extreme age), rather 
short pointed posteriorly, passing into sagittal crest in fully adult 
skulls anterior to fronto-parietal suture; fronto-nasal region dished; 
postorbitals decurved and weak, small for size of skull; rostrum 
rather short and of medium breadth; anterior nares flaring; zygo- 
mata extraordinarily outstanding and strongly outbowed ; palate 
rather broad; postpalatal shelf rather narrow; mastoids long and 
divergent Underjaw large, massive, and rather short; coronoid 
blade high and moderately recurved; molars small for size of skull. 

Cranial comparisons. — Old male compared with old male gyas of 
essentially same size: Vault of cranium much more highly arched and 
usually domed; postorbital processes weak and decurved; zygomatic 
arches much more widely outstanding and far more strongly bowed 
(ratio of zygomatic breadth to basal length much greater) : anterior 
nares strongly flaring (in gyas not flaring) ; coronoid blade higher. 
Last upper molar with heel more broadly rounded; lower molars 
narrower. 

Flesh measurements. — Young-adult male killed by J. H. Kidder 
on Shuyak Island, off Afognak, Alaska, July, 1901 : Total length, 
nose to end of tail vertebrae, 8 ft.; nose to base of tail. 7 ft. 8 % in.; 
height at shoulders, 4 ft. 34 in.; length of forefoot, including claws, 

I ft. 2£ in.; hind foot, 1 ft. 4 in.; width of forefoot, 84, in.: width of 
hind foot, 7| in.; length of fore claws. 44, in.; girth of body behind 
shoulders, 5 ft. If in.; girth of neck, 3 ft. 2 in.; girth of head at ears, 
3 ft. If in.; nose to tip of hind foot (animal lying on one side), 9 ft. 

II in. The body after the skin was removed was the size of a big ox. 
Skull measurements. — No. 134407, largest of the males: Basal 

length, 392; occipito-nasal length, 369; palatal length, 211; zygomatic 
breadth, 306; interorbital breadth, 101. 

Kenaiensis Group. 

URSUS KENAIENSTS Mekriam. 

Kenai Giant Beau. 

(PI. IV.) 

Ursus lenaiensis Merriam, Proc. Biol. Soc, Washington, XVII, p. 154, October 6, 
1904. 

Type locality.— Cape Elizabeth, at extreme west end of Kenai 
Peninsula, Alaska. 

Type specimen— No. 128672, 9 ad., U. S. National Museum, Bio- 
logical Survey collection. Collected in 1903 by C. A. Lambert. 

Range. — Kenai Peninsula, 
C4854°— 18 9 



128 NORTH AMERICAN FAUNA. [No. 41. 

Characters. — Size large; appearance that of a big grizzly; colora- 
tion rather dark; claws moderately curved, dark, usually marked 
with whitish streaks on sides and near tip; longest claw in three 
adults 82-90 mm. Skull broad and massive, that of male strikingly 
larger than female. 

Color.— Old male killed in October, 1912, by Wilson Potter, of 
Philadelphia (skull 1S1099) : Muzzle pale fulvous-brown; cheeks 
and forehead similar but hairs longer and with pale tips; ground 
color of top of head, neck, and back much darker but deeply washed 
on tips with buffy or buffy whitish, giving these parts the look of a 
grizzly; legs and feet (but not belly) much darker. 

Cranial characters. — Adult mate: Skull large, broad, and massive, 
with broad frontal shield and rostrum, widely outbowed zygomata, 
large outstanding bluntly rounded postorbitals, and small anterior 
nares. Frontals broad throughout (interorbitally, postorbitally, and 
across postorbital processes) ; frontal shield well elevated above plane 
of rostrum, shallowly sulcate medially between orbits, slightly swollen 
over orbits, long pointed posteriorly ; sagittal crest short, not reach- 
ing anteriorly beyond f ronto-parietal suture ; braincase broad anteri- 
orly; rostrum broad throughout but much broader posteriorly than 
anteriorly; zygomata abruptly and widely outbowed, the squamosal 
root vertically expanded; palate very broad; underjaw large and 
massive, coronoid blade large and high, scarcely recurved. Canines 
small and short. Adult female: Skull broad, flat, and massive, with 
exceedingly broad rostrum, broadening posteriorly; zygomata 
broadly spreading; jugal broad anteriorly; frontals flattened, de- 
pressed, low posteriorly ; postorbital processes large, blunt, and hori- 
zontally outstanding; palate exceedingly broad; nasals large and 
broad; anterior nares small. Canines small. 

Cranial comparisons. — Adult male compared with adult male 
alescandrce : Length essentially the same, but henaiensis much broader 
and more massive throughout; frontal shield broader throughout 
(interorbitally, postorbitally, and across postorbital processes), more 
elevated, convex (not flat) antero-posteriorly, shallowly sulcate inter- 
orbitally, slightly swollen over orbits; longer pointed posteriorly, 
with correspondingly shorter sagittal crest ending anteriorly at 
f ronto-parietal suture; postorbital processes more massive and out- 
standing; frontal part of braincase not keeled or compressed; rostrum 
much broader, especially posteriorly; nasals shorter anteriorly; zygo- 
mata much more widely and abruptly spreading posteriorly, outbowed 
instead of angular; squamosal part rising abruptly and broadly ex- 
panded vertically, differing strikingly from the more slender and 
gently curving form in alexondrce; palate broader; mastoids more 
outstanding; ramus longer and more massive; coronoid blade more 



1918.] KENAIENSIS GROUP. 129 

nearly vertical, higher, broader in upper third, less recurved, not 
ending in posterior point. Canines, both upper and lower, smaller 
and shorter. 

Adult female compared with adult female alexandrm: Skull 
shorter, both basall y and on top ; vault of cranium and rostrum lower ; 
fronto-nasal region more dished; rostrum very much broader basally, 
its sides sloping anteriorly; braincase shorter (less occipital over- 
hang) ; sagittal crest lower; occipito-sphenoid and base of skull 
shorter; palate broader; ramus more massive. Canines smaller (the 
lower notably more slender). 

Adult and old males compared with adult and old male dalli: 
Size the same or somewhat larger; skull much more massive; brain- 
case conspicuously broader; frontal shield interorbitally broader, 
longer pointed posteriorly; fronto-nasal region normally less 
strongly dished; postorbital processes more broadly rounded; ros- 
trum more massive and more elevated; sagittal crest much shorter; 
occipito-sphenoid longer; mastoids usually more outstanding. 

Skull measurements. — Average of 2 old males from Kenai Pen- 
insula: Basal length, 3G7; occipito-nasal length, 3G0; palatal length, 
197; zygomatic breadth, 2G3.5; interorbital breadth, 205. Adult 
female (type) : Basal length, -288.5 ; occipito-nasal length, 285; palatal 
length, 158; zygomatic breadth, 214; interorbital breadth, 78. 

UKSUS SHELDONI * Mekkiaji. 

Montague Island Bear. 

<ri.v.) 

Ursus slicldoni Merriam, Proc. Biol. Soc Washington, XXIII, pp. 127-130, 
September 2, 1910. 

Type locality. — Montague Island, Prince "William Sound, Alaska. 

Type specimen. — No. 137318, $ young-adult, U. S. National 
Museum, Biological Survey collection. Collected May, 1905, by 
Charles Sheldon. 

Characters. — Size large; teeth and claws of the grizzly type; color 
variable, from dark to light brown. Skull broad and massive; vault 
of cranium domed ; hairs over shoulders elongated to form a small 
but distinct hump. 

Color. — General color brownish, varying from pale to dark, the 
hairs of the back sometimes yellowish tipped, those of the head griz- 
zled; color darkest (almost blackish) on belly, legs, and feet; ears 
dark with whitish tips. An old she-bear killed by Sheldon, May 18, 

1 Named for Charles Sheldon, of New York, who collected and presented the type and 
Other specimens. 



130 NOBTH AMERICAN FAUNA. I No. 41. 

1905, is very pale grizzled gray on the upperparts, and only moder- 
ately darker on the legs and feet. The cub of this bear, killed 
the same day, Avas in its second year (about 1G months old) and 
is very pale — almost buffy gray — with dark feet and legs, and a 
strongly marked hump. 

Cranial characters. — Adult male (type) : Large, massive, excep- 
tionally broad, Avith broadly outboAved zygomata. Frontal shield 
long and broad, nearly horizontal from postorbitals posteriorly (act- 
ually sloping downAvard posteriorly) ; broadly sulcate medially be- 
tAveen postorbital processes, moderately sAvollen on each side, th? pos- 
terior part long and broad (temporal ridges in type specimen not 
curving inward as in most skulls), the point falling over middle of 
parietals (in old age doubtless more anterior) ; sagittal crest excep- 
tionally short, confined in type skull to posterior half of parietals; 
fronto-nasal region elevated, forming a convexity instead of the usual 
depression b?tAveen plane of rostrum and that of frontal shield; 
rostrum broad and short; nasals strongly sloping, dished at junction 
of anterior and middle thirds, the anterior third horizontal or 
slightly upturned ; palate and postpalatal shelf very broad : notch 
rather broad; zygomata broadly outstanding, rounded posteriorly, 
someAvhat bowed, not much expanded vertically (doubtless more 
broadly expanded in old age) ; squamosal shelves exceptionally 
broad; braincase narroAvest on anterior part of parietals instead of 
on frontals. Und^rjaAv large and massive; ramus broad vertically, 
broadest under posterior molars; coronoid blade elevated, moderate, 
vertical, the apex only slightly recurved. Dentition light; canines 
and molars small for size of skull. 

Old female (No. 137316, mother of cub No. 137315) from Mon- 
tagu? Island, May 18, 1905, collected and presented by Charles Shel- 
don : Skull of medium size, broad, flattish on top. Frontal shield flat, 
nearly horizontal, the posterior part broad and long, reaching past 
middle of parietals; postorbital processes strongly developed, almost 
peglike, horizontally outstanding; fronto-nasal region elevated in 
fronto-facial plane; rostrum short, rather broad; palate and post- 
palatal shelf broad; zygomata broadly outstanding, subtriangular, 
rounded posteriorly. Underjaw rather massive; coronoid moderate, 
the apex only slightly recurved; teeth small for size of skull; brain- 
case very broad. 

Cranial comparisons. — Skull in general similar to that of kenaien- 
sis but basisphenoid broader and flatter, its length nearly equal to 
that of basioccipital; posterior roots of interpterygoid fossa more 
Avidely spreading; condyle of jaw more exserted (in kenaiensis ses- 
sile), reaching so far back that a line dropped from peak of coronoid 
to tip of angle touches or traverses it (in kenaiensis this line passes 
freely behind the condyle) ; coronoid, in females of same age, smaller 



1018.] GENUS VETULARCTOS. 131 

and lower, its area for muscular attachment less; ramus of jaw 
strongly bellied posteriorly, its inferior border below the coronoid 
strongly convex downward and curving evenly, with only a very 
slight break, to angular process. (In kenaiensis the inferior border 
of ramus is nearly straight, not appreciably bellied under coronoid, 
and ends abruptly in a step or jog at some distance behind the angle.) 

In general form and appearance skulls of females closely resemble 
those of female kenaiensis, differing chiefly in the characters above 
mentioned and in certain dental peculiarities, notably the smaller 
size and more pointed heel of the last upper molar, and the oblique 
truncation of M^. 

Dental characters. — Teeth in general of the grizzly type. Last 
(fourth) lower premolar normally with horizontal heel, slightly up- 
turned at posterior end, the shallow median sulcus reaching from 
cusp to end of heel, its defining ridges ending in slightly developed 
posterior cusplets. (In kenaiensis the last lower premolar is more 
conical, the heel sloping, the sulcus incomplete, with only a single 
posterior cusplet — on inner side of main cusp posteriorly.) First 
upper molar peculiar, having both ends obliquely truncate and paral- 
lei, sloping strongly from outer angles backward and inward; in- 
ner row of cusps pushed back so that each falls behind plane of 
corresponding cusp on outer side; tooth as a whole more nearly rec- 
tangular, its inner corners more nearly square (less rounded), and 
inner side more flattened and much less convex than in kenaiensis. 

In the females the last lower molar is conspicuously smaller than in 
kenaiensis, and the last upper molar is smaller, narrower, more wedge- 
shape, and more pointed posteriorly. In one of the males it is simi- 
lar. In the other three males the last upper molar is larger and less 
acute posteriorly than in the females, and the third cusp on the inner 
side is better developed. 

Skull measurements. — Adult male (type) : Basal length, S r >9 ; occip- 
ito-nasal length, 315; palatal length, 198; zygomatic breadth, 270; 
intcrorbital breadth, 102.5. 

VETULARCTOS, A NEW GENUS RELATED TO URSUS. 

Generic characters. — Skull like that of Ursus. 1 Teeth in the main 
like those of Ursus, but Mj quite different, presenting a broad flat 
squarish grinding surface with suppression of the hypoconid and 
entoconid, and absence of the usual posterior cusp and notch on 
outer side. M- with outer cusps (paracone and metacone) normal ; 
inner cusps (protocone and hypocone) obsolete. Mj with anterior 
part (protoconid and metaconid) normal but rather low; posterior 

1 The type is a young-adult female. Skulls of adult males may show cranial differences. 



132 NORTH AMERICAN FAUNA. lNo.41. 

part occupying more than half of crown, a flat rectangular platform, 
truncate posteriorly, and bordered by a low marginal rim bearing a 
single small cusp on inner side immediately behind the metaconid; 
hypoconid absent ; entoconid obsolete ; outer side of tooth nearly flat, 
without sulcus or reentrant angle and without notch between pro- 
toconid and hypoconid. 

In the reduction or suppression of the inner cusps of M 1 and M a , 
and of both inner and outer cusps of the large posterior moiety of 
M^, together with the absence of a reentrant angle and notch on 
the outer side of M 7 , V etularctos resembles Arctotherium and Trem- 
arctos. The details of these and other teeth in the three genera, 
however, are quite different and ,ihe genera have little in common. 
Nevertheless the resemblances are sufficient to suggest that V etularctos 
may claim a rather ancient line of descent, from which Arctotherium, 
and Tremarctos also arose — a line quite different from the one 
culminating in Ursxis proper. 

Type species. — V etularctos inopinatus. 

VETULARCTOS INOPINATUS sp. NOV. 
Patriakchal Beak 

Type, skull No. 7149 (skin No. 870G), 9 nearly adult, U. S. 
National Museum, from Rendezvous Lake, northeast of Fort Ander- 
son, Mackenzie. Collected June 24, 1864, by R. MacFarlane. Origi- 
nal No. 1979. Teeth practically unworn. 

Color. — General color varying from whitish buff to pale yellowish 
buff (yellowest on back of head and neck), darkening to dull reddish 
brown on ankles, feet, and median line of belly. The pale body color 
covers the entire body from between eyes to base of tail and reaches 
down over thighs and upper part of legs. Muzzle golden brown, 
becoming dull fulvous-brown around eyes; top of head from be- 
tween eyes posteriorly soiled buff; long hairs of cheeks washed with 
buffy; ears pale buffy. Fur everywhere full, soft, and woolly; 
basal fur of upperparts varying from grayish to grayish brown, but 
distal half or more than half, pale buffy, so the animal as a whole 
appears to be buffy whitish. 

Cranial and dental characters. — Skull small, moderately arched 
above; basicranial axis arched, palate strongly arched and slightly 
concave longitudinally. Teeth peculiar, presenting a combination of 
long canines and well-developed cusps for seizing (main cusp of 
PM-* and PM4, outer cusps of M- 1 - and M^ and anterior cusps of M T ) 
with broadly flattened surfaces for crushing (in M- 3 -, M 7 , and M 7 ). 
Incisors, canines, and premolars as in JJrsus; posterior molars 
peculiar, showing a strong tendency toward the suppression of <*usps, 
particularly those of the inner side. Crown of M A with outer cusps 



1918.1 GENUS VETULAECTOS. 133 

(paracone and metacone) normal; inner cusps (protocone and hypo- 
cone — slightly worn) low, scarcely rising above level of median 
part of tooth ; no middle cusplet. Crown of M- 2 - with outer cusps 
(paracone and metacone) normal; inner cusps (protocone and 
hypocone) obsolete, represented by a low ridge scarcely projecting 
above the granular surface of the interior of the crown. Crown of 
My normal, but anterior part higher than posterior. Crown of 
M^ very remarkable (see description of genus). Crown of M, flat, 
with only insignificant marginal thickenings to represent protoconid 
and metaccnid, the flat crown forming a direct continuation of the 
large crushing platform of the preceding tooth. 

/Skull measurements. — Young-adult female (type) : Basal length, 
268; cccipito-nasal length, 255; palatal length, 148; zygomatic 
breadth, 181; interorbital breadth, 70. 

Tooth measurements.— PU± 17 x 14; M*, 22.5 x 17; M^, 35.5 x 19; 
PM T , 14x8; M T ,24.5xl2;M 7 , 27x17; M3, 21x16; upper molariform 
series, 73; upper molars, 59; lower molars, 72; diameter of lower 
canine, 13; upper incisors (series), 40. 



North Arr <:• eai U. S. Dept. Aj 



Plate II. 




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North American Fauna No. 41, U. S. Dept. Agr. Biological Survey. 



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North American Fauna No. 41. U. S. Dept. Agr. Biological Survey. 



Plate IV. 




North American Fauna No. 41, U. S. Dept. Agr. Biological Survey. 



Plate V. 




North American Fauna No. 41. U. S. Dept. Agr. Biological Survey. 



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North American Fauna No. 41, U. S. Dept. Agr. Biological Survey. 



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North American Fauna No. 41, U. S, Dept. Agr. Biological Survey. 



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North American Fauna No. 41, U. S. Dept Agr. Biological Survey. 



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North American Fauna No. 41, U. S. Dept. Agr. Biological Survey. 



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North American Fauna No. 41, U. S. Dept. Agr Biological Survey. 



Plate XIII. 




North American Fauna No. 41, U. S Dept. Agr. Biological Survey 



Plate XIV. 




North American Fauna No. 41, U. S. Dept. Agr. Biological Survey. 



Plate XV. 




North American Fauna No. 41, U. S, Dept. Agr. Biological Survey. 



Plate XVI, 




2 £ 



INDEX. 



[New species in bold-faced type; synonyms in italics; principal references In bold-faced 

figures.) 

A. 

absarokus, 18, 21, 41, 00, CI, C7, 85, 80, 

93-94, 103, 100. 
Acknowledgments. 10-11. 
alascensis. 94-05, 0G, 100, 10S. 110. 
alexandra?, 75. Ill, 114-115, 128, 129. 
andersonl, 83-84. 
apache, 54. 84-85, 80. 
Arctotbei-ium, genus, 132. 
arizona*. 53-54, 54, G4, 00, G7, 85. 
atuarko, 22-23. 

B. 

bairdi. 13. 18, 19-20, 21, 37 (footnote), G3, 

04, 05, 60, 07, 70. 
blsonophiigus, 54, 60, G7-CS. 

C. 

callfornicus. 29-30, 30, 31, 32, 73, 74, 75, 70. 
canadensis, 43, 52-53, 08, 103. 
candescent, 17. 
caurlnus, 0, CS, 69-71. 
cbelan, 57, 58, 58-60, 78. 
chelidonias, 13, 21-22. 
cinereus, 17. 
Classification, 12-13. 
colusus, 30, 32-33, 74. 
Crama! characters, 13. 
crassodon, 90, 01. 
crassos, 90-92, 119. 

cressonus, 111, 112, 112-113, 117, 118, 123, 
124. 

D. 

dalH, 9, 98, 301, 102, 113, 11G-118, 120, 

122, 123, 129. 
Danis, 86. 

Dental characters, 13. 
dnsorgus, 33-34. 



eltonclarki, 41-43, 44, 45. 
creosotes, 55, 56, 57, 58. 
Euarctos, 04. 

eulophus. 9, 6!>. 70, 71-72, 121. 
eximius, 109-110. 



F. 



ferox, 17. 



Geographic distribution, 9-10. 
grisevs, 17. 
Groups, 12-13. 

absarokus. 10. 

alascensis. 94-99. 

arizona?,, 53-76. 

arizona?-utahensis. Gl. 

californicus-kwakiutl, 25. 

chelan. 10. 

dalli, 116-124. 

g.vas, 124-127. 

horria>us, 84-88. 

horribilis, 10. 13. 17-34. 

bylodromus. 77-84. 

innuitus 110-115. 

kenaiensis. 127-131. 

kidderi. 106-110. 

planiceps. 34-53. 

richardsoni. 99-100. 

shoshone, 57. 

shoshone-idahoensis, 73. 

stikeenensis. 88-94. 

tahltanicus, 10, 42. 

townsendi, 115-116. 
gyas, 106, 107, 108, 124-126, 127. 



64S54 — 1S- 



-10 



henshawl. 31, 73, 87-88. 

hoots, 44, 91, 118-119. 

horr,aceu8, 80. 

borriaeus, 7, 35, 36, 61, 62, 84, 85, 85-87, 

87, 104. 
horribllis, 7, 33, 17-19, 19, 20, 41, 54, 64, 

05. Go. 07. 75, 93, 111. 
hylodrumus, 50, 59, 77-78, 81, 82. 

I. 

Idahoensls, 41, 54-65, 57, 58, 64, 65, 60, 72, 

73, 78. 
imperator, 13, 20-21, 21, 22, 34, 41, 00, 
iraplger, 81-82. 
innuitus. 110-112. 
inoplnatus, Vehilarrtos, 132-133. 
insularis, 9, 44-45. 
internationaiis, 103. 

K. 

kadiaki, 126. 

kenaiensis, 114, 115, 118, 124, 127-129, 

130, 131. 
kennerlyl. 61-62, 85. 
kidderi, 10, 106-108, 5 08. 109, 110. 
klamathensis, 29, 33, 72-74, 74, 75. 
kluane, 10, 48, 55, 78, 79-80, 81, 82, 92, 96, 

1)8, 102. 
kwakiutl, 9, 21, 23, 24-25, 26, 27, 28, 45. 

U 

latifrons. 78, 97-99. 

List of species and subspecies, 14-16. 

M. 

macfarlani, 49, 51-52. 

macrodon, 37 (footnote), 38, 38-40, 53. 

maglster, 10, 31, 75-76. 

Material examined, 10-11. 

mendocinensis, 74-75. 

mcrriami, 124. 126. 

middendorffl. 10, 124, 125, 126, 126-127. 

mirabilis. !), 10, 92. 

mirus, 40-41, 42. 

N. 

navaho. 37. 
neglectus, 9, 28-29. 
nelsoni, 34-35, 87. 
nortoni, 24, 25-27. 
nuchek, 113, 117, 122-124, 

O. 

ophrns, 49, 50, 103-104, 105. 
ors'.loides, 46. 46-47, 48. 
orgilos, 42, 44, 45, 45-46, 47, 48, 51. 
oribasus, 56-58. 

P. 

pallasi. 4G. 47. 47-49, 55, 79, 80, 102. 

pellyensls, 82-83, 83. 84. 99. 

prrturbans, 54, 64-65, 6~>. 

pervagor, 24, 25, 58. 50. 68-69, 70, 71, 73. 

phjeonvx. 98. 98, 101-102. 

planiceps, 36. 37. 37-38, 39. 40, 41. 

pulchcllus, 48, 55-56, 56, 58, 80, 83, 102. 

R. 

richardsoni. 83. 99-100, 100. 
rogersl, 54. 65-66, 67. 
riinsrlnsl, 49-50, 50, 51. 
russelli, 100-101. 



135 



136 



NOBTH AMERICAN FAUNA. 



(No. 41. 



taglttalls, 50-51. 

aaribur, 14 (footnote). 

selkirki, 77, 78. 

Sexual differences, 10. 

sheldoni. 00. 115, 129-181. 

shirasi, 0, 1 20-1 22. 

Shoshone, 35, 30, 41, 43, 58, 60-61, 86, 93, 

104. 
sitkensis, 119, 119-120. 
stikeenensis, 9, 10, 28, 43, 44, 88-90, 92. 

T. 

tahltanicus, 9, 40, 41, 42, 43-44, 44, 45, 4G, 

48. 83. 84, 89. 
Technical terms, 11-12. 
texensis. 35, 35-36, 37, 38, 39, 40. 
toklat, 80. 83, 95, 96-97, 97, 98, 102. 
townsendi, 115-116. 

Tremarctos, genus, 132. / 

tularensis, 30-32, 88. 
tundrensis, 95, 108. 

V. 

Ursus, genus, 13, 14-16, 17-131, 181, 132. 
absarokus, 18, 21. 41, 60, 01, 67, 85, 

89, 93-94, 105, 106. 
alascensis, 94-95, 96, 100, 108, 110. 
alexandra;, 75, 111, 114-115, 128, 129. 
andcrsonl, 83-84. 
apache, 54, 84-85, 86. 
arizonae, 53-54, 54, 64, 66, 67, 85. 
atnarko, 22-23. 
bairdi, 13, IS, 19-20, 21, 37 (footnote). 

63, 64. G5, 06, 07, 76. 
blsonophagUH, 54, 66, 67-68. 
californicus, 29-30, 30, 31, 32, 73, 74, 

75. 76. 
canadensis, 43, 52-63, 68, 105. 
candescens, 17. 
eaurlnus, 9, 68, 69-71. 
chelan, 57, 58, 68-60, 78. 
chelidonia*. 13, 21-22. 
cinereus, 17. 
colusus, 30, 32-33, 74. 
crassodon, 90, 01. 
crassus, 90-92, 119. 
cressonus, 111, 112, 112-113, 117, 118, 

123, 124. 
dalli, 9, 98, 101. 102, 113, 116-118, 

120, 122 123, 129. 
dnsorsus, 33-34. 
eltonclarkl. 41-43, 44, 45. 
erfunetes, 55, 56, 57, 58. 
eulophus, 9, 69, 70, 71-72, 121, 
eximius. 109-110. 
ferojr, 17. 
griseus, 17. 

gyas. 106, 107, 108, 124-126, 127, 
henshawi. 31, 73, 87-88. 
hoots, 44, 91, 118-119, 
horriaceu8, 86. 
horriaMis, 7. 35. 36, 61, 62, 84, 85, 

85-87, 87, 104. 
horribilis, 7, 13. 17-19, 19, 20, 41, 54, 

64. 65, 66, 67, 75, 93, 111, 
hylodrorous. 50. 59, 77-78, 81, 82. 
Idahoensis, 41. 54-65, 57, 58, 64, 65, 66, 

72, 73, 78. 
imperator, 13, 20-21, 21, 22, 34, 41, 66. 



Ursus Implgrr, 81-82. 
Innuitus, 110-112. 

insularis '.), 44-45. 

intemationalis, 103. 

kudiaki, 12U. 

kenaiensis, 114, 115, 118, 124, 127-129, 

130. 131. 
kennerlyi, 61-62, 85. 
kidderi, 10, 100-108, 10S, 109, 110. 
klamatbensis, 20, 33, 72-74, 74, 75. 
khiaae. Id, 48, 55, 7S, 79-80, 81, 82, 

1)2. 96. 08, 102. 
kwakiutl, 9, 21, 23. 24-25, 20, 27, 28, 45. 
latifrons, 78, 97-99. 
macfarlanl, 40, 51-52. 
macrodou, 37 (footnote), 38, 39-40, 53. 
magister. 10, 31, 75-76. 
mendocinensls, 74-75. 
merriami, 124, 126. 
mlddendorffi, 10, 124, 125, 126, 126-12S. 
mirabilis, !». 10, 92. 
mirus, 40-41, 42. 
navabo. 37. 
neglectus, 0, 28-29. 
nelsoni, 34-35, 87. 
nortoni, 24, 25-27. 
nuchek, 113, 117, 122-124. 
opnrus, 49, 50. 103-104, 105. 
orgiloldes, 46, 46-47, 4S. 
orgilos, 42 44, 45, 45-46, 47, 48, 51. 
oribaeus, 56-58. 

pallasi, 46, 47, 47-49, 55. 79, 80, 102. 
pellyensls, 82-83, 83, 84, 09. 
pertnrbans, 54, 64-65, 65. 
pervagor, 24, 25, 58, 59, 68-69, 70, 

71, 73. 
pha?onyx, 96, 98, 101-102. 
planleeps, 36, 37. 37-38, SO, 40, 41. 
pulchellus, 48, 55-56, 56, 58, 80, 83, 102 
ricbardsoni, 83. 99-100, 100. 
rogersl, 54, 65-66, 67. 
rungiusi, 49-50, 50, 51. 
russelli. 100-101. 
•agittalis, 50-51. 
saribur, 14 (footnote). 
telkirki, 77, 78. 
sheldoni. 96. 115. 129-131. 
shirasi. 9, 120-122. 
Shoshone, 35, 36, 41, 43, 58, 60-61, 86, 

03. 104. 
sitkensis, 119. 119-120. 
stikeenensis, 9, 10, 28. 43, 44, 88-90,92. 
tahltanicus, 9, 40. 41, 42, 43-44, 44, 45, 

46. 48. 83, 84, 89. 
texensis. 35, 35-36, 37, 38, 39, 40. 
toklat, 80. 83, 95, 95-97, 97, 98, 102. 
townsendi, 115-116. 
tularensis, 30-82, 88. 
tundrensis, 95, 108. 
utahensis, 61, 02-64, 64, 65, 66, 70. 
warburtoni, 22. 27-28. 28. 
washake, 59, 60. 104-106. 
utahensis, 61, 62-64, 64, 65, 66, 76. 

V. 

Yetulareto*, genus 16. 131-183. 
iBoplnattiB, 132-133. 

W. 

warburtoni, i2. 27-28, 28. 
washake, 59, 60, 104-100. 



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4 



U. S. DEPARTMENT OF AGRICULTURE 

BUREAU OF BIOLOGICAL SURVEY 

E. W. NELSON, Chief 



NORTH AMERICAN FAUNA 



TSTo. 42 



[Actual date of publication, October 3, 1917] 




LIFE ZONE INVESTIGATIONS IN WYOMING 



BY 

MERRITT CARY 

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1917 









LETTER OF TRANSMITTAL. 



United States Department or Agriculture, 

Bureau of Biological Survey, 
Washington, D. C, November 23, 1916. 
Sir: I have the honor to transmit for publication as North 
American Fauna No. 42, a report on life zone investigations in 
Wyoming, by Merritt Gary, Assistant Biologist of the Biological 
Survey. The report is based on the results of natural history ex- 
plorations conducted in recent years by Survey field parties in all 
the important physiographic areas of the State. The first section 
characterizes the five transcontinental life zones represented in 
Wyoming, defines their extent and limits, and discusses their eco- 
nomic possibilities. The second consists of notes on the distribution 
and abundance of conspicuous trees and shrubs observed during the 
progress of the survey. Of particular importance in connection 
with this report, as well as with others yet to be made on the dis- 
tribution of the birds and mammals of Wyoming, is the accompany- 
ing map, which shows in detail the extent and boundaries of the 
life zones which traverse the State. 
Respectfully, 

Henry W. Henshaw, 
Chief, Biological Survey. 
Hon. David F. Houston, 

Secretary of Agriculture. 



CONTENTS. 



Introduction 7 

Physiography and climate 9 

Life zones 12 

Upper Sonoran Zone 13 

Great Plains Division 14 

Great Basin Division 17 

Green River Valley 18 

Bed Desert 19 

Wind River Basin 21 

Bighorn Basin 23 

Characteristic species 24 

Mammals 25 

Breeding birds 26 

Reptiles 27 

Amphibians 27 

Plants 28 

Agricultural utility 30 

Transition Zone 31 

Characteristic species 32 

Mammals 34 

Breeding birds 34 

Plants 36 

Agricultural utility : 37 

Canadian Zone 38 

Characteristic species 42 

Mammals 42 

Breeding birds 43 

Plants 44 

Hudsonian Zone 46 

Characteristic species 47 

Mammals 48 

Breeding birds 49 

Plants 49 

Arctic- Alpine Zone 50 

Characteristic species 51 

Mammals 51 

Breeding birds 51 

Plants 52 

Importance of Boreal zones to Wyoming and adjoining areas 52 

Notes on the distribution of conspicuous trees and shrubs 55 

Index 83 

3 



ILLUSTRATIONS. 



PLATES. 

Page. 

Plate I. Map of Wyoming, showing life zones Frontispiece. 

II. Fig. 1. — Chugwater Valley, showing growth of juniper, mountain 
mahogany, and box elder. Fig. 2.— Western edge of Great Plains 
area, showing scattered yellow pine and juniper 16 

III. Fig. 1. — Plains yucca in flower, Chugwater Valley. Fig. 2. — Nearer 

view of same 16 

IV. Fig. 1. — Bluffs along Green River, showing growth of greasewood. 

Fig. 2.— Desert vegetation, Red Desert, chiefly saltbush 24 

V. Fig. 1. — Yellow pine forest, north base of Laramie Peak. Fig. 2. — 

Sagebrush plain, Wind River Basin 24 

VI. Fig. 1. — Sagebrush in Wind River Valley, lower Transition Zone. 

Fig. 2. — Garfield Peak, Rattlesnake Mountains, showing sagebrush . 32 
VII. Fig. 1.— Narrow-leaved cottonwoods and yellow pines, Rawhide 
Butte, edge of Great Plains, lower border of Transition Zone. 
Fig. 2. — Plain at southwestern base of Wind River Range, showing 

rabbit brush 32 

VIII. Fig. 1.— Snake River Valley, Jackson Hole, showing mixed forest con- 
ditions at lower edge of Canadian Zone. Fig. 2. — Heavy Engel- 

mann spruce forest, Wyoming Range 40 

IX. Fig. 1. — Sylvan Lake, Yellowstone National Park, in Engelmann 
spruce belt. Fig. 2. — Heavy stand of lodgepole pine, north slope 

of Ferris Mountains 40 

X. Fig. 1.— Engelmann spruces, timberline on Whirlwind Peak, Absaroka 

Range. Fig. 2. — White-barked pines, same locality 48 

XI. Fig. 1.— Lower edge of Hudsonian Zone, Wind River Range south of 
Fremont Peak. Fig. 2.— Timberline, east slope of Needle Moun- 
tain, showing growth of spruce, fir, and willow 48 

XII. Fig. 1. — Arctic-Alpine Zone, Wind River Range south of Fremont 

Peak. Fig. 2. — Arctic-Alpine Zone, Absaroka Range 48 

XIII. Fig. 1.-— Teton Range, Mount Moran south to Grand Teton. Fig. 2. — 

Snow in lower Hudsonian Zone, east slope of Bridger Peak, Sierra 
Madre, July, 1911 52 

XIV. Fig. 1.— Heavy forest of Engelmann spruce, Wyoming Range. Fig. 

2. — Rank vegetation, Canadian Zone forest floor, Absaroka Range. . 52 
XV. Fig. 1.— Mountain mahogany, west slope of Bighorn Mountains. 

Fig. 2. — Nearer view of same 68 

TEXT FIGURES. 

Fig. 1 . Map of Wyoming showing routes and collecting localities of Merritt Cary 
and other members of the Biological Survey, mainly from 1909 to 
1915 9 

2. Irrigation canal traversing original sage plain, Bighorn Basin 25 

3. Forest of white-barked pine just below timberline, Absaroka Range. . 55 

5 



6 ILLUSTRATIONS. 

Page. 

Fig. 4. Forest of Engelmann spruce, Ferris Mountains 56 

5. Fringe of blue spruce, base of Wind River Range 57 

6. Forest of Douglas spruce, Ferris Mountains 58 

7. Grove of aspen poplar, base of Laramie Peak 60 

8. Large narrow-leaved cottonwood, upper Wind River Valley 61 

9. Copses of gray-leaved willow at timberline, Absaroka Range 62 

10. Thicket of canoe birch, Bear Lodge Mountains 64 

11. Clumps of mountain birch, base of Shirley Mountains 65 

12. Fringe of alder on Pacific Creek 66 

13. Flowering raspberry, base of Casper Mountains 70 

14. Mountain balm in flower, Sierra Madre 74 

15. Canadian buffaloberry in aspen woods, Wyoming Range 75 

16. Red elderberry in bloom, Sierra/Madre 77 

17. Black sagebrush, upper Wind River Valley 80 






No. 42. NOKTH AMERICAN FAUNA. Oct., 1917. 

LIFE ZONE INVESTIGATIONS IN 
WYOMING. 



By Meeritt Caky. 



INTRODUCTION. 

Wyoming is among the foremost of our States in its wealth of 
natural scenery, culminating in the grandeur of Yellowstone Na- 
tional Park, one of the wonders of the world. In addition to this 
distinction it posseses vast open plains and lofty mountains whence 
flow the headwaters of mighty river systems emptying far away 
to the west into the Pacific Ocean, to the southeast into the Gulf 
of Mexico, and to the southwest into the Gulf of California. The 
various slope exposures of its mountain ranges, the fertility of its 
intervening valleys or basins, and the aridity of its desert spaces 
present a study of geographic and vertical distribution of wild 
life that is in many particulars unique. 

The study of geographic and vertical distribution of life with 
the governing factors and attendant problems is valuable as a matter 
of scientific research and in the attainment of practical knowledge. 
The Biological Survey has been making detailed investigations of 
the transcontinental life belts, or zones, of North America for some 
years, and this work has been carried on with special reference to 
their practical value. It has become increasingly evident that life 
zones furnish a fairly accurate index to average climatic conditions 
and, therefore, are useful as marking the limits of agricultural 
possibilities, so far as these are dependent upon climate. The 
knowledge thus gained has been published and made available as 
the investigations have progressed and the life zones have been 
mapped. 1 

The opening up to agriculture of the arid and semiarid West 
through irrigation and efficient methods of conserving the natural 

*For detailed discussion and classification of the life zones of North America see 
Merriam, C. Hart, Life Zones and Crop Zones of the United States (Bull. 10, Biological 
Survey, U. S. Dept. Agr., 1898) ; also Bailey, Vernon, Biological Survey of Texas (North 
Amer. Fauna No. 25, 1905) ; Fourth Provisional Zone Map of North America, prepared 
by the Biological Survey, 1910 ; Cary, Merritt, Biological Survey of Colorado (North 
Amer. Fauna No. 33, 1911) ; and Bailey, Vernon, Life Zones and CroD Zones of New 
Mexico (North Amer. Fauna No. 35, 1913). 

7 



8 NORTH AMERICAN FAUNA. [No. 42. 

rainfall offers a favorable field for the practical application of this 
knowledge. A wide range of altitude and a correspondingly varied 
climate and physiography include from two to six of the major life 
zones in each of the several States, and the zonal boundaries are 
on the wdiole well marked by reason of the usually rapid or abrupt 
changes in elevation. New areas are continually being reclaimed. 
while in practically all the Western States large districts await 
future development. This is especially true of the Rocky Mountain 
States, where general agriculture has been least developed. 

In Wyoming, agriculture has made rapid strides during the past 
few years, but it has not yet advanced much beyond the experimental 
stage, and the possibilities are somewhat limited by a cool climate 
due to high average base level. It appears unlikely that crop pro- 
duction will in future greatly exceed the local demand. There are, 
however, certain restricted areas of low elevation and moderate 
climate where a variety of crops and some of the hardier fruits have 
proved decidedly successful. A special value attaches to these low- 
lying districts inasmuch as they are immediately surrounded or 
bordered by extensive nonagricultural areas where mining, lumber- 
ing, and stock raising are the principal industries. The melting 
snows of Wyoming mountains furnish an unfailing supply of water 
for irrigation purposes, and Federal and private irrigation projects 
have already reclaimed considerable sections. Much valuable agri- 
cultural land in the valleys and basins awaits future development. 
Useless experimentation might be avoided or a more favorable loca- 
tion secured if the prospective as well as the resident agriculturist, 
and especially the horticulturist, would become familiar with the 
groups of native species of mammals, birds, reptiles, and plants 
which have proved to be closely associated with the areas of success- 
ful production of particular crops in other parts of the arid West. 

Natural history explorations carried on in recent years by the 
Biological Survey in all the important physiographic areas of Wyo- 
ming warrant the present report on the life zones with the accom- 
pan} T ing map (PI. I). Sufficient material has been gathered also 
for inclusion of notes on the distribution of conspicuous trees and 
shrubs and for later reports on the mammals and birds of the State. 1 

*The present report combines the results of field investigations for the Biological Sur- 
vey conducted at various times by Dr. C. Hart Mcrriam, Vernon Bailey, B. II. Dutcher, 
J. Alden Loring, Edward A. Preble, Alexander Wetmore, II. E. Anthony, Stanley G. 
Jewett, and D. D. Streeter, jr. ; besides those made by the author since 1909 (see map 
of Wyoming, showing routes of field parties, fig. 1). Lists and other publications bearing 
cm the distribution of the Wyoming fauna and flora, although few in number, have been 
freely consulted. For identification of many of the plants collected in the survey the 
author is indebted to Dr. J. N. Rose and Paul C. Standley, of the F. S. National 
Herbarium, and to F. V. Coville, curator of the National Herbarium, who has named the 
Ribes. The few reptiles and amphibians collected have been identified by Dr Leonhard 
Stejneger, of the U. S. National Museum. 



1917.] 



PHYSIOGRAPHY AND CLIMATE. 



PHYSIOGRAPHY AND CLIMATE. 

In common with other States of the Rocky Mountain region, 
Wyoming has a varied physiography and climate and great natural 
resources. The surface features may be classified broadly as moun- 
tains, plains, and valleys or basins. 

The continental watershed formed by the main chain of the 
Rockies enters the State through Yellowstone Park near the mid- 
western boundary of the forested plateau and maintains a general 
southeasterly trend along the lofty crests of the Absaroka and Wind 
River Ranges, lowering in the Red Desert region to arid plains and 




■ &4CK, HtfGOtySWO SK4G£- 7XY/RS 



•ftt/ufo^o ^/ou/^ers. 



+ COlL£Cr/A/G £OCy4l/rK 



Fig. 1. — Map of Wyoming showing routes and collecting localities of Merritt Cary and 
other members of the Biological Survey, mainly from 1909 to 1915. 

alkaline basins at 7,000 feet elevation, but again rising to the sum- 
mits of the Sierra Madre, where it crosses the southern boundary 
of the State near its middle point. This watershed divides the 
Great Plains from the Great Basin, and on its slopes in northwest 
Wyoming rise the headwaters of the Colorado in Green River, the 
Columbia in Snake River, and the Missouri in Yellowstone and 
Madison Rivers. 

The mountains of Wyoming, massed largely in the northwest, 
occupy approximately a fourth of the total area. Chief among them 
are the Absaroka, Wind River, Gros Ventre, and Teton Ranges in 



10 NORTH AMERICAN FAUNA. [No. 42. 

the northwest; the Bighorn Mountains in the central northern por- 
tion; and the Sierra Madre and the Medicine Bow Ranges at the 
south. Most of these are heavily forested groups of great elevation, 
whose summits and crests reach far above timberline and are 
usually snow capped even in midsummer. All belong to the Rocky 
Mountain system except the Bighorn Mountains, which alone are 
detached from the main chain. Gannett Peak in the Wind River 
Mountains, at 13,785 feet, is the highest point in the State, exceeding 
the height of its close neighbor, Fremont Peak (13,730 feet) ; and of 
the Grand Teton (13,717 feet) in the Teton Range. 

The Ferris, Green, Seminole, Shirley, and Rattlesnake Ranges are 
small separated groups along the upper Platte and Sweetwater 
Valleys and in the northern borders of the Red Desert, lying a lit- 
tle east of the continental watershed but indicating the general 
course of the Rockies. These differ greatly in configuration, but 
are usually characterized by densely forested northern slopes and 
summits and abrupt barren southern exposures facing the desert. 
In this region also are the huge bare granite heaps and domes rising 
from the sandy plain north of the Sweetwater, known collectively 
as the Granite Mountains. Together, these small ranges are a pleas- 
ant relief to a generally barren landscape. 

The southwest corner of Wyoming is much broken by the northern 
timbered shoulders of the Uinta Mountains and by barren elevated 
ridges and mesas on either side of Green River and east of Bear 
River. In the southeast the Laramie Mountains are a foothill spur 
of the Medicine Bows continuous north to the Laramie River Gap 
and thence in more broken formation extending to the Platte Val- 
ley near Casper. Conspicuous groups of outlying hills at the east 
and northeast are the pine-clad Hartville Mountains north of Guern- 
sey and the densely forested Black Hills and Bear Lodge Mountains. 

Although well supplied with mountains, Wyoming is perhaps bet- 
ter known for its vast open plains. These are either level or rolling, 
lying mainly between 4,500 and 7,000 feet elevation, and are dis- 
tinguished by characteristic types of vegetation, as the sage plains of 
the high, arid, interior plateaus; and the grassy plains to the east 
and northeast, which are part of the Great Plains. These treeless 
expanses were ranged long before historic times by great bands of 
buffalo, and, succeeding these, by countless herds of cattle and sheep, 
and their great grazing value is well attested by the long and bitter 
warfare for their possession between cattle barons and flockmasters 
which marked the days of the open range. At present dry farming 
is greatly restricting the cattle range on the Cheyenne and Lusk 
Plains and elsewhere along the eastern edge of the State. The ex- 
tensive arid sage plains farther west, however, are mainly utilized 
for sheep grazing, to which they are peculiarly adapted. 



1917.] PHYSIOGRAPHY AND CLIMATE. 11 

The numerous valleys of Wyoming are "well watered, and with 
their rich soils and low elevations (chiefly below 5,000 feet) include 
the areas of greatest agricultural importance and promise. Most 
important are those of the North Platte, Laramie, Cheyenne, Belle 
Fourche, Powder, Bighorn, Wind, Sweetwater, and Green Rivers. 1 
The Bighorn and Wind River Valleys are extensive basins of low 
altitude and mild climate, well suited to the production of certain 
fruits and other crops. The more elevated valleys of the Sweetwater 
and Green Rivers are mainly devoted to stock raising. 

The Red Desert is an extensive barren alkaline plain or basin of 
great aridity lying mainly west of the continental watershed in the 
southern part of the State. Without perennial streams and with 
soils strongly alkaline, it would appear to have no agricultural 
future. Alkali-resistant desert shrubbery and the moderate winter 
climate of this region nevertheless combine to furnish an excellent 
winter range for sheep, and it has long been thus utilized by flock- 
masters. 

Wyoming's lowest elevation is in the extreme northeast, and its 
highest is in the northwest. Plains and plateaus occupy much of its 
southern half. The altitudinal extremes are 3,100 feet (approxi- 
mate), where the Belle Fourche River crosses the eastern boundary; 
and 13,785 feet, on the summit of Gannett Peak in the Wind River 
Range. 

The climate of the State is mainly arid, the rainfall ranging from 
12 to 15 inches in the semiarid eastern Great Plains area to under 
10 inches in the extreme arid central desert region (Bighorn Basin 2 
and Red Desert). A heavier precipitation in the Bear Lodge and 
Black Hills districts at the northeast (15 to 20 inches) admits of 
tolerably successful agriculture without irrigation. The high tim- 
bered mountain ranges receive a great deal of moisture, not only as 
winter snows, but also during summer as frequent heavy, dashing 
rains. 3 

The elevated base level of Wyoming (about 6,000 feet) insures a 
generally cool climate. Warm summers (mean summer temperature 
about 65° F.) with a long growing season and moderate winters 
with light snowfall are the rule only at the lower levels in the north 
and east. The high interior valleys, plains, and plateaus are marked 
by short, cool summers (mean summer temperature about 55° F.), 
with prevalent late spring frosts, and by long winters with tolerably 
heavy snowfall and frequent cold winds. The snowfall is excessive 
in the mountainous country of the northwest, where occasionally very 

1 Most of the valleys are treated in some detail under their respective zones. 

2 The lowest parts of the Bighorn Basin often receive less than G inches of annual 
rainfall. 

3 The precipitation usually given for the mountains is over 18 inches, but data are 
lacking for the higher altitudes, where it must be much greater. 



12 NORTH AMERICAN FAUNA. [No. 42. 

low temperatures are recorded, 1 but the winter season as a whole is 
perhaps less severe than on the high wind-swept plains. 

For a State with an arid climate Wyoming is exceptionally well 
watered, and among its natural resources none is more essential to 
its future development than its rivers and streams. The Snake, 
Yellowstone, Bighorn, and Green Rivers rise in the mountains of 
the northwest; the Tongue, Powder, Belle Fourche, and Cheyenne 
Rivers, with their numerous tributaries, head in the Bighorn Moun- 
tains and the elevations of the northeast ; while the North Platte and 
Laramie Rivers, which describe long, circuitous courses in the south- 
eastern part of the State, have their sources in the high ranges of 
Colorado. 

LIFE ZONES OF WYOMING. 

Wyoming has a generous representation of animal and plant life. 
This is largely due to the varied climate resulting from a difference 
in altitude within its borders of nearly 10,700 feet; and in a lesser 
degree to a difference in latitude of 4 degrees, and a wide range of 
local physiographic conditions. 

The life zones range from Upper Sonoran (the western arid 
subdivision of the Upper Austral Zone) at the lowest and warmest 
elevations, through the Transition, Canadian, and Hudsonian, to the 
Arctic-Alpine Zone on the crests of the highest mountain ranges. 
Of the seven North American transcontinental life zones, only the 
Lower Sonoran and the Tropical are unrepresented ; and the Upper 
Sonoran Zone, while covering large areas, is represented only by its 
upper, cooler part. 

The five zones present in Wyoming are briefly characterized as 
follows: Upper Sonoran,, the zone of broad-leaved cottonwood, 
juniper, saltbush, and yucca, occupying most of the valleys and low 
plains; Transition, the zone of yellow pine, narrow-leaved cotton- 
wood, and pure sagebrush, embracing the high plains, the basal 
slopes of the mountains, and all except the highest foothills; Cana- 
dian, the Boreal forest belt of spruce, fir, lodgepole pine, and aspen, 
covering the middle mountain slopes and highest foothill ranges; 
Hudsonian, the narrow zone or belt of white-barked pine and 
dwarfed spruce and fir, in the timberline region; and Arctic -Alpine, 
the treeless zone, on mountain crests and peaks above timberline. 

Zonal boundaries and sequence usually are well marked on ranges 
rising abruptly from a low base, as on the western slope of the 
Bighorn Mountains, where a vertical interval of about 9,500 feet 
may be traversed in 15 or 20 miles; and on the Wind River Range 
southwest of Lander. Under gradual change in altitude, however, 

* — 45° F. is sometimes recorded in Jackson Hole, and though data are lacking, still 
lower temperatures undoubtedly are reached on the high ranges. 



1917.] UPPEE SONORAN ZONE. 13 

as on the open plains, the passage from the Upper Sonoran to the 
Transition Zone is often scarcely noticeable, and in mapping zones 
generally the limits in many places must be more or less arbi- 
trarily fixed. In a survey of native mammals, birds, reptiles, and 
plants made at a given locality, a marked preponderance of charac- 
teristic species of a zone decides the zonal position ; while a nearly 
equal representation, or a marked absence, of species peculiar to two 
adjoining zones is indicative of an intermediate position, or the ap- 
proximate boundary. 

The several zones occupy areas of great irregularity in "Wyoming 
owing to the very broken configuration, and their vertical boundaries 
are subject also to the usual variation resulting chiefly from differ- 
ences in latitude, base level, and slope exposure. In general, zone 
levels are lowest in the north because of lower base and higher 
latitude, and highest in the south where the base level is more ele- 
vated. 1 

UPPER SONORAN ZONE. 

The arid subdivision of the Upper Austral Zone, the Upper 
Sonoran, occupies nearly a third of Wyoming (about 30,000 square 
miles), and covers all the lower levels. Fully half this area lies 
below 5,000 feet altitude in the eastern and northern sections, and 
most of it is well characterized. On the high plains and deserts of 
the south the Sonoran element is present between 5,500 and 6,500 feet 
elevation, mainly in dilute form. Low altitude, a warm climate and 
long growing season, and extensive open, level, or gently rolling 
areas of rich soils combine to make the Upper Sonoran the chief 
zone of crop production, dependent as in other sections of the arid 
West upon careful conservation and distribution of the natural water 
supply. All areas adapted to any extent to horticulture lie within 
this zone and, because of its agricultural importance, the limits and 
characterization are somewhat detailed. 1 

The Upper Sonoran areas of Wyoming are mainly broad exten- 
sions of the zone from lower elevations on the south, east, and north. 
Those entering from the south comprise a narrow strip of desert valley 
along both sides of Green River north nearly to Labarge; and in 
the Red Desert a broad region of barren plains and alkaline de- 
pressions which reaches eastward to include a small area in the 
upper Platte Valley both north and south of Fort Steele. The above 
areas are narrowly connected along Bitter Creek, and more broadly 
over the Snake River Valley in northwest Colorado, and together 
they form the northernmost extensions of the important Green River 
Upper Sonoran area. 

1 For detailed boundaries of the zonal areas of Wyoming see the zone map (frontis- 
piece). 



14 NORTH AMERICAN FAUNA. [No. 42. 

Two large tracts of Upper Sonoran country in the east are part 
of the Great Plains area. The southernmost of these is approxi- 
mately bounded on the west by the 5,000 to 6,000 foot basal plains 
flanking the Laramie and Hartville Mountains, and extends north 
to the narrow strip of Transition Zone along the northern escarp- 
ment of the Lusk Plains. In the North Platte Valley the zone con- 
tinues narrowly through the canyons above Guernsey, then in greater 
width to Casper, and in dilute character to the Seminole Mountains 
and to Splitroek in the Sweetwater Valley. North of the Lusk Plains 
the Upper Sonoran Zone includes most of the Cheyenne River 
drainage lying between the pure sage plains of the northern central 
section and the yellow pine country of the Black Hills. The Belle 
Fourche Valley in the northeast also carries a narrow Upper Sonoran 
strip around the northern and western bases of the Bear Lodge Moun- 
tains, which widens above Moorcroft and extends nearly to the Pump- 
kin Buttes. 

A broad band of this zone entering the State from the north along 
the Bighorn River and Clarks Fork covers a large extent of low- 
altitude country in the arid Bighorn and Wind River Basins below 
5,500 or 6,000 feet elevation. These two areas, of which the Bighorn 
Basin is the larger and agriculturally the more important, are nar- 
rowly connected through the canyon south of Thermopolis, but are 
otherwise separated by the elevated Transition Zone ridge of the Owl 
Creek Mountains. East of the Bighorn Mountains, the valleys of 
the Tongue, Powder, Little Powder, and Little Missouri Rivers carry 
narrow tongues of Upper Sonoran Zone some distance into the State, 
separated more or less widely by low, open or pine-clad Transition 
Zone watersheds. 

In Wyoming, as in other States traversed by the continental 
watershed, the Upper Sonoran Zone is best treated under its two 
main subdivisions, the Great Plains and the Great Basin. 

Great Plains Division — Upper Sonoran Zone. 

Great uniformity of surface features and characterization marks 
the Great Plains area from the Dakotas to Texas and west to the 
eastern foothills of the Rocky Mountains. It is a vast level or undu- 
lating region of abundant grasses and moderate rainfall, entirely open 
except along streams, which are usually fringed with deciduous trees 
and shrubbery, and also in the rougher parts near the foothills where 

1 The subdivisions are based upon differences in climate, configuration, and native 
species, and are not to be confused with the physical Great Plains and Great Basin 
areas as generally understood and with which they are not coextensive. The imperfect 
characterization of the Great Basin division in Wyoming, particularly as regards its 
mammal and bird life, is due to high altitude, and in the Red Desert region in part to the 
iu fusion of Great Plains species as a result of the continuity of the Sonoran areas on 
either slope of the Continental Divide. 



1917.] UPPER SONORAN ZONE. 15 

junipers and pines often occur in scattered growth. Areas of firm 
soils alternate with tracts of sandhills or rolling sandy country, and 
as the foothills are approached there are scattered areas of rough 
bad lands, the bluffs along streams become rocky, while numerous 
talus ridges, and clay, chalk, or rock buttes of usually irregular but 
in some places of strikingly symmetrical outline, stand up from the 
plain (PL II). The important streams of the Great Plains have 
their sources in the Rocky Mountains and course through val- 
leys usually shallow and often sandy, which show little erosion. This 
type of country is well marked along the eastern edge of Wyoming 
as far north as Lusk. 

The greatest elevation of the Upper Sonoran Zone is in the south, 
where on the firm-soil plains of the Cheyenne region it becomes dilute 
at 5,500 feet elevation, and its upper limits are indicated at approxi- 
mately 6,000 feet, chiefly by absence of characteristic species. Exact 
delimitation is difficult on these open grassy plains, which extend 
on a gradual incline to the 7,000-foot base of the Laramie Moun- 
tains. The Chej^enne Plains descend steadily northward to the 
Platte Valley at a little over 4,000 feet, the surface meanwhile be- 
coming broken and the soil sandy. The Sonoran element is very 
pronounced in the warm valleys of the Platte drainage, as in the 
Chugwater Valley below Bordeaux ; in the Laramie Valley at Wheat- 
land, Jetsam, and Uva ; in Rawhide Valley below Patrick ; in Goshen 
Hole; and along the North Platte below Guernsey; and agriculture 
under irrigation in these districts is correspondingly successful and 
varied. 

North of the Platte Valley, and jutting squarely against the eastern 
bases of the Hartville Mountains and the Rawhide Butte, is an exten- 
sive grassy plateau with an elevation of from 4,800 to 5,000 feet, 
extending east into Nebraska and breaking sharply at the north 
toward the Cheyenne River. Upper Sonoran species predominate 
up to 5,000 feet, but the region is near the upper edge of the zone, 
since rocky buttes, ridges, gulches, and cool northern declivities 
carry a scrubby growth of yellow pine, Rocky Mountain juniper, 
red currant, mountain mahogany, and other Transition Zone vege- 
tation. This plateau, often known as the Lusk Plains, is character- 
ized mainly by its luxuriant growth of nutritious grasses a>nd has 
long been noted as choice cattle range. Extensive areas on both the 
Cheyenne and Lusk Plains are now utilized in dry farming, to which 
the soil and climate are well adapted. 

Characteristic associations of Upper Sonoran species mark the 
lower portions of the Cheyenne and Lusk Plains and the North 
Platte Valley. Large groves of broad-leaved cottonwoods (chiefly 
Populus occidentalis) are on the North Platte and Laramie Rivers 



16 NORTH AMERICAN- FAUNA. [No. 42. 

and especially on Sibylee Creek, while in addition to cottonwoods 
the usual fringe along streams consists of willows, box elder, ash, 1 
flowering currant, and wolf berry. Common shrubs or shrubby plants 
on dry flats, in gulches, and on rocky or gravelly, slopes are saltbush, 
rabbit brush, narrow-leaved sagebrush, yucca (PI. Ill), bush morn- 
ing-glory, sand cherry, and skunk bush. 

Some of the most conspicuous and characteristic flowering plants 
are Eriogonwm annuum, Eumex venosus, Abronia elliptica, Arge- 
mone intermedia, Cleome serrulata, Polanisia trachysperma, Luplnus 
plattensis, Astragalus crassicarpus and A. mollissimus, Psoralea 
(spp.), P etalostemon (spp.), Linum rigidum, Croton texensis, Ment- 
zelia decapetala, Opuntia polyacantha, Marmllaria vimpara and M. 
missouriensis, Anogra albicaulis, Meriolix serrulata, Lithospermum 
gmelini, Llppla cuneifolla, Verbena hastata and V. bracteosa, Phy- 
salis lanceolata, Solanum rostratum, Pentstemon angustifolius, Plan- 
tago purshl, Liatrls punctata, Grindclia squarrosa, Ratlbida colum- 
narts, Ilelianthus annuus and H. petiolarls, Hymenopappus ftlifolius, 
Carduus plattensis, and Lygodesmia rostrata. 

Mammals which especially mark this region as Upper Sonoran 
are the Kennicott ground squirrel, prairie-dog, Great Plains grass- 
hopper mouse, prairie harvest mouse, Colorado bushy-tailed wood 
rat, Hayden field mouse, yellow pocket gopher, sage pocket gopher, 
Wyoming kangaroo rat, Kansas pocket mouse, Bailey cottontail, 
black-footed ferret, northern plains mole, and California bat. 

Characteristic breeding birds of the plains are the mourning dove, 
burrowing owl, Arkansas kingbird, Bullock oriole, bronzed grackle, 
lazuli bunting, lark bunting, western grasshopper sparrow, western 
lark sparrow, white-rumped shrike, yellow warbler, long-tailed chat, 
western mockingbird, catbird, and brown thrasher. 

Reptiles are poorly represented on the plains of eastern Wyoming. 
The few conspicuous snakes and lizards include the plains rattle- 
snake, prairie bull snake, hog-nosed snake, blue racer, garter snakes, 
desert horned lizard, sand swift, scaly lizard, six-lined lizard, and 
many-lined skink. 

The Upper Sonoran area north of the Lusk Plains is open, but 
much rougher in configuration, and the shrubby type of vegetation 
becomes increasingly prominent, especially in the valleys below 
4,500 feet elevation, where the Sonoran element is strongest. Grass 
and cactus flats alternate with tracts of sagebrush' rabbit «brush, and 
greasewood over much of the drainage basin of Cheyenne River, and 
in the low Belle Fourche and Little Missouri Valleys in northeast 
Wyoming. The watersheds between- the valleys are either ranges 
of rolling grassy hills or abrupt barren ridges of bad lands of about 

1 Uncommon. 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate II. 




Fig. 1.— Chuqwater Valley below Chugwater. 
Juniper and mountain mahogany ( Cercocarpus parvifolius) on bluffs, and box elders in valley. 




Fig. 2.— Western Edge of Great Plains Area. 
Scattered yellow pines and junipers in butte country southwest of Guernsey (4,800 feet). 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate III. 




Fig. 1 .—Plains Yucca (Yucca glauca) in Flower in Chuqwater Valley near 
Bordeaux, July 1, 1909. 




Fig. 2.— Nearer View of Same. 



1917.] UPPER SONORAN ZONE. 17 

5,000 feet elevation. Extensive breaks with more or less yellow pine, 
juniper, and Transition Zone shrubbery margin the Cheyenne River 
basin, especially on the south, while the watercourses usually are 
bordered narrowly with gnarled cottonwoods, willows, box elders, 
and other deciduous trees and shrubs, as along plains streams gen- 
erally. 

Sagebrush country rapidly takes the place of grassy plains west of 
Lusk, varied in the upper valley of the Platte with greasewood, 
rabbit brush, saltbush, and other desert shrubbery, which becomes 
increasingly common westward. Shrubby Sonoran vegetation fills 
also most of the narrow extensions of the Yellowstone Valley Upper 
Sonoran Zone in the valleys of the Little Powder, Powder, and 
Tongue Rivers east of the Bighorn Mountains. 

Although the shrubby type of desert vegetation, more barren 
surface, and greater aridity of the upper Platte Valley, and the 
Upper Sonoran areas between the Black Hills and Bighorn Moun- 
tains, would seem to place them with the Great Basin division of 
the zone, their mammals, birds, and herbaceous plants are mainly 
those of the Great Plains. Species which are absent or rare on the 
plains farther south include among plants the greasewood, black 
sagebrush, several species of rabbit brush, white sage, saltbushes, 
and stanleya; and among mammals, the Maximilian and Sweet- 
water pocket mice. 1 

Upper Sonoran areas of eastern Wyoming are within the semi- 
arid region and receive in most sections a mean annual rainfall of 
from 12 to 15 inches. This is sufficient for a luxuriant growth of 
the best varieties of range grasses in all sections with suitable soils, 
but will not admit of agriculture apart from irrigation or dry- 
farming methods, except possibly around the base of the Black 
Hills, where the precipitation is a little greater. Under irrigation 
the rich alluvial soils of the valleys yield abundant crops, while the 
moderate returns from the soil secured most years in the dry- 
farming communities scattered over the plains are inducing a 
steadily increasing settlement of these districts. 

Great Basin Division — Upper Sonoran Zone. 

The Green River Valley, Red Desert, and the Bighorn and Wind 
River Basins have a barren surface which shows everywhere much 
erosion, especially along the larger streams, which in many places 
flow through rugged canyons. These open interior areas of from 
4 ; 000 to 6,500 feet elevation lie within the arid region of slight 
rainfall, and the scanty vegetation is of the shrubby, bunchlike, 

1 A number of mammals and birds and a few plants found commonly on the Lusk Plains 
and southward do not reach this region. 

74440°— 17 2 



18 NORTH AMERICAN FAUNA. [No. 42. 

desert type so characteristic of the Great Basin region as a whole 
(PI. IV, fig. 2), in marked contrast with the grassy plains of eastern 
Wyoming. A narrow fringe of scattering junipers usually marks 
the upper border of the zone on the rough lower margins of these 
desert valleys and basins, while considerable areas -of junipers with 
scattering pinyons partly fill the upper part of the zone in lower 
Green River Valley, and in the extreme southwestern borders of the 
Red Desert. 1 The Great Basin division is better characterized in 
Wyoming by vegetation and climate than as a faunal region, 2 al- 
though a few Great Basin species of both mammals and breeding 
birds occur in the Green River Valley and on the Red Desert. The 
mammals and birds of the Wind River and Bighorn Basins are 
mainly those common to the Great Plains. 

The great dearth of rainfall in these desert areas precludes ordi- 
nary agriculture and even dry farming. Effective agriculture is 
therefore possible through irrigation alone. 

Green River Valley — Upper Sonoran Zone. 

The Upper Sonoran area extending north along Green River nearly 
to Labarge is considered apart from the Red Desert, with which it 
is connected, on account of different topographic features and a 
stronger characterization in its lower part, near the Utah boundary, 
where the elevation is only 5,800 feet in the river valley. In this 
section it is a rough incised region of rocky, juniper-clad ridges and 
dry, open canyons or narrow valleys reaching gradually down to 
Green River from high, bordering Transition Zone hills and mesas; 
and of precipitous, brightly colored canyons whose various shades 
of red and yellow are in strong contrast with the intense black of the 
juniper ridges. The steep, hot, protected slopes of this broken can- 
yon region carry the Upper Sonoran Zone regularly as high as 7,000 
feet. 

Its species are characteristic of the juniper and pinyon belt of the 
Great Basin, but are in less variety than in this part of the zone 
farther south, and some of them reach but a short distance into the 
State. Junipers, scattering pinyons, mountain mahogany, shadscale, 
syringa, and cactuses comprise the principal vegetation on ridges and 
dry slopes ; saltbushes, grayia, yellow cleome, eriogonum, and cactuses 
are common shrubs and plants on the open sand or adobe flats in the 
valleys between the ridges; the skunk bush and flowering currant 
form characteristic shrubbery on the bluffs immediately along Green 
River ; while extensive flats in the wider parts of the river valley are 

1 The belt of junipers and pinyons, which in regions farther south is usually present in 
good width and characterizes the upper part of this division of the zone, is but poorly 
indicated in Wyoming, where the zone is more often open throughout. 

2 See footnote, p. 14, 



1917.] UPPER SONORAN ZONE. 19 

densely covered with grease wood (PL IV, fig. 1) and glasswort in 
damp, alkaline spots. ' 

The gray titmouse, Baird wren, and piny on jay are common birds 
in the juniper growth, and probably breed, as does also the sage spar- 
row, on the greasewood flats. Among the characteristic Upper So- 
noran mammals of this region are the least and rock chipmunks, 
golden-breasted canyon mouse, True's cliff mouse, Green River pocket 
gopher, and Great Basin spotted skunk. The reptiles and amphibians 
include the plains rattlesnake, scaly rock lizard, Stansbury sand liz- 
ard, short-horned lizard, and the spadefoot toad. 

North of the canyon region, which extends to within 15 or 20 
miles of the town of Green River, the Upper Sonoran Zone spreads 
out in dilute form to include the lower valleys of the Black Fork and 
Big Sandy, connecting eastward through the Bitter Creek drainage 
with the larger area on the Red Desert. 1 As a narrowing strip along 
Green River it continues to the warm valley flats between Fontenelle 
and Labarge. This is an open, deeply eroded region of barren val- 
leys and bench lands, of bare mesas and variously colored bad lands, 
buttes, and bluffs, whose zonal position is best defined by its conspicu- 
ous vegetation, chiefly that common to the Red Desert. Upper 
Sonoran species of mammals and breeding birds are comparatively 
few in number in this high part of the zone, where saltbushes, grayia, 
small brown sagebrush, rabbit brush, and other desert shrubbery, 
and such plants as yellow cleome and stanleya give way to the pure 
sage and rabbit brush plains of the Transition Zone at about 6,500 
feet elevation. 

The Green River Valley is rather bleak and inhospitable and does 
not seem to invite agricultural development. In character it resem- 
bles the Red Desert in many ways, but is less intensified, particularly 
in the aridity and the alkalinity of its soils. Owing to erosion, 
there is little valley land, and this little is partly used in the cultiva- 
tion of forage crops. At the north a limited agriculture is possible, 
wherever lands can be brought under irrigation, on the small areas of 
bench lands which lie along Green River and its few permanent side 
streams. The rainfall is insufficient for a good growth of forage 
grasses, yet large numbers of sheep subsist upon the desert shrub- 
bery and scanty grass. Sheep grazing appears to be the most prac- 
ticable industry, and the rough character of lower Green River Val- 
ley, with its many sheltered canyons, warm protected slopes, and 
mild climate, peculiarly fits it for a winter range. 

Red Desert — Upper Sonoran Zone. 

Fairly constant surface features obtain over the open elevated 
region of undulating plains and alkaline depressions or basins, known 

1 See frontispiece (map). 



20 NORTH AMERICAN FAUNA. [No. 42. 

as Red Desert, which is particularly characterized by great aridity, 
saline or other strongly alkaline soils, and dearth of permanent sur- 
face water. The only relief to the general monotony and barrenness 
is in scattering buttes, occasional ranges of low hills, bluffs along dry 
washes, or summits of distant bordering hills and mountains. The 
impression of barrenness is only intensified by the prevalent dull 
greenish gray or occasional light-gray hues of desert shrubbery. 1 
The continental watershed extends across the north-central portion 
at an elevation of less than 7,000 feet where lowest, being crossed by 
the Union Pacific Railroad at Creston at 7,000 feet. Eastward the 
altitude decreases slightly towapd the North Platte Valley (6,500 
feet), and westward slopes through its only conspicuous drainage 
area, Bitter Creek Valley, to 6,000 feet on Green River. The climatic 
features are hot sunny days, cool nights, very slight rainfall during a 
short summer, and a long, moderately open winter. The heaviest 
snows are often in spring and are more beneficial to vegetation than 
summer rains, which are either largely lost through rapid evapora- 
tion or, in the case of the occasional heavier showers, quickly run off 
the barren slopes and fill the dry gulches with muddy torrents. 
Frosts are not uncommon during the height of the growing season. 

Dilute Upper Sonoran Zone, poorly characterized apart from vege- 
tation, covers the lower portion of the Red Desert to 6,500 or 6,800 
feet altitude, a total area of between 5,000 and 6,000 square miles, in- 
cluding the North Platte Valley from the Platte Canyon to above 
Fort Steele. The conspicuous and dominant vegetation is Upper 
Sonoran desert shrubbery — the various species of saltbush, the white 
sage, greasewood, grayia, kochia, rabbit brush, the black sagebrush, 2 
and low desert sages; with scattering desert junipers on many of the 
bluffs. A willow (Salix fluviatilis) is not uncommon on creeks of the 
Bitter Creek drainage. Such plants as prickly-pear cactus, yellow 
cleome, stanleya, many alkali-resistant members of the goosefoot 
family, and scattering grasses, conspicuous among which are wheat 
grasses and giant rye-grass, are abundant and characteristic. 

The small striped ground squirrel, pocket mice, long-eared bat, 
least chipmunk, Green River pocket gopher, kangaroo rat, Bailey 
cottontail, and Great Basin spotted skunk are Upper Sonoran mam- 
mals inhabiting the Red Desert. 

The variety of bird life is very limited, and few characteristically 
Sonoran birds breed here. The sage sparrow and western lark spar- 
row are perhaps most nearly restricted to the zone. Other birds 
found in abundance during the breeding season, the Brewer spar- 

1 The name " Red Desert," originally applied to a restricted area of reddish soil along 
the Union Pacific Railroad, does not convey a correct impression of this region as a 
whole, where desert vegetation instead of soil lends the characteristic colors. 

2 Abundant also in Transition Zone. 



1917. J UPPER SONORAN ZONE. 21 

row, thick-billed redwing, western meadowlark, western nighthawk, 
sage thrasher, and white-rumped shrike, breed also on the surround- 
ing sage plains of the Transition Zone. 

This arid, barren waste is naturally unsuited to human habitation. 
Not only do the extensive alkaline deposits in the desert basins 
tend to make soil conditions over large areas unsuitable for crops, 
but even the underground water over practically the entire region 
is strongly alkaline and unfit for use. These conditions, combined 
with a deficient rainfall and absence of perennial streams, give no 
promise of an agricultural future. The few ranches are situated 
mainly along the skirts of the desert, where a number of small creeks 
coursing down from higher country furnish sufficient water for small 
fields and garden patches before being lost in the sand. Others are 
found at the few widely separated springs which rise in different 
parts of the desert. A ready and remunerative market for garden 
truck is furnished by the large coal-mining town of Eock Springs, 
and trucking is carried on in that district wherever there is sufficient 
water for irrigating a small garden patch. 

The chief value of the Red Desert, aside from the extensive coal 
fields in its western part, is as a winter range for the hundreds of 
thousands of sheep which spend the summer on the higher plains 
and in the hills and mountains of central and western Wyoming. 
The abundant sagebrush, greasewood, and saltbushes, particularly 
the Nuttall saltbush (see PL IV, fig. 2) and other alkali-resistant 
shrubs and plants, afford an abundance of winter forage, 1 while 
plenty of water for stock is insured by the snows which drift before 
the frequent winds and permit browsing in the cleared spaces. Its 
peculiar adaptation to the winter feeding of sheep, long appreciated 
by the flockmasters, gives the Red Desert region a special importance 
as a necessary complement to one of Wyoming's great industries. 

Wind River Basin — Upper Sonoran Zone. 

The Upper Sonoran area drained by Wind River and its affluents 
lies mainly between 5,000 and 6,000 feet elevation. It extends from 
the southern escarpment of the Owl Creek Mountains to the north- 
eastern base of the lofty Wind River Range, and at the east and 
southeast to the broad tract of high, rolling, sagebrush plains which 
separate it from the Upper Sonoran areas along the Platte and on 
the Red Desert. In its surface features, climate, vegetation, and 
animal life this region is generally similar to the Bighorn Basin, 
with which it is connected narrowly through the rugged canyon 
which cleaves the Owl Creek Mountains and carries the waters of the 

1 For a full discussion of the pasture value of the alkaline desert basins see Nelson, 
The Red Desert of Wyoming and its Forage Resources, Bull. 18, Div. of Agrostology, 
U. S. Dept. Agr., 1898. 



22 NORTH AMERICAN FAUNA. [No. 42. 

Bighorn. The greater elevation of the Wind Kiver Basin results, 
however, in a weaker characterization of the zone, while extensive 
areas of rough bad lands, which fill much of the upper (western) 
part between Wind Eiver and the Owl Creek Mountains and form 
in many places the watersheds between streams elsewhere, greatly re- 
strict the agricultural lands and confine them largely to the valleys. 
Owing to its proximity to the mountain mass of the Wind River 
Range, the western edge of this area receives more moisture than the 
central and eastern portions, where the rainfall seldom exceeds 10 
inches. 

The Upper Sonoran element is most pronounced in the lower val- 
leys below Riverton, and on the plains and bad lands which jut 
against the southern bases of the Owl Creek Mountains. The vege- 
tation on the sandy plain north of Shoshoni is typical of the lower 
elevations generally, consisting of saltbush, greasewood, rabbit brush, 
small brown sagebrush, spiny sagebrush, yucca, and prickly-pear 
cactus, with skunk bush and juniper on bluffs, and broad-leaved 
cottonwood, buffaloberry, flowering currant, and wolfberry along 
streams. Along the eastern base of the Wind River Range and on 
the barren slopes and gulches of the higher bad lands dilute Upper 
Sonoran Zone reaches to nearly 6,000 feet altitude, and here, as well 
as on the southern side of the Owl Creek Mountains, includes the 
fringe of scattering junipers which is more or less evident on the 
margins of the open basin. The effect of slope exposure on zone level 
is especially noticeable on the warm southern slopes of the Owl Creek 
Mountains, where the zone is carried regularly to 6,500 feet, at least 
500 feet higher than on the cooler basal slopes of the Wind River 
Mountains, on the opposite side of the basin. 

The generally weak character of the Upper Sonoran Zone over the 
Wind River Basin is evident from the paucity of characteristic zone 
species of mammals and breeding birds, which include, among mam- 
mals, the pale chipmunk, Great Plains grasshopper mouse, kangaroo 
rat, Bailey cottontail, Great Basin spotted skunk, and California 
bat ; and among birds, the mourning dove, burrowing owl, Arkansas 
kingbird, Bullock oriole, western lark sparrow, white-rumped shrike, 
and long-tailed chat. The plains rattlesnake and the desert short- 
horned and scaly rock lizards are the most noticeable reptiles. 

Extensive coal and oil fields are among the natural resources of 
the Wind River Basin. Agriculture is largely supplemental to cattle 
and sheep raising. Under irrigation the arable valley lands and some 
of the lower bench lands are producing fine crops of alfalfa and other 
forage, and also cereals. The growing season is short for tomatoes 
and tender vegetables, but most kinds do well and are raised exten- 
sively near Lander, Fort Washakie, Riverton, and elsewhere. Apple 



1917.] UPPER SONORAN ZONE. 23 

growing has proved decidedly successful in certain protected valleys 
at the base of the mountains southwest of Lander, and small fruits 
are grown on a considerable scale at Lander and elsewhere. With an 
abundant supply of water in the Wind River Range, the reclamation 
and productiveness of considerable areas of both bench and valley 
lands await the construction of additional and especially higher irri- 
gation canals. 

Bighorn Basin — Upper Sonoran Zone. 

The drainage basin of the Bighorn River is a large open area of 
3,500 to 5,500 feet elevation lying between the high Bighorn and 
Absaroka Ranges in northwestern Wyoming; extending from the 
open Transition Zone ridge of the Owl Creek Mountains northward 
beyond the Montana line. It is a warm, protected section of low 
altitude and extreme aridity, but with an abundance of permanent 
streams, while the generally barren surface of its valley and low 
bench lands or scattered tracts of bad lands supports a scanty desert 
vegetation. The rainfall is often as light as 5 or 6 inches at Basin 
and Lovell, in the low central and northern sections, and less than 
10 inches elsewhere. The Upper Sonoran Zone in this region covers 
an area of fairly regular outline about 100 miles in length from 
north to south by 60 miles in breadth, approximately 6,000 square 
miles. The zone is strongly characterized in the valleys and over 
the lower portions generally, reaching its upper limits at a little 
over 5,000 feet on the bordering sage slopes west and south, and east- 
ward at anywhere from 5,500 to 6,000 feet on the abrupt hot slopes 
along the western bases of the Bighorn Mountains, where it includes 
an irregular belt of scattered junipers varying from 500 to 1,000 feet 
in vertical breadth. 

The fauna and flora of the Bighorn Basin are derived alike from 
the Great Basin and Great Plains regions, and include few if any 
species not common to some of the other Sonoran areas of Wyoming. 
Over this region the Upper Sonoran Zone is variously characterized 
as to vegetation by a rank growth of broad-leaved cottonwood, wil- 
low, buffaloberry, skunk bush, and flowering currant along most of 
the streams; greasewood, rabbit brush, and Suceda on adobe river 
flats; saltbushes, rabbit brush, spiny sagebrush, prickly-pear cactus, 
and such plants as Cleome lutea, Psoralea tenuiflora, and Plantago 
purshi on firm-soil benches, with Grayia spinosa, Polarmia trachys- 
perma, Lupinus pusillus, yucca, sand dock, and a small yellow- 
flowered Malacothrix added in sandy areas; and by a scattering 
growth of juniper and skunk bush on bad lands bluffs and on the 
rough southern and especially eastern margins of the basin. Ex- 
tensive barren flats midway between Greybull and Cody have an 



24 NORTH AMERICAN FAUNA. [No. 42. 

almost pure growth of prickly-pear cactus, while similar tracts near 
Frannie are likewise clothed with the Nuttall saltbush. 

Representative breeding birds include the mourning dove, bur- 
rowing owl, Arkansas kingbird, Bullock oriole, bronzed grackle, 
western lark sparrow, sage sparrow, house finch, lazuli bunting, 
white-rumped shrike, yellow warbler, western yellow-throat, long- 
tailed chat, catbird, brown thrasher, and western marsh wren. 

A few characteristic mammals of the Bighorn Basin are the pale 
chipmunk, black-tailed prairie-dog, Great Plains grasshopper mouse, 
Hayden field mouse, sage pocket gopher, kangaroo rat, Maximilian 
pocket mouse, Bailey cottontail, and the brown and California bats. 
Several reptiles, the prairie bull snake, plains rattlesnake, horned 
lizard, and scaly rock lizard, are common over much of the region. 

This huge depression between lofty mountain ranges is a highly 
favored region of great promise. While physically and climatically 
the best suited to general agriculture of any of the low-altitude areas 
of western Wyoming, it is, perhaps, best known for its adaptation 
to horticulture and for the rapid strides already made in the success- 
ful production of high-grade apples and other fruits. The warm, 
sheltered valleys and hot Upper Sonoran slopes along the western 
bases of the Bighorn Mountains, especially toward the northern end 
of the basin, are highly favorable to fruit culture. It is in these 
situations that the older bearing orchards are chiefly located and the 
best results have thus far been obtained. Young orchards are now 
extensively planted throughout the lower open portions of the basin 
and, although more exposed than nearer the mountains, nevertheless 
give promise of handsome returns under proper care and due atten- 
tion to local conditions. 

The Bighorn River and its principal tributaries, the Shoshone and 
Greybull Rivers, and Shell, No Wood, and Owl Creeks, fed by the 
melting snows of high mountain ranges, carry an abundance of water, 
amply sufficient under proper storage control for watering all ir- 
rigable lands in their drainages (fig. 2). Private irrigation projects 
have already reclaimed considerable portions of the broad and fertile 
stream valleys, together with the lowest of the adjoining bench 
lands, and the Federal Shoshone project has opened large tracts in 
Shoshone Valley. The higher bench lands, at present utilized in 
sheep grazing, await the construction of more storage dams and 
higher irrigation canals before they can be made productive. 

Characteristic Species — Upper Sonoran Zone. 

The delimitation of life zones is based upon the combined ranges of 
characteristic species of mammals, breeding birds, reptiles, and 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate IV. 




Fig. 1.— Bluffs alonq Green River near Utah Boundary. 

Dense growth of greasewood (Sarcobatus vermiculatus) at left. 




12c~ 



■ ^-lOi '■•'»-"" 






■i 




Fiq. 2— Desert Vegetation, Red Desert, South of the Ferris Mountains. 

Chiefly saltbush (Atriplex nuttalli). 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate V. 




Fig. 1.— Yellow Pine Forest near Springhill, Northern Base of Laramie Peak 

(6,500 Feet). 




Fig. 2.— Sagebrush Plain near Fort Washakie, Wind River Basin. 



1917.] 



UPPER SONORAN ZONE. 



25 



plants. x Species which in Wyoming best mark the Upper Sonoran 
Zone throughout. 2 or in restricted areas of its Great Plains or Great 




Fig. 2. — Irrigation canal traversing original sage plain on bench north of Burlington, 

in the Bighorn Basin. 

Basin subdivisions, are included in the following lists. Many species 
of wide zonal dispersion occur commonly in the Upper Sonoran Zone 
in Wyoming, but as these give no definition to the zone they are not 
listed. 

Mammals — Upper Sonoran Zone. 

[Species marked T. occur also in the Transition Zone.] 



Antilocapra americana americanu, An- 
telope. T. 

Eutamias pallidus pallidas, Pale Chip- 
munk. T. 

Eutamias minimus minimus, Least 
Chipmunk. 

Eutamias minimus pictus, Sagebrush 
Chipmunk. 

Eutamias dorsalis utahensis, Utah 
Rock Chipmunk. 

Callospermophilus lateralis ivortmani, 
Wortman Mantled Ground Squir- 
rel. T. 

Citellus obsoletus, Kennicott Ground 
Squirrel. 

Citellus tridecemlineatus pallidus, Pale 
Striped Ground Squirrel. 



Citellus tridecemlineatus parvus, Small 

Striped Ground Squirrel. 
Cynomys ludovicianus ludovicianus, 

Black-tailed Prairie-dog. 
Onychomys leucogaster arcticeps, Great 

Plains Grasshopper Mouse. 
Peromyscus manieulatus o s g o o di , 

Black-eared White-footed Mouse. T. 
Peromyscus manieulatus nebrascensis, 

Yellow White-footed Mouse. 
Peromyscus crinitus auripectus,Go\i\e\\- 

breasted Canyon Mouse. 
Peromyscus truei truei, True's Cliff 

Mouse. 
Reithrodontomys megalotis dyehei, 

Prairie Harvest Mouse. 



1 Species of insects and other groups might be equally useful in the determination of 
zones if their ranges were better known. 

2 See frontispiece (map). 



26 



NORTH AMERICAN FAUNA. 



so. 41 



Mammals — Upper Sonoran Zone — Continued. 



Neotoma cincrca orolestes, Colorado 
Bushy-tailed Wood Rat. T. 

Microtus ochrogaster haydeni, Hayden 
Field Mouse. 

Fiber eibethicus tinnamominus, Great 
Plains Muskrat. T. 

Geomys lutescens, Yellow Pocket 
Gopher. 

Thomomys otitis, Green River Pocket 
Gopher. 

Thomomys talpoides bullatus, Sage 
Pocket Gopher. 

Perodipns ordii luteolus, Wyoming 
Kangaroo Rat. 

Perognathus Mspidus paradoxus, Kan- 
sas Pocket Mouse. 

Perognathus fasciatus fasciatus, Maxi- 
milian Pocket Mouse. T. 

Perognathus fasciatus Vitus, Sweetwa- 
ter Pocket Mouse. 

Perognathus flainis piperi, Cheyenne 
Pocket Mouse. 



Perognathus eallistus, Red Desert 
Pocket Mouse. 

Lepus californieus melanotis, Black- 
tailed Jack Rabbit. 

Sylvilagus auduboni baileyi, Bailey 
Cottontail. 

Sylvilagus floridanus si m His, Nebraska 
Cottontail. 

Canis nebracensis, Plains Coyote. T. 

Vulpes velox velox, Kit Fox, Swift. T. 

Procyon lot or lot or, Raccoon. T. 

Taxidea taxus taxus, Badger. T. 

Spilogale gracilis saxatilis. Great Ba- 
sin Spotted Skunk. 

Mustela nigripes. Black-footed Ferret. 

Scalopus aquaticus caryi, Northern 
Plains Mole. 

Myotis californieus caHfornicus, Cali- 
fornia Bat. 

Myotic evotis, Long-eared Bat. 

Myotis longicrus interior, Long-legged 
Bat. T. 

Eptesicus fuscus fusciis. Brown Bat. T. 



Breeding Birds — Upper Sonoran Zone. 
[Species marked T. breed also in the Transition Zone.] 



Querquedula discors, Blue-winged Teal. 
T. 

Querquedula ci/anoptera, Cinnamon 
Teal. T. 

Botaurus lentiginosus, Bittern. T. 

Ardea herodias. Great Blue Heron. T. 

Nycticorax nycticorax narius, Black- 
crowned Night Heron. T. 

Rail us Virginian us, Virginia Rail. 1 

Porzana Carolina, Carolina Rail, Sora. 
T. 

Fulica americana, Coot. T. 

Bartramia longicauda, Upland Plover. 
T. 

Numenius americanus, Long-billed Cur- 
lew. T. 

Zenaidura macroura carolinensis, 
Mourning Dove. 

Archibuteo ferrugi/neus, Ferruginous 
Rough-legged Hawk. T. 

SiJeotiito cunicularia hypugcea, Burrow- 
ing Owl. 

Chordeiles rirginianus hcnriji. Western 
Nighthawk. T. 

1 Observed during 



Tyrannus tiirannus. Kingbird. T. 

Tyrannus rerticalis, Arkansas King- 
bird. 

Tyrannus vociferans, Cassin Kingbird. 

Corrus brachyrhynchos hesperis, West- 
ern Crow. T. 

Cyanocephalus cyanocephalus, Pinyon 
Jay. 

Xanthocephalus xanthoeephalus, Yel- 
low-headed Blackbird. T. 

Agelaius phoerdceus fortis, Thick-billed 
Redwing. T. 

Sturnclla neglecta, Western Meadow- 
lark. T. 

Icterus bullocki, Bullock Oriole. 

Quiscalus quiscula aneus, Bronzed 
Grackle. 

Carpodacus mexieanus frontalis. House 
Finch. 

Annnodrainus sarannaruiih bimacula- 
tus, Western Grasshopper Sparrow. 

Chondestes grammacus strigatus,West- 
ern Lark Sparrow. 

Spisella breweri, Brewer Sparrow. T. 

breeding season. 



1917.] 



UPPER SONORAN ZONE. 



27 



Breeding Birds — Dppeb Sonoran Zone — Continued. 



AmpMspiza neradensis, Sage Sparrow. 

Passerina amcena, Lazuli Bunting. 

Spiza americana, Dickcissel. 

Calamospiza melanocorys, Lark Bunt- 
ing. T. 

Lanius ludovicianus excubitorides, 
White-rumped Shrike. T. 

Dendroica (estiva ce&tiva, Yellow 
Warbler. T. 

Oeothlypis trichas Occident alis, West- 
ern Yellow-throat. T. 

Icteria rirens longicauda, Long-tailed 
Chat. 

Oreoscoptes montanus. Sage Thrasher. 
T. 



Mimus polyglottos leucopterus, West- 
ern Mockingbird. 

Dumetella carolinensis, Catbird. 

Toxostoma rufum, Brown Thrasher. 

Thryomanes betoicM bairdi, Baird 
Wren. 1 

Telmatodytes palustris plexitis. West- 
ern Marsh Wren. 

Boeolophus inomatus griseus, Gray 
Titmouse. 1 

Psaltriparus plumbeus, Lead-colored 
Bush-tit. 1 

Polioptila ccBrulea obscura, Western 
Gnatcatcher. 1 



Reptiles — Upper Sonoran Zone. 

[Species marked T. occur also in the Transition Zone.] 

Lizards. 



Holbrookia maculata. Sand Swift. 
Uta stansburiana, Stansbury Lizard. 
Sceloporus consobrinus, Scaly Lizard. 
Sceloporus graciosus, Scaly Rock 

Lizard. 
Phrynosoma ornatissimum, Desert 

Short-horned Lizard. T. 



Cnemidophorus sexlineatus, Six-lined 

Lizard. 
Eumeces multivirgatus, Many-lined 

Skink. 



Snakes. 



Thamnophis sirtalis parietalis, Red- 
barred Garter Snake. T. 
Thamnophis radi.v. Garter Snake. T. 
Bascanion constrictor. Blue Racer. 
Pituophis sayi, Prairie Bull Snake. 



Liopeltis vemalis, Smooth Green 
Snake. T. 

Heterodon nasicus, Hog-nosed Snake. 

Crotalits confluentus, Plains Rattle- 
snake. 



Amphibians — Upper Sonoran Zone. 

[Species marked T. occur also in the Transition Zone.] 

Toads and frogs. 



Scaphiopus hammondi bombifrons, 

Spadefoot Toad. 
Bufo lentiginosus woodhousei, Toad. 

T. 



Bufo cognatus. Toad. 

Rana pipiens, Leopard Frog. T. 



Salamanders. 



Ambystoma tigrinum. Tiger Salaman- 
der. T. 



Probably breeds. 



28 



NORTH AMERICAN FAUNA. 



[No. 42. 



Plants — Upper Sonoran Zone. 

[Species marked T. occur also in the Transition Zone.] 

Trees and shrubs. 



Pinus edulis, Pinyon, Nut Pine. 

Junipcrus knighti, Desert Juniper. 

Junipcrus monosperma, One-seeded 
Juniper. 

Populus occidentalis, Broad-leaved 
Cottonwood. 

Populus acuminata, Lance-leaved Cot- 
tonwood. 

Salia amygdaloides, Peach-leaved Wil- 
low. 

Salix fluviatilis, Sand-bar Willow. 

Ulmus americana? Elm. T. 

Acer negundo, Box Elder. T. 

Fraxinus lanceoiata, Ash. T. 

Cercocarpus parvifolius, Mountain Ma- 
hogany. T. 

Sarcobatus vermiculatus, Greasewood. 

Atriplex canescens, Saltbush, Gray 
Shadscale. 

Atriplex confertifoUa, Spiny Saltbush. 

Atriplex nuttalli, Nuttall Saltbush. 

Atriplex pabularis, Nelson Saltbush. 

Atriplex argentea, Silvery Saltbush. 

Eurotia lanata, White Sage, Winter 
Fat. 

Grayia spinosa, Grayia. 

Kochia americana, Kochia, White 
Sage. 

Philadelphus occidentalis, Western 
Syringa. 

Ribes longiflorum, Flowering Currant. 

Prunus americana, Wild Plum. T. 

Prunus besseyi, Sand Cherry. 



Amorpha canescens, False Indigo, 

Shoestring. T. 
Amorpha nana, False Indigo. 
Schmaltzia trilobata, Skunk Bush. 
Sell maltzia glabra, Smooth Sumac. T. 
Rhus rydbergi, Western Poison Ivy. T. 
Vitis vuipina, Wild Grape. T. 
Paj-thenocissus vltacea, Virginia 

Creeper. T. 
Lepargyrea argentea, Buffaloberry. T. 
Symphoricarpos occidentalis, Wolf- 
berry. T. 

Gutierrezia sarothrw, Rabbit Brush. 
Chrysothamnus graveolens, , Rabbit 

Brush. 
Chrysothamnus plattensis, Rabbit 

Brush. 
Chrysothamnus linifolius, Rabbit 

Brush. 
Chrysothamnus stcnophyllus, Rabbit 

Brush. 
Tctradymia inermis, Rabbit Brush. 
Tctradymia spinosa, Rabbit Brush. 
Tetradymia nuttalli, Rabbit Brush. 
Artemisia fllifolia, Narrow - leaved 

Sagebrush. 
Artemisia spinescens, Spiny Sagebrush, 

Budbrush. 
Artemisia pedatiflda, Small Brown 

Sagebrush. 
Artemisia, tridentata, Black Sage- 
brush. T. 



Herbaceous plants. 



Tradescantia occidentalis, Spiderwort. 
Yucca glauca, Plains Yucca. 
Eriogonum effnsum, Eriogonum. 
Eriogonum annuum, Eriogonum. 
Eriogonum campanulatum, Eriogonum. 
Eriogonum corymbosum, Eriogonum. 
Eriogonum multiccps, Eriogonum. 
Rumex venosus, Sand Dock. 
Suwda diffusa, Sea Blite. 
Suceda moquini, Shrubby Blite. 
Endolepsis sucTcleyana. 
Salicornia rubra, Glasswort. T. 



Abronia fragrans. 

Abronia elliptica. 

Argemonc intermedia, Prickly Poppy. 

Argemone hispida.. Prickly Poppy. 

Stanleya tomentosa, Stanleya. 

Stanleya intcgrifolia, Stanleya. 

Cleome lutea, Yellow Cleome. 

Cleome serrulata, Red Cleome, Honey 

Plant. 
Polanisia trachysperma. Clammy-weed. 
Ijiipinus plattensis. Lupine. 
Lupiwus pusillus, Small Lupine. 



> Northeast Wyoming only. 



1917.] 



UPPER SONORAN ZONE. 



29 



Plants — Upper Sonokan Zone — Continued. 
Herbaceous plants — Continued. 



Astragalus crassicarpus, Buffalo Bean, 
Ground Plum. 

Astragalus mollissimus, Milk Vetch. 

Astragalus missouricnsis, Milk Vetch. 

GlycyrrMza lepidota, Wild Licorice. T. 

Psoralea tenuiflora, Psoralea. 

Psoralea Unearifolia, Narrow-leaved 
Psoralea. 

Psoralea lanceolata, Psoralea. 

Psoralea hypogca, Psoralea. 

Psoralea esculcnta, Psoralea. 

Parosela enncandra, Dalea. 

Pctalostemon oligophyllus, White Prai- 
rie Clover. 

Petalostcmon eandidus, Prairie Clover. 

Petalostcmon purpureas, Purple Prai- 
rie Clover. 

Petalostcmon villosus, Silky Prairie 
Clover. 

Lathyrus ornatus, Wild Pea, Vetch- 
ling. 

Linum rigidum, Wild Flax. 

Euphorbia margiiiata, Snow-on-the- 
inountain. 

C rot on texensis, Croton. 

Malvastrum coccincum, False Mallow. 

Malvastrum dissection, False Mallow. 

Mentzclia decapetala, Loasa. 

Mentzclia nuda, Loasa. 

Mentzelia albicanlis, Loasa. 

Mentzclia he ricaulis. Yellow Loasa. T. 

Ma miliaria missouricnsis, Ball Cactus. 
T. 

Mamillaria vivipara, Ball Cactus. 

Eehinocereus viridiflorus, Green- 
flowered Petaya. T. 

Opuntia polyacantha. Prickly Pear. 

Opuntia rutila, Prickly Pear. 



Anogra albicaulis, White Evening 
Primrose. 

Galpinsia la v and ulw folia.. 

Meriolix serrulata. 

Oaura coccinca. 

Asclepias speciosa, Milkweed. 

Asclepias pumila, Milkweed. 

I pom oca leptophylla, Bush Morning- 
glory. 

Gilia polycladon. 

Lithospcrmum angustifolium, Grom- 
well. T. 

Lithospcrmum gmcliiii, Gromwell. 

Oreocarya flava. 

Lippia cuneifolia. 

Verbena hastata, Blue Vervain. 

Verbena bractcosa, Low Vervain. 

Physalis lanceolata, Ground Cherry. 

Solarium rostratum, Buffalo Bur. 

Pcntstcmon albidus, Beard-tongue. 

Pentstemon angustifolius, Beard- 
tongue. 

Plant a go purshi, Plantain. 

Liatris punctata, Blazing Star. 

Grindelia squarrosa, Gum Plant. 

Sideranthns spinulosus. 

Solid a go mollis, Goldenrod. 

Solid a go canadensis gilvocanescens, 
Goldenrod. 

Solidago rigida, Goldenrod. 

Ratibida columnaris, Cone Flower. 

Helianthus annuiis. Sunflower. 

Helianthus petiolaris, Sunflower. 

Hymenopappus fllifolius. 

Cardans plattensis, Thistle. 

Malacothrix sonehoides. 

Lygodesmia juncea. 

Lygodesmia rostrata. 



Grasses. 



Andropogon scoparius, Bluestem. 
Andropogon halli, Bluestem. 
Panieum virgatum, Panic Grass. 
Aristida longiscta, Wire Grass. T. 
Stipa comata. Feather Grass. 
Stipa viridula, Feather Grass. T. 
Oryzopsis micrantha, Rice Grass. 
Eriocoma cuspidata, Indian Millet. 



Muhlenbergia pungens, Dropseed Grass. 
Sporobolus airoides, Fine-top Salt 

Grass. 
Sporobolus cryptandrus, Bunch Grass. 
Calamovilfa longifolia, Reed Grass, 

Sand Grass. 
Sehedonnardus paniculatus, Crab 

Grass. 



30 



NORTH AMERICAN FAUNA. 



[No. 42. 



Plants — Upper Sonoran Zone — Continued. 
Grasses — Continued. 



Spartina gracilis. Marsh Grass. 
Bouteloua oligostachya, Grama 

Grass. T. 
Bouteloua hirsuta, Grama Grass. 
Atheropogon curtipendula, J?all 

Grama. T. 
Bulbilis dactyloides, Buffalo Grass. 
Munroa squarrosa, False Buffalo 

Grass. 
Eatonia obtusata. 
Distichlis spicata, Salt Grass. 
Poa fendlcriana, Spear Grass. T 



/ 



Poa sheldoni, Spear Grass. T. 
Festuca octoflora, Fescue Grass. 
Agropyron spicatum, Wheat Grass. 
Agropyron smithi, Wheat Grass. T. 
Sitanion longifolium, Long - bearded 

Rye Grass. T. 
sitanion Itystri.r. T. 
Ely in us canadensis, Wild Rye. T. 
Elymus condcnsatus, Giant Rye 

Grass. T. 
Elymus salinus, Desert Rye Grass. 



Agricultural Utility of the Upper Sonoran Zone. 

The growing of lm 7 and forage crops as supplemental to stock 
raising and a limited planting of cereals and vegetables for ranch 
use long constituted the chief agricultural endeavors in even the 
most favored sections of Wyoming. The establishment of agricul- 
ture as a separate industry in this arid region is comparatively recent, 
the result of greatly increased irrigation facilities and of more effi- 
cient methods of conserving the rainfall in the less arid dry-farming 
districts. 

The Upper Sonoran areas of Wyoming lie mainly in the upper 
and cooler parts of the zone and are therefore climatically unsuited 
to a variety of tender crops, and especially fruits, which are suc- 
cessfully grown elsewhere in its lower and warmer portions. They 
are proving well adapted, how T ever, to most of the standard varieties 
of wheat, oats, rye, emmer, and other cereals, which yield abundantly 
under irrigation and moderately under dry farming. None but the 
early varieties of corn mature, and these are raised to a small extent 
only in the warmest parts, as in the lower Platte and Bighorn 
Valleys. Alfalfa, the staple forage crop, produces regularly two 
and at the lowest elevations often three or even four crops a year. 
Sugar beets are raised extensively in the Bighorn Basin, the lower 
Platte Valley, and near Wheatland. The sugar content of Wyo- 
ming beets is high, and with the construction of local sugar factories 
this crop should become one of the most important and profitable. 
Potatoes yield abundantly and are well adapted to the dry-farming 
districts along the eastern edge of the State, where they are pro- 
duced on a large scale. Vegetables of most sorts thrive under irri- 
gation, and trucking is profitably carried on in many valleys. The 
tender sorts usually succeed in the warmest localities, but can not be 
raised generally, because of the short growing season. 



7.] TRANSITION ZONE. 31 

Fruit growing has been attempted on a commercial scale mainly 
the Bighorn Basin, near Lander, and in the Wheatland district, 
Lere many of the hardier varieties of apples, cherries, and plums, 
well as the small fruits, raspberries, strawberries, grapes, currants, 
oseberries, and dewberries, are moderately successful under irri- 
tion. Even in these sheltered and favored districts the physical 
d climatic conditions are so localized that successful horticulture 
11 perforce be limited to comparatively small areas where there is 
lple protection and the greatest freedom from the destructive late 
osts, which kill so much of the fruit blossom in this zone through- 
t the Eocky Mountain region. Late-flowering and frost-resistant 
rieties succeed best. 

TRANSITION ZONE. 

Fully half of Wyoming, or about 50,000 square miles, is in the 
■ansition Zone, the zone intermediate between the Boreal and Aus- 
al regions, which is here in its greater part open and treeless, and 
us less conspicuously characterized than in the southern Rocky 
Mountain region, where it is very generally marked by extensive for- 
ests of yellow pine. In Wyoming the zone comprises, broadly, vast 
interior sagebrush plains (see PL V, fig. 2) and watersheds, plateaus, 
and high-altitude basins in the central and southwest sections; and 
elevated grassy plains to the east and southeast. It also includes all 
except the higher summits of the pine-clad foothill ranges in the 
eastern part of the State, and the open basal sagebrush slopes of 
the high mountain ranges farther west. In greater detail, 1 the 
Transition Zone in Wyoming embraces the following important 
areas : 

The Cheyenne Plains above 5,500 feet elevation ; Lusk Plains above 
5,000 feet ; all of the Laramie Plains and Shirley Basin ; upper valley 
of the North Platte above 6,500 feet; upper Green River Basin, 
Bear River region, and extensive tracts on the borders of the Red 
Desert above 6,500 feet; sage plains between Casper and Lander 
above 6,000 feet; Wind River Valley above 6,000 feet; mainly open 
borders of the Bighorn Basin, 5,500 to 7,000 or 7,500 feet; all open 
or partly timbered watersheds between the Bighorn and Bear Lodge 
Mountains above 4,500 feet; basal and middle slopes of Bear 
Lodge Mountains and northern Black Hills, 4,000 to 5,500 feet; 
and southern Black Hills region, 4,500 to 6,000 feet. Limited areas 
in Jackson Hole and Salt River Valley are eastward continuations 
of the Snake River Transition Zone, and narrow strips of the zone 
enter Yellowstone Park for a short distance along the Yellowstone, 

i See zone map (frontispiece). 



32 NORTH AMERICAN FAUNA. [N- 

penetrating the valleys of the Gardiner, Lamar, and other tributa 
and also along the Gallatin, Madison, and Fall River valleys. 

The foregoing well illustrates the effect of both base level 
latitude on the vertical position of the Transition Zone. A 
level of over 6,000 feet in the southwest elevates the zone i 
of the region contiguous to the Red Desert and Green River ~\ 
to between 7,000 and 8,500 feet on warmest (southwest) slope 
0,000 to 7,500 feet on coldest (northeast) slopes. In extreme 
eastern Wyoming, adjacent to the low-altitude northern 
Plains, the zone level attains only 4,500 to 6,000 feet on sou 
slopes and 4,000 to 5,500 feet on northeast slopes. 1 Much 
2,000-foot variation in zone level results from difference i 
elevation, while probably 800 feet is normal depression in th( 
east due to higher latitude. Locally, mountain slopes of unuo,. 
exposure and warmth carry zones abnormally high, and if very 
steep and abrupt the horizontal contraction is also very marked. 
This is well shown in the Transition belt on the abrupt, hot, south- 
west slopes of the Bighorn Mountains east of Greybull and Ionia. 
Varied physical conditions, and, to a certain extent, deforestation, 
affect both the elevation and the horizontal as well as the vertical 
width of life zones, especially in mountainous districts. In the 
main, however, the Transition Zone is maintained in fairly uniform 
elevation and width along the bases of the Wyoming ranges. 

Characteristic Species — Transition Zone. 

The Transition Zone in Wyoming is conspicuously marked only 
along its upper border, where mostly open sage slopes give way to 
the aspen and coniferous forest belt of the Canadian Zone. Incon- 
spicuous vegetation is characteristic of the lower part on the open 
plains, where the zonal position is further indicated by either the 
absence or a marked paucity of Sonoran species. As elsewhere in 
the Rocky Mountain region, its fauna and flora are fairly constant 
throughout, but include many species, both Boreal and Austral, from 
the adjoining zones. 

Sagebrush, yellow pine, and grasses are prominent types of vege- 
tation in the Wyoming Transition area. The sagebrush (PI. VI), 
the most widely distributed shrub, usually occurs in pure growth, 
while the yellow pines are restricted largely to the lower mountains, 
foothills, and rough tracts in the eastern half of the State. Con- 
siderable Douglas spruce and scattering Rocky Mountain white pines 

1 On exposed slopes of mountains the difference in zone level on the warm and cold 
sides is usually as much as 1,000 feet. So much variation is not found in the low hill 
country of northeastern Wyoming. 



North American Fauna No. 42, U. S. Dept Agr. Biological Survey. 



Plate VI. 




Fig. 1.— Sagebrush in Wind River Valley. 
Lower part of Transition Zone, near Circle. 




Fig. 2.— Garfield Peak, Rattlesnake Mountains. 
Sagebrush covers much of this barren 8,000 to 9,000 foot range. 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate VII. 



'■-*ts£SBSi 




~ ! y*&$& 




Fig. 1.— Rawhide Butte, Lower Border of Transition Zone, at Edge of Great 

Plains. 

Narrow-leaved cottonwoods on Rawhide Creek, southern base of the butte, and yellow pines on 

hills (5,000 feet). 




Fig. 2.— Southwestern Base of Wind River Range near Big Sandy. 

Rabbit brush ( Chrysotltamnus and Tetradymia inermis) on the plain (7,500 feet). 



1917.] 



TRANSITION ZONE. 33 



occur in this zone in the mountains of western Wyoming, and the bur 
( • ,k is a common tree on the Bear Lodge Mountains and elsewhere in 
northern Crook County. On streams along the bases of the moun- 
tains generally the zone is marked by narrow-leaved cotton wood (PL 
VII, fig. 1), diamond willow, and usually by a dense shrubbery of 
Rocky Mountain birch, black and red haws, cornel, wild gooseberry 
and currant, serviceberry, and silverberry; on foothill and lower 
mountain slopes both in the forest as undershrubs and in the open, 
by Rocky Mountain and creeping junipers, Bebb willow, barberry, 
wild red currant, mountain mahogany, kunzia, ninebark, wild 
cherry, mountain and large-toothed maples, mountain balm, bear- 
berry, mountain snowberry, and several high plains species of sage- 
brush and rabbit brush (PL VII, fig. 2) ; and throughout by a great 
many herbaceous plants. 

A considerable number of birds of both general and restricted 
breeding range within the zone characterize this area in Wyoming 
during the nesting season. Representative species are the sage hen, 
sharp-shinned hawk, saw-whet owl, Lewis woodpecker, white- 
throated swift, Wright flycatcher, magpie, piny on jay, McCown 
iongspur, white- winged junco, mountain song sparrow, arctic and 
, ireen-tailed towhees, western tanager, plumbeous vireo, Macgillivray 
warbler, Rocky Mountain and pygmy nuthatches, and willow thrush. 

Mammals wholly or chiefly restricted to the Transition Zone in 
different parts of Wyoming include the plains white-tailed deer; 
Black Hills red squirrel; Wyoming and Uinta ground squirrels; 
white-tailed prairie-dog; bushy-tailed wood rat; pygmy and Uinta 
field mice; Coues, Black Hills, Fort Bridger, and pygmy pocket 
gophers; Uinta pocket mouse; prairie jumping mouse; white-tailed 
lack rabbit; Black Hills cottontail; northern plains skunk; and long- 
legged bat. 

The following reptiles and amphibians are apparently more abun- 
dant in the Transition than in the Upper Sonoran Zone in Wyoming: 
Western garter snake (Thamnophis ordinoides vagrans), a toad 
(Bufo boreas), and a frog (Rana pretiosus). Others of regular oc- 
currence in at least the lower part of the zone include the horned 
lizard (Phrynosoma ornatissimum) , scaly rock lizard (Sceloporus 
grariosus), garter snakes (Thamnophis sirtalis parietalis and T. ra- 
dli'), smooth green snake (Liopeltis vernalis), western toad {Bufo 
lentiginosus woodhousei) , frogs (Rana pipiens and Chorophilus tri- 
seriatus), and tiger salamander (Ambystoma tigrinum). 
74440°— 17 3 



34 



NORTH AMERICAN FAUNA. 



[No. 42. 



Mammals — Transition Zone. 

[Species marked U. occur also in the Upper Sonoran Zone; those marked C, also in the 

Canadian.] 



Cervus canadensis canadensis, Elk. C. 
Odocoilcns virginiamts niacrourus. 

Plains White-tailed Deer. 
Odocoileus hemionus hemionus, Mule 

Deer. U., C. 
Ovis canadensis auduboni, Bad Lands 

Sheep. 1 U. 
Sciurus hudsonicus dakotensis, Black 

Hills Red Squirrel. 
Glaucomys sabrinus canescens, Flying 

Squirrel. 
Callospermophilus lateralis lateralis, 

Say Ground Squirrel. C. 
Callospermoph Hits lateralis wort mani, 

Wortman Ground Squirrel. U. 
Citellus richardsoni clcgans, Wyoming 

Ground Squirrel. U. 
Citellus armatus, Uinta Ground Squir- 
rel. C. 
Cynomys leucurus, White -tailed 

Prairie-dog. U. 
Onychom ys h ucogaster brevicaudus, 

Idaho Grasshopper Mouse. U. 
Onychomys leucogaster tnissouriensis, 

Northern Grasshopper Mouse. U. 
Peromyseus leucopus aridulus, White- 
footed Mouse. 
Ncotoma- cinerea cinerea, Bushy-tailed 

Wood Rat. 
Neotoma cinerea, orolestes, Colorado 

Bushy-tailed Wood Rat. 17. 
Microtus pennsylvanicus modestus, 

Saguache Meadow Mouse. U. 
Microtus montanus caryi, Uinta 

Meadow Mouse. 
Microtus pan per rim us, Pygmy Field 

Mouse. 
Fiber zibethicus osoyoosensis. Rocky 

Mountain Muskrat. C. 



('((■star canadensis, Beaver. C. 
Thomomys talpoides nebulosus, Blark 

Hills Pocket Gopher. C. 
Thomomys talpoides clusius, Coues 

Pocket Gopher. 
Thomomys bridgeri, Fort Bridger 

Pocket Gopher. 
Thomomys pygmceus, Pygmy Pocket 

Gopher. 
Perognathvs parvus clarus, Uinta 

Pocket Mouse. 
Zapus hudsonius campestris, Prairie 

Jumping Mouse. 
Erethizon epixanthum, Yellow-haired 

Porcupine. C. 
Lcpus toiniscndi eampanius, White- 
tailed Jack Rabbit. 
Lcpus townsendi townsendi, Western 

White-tailed Jack Rabbit. 
Sylvilagus nuttalli grangcri, Black 

Hills Cottontail. 
Felis hippolestes, Mountain Lion. C. 
Lynx uinta. Mountain Wildcat. 
Cants nubilus, Buffalo Wolf. U. 
Canis testes, Mountain Coyote. 
Taxidca taxus taxus. Badger. U. 
Spilogale tenuis, Rocky Mountain 

Spotted Skunk. 
Mephitis hudsonica, Northern Plains 

Skunk. 
Mustela arizonensis, Arizona Weasel. 

C. 
Mustela vison encrgumenos, Mink. C. 
Ursus horrlbilis, Grizzly Bear. 
Corynorhinus macrotis palleseens. Big- 
eared Bat. U. 
Myotis longicrus interior, Long-legged 

Bat. U. 



Breeding Birds — Transition Zone. 

[Species marked U. breed also in the Upper Sonoran Zone; those marked C, also in the 

Canadian.] 



Colymbus nigricollis californicus, 
American Eared Grebe. U. 

Anas platyrynchos. Mallard. U. 

Cliaulelasmus streperus, Gadwall. U. 

Ncttion carolinense^ Green-winged 
Teal. 



Steganopus tricolor, Wilson Phalarope. 

U. 
Recurvirostra americana, Avocet. U. 
Gallinago delicata, Wilson Snipe. C. 
Ca top troph or us semi pa I ma tus inorna- 

tus. Western Willet. 
1 Apparently nearing extinction. 



1917.] 



TRANSITION ZONE. 



35 



Breeding Birds — Transition Zone — Continued. 



Podasocys montanus, Mountain Plover. 
V. 

Pedioscetes phasianellus columbianus, 
Columbian Sharp-tailed Grouse. 

Centrocercus urophasiarms, Sage Hen. 

Accipiter velox, Sharp-shinned Hawk. 
C. 

Accipiter cooperi, Cooper Hawk. 

Butco swainsoni, Swainson Hawk. 

Cryptoglaux acadica, Saw-whet Owl. 

Otus asio maxwelliw, Rocky Mountain 
Screech Owl. U. 

Asio tcilsonianus, Long-eared Owl. 

Coccyzus erythropthalmus, Black- 
billed Cuckoo. U. 1 

Dryobates villosus monticola, Rocky 
Mountain Hairy Woodpecker. G. 

Dryobates pubcscens homorus, Batch- 
elder Woodpecker. 

Melanerpes erythroccphalus, Red- 
headed Woodpecker. U. 

Asyndesmus lewisi, Lewis Wood- 
pecker. 

Phalwnoptilus nuttalli nuttalli, Poor- 
will. U. 

Aeronautes melanoleucus, White- 
throated Swift. U. 

Stellula calliope, Calliope Humming- 
bird. 2 

Sayornis sayus, Say Phoebe. U. 

Myiochanes richardsoni, Western Wood 
Pewee. 

Empidonax minimus, Least Flycatcher. 

Empidonax icriyliti, Wright Fly- 
catcher. 

Empidonax hammondi, Hammond Fly- 
catcher. C. 

Otocoris alpestris leucolcema, Desert 
Horned. Lark. U. 

Pica pica hudsonia, Magpie. 

Cyanoceplwlus eyanocepliaius, Pinyon 
Jay. V. 

Dolichonyx oryzirorus. Bobolink. U. 

Euphagus eyanocephalus, Brewer 

Blackbird. U. 
Calcarius ornatus, Chestnut-collared 
Longspur. U. 

Rhynchoplianes mccowni, McCown 
Longspur. 



Pooecetes gramineus confinis, Western 
Vesper Sparrow. U. 

Passerculus sandwichensis alaudimis, 
Western Savannah Sparrow. U. 

Spisella passerina arizonw, Western 
Chipping Sparrow. 

Spizella breweri, Brewer Sparrow. U. 

Junco aikewi, White-winged Junco. 

Melospiza melodia montana, Mountain 
Song Sparrow. 

PassereUa iliaca schistacca, Slate-col- 
ored Fox Sparrow. 

Pipilo maculatus arcticus, Arctic 
Towhee. 

Oreospiza chlorura, Green - tailed 
Towhee. 

Zamelodia melanoeeplxala, Black- 
headed Grosbeak. U. 

Piranga ludoviciana, Western Tana- 
ger. 

Tachycineta thalassina lepida. North- 
ern Violet-green Swallow. C. 

Vireosylva gilva sioainsoni, Western 
Warbling Vireo. 

Lanirireo solitarius pi umbras. Plum- 
beous Vireo. 

Dendroica auduboni, Audubon War- 
bler. C. 

Dendroica nigrescens, Black-throated 
Gray Warbler. 3 

Seiurus aurocapillus. Oven-bird. C. 

Oporornis tolmiei, Macgillivray War- 
bler. C. 

Oreoscoptes montanus, Sage Thrasher. 
U. 

Salpinctes obsoletus, Rock Wren. U. 

Troglodytes aedon parkin a ni. Western 
House Wren. U. 

Sitta carolinensis nelsoni. Rocky 
Mountain Nuthatch. 

Sitta pygmwa, Pygmy Nuthatch. 

Penthestes atricapillus septentnonalis. 
Long-tailed Chickadee. U. 

Hylocichla fuscescens salieicola, Wil- 
low Thrush. 

Planesticus m i<ira$orius propinqv »••>•. 
Western Robin. C. 

Sialia currucoides, Mountain Blue- 
bird. C. 



1 Observed during breeding season. 

2 Taken during breeding season. 

3 Probably breeds. 



36 



NORTH AMERICAN FAUNA. 



[No. 42. 



Plants — Transition Zone. 

[Species marked 17. occur also in the Upper Sonoran Zone; those marked C, also in the 

Canadian.] 

Trees and shrubs. 



Pinus scopulorum, Yellow Pine, Rock 
Pine. 

Junipcrus scopulorum, Rocky Moun- 
tain Juniper. V. 

Junipcrus sabina, Creeping Juniper, 
Trailing Savin. 

Populus angustifolia, Narrow-leaved 
Cottonwood. 

Salix bebbiana, Bebb Willow. C. 

Salix cordata watsoni, Diamond Wil- 
low. U. 

Salix mackenziana, Diamond Willow. 

Salix scoulcriana, Willow. 

Betula fontinalis, Rocky Mountain 
Birch. 

Quercus macrocarpa, Bur Oak. 

Herberts aquifolium, Barberry, Ore- 
gon Grape. 

Grossularia incrmis, Gooseberry. 

Ribes inebrious, Red Currant. 

Ribes americanum, Currant. 

Edwinia americana. 

Ccrcocarpus ledifolius, Mountain Ma- 
hogany. 

Ccrcocarpus intricatus, Mountain Ma- 
hogany. 

Ccrcocarpus parvifolius, Mountain Ma- 
hogany. U. 

Kunzia tri&entaia. 

Holodiscus dumosus. 

Opulaster monoyynus, Ninebark. 

Opulaster pubescens, Ninebark. 

Opulaster malvaceus, Ninebark. 

Rubus deliciosus, Flowering Rasp- 
berry. 

iclanchier alni folia, Serviceberry. 
iclanchier elliptica, Serviceberry. 
nelanchier oreophila, Serviceberry. 
•atcegus rivularis, Black Hawthorn. 

i "atcegus cerronis, Hawthorn. 



Crataegus sheridana, Red Hawthorn. 

Prunus melanocarpa, Chokecherry. 

Prunus poinsylvanica, Wild Red 
Cherry. C. 

Acer glabrum, Mountain Maple. C. 

Acer grandidentatum, Barge-toothed 

Maple. 
jDeanothus velutinus, Mountain Balm. 

Ceanothus fendleri, Wild Tea Bush. 

Ceanothus mollissimus. 

Elwagnus argentea, Silverberry. 

Cornus stolonifera, Cornel. 

Cornus instolonea, Cornel. 

Arctostaphylos uva-ursi, Bearberry. 

Sambucus canadensis, Elderberry. 

Sambucus melanocarpa, Mountain 
Black Elderberry. C. 

Viburnum lentago, Sweet Viburnum. 

Symphoricarpos rotundifolius, Snow- 
berry. 

Symphoricarpos orcophilus, Mountain 
Snowberry. 

Symphoricarpos pauciflorus, Snow- 
berry. 

Lonicera glaucescens, Douglas Honey- 
suckle. 

Chrysothamnus wyomingensis. Rabbit 
Brush. 

Chrysothamnus pulcherrimus, Rabbit 
Brush. 

Chrysothamnus parryi, Rabbit Brush. 

Chrysothamnus frigidus, Rabbit Brush. ' 

Artemisia tridentata, Black Sage- 
brush. U. 

Artemisia cana, Gray Sagebrush. U. 

Artemisia trifida, Sagebrush. 

Artemisia arbuscula, Brown Sage- 
brush. 

Artemisia frigida. Sagebrush. 

Artemisia ludoviciana, Sagebrush. V. 



Calochortus gnnnisoni, Mariposa Lily. 
Calochortus nuttalli, Mariposa Lily. 
Zygadenus venenosus, Poison Ca- 
mas. U. 
Iris missouriensis, Blue Flag. 
Corallorhiza multiflora. Coral Root. C. 
Humulus lupulus, Wild Hop. 



Herbaceous plants. 

Letcisia rediriva. Bitter Root. 
Arenaria congesta, Sandwort. C. 
Clematis ligust id folia, White Virgin's 

Bower. 
Clematis douylasi. Purple Virgin's 

Bower. 
Clematis occidentalis, Virgin's Bower. 



1917.] 



TRANSITION" ZONE. 



37 



Plants — Transition Zone — Continued. 
Herbaceous plants — Continued. 



Anemone cylindrica, Anemone. U. 

Cyrtorhyncha ranunculina, Nuttall 
Buttercup. 

Physaria didymocarpa, Double Blad- 
der-pod. 

Drymocallis glandulosa, Glandular 
Cinquefoil. C. 

Potentilla effusa, Cinquefoil. 

Thermopsis rhombifolia, Yellow Ther- 
mopsis. 

Lupinus argenteus, Silvery Lupine. 

Astragalus succulentus, Ground 
Plum. U. 

Astragalus drummondi, Drummond 
Milk Vetch. 

Astragalus hypoglottis, Milk Vetch. U. 

Astragalus flexuosus, Milk Vetch. U. 

Astragalus nitidus, Milk Vetch. 

Astragalus calycosus, Milk Vetch. 

Aragallus lamberti. U. 

Psoralea urgophylla, Silvery Psoralea. 

Hcdysarum cinerascens, Hedysaruin. 

Hcdysarum uintahense, Uinta Hedy- 
sarum. C. 

Ticia americana, Vetch. U. 

Geranium fremonti, Geranium. C. 

Geranium richardsoni, Geranium. C. 

Linum lernisi, Wild Flax. 



Savastana odorata, Holy Grass. 
Stipa nelsoni, Feather Grass. 
Muhlenbergia comata, Dropseed Grass. 
Agrosiis hiemalis, Bent Grass. C. 
Koeleria cristata, June Grass. U. 
Poa longipedunculata, Long-stalked 

Spear Grass. C. 
Poa lucida, Spear Grass. 
Particular ia nervata, Manna Grass. 



Opuntia fragilis, Small-jointed Cactus. 
V. 

Echinocactus simpsoni, Simpson Ball 
Cactus. C. 

Pachylophus montanus, Evening Prim- 
rose. U. 

Harbouria trachy < pleura, Water Hem- 
lock. C. 

Aralia nudicaulis, Wild Sarsaparilla. 

Gentiana affinis, Gentian. C. 

Ap'ocynum androswm i folium, Indian 
Hemp. 

Phlox depressa, Wild Phlox. 

Phlox hoodi, Wild Phlox. 

Gilki congesta, Gilia. 

Phacelia linearis, Phacelia. 

Phacelia heterophylla, Phacelia. 

Oreocarya virgata, Oreocarya. 

Monarda menthwfolia, Horse Mint. U. 

Pentstemon laricifolius, Beard-tongue. 

Adenostcgia ramosa. U. 

Scutellaria brittoni, Skullcap. U. 

Mimulus floribundus, Monkey Flower. 

Orthocarpus luteus, Yellow Orthocar- 
pus. 

Campanula rotundifolia, Harebell. C. 

Antennaria rcflexa, Everlasting. 

Balsamorrhiza sagittata, Balsam Boot. 

Balsamorrhiza iticaua, Balsam Boot. 

Chwnactis douglassi. 



Festuca ovina duriuscula, Fescue 
Grass. 

Bromus marginatus, Brome Grass. 
Bromus porteri, Brome Grass. 
Agropyron pscudorcpcns, Wheat Grass. 

C. 
Agropyron caninum, Wheat Grass. 
Agropyron spicatum, Wheat Grass. U. 



Agricultural Utility of the Transition Zone. 

The elevated Transition area with its vast extent of grazing lands 
is now, as in the past, the center of the sheep and cattle industries of 
Wyoming, and there are also extensive coal and oil fields in various 
stages of development. The principal timberlands in the Transition 
Zone are of small extent and lie to the east and northeast. They 
comprise a moderate growth of yellow 7 pine on the Laramie (see 



38 NORTH AMERICAN FAUNA. [No. 42. 

PL V, fig. 1), Casper, and Bear Lodge Mountains, and heavy forests 
of this valuable timber on the western slopes of the Black Hills. 1 

The climate is cool but dry and healthful, with a rainfall varying 
from 10 to 12 inches in the arid parts, 12 to 15 inches in the east-cen- 
tral and southeast sections, and 17 to 21 inches in the Black Hills and 
Bear Lodge districts at the northeast where crops succeed moderately 
in all except the sagebrush lands. The snowfall is rather heavy 
throughout. This quantity of moisture, though not large, in a meas- 
ure counterbalances the cool climate of the Transition Zone and, com- 
bined with the much greater area, gives this zone an agricultural 
value in Wyoming comparable to that of the warmer Upper Sonoran. 
Large areas of excellent grass land in the mountain valleys and on 
the higher plateaus in the upper part of the zone are ideal either as 
cattle range or for the summer grazing of sheep, while a fair growth 
of range grasses generally obtains even in the lower, more arid sec- 
tions. The rich soils yield abundantly either under dry farming 
or irrigation wherever there has been a proper selection of hardy, 
quick-maturing crops adapted to a usually short growing season. 

High-altitude farming has been thoroughly tested at the Wyo- 
ming Experiment Station at Laramie for a series of years. Very 
favorable results 2 were obtained with a great variety of vegetable and 
cereal crops and even certain fruits, in experiments conducted in the 
heart of the Transition Zone at over 7,000 feet elevation, where bleak 
winds are unusually prevalent. Apples and small fruits which are 
grown with considerable difficulty on the Laramie Plains succeed 
admirably in many localities with greater natural protection. Suc- 
cessful farming districts in the Wyoming Transition include sections 
of the Laramie Plains, the Platte Valley above Saratoga, Bear Val- 
ley and Fort Bridger region in Uinta County, Salt River Valley in 
Lincoln County, the eastern base of the Bighorn Mountains in Sheri- 
dan and Johnson Counties, and the Sundance region in Crook County. 
In most of these, oats and hardy cereals, alfalfa (two cuttings), field 
peas, potatoes, and hardy vegetables are raised to great perfection. 
Hay, forage, garden vegetables, and a limited crop of small grain are 
very generally grown on stock ranches even in the colder parts of 
the zone. 

CANADIAN ZONE. 

The Canadian Zone, the region of coniferous Boreal forest, is the 
most important of the Boreal transcontinental life areas. It extends 
far southward in the principal mountain masses of the Western 

1 Open sage country usually fills the zone on the basal slopes of the high ranges in 
western Wyoming. The growth of Douglas spruce along its upper border in this region 
is generally scattering, while the yellow pine is of rare occurrence. 

2 Discussed in bulletins of the Wyoming Experiment Station, which contain also valu- 
able cultural and other data based on tests made at the station at Laramie and at the 
experimental farms at Sundance and Saratoga. 



1917.] CANADIAN ZONE. 39 

States, and over much of the Rocky Mountain region covers the 
middle slopes of the high ranges and the summits and upper slopes 
of mountains of medium elevation. It is uniformly and conspicu- 
ously characterized from Montana to Colorado by forests of spruce, 
fir, lodgepole pine, and aspen, and by a large variety of Boreal under- 
shrubs and plants. 

In Wyoming the greatest extent of Canadian Zone country is in 
the mountainous northwest. Here the zone includes most of the 
extensive undulating forested plateau of Yellowstone Park; large 
rolling or hilly tracts of mixed forest and open country on the 
borders of Jackson Hole (PI. VIII, fig. 1), in the basin of Hoback 
River, at the head of Green River, and on the southern end of the 
Wyoming Range ; and the forested slopes of the Wind River, Absa- 
roka, Teton, Gros Ventre, Salt River, Snake River, and Wyoming 
Ranges (PL VIII, fig. 2) from near their bases to the upper limit 
of large tree growth at 9,000 to 10,000 feet elevation. Elsewhere 
the Bighorn and Medicine Bow Ranges and the Sierra Madre are 
extensively Canadian, and the Laramie Mountains and Casper Range 
have considerable areas on their summits. Elevations of medium 
altitude which are capped with Canadian Zone forests and also have 
small areas on their cool slopes are the Black Hills, the Bear Lodge, 
Rattlesnake, Green, Ferris, Seminole, and Shirley Mountains, the 
northern shoulders of the Uinta Mountains, and high plateaus along 
the southern boundary of the State between Green River and the 
Red Desert. Traces of the zone, indicated usually by aspens and 
lodgepole pines, or merely by dense, scrubby thickets of aspens, are 
on the upper, cool slopes of the Aspen Mountains, the Bear River 
Divide, a few desert peaks along the continental watershed between 
South Pass and Steamboat Mountain, on Heart Mountain north of 
Cody, and on Pyramid and Heaths Peaks along the upper Platte. 

The lofty Wind River Range of comparatively straight axis pre- 
sents a graphic view of the Canadian forest belt, which is main- 
tained usually at uniform elevation and in full vertical width (about 
2,000 feet) on both slopes, its lower border sharply defined where 
the forests meet the open basal sagebrush country of the Transition 
Zone. 1 The upper border of the Canadian Zone is obscure in these 
mountains, as elsewhere, the change from heavy forest growth to 
the narrow Hudsonian timberline belt of dwarfed forest being 
gradual and almost imperceptible. 

The main forest composition on the Wyoming mountains is very 
uniform, with forests of lodgepole pine and aspen in the lower half 
of the zone and a heavy stand of Engelmann spruce, or more often 

1 The lower edge of the Canadian Zone is less clearly marked on the eastern slopes of 
the Bighorn Mountains and elsewhere in eastern Wyoming where forests of yellow 
pine fill much of the upper Transition. 



40 NORTH AMERICAN FAUNA. [No. 42. 

a mixed forest of spruce and fir, higher up. Other trees of less 
extensive growth and more restricted distribution are the Douglas 
spruce in the lower part of the zone, chiefly in the northwestern 
mountains; the blue spruce and balsam poplar fringing streams 
along the lower edge of the zone in the south and west, and in the 
northwest, respectively ; and the canoe birch in the Black Hills and 
Bear Lodge Mountains. Fire-swept tracts usually are first covered 
with a dense growth of young aspens, which are of more rapid 
growth than conifers. In the Sierra Madre the fresh growth on 
burned-over areas is occasionally of fir over original lodgepole pine 
forest. 

In the lower part of the Canadian forest belt are considerable 
areas of partly open mountain meadows and parks, and more rarely, 
of open slopes. Characteristic tracts are the 8,000-foot watershed 
between the Hoback and Green Eivers, where groves of aspen inter- 
mixed with a little lodgepole pine and fir alternate over a gently 
rolling country with open parks covered with low matted sagebrush, 
Frasera, and Balsamorrhiza, or with beautiful grassy meadows bril- 
liantly colored in summer with flowers of shrubby cinquefoil, lark- 
spur, lupine, geranium, iris, and painted cup; the grassy meadows 
and bordering sage benches of the Du Noir Valley at the head of 
Wind River; extensive willow-grown meadows and flats at the north- 
ern end of Jackson Hole and in Yellowstone Park; and open grass 
or sage slopes on the western side of the Bighorn Mountains be- 
tween 8,000 and 9,000 feet elevation. On the moderately inclined 
eastern side of the Wyoming Range southwest of Bigpiney are 
unusually open mountain slopes. Here grass and sage country ex- 
tend in many places on ridges and south slopes to the 10,000-foot 
crest of the range, alternating regularly with dense tracts of Douglas 
spruce, lodgepole pine, and Engelmann spruce forest on all north 
slopes and in gulches. 

The usual factors of base level, latitude, slope incline and ex- 
posure, and, to a certain extent, air currents, affect in varying degree 
the altitude of the Canadian Zone. Base level appears to be more 
potent than latitude in Wyoming. The variation from 8,500 to 10,- 
500 feet on southwest slopes and 7,500 to 9,500 feet on northeast 
slopes in the mountains along the southern boundary and bordering 
Green River Basin, to 8,000 to 9,500 feet on southwest slopes and 
7,000 to 9,000 feet on northeast slopes in the northern mountains, is 
largely latitudinal. The depression to 6,000 feet on the cap of the 
Black Hills and Bear Lodge Mountains, however, is probably due 
to the low base level of the adjacent plains on the north and east. 

The high level of the Canadian Zone (above 8.000 feet) on the 
warm western slope of the Bighorn Mountains east of Ionia is due 
to bold southwest exposure, which more than offsets the lowering 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate VIII. 




Fig. 1.— Snake River Valley (6,500 Feet) near Moran, Jackson Hole. 

Mixed forest conditions at lower edge of Canadian Zone are shown. Scattered groves of aspen, 
balsam poplar, and blue spruce on valley flats; aspen and lodgepole pine forests on hills. 




Fig. 2.— Heavy Engelmann Spruce Forest Wyoming Range West of Merna 

(9,000 Feet). 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate IX. 




Fig. 1.— Sylvan Lake, Yellowstone National Park, in the Enqelmann Spruce 

Belt. 




Fiq..2.— Heavy Stand of Lodgepole Pine, North Slope of Ferris Mountains 

(8,500 Feet). 



1917.] CANADIAN ZONE. 41 

influence of the low altitude of the Bighorn Basin at their western 
bases. The Ferris and Green Mountains, small isolated ranges on 
the northern border of the Red Desert, show graphically the effect 
of slope exposure on zone level. The Ferris Range, of 9,500 feet 
elevation, is narrow and sharp-crested, with very abrupt southwest 
slopes facing the desert. These hot, exposed slopes are open Transi- 
tion Zone, and crowd the Canadian element above 9,000 feet, where 
it is indicated just below the summit by thickets of aspen and lodge- 
pole pine in gulches, and by Rocky Mountain white pine on exposed 
slopes and ridges. Just over the 9,500-foot crest on the cool exposure 
are the Engelmann spruces and firs of the Canadian Zone, somewhat 
dwarfed at first but soon encountered in large and dense growth. 
The spruces and firs, with lodgepole pines and aspens lower down, 
form a heavy forest on the northeast slopes down to 7,800 feet, while 
descending tongues in gulches along cold streams extend 500 feet 
lower. The Canadian Zone on the Ferris Range thus exhibits about 
the maximum variation in zone level due to slope exposure — fully 
1,500 feet. 

In Jackson Hole, the Yellowstone Valley, along both forks of the 
Shoshone River, and at the heads of many other narrow valleys 
deeply penetrating the mountain mass of northwest Wyoming the 
Canadian element reaches a low elevation, and on cold slopes is 
often unmixed with Transition species at 6,000 feet. Over this re- 
gion the mean summer temperature is low, the cold air of the sur- 
rounding mass of Boreal country settling into the valleys and can- 
yons at night, and frequent frosts occur during the warmest months. 
Furthermore, these mainly steep-walled valleys receive a minimum of 
sunlight, and many slopes are shaded during the warmest part of the 
day. Under these conditions the zone occasionally has a vertical 
breadth of nearly 3,000 feet, since the factors which cause the abnor- 
mal depression of its lower boundary do not appreciably affect the 
upper limit. 

The cooling influence of cold streams and of descending cold air 
currents which flow down gulches and canyons regularly carries nar- 
row tongues of Canadian Zone far below the average level on moun- 
tain slopes. This is very noticeable on some of the streams at the 
southwestern base of the Wind River Range. The clear icy waters 
of Pine Creek, the outlet of Fremont Lake, carry a broad fringe of 
lodgepole pines, aspens, and a pure Canadian undergrowth almost to 
its junction with the New Fork at 7,000 feet elevation, fully 1,000 
feet below the mean lower border of the zone on the southwest slope 
of these mountains. The waters of the Big Sandy, 40 miles to the 
southeast, also carry Canadian species far down into open Transition 
sagebrush country. Streams which break out of the steep northern 
escarpment of the Casper Range through deep, shaded gulches are 



42 NORTH AMERICAN FAUNA. [No. 42. 

heavily fringed with aspen groves and Canadian undershrubs on the 
otherwise open 6,000-foot basal slopes, 1,000 feet below the usual 
altitude of these species on the northern slopes of this range. 

The Canadian Zone is marked in Wyoming as a cool region of con- 
siderable humidity, but the percentage of bright sunny days through- 
out the year is high. Much of the precipitation is in the form of 
frequent sharp showers in summer and heavy snows in winter, but 
there are occasional rains of greater duration in spring and fall. 
The chief value of this zone is its natural adaptation, in abundant 
forest and plant cover and cool summer temperature, to moisture 
retention. 1 The extensive forests,, its greatest natural resource, are 
now largely under Federal control. These are of great value, espe- 
cially the forests of lodgepole pine and Engelmann spruce (PI. IX), 
but large areas are as yet inaccessible. Winter temperatures are low, 
occasionally reaching — 45° F., and in the small areas physically 
adapted to agriculture the prevalent summer frosts preclude the 
growing of more than a few of the hardiest crops and vegetables for 
ranch use. 

Characteristic Species — Canadian Zone. 

A rich fauna and flora uniformly characterize the Canadian Zone 
in Wyoming. Forest species predominate, many of them identical 
with or closely related to Boreal types of transcontinental range from 
Labrador to Alaska ; others are peculiar to the Rocky Mountain re- 
gion ; and a small number have a restricted range. The canoe birch, 
beaked hazelnut^ Douglas honeysuckle, northern chipmunk, and a 
few other species of wide Boreal dispersion occur only in the out- 
lying Black Hills and Bear Lodge Mountains at the northeast, and 
the northern varying hare only in the Bighorn Mountains, but 
elsewhere the characteristic species are very uniformly distributed in 
the different mountain ranges. With the exception of many of the 
breeding birds, which merely make their summer home in the Cana- 
dian Zone, all are species adapted to a region of long, cold winters 
and short, cool summers. 

Mammals — Canadian Zone. 

[Species marked T. occur also in the Transition Zone; those marked H., also in the 

Hudsonian.] 



Cervus canadensis canadensis, Elk. 

T, H. 
Aices americanus shirasi, Shiras 

Moose. 
Schtrits frcnionti frcmonti, Fremont 

Spruce Squirrel. 
Sciurus hudsonims baileyi, Bighorn 

Red Squirrel. 



Sciurus hudsonicus rentonan, Rocky 
Mountain Red Squirrel. 

Glaucomys sabrinus bangsi, Rocky 
Mountain Flying Squirrel. 

Eutamias borealis, Northern Chip- 
munk. 

Eutamias operariits, Colorado Chip- 
munk. 



i See pp. 52-53. 



1917.] 



CANADIAN ZONE. 



43 



Mammals — Canadian Zone — Continued. 



Eutamias umbrinus, Uinta Chipmunk. 
Eutamias liiteiventris, Buff -bellied 

Chipmunk. T. 
Callospermophilus lateralis lateralis, 

Say Ground Squirrel. T. 
Callospermophilus lateralis caryi, Wind 

River Mantled Ground Squirrel. 
Callospermophilus lateralis castanu- 

rus, Chestnut-tailed Ground Squir- 
rel. 
Marmata flaviventris nosophora, 

Golden-mantled Marmot. H. 
Marmot a flaviventris luteola, Park 

Marmot. H. 
Marmota flaviventris dacota, Black 

Hills Marmot. T. 
Peromyseus maniculatus artemisiw, 

Sagebrush White-footed Mouse. '/. 
Peromyseus maniculatus ruflnus, 

Tawny White-footed Mouse. 
Phenacomys orophilus, Mountain 

Phenacomys. 
Evotomys gapperi galei, Gale Red- 
backed Mouse. 
Evotomys brevicaudus, Black Hills 

Red-backed Mouse. 
Mierotus mordax mordax, Rocky 

Mountain Meadow Mouse. 
Mierotus longicaudus, Long-tailed 

Meadow Mouse. T. 
Mierotus richardsoni macropus, Big- 
footed Meadow Mouse. 
Mierotus nanus nanus, Dwarf Field 

Mouse. T, H. 
Castor canadensis, Beaver. T. 
Thomomys fossor, Colorado Pocket 

Gopher. 
Thomomys uinta, Uinta Pocket 

Gopher. H. 
Thomomys fuscus fuscus, Brown 

Pocket Gopher. 



Thomomys talpoides caryi, Bighorn 
Pocket Gopher. II. 

Zapus princeps princeps, Rocky Moun- 
tain Jumping Mouse. 

Erethizon epixanthum, Yellow-haired 
Porcupine. T. 

Lepus americanus americanus, North- 
ern Varying Hare. 

Lepus bairdii bairdii, Snowshoe Rab- 
bit. 

Fells hippolestes, Mountain Lion. T. 

Lynx canadensis canadensis, Canada 
Lynx. 

Vulpes maerourus. Mountain Red Fox. 

Mustela arizonensis, Arizona Wea- 
sel. T. 

Mustela cicognanii leptus, Dwarf Wea- 
sel. 

Mattes caurina origenes, Rocky Moun- 
tain Marten. 

Lutra canadensis canadensis, Otter. T. 

Gulo luscus, Wolverene. 

Ursus americanus, Black Bear. T. 

Ursus imperator, Yellowstone Park 
Grizzly Bear. T. 

Ursus washake, Washakie Grizzly 
Bear, Silver-tip. T. 

Sorex personatus personatus, Masked 
Shrew. H. 

Sorex obscurus obscurus, Rocky 
Mountain Shrew. H. 

Sorex vagrans dobsoni, Dobson 
Shrew. 

Neosorex palustris navigator, White- 
bellied Water Shrew. 

Nycteris cinerea. Hoary Bat. T. 

Lasionycteris noctivagans, Silver- 
haired Bat. T. 

Myotis lucifugus carissima. Little 
Brown Bat. T. 



Breeding Birds — Canadian Zone. 

[Species marked T. breed also in the Transition Zone; those marked H., also in the 

Iludsonian.] 



Pelecanus erythrorhynchos, White Pel- 
ican. 

Mergus americanus, Merganser. T. 

Clangula islandica, Barrow Golden- 
eye. H. 

Charitonetta albeola, Buffle-head. 



Rranta canadensis, Canada Goose. T. 
Olor buccinator, 1 Trumpeter Swan. T. 
Dendragapus ob s cur u s richardsoni, 

Richardson Dusky Grouse. T. 
Picoides arcticus, Arctic Three-toed 

Woodpecker. 



1 Probably no longer breeding in Wyoming. 



44 



NORTH AMERICAN FAUNA. 



[No. 42. 



Breeding Birds — Canadian Zone — Continued. 



Picoides americanus dorsalis, Alpine 
Three-toed Woodpecker. H. 

Sphyrapicus varius nuchalis, Red- 
naped Sapsucker. T. 

Sphyrapicus thyroideus, Williamson 
Sapsucker. 

Selasphorus platycercus, Broad-tailed 
Hummingbird. T. 

Kuttallornis borealls, Olive-sided Fly- 
catcher. 

Empidonax difflcilis, Western Fly- 
catcher. T. / 

Cyanocitta stelleri annectens, Black- 
headed Jay. 

Perisorcus canadensis capitalis, Rocky 
Mountain Jay. H. 

Carpodacus cassini, Cassin Purple 
Finch. 

Loxia curvirostra minor. Crossbill. 

Spinus pinus, Pine Siskin. 

Zonotrichia leucophrys, White-crowned 
Sparrow. H. 

J unco hyemalis mearnsi, Pink-sided 
Junco. H. 



J unco phceonotus caniceps, Gray-head- 
ed Junco. H. 

Mclospiza lincolni, Lincoln Sparrow. 

Wilsonia pusilla pileolata, Pileolated 
Warbler. H. 

Cinclus mexicanus unicolor, Dipper, 
Water Ouzel. T. 

N annus hiemalis pacificus, Western 
Winter Wren. 

Sitta canadensis, Red-breasted Nut- 
hatch. 

Penthestes gambeli, Mountain Chicka- 
dee. T. 

Regulus satrapa, Golden - crowned 
Kinglet. H. 

Reyulus calendula, Ruby -crowned 
Kinglet. 

Myadestes townsendi, Towusend Soli- 
taire. T. 

Ilylocichla ustulata swainsoni, Olive- 
backed Thrush. T. 

HylocicJila guttata auduboni, Audubon 
Hermit Thrush. 

Sialia currucoides, Mountain Blue- 
bird. T. 



Plants — Canadian Zone. 

[Species marked T. ooeur also in the Transition Zone; those marked IT., also in the 

Hudsonian.] 

Trees and shrubs. 



Pinus murrayana, Lodgepole Pine. 

Pinus flexilis. Rocky Mountain White 
Pine. T, H. 

Picea engelmanni, Engelmann Spruce. 

Picea parry ana, Blue Spruce. 

Abies (concolor ?), White Fir. 

Pseudotsuga mueronata, Douglas 
Spruce. T. 

Juniperus sibirica, Low Juniper. H. 

Populus tr< tiiuloides, Aspen. 

Populus balsamifera, Balsam Poplar. 

Salix pyrifolisi obscura, Willow. 

Salix nelsoni, Nelson Willow. H. 

Betula papyrifera, Canoe Birch. 

Betula glandulosa, Dwarf Birch. 

Corylus rostra ta, Hazelnut. T. 

Alnus tenuifolia, Alder. 

Ribes petiolare, Mountain Black Cur- 
rant. 



Ribes wolfl, Blue Currant. 

Ribes lacustre, Currant. 

Ribes viscosissimuin. Currant. 

Spircea lucida, Meadowsweet. T. 

Rubus parviflorus, Thimbleberry. 

Rubus strigosus. Red Raspberry. T. 

Dasiphora fruticosa. Shrubby Cinque- 
foil. H. 

Rosa sayi, Rose. T. 

Morbus scopulina. Mountain Ash. 

Pachystima myrsin ites. 

Rhamnus alnifolia. Buckthorn. 

Lepargyrea canadensis, Canadian Buf- 
faloberry. 

Ledum glandulosum, Labrador Tea. 

Mensiesia ferrugin ra. 

Vaccinium oreophilum, Mountain Blue- 
berry. 



1917.] 



CANADIAN ZONE. 



45 



Plants — Canadian Zone — Continued. 
Trees and shrubs — Continued. 



Vaccinium erythrococcum, Red Bil- 
berry. H. 

Vaccinium occidentale, Blueberry. 

Vaccinium cwspitosum, Blueberry. 

Sambucus microbotrys, Red Elder- 
berry. 



Lonicera glaucescens, Douglas honey- 
suckle. 

Lonicera involucrata, Involucred Fly- 
honeysuckle. 

Lonicera utahensis, Honeysuckle. 



Herbaceous plants. 



Veratrum tenuipetalum, White Helle- 
bore. 

Zygadenus elegans, Beautiful Camas. 
H. 

Calochortus pavonaccus, Yellow Mari- 
posa. 

Erytlironium parriflorum, Dog-tooth 
Violet. H. 

Streptopus amplexifolius, Twisted- 
stalk. 

Disporum trachyearpum. T. 

Limnorchis borealis, Bog Orchid. 

Calypso bulbosa, Calypso. 

Rum ex paucifolius, Dock. H. 

Claytonia rosea, Spring Beauty. H. 

Aetata arguta, Baneberry. 

Aetata rubra, Baneberry. 

Aquilegia cwrulea, Blue Columbine. 

Aquilegia flavescens, Yellow Colum- 
bine. 

Aquilegia oreophila, Columbine. H. 

Anemone lithophila, Anemone. 

Aconitum columhianum. Monkshood. 

Clematis pseudalpina, Purple Virgin's 
Bower. 

Ranunculus inamemus, Crowfoot. 

Thlaspi glaucum, Penny Grass. H. 

Parnassia flmbriata, Grass-of-Parnas- 
sus. 

lleuehera parvifolia, Alum Root. II. 

Micranthes arguta, Saxifrage. 

Fragaria paueiflora. Strawberry. T. 

Sieversia cUiata., Mountain Avens. 

Astragalus alpinus. Milk Vetch. 

Trifolium rydbergi, Clover. T. 

Trifolium ancmophilum. Clover. T. 

Geranium parryi, Geranium. 



T. 



T. 



Geranium cwspitosum, Geranium. 

Viola canadensis rydbergi, Violet. 

Viola, bellidifolia, Violet. H. 

Epilobium spp. 

Heracleum lanatum. 

Pyrola secunda, Wintergreen. 

Pyrola uliginosa, Wintergreen. 

Pyrola. chlorantha, Wintergreen. 

Pyrola elliptica, Wintergreen. 

Pyrola pieta, Painted Wintergreen. 

Pterospora andromeda, Pinedrops. 

Chimaphila umbellata, Pipsissewa. 

Moneses uniflora, One-flowered Win- 
tergreen. 

Gentiana forwoodi, Closed Gentian. 

Gentiana elegans, Mountain Fringed 
Gentian. 

Frasera speeiosa, Frasera. 

Pentstemon glaucus. Beard-tongue. 

Pentstemon fruticosus, Beard-tongue. 
T. 

Mimulus langsdorfii, Monkey Flower. 

Mini til us letcisi, Crimson Monkey 
Flower. 

Elephant clla groenlandica, Elephant- 
head. H. 

Pedieularis raeemosa, Purple Louse- 
wort. 

Castilleja sulphured. Painted Cup. 

Linnwa americana, Twinflower. 

Valeriana aeutiloba, Valerian. If. 

Anaphalis snbalpina. Pearly Everlast- 
ing. 

Antennaria puleherrima, Everlasting. 
T. 

Wyethia ample.ricaulis. 

Arnica cordifolia, Heart-leaved Arnica. 



46 



NORTH AMERICAN FAUNA. 



[No. 42. 



Plants — Canadian Zone — Continued. 
Grasses. 



Phleum alpinum, Alpine Timothy. H. 
Xlopecurus occidental is, Mountain 

Foxtail. 
Catamagrostis canadensis acuminata, 

Mountain Reed Grass. 
Deschampsia ccespitosa, Tufted I lair 

Grass. T. 



Danthonia intermedia, Mountain Oat 

Grass. 
Poa reflecea, Mountain Bluegrass. II. 
Festuca thurberi, Thnrber Fescue. 
Agropyron violaceum, Wheat Grass. 

H. 
Elymus glaueus, Rye Grass. T. 



HUDSONIAN ZONE. 

On the high mountain ranged of the Western States the Canadian 
forest belt is fringed along its upper edge by the narrow Hud- 
sonian strip of dwarfed forest and depauperate vegetation, which 
gives way to bare Alpine slopes along a sharply defined climatic 
boundary known as timberline, corresponding to the transconti- 
nental limit of trees at the southern edge of the Arctic tundras. 
The Hudsonian Zone occupies a relatively small area in Wyoming. 
It encircles the Alpine summits and crests of the northwestern 
ranges, including the Bighorn Mountains, and small areas cap all 
ranges or isolated peaks which approximate timberline altitudes. A 
belt bounds the Alpine cap of the Snowy Mountains at the northern 
end of the Medicine Bow Range, and small, widely separated tracts 
or islands are on Bridger Peak and neighboring summits of the 
Sierra Madre, on Elk Mountain, and on Laramie Peak. 

In the southern mountains the forests commence to dwarf almost 
imperceptibly at from about 10,000 to 10,500 feet altitude, accord- 
ing to slope exposure, the elevation decreasing with higher latitude 
to 9,000 or 9,500 feet in northern Wyoming. The vertical breadth 
of the Hudsonian belt varies with slope and soil conditions from a 
few hundred to 1,000 feet. The peculiar ruggedness and broken, 
incised character of many mountains greatly obscures this belt, 
while in places sheer, perpendicular cliffs and avalanches greatly 
contract it or even narrowly interrupt continuity. Avalanches 
sweep away every vestige of tree growth and in many places replace 
the original soil with extensive fields of slide rock. The result is 
not a climatic change, however, and can not be considered in delimit- 
ing the zone. The Hudsonian strip is usually widest in cold gulches 
with abundant soil, and is narrow on exposed, scantily soiled ridges. 
In fact, soil conditions often counterbalance the elevating influence 
of slope exposure, and tree growth may be found as high on cold 
well-soiled slopes as on warm rocky inclines. 

An almost Arctic climate prevails in the Hudsonian Zone, which 
in winter is buried under deep snow and in summer is flecked with 
huge drifts, many of which never entirely leave protected gulches. 
The deep-soiled slopes are thoroughly saturated in summer by 



1917.] HUDSONIAN ZONE. 47 

melting snow and frequent showers and squalls of snow or sleet, and 
bogs and small lakes abound in all level situations and natural 
basins. Exposed to high winds throughout the year, the conifers 
are in ragged, fantastic, and usually one-sided growth, and, along 
with the shrubs and many larger plants, evidence an adverse cli- 
mate in stunted and otherwise deficient development. (PI. X.) The 
middle of May found the timberline region on the Wind River 
Range near Dubois still in the grasp of wintry weather, with few 
plants in flower and the low willows and shrubby cinquefoil not 
yet in leaf; while on the Bighorn Mountains east of Hyattville the 
alpine willows had not put forth leaves by June 5. Many plants 
were past flowering, and herbaceous vegetation was partly dried 
up on the Wyoming Range at 10,400 feet altitude, August 9; and 
the timberline slopes on the Tetons were sere and brown on August 
30 except for the hardy, late-flowering blue gentians. 

Although the climate is rigorous for eight months of the year 
and myriads of mosquitoes greet one in summer, this highly inter- 
esting region well repays the arduous climb entailed to reach its 
confines. The gently rounding crests of the Wyoming Range south 
of Hoback Peak are peculiarly attractive. Grassy openings and 
parks mingle with scattered clumps and mats of Engelmann spruce 
and alpine fir; fields of scarlet painted cup, blue larkspur, white 
columbine, and purple lupine enliven a landscape flecked with white 
banks of melting snow; and a moderate incline adds to the beauty 
of the region. The Hudsonian area on the southwest side of the 
Wind River Range is of very different character. South of Fre- 
mont Peak (PI. XI, fig. 1) it occupies a sloping granitic plateau a 
mile or two wide, and between 10.500 and 11.200 feet altitude, at the 
base of abrupt Alpine peaks of nearly 14,000 feet elevation. Its ex- 
ceedingly rough surface of a seemingly endless succession of bare 
granite hummocks, studded with countless clear, snow-fed lakes in 
rocky basins, makes travel extremely arduous. Vegetation is scanty, 
and coniferous growth is very scattering over this unusually rocky 
region. Hudsonian vegetation is most abundant on deep-soiled 
slopes, as on Whiskey Mountain in the Wind River Range south of 
Dubois, and along the eastern slope of Needle Mountain in the 
Absaroka Range (PI. XI, fig. 2). 

Characteristic Species — Hudsonian Zone. 

Trees of the timberline belt in Wyoming are the alpine fir, white- 
barked pine, and Engelmann spruce. The first two are characteristic 
Hudsonian trees, but the spruce extends up from the Canadian Zone. 
The spruces and firs, mainly on cool exposures, exhibit to a marked 
degree the depressed growth due to high altitude and are usually 
prostrate mats at extreme timberline. The white-barked pines of the 
northwest ranges show less dwarfing, but usually are very ragged 



48 NORTH AMERICAN FAUNA. [No. 42. 

and one-sided as a result of the prevalent winds. They occur in 
scattering growth or as compact tongues push up warm exposed 
slopes and ridges, and even form forests of considerable size a little 
below timberline, as on Whirlwind Peak, in the Absaroka Range. 
The Rocky Mountain white pine, although a tree usually found only 
at a lower altitude, was found near timberline on the Bighorn Moun- 
tains, and also on the exposed 10,000-foot summit of Laramie Peak, 
where its characteristic growth is in depressed mats at the base of a 
low central bole 6 feet or more in height. Prostrate clumps of low 
juniper are common in slide rock at timberline on the Sierra Mad re, 
on the Bighorn and Wind River Ranges, and elsewhere. 

Although prominently characterized by depauperate vegetation, 
the grassy slopes between the scattered clumps of conifers are hand- 
somely carpeted with a wealth of small flowering plants. Conspicu- 
ous flowers of spring and early summer, as observed on the Wind 
River and Bighorn Ranges, include globe flower, mountain cowslip, 
shooting star, columbine, spring beauty, and various mertensias, 
Jacob's ladder, forget-me-nots, buttercups, saxifrages, and drabas. 
In early autumn the timberline region on the Wyoming and Ab- 
saroka Ranges was brilliant with flowering mats of lupine, larkspur, 
painted cup, mountain heath, and mountain laurel, with the more 
scattering Parry primroses, harebells, gentians, phloxes, and ryd- 
bergias. 

Many of the plants of the Hudsonian Zone and most of the birds 
and mammals occur also in the adjoining Canadian or Arctic- Alpine 
Zones. Comparatively few species are closely restricted to this 
narrow area. 

Mammals — Hudsonian Zone. 

Mammals having their center of abundance at or near timberline 
are the mountain sheep (Oris canadensis canadensis) , the timberline 
chipmunk (Eutamias oreocetes), marmots (Marmota flaviventris 
nosophora and M. f. luteola), and the pika. or coney (Ochofona 
uinta). A number of species range into this region from the Cana- 
dian Zone, or make their homes in both areas. Those occurring thus 
with some regularity are : 



Sciurus hudsonievs ventorum, Rocky 
Mountain Red Squirrel. 

Callospermophilns lateralis: eargi. Wind 
River Mantled Ground Squirrel. 

Evotoings gapperi galei, Gale Red- 
backed Mouse. 

Microtus mordax mordax, Rocky Moun- 
tain Meadow Mouse. 

Microtus nanus nanus. Dwarf Field 
Mouse. 

Thomomys uinta, Uinta Pocket Gopher. 



Thomomys talpoidcs caryi. Bighorn 
Pocket Gopher. 

Lepus bairdii bairdii, Snowshoe Rab- 
bit. 

Canis testes, Mountain Coyote. 

Vulpes macrourus, Mountain Red Fox. 

Mattes catirina origenes, Rocky Moun- 
tain Marten. 

Boreas obscurus obscurus, Rocky Moun- 
tain Shrew. 

Sorex personatus personatus. Masked 
Shrew. 



North American Fauna No. 42, U. S. Dept Agr. Biological Suivey, 



Plate X. 




Fig. 1.— Engelmann Spruces at Timberline, West Slope of 
Whirlwind Peak, Absaroka Range (10,000 Feet). 




Fig. 2.— White-Barked Pines, Same Locality. 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate XI. 




Fig. 1.— Lower Edge of Hudsonian Zone, Wind River Range South of Fremont 

Peak (10,500 Feet). 




Fig. 2.— East Slope of Needle Mountain, Absaroka Range. 

Growth of Engelinann spruce, alpine fir, and grav-leaved willow (Salix glaucops) &t timberline 

(10,000 feet). 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate XII, 




Fig. 1.— Arctic-Alpine Zone, Wind River Range. 
South of Fremont Peak, from 11,000 feet elevation, July 17, 1911. 




Fig. 2.— Arctic-Alpine Zone, Absaroka Range. 

Between the Greybull and the South Fork of Shoshone River, from east slope of Needle Mountain 

(10,500 feet), July 11, 1910. 



1917.] 



HUDSON' I AX /ONE. 



41) 



BBEEDING BlKDS — HUDSONIAN ZONE. 

[Species marked C. breed also iu the Canadian Zone.] 



Picoides americawus dorsaliSj Alpine 
Three-toed AYoodpecker. C. 

Perisoreus canadensis capitalis, Rocky 
Mountain Jay. C. 

Nucifraga colutnbiana, Clark Nut- 
cracker. 1 

Pinicola enucleator montana, Rocky 
Mountain Pine Grosbeak. 

Zonotrichia leucophrys, White-crowned 
Sparrow. C. 



Junco hyemalis meamsi, Pink-sided 
Junco. C. 

Junco phwonotus caniceps, Gray-headed 
Junco. C. 

Wilsonia pusilla pileolata, Pileolated 
Warbler. C. 

Certhia. familiaris molilalia. Rocky 
Mountain Creeper. C. 

Regulus satrapa, Golden-crowned King- 
let. C. 



Plants — Hudsonian Zone. 

[Species marked ('. occur also in the Canadian Zone; those marked A., also in the 

Arctic-Alpine.] 



Vuius albicaulis, White-barked Pine. 

ricea engelmanni, Engelmann Spruce 
(dwarf). C. 

Abies lasiocarpa, Alpine Fir (dwarf). 
C. 

Juniperus sibirica, Low Juniper. G. 

Salix glaueops, Gray-leaved Willow. 

Salix chlorophylla, Willow. C. 

Salix sarimontana, Willow. A. 

Ribes montii/cnum, Bristly Red Cur- 
rant. 

Rubus strigosus, Red Raspberry. C. 

Polygonum bistortoides, Twisted Po- 
lygonum. C. 

Claytonia rosea, Spring Beauty. 

Calandrinia pygmcea, Alpine Bitter 
Root. 

Cerastium beeringianum, Mouse-ear 
duckweed. 

Caltha leptoscpala, Mountain Cow- 
slip. 

Trollius albiflorus, Globe Flower. 

Aquilegia saximontana, Columbine. 

Delphinium subalpinum, Larkspur. 

A lie hi one tetouensis, Anemone. 

A n cm one globosa, Anemone. G. 

Ranunculus alpeophilus, Buttercup. 

Ranunculus calthcefolius, Buttercup. 

Thlaspi glaucum, Penny Grass. 

Draba luteola, Whitlow Grass. 

Clementsia rhodantha, Red Orpine. .4. 

Leptasea hirculus, Saxifrage. 

Potentilla glaucophylla, Cinquefoil. 
C. 



Drymocallis pseudorupestris, Avens. 
Lupinus ccespitosus, Lupine. G. 
Lupinus laxiflorus, Lupine. C. 
Trifolium dasyphyllum, Dwarf Clover. 

A. 
Trifolium parryi, Parry Dwarf Clover. 

A. 
Hedysa ru m s ulphun set m s. 
Angelica roseana. 
Phyllodoce empetriformis, Mountain 

Heath. 

Kahnia polifolia, Mountain Laurel. 
Primula parryi, Parry Primrose. 
. [iid rosace su b u m bell a ta. 
Dodecatheon radicatum, Shooting Star. 
C. 

Gcntiana calycosa, Gentian. 
fit ut in mi st rift i flora. Gentian. 
Sircrtia congesta, A. 
Swertia palustris, G. 
Phlox qwstritosa, Phlox. ('. 
Polemonium viscosum, Jacob's Ladder. 
Polemonium mellitum, Jacob's Ladder. 
Phacelia sericea, Silky Phacelia. A. 
Myosotis alpestris, Forget-me-not. A. 
Uertensia tweedyi, Lungwort. G. 
Pentstemon alpinus, Beard-tongue. 
Veronica toormskjoldi, Alpine Speed- 
well. 

Veronica serpyllifolia, Speedwell. C. 
Castilleja spp., Painted Cup. 
Campanula parryi, Parry Harebell. 
Rydbergia grandiftora, Rydbergia. A. 



Young noted in summer. 



74410° 



50 



NORTH AMERICAN FAUNA. 



[ No. 42. 



Plants — Hudson i an Zonk — Continued. 



Townsendia parryi, Parry Townsendia. 

A. 
Senecio frembnti, Paintbrush. A. 
Senedo crassulus, Paintbrush. 
Antennaria rcfU.ra. Everlasting. 
Erigeron compositus, Fleabane. C. 
Erigeron salsuginosus, Fleabane. 



Chwnactis alpina, Alpine Chsenactis. 

Trisetum subspicatum, Oat Grass. 

Poa epilis, Bluegrass. A. 

Phleum alpinum, Alpine Timothy. C 

Carex nigricans, Sedge. 

Carex nova, Sedge. 

Juncus subtriflorus, Rush. 



ARCTIC-ALPINE ZONE. 

The area above timberline on the highest mountains, the Arctic- 
Alpine Zone, corresponds in climate, and in plant species especially, 
to the barren grounds of the Arctic. It is not continuous from one 
range to another in Wyoming, for even in the mountainous northwest 
the areas are separated, often widely, by Canadian Zone valleys and 
forested divides of medium elevation. 

A wide, almost unbroken stretch of this zone caps the massive 
Wind River Range for its entire length (PL XII, fig. 1), and there 
is a broken, irregular area of equal extent on the main crest and 
primary spurs of the rugged Absaroka Range (PL XII, fig. 2). 
The Gros Ventre Range is capped by a succession of Alpine plateaus 
sloping moderately toward the north, while the lofty peaks and 
jagged crests of the Tetons form a narrow strip of Alpine country. 
Broad Alpine areas on the Bighorn Mountains lie south of the gap 
at the head of Tongue River, and include all elevated summits of 
the Cloud Peak group. There are traces of the zone, too small for 
plotting on the map (see frontispiece), on a few mountains in the 
eastern and northwestern borders of Yellowstone Park. In southern 
Wyoming the Arctic-Alpine Zone is restricted to the lofty plateau 
on the Snowy Mountains, at the northern end of the Medicine Bow 
Range. 

This high-altitude area is a bleak, wind-swept region of excessive 
snowfall in winter and frequent squalls of rain, sleet, or snow in the 
short summer, and arctic temperatures prevail throughout the year. 
On all the ranges snow fills the gulches and partly covers cold slopes 
and declivities even in the warmest months, while such massive, 
elevated ranges as the Wind Rivers. Absarokas, and Tetons carry ex- 
tensive snow fields, and even a few perennial ice fields or glaciers in 
protected Alpine valleys. 

The Arctic- Alpine Zone is conspicuously marked by the absence of 
tree growth, which ceases at its lower border. The altitude of 
timberline varies with latitude and slope exposure from 10.500 or 
11,000 feet in the Sierra Mad re and the Medicine Bow Range at the 
south, to 9,500 or 10,000 feet in the Bighorn Mountains and in the 
Yellowstone Park region. 



1917.] ARCTIC-ALPINE ZONE. 51 

Characteristic Species — Arctic-Alpine Zone. 

The season of plant growth is from the middle of May until August. 
but during this brief period the bleak slopes and even the fields 
of slide rock for 1,500 or 2,000 feet above timberline are bedecked 
with a profusion of bright-hued Alpine flowers. A luxuriant growth 
of Alpine grasses and sedges obtains wherever there is any depth of 
soil, and furnishes rich pasturage for mountain sheep and a few other 
mammals which spend the summer in this usually forbidding region. 
In Wyoming plant growth rapidly decreases in size above 12,000 feet. 

The low shrubby or matted growth usually extending from 500 to 
1,000 feet above the limit of trees consists chiefly of dense thickets of 
willow (Salix glaucops) and copses of shrubby cinquefoil (Pasi- 
phora frutiaosa) which push up the bottoms and along the margins 
of wet gulches and basins; mats of mountain heath (Phyllodoce em- 
petriformis) and alpine avens on rocky slopes and ridges; and spiny 
red currant {Ribes montigenum) and dwarfed raspberry (Rubus 
strigosns) in slide rock. Dwarf alpine willows mat the ground in 
places for an indefinite distance above timberline, and may even 
reach the highest summits with the mosses and lichens. 

A large number of characteristic Arctic- Alpine herbaceous plants 
mark the zone in Wyoming, but only four species of breeding birds 
are peculiar to it, and no mammals. 

Mammals — Arctic-Alpine Zone. 

The few mammals found in the Arctic- Alpine Zone in Wyoming 
belong to lower zones. Among those attracted in summer to its 
grassy slopes and crests are the elk {Cervu-s canadensis canadensis), 
mountain sheep (Ovis canadensis canadensis), timberline chipmunk 
(Eutamias oreoeete*), and Wind River mantled ground squirrel (Cal- 
lospermophilus 1. caryi). Marmots (Marmota f. noso/>hora), pocket 
gophers (Thonwniys uinta and T. f. fuscus), meadow mice (Microtus 
m. mordax), coneys {Ochotona uinta). and rarely the Uinta spermo- 
phile (('/fellas armatus), apparently are resident in the lower part of 
the zone in different localities. The coyote (Canis It stes?) and moun- 
tain red fox (Vulpes maerourus) range at various times into Alpine 
country in search of prey. On the Wyoming mountains mammals are 
rarely met with above 12.000 feet. 

Breeding Birds— Arctic-Alpine Zone. 



Lciteosticte atrata,' Black Rosy Finch. 
Anthus rubesccns? Pipit, Titlark. 
Otocoris alpestris leucolwma* Deserl 
Horned Lark. 



Lagopus lencurus leucurus, 1 White- 
tailed Ptarmigan. 

Leucosticte australis? Brown-capped 
Rosy Finch. 

1 On the Medicine Bow Range. 

2 On the Teton, Wind River, and Absaroka Ranges. 

3 Throughout the mountains. 

4 One of the few nesting birds of the Arctic-Alpine, but belonging chiefly to the Transi 
tion and Upper Sonoran Zoms. 



52 



NORTE AMERICAN FAUNA. 



[No. 12 



Plants — Abctic- Alpine Zone. 

[Species marked //. occur also in the Hudsonian Zone.] 
Aragallus nanus, Loco. 



Salix petrophila, Rock Willow. 

Salix latent. Rock Willow. 

Salix nivalis, Alpine Willow. 

Salix saximontdna, Creeping Wil- 
low. //. 

Oxyria digyna, Mountain Sorrel. IT. 

Polygonum viviparum, Alpine Knot- 
weed. 

Claytonia megarrhiza, Arctic Spring 
Beauty. 

Spraguea multiceps. / 

Silt lie acaulis, Stemless Catchfly. II. 

Alsinopsis obtusiloba, Sandwort. 

Paronychia pulvinata, Whitlowwort. 

Ranunculus adoneus, Buttercup. II. 

Thalictrum alpinum, Alpine Meadow 
Rue. 

\ni his lyalli. 

Smelowskia americana. 

Draba caria, Whitlow Cress. 

Draba crassifolia, Whitlow Cress. 

Draba densifolia; Whitlow Cress. 

Parrya nudicaulis, Purple Parrya. 

Rhodiola integrifolia, Rosewort. 

Saxifraga cernua, Arctic Saxifrage. 

Leptasea flagellaris, Saxifrage. 

Boykinia heuclieriformis, Saxifrage. 
Boykinia. //. 

Dryas octopetala, Alpine Avens. 

Sibbaldia procumbens, Sibbaldia. H. 

Sievcrsia t u r b i n a t a , Mountain 
Avens. //. 

Lupinus m o n ticola, Mountain Lu- 
pine. IF. 



Aragallus lagopus, Loco. 

Bupleurum americanum, Thorough- 
wax. 

(Jreoxis alpina. 

Androsace carinata. 

Gentiana romanzovi, Dwarf Closed 
Gentian. 

Polemonium confertum, Jacob's Lad- 
der. H. 

Eritrichium argenteum, Alpine Forget- 
me-not. 

Mertensia brevistyla and others. 

Pellicula ris parryi, Parry Louse- 
wort. H. 

Besseya alpina. 

Campanula tttti flora, Arctic Harebell. 

Solidago decumbens, Goldenrod. 

Erigcron pinnatisectus, Fleabane. //. 

Erit/cron radicatus, Fleabane. 11. 

Toncstus pygmcBUS. 

Achillea alpicola, Alpine Yarrow. 

Artemisia scopulorum, Alpine Sage- 
brush. 

Festuca brachyphylla, Alpine Fescue. 

Poa arctica, Arctic Bluegrass. 

Poa lettermanni, Bluegrass. 

Poa alpina, Alpine Bluegrass. II. 

Carex engelmanni, Sedge. 

Carer albo-nigra, Sedge. 

Caret- nubicola, Sedge. 

Carer at rata. Black Sedge. 

Carex ph&ocephala, Sedge. 

Juncoides spicatum, Wood Rush. II. 



IMPORTANCE OF BOREAL ZONES TO WYOMING AND ADJOINING 

AREAS. 

Climatically and physically unsuited to agriculture, the high alti- 
tude Canadian, 1 Hudsonian, and Arctic- Alpine Zones nevertheless are 
not only a valuable but an essential complement to the lower agricul- 
tural areas of Wyoming and most adjoining States. As the chief 
sources of three great river systems — the Columbia, the Missouri, and 
the Colorado — their importance is far from local. The great value of 
the Boreal zones lies in their peculiar adaptation to moisture con- 
servation. This is accomplished climatically on the bleak and bar- 
ren Alpine slopes and summits, and by plant and forest cover and 
climate combined in the Hudsonian and Canadian Zones. 



1 See p. 42. 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate XIII. 




Fig. 1.— Teton Range, Mount Moran South to Grand Teton. 
Photograph taken from foot of Jackson Lake, June 5, 1911, by Edward A. Preble. 




Fig. 2.— Snow in Lower Part of Hudsonian Zone (10,800 Feet). 
East slope of Bridger Peak, Sierra Madre,July 7, 1911. 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate XIV. 







Fig. 1.— Heavy Forest of Engelmann Spruce, Canadian Zone, Wyoming Range 
West of Merna (9,000 Feet). 




Fig. 2.— Rank Vegetation on Floor of Canadian Zone Forest, Grinnell Creek, 
Absaroka Range (7,500 Feet). 



1917.] BOREAL ZONES. 53 

A heavy mantle of snow covers the-entire Arctic-Alpine cap of the 
main ranges for seven or eight months of the year ( PI. XIII, fig. i), 

while huge drifts and snow fields remain in protected spots through- 
out the summer. In the timberline region shaded gulches are banked 
with snow until August, and it is not uncommon to find snowdrifts 
in the forest depths of the Canadian Zone until midsummer (PI. 
XTII 5 fig. 2), although most of the snow on the lower slopes goes off 
in flood in May and early June. The summer precipitation is heavy, 
consisting of rain in the forest belt, and frequent showers and squalls 
of rain, sleet, and snow on the peaks and higher slopes. 

The rotting vegetation and mellow soil of the cool mountain 
forests are specially adapted to the retention of this moisture, much 
of which works down from the slowly melting snow banks in the 
Alpine area. The shaded, mossy forest floor soon becomes saturated, 
and the water, percolating through the leaves and loose soil, finds its 
way gradually through rock crevices into ravines and depressions, 
finally flowing clear and sparkling into the streams lower down. Thus 
is insured an abundance of pure mountain water to the arid but fertile 
valleys and plains, and, what is of greatest importance, a fairly uni- 
form volume in the streams toward the end of the growing season, the 
period when most required by crops. The conversion through irriga- 
tion of portions of the valleys of the Snake. Yellowstone, Bighorn, 
Green, and North Platte Rivers in Wyoming, and especially in 
neighboring States, as well as of vast tracts as yet undeveloped, into 
rich agricultural districts is made possible through the combined 
agency of the climate and the forest and plant cover of the Boreal 
zones of Wyoming. 

The Canadian Zone has large tracts of forest, mainly of lodgepole 
pine and Engelmann spruce (PL XIV, fig. 1), with considerable 
Douglas spruce on the lower slopes. While these are useful for lum- 
ber and other utilities, the intrinsic value is small in comparison with 
the permanent service they are naturally fitted to perform in connec- 
tion with the agricultural utilization of the arid regions. Fortun- 
ately, most of the timberlands of the Canadian Zone in Wyoming are 
already included in national forests. Forest control with a view to 
their conservation is therefore most timely. 

While the higher mountain slopes of Wyoming afford a rich pas- 
turage during the summer months for many hundreds of thousands 
of sheep, careful regulation of sheep grazing is of the utmost impor- 
tance, as the natural plant cover is a vital factor in catching and 
holding moisture. (PL XIV, fig. 2.) Once this is badly broken up 
or removed by overgrazing the rains go off with a rush, carrying 
much of the soil with them. This results in dry, barren slopes, dirty 
streams, and a greatly diminished flow of water in the lower country 
during much of the growing season, unless there are adequate water- 



54 NORTH AMERICAN FAUNA. [No. 42. 

storage facilities. Mountains which hare been extensively grazed in- 
clude the Wyoming and Salt River Ranges, the southern end of the 
Wind River Range below the Big Sandy. Sierra Madre, and the 
mountains on either side of the Bighorn Basin at its southern end. 
The mountain meadows and parks with their luxuriant grasses con- 
stitute an ideal summer range for cattle, and are extensively utilized 
for this purpose. 

Wyoming offers many attractive regions to the tourist, the sports- 
man, and to those in search of health or recreation. As a permanent 
pleasure ground the mountainous region at the northwest is a val- 
uable asset, and is perhaps unsurpassed in extent and rugged 
grandeur. Dashing trout-filled streams add to the attractiveness 
of a section full of wild charm and beauty, while the dense forests 
of Yellowstone Park and the northern end of Jackson Hole afford a 
safe retreat and breeding range, under Federal and State protection, 
for many thousands of elk and other large game animals, and insure 
the best of hunting in season in districts adjacent to these protected 
areas. 



NOTES ON THE DISTRIBUTION OF CONSPICUOUS TREES AND 
SHRUBS OF WYOMING. 

The following annotated list of Wyoming trees and shrubs, while 
very incomplete, includes principally the more conspicuous and char- 
acteristic zone species, and should add to the knowledge of their dis- 
tribution within the State. It is based chiefly on notes and specimens 
collected by Biological Survey field parties. 

Pinus albicaulis Eugelmann. White-Barked Pine. (Fig. 3.) 

The small white-barked pine in Wyoming is peculiar to the high altitudes of 
the northwest, where it is a characteristic tree of the Hudsonian Zone just 
below timberline. It oc- 
curs on all the lofty 
mountains, finding its 
southern limits in the 
main chain of the Rock- 
ies on the Wind River 
and Salt River Ranges. 
At timberline it is often 
the most abundant tree, 
especially on parts of the 
Absaroka Range, but usu- 
ally shares this bleak 
region with dwarfed al- 
pine firs and Engelmann 
spruces, pushing up dry 
slopes and crests of ex- 
posed ridges in ascend- 
ing tongues, while the 
spruces and firs occupy 
wet gulches and the 
deeper-soiled slopes. 

Pinus flexilis James. 

Rocky Mountain White 

Pine. 

The Rocky Mountain 
white pine has a general 
dispersion in upper 
Transition and Canadian 
Zones in all except the 
northeast corner of the 
State. It has its center 
of abundance with the Douglas spruce along the lower edge of the Cana- 
dian forest belt, occupying the ridges and dry slopes, while the spruces are 
in gulches and on steep, cold exposures. There are few gravel or rocky ridges 
on the high central and western plains and deserts that do not have more or 
less of this pine in scattered and usually somewhat ragged growth. In 
Wyoming it is a small tree, rarely attaining a height of more than 30 or 40 
feet or a diameter above 1* feet under most favorable conditions. Its usual 

55 




Fig. 3. — Forest of white-barked pine (Pinus albicaulis) 
just below timberline on west slope of Whirlwind Peak, 
Absaroka Range (9,800 feet). 



56 



NORTH AMERICAN FAUNA. 



[No. 42. 



habit of growth is scattering and patchy, lmt hi deep-soiled mountain districts 
it often produces groves <>(' considerable extent. 

Pinus scopulorum (Engelmann) Lemmon. Rocky Mountain Yellow Pine. 

The distribution in Wyoming of the Rocky Mountain yellow pine, a well- 
known Transition Zone tree, is mainly west to the eastern slopes of the 
Bighorn Mountains and " the region of the upper Platte, as follows : Black 
Hills, he-ivy open forest to 6,000 or 6,500 feet altitude; Bear Lodge Moun- 
tains, moderate growth with oaks, 3,500 to 6.000 feet; Colony, low ridges; 
throughout the borders of Cheyenne River drainage, in scattering pockets and 
fringe; watersheds between Belle Fourche and Tongue Rivers, in thin forest 

over roughest sections ; 
lower eastern slopes of 
Bighorn Mountains, tol- 
erably wide belt up to 
6,000 or 7,000 feet; foot- 
hill region of Casper 
Mountains south through- 
out the length of the 
Laramies, including the 
Hartville group east of 
the Platte, in usually 
good growth ; divide 
southeast of Efell ; Pine 
Mountain south of Na- 
trona ; borders of North 
Platte Valley, between 
Alcova and Leo. and 
along north base of Shir- 
ley Mountains, scatter- 
ing trees ; Seminole Can- 
yon on the Platte, heavier 
growth ; Roc k Rive r, 
ridges ; Woods, Medicine 
Bow Range, some at 
lower edge of coniferous 
forest ; basal slopes of 
Sierra Madre south of 
Downington ; and can- 
yons near the mouth 
of Grand Encampment 
River. This pine was not found in western Wyoming, although doubtless there 
are scattering trees on the basal slopes of some ranges. 

The yellow pine yields valuable lumber mainly in the Black Hills, Bear 
Lodge and Bighorn Mountains, and on the Laramies in the region north of 
Laramie Peak. Elsewhere its growth is generally more or less scrubby. 
Pinus murrayana Balfour. Lodgepole Pine. 

The coniferous element of the Canadian forest belt in Wyoming is chiefly 
of lodgepole pine and Engelmann spruce. The pines, although occurring 
throughout the full width of the Canadian Zone, reach their greatest abundance 
and heaviest and purest stand in its lower half. The best forests are on the 
main ranges, varying in elevation from 8.500 to 10,000 feet on the Sierra 
Madre at the south; 8,000 to 9,500 feet on the Wind River Range; and 7,000 
to 9,000 feet in Yellowstone Park, the northern Absarokas, and on the eastern 
slopes of the Bighorns. 




BIIG60M 

Fia. 4. — Forest of Engelmann spruce (Picea engelmanni) 

on north slope of Ferris Mountains (9,000 feet). 



1917.] 



TREES AND SHRUBS. 



57 



The lodgepole pine forests of Wyoming are of gveal value, bul have been 
extensively levied upon for ranch fences, railroad ties, mine lagging, and 
lumber, and extensive tracts have been destroyed by fire. The deforested 
areas are now largely grown up with second-growth pine and aspen. 

Pinus edulis Engelmann. Pinyon ; Nut Pine. 

The pinyon, a small representative of the Upper Sonoran Zone, barely 
enters Wyoming in the lower valley of Green River. Scattering trees were 
found at 7,000 feet elevation on the north face of a juniper ridge 3 miles 
north of the Utah boundary and the same distance east of Green River. The 
pinyon may possibly occur elsewhere in this rough juniper-clad ridge and 




Fig. 5.— Frini 



of blue spruce (Picea parryana) on the Big Sandy, southwest base of 
Wind River Range (7,500 feet). 



mesa country, but it was not detected from Green River east to Red Creek, 
and thence north to Rock Springs. 

Picea engelmanni (Parry) Engelmann. Engelmann Spruce. (Fig. 4.) 

The Engelmann spruce is the principal conifer in the upper part of the 
Boreal forest belt on the high ranges of northwest Wyoming, and also on the 
Bighorn, Sierra Madre, and Medicine Bow Ranges. Next to the lodgepole 
pine it is the most abundant forest tree of Yellowstone Park. Although of 
regular occurrence at timberline in a dwarfed state, this spruce belongs to 
the Canadian Zone, attaining its maximum growth on cold, damp slopes and 
in bogs between 8,000 and 9,000 feet. It does not attain its best development 
in dry situations or on warm slopes, where it is found at a somewhat higher 
level. 

Picea parryana (Andree) Sargent. Blue Spruce. (Fig. 5.) 

The blue spruce occurs chiefly in western Wyoming north to Jackson Lake 

and the head of Wind River, but was also noted at Woods and in gulches south 



58 



NORTH AMERICAN FAUNA. 



[No. 42. 



of Sherman, and should be present elsewhere in the southern mountains. It 
inhabits the margins of cold streams in the lower border of the Canadian forest 
belt between 7.000 and 8.000 feet elevation (6.500 to 7.500 feet in Jackson Hole), 
forming usually a most attractive fringe of scattering symmetrical trees. 

Abies lasiocarpa (Hooker) Nuttall. Alpine Fir. 

The alpine fir has much the same distribution and vertical range as the Engel- 
mann spruce, although generally less abundant and growing in scattered thicket 
formation, rarely forming a heavy forest. With the spruce it extends regularly 
to timberline on the Wyoming ranges, where it is the more common of the two 
and forms dense prostrate mats on the bleak, wind-swept slopes. This fir is 

partial to cold gulches, 
stream banks, and damp 
spots generally. It may 
be seen to best advantage 
in typical growth as dark 
scattered clumps in aspen 
woods. While usually a 
small tree, it attains large 
size in gulches and on 
streams in the higher 
portions of Yellowstone 
Park, where it forms for- 
ests of considerable ex- 
tent above 8,000 feet 
altitude. It was observed 
on all the mountains ex- 
cept the Bear Lodge and 
Black Hills groups at the 
northeast. 

Abies (concclor?) Lind- 
ley. White Fir. 
The white fir (appar- 
ently Abies ooncolor) 
forms small forests be- 
low 9,000 feet elevation 
on the northern shoul- 
ders of the Uinta Moun- 
tains west of Lonetree, 
and is present also in 
fairly good stand with 
aspens on the summit of the plateau west of Maxon. Firs which were com- 
mon at 8,000 feet on the western slopes of the Salt River Range near 
Afton also had smooth, dark-gray bark and were unquestionably distinct 
from the light-barked A. lasiocarpa on the upper slopes of the same mountains 
above 9,000 feet. Unfortunately no specimens were preserved from the above 
localities. 

Pseudotsuga mucronata (Rafinesque) Sudworth. Douglas Spruce. (Fig. 6.) 
The Douglas spruce has a wide distribution in upper Transition and lower 
Canadian Zones from the Laramie and Bighorn Mountains westward. It was 
not observed on the Black Hills or Bear Lodge Mountains. This spruce grows 
chiefly at the lower edge of the lodgepole pine belt on the main ranges of 




Bl 1631 

p IG- c». — Forest of Douglas spruce {Pseudotsuga mucro 
naia), north slope of Ferris Mountains (8,500 feet). 



1017.1 TREKS AND SHRUBS. 59 

western Wyoming, teaching down t<> open sagebrush slopes on their basal 
Hanks, and on (he Bighorns and La ramies, extending down into the upper pari 
of the yellow pine belt. II varies locally as to abundance from a thin fringe 
of scattering trees and thickets on cool declivities and in gulches (which is 
usual), to a wide belt of heavy pure forest of lumber size. 

Juniperus scopulorum Sargent. Rocky Mountain Juniper. 

The most conspicuous and widely distributed juniper in the State is the 
Rocky Mountain juniper, occurring from the barren rocky ridges, canyons, 
gulches, and badlands bluffs on the Sonoran plains and deserts, up through 
the Transition Zone, where it mingles with yellow pines, Douglas spruces, and 
Rocky Mountain white pines on the lower mountain slopes. This juniper is of 
scattering growth along the bases of mountains and on the margins of the 
desert basins and valleys, and does not form a well-defined belt along the 
upper edge of the Upper Sonoran Zone as it often does in the southern Rocky 
Mountains. 

Juniperus knighti A. Nelson. Desert Juniper. 

The distribution of the desert juniper is imperfectly known, owing to very 
scattering held work in the difficult Red Desert region which it inhabits. The 
few localities from which there are specimens indicate a restricted range, 
mainly in the Upper Sonoran Zone. S. G. Jewett collected the species at 
Mountainview, on May 27, 1913, and it appears to be the dominant juniper 
at Rock Springs, near Carter, and in the badlands to the south and east of 
Lyman. < >n the Green River bluffs near the Utah boundary Juniperus mono- 
sperma is not uncommon, and on the higher borders of the Red Desert 
J. scopulorum is the common species. The desert juniper is usually of shrubby 
stature, branching from the base, and scarcely attains the dignity of a tree. 
Nelson found the species on the sandstone bluffs of the Bitter Creek drainage, 
and records specimens from Point of Rocks and Rock Springs. 1 

Juniperus monosperma Engelmann. One-Seeded Juniper. 

The one-seeded juniper is found at the northern base of Owl Creek Moun- 
tains west and southwest of Thermopolis ; at Hailey. southeast of Lander: 
along the Platte near Alcova in canyons and on dry slopes up to 0,800 feet 
elevation; and on the rough breaks along the lower Green River Valley, par- 
ticularly on the east side between Sage Creek and the Utah boundary. It is 
tolerably common on the Snake River bluffs near Baggs and is probably the 
species which forms a considerable belt along the western bases of the Sierra 
Madre. It was noted only in the Upper Sonoran Zone, extending up to 7,500 
feet on the hot slopes east of Green River near the State line. 

Juniperus sibirica Burgsdorff. Low Juniper. 

The low juniper, a graceful evergreen, is a characteristic undershrub in the 
Boreal forest belt throughout the Wyoming mountains. On most of the loftier 
ranges it extends to timberline, where it forms dense prostrate mats among 
lo-ks. Its center of abundance is in the Engelmann spruce and lodgepole pine 
forests of the Canadian Zone. 

Juniperus communis Linnaeus. Mountain Juniper. 

Not infrequent in mountain forests at lower elevations than the low juniper. 
The shrubby mountain juniper was noted as follows: Foothills west of Wheat- 
land, 5,500 feet; Springhill; near Sundance; Wolf, north base of Bighorn 
Mountains. 0,700 feet: head of Pat O'Hara Creek, northwest of Cody. 

1 Bull. 13, Div. of Agrost., T\ S. Dept. Agr., p. 54, 1898. 



00 



NORTH AMERICAN FAUNA. 



[No. 42. 



Juniperus sabina Liiuueus. Creeping Juniper; Trailing S:ivin. 

Mats of the creeping juniper, or trailing savin, are conspicuous <>n dry gravel 
ridges and exposed points and summits of high hills along the northern edge 
of Wyoming between the Bighorn and Bear Lodge Mountains. The species 
extends west at least to the eastern base of the Absaroka Range, but is of more 
general distribution in the northeast. It was not found in other sections of 
the State. The habitat of this species is in the Transition and lower Canadian 
Zones. 

Populus tremuloides Michaux. Aspen Poplar. (Fig. 7.) 

The Boreal aspen poplar is generally dispersed at suitable elevations, but 

does not, as a rule, reach large size in the Wyoming mountains. It commonly 

occurs in thickets 10 to 
20 feet high on damp, 
cold slopes, as under- 
growth in the coniferous 
forests, or as first growth 
on burned-over tracts. 
Low, scraggy thickets of 
aspen are on most of the 
cold slopes and draws 
above 7,500 feet altitude 
on barren elevations of 
the southwest, as the 
Aspen Mountains, Bear 
River Divide, and the 
high plateaus east of 
Green River near the 
southern boundary of the 
State. Beautiful aspen 
groves were noted, how- 
ever, between 8,000 and 
9.000 feet on the lower 
southern slopes of the 
Wind River Range east 
of Leckie, on the divide 
between Hoback and 
Green Rivers, and on the 
northern shoulders of the 
Uinta Mountains south- 
east of Hilliard. The 







Bl 1622 

Fig. 7. — Grove of aspen poplar (Populus tremuloides) 
near Springhill, north base of Laramie Peak (0,000 feet) 



aspen is perhaps the best characterizing tree of the Canadian Zone. 

Populus balsamifera Linnaeus. Balsam Poplar. 

The balsam poplar inhabits the borders of many of the larger streams in the 
northwestern mountains, being largely confined to the Canadian Zone between 
6,500 and 7,500 feet elevation. It occurs south in the Jackson Hole country at 
least to the Gros Ventre River. Scattering trees are found also in wet gulches 
at the southern base of the Bear Lodge Mountains near Sundance at about 
5,000 feet elevation. 
Populus occidentalis ( Rydberg > Britton. Broad-Leaved Cottonwood. 

The broad-leaved Cottonwood, characteristic of the Upper Sonoran Zone, 
forms the principal fringe on the streams of eastern and northern Wyoming, 



1917.] 



TBEES AND SHRUBS. 



01 



attaining perfection of growth in the lowest and warmest valleys. Large groves 
border the Bighorn, Belle Fourche, and especially the streams of the lower 
Platte drainage" There is a great deal of cottonwood growth on the Cheyenne 
River and other streams southwest of the Black Hills, hut in this section the 
species grows in a very stunted state, low and irregular and very thick at the 
the base. In the Wheatland district it occurs in places in heavy growth with 
Populus acuminata, and along the base of the mountains generally meets and 
commingles with the narrow-leaved Transition species, /'. angustifolia. 

Populus acuminata Rydberg. Lance-Leaved Cottonwood. 

On the Chugwater, Sibylee, and other tributary streams of the Laramie ami 
North Platte Rivers east of the mountains the lance-leaved cottonwood forms 
in many places a heavy 
fringe with Populus occi- 
dentalis, though by no 
means so generally dis- 
tributed as the latter spe- 
cies. A splendid growth 
of the lance-leaved cot- 
tonwood is on Sibylee 
Creek southwest of 
Wheatland. The species 
was not noted in north- 
ern Wyoming, but at the 
west scattering trees 
are on Green Rive r, 
just north of the Utah 
line. 

Populus angustifolia 

James. Narrow-Leaved 

Cottonwood. ( Fig. 8. ) 

The narrow-leaved cot- 
tonwood inhabits the bor- 
ders of mountain streams 
in the Transition Zone, 
mainly at elevations from 
G.(i(Mi to 7,500 feet at the 
west, extending down to 
5,000 feet in the central 
districts, and to 4,500 
feet at the eastern base 
of the Bighorn Moun- 
tains. It is apparently absent from the northeastern part of the State. Impor- 
tant streams bordered with good growth include the Bear, Green. Wind, and 
Greybull Rivers, the north and south branches of the Shoshone River above the 
forks, Snake River below the mouth of the Gros Ventre, and the upper North 
Platte down to 18 miles above Casper. Unusually tine groves are on Wolf and 
Big Goose Creeks, at the eastern base of the Bighorns. 




Fig. S. — Large narrow-lea ved cottonwood (Puiiiilits angus- 
tifolia) on sage flat in upper Wind River Valley. 



Salix amygdaloides Anderson. Peach-Leaved Willow. 

The large peach-leaved willow occurs in scattered clumps along streams in 
the Upper Sonornn Zone, chiefly at the east and north, as follows: Chugwater 
Creek and affluents; Little Bear Creek northeast of Meadow; Sibylee Creek 
west of Wheatland ; Rawhide Creek to base of Rawhide Butte ; near Lusk ; Hay 



62 



NORTH AMERICAN FAUNA. 



[No. l: 



Creek east of Aladdin; Wind Creek northeast, of Moorcroft : Sheridan : Arvada : 
Clear Creek, Clearmont to Buffalo; Bighorn River and tributaries near Ther- 
mopolis; and Green River near Utah boundary. 

Salix bebbiana Sargent. Bebb Willow. 

The Bebb willow is characteristic on foothill and lower mountain slopes up 
to about 9,000 feet altitude, and there are few if any ranges in Wyoming where 
it is not present. The usual growth is in scattered clumps about springs and 
bogs, but occasionally there is a heavier stand on the margins of mountain 
streams. The height attained seldom exceeds 12 or 15 feet, and 8 or 10 feet is 
usual. Specimens were taken on the Laramie. Bear Lodge, Bighorn, and Wy- 




FiGi 9. — Terraced copsca of gray-leaved willow {Salix glaucops) at tiuibcrline, Needle 

Mountain, Abwuroka Range. 



oming Ranges, and at Evanston, while it was observed at a wide range of 
localities. Nelson records it from Creston, on the open Red Desert. 1 

Salix pyrifolia obscura Anderson. Willow. 

Another willow, Salix pyrifolia obscura, was collected in the upper forests 
on the Wyoming Range west of Alerna, between 9.000 and 10.000 feet altitude, 
in the Canadian Zone. It is tolerably common at this locality. 

Salix nelscni Ball. Nelson Willow. 

Vernon Bailey collected the Nelson willow along the Iludsonian crest of the 
Salt River Range at 10,000 feet elevation, August 20. 1911. 

Salix glaucops Anderson. Gray-Leaved Willow. (Fig. 9; Rl. XI, fig. 2. i 

The gray-leaved willow is a low species of Alpine bogs, growing in dense 

copses about 2 feet high near timberline on the Bighorn. Wind River, and 



1 Bull. 13, Div. of Agrost., U. S. Dept. Agr., p. 59, 1898. 



1917.] TREES AND SHRUBS. 63 

Absaroka Ranges, and doubtless elsewhere. On Whiskey Mountain, south of 
Dubois, a few blossoms were out on May 14, 1910, but the leaves had not un- 
folded. The species was not in leaf by June 5 at timberline on the Bighorn 
above Hyattville. It was very abundant on Needle Mountain, Absaroka Range, 
in beautiful terraced copse formation, and likewise on the Wind River Moun- 
tains south of Fremont Peak. 

Salix petrophila Rydberg. Rock Willow. 

The rock willow was collected on the Alpine slopes above timberline at the 
head of Bull Creek, Wind River Range, in August, 1893, by Vernon Bailey. 

Salix tenera Anderson. Alpine Rock Willow. 

Low dense mats of the Alpine rock willow, which is very abundant on Whirl- 
wind Peak, in the Absaroka Range, cover portions of these slopes between 10,000 
and 11,000 feet elevation. 

Salix nivalis Hooker. Alpine Willow. 

The dwarf alpine willow is tolerably common on the bleak slopes above 
timberline on the Wind River Range, occurring among the rocks in dense 
creeping mats a few inches high. It is abundant south of Fremont Peak from 
11,500 feet upward. 

Salix saximontana Rydberg. Net-Veined Willow. 

More generally dispersed on the Wyoming ranges than the other Alpine wil- 
lows, but similar in habit of growth, Salix saximontana occasionally extends 
a little below timberline. It is especially abundant at the northern end of 
the Teton Range, where the creeping mats are very extensive, and push down 
the cool Hudsonian slopes to 9,500 feet altitude. 

Ostrya virginiana (Miller) Willdenow. Ironwood. 

Vernon Bailey reports the ironwood as abundant in Sand Creek canyon above 
Beulah, at the northern base of the Black Hills. The species <-losely approaches 
the Wyoming boundary in northwestern Nebraska, 8 or 10 miles east of Kirt- 
ley, Wyo., where it is not infrequent in wooded canyons along the northern 
escarpment of Pine Ridge. 

Corylus rostrata Alton. Beaked Hazelnut. 

The beaked hazelnut is abundant on the upper slopes of the Bear Lodge Moun- 
tains and Black Hills, forming dense undergrowth in. aspen and birch thickets 
between 5,500 and 6,000 feet altitude. It appears to be absent from the mairi 
ranges in Wyoming. 

Betula papyrifera Marshall. Canoe Birch. (Fig. 10.) 

The canoe birch apparently reaches its southern limits on the Bear Lodge 
Mountains and northern Black Hills. Over this region it grows to medium size, 
and with the aspen occurs in dense thickets on cool, shaded slopes and in damp 
spots as low as 5,000 feet altitude, and on Sundance Creek scattering trees of 
good size are found at 4,700 feet. On the dry upper slopes of the Bear Lodge 
Mountains this birch becomes very scrubby. It is apparently absent from the 
Bighorn Mountains and the ranges of western Wyoming. 

Betula fontinalis Sargent. Rocky Mountain Birch. (Fig. 11.) 

The Rocky Mountain, or black, birch borders often in dense growth most of 
the streams on the basal slopes of the mountains, and under the cooling influ- 
ence of the mountain water extends some distance out onto the plains. At the 
base of the Bear Lodge Mountains it is common at 4,000 feet altitude, but farther 



64 



NORTH AMERICAN FAUNA. 



[No. 42. 



west it is usually present between 6,000 and 7,500 feet. On warm exposed slopes 
of the Wind River Range north of Big Sandy it extends to 8,000 feet. 

Betula glandulosa Michaux. Dwarf Birch. 

The dwarf birch was noted only in the mountain valleys of the northwest, 
from Yellowstone Park (West Gallatin and Lewis River meadows) south to 
the Wyoming Range (South Piney canyon at 7,500 feet altitude), in the Canadian 
Zone. This little birch is especially abundant in the extensive willow bogs and 
swamps bordering Jackson Lake, where it occurs in dense thickets 3 or 4 feet 
high. Its leaves had mostly turned to a deep Indian red and a few were falling 
in Jackson Hole by September 13, 1910. It was noted also as follows: Head of 

Pacific Creek, near Two 
Ocean Pass; and Horse 
Creek meadows, at 
Merna, 8,000 to 9,000 
feet. 

Alnus tenuifolia Nut- 
tall. Alder. (Fig. 12.) 
The alder is found in 
a growth of varying 
density on the upper 
reaches of cold mountain 
streams, and with many 
other Canadian Zone 
species follows the cold 
conditions on their mar- 
gins down for some dis- 
tance into the Transition 
Zone. It is most abun- 
dant in the mountainous 
northwest, and was not 
noted in the Bear Lodge 
Mountains or on the 
northern groups of the 
Laramie Range. 




Ulmus 
nseus. 
The 



Fig. 



10. — Thicket of canoe birch (Betula papyrifera) in 
gulch near Sundance, Bear Lodge Mountains. 



americana Lin- 
Elm, 
elm penetrates 



Wyoming for a short 
distance in the low val- 
leys at the northeast. It was common and growing to large size on Sand Creek 
to 10 miles above Beulah at the northern base of the Black Hills, and on Hay 
Creek west to Aladdin and Eothen, while it is reported at Hulett in the Belle 
Fourche Valley. A'ernon Bailey found it on Little Powder River near Morse. 

Quercus macrocarpa Michaux. Bur Oak. 

The bur oak extends into Wyoming from the northeast and is found in 
abundance over a small area in Crook County, principally east of the Belle 
Fourche River and north of Linden and Inyankara. It occurs in scattered 
groves on the partially open basal flanks of the Bear Lodge Mountains and at 
the northern base of the Black Hills, extending to the dry Bear Lodge summits 
at 0,000 feet elevation, where Vernon Bailey found scrubby thickets 4 or 5 feet 



1917.] 



TREES AND SHRUBS. 



65 



high, loaded with acorns, in August, 1913. In some of the stream valleys the 
bur oak grows to large lumber size, notably on Sand Creek above Beulah, and 
many trees were there noted with clean straight trunks of good height from 3 
to 4 feet in diameter at the base. The vertical range of this oak is about the 
same as that of the yellow pine, with which it usually commingles in this 
region. Vernon Bailey noted a little oak growth near the head of the Little 
Missouri River, apparently its western limit in the State. 

Atriplex canescens James. Saltbush ; Gray Shadscale. 

The various saltbushes are characteristic Sonoran species of the arid Great 
Basin region, and barely enter the Great Plains area on some of the dry valley 

























:M 


n Jfi * 


■ 










■jii. 




















■>£ ■ I ^fl^iPS 


















Uti 




■ 
































' 




: • ■ 














-^. 





Fig. 11. 



-Clumps of Rocky Mountain birch (Bctula fontinalis) 15 feet high, at north 
base of Shirley Mountains. 



flats at the eastern base of the foothills. They are mostly alkali-resistant and, 
with the possible exception of Atriplex canescens, furnish valuable winter forage 
for sheep in the central desert sections. Of the three principal shrubby species 
found in Wyoming, .4. canescens is the least abundant. It is apparently absent 
from the Red Desert proper where A. confertifoMa and .4. nuttalli abound, but 
extends farther to the east than either of the foregoing. Atriplex canescens oc- 
curs in very dense growth 3 feet or more high on dry flats along the Chugwater 
at Bordeaux, and also in the sand along the Laramie at Uva, but is in scatter- 
ing growth elsewhere. 

Atriplex confertifolia S. Watson. Round-Leaved Saltbush. 

The round-leaved saltbush is a low stocky shrub 1 or 2 feet high, very 
abundant on sandy and alkaline soils up to 7,500 feet elevation on warm slopes 
east of Green River near the Utah line ; bad lands south of Lyman, to 7,000 feet ; 
74440°— 17 5 



66 



NORTH AMERICAN FAUNA. 



[No. 42. 



Bear River below Evanston, to 6.600 feet ; Fossil ; Bigpiney ; Dubois ; Trout 
Creek, north Shoshone Valley ; Bighorn slopes above Hyattville, to 5,500 feet ; 
north base of Rattlesnake Mountains, to 6,500 feet ; Splitrock, Sweetwater Val- 
ley ; east to Fort Steele, Shirley, Old Fort Fetterman, and Arvada. 

Atriplex nuttalli S. Watson. Nuttall Salthush. (PI. IV, fig. 2.) 

The Nuttall saltbush, a low-spreading species, commonly known as "salt 
sage " to the sheepmen, is of great economic value, as it affords the chief winter 
food to the flocks on the Red Desert. It has about the same range as Atriplex 
confertifolia, but is seldom found in sand, being partial to dry adobe and 
saline flats, where it forms often the dominant plant growth. Extensive flats 

at Frannie and else- 
where in the Bighorn 
Basin are covered with 
a pure and uniform 
growth of this saltbush. 
The observed eastern 
limits are Walcott ; Lit- 
tle Medicine Bow River 
west of Marshall ; Indian 
Creek at State line north 
of Kirtley ; Newcastle ; 
and Colony. 

Grayia spinosa Moquin. 
Grayia. 

A characteristic spiny 
shrub of the Sonoran 
desert tracts from the 
Sweetwater Valley west- 
ward, especially abun- 
dant in sandy or adobe 
soils at the lowest levels. 
On the sandy hummocks 
between Frannie and 
Garland at the end of 
June dense clusters of 
flat, winged seeds were 
borne in great profusion 
on the grayia bushes. 
These varied from green- 
ish to pink or purple- 
brown and lent a peculiar hue to the landscape. This shrub has considerable 
forage value, as its thick leaves and seeds, gathered into drift piles under the 
bushes, are eagerly eaten by sheep in winter. 

Grayia spinosa was common from Independence to Splitrock; Dry Lake to 
Lorey ; on first and second benches above the Platte at Fort Steele ; Worland ; 
Manderson ; Bonanza ; Greybull west nearly to Cody ; Rock Springs region ; 
Green River near Utah line; Carter to Lyman; bad lands south of Lyman; 
Cumberland ; Fontenelle to Opal ; and Green River flats north nearly to 
Labarge. 

Sarcobatus vermiculatus Torrey. (Jreasewood. (PI. IV, ivj;. 1.) 

This common desert shrub is of wide dispersion in the more arid por- 
tions of the Upper Sonoran Zone and was noted at numerous localities. It 




Fig. 12. 



BIISC9 

-Fringe of alder (Alnns tenuifolia) on Pacific 
Creek below Two Ocean Pass. 



1017.] TREES AND SHRUBS. 07 

forms a very rank growth on the dry alkaline lake basins and flats of central 
Wyoming, especially in the Red Desert region, where its dark-green foliage 
and large size make it the most conspicuous of the alkali-resistant shrubs. 
The greasewood follows dry adobe valley flats to about 7,000 feet elevation. 

Berberis aquifolium Pursh. Barberry; Oregon Grape. 

The Oregon grape, a low undershrub, is characteristic of dry forested or 
partly wooded slopes throughout the State, mainly in the Transition Zone. It 
is commonly associated with the bearberry, with which it penetrates to the 
lower edge of the Canadian forest belt. On the Ferris Mountains it was noted at 
8,700 feet altitude, and on the west side of the Salt River Range was common 
to 9,000 feet. 

Ribes longiflorum Nuttall. Flowering Currant. 

In early summer the brush fringe along streams at the lower elevations is 
enlivened by the yellow bloom of the flowering currant. This shrub is a char- 
acteristic Sonoran species, but is perhaps most conspicuous at the base of the 
mountains, where it penetrates the foothills for some distance in warm stream 
valleys. Its upper limits were observed as follows: Foothills west of Wheat- 
land, to 5,300 feet altitude; eastern base of Bighorn Mountains; Greybull River 
at Meeteetse; Pat O'Hara Creek to 6,000 feet; south slope of Owl Creek Moun- 
tains to 7,000 feet; Bull Creek, Wind River Valley; streams of Salt River 
Valley; Evanston and west slope of Bear River Divide to 7,500 feet; Mountain- 
view ; and Henry's Fork of Green River to Burntfork P. O. 

Ribes inebrians Lindley. Red Currant. 

The red currant is widely distributed in the Transition Zone over most of 
Wyoming, extending regularly to the lower edge of the Canadian Zone. Flower- 
ing specimens were taken in the Laramie foothills west of Islay on June 16, 
1909, and near Merna, Wyoming Range, at 8,000 feet altitude, as late as Au- 
gust 10, 1911. This species is partial to rocky situations. 

Ribes montigenum McClatchie. Bristly Red Currant. 

A high-altitude species, the bristly red currant has a general distribution on 
the higher Wyoming ranges. It is most abundant near timberline, where it 
occurs either as scattering bushes in slide rock, or in dense patches a foot 
or two high on deep-soiled slopes, as on the Wyoming Range west of Merna. It 
was in flower on the summit of Bridget' Peak in the Sierra Madre, on July 
7, 1911, and still so on the high Wind River Range south of Fremont Peak on 
July 18. The red, edible fruit was abundant and fully ripe in the timberline 
region on the Teton Range on August 30, 1910. It was found in abundance on 
the high ridge extending north from Needle Mountain, in the Absaroka Range, 
between 10,000 and 11,000 feet elevation ; and also on the Salt River Range, 
above 9,000 feet. 

Ribes lacustre (Persoon) Poiret. 

This species occurs in Canadian Zone forests in northwestern Wyoming. It 
was still in flower near the upper end of Fremont Lake on July 15, 1911, but 
was bearing fruit on Grinned Creek. Absaroka Range (8,000 feet altitude), 
July 30, 1910. Specimens were collected al both localities. 

Ribes petiolare Douglas. Mountain Black Currant. 

The mountain black currant, a wide-ranging Boreal species, was collected 
on the summit of the Bear Lodge Mountains on June 20, 1912, where it was not 
uncommon on the dry crests at 6,000 feet elevation. Flowering specimens were 



68 NORTH AMERICAN FAUNA. [No. 42. 

also taken in the Canadian Zone forest near the upper end of Fremont Lake, 
Wind River Range, July 15, 1911, and fruiting specimens at Tower Falls, Yel- 
lowstone Park, August 11, 1910. It is especially abundant in the Yellowstone 
Park forests. 

Ribes viscosissimum Pursh. Currant. 

This currant is characteristic of the Canadian Zone forest belt in northern 
Wyoming. Flowering specimens were taken at 8,500 feet elevation on the 
head of Shell Creek, Bighorn Mountains, June 5, 1910; near the upper end of 
Fremont Lake, Wind River Range, July 15, 1911 ; and at 8,000 feet on Grinned 
Creek, Absaroka Range, as late as July 30, 1910. Vernon Bailey collected the 
species on Wolf Creek, northern slope of the Bighorns, August 10, 1913. 

Edwinia americana (Torrey and Gray) Heller. Edwinia. 

The low flowering edwinia was found only on the crest of the Laramie 
Range east of Laramie, between 8,500 and 9,000 feet elevation. Its handsome 
white cyinous bloom enlivened the rock ledges and cliffs on June 18, 1909. 

Cercocarpus ledifolius Nuttall. Mountain Mahogany. 

The mountain mahogany is a small evergreen tree or stout shrub peculiar 
to rocky plateaus and ridges and warm exposed basal slopes of mountains. 
In Wyoming a scattering distribution is indicated from the lower Green River 
Valley north to Jackson Hole, in the Transition Zone. Most of the rocky 
ridges adjacent to Green River near the Utah boundary are clothed up to 
7,500 feet elevation with dense scrubby thickets from 3 to 6 feet in height. 
The species was not found farther north in the Green River country, btit 
enters the State from the west along Snake River, and covers in good growth 
the more exposed of the lower western slopes of the Salt River Range east 
of Afton and Smoot up to 7,500 feet. Its upper limits in the Snake River 
drainage are reached apparently near Jackson, where Edward A. Preble found 
a considerable growth on a warm slope at 7,000 feet in a tributary gulch of 
Cache Creek. 

Cercocarpus intricatus Watson. Mountain Mahogany. (PI. XV, figs. 1 and 2.) 
An abundant evergreen shrub on warm open slopes of the Bighorn Mountains. 
(in the eastern side scattering bushes dot t lie bare Tongue River bluffs at 
Rancbester. The species forms dense thickets 3 feet high at about 5,000 feel 
altitude near Eaton's Ranch, and thence it ascends dry, rocky ridges to 6,500 
feet on bare exposed points in the mountains south of Wolf. On the warm 
western side of the Bighorns above Hyattville Cercocarpus intricatus was not 
seen below 5.800 feet, but between 6,000 and 6,500 feet the dry, hot, reddish 
slopes were dotted with this intricately branched, steely gray shrub. 

Cercocarpus parvifolius Nuttall. Mountain Mahogany. (PI. II, fig. 1.1 

This mountain mahogany occurs mainly in the Transition Zone in Wyoming, 
although in the southern Rocky Mountains its center of abundance is in the 
juniper and pinyon belt of the Upper Sonoran Zone. It is partial to warm, 
cocky situations, either partly open foothill slopes or outlying ridges and 
buttes on adjacent plains or deserts. The largest growth of mountain mahogany 
observed was fully 9 feet high, on the pine-clad foothills southwest of Wheat- 
land, at 5,300 feet altitude. The usual height attained is from 4 to feet. 
Unusually extensive thickets grow on open ridges paralleling I he Horse Creek 
valley between l>a\is Ranch and Meadow. This shrub flowers early in 
.lime, but occasionally a little later. A flowering specimen from Steamboat 
Mountain was collected June 26, 1913. The distribution of Cercocarpus parvi- 



North American Fauna No. 42, U. S. Dept. Agr. Biological Survey. 



Plate XV. 






■*».«!«:-. 



♦9i 



^.* i * : .vl^.'f >\^/, ^i^**^' 




Fig. 1.— Mountain Mahogany (Cercocarpus intricatus). 
• West slope of Bighorn Mountains above Hyattville (6,000 feet). 




Fig. 2.— Nearer View of Same. 



1017.] TREES AXD SHRUBS. 69 

f alius as noted by Survey parties is entirely south of ;i line from Newcastle 
in the northeast to Evanston in the southwest. 

Kunzia tridentata (Pursh) Sprengel. Antelope Brush. 

The antelope brush is a common shrub on dry open Transition slopes in 
southern and western Wyoming, being especially abundant on arid hills, pla- 
teaus, and occasional sand ridges in the borders of the Red Desert at eleva- 
tions between 7,000 and 8,500 feet. It was not noted north of the Wind River, 
Rattlesnake, and Casper Ranges, but it is not uncommon on dry slopes and 
sagebrush flats south of Buffalo Creek and the Gros Ventre River in Jackson 
Hole. With the sagebrush {Artemisia tridentata) this shrub regularly ascends 
warm slopes and occurs in dry parks and openings to the lower edge of the 
Canadian Zone. Its vertical range varies from 4,500 feet on ridges along 
the North Platte near Glendo to 9,000 feet on the summits of the Laramie 
and Ferris Ranges, and also on the warm southern slopes of the Wind Rivers 
north of Big Sandy. 

Holodiscus dumosus (Nuttall) Heller. 

Common locally on rocky slopes in some of the dry desert mountains and 
in the rougher borders of the Red Desert, mainly in the Transition Zone. 

Opulaster pubescens Rydberg. Ninebark. 

On dry, steep, basal mountain slopes at the head of Pat O'Hara Creek, north- 
west of Cody, the ninebark forms a low but exceedingly dense chaparral 
among partly dead Douglas spruce and white pine forests up to 6,500 feel 
elevation. Along the northern base of the Bighorn Mountains near Wolf it 
was likewise partial to dry slopes between 5.000 and 7,000 feet. 

Opulaster monogynus (Torrey) Kuntze. Western Ninebark. 

The western ninebark, a Transition Zone shrub, is tolerably common on the 
Laramie Mountains and on outlying ridges of the plains region to the eastward, 
at elevations between 5,000 and 7,000 feet. 

Opulaster malvaceus (Greene) Kuntze. Ninebark. 

This ninebark is very abundant and conspicuous on the warm western 
slopes of the Salt River Range near Afton, but was not found elsewhere. Its 
vertical range in these mountains is from their bases up to 7,100 feet in 
canyons and to 7.500 feet on warm slopes, and like the other Wyoming species 
of the genus it is mainly restricted to the Transition Zone. 

Spiraea lucida Douglas. Meadowsweet. 

The handsome flowering meadowsweet is sparingly present across Wyoming 
at the north in the lower part of the Canadian forest belt. Alexander Wetmore 
collected it at 7.000 feet elevation on the east side of Teton Pass on September 
9. 1910, and I have observed it down to 5,800 feet in the Bighorn Mountains 
near Wolf, and in abundance on the Bear Lodge Mountains above 5,500 feet. 

Vernon Bailey reports a pink-flowered species, probably Spiraea densiflora, 
between 7,000 and 8,000 feet on the eastern slope of the Bighorns, above Wolf. 

Dryas octopetala Linnaeus. Alpine Avens. 

The low alpine avens is restricted to Arctic and Alpine regions. In Wyoming 
I collected it on the Bighorn Mountains, where it was in characteristic pros- 
trate matted growth on the rocky slopes above timberline at the head of 
Trapper Creek, being not yet in flower on June 10, 1910. Vernon Bailey 
collected it in August, 1893, on the high Wind River Range at the head of 
Bull Creek. 



70 



NORTH AMERICAN FAUNA. 



[No. 42. 



Rubias deliciosus James. False Raspberry. 

The handsome false raspberry was abundant and still flowering on the pine- 
clad foothill ridges southwesl of Wheatland on July 4. 1909, at elevations be- 
tween 5,000 and 5,500 feet. This is probably near its northern limit, as it was 
not noted in the Laramie Peak region nor on any of the ranges farther west. 

Rubus parviflorus Nuttall. Flowering Raspberry ; Thimbleberry. ( Fig. 13. ) 

The flowering raspberry is a tolerably common fruiting species in Cana- 
dian Zone woods on most of the Wyoming ranges, especially at the north. 
The large leaves, showy white flowers in June and early July, and in most years 
an abundance of handsome red berries in late August, make the thimbleberry 
very conspicuous in the woodlands between 7,000 and 0,000 feet elevation. The 




BII64S 

Fn;. 13. — Flowering raspberry (Rubua parviflorus) In cool gulch, north base of Casper 

Mountains ((1.500 feet). 

large berries, which are pleasantly flavored, though somewhat dry and seedy, 
are eaten by many native birds and mammals. 

Rubus strigosus Michaux. Red Raspberry. 

The red raspberry is abundant at numerous localities in the mountain dis- 
tricts from the lower edge of the forest belt to timberline. Its observed vertical 
range is from 4,500 feet at Wolf, at the north base of the Bighorn Mountains, 
to 11,700 feet, above timberline, south of Fremont Peak, Wind River Range. 
It reaches its best growth and bears most abundantly on rocky, partially for- 
ested slopes in upper Transition and lower Canadian Zones on the northwestern 
ranges. In the heavy forests between 7,000 and 8,000 feet elevation on Grin- 
ned Creek, in the Absaroka Range, the fruit was just ripening on August 1, 
1910, while on the western slope of the Salt River Range the bushes were full 



1917.] TREKS AND SHRUBS. 71 

of berries on August 20, 1911. in the timberline region the species occurs in 
a dwarfed state, usually uol over a foot nigh in fields of slide rock. Rock- 
strewn paths of avalanches throughout the mountains support more or less 
raspberry growth. 

Dasiphora fruticosa (Linnaeus) Rydberg. Shrubby Cinquefoil. 

The low shrubby cinquefoil inhabits cold mountain bogs and meadows and 
is more common on the ranges aleag the western edge of the State than farther 
oast. It is especially abundant in the cold stream meadows along the northern 
base of the Uinta Mountains, in the upper Green River Basin, and at the head 
of Wind River; in the willow swamps of Jackson Hole and Yellowstone Park; 
and in the timberline region on the Wind River and Absaroka Ranges. It 
extends above timberline on deep-soiled slopes, but at the higher altitudes is 
depauperate, from a few inches to a foot in height. 

Amelanchier alnifolia Nuttall. Serviceberry. 

Several species of serviceberry are characteristic of the Transition Zone of 
Wyoming, but Amelanchier alnifolia appears to lie of widest range. On the less 
arid foothills and lower mountain slopes at the east and north, where it is most 
abundant, it is usually the only species present, but in the central and southern 
districts it occurs in places with .1. oreophila, .1. elliptica, and possibly others. 1 
Typical specimens are from the Bear Lodge Mountains. It was noted at locali- 
ties too numerous for inclusion. 

Amelanchier elliptica A. Nelsou. Serviceberry. 

Taken only in the southwest, where it is the predominating species of Ame- 
lanehier in the western Transition borders of the Red I >esert region at elevations 
up to 8,000 feet. It was observed at a number of points on the Bear River Divide 
from Hilliard north to Cokeville. and also in (he badlands south of Lyman. 
S. G. Jewett collected flowering specimens at Spring Valley on June 9, 1912, and 
also on Steamboat Mountain on June 2.1. 

Crataegus cerronis A. Nelson. Hawthorn. 

Several species of hawthorn are represented in Wyoming, where they form a 
characteristic fringe on streams in the Transition Zone. Crataegus cerronis is 
apparently the predominating form in the Bighorn Mountains, and possibly 
east to the Black Hills. It was blooming profusely on .lime 0. 1912, in the 
heavy deciduous fringe along Wolf Creek, at the northern base of the Bighorns, 
up to 4,800 feet. 2 Flowering examples were collected at this point. 

Crataegus rivularis Nuttall. Black Hawthorn. 

The black hawthorn is common in southwestern Wyoming north to the Salt 
River Valley and Green River Basin and east at least to the upper Platte, at 
elevations from 6.000 to 8,000 feet. 

Sorbus scopulina Greene. Mountain Ash. 

In the Wyoming ranges the mountain ash is usually ft low or medium-sized 
shrub 3 or 4 feet high, growing in scattered clumps or occasionally forming 
small thickets, although in the Shirley Mountains a few clumps feet or more 
in height were noted. It is very generally distributed in the forests of the 
Canadian Zone up to 8.500 or 9,000 feet altitude. 

1 Data collated under Amelanchier alnifolia may include some related to other species, 
being based mainly on field identifications. Unfortunately, tew specimens of Amelan- 
chier have been preserved. 

2 Vernon Bailey found the hawthorn up to 0,000 feet near Wolf. 



72 NORTH AMERICAN FAUNA. 



12. 



Primus americana Marshall. Wild Red Plum. 

Scattered scrubby thickets of wild red plum are met with in gulches and on 
streams of tbe northeastern counties e;ist of tbe Bighorn Mountains, the species 
being perhaps most abundant on the basal slopes of the Bear Lodge Mountains 
and at the northern base of the Black Hills. 

Prunus pennsylvanica Linmeus f. Wild Red Cherry. 

The wild red cherry is sparingly present in northern Wyoming from the up- 
per slopes of the Black Hills and Bear Lodge Mountains west nearly to Yellow- 
stone Park. On the dry, partially open summit of the Bear Lodge Mountains 
it grows in a dwarfed state. Shrubs not over 2 feet high were flowering pro- 
fusely at 6,000 feet altitude on June 20, 1912, and considerable fruit was found 
by Vernon Bailey near Welcome late in August, 1913. Fruit was reddening at 
the end of July, 1910, on the North FofK of the Shoshone River. 

Prunus melanocarpa (A. Nelson) Rydberg, Chokecherry. 

The chokecherry is the predominating species of Prunus, being omnipresent 
in the foothills and mountains below 8,000 feet elevation, and on the Laramie 
Range reaching 9.000 feet. It is most abundant on dry, open, or partly forested 
Transition Zone slopes, where, however, it occurs usually in low scraggy 
thickets. It attains its perfection in damp gulches and along streams. The 
fragrant white bloom is conspicuous in May and early in June. 

Amorpha canescens Pursh. False Indigo. 

The low silvery false indigo, a shrub of the open Great Plains area, barely 
enters the State at the northeast in the Belle Fourche Valley. In June, 191:2. 
it was common near Aladdin on open grassy slopes bordering Hay Creek 
valley. 

Rhus rydbergi Small. Western Poison Ivy. 

The western poison ivy was occasionally noted at the eastern base of the 
mountains: base to summit of Rawhide Butte, near Lusk; and Wolf Creek, 
Bighorn Mountains, below Eaton's Ranch. 

Schmaltzia glabra (Linmeus) Small. Smooth Sumac. 

The smooth sumac was noted by Vernon Bailey on the lower eastern slopes 
of the Bighorn Mountains near Wolf. August 10, 1913. It was sparingly present 
between 4,000 and 6.000 feet elevation in this locality, but has not been noted 
elsewhere in Wyoming by Survey tield parties. 

Schmaltzia trilobata (Nuttall) Small. Skunk Bush. 

The skunk bush, characteristic of the Upper Sonoran Zone, fringes water- 
courses at the lower elevations up to 5,000 or (i.OOO feet, and on especially warm 
slopes has been noted at a little over 7,000 feet. The heaviest growth is along 
streams, as at Greybull. where there are dense thickets S or 10 feet high in 
the cottonwood growth along the Bighorn. A scrubby growth clothes the hot 
slopes in the bad lands between Greybull and Ionia to a considerable elevation. 
On Horse Creek north of Cheyenne, and also near Wheatland and Cassa, it 
grows on valley flats with sagebrush. In the warm districts generally it oc- 
curs in scattering growth in dry gulches and on exposed faces of bluffs. 

Pachystima myrsinites (Pursh) Rafinesque. 

A low evergreen shrub of the Canadian forest belt in Wyoming, chiefly in the 
western and southwestern mountains. Its vertical limits are approximately 
6,000 and 10,000 feet, but it is most abundant between 7,000 and 9,000 feet. 
« >n the Salt River Range near timberline it was growing abundantly in a 



1917.] TREES AND SITRUBS. 73 

dwarfed state a few inches high. The small green flowers appear quite early, 
the flower buds being evidenl on a specimen collected al Jackson, May 4, L911, 
by Edward A. Preble. 

Acer negundo Linnaeus. Box Elder. 

The box elder was noted chiefly along streams and in wet draws of the 
Great Plains area, but also at Bonanza in the Bighorn Basin, and between 
Dixon and Baggs along the southern boundary of the State. 

Acer glabrum Torrey. Mountain Maple. 

The handsome mountain maple inhabits cool slopes, gulches, and damp spots 
generally from the base of the mountains to 8,000 or 9, (too feet elevation, being 
most abundant in southeastern Wyoming. It was not noted in the Black Hills 
region, although found sparingly in cool north gulches at Squaw Butte, on the 
Wyoming-Nebraska line east of Kirtley. Its usual height is 5 or 6 feet in the 
mountains, and it rarely exceeds 8 feet even along streams. 

Acer grandidentatum Nuttall. Large-Toothed Maple. 

Scattered clumps of the large-toothed maple, 10 or 12 feet in height, are on 
the warm lower western slopes of the Salt River Range near Afton. The verti- 
cal range in these mountains is from their 6,300-foot bases up to 7.300 feet, in 
the Transition Zone, and it does not here attain its maximum growth. On the 
Salt River Range it is associated with the mountain maple, but does not occur as 
high as the latter. 

Rharnnus alnifolia L'Heritier. Buckthorn. 

The buckthorn reaches a short distance into Wyoming at the west, in the 
region contiguous to Snake River, but was not detected elsewhere. It is a 
conspicuous shrub on stream margins and in wet willow bottoms in the borders 
of Jackson Hole, and was abundant on Pacific Creek, 15 miles northeast of 
Moran. It was bearing its large ripe black berries in Webb Canyon in the 
Teton Mountains, at 6.700 feet elevation, September 1, 1910, and was collected 
by Alexander Wetmore on Trail Creek, near Teton Pass, late in September. 

Ceanothus velutinus Douglas. Mountain Balm. (Fig. 11.) 

Throughout the mountains one of the most characteristic shrubs is the moun- 
tain balm, in many places forming a dense chaparral 2 or 3 feet high on dry. 
open or partly forested slopes mainly in the Transition Zone. It was found in 
greatest abundance on dry summits of medium elevations, as the Bear Lodge. 
Casper. Shirley, Ferris, and Rattlesnake Mountains, on the 8,000-foot divide 
between the head of Salt River and Smiths Fork, and on Little Mountain, the 
elevated plateau between Maxon and the Green River Valley. The vertical 
limits are from 5,000 feet near AVolf, at the eastern base of the Bighorn Moun- 
tains, to 9,500 feet on Bridger Peak in the Sierra Madre. The white, sweet- 
scented flowers were conspicuous in Sierra Madre forests west of Grand En- 
campment between 8,500 and 9,500 feet, July 7. 1911, and the species was still in 
partial flower on the Wind River Range north of Rig Sandy July 26, although 
past flowering near Fremont Lake on the same range by July 20. 

Ceanothus fendleri Gray. Wild Tea Bush. 

The low, much-branched wild tea bush is common on dry, warm slopes near 
Springhill, north of Laramie Peak. It was found in open yellow-pine forest up 
to 7,400 feet elevation, but was not noted elsewhere within the State. 

Ceanothus mollissimus Torrey. Wild Tea Bush. 

Vernon Bailey found this wild tea bush on Big Goose Creek, at the eastern 
base of the Bighorn Mountains. It is apparently rare in Wyoming. 



74 



NORTH AMERICAN FAUNA. 



[No. 42. 



Elseagnus argentea Pursh. Silverberry. 

The silverberry occurs across Wyoming ;it the west, where it forms a scatter- 
ing silvery fringe on stream banks mainly in the Transition Zone. It is espe- 
cially abundant on streams of the upper Green River Basin and on tributaries 
of Wind and Snake Rivers. Elsewhere it was observed at Maxon, Lone- 
tree, Cokeville, Meeteetse, on the South Fork of the Shoshone River, Pat O'Hara 
Creek, and on Gardiner River, near Mammoth Hot Springs. 

Lepargyrea argentea (Nuttall) Greene. Buffaloberry. 

While abundant locally on many streams at the base of the mountains, espe- 
cially at the north, the buffaloberry is by no means generally distributed over 
the State, and was not detected in the southwest. The localities where it was 




Mountain balm iCeanothux velutinus) in flower 
(9,000 feet). 



B I 9636 

Siena Madre, July 7, 1911 



found are mainly Upper Sonoran : Sibylee Creek, Casper, Belle Fourche River, 
Little Missouri River, Powder River, Clear Creek, streams near Sheridan and 
Ranchester, up Wolf Creek to 4,500 feet elevation, Alcova, Casper Creek, north 
slope Rattlesnake Mountains, Snake River at Baggs, Wind River Basin (mainly 
below 6,000 feet), and streams of Bighorn Basin. 

Lepargyrea canadensis (Linnaeus) Green. Canadian Buffaloberry. (Fig. 15.) 
The Canadian buffaloberry is a characteristic and widely distributed under- 
shrub in the forests of the Canadian Zone between 8,000 and 10,000 feet eleva- 
tion. Although usually found on mountain slopes, it is abundant in balsam 
poplar growth along Snake River and other streams of Jackson Hole. 

Phyllodoce empetriformis (Smith) Don. Mountain Heath. 

The mountain heath, a characteristic evergreen of the Hudsonian Zone, was 
found locally abundant at widely separated localities in the high ranges of 



1917.] 



TREES AXD SHRUBS. 



75 



northwestern Wyoming. Near Fremont Peak it was abundant in bogs and 
rocky situations from 10,500 feet altitude to a little above timberline, and 
occasional at 10,000 feet, on July 20 the dainty rose-colored flowers were 
still open on the Wind River Mountains, bid on August 1 it was nearly through 
flowering on Whirlwind Peak, in tbe Absaroka Range. At timberline on the 
Tetons no flowers remained on the branches by August 30. 

Menziesia ferruginea Smith. Rustyleaf. 

This handsome and conspicuous shrub was observed only in the dense 
coniferous forests of Yellowstone Park, where it is common near Sylvan Pass, 




1'ic. 15. — Canadian buffaloberry {Lepargyrea canadensis) 
Range near Merna C8.000 feet). 



B 1 3685 

aspen woods, Wyoming 



Loin' Star Geyser, below Norris Basin, Thumb, Lewis ttiver, and Snake River 
near Soldier Station. 

Kalmia polifolia Wangenheim. Laurel. 

The wet bogs and mossy margins of the numerous snow-fed lakes on the 
Wind River Range south of Fremont Peak were bright with pretty pink- 
purple flowering mats of laurel in mid-July, 1911. The species was in flower 
on July 18 in the Hudsonian Zone from 10,000 to 11,000 feet elevation, but was 
not detected above timberline. With Phyllodoce em pet ri for mis it often formed 
dense mats, from a few inches to nearly a foot in height. Vernon Bailey col- 
lected the laurel in the timberline region on the Wind River Mountains above 
Bull Lake in 1893, and at the head of Raven Creek, in Yellowstone Park, 
in 1915. 



76 NORTH AMERICAN FAUNA. [No. 42. 

Arctostaphylos uva-ursi (Linnaeus) Sprengel. Red Bearberxy. 

The low trailing red bearberry, with its leathery evergreen leaves, dainty 
rose-hued blossoms, and red berries, forms beautiful mats in the open yellow- 
pine forests of the Black Hills, Laramie Peak region, and on the eastern 
slopes of the Bighorn Mountains. While most abundant in the Transition Zone. 
it occurs occasionally in the coniferous forests higher up, as on the upper 
slopes of Laramie Peak and on the Wind River Range south of Dubois. It is 
very abundant in the woodlands of Jackson Hole, and also on the Bear Lodge 
Mountains ; at from 5,000 to 8,000 feet elevation on the eastern slope of the 
Bighorn Mountains near Wolf ; and from 8,000 to 9,500 feet on their western 
slope above Hyattville; on the Rattlesnake, Shirley, and Green Mountains; 
south of Fremont Peak and on the head of the Big Sandy, Wind River Range : 
Needle Mountain and Grinned Creek, £bsaroka Range; Merna; Evanston ; and 
north slopes of Uinta Mountains west of Lonetree. 

Vaccinium oreophilum Rydberg. Mountain Blueberry. 

The mountain blueberry reaches perfection of growth in the cool, damp 
forests of the Canadian Zone at about 8,000 feet elevation. In many localities 
it occurs with Vaccinium erythrococcum, but it is less abundant and usually 
not found so high up. It was observed oh the Bear Lodge, Shirley, Casper, 
Ferris, and Bighorn Mountains ; Grinned Creek, Absaroka Range ; forests of 
Yellowstone Park; Teton Pass and Moose Creek, Teton Range; wooded hills 
bordering Jackson Lake; head of Pacific Creek; Snake River west of Jackson; 
Salt River Range near Afton ; and on the Wyoming Range near Merna. 

Vaccinium erythrococcum Rydberg. Red Bilberry. (PI. IX, fig. 2.) 

The low, small-leaved red bilberry was noted on all the higher elevations 
of Wyoming. It is most abundant in the upper Canadian Zone, where it 
densely carpets the coniferous forest floor, and it is the only Vaccinium present 
in the Hudsonian belt of dwarfed forest higher up. In early August the small 
juicy red berries are greatly relished by dusky grouse and other fovest birds. 

Vaccinium occidentale Gray. Western Blueberry. 

The shrubby western blueberry was encountered in northwestern Wyoming, 
from Lewis Lake, Yellowstone Park, south to Jackson Lake, and east on the 
Wind River Range to Fremont Peak. It attains a height of nearly 2 feet in 
Yellowstone Park, growing in dense clumps in lodgepole pine forests between 
7,000 and 8,000 feet elevation. Its small blue berries were ripe south of Lewis 
Lake on August 18, 1910. South of Fremont Peak it was growing in bogs a 
little below timberline, at 10,500 feet. 

Fraxinus lanceolata Borekhausen. Green Ash. 

The green ash occurs sparingly in the low stream valleys of extreme eastern 
and northern Wyoming west to the base of the Bighorn Mountains. It was 
found at Newcastle; Sand Creek to 10 miles above Beulah; Hay Creek to 
Aladdin and Eothen; groves on the Belle Fourche flats at Devils Tower; Little 
Missouri River; gulches near Morse; Powder River near Montana boundary; 
Prairie-dog Creek; Sheridan; Big Goose Creek; Wolf Creek, to 4,500 feet eleva- 
tion, near Eaton's Ranch. 

Sambucus microbotrys Rydberg. Red Elderberry. (Fig. 16.) 

The red elderberry is a characteristic shrub of the Canadian Zone coniferous 

forest on the main ranges of Wyoming. Usually it is the only elderberry present 

on the higher slopes, but lower down occurs with its black-fruited relative, 



l'.KT.I 



TREES AND SHRUBS. 



77 



Sambucus melanocarpa. The red elder was not taken in the foothill ranges of 
eastern Wyoming nor in the Bighorn Mountains, but it has a general distribu- 
tion from 7,0(10 to 9,."i00 feet elevation in the mountains farther west. 

Sambucus melanocarpa (J ray. Black Elderberry. 

The black elderberry is of lower vertical range than the red species, occurring 
mainly in the Transition Zone, and from our observations was not so generally 
distributed. It was noted on Laramie Peak, north slope, rare at 8,000 feet 
elevation; Shirley Mountains, north escarpment, 7,600 feet to summit; Ferris 
Mountains, in dense coniferous forest, 8,000 to 9,000 feet ; Salt River Range, 
7.000 feet. Vernon Bailey found it bearing ripe berries on August 10, at 7,500 
feet, on a southwest slope in the Bighorn Mountains near Wolf. 




Pig. 10. — Red elderberry (Sambucus microbotrys) in bloom, Sierra Madie, July 7, 1911 

(10,000 feet). 

Sambucus canadensis Linnaeus. Elderberry. 

The large elderberry, Sambucus canadensis, was detected only at the southern 
base of the Bear Lodge Mountains near Sundance, in the Transition Zone. Its 
handsome flat-topped cymes of white flowers were very conspicuous in the 
canyon along Sundance Creek, at 4,700 feet elevation, on June 20, 1912. 

Viburnum lentago Linnaeus. Sweet Viburnum. 

Vernon Bailey found the sweet viburnum not uncommon in August, 1913. in 
Hie shaded canyons and gulches along Sand Creek above Beulah, at the northern 
base of the Black Hills. He also reports it from the Belle Fourche Valley near 
Devils Tower, and along Big Goose Creek at the cast base of the Bighorn 
.Mountains, 



78 NORTH AMERICAN FAUNA. [No. 42. 

Symphoricarpos occidentalis Hooker. Wolfberry. 

The wolfberry occurs over much of the Upper Sonoran Great Plains area, 
forming dense thickets 2 feet in height in gulches and along si reams, and in 
many localities constituting the principal shrubby growth. The species regu- 
larly extends west to the bases of the Laramie and Bighorn Mountains, and 
was found also in Wind River Basin, and at the eastern base of the Absaroka 
Range northwest of Cody. 

Symphoricarpos pauciflorus (Robbins) Britton. Few-Flowered Snowberry. 

The few-flowered snowberry is higher ranging than the wolfberry (Symphori 
earpos occidentalis) and occurs mainly in the Transition Zone in northern 
Wyoming. It was recorded at various altitudes from 4,700 feet to 8,500 feet 
in the Bear Lodge, Bighorn, Absaroka, and Wyoming Ranges. 

Symphoricarpos oreophilus Gray. Mountain Snowberry. 

The mountain snowberry is the predominating species of its genus in Wyoming, 
where it is rarely absent from dry, rocky, open, or partly forested slopes and 
ridges in the Transition Zone. On some of the southern and western ranges it 
is common on exposed points in dry forest openings up to 9,000 feet elevation. 
This shrub enters the lower portions of Yellowstone Park at the north and 
south, and occurs in the open sections of Jackson Hole. 

Symphoricarpos rotundifolius Gray. Round-Leaved Snowberry. 

The round-leaved snowberry is apparently uncommon. It is represented by a 
specimen from Rawhide Butte, taken July 23, 1000, where scattering bushes 
were noted from base to summit in the Transition Zone. 

Lonicera glaucescens Rydberg. Douglas Honeysuckle. 

The Douglas honeysuckle reaches Wyoming in the extreme northeast, where 
it is found sparingly along streams and on damp slopes above 4,700 feet eleva- 
tion near Sundance, in the Bear Lodge Mountains. 

Lonicera utahensis Watson. Utah Honeysuckle. 

The Utah honeysuckle abounds in the mountain forests of extreme western 
and northwestern Wyoming. From the forests of Yellowstone Park, where it 
is especially abundant, it occurs south at least to the Salt River Mountains, 
reaching its upper limits on the western slope of this range at 9,000 feet eleva 
tion, and on the eastern slope of the Teton Range at about 8,000 feet. 

Lonicera involucrata Banks. Involucred Fly Honeysuckle. 

In Wyoming the involucred fly honeysuckle occurs commonly in all the prin- 
cipal ranges with the possible exception of the Bighorn Mountains. It becomes 
increasingly abundant westward, and is one of the most characteristic shrubs 
on the banks of cold streams up to 8.000 or 9.000 feet elevation in the Canadian 
forest belt, ripening its blackish berries about the first of August. 

Chrysothamnus graveolens (Nuttall) Greene. Rabbit Brush. 

The plains, deserts, and basal mountain slopes of Wyoming are rich in species 
of Chrysothamnus, some of them among the best characterizing shrubs of the 
Upper Sonoran or Transition Zones. The meager data secured by Survey par- 
ties relates mainly to species which are of importance in determining life zones. 
€. graveolens is a conspicuous shrub, often several feet high, growing in Upper 
Sonoran gulches, desert arroyos, and on dry slopes, principally in eastern and 
northern Wyoming. It is especially abundant on flats along Bighorn River iv 
the lower portions of the Bighorn Basin. The species occurs at Cassa ; Fori 
Fetterman; Casper; along the North Platte near Alcova ; Hay Creek east of 



1917.] TEEES AND SHRUBS. 79 

Aladdin; Wind < 'reek northeast of Moorcroft; Buffalo; gulches ;it south base <>f 
Owl Creek Mountains; Badwater River north of Shoshoni; Worland; Mander- 
son ; Greybull ; Frannie to Garland ; and east of Cody. 

Chrysothamnus linifolius Greene. Rabbit Brush. 

Like Chrysothamnus graveolcns, this species is a fair-sized shrub, inhabiting 
principally the cut banks of adobe along desert washes and streams in the 
Upper Sonoran Zone, as along Bear Creek, south of Cassa; Rock Springs; and 
warm pockets along Green River between Fontenelle and Labarge. Vernon 
Bailey collected a specimen at Arvada, on Powder River, August 13, 1913. 
Nelson found the species at Point of Rocks and Granger. 1 

Chrysothamnus plattensis Greene. Rabbit Brush. 

Chrysothamnus plattensis is a low, spreading Upper Sonoran species of rabbit 
brush of the North Platte and tributary valleys at the southeast. It was noted 
up to 6,200 feet elevation on Horse Creek, near Meadow, and also at Snow's 
Ranch, on Rawhide Creek below Patrick. 

Chrysothamnus stenophyllus Greene. Rabbit Brush. 

This is a common species of rabbit brush over most of the sandy plains and 
ridges of central Wyoming, in the Upper Sonoran Zone, as al Casper, Bfell, and 
in Sweetwater Valley from Independence to Splitrock. Nelson records it from 
Point of Rocks, on the Red Desert. 1 

Chrysothamnus stenophyllus Greene. Rabbit Brush. 

This species was taken in the Upper Sonoran Zone in the Belle Fourche 
Valley at Moorcroft, August 15, 1913, by Vernon Bailey. 

Chrysothamnus howardi (Parry) Greene. Rabbit Brush. 

A specimen of Chrysothamnus howardi was collected by Vernon Bailey at 
Arvada, on Powder River, August 13, 1913. It is usually of the higher plains. 

Chrysothamnus wyomingensis A. Nelson. Rabbit Brush. 

Mainly of the high Transition plains and basal mountain slopes at the north- 
It is very abundant at the northern base of the Bighorn Mountains at Wolf, 
where a specimen was collected on June 6, 1912, and is apparently the species 
so abundant on the basal slopes of the Bear Lodge Mountains. Vernon Bailey 
collected it at Arvada, on Powder River. 

Chrysothamnus pulcherrimus A. Nelson. Rabbit Brush. 

This rabbit brush is common in the Transition Zone on the high central 
plains and in the upper Green River Basin, growing on open slopes with sage- 
brush regularly to 8.000 feet altitude. It is especially abundant on both slopes 
of the mountains near Laramie Peak, on the northern Laramie Plains, and 
in Shirley Basin. It was noted along Little Medicine Bow River west of Mar- 
shall at 7,000 feet. 

Artemisia tridentata Nuttall. Black Sagebrush. (Fig. 17.) 

The most widely distributed shrub in Wyoming is the black sagebrush. It 
is omnipresent in open country east to the edge of the Great Plains, where its 
eastern limits are marked by Grin Junction (North Platte Valley), Lost Spring. 
Indian Creek north of Kirtley at State line, Clifton, Newcastle, Wind Creek 
northeast of Moorcroft, and Colony. It was not found east of the Laramie 
Mountains in southeast Wyoming. From 4.000 feet elevation at the east this 

1 Bull. 13, Div. <>f Agrost., U. S. Dopt. Agr., p. 66, 1898. 



80 



NORTH AMERICAN FAUNA. 



[No. 42. 



sagebrush extends to timberline on some of the ranges, attaining 9.800 feet 
on the Bighorns above Hyattville, and 10,000 feet on the Wind Rivers south 
of Dubois and on the Wyoming Range west of Bigpiney and Merna. 

Artemisia cana Pursh. Gray Sagebrush. 

The gray sagebrush is generally dispersed and especially common in the 
mountain valleys and parks, but less abundant than the black species. It ex- 
tends into the edge of the Great Plains area. In the valleys east of the Laramie 
Mountains this is usually the only shrubby sage present, and it forms much 
of the scattering growth on the open stretches between the Black Hills and 
Bighorn Mountains. Westward it rapidly gives way to the black sagebrush. 




Fig. 17. — Black sagebrush (Artemisia tridentata) 10 feet high, upper Wind River Valley. 



Artemisia trifida Nuttall. Sagebrush. 

A small shrubby sage, sparingly present in open Transition country chiefly 
at the west, as on the summit of the Owl Creek Mountains; Bear River Divide 
north of Evanston, 7,500 feet elevation; benches above Cokeville; and Salt 
River Valley. Vernon Bailey found the species in open parks in the mountains 
south of Douglas, near Valley, and in Jackson Hole. 

Artemisia fllifolia Torrey. Narrow-Leaved Sagebrush. 

The narrow-leaved sagebrush is a characteristic shrub of the Great Plains 
Sonoran area, occurring in the North Platte drainage chiefly in sandy tracts as 
far west as Guernsey. A very dense growth 2 or 3 feet high covers the Rawhide 
flats at Snow's Ranch south of Patrick, and it is abundant in the sandy valley 
of the Chugwater east of Wheatland, 



1917.] TREES AND SHRUBS. 81 

Artemisia arbuscula Nuttall. Brown Sagebrush. 

The brown sagebrush is uncommon, according to the author's observations. 
Vernon Bailey reports it at Hams Fork Station (Moyer Junction) and Valley, 
and collected specimens on the Owl Creek Mountains in 1S93 and in Salt River 
Valley on August 15, 1911, all Transition Zone localities in western Wyoming. 

Artemisia spinescens Eaton. Spiny Sage ; Bud Brush. 

A low spinescent shrubby sage, abundant at many localities in the Upper 
Sonoran desert region at the southwest. It is common, also, on sandy benches 
near Shoshoni and in the lower parts of the Bighorn Valley near Greybull and 
Manderson. It occurs at many stations with the less shrubby Artemisia 
ijedati/ida, with which it is sometimes confused. 

Artemisia frigida Willdenow. Pasture Sage. 

The pasture sage, a low silvery sage, is scarcely a shrub, but is included as 
a characteristic Transition species of the genus. It has a very wide range 
over Wyoming on high plains, bare ridges and plateaus, and open mountain 
slopes generally. 

Tetradymia spinosa Hooker and Arnott. Spiny Rabbit Brush. 

The members of the genus Tetradymia are characteristic shrubs of the Great 
Basin Division of the Upper Sonoran Zone, and do not enter the Great Plains 
area. They occur in varying abundance and scattered bunchlike growths over 
the dry hills and plains of the arid central desert section, from the Green River 
Valley and Red Desert north to the Bighorn Basin. 

Tetradymia spinosa is the more widely distributed species below 6,500 feet 
altitude. It is common at Green River, Superior, Rock Springs region, warm 
river flats (Fontenelle to Labarge), sand flats north of Opal, Shoshoni, Wor- 
land, Bonanza, Manderson, Greybull to Cody, and Frannie to Garland. 

Tetradymia nuttalli Torrey and Gray. Nuttall Rabbit Brush. 

The Nuttall rabbit brush is more spiny than the preceding species. It was 
collected only at Fort Steele, where it was abundant in the warm, lower parts 
of the North Platte Valley. Nelson mentions specimens from Bitter Creek 
and Green River. 1 

Tetradymia inermis Nuttall. Rabbit Brush. (PI. VII, fig. 2.) 

This high-ranging rabbit brush grows abundantly on the sandy plains of 
the upper Green River Basin to an elevation of 7,500 feet. It was not found 
at the lower levels, but was common on sand flats at Eden and Big Sandy and 
also on Little Piney Creek. Nelson collected specimens at Bitter Creek on the 
Red Desert.' 



1 Bull. 13, Div. of Agrost., U. S. Dept. Agr., p. 67, 1898. 
74440°— 17 Q 



INDEX 



[Pages containing the principal reference to a species in bold-faced figures.] 



A. 



Abies (concolor?), 44, 58. 

lasiocarpa, 49, 58. 
Abronia elliptica, 16, 28. 

fragrans, 28. 
Accipiter cooperi, 35. 

velox, 35. 
Acer glabrum, 36, T3. 

grandidentatuni, 36, 73. 

negundo, 28, 73. 
Achillea alpicola, 52. 
Acknowledgments, 8 (footnote). 
Aconitum columbianum, 45. 
Actsea arguta, 45. 

rubra, 45. 
Adenostegia ramosa, 37. 
Aeronautes melanoleucus, 35. 
Agelaius phceniceus fortis, 26. 
Agriculture, Arctic-Alpine Zone. See Boreal 
Zones. 

Boreal Zones, 52-54. 

Canadian Zone, 42. See also Boreal 
Zones. 

Hudsonian Zone. See Boreal Zones. 

Transition Zone, 37-38. 

Upper Sonoran Zone, 30-31. 
Great Basin Division, 18. 
Bighorn Basin, 24. 
Green River Valley, 19. 
Red Desert, 21. 
Wind River Basin, 22-23. 
Great Plains Division, 17. 
Agropyron caninum, 37. 

pseudorepens, 37. 

smithi, 30. 

spicatum, 30, 37. 

violaceum, 46. 
Agrostis hiemalis, 37. 
Alces americanus shirasi, 42. 
Alder, 44, 64. 
Alnus tenuifolia, 44, 64. 
Alopecurus occidentalis, 46. 
Alsinopsis ohtusiloba, 52. 
Alum root, 45. 
Ambystoma tigrinum, 27, 33. 
Amelanchier alnifolia, 36, 71. 

elliptica, 36, 

oreophila, 36. 
Ammodramus savannarum bimaculatus, 26. 
Amorpha canescens, 28, 72. 

nana, 28. 
Amphibian lists, 19, 27, 33. 
Amphispiza nevadensis, 27. 



Anaphalis suhalpina, 4."i. 
Anas platyrhynchos, 34. 
Andropogon halli, 29. 

scoparius, 29. 
Androsace carinata, 52. 

subumbellata, 49. 
Anemone, 37, 45, 49. 
Anemone cylindrica, 37. 

globosa, 49. 

lithophila, 45. 

tetonensis, 49, 
Angelica roseana, 49. 
Anogra albicaulis, 16, 29. 
Antelope, 25. 
Antennaria pulcherrima, 45. 

reflexa, 37, 50. 
Anthus rubescens, 51. 
Antilocapra americana americana, 25. 
Apocynum adrosa^mifolium, 37. 
Aquilegia cserulea, 45. 

flavescens, 45. 

oreophila, 45. 

saximontana, 49. 
Arabia lyalli, 52. 
Aragallus lagopus, 52. 

lamberti, 37. 

nanus. 52. 
Aralia nudicaulis, 37. 
Archibuteo ferrugineus, 26. 
Arctostaphylos uva-ursi, 36, 76. 
Ardea herodias, 26. 
Arenaria congesta, 36. 
Argemone hispida, 28. 

intermedia, 16, 28. 
Aristida longiseta, 29. 
Arnica, heart-leaved. 45. 
Arnica cordifolia, 45. 
Artemisia arbuscula, 36, 81. 

cana, 36, 80. 

filifolia, 28, 80. 

frigida, 36, 81. 

ludoviciana, 36. 

pedatifida, 28, 81. 

scopulorum, 52. 

spinescens, 28, 81. 

tridentata, 28, 36, 79-SO 

trifida, 36. 80. 
Asclepias pumila, 29. 

speciosa, 29. 
Ash, 16, 28. 

green, 76. 

mountain, 44, 71. 
Asio wilsonianus, 35. 
Aspen, 39, 40, 41, 42, 44, 60. 

83 



84 



NORTH AMERICAN FAUNA. 



[No. 4: 



Astragalus alpinus, 45. 

calycosus, 37. 

crassicarpus, 16, 29. 

drummondi, 37. 

flexuosus, 37. 

hypoglottis, 37. 

missouriensis, 29. 

mollissimus, 16, 29. 

nitidus. 37. 

succnlentus, 37. 
Asyndesmus lewisi, 35. 
Atheropogon curtipendula, 30. 
Atriplex argentea, 28. 

canescens, 28, 65. 

confertifolia, 28, 65-66. 

nuttalli, 28, 66. 

pabularis, 28. 
Avens, 49. 

alpine, 51, 52, 69. 

mountain, 45, 52. 
Avocet, 34. 

B. 

Badger, 26, 34. 

Baeolophus inornatus griseus, 27. 
Balm, mountain, 33, 36, 73. 
Balsamorrhiza, 40. 
incana, 37. 
sagittata, 37. 
Balsam root, 37. 
Baneberry, 45. 
Barberry. 33, 36, 67. 
Bartramia longicauda, 26. 
Bascanion constrictor, 27. 
Bat, big-eared. :'.4. 
brown, 24, 26. 
little, 43. 
California, 16, 22, 24, 26. 
boary, 43. 
long-eared, 20, 26. 
long-legged, 26, 33, 34. 
silver-haired, 43. 
Bean, buffalo, 29. 
Bear, black, 43. 
grizzly, 34. 

Washakie, 43. 
Yellowstone Park, 43. 
Bearberry. 33, 36. 

red, 76. 
Beard-tongue, 29, 37, 45, 49. 
Beaver, 34, 43. 
Berberis aquifolium, 36, 67. 
Besseya alpina, 52. 
Betula fontinalis, 36, 63-64. 
glandulosa, 44, 64. 
papyrifera, 44, 63. 
Bilberry, red. 45. 76. 
Birch, canoe, 40, 42, 44, 63. 
dwarf, 44, 64. 

Rocky Mountain, 33, 36, 03-64. 
Bird lists, 16, 19, 20-21, 22, 24, 26-27, 

34-35, 43-44, 49, 51. 
Bitter root, 36. 
alpine, 49. 
Bittern, 26. 
Blackbird, Brewer, 35. 
yellow-headed, 26. 



33, 



Bladder-pod, double, 37. 
Blazing star, 29. 
Blite, sea, 28. 

shrubby, 28. 
Blueberry, 45. 

mountain, 44, 76. 
western, 76. 
Bluebird, mountain, 35, 44. 
Bluegrass, 50, 52. 
alpine, 52. 
arctic, 52. 
mountain, 46. 
Bluestem, 29. 
Bobolink, 35. 

Botaurus lentiginosis, 26. 
Bouteloua hirsuta, 30. 

oligostachya, 30. 
Boykinia heucheriformis, 52. 
Branta canadensis, 43. 
Bromus marginatus, 37. 

porteri, 37. 
Brush, antelope, 69. 
bud, 81. 

rabbit, 16, 17, 19, 20, 28, 36, 78-79, 
79, 81. 

Nuttall, 81. 
spiny, 81. 
Buckthorn, 44, 73. 
Budbrush, 28. 
Buffaloberry, 22, 28, 74. 

Canadian, 44, 74. 
Buffle-head, 43. 
Bufo boreas, 33. 
cognatus, 27. 

lentiginosis woodhousei, 27, 33. 
Bulbilis dactyloides, 30. 
Bunting, lark, 16, 27. 
lazuli, 16, 24, 27. 
Bnpleurum americanum, 52. 
Bur, buffalo, 29. 
Bush, skunk, 28. 
Bush-tit, lead-colored, 27. 
Buteo swainsoni, 35. 
Buttercup, 48, 49, 52. 
Nuttall, 37. 



C. 



Cactus, 18. 
ball, 29. 

Simpson, 37. 
prickly-pear, 20, 22, 23. 
small-jointed, 37. 
Calamagrostis canadensis acuminata, 46. 
Calamospiza melanocorys, 27. 
Calamovilfa longifolia, 29. 
Calandrinia pygmsea. 49. 
Calcarius ornatus, 35. 
Calochortus gunnisoni, 36. 
nuttalli, 36. 
pavonaceus 45. 
Callospermophilus lateralis caryi, 43, 48, 51. 
castanurus, 43. 
lateralis, 34, 43. 
wortmani, 25, 34. 
Caltha leptosepala, 49. 
Calypso bulbosa, 45. 



1917.] 



INDEX. 



85 



Camas, beautiful, 45. 

poison, 36. 
Campanula parryi, 49. 

rotundifolia, 37. 

uniflora, 52. 
Canis lestes, 34, 48, 51. 

nebracensis, 26. 

nubilus, 34. 
Canyon mouse. See Mouse. 
Carcluus plattensis, 16, 29. 
Carex albo-nigra, 52. 

atrata, 52. 

engelmanni, 52. 

nigricans, 50. 

nova, 50. 

nubicola, 52. 

phseocephala, 52. 
Carpodacus cassini, 44. 

mcxicanus frontalis, 26. 
Castilleja, 49. 

sulphurea, 45. 
Castor canadensis, 34, 43. 
Catbird, 16, 24, 27. 
Catchfly, stemless, 52. 

Catoptrophorus semipalmatus inornatus, 34. 
Ceanotbus fendleri, 36, 73. 

mollissimus, 36, 73. 

velutinus, 36, 73. 
Centrocercus urophasianus. 35. 
Cerastium beeringianum, 49. 
Cercocarpus intricatus, 36, 68. 

ledifolius, 36, 68. 

parvifolius, 28, 36, 68-69. 
Certbia familiaris montana. 49. 
Cervus canadensis canadensis, 34, 42, 51. 
Chaenactis alpina, 50. 

douglassi, 37. 
Charitonetta albeola, 43. 
Cbat, long-tailed, 16, 22, 24, 27. 
Chaulelasmus streperus, 34. 
Cberry, ground, 29. 

red, wild, 36, 72. 

sand, 16, 28. 

wild, 33. 
Chickadee, long-tailed, 35. 

mountain, 44. 
Chickweed, mouse-ear, 49. 
Chimaphila umbellata, 45. 
Chipmunk, buff-bellied, 43. 

•Colorado, 42. 

least, 19, 20, 25. 

northern, 42. 

pale, 22, 24, 25. 

rock, Utah, 19, 25. 

sagebrush, 25. 

timberline, 48, 51. 

Uinta, 43. 
Chokecherry, 36, 72. 
Chondestes grammacus strigatus, 26. 
Chordeiles virginianus henryi, 26. 
Chorophilus triseriatus, 33. 
Chrysothamnus frigidus, 36. 

graveolens, 28, 78-79. 

howardi, 79. 

lanceolatus, 79. 

linifolius, 28, 79. 

parryi, 36. 



Chrysothamnus plattensis, 28, 79. 

nulcherrimus, 36, 79. 

stenophyllus, 28, 79. 

wyomingensis, 36, 79. 
Cinclus mexicanus unicolor, 44. 
Cinquefoil, 37, 49. 

glandular, 37. 

shrubby, 40, 44, 47, 51, 71. 
Citellus armatus, 34, 51. 

obsoletus, 25. 

ricbardsoni elegans, 34. 

tridecemlineatus pallidus, 2.~>. 
parvus, 25. 
Clammy-weed, 28. 
Clangula islandica, 43. 
Claytonia megarrhiza, 52. 

rosea, 45, 49. 
Clematis douglasi, 36. 

ligusticifolia, 36. 

occidentalis, 36. 

pseudalpina, 45. 
Clementsia rhodantba, 49. 
Cleome. red, 28. 

yellow, 18, 19, 20, 28. 
Cleomelutea, 23. 28. 

serrulata, 16. 28. 
Cliff mouse. See Mouse. 
Climate, 11-12. 
Clover, 45. 

dwarf, 49. 

Parry, 49. 

prairie, 29. 

purple, 29. 
silky, 29. 
white, 29. 
Cnemidophorus sexlineatus, 27. 
Coccyzus erythropthalmus, 35. 
Columbine, 45. 48, 49. 

blue, 45. 

white, 47. 

yellow, 45. 
Colymbus nigricollis californicus, 34. 
Cone flower, 29. 
Coney, 48, 51. 
Coot, 26. 

Corallorhiza multiflora, 36. 
Coral root, 36. 
Cornel. 33, 36. 
Cornus instolonea, 36. 

stolonifera, 36. 
Corvus brachyrhynchos hesperis, 26. 
Corylus rostrata, 44, 63. 
Corynorhinus macrotis pallescens, 34. 
Cottontail, Bailey, 16, 20, 22, 24, 26. 

Black Hills. 33, 34. 

Nebraska, 26. 
Cottonwood, broad-leaved, 15, 22, 28, 
60-61. 

lance-leaved, 28, 61. 

narrow-leaved, 33, 36, 61. 
Coyote, 51. 

mountain, 34, 48. 

plains, 26. 
Cowslip, mountain, 48, 49. 
Crataegus cerronis, 36, 71. 

rivularis, 36, 71. 

sheridana, 36. 



86 



NORTH AMERICAN FAUNA. 



[No. 42. 



Creeper, Rocky Mountain, 49. 

Virginia, 28. 
Cross, Whitlow, 52. 
Crossbill, 44. 
Crotalus confluentus, 27. 
Croton, 29. 

texensis, 16, 29. 
Crow, western, 26. 
Crowfoot, 45. 
Cryptoglaux acadica, 35. 
Cuckoo, black-billed, 35. 
Curlew, long-billed, 26. 
Currant, 36, 44, 68. 

black, mountain, 44, G7-G8. 

blue, 44. 

flowering, 16, 18, 22, 28, G7. 

red, 15, 36, 67. 

bristly, 49, G7. 
spiny, 51. 
wild, 33. 
Cyanocephalus cyanocephalus, 26, 35. 
Cyanocitta stelleri annectens, 44. 
Cynomys leucurus, 34. 

ludovicianus ludovicianus, 25. 
Cyrtorbyncha ranunculina, 37. 

D. 

Dalea, 29. 

Danthonia intermedia, 46. 
Dasiphora fruticosa, 44, 51, 71. 
Deer, mule, 34. 

white-tailed, plains, 33, 34. 
Delphinium subalpinum, 49. 
Dendragapus obscurus riehardsoni, 43. 
Dendroica sestiva sestiva, 27. 

auduboni, 35. 

nigrescens, 35. 
Deschampsia csespitosa, 46. 
Dickcissel, 27. 
Dipper. 44. 

Disporum tracbycarpum, 45. 
Distichlis spicata, 30. 
Dock, 45. 

sand, 23, 28. 
Dodecatheon radicatum, 49. 
Dolichonyx oryzivorus, 35. 
Dove, mourning, 16, 22, 24, 26. 
Draba, 48. 

cana, 52. 

crassifolia, 52. 

densifolia, 52. 

luteola, 49. 
Dryas octopetala, 52, 69. 
Drymocallis glandulosa, 37. 

pseudorupestris, 49. 
Dryobates pubescens bomorus, 35. 

villosus monticola. 35. 
Dumotella carolinensis. 27. 

E. 

Eatonia obtusata, 30. 
Echinocactus simpsoni, 37. 
Echinocereus viridiflorus, 29. 
Edwinia, 68. 

americana, 36. 68. 
Elseagnus argentea, 36, 74. 
Elder, box. 16, 28, 73. 



Elderberry, 36, 77. 
black, 77. 

mountain, 36. 
red, 45, 76-77. 
Elephantella groenlandica, 45. 
Elephant-head, 45. 
Elk, 34, 42, 51, 54. 
Elm, 28, 64. 
Elymus canadensis, 30. 
condensatus, 30. 
glaucus, 46. 

salinus, 30. 
Empidonax difficilis, 44. 

hammondi, 35. 

minimus, 35. 

wiighti, 35. 
Endolepsis suckleyana, 28. 
Epilobium, 45. 
Eptesicus fuscus fuscus, 26. 
Erethizon epixanthum, 34, 43. 
Erigeron compositus, 50. 

pinnatisectus, 52. 

radicatus, 52. 

salsuginosus, 50. 
Eriocoma cuspidata, 29. 
Eriogonum, 18, 28. 

annuum, 16, 28. 

campanulatum, 28. 

corymbosum, 28. 

effusum, 28. 

multiceps, 28. 
Eritricbium argenteum, 52. 
Erythronium parviflorum, 45. 
Eumeces multivirgatus, 27. 
Euphagus cyanocephalus, 35. 
Euphorbia marginata, 29. 
Eurotia lanata, 28. 
Eutamias borealis, 42. 

dorsalis utahensis, 25. 

luteiventris, 43. 

minimus minimus, 25. 
pictus, 25. 

operarius, 42. 

oreocetes, 48, 51. 

pallidus pallidus, 25. 

umbrinus, 43. 
Everlasting, 37, 45, 50. 

pearly, 45. 
Evotomys brevicaudus, 43. 

gapperi galei, 43, 48. 



Felis hippolestes, 34, 43. 
Ferret, black-footed, 16, 26. 
Fescue, alpine, 52. 

Thurber, 46. 
Festuca brachyphylla, 52. 

octoflora. 30. 

ovina duriuscula, 37. 

tburberi. 46. 
Fiber zibethicus cinnamominus, 26. 

osoyoosensis. 34. 
Field mouse. See Mouse. 
Finch, house, 24. 26. 

purple, Cassin. 44. 

rosy, black, 51. 

brown-capped, 51. 



1917.] 



INDEX. 



87 



Fir, 39, 40, 41. 

alpine, 47, 58. 
(dwarf), 49. 

white, 44, 5S. 
Flag, blue. 36. 
Flax, wild, 29, 37. 
Fleabane, 50, 52. 
Flycatcher, Hammond, 35. 

least, 35. 

olive-sided, 44. 

western, 44. 

Wright, 33, 35. 
Forget-me-not, 48, 49, 52. 
Fox, kit, 26. 

red, mountain, 43, 48, 51. 
Foxtail, mountain, 46. 
Fragaria pauciflora, 45. 
Frasera, 40, 45. 

speciosa, 45. 
Fraxinus lanceolata, 28, 70. 
Frog, 33. 

leopard, 27. 
Fulica amevicana, 26. 

G. 

Gadwall, 34. 
Gallinago delicata, 34. 
Galpinsia lavaudulcefolia, 29. 
Gaura coccinea, 29. 
Gentian, 37, 48, 49. 

blue, 47. 

closed, 45. 

dwarf, 52. 

fringed, mountain, 45. 
Gentiana afflnis, 37. 

calycosa, 49. 

elegans. 45. 

forwoodi, 45. 

romanzovi, 52. 

strictiflora, 49. 
Geomys lutescens, 26. 
Geothlypis trichas occidentalis, 27. 
Geranium, 37, 40, 45. 

coespitosum, 45. 

fremonti, 37. 

parryi, 45. 

richardsoni, 37. 
Gilia, 37. 

congesta, 37. 

polycladoh, 29. 
Glasswort, 19, 28. 
Glaucomys sabrimis bangsi, 42. 

canescens, 34. 
Globe flower, 48, 49. 
Glycyrrhiza lepidota, 29. 
Gnatcatcher, western, 27. 
Golden-eye, Barrow. 43. 
Goldenrod, 29, 52. 
Goose, Canada, 43. 
Gooseberry, 36. 

wild, 33. 
Gopher. See Pocket gopher. 
Grackle, bronzed, 16, 24, 26, 
Grama, tall, 30. 
Grape. Oregon, 36, 67, 

wild, 28. 



Grass, bent, 37. 

brome, 37. 

buffalo, 30. 
false, 30. 

bunch, 29. 

crab, 29. 

dropseed, 29, 37. 

feather, 29, 37. 

fescue, 30, 37. 

grama, 30. 

hair, tufted, 46. 

holy, 37. 

June, 37. 

manna, 37. 

marsh, 30. 

oat, 50. 

mountain, 46. 

panic, 29. 

penny, 45, 49. 

reed, 29. 

mountain, 46. 

rice, 29. 

rye, 46. 

desert, 30. 
giant, 20, 30. 
long-bearded, 30. 

sand, 29. 

salt, 30. 

fine-top, 29. 

spear, 30, 37. 

long-stalked, 37. 

wheat, 20, 30, 37, 46. 

whitlow, 49. 

wire, 29. 
Grasshopper mouse. See Mouse. 
Grass-of-Parnassus, 45. 
Grayia, 18, 19, 20, 28, 66. 

spinosa, 23, 28, 66. 
Greasewood, 17, 19, 20, 21, 28, 66-67. 
Grebe, eared, American, 34. 
Grindelia squarrosa, 16, 29. 
Gromwell, 29. 
Grosbeak, black-headed, 35. 

pine, Rocky Mountain, 48. 
Grossularia inermis, 26. 
Ground squirrel. See Squirrel. 
Grouse, dusky, 76. 

Richardson, 43. 

sharp-tailed, Columbian, 35. 
Gulo luscus, 43. 
Gum plant, 29. 
Gutierrezia sarothrae, 28. 



Harbouria trachypleura, 37. 
Hare, varying, northern, 42, 43, 
Harebell, 37, 48. 

arctic, 52. 

Parry, 49. 
Harvest mouse. See Mouse. 
Haw, black, 33. 

red, 33. 
Hawk, Cooper, 35. 

rough-legged, ferruginous, 26. 

sharp-shinned, 33, 35. 

Swainson, 35. 



88 



NORTH AMERICAN FAUNA. 



[No. 42. 



Hawthorn, 36, 71. 

black, 36, 71. 

red, 36. 
Hazelnut, 44. 

beaked, 42, 63. 
Heath, mountain, 48, 49, 51, 74-78. 
Hedysarum, 37. 

Uinta, 37. 
Hedysarum cinerascens, 37. 

sulphurescens, 49. 

uintahense, 37. 
Helianthus annuus, 16, 29. 

petiolaris, 16, 29. 
Hellebore, white, 45. 
Hemlock, water, 37. 
Hemp, Indian, 37. 
Hen, sage, 33, 35. 
Heracleum lanatum, 45. 
Heron, blue, great, 26. 

night, black-crowned, 26. 
Heterodon nasicus, 27. 
Heuchera parvifolia, 45. 
Ilolbrookia maculata, 27. 
Holodiscus dumosus, 36, 69. 
Honey plant, 28. 
Honeysuckle, 45. 

Douglas, 36, 42, 45, 78. 

involucred fly, 45, 78. 

Utah, 78. 
Hop, wild, 36. 
Hummingbird, broad-tailed, 44. 

calliope, 35. 
Humulus lupulus, 36. 
Hylocichla fuscescens salicicola, 35. 

guttata auduboni, 44. 

ustulata swainsoni, 44. 
Hymenopappus filifolius, 16, 29. 

I. 

Icteria virens longicauda, 27. 
Icterus bullocki, 26. 
Indigo, false, 28, 72. 
Ipomsea leptophylla, 29. 
Iris, 40. 

missouriensis, 36. 
Ironwood, 63. 
Ivy, poison, western, 28, 72. 

J. 

Jacob's ladder, 48, 49, 52. 
Jay, black-headed, 44. 

pinyon, 19, 26, 33, 35. 

Rocky Mountain, 44. 49. 
Juiico, gray-headed, 44, 49. 

pink-sided, 44, 49. 

white-winged, 33, 35. 
Junco aikeni, 35. 

hyemalis mearnsi, 44, 49. 

phaeonotus caniceps, 44, 49. 
Juncoides spicatum, 52. 
Juneus snbtriflorus, 50. 
Juniper, 17, 18, 22, 23. 

creeping, 33, 36. 60. 

desert, 20, 28, 5J>. 

low, 44, 48, 49, 59. 

mountain, 59. 



Juniper, one-seeded, 28, 59. 

Rocky Mountain, 15, 33, 36, 59. 
Juniperus communis, 59. 

knighti, 28, 59. 

monosperma, 28, 59. 

sabina, 36, 60. 

scopulorum, 36, 59. 

sibirica, 44, 49, 59. 

K. 

Kalmia polifolia, 49, 75. 
Kangaroo rat. See Rat. 
Kingbird, 26. 

Arkansas, 16, 22, 24, 26. 

Cassin, 26. 
Kinglet, golden-crowned, 44, 49. 

ruby-crowned, 44. 
Knotweed, alpine, 52. 
Kochia, 20, 28. 

americana, 28. 
Kceleria cristata, 37. 
Kunzia, 33. 

tridentata, 36, 69. 



Lagopus leucurus leucurus, 51. 
Lanius ludovicianus excubitorides, 27. 
Lanivireo solitarius plumbeus, 35. 
Lark, horned, desert, 35, 51. 
Larkspur, 40, 48, 49. 

blue, 47. 
Lasionycteris noctivagans, 43. 
Lathyrus ornatus, 29. 
Laurel, 75. 

mountain, 48, 49. 
Ledum glandulosum, 44. 
Lepargyrea argentea, 28, 74. 

canadensis, 44, 74. 
Leptasea flagellaris, 52. 

hirculus, 49. 
Lepus americanus americanus, 43. 

bairdii bairdil, 43, 48. 

californicus melanotis, 26. 

townsendi campanius, 34. 
townsendi, 34. 
Leucosticte atrata, 51. 

australis, 51. 
Lewisia rediviva, 36. 
Liatris punctata, 16, 29. 
Lichen, 51. 
Licorice, wild, 29. 
Life zones. See Zones. 
Lily, mariposa, 36. 
Limnorchis borealis, 45. 
Linna?a americana, 45. 
Linum lewisi, 37. 

rigidum, 16, 29. 
Lion, mountain, 34, 43. 
Liopeltis vernalis, 27, 33. 
Lippia cuneifolia, 16, 29. 
Lithospermum angustifolium, 29. 

gmelini, 16, 29. 
Lizard, horned, 24, 33. 
desert, 16. 

rock, scaly, 19, 22, 24, 27, 33. 

scaly, 16, 27. 



1917.] 



INDEX. 



89 



Lizard, short-horned, 19. 
desert, 22, 27. 

six-lined, 16, 27. 

Stansbury, 19, 27. 
Loasa, 29. 

yellow, 29. 
Loco, 52. 
Longspur, chestnut-collared, 35. 

McCown, 33, 35. 
Lonicera glaucescens, 30, 45, 7S. 

involucrata, 45, 78. 

utahensis, 45, 7S. 
Lousewort, Parry, 52. 

purple, 45. 
Loxia curvirostra minor, 44. 
Lungwort, 49. 
Lupine, 28, 40, 48, 49. 

mountain, 52. 

purple, 47. 

silvery, 37. 

small, 28. 
Lupinus, 16. 

argenteus, 37. 

esespitosus, 49. 

laxiflorus, 49. 

montieola, 52. 

plattensis, 28. 

pusillus, 23, 28. 
Lutra canadensis canadensis, 43. 
Lygodesmia juncea, 29. 

rostra ta, 10, 29. 
Lynx, Canada, 43. 
Lynx canadensis canadensis, 43. 

uinta, 34. 

M. 

Magpie, 33, 35. 

Mahogany, mountain. 15, 18, 28, 33, 36, 

68, 68-69. 
Malacothrix, 23. 

sonchoides, 29. 
Mallard, 34. 
Mallow, false, 29. 
Malvastrum coccineum, 29. 

dissectum, 29. 
Mamillaria missouriensis, 10, 29. 

vivipara, 16, 29. 
Mammal lists, 16, 19, 20, 22, 24, 25-26, 33. 

34, 42-43, 48, 51. 
Maple, large-toothed, 33, 36, 73. 

mountain, 33, 36, 73. 
Mariposa, yellow, 45. 
Mariposa lily, 36. 
Marmot, 48, 51. 

Black Hills, 43. 

golden-mantled, 43. 

park. 43. 
Mar m ota flaviventris dacota, 43. 
luteola, 43, 4S. 
nosophora, 43, 48, 51. 
Marten, Rocky Mountain, 43, 48. 
Martes caurina origenes, 43, 48. 
Meadowlark, western, 21, 26. 
Meadow mouse. See Mouse. 

rue, alpine, 52. 
Meadowsweet, 44, 69. 
Mi lanerpes erythrocephalus, 35. 



Melospiza lincolni, 44. 

melodia montana, 35. 
Mentzelia albicaulis, 29. 
decapetala, 16, 29. 
lsevieaulis, 29. 
nuda, 29. 
Menziesia ferruginea, 44, 75. 
Mephitis hudsonica, 34. 
Merganser, 43. 
Mergus americanus, 43. 
Meriolix serrulata, 10, 29. 
Mertensia, 48. 

brevistyla, 52. 
tweedyi, 49. 
Micranthes arguta, 45. 
Microtus longicaudus, 43. 
montanus caryi, 34. 
mordax mordax, 43, 4S, 51. 
nanus nanus, 43, 48. 
ochrogaster haydeni, 20. 
pauperrimus, 34. 
pennsylvanicus modestus, 34. 
richardsoni macropus, 43. 
Milkweed, 29. 
Millet, Indian, 29. 
Mimulus floribundus, 37. 
langsdorfii, 45. 
lewisi, 45. 
Mimus polyglottos leucopterus, 27. 
Mink, 34. 
Mint, horse, 37. 
Mockingbird, western, 16, 27. 
Mole, northern plains, 16, 20. 
Monarda rnenthsefolia, 37. 
Moneses uniflora, 45. 
Monkey flower, 37, 45. 

crimson, 45. 
Monkshood, 45, 
Moose, Shiras, 42. 
Morning-glory, bush, 16, 29. 
Moss, 51. 

Mouse, canyon, golden-breasted, 19, 25. 
cliff, True's, 19, 25. 
field, dwarf, 43, 48. 

Hayden, 16, 24, 26. 
pygmy, 33, 34. 
Uinta, 33. 
grasshopper, Great Plains, 16, 22, '..'4, 
25. 

Idaho, 34. 
northern, 34. 
harvest, prairie, 10, 25. 
« jumping, prairie. 33, 34. 
Rocky Mountain, 43. 
meadow, 51. 

big-footed, 43. 
long-tailed, 43. 
Rocky Mountain, 43, 4S. 
Saguache, 34. 
Uinta, 34. 
pocket, 20. 

Cheyenne, 20. 
Kansas, 16, 26. 
Maximilian, 17, 24, 26. 
Red Desert. 26, 
Sweetwater, 17, 26. 
Uinta, 33, .14. 



90 



NORTH AMERICAN FAUNA. 



[No. 42. 



Mouse, red-backed, Black Hills, 43. 
Gale, 43, 48. 
white-footed, 34. 

black-oared, 25. 
sagebrush, 43. 
tawny, 43. 
yellow, 25. 
Muhlenbergia comata, 37. 

pungens, 29. 
Munroa squarrosa, 30. 
Muskrat, Great Plains, 26. 

Rocky Mountain, 34. 
Mustela arizonensis, 34, 43. 
cicognanii leptus, 43. 
nigripes, 26. 
vison energumenos, 34. 
Myadestes townsendi, 44. 
Myiochanes richardsoni, 35. 
Myosotis alpestris, 49. 
Myotis californicus californicus, 26. 
evotis, 26. 

longicrus interior, 26, 34. 
lucifugus carissima, 43. 



X. 



Nannus hiemalis pacificus, 44. 
Neosorex palustris navigator, 43. 
Neotoma cinerea cinerea, 34. 

orolestes, 26, 34. 
Nettion carolinense, 34. 
Nighthawk, western, 21, 26. 
Ninebark, 33, 36, 69. 

western, 69. 
Nucifraga columbiana, 49. 
Numenius americanus, 26. 
Nutcracker, Clark, 49. 
Nuthatch, pygmy, 35. 

red-breasted, 44. 

Rocky Mountain, 35. 
Nuttallornis borealis, 44. 
Nycteris cinerea, 43. 
Nycticorax nycticorax nsevius, 26. 



Oak, bur, 33, 36, 64-65. 

Ochotona uinta, 48, 51. 
Odocoileus hemionus hemionus, 34. 

virginianus macrourus, 34. 
Olor buccinator, 43. 
Onychomys leucogaster arcticeps, 25. 
brevicaudus, 34. 
missouriensis, 34. 
Oporornis tolmiei, 35. 
Opulaster malvaceus, 36, 69. 

monogynus, 36, 69. 

pubescens, 36, 69. 
Opuntia fragilis, 37. 

polyacantha, 16, 29. 

rutila, 29. 
Orchid, bog, 45. 
Oreocarya flava, 29. 

virgata, 37. 
Oreoscoptes montanus, 27, 35. 
Oreospiza chlorura, 35. 
Oreoxis alpina. 52. 



Oriole, Bullock, 16, 22, 24, 26. 

Orpine, red, 49. 

Ortbocarpus, yellow, 37. 

Orthocarpus luteus, 37. 

Oryzopsis micrantha, 29. 

Ostrya virginiana, G3. 

Otocoris alpestris leucolsema, 35, 51. 

Otter, 43. 

Otus asio maxwellise, 35. 

Ouzel, water, 44. 

Oven-bird, 35. 

Ovis canadensis auduboni, 34. 

canadensis, 48, 51. 
Owl, burrowing, 16, 22, 24, 26. 

long-eared, 35. 

saw-whet, 33, 35. 

screech, Rocky Mountain, 35. 
Oxyria digyna, 52. 

P. 

Pachylophus montanus, 37. 
Pachystinia myrsinites, 44, 72-73. 
Paintbrush, 50. 
Painted cup, 40, 45, 48, 49. 

scarlet, 47. 
Panicularia nervata, 37. 
Panicum virgatum, 29. 
Parnassia flmbriata, 45. 
Paronychia pulvinata, 52. 
Parosela enneandra, 29. 
Parrya, purple, 52. 
Parrya nudicaulis, 52. 
Parthenocissus vitacea, 28. 
I'asserculus sandwichensis alaudinus, 35. 
Passerella iliaca schistacea, 35. 
Passerina amoena, 27. 
Pea, wild, 29. 

Pear, prickly, 29. (See also Cactus.) 
Pedicularis parryi, 52. 

racemosa, 45. 
Pedicecetes phasianellus columbianus, 35. 
Pelecanus erythrorhynchos, 43. 
Pelican, white, 43. 
Pentbestes atricapillus septentrionalis, 35. 

gambeli, 44. 
Pentstemon albidus, 29. 

alpinus, 49. 

angustifolius, 16, 29. 

fruticosus, 45. 

glaucus, 45. 

laricifolius, 37. 
Perisoreus canadensis capitalis, 44, 49, 
Perodipus ordii luteolus, 26. 
Perognathus callistus, 26. 

fasciatus fasciatus, 20. 
litus. 26. 

fiavus piperi, 26. 

hispidus paradoxus, 26. 

parvus clarus, 34. 
Peromyscus crinitus auripectus, 25. 

leucopus aridulus, 34. 

maniculatus artemisi», 43. 
nebrascensis, 25. 
osgoodi, 25. 
rufinus, 43. 

truei truei, 25. 



1017.] 



INDEX. 



91 



Petalostemon, 16. 

candidus, 29. 

oligopbyllus, 29. 

purpureus, 29. 

villosus, 29. 
rptaya, green-flowered, 29. 
Pewee, wood, western, 35. 
Phacelia, 37. 

silky, 49. 
Phacelia heterophylla, 37. 

linearis, 37. 

sericea, 49. 
Phalaenoptilus nuttalli nuttalli, 35. 
Phalarope, Wilson, 34. 
Phenacomys, mountain, 43. 
Phenacoruys orophilus, 43. 
Pbiladelphus occidentalis, 28. 
Phleum alpinum, 46, 50. 
Phlox, 48, 49. 

wild, 37. 
rhlox caespitosa, 49. 

dcpressa, 37. 

hoodi, 37. 
Phoebe, Say, 35. 

Phrynosoma ornatissimum, 27, 33. 
Phyllodoce ernpetriformis, 49, 51, 74-75. 
Pbysalis lanceolata, 16, 29. 
Pbysaria didymocarpa, 37. 
Physiography, 9-12. 
Pica pica hudsonia, 35. 
Picea engelmanni, 44, 49, 57. 

parryana, 44, 57-58. 
Picoides americanus dorsalis, 44, 49. 

arcticus, 43. 
Pika, 48. 

Pine, lodgepole, 39, 40, 41, 42, 44, 53, 
56-57. 

nut, 28, 57. 

rock, 36. 

white. Rocky Mountain, 32, 41, 44, 48, 
55-56. 

white-barked, 47, 49, 55. 

yellow, 15, 17, 36, 38, 39 (footnote). 
Rocky Mountain, 56. 
Pinedrops, 45. 

Pinicola enucleator montana, 49. 
Pinus albicaulis, 49, 55. 

edulis, 28, 57. 

flexilis, 44, 55-56. 

murrayana, 44, 56-57. 

scopulorum, 36, 56. 
Pinyon, 18, 28, 57. 
I'ipilo maculatus arcticus, 35. 
Pipit, 51. 
Pipsissewa, 45. 
Piranga ludoviciana, 35. 
1'ituophis sayi, 27. 

Planesticus migratorius propinquiiS4-35. 
Plant lists, 15-16, 17, 18, 19, 20," 22, 23, 
28-30, 32-33, 36-37, 39-42, 44-46, 47- 
48, 49-50, 51, 52, 55-81. 
Plantago purshi, 16, 23; -0. 
Plantain, 29. 
Plover, mountain, 35. 

upland, 26. 



Plum, ground, 29, 37. 

red, wild, 72. 

wild, 28. 
Poa alpina, 52. 

arctica, 52. 

epilis, 50. 

fcndleriana, 30. 

lettermanni, 52. 

longipedunculata, 37. 

lucida, 37. 

reflexa, '46. 

sheldoni, 30. 
Pocket gopher, 51. 

Bighorn, 43, 48. 

Black Hills, 33, 34. 

brown, 43. 

Colorado, 43. 

Coues, 33, 34. 

Fort Bridger, 33, 34. 

Green River, 19, 20, 26. 

pygmy, 33, 34. 

sage, 16, 24, 26. " 

Uinta, 16, 43, 48. 

yellow, 26. 
Pocket mouse. See Mouse. 
Podasocys montanus, 35. 
Polanisia trachyspeima, 10, 23, 28. 
Polemonium confertum, 52. 

mellitum, 49. 

viscosum, 49. 
Polioptila ca?rulea obscura, 27. 
Polygonum, twisted, 49. 
Polygonum bistortoides, 49. 

viviparum, 52. 
Pocecetes gramineus conflnis, 35. 
Poor-will, 35. 
Poplar, aspen. 60. 

balsam, 40, 44, 60. 
Poppy, prickly, 28. 
Populus acuminata, 28. 61. 

augustifolia, 36, 61. 

balsamifera, 44. 60. 

occidentalis, 15, 28,' 60-61. 

tremuloides, 44, 60. 
Porcupine, yellow-haired, 34, 43. 
Porzana Carolina, 26. 
Potentilla effusa, 37. 

glaucophylla, 49. 
Prairie-dog, 16. 

black-tailed, 24, 25. 

white-tailed, 33, 34. 
Primrose, evening, 37. 
white, .28. 

Parry, 48, 49. 
Primula parryi, 49. 
Procyon lotor lotor, 26. 
Primus americana, 28, 72. 

besseyi, 28. 

melanocarpa, 36, 72. 

pennsylvanica, 36,' -72. 
Psaltriparus plumbeus, 27. 
Pseudotsuga mucronata, 44, 58—59. 
Psoralea, 16, 29. 

narrow-leaved, 29. 

silvery, 37. 



92 



NORTH AMERICAN FAUNA. 



[No. 42. 



Psoralea argophylla, 37. 

esculenta, 29. 

hypogea, 29. 

laneeolata, 29. 

linearifolia, 29. 

tenuiflora, 23, 29. 
Ptarmigan, white-tailed, 51. 
Pterospora andromeda, 45. 
Pyrola chlorantha, 45. 

elliptica, 45. 

picta, 45. 

secunda, 45. 

uliginosa, 45. 

Q. 

Quercus macrocarpa, 36, 64—65. 
Querquedula cyanoptera, 26. 

discors, 26. 
Quiscalus quiscula seneus, 26. 

R. 

Rabbit, jack, black-tailed, 26. 
white-tailed, 33, 34. 
western, 34 

snowshoe, 43, 48. 
Raccoon, 26. 
Racer, blue, 16, 27. 
Rail, Carolina, 26. 

Virginia, 26. 
Rallus virginianus, 26. 
Rana pipiens, 27. 33. 

pretiosus, 33. 
Ranunculus adoneus, 52. 

alpeophilus, 49. 

calthaefolius, 49. 

inamoenus, 45. 
Raspberry, dwarfed, 51. 

false. 70. 

flowering, 36, 70. 

red, 44, 49, 70-71. 
Ratibida columnaris, 16, 29. 
Rat, kangaroo, 20, 22, 24. 
Wyoming, 16, 26. 

wood, bushy-tailed, 33, 34. 

Colorado, 16, 26, 34. 
Rattlesnake, plains, 16, 19, 22, 24, 27. 
Recurvirostra americana, 34. 
Red-backed mouse. See Mouse. 
Redwing, thick-billed, 21, 26. 
Regulus calendula, 44. 

satrapa, 44, 49. 
Reithrodontomys megalotis dychel, 25. 
Reptile lists, 16, 19, 24. 27, 33. 
Rhamnus alnifolia, 44, 73. 
Rhodiola integrifolia, 52. 
Rhus rydbergi, 28. 72. 
Rhynchophanes mccowni, 35. 
Ribes americanum. 36. 

inebrians, 36, 67. 

lacustre. 44, 67. 

longiflorum, 28, 67. 

montigenum, 49, 51, 67. 

petiolare, 44, 67-68. 

viscosissimum, 44, 68. 

wolfi, 44. 
Robin, western, 35. 



Rosa sayi, 44. 

Rose, 44. 

Rosewort, 52. 

Rubus deliciosus, 36, 70. 

parviflorus, 44. 70. 

strigosus, 44, 49, 51, 70-71. 
Rumex paucifolius, 45. 

venosus, 16, 28. 
Rush, 50. 

wood, 52. 
Rustyleaf, 75. 
Rydbergia, 48, 49. 

grandiflora, 49. 
Rye, wild, 30. 

S. 

Sage, desert, low, 20. 

pasture, 81. 

spring, 81. 

white, 17, 20, 28. 
Sagebrush, 21, 36, 80. 

alpine, 52. 

black, 17, 20, 28, 36, 79-SO. 

brown, 36, 81. 

small, 19, 22, 28. 

gray, 36, 80. 

narrow-leaved, 16, 28, 80. 

spiny, 22, 28. 
Salamander, tiger, 27, 33. 
Salicornia rubra, 28. 
Salix amygdaloides, 28, 61-62. 

bebbiana, 36, 62. 

chlorophylla, 49. 

cordata watsoni, 36. 

fluviatilis, 20, 28. 

glaucops, 49, 51, 62-63. 

mackenziana, 36. 

nelsoni, 44, 62. 

nivalis, 52, 63. 

petrophila, 52, 63. 

pyrifolia obscura, 44, 62. 

saximontana, 49, 52, 63. 

scouleriana, 36. 

tenera, 52, 63. 
Salpinctes obsoletus, 35. 
Saltbush, 16, 17, 18, 19, 20, 21, 28, 65. 

Nelson, 28. 

Nuttall, 21, 24, 28, 66. 

round-leaved, 65—66. 

silvery, 28. 

spiny, 28. 
Sambucus canadensis, 36, 77. 

melanocarpa, 36, 77. 

microbotrys, 45, 76-77. 
Sandwort, 36, 52. 
Sapsucker, red-naped, 44. 

Williamson, 44. 
Sarcobatus vermiculatus, 28, 66-67. 
Sarsaparilla, wild, 37. 
Savastana odorata, 37. 
Savin, trailing, 36. 60. 
Saxifraga cernua, 52. 
Saxifrage, 45, 48, 49, 52. 

arctic, 52. 
Sayornis sayus, 35. 
Scalopus aquaticus caryi, 26. 
Scaphiopus hammondi bombifrons, 27. 



1917.] 



INDEX. 



93 



Sceloporus consobrinus, 27. 

graciosus, 27, 33. 
Schedonnardus paniculatus, 29. 
Schmaltzia glabra, 28, 72. 

trilobata, 28, 72. 
Sciurus fremonti fremonti, 42. 

hudsonicus baileyi, 42. 
dakotensis, 34. 
ventorum, 42, 48. 
Scutellaria brittoni, 37. 
Sedge, 50, 52. 

black, 52. 
Seiurus aurocapillus, 35. 
Selasphorus platycercus, 44. 
Senecio crassulus, 50. 

fremonti, 50. 
Serviceberry, 33, 36, 71. 
Shadscale, 18. 

gray, 28, 65. 
Sheep, bad lands, 34. 

mountain, 48, 51. 
Shoestring, 28. 
Shooting star, 48, 49. 
Shrew, Dobson, 43. 

masked, 43, 48. 

Rocky Mountain, 43, 48. 

water, white-bellied, 43. 
Shrike, white-rumped, 16, 21, 22, 24, 27. 
Shrubs, distribution, 55-81. 
Sialia currucoides, 35, 44. 
Sibbaldia procumbens, 52. 
Sideranthus spinulosus, 29. 
Sieversia ciliata, 45. 

turbinata, 52. 
Silene acaulis, 52. 
Silverberry, 33, 36, 74. 
Silver-tip, 43. 
Siskin, pine, 44. 
Sitanion hystrix, 30. 

longifolium, 30. 
Sitta canadensis, 44. 

carolinensis nelsoni, 35. 

pygmsea, 35. 
Skink, many-lined, 16, 27. 
Skullcap, 37. 
Skunk, northern plains, 33, 34. 

spotted, Great Basin, 19, 20, 22, 26. 
Rocky Mountain, 34. 
Skunk bush, 16, 18, 72. 
Smelowskia americana, 52. 
Snake, bull, prairie, 16, 24, 27. 

garter, 16, 27, 33. 
red-barred, 27. 
western 33. 

green, smooth, 27, 33. 

hog-nosed, 16, 27. 
Snipe, Wilson. 34. 
Snowberry, 36. 

few-flowered, 78. 

mountain, 33, 36, 78. 

round-leaved, 78. 
Snow-on-the-mountain, 29. 
Solanum rostratum, 16, 29. 
Solidago canadensis gilvocanescens, 29. 

decumbens, 52. 

mollis, 29. 

rigida, 29. 



Solitaire, Townsend, 44. 
Sora, 26. 

Sorbus scopulina, 44. 
Sorex obscurus obscurus, 43, 48. 
personatus personatus, 43, 48. 
vagrans dobsoni, 43. 
Sorrel, mountain, 52. 
Sparrow, Brewer, 20, 26, 35. 
chipping, western, 35. 
fox, slate-colored, 35. 
grasshopper, western, 16, 26. 
lark, western, 16, 20, 22, 24, 26. 
Lincoln, 44. 
sage, 19, 20, 24, 27. 
Savannah, western, 35. 
song, mountain, 33, 35. 
vesper, western, 35. 
white-crowned, 44, 49. 
Spartina gracilis, 30. 
Speedwell, 49. 

alpine, 49. 
Speotyto cunicularia hypugsa, 26. 
Spermophile, Uinta, 51. 

See also Squirrel. 
Sphyrapicus thyroideus, 44. 

varius nuchalis, 44. 
Spiderwort, 28. 
Spilogale gracilis saxatilis, 26. 

tenuis, 34. 
Spinus pinus, 44. 
Spiraea lucida, 44, 6J>. 
Spiza americana, 27. 
Spizella breweri, 26, 35. 

passerina arizona?, 35. 
Sporobolus airoides, 29. 

cryptandrus, 29. 
Spraguea multiceps, 52. 
Spring beauty, 45, 48, 49. 

arctic, 52. 
Spruce, blue, 40, 44, 57-58. 

Douglas, 32, 38 (footnote), 40, 44, 53, 

58-59. 
Engelmann, 39, 40, 41, 42, 44, 47, 
53, 57. 

(dwarf), 49. 
Squirrel, flying, 34. 

Rocky Mountain, 42. 
ground, chestnut-tailed, 43. 
Kennicott, 16, 25. 
mantled, Wind River, 4.'!, 4S, 51. 

Wortman, 25, 34 
Say, 34, 43. 
striped, 20. 
pale, 25. 
small, 25. 
Uinta, 33, 34. 
Wyoming, 33, 34. 
red, Black Hills, 33, 34. 
Bighorn, 42. 
Rocky Mountain, 42, 48. 
Spruce, FremoDt, 42. 
Stanleya, 17, 19, 20, 28. 
integrifolia, 28. 
tomentosa, 28. 
Steganopus tricolor, 34. 
Stellula calliope, 35. 



94 



NORTH AMERICAN FAUNA. 



[No. 42. 



Stipa comata, 29. 

nelsoni, 37. 

viridula. 29. 
Strawberry, 4."). 
Streptopus amplexifolius, 45. 
Sturnella neglecta, 26. 
Suseda, 23. 

diffusa, 28. 

moquini, 28. 
Sumac, smooth, 28, 72. 
Sunflower, 29. 

Swallow, violet-green, northern, 35. 
Swan, trumpeter, 43. 
Swertia congesta, 49. 

palustris. 49. 
Swift, 26. 

sand. 16, 27. 

white throated, 33, 35. 
Sylvilagus auduhoni baileyi, 26. 

floridanus similis, 26. 

nuttalli grangeri, 34. 
Symphoricarpos occidentalis, 28, 7S. 

oreophilus, 36, 78. 

pauciflorus, 36, 7S. 

rotundifolius, 36, 78. 
Syringa, 18. 

western, 28. 



Taehycineta thalassina lepida, 35. 
Tanager, western, 33, 35. 
Taxidea taxus taxus, 26, 34. 
Tea, Labrador, 44. 
Tea bush, wild, 36, 73. 
Teal, blue-winged, 26. 

cinnamon, 26. 

green-winged, 34. 
Telmatodytes palustris plesius, 27. 
Tetradymia inermis, 28. 81. 

nuttalli, 28, 81. 

spinosa, 28, 81. 
Thalictrum alpinum, 52. 
Thamnophis ordinoides vagrans, 33. 

radix, 27, 33. 

sirtalis parietalis, 27, 33. 
Thermopsis, yellow, 37. 
Thermopsis rhombifolia, 37. 
Thimbleberry, 44, 70. 
Thistle, 29. 

Thlaspi glaucum, 45, 49. 
Thomomys bridgeri, 34. 

fossor, 43. 

fuscus fuscus, 43, 51. 

ocius, 26. 

pygmaeus, 34. 

talpoides bullatus, 26. 
caryi, 43, 48. 
clusius, 34. 
nebulosus, 34. 

uinta, 43, 48, 51. 
Thoroughwax, 52. 
Thrasher, brown, 16, 24, 27. 

sage, 21. 27, 35. 
Thrush, hermit, Audubon, 44. 

olive-backed, 44. 

willow, 35. 
Thryomanes bewicki bairdi, 27. 



Timothy, alpine, 46, 50. 
Titlark, 51. 
Titmouse, gray, 10, 27. 
Toad, 27, 33. 

spadefoot, 19, 27. 

western, 33. 
Tonestus pygmanis, 52. 
Towhee, arctic. 33, 35. 

green-tailed, 33, 35. 
Townsendia. Tarry, 50. 
Townsendia parryi, 50. 
Toxostoma rufum, 27. 
Tradescantia occidentalis, 28. 
Trees and shrubs, distribution, 55-81. 
Trifolium anemophilum, 45. 

dasyphyllum, 49. 

parryi, 49. 

rydbergi, 45. 
Trisetum subspicatum, 50. 
Troglodytes aedon parkmani, 35. 
Trollius albiflorus, 40. 
Twinflower, 45. 
Twistedstalk, 45. 
Tyrannus tyrannus, 26. 

verticalis, 20. 

vociferans, 26. 

U. 

Ulmus americana, 28, 64. 
Ursus americanus, 43. 

horribilis, 34. 

imperator, 43. 

washake, 43. 
Uta stansburiana, 27. 



Vaccinium cn?spitosum, 45. 

erythrococcum, 45, 76. 

occidentale, 45, 76. 

oreophilum, 44, 76 
Valerian, 45. 
Valeriana acutiloba. 45. 
Veratrum tenuipetalum, 45. 
Verbena bracteosa, 16, 29. 

hastata, 16, 29. 
Veronica serpyllifolia, 49. 

wormskjoldi, 40. 
Vervain, blue, 29. 

low, 29. 
Vetch, 37. 

milk, 29, 37, 45. 
Drummond, 37. 
Vetchling, 20. 
Viburnum, sweet. 36, 77. 
Viburnum lentago, 36. 77. 
Vicia americana, 37. 
Viola bellidifolia. 4.",. 

canadensis rydbergi, 45. 
Violet, 45. 

dog-tooth, 45. 
Vireo, plumbeous, 33, 35. 

warbling, western, 35. 
Vireosylva gilva swainsoni, 35. 
Virgin's bower, 36. 

purple, 36, 45. 

white, 36. 



1917.] 



INDEX. 



95 



Vitis vulpina, 28. 

Vulpes macrourus, 43, 4S, 51. 

velox velox, 26. 

W. 

Warbler, Audubon, "5. 

gray, black-throated, 35. 
Macgillivray, 33, 35. 
pileolated, 44, 49. 

yellow, 16, 24. 27. 
Weasel, Arizona, 34, 43. 

dwarf, 43. 
White-footed mouse. See Mouse. 
Whitlowwort, 52. 
Wildcat, mountain, 34. 
Willet, western, 34. 
Willow, 16, 20, 36, 44, 40, 51, 62. 

alpine, 47, 51, 52, 63. 

Bebb, 33, 36, 62. 

creeping, 52. 

diamond, 33, 36. 

gray-leaved, 49, 62-63. 

low, 47. 

Nelson, 44, 62. 

net-veined, 63. 

peach-leaved, 28, 61-62. 

rock, 52, 63. 
alpine, 63. 

sand-bar, 28. 
Wilsonia pusilla pileolata, 44, 49. 
Winter fat, 28. 
Wintergreen, 45. 

one-flowered, 45. 

painted, 45. 
Wolf, buffalo, 34. 
Wolfberry, 16, 22, 28, 78. 
Wolverene, 43. 
Woodpecker, Batchelder, 35. 

hairy, Rocky Mountain, 35. 



Woodpecker, Lewis, 33, 35. 

red-headed, 35. 

three-toed, alpine, 44, 49. 
arctic. 4:'.. 
Wood rat. See Rat. 
Wren, Baird, 19, 27. 

house, western, 35. 

marsh, western, 24, 27. 

rock, 35. 

winter, western, 44. 
Wyethia amplexicaulis, 45. 



Xanthocephalus xanthoccphalus, 26. 



Yarrow, alpine, 52. 
Yellow-throat, western, 114, 117. 
Yucca, 16, 22. 2::. 

plains, 28. 
Yucca glauca, 28. 



Z. 



Zamelodia melanocephala, 35. 
Zapus hudsonius campestris, 34. 

princeps princeps, 43. 
Zenaidura macroura carolinensis, 26. 
Zone, Arctic-Alpine, 12, 50-52. 

Canadian, 12, 38-46. 

Hudsonian, 12, 46-50. 

Transition, 12, 31-38. 

Upper Sonoran, 12, 13-31. 
Zones, Boreal, 52-54. 

life, 12-13. 
Zonotrichia leucophrys, 44, 49. 
Zygadenus elegans, 45. 

venenosus, 36. 



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NORTH AMERICAN FAUNA 

ISTo. 43 

[Actual date of publication, September 23, 1918] 




THE RICE RATS OF NORTH AMERICA 



(Genus ORYZOMYS ) 



BY 

EDWARD A. GOLDMAN 

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



t^ 3 .Ulf 

LETTER OF TRANSMITTAL. 



United States Department of Agriculture, 

Bureau of Biological Survey, 

Washington, D. C, April 26, 1917. 
Sir: I have the honor to transmit for publication as North American Fauna No. 43 the results of a 
study of North American forms of the rice rats (genus Oryzomys), by Edward A. Goldman, assistant 
biologist of this bureau. This constitutes a revision of this group, based chiefly upon material in the 
collection of the Biological Survey. Rice rats are distributed from the latitude of Maryland and Delaware 
south through parts of the Southeastern States, Mexico, and Central America to South America, where 
they reach their highest development. While not so injurious to agriculture as some other rodents, they 
consume in the aggregate large quantities of forage when, like cottonrats and meadow mice, they increase 
locally to excessive numbers. A knowledge of their distribution, as presented in thisreport and its accom 
panying maps, will aid in studies made to control the depredations of rice rats. Owing to their nocturnal 
habits the animals are little known, and their economic relations should be better understood. 
Respectfully, . 

E. W. Nelson, 
Chief, Biological Survey. 
Hon. D. F. Houston, 

Secretary of Agriculture. 



CONTENTS. 



Page. 

Introduction 3 

Genus Oryzomys 11 

Subgenus Oryzomys 17 

Subgenus Oligoryzomys 87 

Subgenus Melanomys 94 

Index 99 



ILLUSTRATIONS. 



[Following page 98. 

Plate I. Skulls (dorsal and central views) of Oryzomys. 
II. Skulls (dorsal and ventral views) of Oryzomys. 

III. Skulls (dorsal and ventral views) of Oryzomys. 

IV. Skulls (dorsal and ventral views) of Oryzomys. 

V. Skulls and mandibles (lateral views) of Oryzomys. 
VI. Upper and lower molars (partially worn crowns) of Oryzomys. 

TEXT FIGURES. 

Page. 

Fig. 1. Upper and lower molar crowns of Oryzomys, showing nomenclature of cusps 11 

2. Distribution of subspecies of Oryzomys palustris 22 

3. Distribution of Oryzomys couesi and closely related species 28 

4. Distribution of the Oryzomys meianotis group ." 47 

5. Distribution of the Oryzomys alfaroi group 56 

6. Distribution of Oryzomys talamancx 72 

7. Distribution of subspecies of Oryzomys bombycinus 75 

8. Distribution of the Oryzomys devius group 79 

9. Distribution of subspecies of Oryzomys tcctus 83 

10. Distribution of subspecies of Oryzomys fulvescens 89 

11. Distribution of subspecies of Oryzomys caliginosus (exclusive of South America) 96 

2 



No. 43. NORTH AMERICAN FAUNA. Sept. 23, 1918. 

THE RICE RATS OF NORTH AMERICA. 

(Genus Oryzomys.) 



By Edward A. Goldman. 



INTRODUCTION. 

The rice rats of the genus Oryzomys belong to the murine sub- 
family Cricetina?, which includes also a number of closely allied 
generic groups of American rodents, as the vesper rats (Nyctomys) , 
deer mice (Peromyscus) , grasshopper mice (Onychomys) , and others. 
By reason of their small size and nocturnal habits all these rodents 
largely escape observation, and the economic importance resulting 
from their excessive numbers and wide distribution is not generally 
realized. The vernacular name "rice meadow-mouse," bestowed 
on Oryzomys palustris by Bachman, because originally it was found 
in the rice fields of South Carolina and Georgia, has led to the 
appellation "rice rat" commonly applied to all species of the genus, 
although many inhabit regions where rice is not cultivated. 

Owing to similarity in ratlike form and general appearance, most 
of the genera with which Oryzomys is allied, including species with 
widely differing habits, are scarcely recognizable by external char- 
acters alone, and reliance must be placed on distinguishing features 
exhibited by skulls and teeth. The genus Oryzomys comprises a 
somewhat composite assemblage of species presenting considerable 
diversity in general characters, but having the essential dental 
arrangement repeated with remarkable fidelity throughout the 
series. 

The general range of the genus is from New Jersey and the central 
part of the Mississippi Valley southward, the group being represented 
in nearly every part of South America and apparently reaching its 
greatest development there. More than 150 species and subspecies 
have been described, of which number, however, some have been seg- 
regated in the erection of closely allied genera, and others are doubt- 
fully allocated. Many regions remain unexplored and the number 
of forms assignable to Oryzomys as now restricted will probably far 
outnumber those of any other genus of American rodents. In the 
present revision are treated the North American continental species 

3 



4 NORTH AMERICAN FAUNA. [No. 43. 

as far as the eastern border of Panama, and those of outlying islands. 
Fifty-one forms of 21 species are recognized, two of which are char- 
acterized for the first time. These are comprised in the three sub- 
genera Oryzomys, Oligoryzomys, and Melanomys. 

HABITS AND ECONOMIC RELATIONS. 

The habits of all the rice rats are somewhat similar, but differ in 
details in conformity with varying environmental conditions. In 
general, a preference is shown for meadows or marshy areas, com- 
monly in the vicinity of water at rather low elevations; but some 
species have ascended, especially along the courses of streams, to 
high altitudes; others, departing farther from the semiaquatic 
environment, have entered the forest and become partially scan- 
sorial. They are nocturnal and, like many other small rodents, thus 
escape ordinary observation, most species coming out and wander- 
ing here and there through marsh, meadow, and herbaceous or 
even shrubby vegetation. Fairly well-worn runways are sometimes 
made along the edges of water or form general routes through dense 
vegetation, but these are not so well defined as those of the cotton 
rats (Sigmodon), which often share the same local habitat. The 
more aquatic species readily enter the water and swim and dive 
freely. 

The nests, made of plant fibers of many kinds, are placed in shallow 
burrows or sheltered places, commonly under massed vegetation on 
or near the ground, but the site chosen may be in a tangled clump of 
flags or marsh grasses standing in the water. From 3 to 7 young 
are produced at a birth, 4 or 5 being the usual number. In Ory- 
zomys palustris many young are brought forth in the months of 
April and May, but in this and other species their appearance at all 
seasons is ample evidence that there is no definite breeding season, 
except possibly near the extreme northern limit of the group. 

The rice rats feed extensively on green or succulent plants, food 
habits shared with the cottonrats (Sigmodon) and meadow mice ( Mi~ 
crotus). In early morning many freshly cut grass stems in the 
meadows evidence their nocturnal activity in favored places. While 
green food forms the normal ration of most species, drier foods, as 
seeds, also are eaten to some extent, and as with some other small 
rodents meat is relished at times. Specimens may be taken in meat- 
baited traps, and individuals thus caught are sometimes devoured 
by their own kind. Bachman, 1 who studied the habits of Oryzomys 
palustris, records that those kept in captivity " fed on grains of various 
kinds, but always gave the preference to small pieces of meat." In 
their natural habitat he observed them scratching up the recently 

1 Audubon, J. J., and Bachman, John, Quadr. North Amer., Ill, p. 215, 1854. 



1918.] INTRODUCTION. 5 

planted rice. He found that they begin feeding on rice when it is 
in the milky state and continue gathering the scattered grains in 
the fields during autumn and winter. He also observed them feed- 
ing on the large seeds of the gama grass ( Tripsacum dactyloides) , on 
those of the wild rye (Elymus virginicus) , and at certain seasons on 
those of the marsh grass (Spartina glabra). Bachman further states 
that the rice rat " sometimes retires to the shore for food, but has no 
disrelish to the small Crustacea and mollusks that remain on the 
mud at the subsiding of the tide." 

Species inhabiting forested areas usually become very abundant in 
clearings where a ground cover is allowed to grow. Rice rats appar- 
ently are not so injurious to crops as some other rodents, but like 
the cotton rats and meadow mice increase locally to excessive num- 
bers and then consume, in the aggregate, very large quantities of 
forage. Methods of poisoning that have proved effective in checking 
the ravages of meadow mice could probably be utilized with similar 
success in the control of rice rats. Owing to their nocturnal habits 
these mice are preyed upon by owls and doubtless by many carnivor- 
ous mammals. The rice rats and allied members of the great murine 
family to which they belong are the most numerous of American 
mammals, and their economic relations should be better known. 

GENERAL CHARACTERS. 

The rice rats as a whole present a wide range of variation in external 
appearance. Some of the larger, more robust species, including 
Oryzomys palustris, have not infrequently been mistaken for Old 
World rats of the genus Rattus, which have followed civilized man 
in his migrations and now not only universally infest his structures, 
but in favorable districts invade the fields in close competition with 
native rodents. Superficial resemblances are often striking, the 
bodily proportions and color and texture of pelage being very similar. 
On the other hand, many species are widely different from the Old 
World rats, in rich tawny coloration or character of pelage, and are 
more likely to be confused with allied American murine genera. 
The smallest North American species (subgenus Oligoryzomys) simu- 
late in color, delicate structure, and great length of tail some of the 
harvest mice (Beithrodontomys) ; another group (subgenus Melanomys) 
is more robust, the tail very short, and general proportions Akodon- 
like. 

In Oryzomys and allied genera superficial resemblances frequently 
mask the more essential features, and external characters, especially 
size and color, are less dependable than cranial modifications in tracing 
relationships of species. The group alignment, however, is often 
indicated by proportions of body and limbs; color and length of 
general pelage; length of vibrissa?; hairiness of ears, feet, and tail; 



6 NORTH AMERICAN FAUNA. [No. 43. 

presence or absence of prominent tufts of digital bristles; and form 
of claws. Aside from the general form and angularity of the skull, 
the principal characters of taxonomic value are the following: Size 
and form of incisors and molars; depth and arrangement of reentrant 
angles, form and position of enamel islands, and development of 
cusps in molar crowns ; length and form of anterior palatine foramina ; 
length of palatal bridge; position of lachrymals; form of maxillae, 
premaxillae, frontals, parietals, and interparietal; and size and form 
of audital bullae. 

PELAGE. 

The pelage is rather harsh in" texture throughout the genus, but 
varies greatly in length in the component groups. The overfur is 
longer, the vibrissas shorter, and the underfur denser and more woolly 
in semiaquatic species. Groups in which the pelage is normally short 
tend to develop longer fur in the forms which range at high altitudes : 
but several species, some with pelage much longer than others, may 
occur together at the same elevations. The ears are rather small and 
in typical Oryzomys and various groups, including the subgenus Oligo- 
ryzomys, the hairy eovering is moderately long, coarse, and not 
sharply differentiated from the body fur; in the 0. talamancse and 
other groups the ears are minutely pilose, in marked contrast with the 
general pelage. The claws on the toes of the hind feet in the semi- 
aquatic or more strictly terrestrial species are relatively long and 
straight, broad, and obtuse, and only partially overlapped by the 
digital bristles, while in the less aquatic or scansorial species they are 
short and recurved, compressed and sharp pointed, and prominent 
digital tufts project beyond the ends. In 0. palustris the vibrissas 
scarcely reach from the muzzle to the ears; in 0. pirrensis and 0. 
hombycinus they extend over the shoulders, while in various species 
they are intermediate in length. 



In coloration of upperparts the range of variation in the rice rats is 
from pale shades of buff or gray to rich tawny or russet, more or less 
mixed with black, especially over the median part of the dorsum. 
The underparts usually are dull white or buff, without a sharp line 
of demarcation along the sides, the plumbeous basal color showing 
through (a few species exhibiting basally white areas). The tail is not 
very sharply bicolor, but usually is brown above and lighter below, 
at least basally, the epidermis and scanty investing hairs of about the 
same tone. 

MOLT. 

,As breeding begins during adolescence and is continuous through- 
out the year, and as individuals arrive at maturity at all seasons, 
there is no very definite period for molting, although the more 



1918.] INTRODUCTION. 7 

northerly forms tend in winter to acquire a longer pelage, which be- 
comes abraded in summer. As a rule the new coat seems to replace 
the old almost imperceptibly, but adults in apparently fresh and 
in obviously worn pelages may often be seen together. 



The age of individuals is indicated approximately by the degree 
of wear on the molar crowns. The shearing of the slopes, beginning 
early in life, becomes distinctly noticeable as full growth is attained, 
and, progressing rapidly across the summits of the tubercles, in 
advanced age results in the obliteration of all trace of enamel 
arrangement. 

VARIATION. 

Variation in the rice rats is assignable to several categories, of 
which perhaps the most obvious are individual and geographic. 

INDIVIDUAL VARIATION. 

By individual variation is meant all the degrees of divergence 
from a typical mean exhibited by large series of conspecific skins 
and skulls from any given locality. The range of this variation, 
especially in general size, is extraordinary; in many species of corre- 
sponding age and sex it exceeds 10 per cent both in external and 
cranial dimensions. The typical mean, therefore, may be difficult 
to determine when a small series of examples exhibits preponder- 
ance toward either of the extremes; and conclusions based on the 
dimensions of a small number of individuals are likely to be mis- 
leading. While males average slightly larger than females, sexual 
differences in size appear to be negligible. Cranial variations in 
proportions and in the form of individual bones are noticeable, but 
usually within rather circumscribed limits. Some skulls are decid- 
edly broader and more massive than others of the same age and sex. 
Thickness of rostrum is usually, but not always, associated with 
breadth of frontal region and braincase. General expansion of the 
braincase commonly results in increased breadth across the posterior 
part of the frontals. The interparietal is variable in form, as are the 
parietals in the extent of encroachment of the lateral wings on the 
squamosals. The size of the molar teeth and of the audital bullae 
is fairly constant. 

Individual variation in color is much less than in size. Much of 
the variation in color observable is due to age or condition of pelage. 
The older adults tend at all seasons to exhibit more rufescent tones 
than the younger. A rusty reddish appearance is often due to much- 
worn pelage. There are no distinct color phases. 



NORTH AMERICAN FAUNA. [No. 43. 



GEOGRAPHIC VARIATION. 



Geographic variation, or the tendency of species to subdivide into 
regional or more or less localized forms, is very great in the genus 
Oryzomys. Of the numerous forms first described as distinct species 
a considerable number prove when better known to be geographic 
representatives of widely ranging specific types, presenting differ- 
ential characters associated in part with environmental conditions. 
The intergradation and subspecific position of many such forms can 
be demonstrated beyond any reasonable doubt in some instances, 
and in others may be safely assumed, in the light of knowledge of 
the essential characters of the /particular group. Since the distri- 
bution of the rice rats is mainly at low elevations, it is not surprising 
to find that species maintaining the same characters over extensive 
areas near sea level, where nearly uniform topographic and climatic 
conditions prevail, become locally modified on ascending to high 
altitudes. Thus, 0. couesi and 0. alfaroi are represented by widely 
ranging coastal forms and by more localized high-mountain races. 
That forms inhabiting mainly open, arid regions are paler than those 
inhabiting humid or heavily forested areas is well illustrated by the 
distribution of the races of 0. couesi in Mexico. The dark typical 
form occupies the relatively humid area near the Gulf of Mexico and 
the Caribbean Sea, while the pale subspecies, 0. c. mexicanus, 
pushes far northward along the arid Pacific coast, the point of diver- 
gence being near the Isthmus of Tehuantepec. Species reaching 
high altitudes tend to develop structural as well as color differences, 
while varying climatic conditions at low elevations are apt to result 
mainly in color modifications. Insular species are usually related to 
those inhabiting the adjacent mainland, but may exhibit very dis- 
tinctive characters, unless the islands are very near the coast. 

HISTORY AND MATERIAL. 

A species of Oryzomys was first described under the name Mus palus- 
tris from New Jersey, by Richard Harlan, 1 in 1S37. He compared the 
animal with the Norway rat, and owing to superficial resemblances 
regarded it as congeneric with the rats of the Old World. It was 
erroneously referred to the genus Arvicola in 1854 by Bachman, 2 who 
mentions having obtained specimens as early as the winter of 1816. 
According to Bachman, these specimens were described by him (but 
the description not published) in May, 1836. One was sent to the 
Academy of Natural Sciences of , Philadelphia for comparison with 
material there, and on the basis of this and an example in the Academy 
collection Dr. Harlan felt authorized to publish his Mus palustris. 

i Silliman's Amer. Journ. Sci., XXXI, p. 385, 1837. 

* Audubon, J. J., and Bachman, John, Quadr. North Amer., Ill, p. 214, 1854. 



1918.] INTRODUCTION". 9 

In renaming the species Arvieola oryzivora, Bachman assumed that 
Mus palustris Harlan was preoccupied by Arvieola palustris Harlan, 1 
an obvious error, as the two were not congeneric. 

In 1857 Baird, 2 recognizing distinctive characters, used Oryzomys 
as a full generic name for the group, with Mus palustris Harlan as 
type, apparently inadvertently, however, as on a later page of his 
publication 3 he accorded it only subgeneric value under Hesperomys 
of Waterhouse. In this course he was followed by Coues in 1877. 4 
Thirteen years later the same author 5 raised the name to generic 
rank. Meanwhile Hesperomys fulvescens Saussure 6 and Hesperomys 
couesi Alston 7 had been described, but their real generic position 
was not determined until later. New species were added at inter- 
vals, but of the relationships of the North American members of the 
genus little was known until 1901, when, as a result of study 
mainly of material which had accumulated in the collection of the 
Biological Survey, Merriam 8 published a synopsis of the forms 
inhabiting the United States and Mexico. Thirty-five species and 
subspecies were recognized by him, of which 20 were new. The 
species were divided into natural groups for the first time, and their 
salient characters pointed out. Short papers, largely descriptive of 
new species, by Thomas, 9 Allen, 10 Bangs, 11 Elliott, 12 and Goldman, 13 
have since appeared. The larger collections now available render it 
possible to determine the status of nearly all names, and the relation- 
ship, especially of the more austral species, to South American forms. 

The present revision is the result of a study of the rice rats in the 
Biological Survey, the Merriam, and other collections in the United 
States National Museum, now numbering 1,050 specimens, aug- 
mented by 563 from other American museums, 14 the assemblage 
including the types or topotypes of most of the species. The location 

1 Harlan, Richard, Fauna Americana, p. 13G, 1825. 

2 Baird, S. F., Mamm. North Amer., p. 459, 1857. 

* Op. cit., p. 482. 

* Coues, Elliott, Monogr. North Amer. Rodentia, p. 113, 1877. 
» Coues, Elliott, Century Diet., IV, p. 4165, 1890. 

« Saussure, H.de, Rev. et Mag. de Zool., ser. 2, XII, p. 102, March, 1860. 

i Alston, E. R., Proe. Zool. Soc, London, p. 756, 187G. 

8 Merriam, C. Hart., Proc. Washington Acad. ScL, III, pp. 273-295, July 26, 1901. 

» Thomas, O., Ann. Mag. Nat. Hist., ser. 7, VIII, pp. 251-253, Sept., 1901. 

io Allen, J. A., Bull. Amer. Mus. Nat. Hist., XXIV, pp. 654-657, Oct. 13, 1908 (including Oryzomys ochra- 
ceus [=Nectomys alfari],see Goldman, Proc. Biol. Soc, Washington, XXIX, p. 127, June 6, 1916); ibid., 
XXXIII, pp. 99-100, Apr. 30, 1910; ibid., XXXII, pp. 533-554, Nov. 17, 1913. 

U Bangs, O., Bull. Mus. Comp. Zool., XXXIX, pp. 33-36, Apr. 1902. 

12 Elliott, D. G., Field Columb. Mus. publ. 71, Zool. ser. Ill, p. 145, Feb., 1903; ibid., Ill, pp. 266-267, 
Mar. 1904. 

is Goldman, E. A., Smiths. Misc. Coll., LVI, no. 36, pp. 5-8, Feb. 19, 1912; ibid., LX., no. 22, pp. 5-6, 
Feb. 28, 1913; Proc. Biol. Soc. Washington, XXVIII, pp. 127-130, June 29, 1915. 

n For the use of material generously loaned and for other courtesies the author's thanks are due to Dr. J. A. 
Allen, American Museum of Natural History; to Mr. Samuel Henshaw and Mr. Outraru Bangs, Museum 
of Comparative Zoology; to Mr. W. H. Osgood, Field Museum of Natural History; to Mr. Witmer Stone, 
Academy of Natural Sciences of Philadelphia; to Mr. W. E. Clyde Todd and Mr. O. P. Murie, Carnegie 
Museum; and to Mr. CD. Bunker, Kansas University Museum. In addition he is indebted to Mr. Oldfield 
Thomas for critical notes and comparisons of specimens with types in the British Museum. 



10 NORTH AMERICAN FAUNA. [No. 43. 

of specimens examined in collections other than those in the United 
States National Museum is indicated by footnotes. 

EXPLANATIONS. 

MEASUREMENTS . 

All measurements of specimens are in millimeters. With a few 
exceptions, usually stated, the external measurements were taken in 
the flesh by the collector, as follows : Total length, nose to end of ter- 
minal tail vertebra; tail vertebrse, upper base of tail to end of terminal 
tail vertebra; hind foot, heel to end of longest claw. While adult 
males average slightly larger tfyan females, the difference is scarcely 
appreciable and in the small series usually available may be ignored. 
The external and cranial measurements given, therefore, are of series 
which may include specimens of both sexes. Of many species and 
subspecies so few nearly typical examples are available that the meas- 
urements given may not represent the normal range of individual va- 
riation, and too broad generalizations, therefore, should not be based 
upon them. The following cranial measurements were taken with a 
vernier caliper by the author : 

Greatest length. — Length from tip of nasals to supraoccipital in 
median line over foramen magnum. 

Zygomatic breadth. — Greatest distance across zygomata. 

Interorbital breadth. — Least distance between orbits. 

Width of braincase. — Distance between outer sides of squamosals 
at the slight constriction over auditor}^ meatus and immediately in 
front of lateral occipital crests. 

Nasals. — Greatest length of nasals. 

Anterior palatine foramina. — Greatest length of large palatal 
foramina. 

Palatal bridge. — Distance from excavated posterior border of palate 
to posterior end of either large palatal foramen. 

Upper molar series. — Greatest length of maxillary toothrow at 
alveolar border. 

COLORS. 

The names of colors used in descriptions are mainly those of 
Ridgway. 1 A few other modifying or comparative terms, however, 
have been employed, usually when some special difficulty was 
encountered in naming an indefinite hue or tone. 

TEETH. 

The nomenclature of the principal tooth elements used in the text 
is given in figure 1 . 2 

1 Ridgway, Robert, Color Standards and Color Nomenclature, 1912. 

2 For the homology and nomenclature of the molar crown divisions the writer is indebted to Messrs. 
Gerrit S. Miller, jr., and James W. Gidley, whose extensive researches, still in progress, in the phylogeny 
of the rodents, enable them to render an authoritative opinion. 



1918.] 



GENUS ORYZOMYS. 



11 



ZONAL DIVISIONS. 



Aside from the well-known extratropical North American zonal 
divisions, all references in the text under "Geographic distribution" 
are to less well-known tropical divisions, the Lower and Upper 
Tropical Zones, which may be roughly denned as follows: The Lower 
Tropical Zone, extending in tropical Middle America from sea level to 
elevations varying mainly in accordance with latitude and local 
topographic conditions. South of the twentieth parallel this zone 
reaches to about 3,000 or 3,500 feet altitude. Above these limits it 
is replaced by the Upper Tropical Zone, which extends to about 7,000 





pre Protoconule . 

pr Protocone. 

hy Hypocone. 



prcd Protoconulid. 

prd Protoconid. 

hyd Hypoconid. 



pad Paraconid. 

med Metaconid. 

mesd Mesostylid. 

end Entoconid. 

ensd Entostylid. 



pas Parastyle. 

sec. pas Secondary 

parastyle. 

pa Paracone. 

ms Mesostyle. 

me Metacone. 

mts Metast yle . 

Fig. 1.— Molar teeth of typical Oryzomys with outlines accentuated to show principal crown elements. 
A, Right upper molars; B, left lower molars. 

or 8,000 feet. Each of these zones is readily separable into sub- 
divisions on the basis of moisture, and are denominated, respectively, 
the Humid and Arid Lower Tropical and the Humid and Arid Upper 
Tropical Zones. 

GENUS ORYZOMYS Baird. 

Oryzomys Baird, Mamm. North Amer., pp. xlii, 458, 482, 1857 (subgenus of Hes- 
peromys Waterhouse). Type Mus palustris Harlan. 

Oryzomys Coues, Century Diet., IV, p. 4164, 1890. 

Oligoryzomys Bangs, Proc. New England Zool. Club, I, p. 94, Feb. 23, 1900 (sub- 
genus). Type Oryzomys navus Bangs. 

Melanomys Thomas, Ann. Mag. Nat. Hist., ser. 7, X, p. 248, Sept. 1902 (subgenus). 
Type Oryzomys phseopus Thomas. 

Melanomys Allen, Bull. Amer. Mus. Nat. Hist., XXXII, p. 533, Nov. 17, 1913 (genus). 

Geographic distribution. — Eastern and central United States from 
New Jersey, Kentucky, Illinois, Kansas, and Texas across Mexico 
to Lower California and south through South America to Cape Horn. 



12 NORTH AMERICAN FAUNA. [No. 43. 

Generic characters. — Form murine; pelage slightly hispid, but not 
bristly or spiny; tail usually long, but varying from about three- 
fourths length of head and body to one-fourth longer, the annulated 
scales distinctly visible through short, sparse investing hairs; ears 
varying from small to rather large, and finely to coarsely haired; 
soles of hind feet naked to heels, normally 6-tuberculate ; mamma? 8: 
pectoral, 2-2; inguinal, 2-2. 

Skull relatively thin and smoothly rounded; supraorbital and 
temporal ridges prominent or obsolescent; interparietal variable in 
size and contour ; zygomata slender, depressed to near level of molars ; 
antorbital foramen subcylindrical above, more or less abruptly con- 
stricted to a narrow slitlike opening below; outer wall of antorbital 
foramen rather broad, but varying in extent of anterior projection, 
the upper border rounded or slightly angular; palate reaching 
posteriorly beyond plane of last molars, the interpterygoid border 
concave or presenting a slight median projection; palatal pits 
present; audital bullas rather slightly inflated, tapering anteriorly 
and oblique in position. 

Molar crowns low (slightly higher in Melanomys than in the 
other subgenera) with prominent cusps, cones, or tubercles, and 
well-developed styles, the principal cusps arranged nearly opposite 
in two longitudinal series; upper molars 3-rooted; lower molars 
2-rooted. First upper molar with parastyle and paraconule strongly 
developed, partially divided near middle, and extended across 
internally for a distance equal to three-fourths or more of the greatest 
transverse diameter of tooth; secondary parastyle usually prominent; 
mesostyles and metastyles in all the upper molars present as small 
cusps or represented by enamel ridges connecting with principal 
inner cusps; inner cusps of first and second upper molars with oblique 
commissures ; third upper molar with posterior portions more or less 
obsolete, the hypocone varying in development and the metacone 
usually indicated only by a low abbreviated enamel fold. Lower 
molar cusp-arrangement similar to upper, but paraconid and para- 
conulid in anterior tooth faintly or not at all notched longitudinally; 
protoconids and hypoconids in first and second molars with oblique 
commissures as in the homologous elements in upper teeth; meso- 
stylids and entostylids joined by transverse enamel ridges with outer 
principal cusps; second lower molar with a moderately developed 
protoconulid; third lower molar with entoconid and entostylid com- 
pletely fused or obsolescent. Front of upper incisors without 
grooves. 

Remarks. — When the molars of Oryzomys are opposed the cusps 
in the upper jaw enter behind the corresponding cusps in the lower 
jaw. The parastyle is overlapped by the paraconid, the trenchant 



1918.] GENUS OKYZOMYS. 13 

anterior border of which shears on its anterior surface. The para- 
conule of the anterior molar and the protocones and hypocones of 
the other molars are broadly beveled or sheared internally, as are 
the protoconids and hypoconids. The paracones and metacones, on 
the other hand, are at first sheared posteriorly, while the metaconids 
and entoconids are sheared anteriorly, but progressive wear extend- 
ing across the summits tends to level the entire series uniformly. 

The genus presents complex relationships to various murine genera. 
Among its nearer relatives is Nectomys, which the more typical 
forms much resemble in external appearance as well as in the general 
form of the cranium. In dentition the two genera are also similar, 
but in typical Oryzomys, while the molar crowns as a whole are 
decidedly lower, the cusps or cones are higher, more conical, with 
summits more fully covered with enamel. In Nectomys, on the 
other hand, the low cusps tend to fuse with the stjdes and dentine is 
exposed at an earlier age. Oryzomys differs from Nectomys in other 
dental details, especially the development of the paraconulid of the 
second lower molar. The subgenus Melanomys, however, exhibits a 
somewhat intermediate condition; the molar crowns as a whole are 
somewhat higher than in typical Oryzomys, and lower than in Nec- 
tomys; the cones are high as in Oryzomys, but in sculpture, especially 
the early exposure of dentine, approach those of Nectomys. Another 
character suggesting gradation of Melanomys toward Nectomys is 
the position of the lachrymal, its attachment being mainly with the 
maxilla as in that genus, instead of about equally with maxilla and 
frontal as in typical Oryzomys. Agreement with Oryzomys is shown 
in the more essential dental details, including the marked develop- 
ment of the paraconulid in the second lower molar, an element 
absent in Nectomys. 

Comparison with various other genera reveals obvious resem- 
blances and points of difference in varying combination. Oryzomys 
agrees closely with Neacomys in cranial and dental characters, but 
the latter genus exhibits a departure in its grooved and bristly or 
spiny pelage. The generic name Nesoryzomys based on rice rats 
inhabiting the Galapagos Islands does not seem well founded, as the 
palate and dentition are Oryzomys-]ik.Q and in the short tail and 
reduced interorbital space, alleged generic characters, it is not widely 
different from some of the continental species of Oryzomys. 

In dentition Oryzomys is similar to RMpidomys, but the molar 
cusps are more crowded and the parastyle and protoconule are less 
distinctly separated by a median notch. The posterior part of the 
palate, however, presents features at variance in the two genera. 
Unlike that of Oryzomys the palate in Rhipidomys is excavated be- 
tween the posterior molars, and the palatal pits are separated by the 



14 NORTH AMERICAN FAUNA. [No. 43. 

interpterygoid fossa. The genus Nyctomys is allied to Rhipidomys, 
but exhibits a wider departure from Oryzomys. The molar crown 
arrangement is similar in all three genera, but in Nyctomys the cusps 
are more angular, the styles and commissures less developed, and the 
enamel covering maintained until later in life than in the others. 
In the posterior excavation of the palate Nyctomys more nearly 
resembles Rhipidomys; but, in the remarkable lateral development of 
the interparietal to a broad line of contact with the squamosals and 
consequent isolation of supraoccipital and parietals, Nyctomys differs 
notably from both Rhipidomys and Oryzomys. 

Some sections of the large genus Peromyscus are not very widely 
different from Oryzomys in deritition, but the principal cusps are 
obliquely placed, the protoconule much more developed, and the 
paracone and metacone shear mainly internally instead of poste- 
riorly as in Oryzomys. The general form of the skull in Peromyscus 
is distinctive; the palate ends near the posterior plane of the molars 
and the palatal pits present in Oryzomys are absent. 

In external appearance the genus Reithrodontomys bears a remark- 
able resemblance to the oryzomine subgenus Oligoryzomys; and in 
the Aporodon section of Reithrodontomys, in which the styles are 
present, an approach in molar pattern is manifested; but in Reith- 
rodontomys generic distinction is lodged in the grooved upper in- 
cisors. The genus Zygodontomys, until recently associated with 
Oryzomys, is similar in outward appearance and in general form of 
skull, but in height of molar crowns, absence of style ridges, and 
longitudinal instead of oblique commissures of cusps it departs 
widely from Oryzomys and more nearly approaches ATcodon. 

SUBGENERA AND MINOR GROUPS. 

Of the three subgenera into which the North American species of 
the genus Oryzomys are here divided, Oligoryzomys and Melanomys each 
contain a single group of closely related forms, while the subgenus 
Oryzomys is divisible into seven rather well-defined minor groups, or 
assemblages, of species or subspecies (excluding 0. victus, whose 
exact relationships are unknown). These assemblages are usually 
distinguishable by external characters, but recourse to cranial struc- 
ture is sometimes necessary in order to make accurate determina- 
tions. 

(1) The 0. palustris group is characterized by large, robust form, 
small, coarsely haired ears, short vibrissas, and the absence of con- 
spicuous tufts of silvery bristles, which in all the other groups pro- 
ject beyond the ends of the longer claws on the hind feet. The skull 
is broad with short rostrum, very long anterior palatine foramina 
(normally reaching anterior plane of first molars), and large audital 



1918.] GENUS ORYZOMYS. 15 

bullae. The principal reentrant angles normally reach less than half- 
way across the molar crowns. In the moderately worn crown of the 
second upper molar a crescentic central enamel island extends along 
the postero-internal base of the paracone. 

(2) The 0. melanotis group comprises slender, medium-sized, rich 
ochraceous-buffy or ochraceous-tawny species with large ears clothed 
externally with short, fine dusky hairs and internally with similarly 
short, fine rufescent hairs. The skull is narrow with elongated ros- 
trum, short anterior palatine foramina, and small audital bullae. 
The dentition departs slightly from the 0. palustris type ; the enamel 
arrangement is very similar but the reentrant angles extend farther 
across the molar crowns. 

(3) The 0. dlfaroi group includes small, dark-colored forms with 
short pelage, comparatively large, conspicuous ears clothed exter- 
nally and internally with short, fine blackish hairs. The skull is 
small and delicate in structure. The dentition is similar to that of 
the 0. melanotis group. 

(4) The 0. talamancse group bears much superficial resemblance 
to the 0. alfaroi group, but the members are usually brighter, more 
tawny in color. More distinctive characters are exhibited by the 
skull and teeth, especially the molar crown arrangement. In the 
grinding surface of the second upper molar the dentine ridge con- 
necting paracone and protocone, owing to more posterior position, 
eliminates the large central enamel island present in the 0. alfaroi 
group, and the crown, of the third lower molar is much more than 
half cleft by the outer reentrant angle (about half clef t in the 0. alfaroi 
group). 

(5) The 0. bombycinus group is easily recognized by very long 
pelage, that of the back measuring about 12 millimeters. The supra- 
orbital vibrissa? reach the remarkable length of 50 to 70 millimeters. 
The dentition is about as in the 0. talamancse, group. 

(6) The 0. devius group is distinguished by very large but rather 
slender form, relatively long tail (much longer than head and body), 
and dark general coloration. The dentition is similar to that of the 
0. talamanCse and 0. bombycinus groups. 

(7) The 0. tectus group may be known by large size, rich tawny 
coloration, small ears clothed with rather coarse hairs of general body 
color, and short, stout hind feet. The skull is broad, with short ros- 
trum and prominently projecting supraorbital ridges. The dentition 
is much as in the 0. talamancse group. 

In the present revision 44 species and subspecies are assigned to 
the typical subgenus Oryzomys, 5 forms are placed in the subgenus 
Oligoryzomys, and 2 in the subgenus Melanomys. 



16 NORTH AMERICAN FAUNA. [No. 43. 

List of North American Species and Subspecies, with Type Localities. 

Subgenus ORYZOMYS. 

Oryzomys palustris group: 

Oryzomys paluslris palustris (Harlan) . . " Fastland, ' : near Salem, New Jersey. 

palustris natator Chapman Gainesville, Florida. 

palustris coloratus Bangs Cape Sable, Florida. 

palustris texensis Allen Rockport, Texas. 

couesi couesi (Alston) Coban , Guatemala. 

couesi richmondi Merriam Escondido River, Nicaragua. 

couesi zygomaticus Merriam Nenton, Guatemala. 

couesi mexicanus Allen Hacienda San Marcos, Jalisco, Mexico. 

couesi aztecus Merriam Yautepec, Morelos, Mexico. 

couesi crinilus Merriam Tlalpam, Federal District, Mexico. 

couesi regillus Goldman /. . . Los Reyes, Michoacan, Mexico. 

couesi albiventer Merriam Ameca, Jalisco, Mexico. 

couesi peragrus Merriam Rio Verde, San Luis Potosi, Mexico. 

couesi aquaticus Allen Brownsville, Texas. 

fulgens Thomas " Mexico. ' ' 

gatuncnsis Goldman Gatun, Canal Zone, Panama. 

cozumelx Merriam Cozumel Island, Mexico. 

antillarum Thomas Jamaica. 

peninsulas Thomas Santa Anita, Lower California, Mexico. 

nelsoni Merriam Maria Madre Island, Mexico. 

Oryzomys melanotis group: 

Oryzomys melanotis melanotis Thomas. Mineral San Sebastian, Jalisco, Mexico. 

melanotis colimcnsis, nobis Armeria, Colima, Mexico. 

rostratus rostratus Merriam Metlaltoyuca, Puebla, Mexico. 

rostratus megadon Merriam Teapa, Tabasco, Mexico. 

rostratus yucatanensis Merriam Chichen Itza, Yucatan, Mexico. 

Oryzomys alfaroi group : 

Oryzomys alfaroi alfaroi (Allen) San Carlos, Costa Rica. 

alfaroi dariensis Goldman Cana, Panama. 

alfaroi angusticeps Merriam Volcan Santa Maria, Guatemala. 

alfaroi rhabdops Merriam Calel, Guatemala. 

alfaroi caudatus Merriam Comaltepec, Oaxaca, Mexico. 

alfaroi palatinus Merriam Teapa, Tabasco, Mexico. 

alfaroi saturatior Merriam Tumbala, Chiapas, Mexico. 

alfaroi cliapmani Thomas Jalapa, Vera Cruz, Mexico. 

alfaroi dilutior Merriam Huauchinango, Puebla, Mexico. 

guerrerensis Goldman Omilteme, Guerrero, Mexico. 

hylocetes Merriam Chicharras, Chiapas, Mexico. 

Oryzomys talamancss group: 

Oryzomys talamancee Allen Talamanca, Costa Rica. 

Oryzomys bombycinus group: 

Oryzomys bombycinus bombycinus 
Goldman Cerro Azul, Panama. 

bombycinus alleni Goldman Tuis, Costa Rica. 

Oryzomys devius group: 

Oryzomys devius Bangs Boquete, Panama. 

pirrensis Goldman Mount Pirre, Panama. 

Oryzomys tectus group : 

Oryzomys tectus tectus Thomas Bugaba, Panama. 

tectus frontalis Goldman Corozal, Canal Zone, Panama. 

Oryzomys victus 1 Thomas St. Vincent, Lesser Antilles. 

iNot examined and group association not determined. 



1918.] SUBGENUS ORYZOMYS. 17 

Subgenus OLIGORYZOMYS. 

Oryzomys fulvescens fulvescens (Saussure).. Orizaba, Vera Cruz. Mexico. 

fulvescens lenis Goldman Loa Reyes, Michoacan, Mexico. 

fulvescens mayensis, nobia Apazote, Campeche, Mexico. 

fulvescens costaricensis Allen El General, Coata Rica. 

fulvescens vegetus Bangs Boquete, Panama. 

Subgenus MELANOMYS. 

Oryzomys caliginosus idoneus Goldman Cerro Azul. Panama. 

caliginosus chrysomelas Allen Suerre, Costa Rica. 

Key to Subgenera. 

a 1 . Lachrymal articulating about equally with frontal and maxilla anteriorly; tail 
about equal to or longerthan head and body. 
b 1 . Second upper molar with central enamel island normally elongated or absent; 
supraorbital and temporal ridgea present; hind foot usually more than 

25 Oryzomys (p. 17). 

b 2 . Second upper molar with central enamel island normally circular; supraorbital 
and temporal ridgea absent; hind foot usually less than 25. 

Oligoryzomys (p. 87). 
a?. Lachrymal articulating almost entirely with maxilla anteriorly; tail about three- 
fourths length of head and body Melanomys (p. 94). 

Subgenus ORYZOMYS Baird. 

Subgeneric characters. — Color of upperparts usually contrasting 
strongly with that of underparts; 1 feet (epidermis and hairs), includ- 
ing toes, whitish, yellowish, or brownish; 1 tail about equal to or 
longer than head and body; 1 anterior border of lachrymal articulating 
about equally with maxilla and frontal; 1 supraorbital and temporal 
ridges usually prominent; 2 secondary parastyle well developed; 1 
slightly worn crown of second upper molar with central enamel island 
elongated or absent; 2 upper incisors decidedly curved backward near 
points. 1 

Key to Species of the Subgenus Oryzomys. 
[Typical adults.] 

a 1 . Habitat North America and Greater Antilles. 
b 1 . Hind foot without prominent tufts of digital bristles projecting beyond ends of 
three median claws. 
c 1 . Habitat Rio Grande Valley south to Panama, and including islands near 
coasts of Mexico and West Indies. 
d 1 . Habitat continental. 

e l . Supraorbital ridges not projecting prominently over orbita. 
f 1 . Head and shoulders not distinctly grayiah. 

g l . Upperparts richer ochraceous-tawny. ("Mexico."). 0.fulgens(p.41). 
g 2 . Upperparts duller ochraceous-tawny or ochraceous-buffy. (Southern 

Texas to Costa Rica. ) O. couesi (p. 28) . 

f 2 . Head and shoulders distinctly grayish. (Lower California.) 

O. peninsulse (p. 45). 

i Contrasting with Melanomys, » Contrasting with Oligoryzomys* 

14521°— 18— 2 



18 NORTH AMERICAN FAUNA. [No. 43. 

e 2 . Supraorbital ridges projecting prominently over orbits. (Panama.) 

O. gatunensis (p. 42). 
d 2 . Habitat insular. 
e 1 . Habitat islands off coasts of Mexico. 
P. TJpperparts ochraceous-buffy . (Maria Madre Island.). 0.nelsoni(p.46). 
P. Upperparts ochraceous-tawny. (Cozumel Island.). O. cozumelae (p. 43). 

e 2 . Habitat Greater Antilles. (Jamaica.)...^ O. antillarum (p. 44). 

c 2 . Habitat United States north of Rio Grande Valley O. palustris (p. 21). 

b 2 . Hind foot with prominent tufts of digital bristles projecting beyond ends of three 
median claws. 
c 1 . Ears clothed externally with fine blackish hairs contrasting with color of head; 
hind foot long and narrow. 
d 1 . Hind foot less than 33. 
e 1 . Supraorbital vibrissse less than 40. 

*. Ears clothed internally with fine blackish hairs. 

g l . Second upper molar with central enamel island present ; third lower 
molar with outer reentrant angle extending about half way across 
crown. 
ft 1 . Upperparts darker ochraceous-buff or ochraceous-tawny. 
i 1 . Zygomata not wider anteriorly than posteriorly, or if wider 
zygomatic breadth more than 14. (Atlantic slope in eastern 
and southern Mexico and south to Panama.) 

O. alfaroi (p. 58). 
i 2 . Zygomata wider anteriorly than posteriorly; zygomatic breadth 
14 or less. (Pacific slope of mountains of southern Chiapas.) 

O. hylocetes (p. 70). 
ft 2 . Upperparts paler ochraceous-buff or ochraceous-tawny. (Pacific 
slope of Sierra Madre in Guerrero and Oaxaca.) 

O. guerrerensis (p. 69). 

g 2 . Second upper molar with central enamel island absent; third lower 

molar with outer reentrant angle extending more than hah way 

across crown. (Costa Rica and Panama. ).0 . talamancse (p. 73). 

P. Ears clothed internally with buffy or rusty reddish hairs. 

g 1 . Size larger; hind foot 30 or more. (Atlantic coast of Mexico.) 

O. rostratus (p. 52). 
g 2 . Size smaller; hind foot less than 30. (Pacific coast of Mexico.) 

O. melanotis (p. 49). 

e 2 . Supraorbital vibrissse more than 50 O. bombycinus (p. 76). 

d 2 . Hind foot 33 or more. 
e 1 . Color paler; supraorbital ridges absent. (Costa Rica and western Panama.) 

O. devius (p. 80). 
e 2 . Color darker; supraorbital ridges present. (Eastern Panama.) 

O. pirrensis (p. 81). 
c 2 . Ears clothed externally with coarse tawny hairs not contrasting with color of 
head; hind foot short and broad. (Costa Rica and Panama.) 

O. tectus (p. 84). 
a 2 . Habitat Lesser Antilles. (St. Vincent.) O. victus (p. 86). 

Oryzomys palustris Group. 

Geographic distribution. — Coastal areas from southern New Jersey 
to southern Texas ; north in the Mississippi Valley to southern Illinois, 
and southward from the Rio Grande Valley on the east, and southern 



1918.] ORYZOMYS PALUSTRIS GROUP. 19 

Sinaloa on the west, through Middle America to Panama, with out- 
lying forms inhabiting southern Lower California, the Tres Marias 
Islands, and Cozumel Island. Confined mainly to the vicinity of 
water at low elevations, but ranging up to over 7,000 feet altitude in 
the marshy bottom of the Valley of Mexico (see maps, figs. 2 and 3). 

General characters. — Size large; form robust; tail usually equal to 
or longer than head and body (shorter in some examples), thinly but 
rather distinctly haired; ears small and inconspicuous, well haired 
internally as well as externally; general pelage long, rather coarse and 
rigid; the underfur somewhat woolly; vibrissas short, the longest 
arising from muzzle scarcely as long as head; hind feet broad, the 
upper surface rather well haired and under surface naked and coarsely 
granular anteriorly, becoming smooth along outer side of large pos- 
terior tubercle ; inner edge of plantar surface overlapped by fringing 
bristles ; toes of hind feet webbed near base ; the claws long, relatively 
straight, and projecting well beyond overlapping bristles. Color of 
upperparts presenting a wide range of variation from grizzled grayish 
brown, or pale buff, to rich ochraceous-buff or ochraceous-tawny, more 
or less heavily overlaid with black; underparts ranging from white to 
light ochraceous-buff. 

Slcull. — Size large, with rostrum short and braincase high and well 
arched ; outer wall of antorbital foramen with superior border extend- 
ing well forward, the foramen appearing as a deep circular notch as 
viewed from above; frontal region rather broad, the lateral margins 
trenchant, somewhat upturned, and projecting as supraorbital ridges, 
frontals usually encroaching in a narrow point posteriorly along the 
median line between the parietals; temporal ridges well developed 
anteriorly along parieto-squamosal borders, usually becoming indis- 
tinct posteriorly in crossing lateral wings of parietals to low lambdoid 
crest; interparietal small, sub triangular, the anterior border a nearly 
straight fine and the posterior with an ill-defined median angle; ante- 
rior palatine foramina narrow and much elongated, about equal in 
length to palatal bridge, normally reaching posteriorly to anterior 
plane of first molars, the median septum with posterior or maxillary 
portion contracted and anterior or premaxillary section expanded 
above; palatal pits large and normally oval in outline; interpterygoid 
fossa moderately broad; sphenopalatine vacuities large in 0. palus- 
tris, absent, or present as very narrow slits, in 0. couesi and related 
forms; audital bullae large, the swollen portion projecting anteriorly 
beyond anterior plane of basioccipital ; basioccipital narrow; angle of 
mandible rather broad and projecting posteriorly; coronoid process 
large, rising high above condyle; dentition moderately heavy; third 
lower molar rather short and broad; mandibular toothrow only 
slightly narrower posteriorly than anteriorly; inner reentrant angles 
in upper molars and outer reentrant angles in lower molars reaching 



20 NORTH AMERICAN FAUNA. [No. 43. 

less than half way across moderately worn crowns; second upper 
molar with a somewhat crescentic enamel island, or furrow, along 
postero-internal base of paracone, becoming restricted and finally 
obliterated through extended wear; tubercle over root of lower incisor 
prominent. 

Remarks. — The 0. palustris group includes 0. palustris of the United 
States, and 0. couesi and nearly related Middle American forms. 
Aside from the darker, more brownish colors which usually charac- 
terize 0. palustris, in contrast with the brighter, more ochraceous 
buffy or rufescent tones of 0. couesi and its relatives, these sections of 
the group also differ notably in cpality of pelage, the overfur being 
longer and projecting farther beyond the underfur in the former than 
in the latter. The forms of the two sections of the group agree closely 
in essential cranial details, but skulls of subspecies of 0. palustris are 
usually recognizable by the large size of the sphenopalatine vacuities. 
Members of the group as a whole are distinguished externally from 
those of other Middle American groups of the same subgenus Ly the 
small size and internal as well as external hairiness of the ears. 

Key to Species and Subspecies of the O. palustris Group. 

[Typical adults.] 

a 1 . Upperparts mainly grayish or brownish, or if ochraceous-buffy or ochraceous- 
tawny underparts not distinctly buffy. (United States north of Rio Grande 
Valley [0. palustris].) 
b 1 . Upperparts mainly grayish or brownish. 
c 1 . Color darker. (Atlantic coast region from southern New Jersey to northern 
Florida; Alabama; southeastern Mississippi; central Tennessee; southern 
Kentucky; southern Illinois, and parts of southeastern Missouri.) 

O. p. palustris (p. 22). 

c 2 . Color paler. (Coast region of Texas from Nueces Bay northward; Louisiana; 

western Mississippi; southern and eastern Arkansas; extreme southeastern 

Missouri; southeastern Kansas.) O. p. texensis (p. 27). 

b 2 . Upperparts mainly clay color or ochraceous-tawny. 

c 1 . Color duller, less distinctly ochraceous-tawny. (North-central Florida.) 

O. p. natator (p. 24). 
c 2 . Color brighter, more distinctly ochraceous-tawny. (Southern Florida.) 

O. p. coloratus (p. 26). 
a 2 . Upperparts mainly ochraceous-buffy or ochraceous-tawny. (Rio Grande Valley 
to Panama and islands near coasts of Mexico and West Indies [0. couesi and 
related forms].) 
b 1 . Habitat continental. 
c 1 . Supraorbital ridges not projecting prominently over orbits. 
d l . Head and shoulders not distinctly grayish. 

e 1 . Upperparts duller ochraceous-tawny or ochraceous-buffy. [0. couesi 
and subspecies.] 
f 1 . Underparts normally white. 

g 1 . Size larger; hind foot usually 35 or more; upper molar series usually 
more than 5. 



1918.] 



OKYZOMYS PALUSTBIS GROUP. 21 



h 1 . Upperparts more intense ochraceous-buff. (Northwestern Michoa- 

can) O. c. regillus(p. 37). 

h 2 . Upperparts less intense ochraceous-buff. (Central Jalisco.) 

O. c. albiventer (p. 38). 
g 2 . Size smaller; hind foot usually less than 35; upper molar series 
usually less than 5. 
h l . Upperparts darker ochraceous-buff. (Pacific coastal region from 
southern Sinaloa to southeastern Oaxaca.) 

O. c. mexicanus (p. 33). 
h 2 . Upperparts paler ochraceous-buff. (Morelos; southern Puebla; 
northern Oaxaca; northeastern Guerrero.). O. c. aztecus (p. 35). 
f 2 . Underparts normally buffy. 
g 1 . Size larger; hind foot usually more than 33; upper molar series 
usually more than 4.8. 
h 1 . Size smaller; hind foot averaging less than 35. 

i K Color darker. (San Luis Potosi.) O. c. peragrus (p. 39). 

i 2 . Color paler. (Rio Grande Valley.). . O. c. aquaticus (p. 39). 
h 2 . Size larger; hind foot averaging about 36. (Valley of Mexico.) 

O. c. crinitus (p. 36). 
g 2 . Size smaller; hind foot usually less than 33; upper molar series 
usually more than 4.8. 
h 1 . Upperparts normally ochraceous-buffy. 

i 1 . Color darker. (Northern Vera Cruz to northwestern Costa 

Rica.) O. c. couesi (p. 29). 

i 2 . Color paler. (Southwestern Gautemala and south-centra] 

Chiapas.) O. c. zygomaticus(p. 32). 

h 2 . Upperparts normally ochraceous-tawny. (Lowlands of eastern 

Nicaragua.) O. c. richmondi (p. 32). 

e 2 . Upperparts richer ochraceous-tawny. (Mexico.) O. fulgens(p. 41). 

cP. Head and shoulders distinctly grayish. (Lower California.) 

O. peninsulee (p. 45). 
c 2 . Supraorbital ridges projecting prominently over orbits. (Panama.) 

O. gatunensis (p. 42). 

b 2 . Habitat insular. 

c 1 . Habitat off east coast of Mexico. (Cozumel Island.). . O. cozumelse (p. 43). 

c 2 . Habitat off west coast of Mexico. (Maria Madre Island.). O. nelsoni (p. 46). 

c 3 . Habitat West Indies. (Jamaica.) O. antillarum (p. 44). 

ORYZOMYS PALUSTRIS (Harlan). 
[Synonymy under subspecies.] 

Geographic distribution. — Atlantic and Gulf coastal areas from 
southern New Jersey (not yet recorded from Delaware and Maryland, 
but doubtless occurs there), to southern Texas, and north through the 
Mississippi Valley to southern Kentucky, southern Illinois, and 
eastern Kansas (fig. 2). Altitudinal range from sea level up along 
streams to about 500 feet altitude (rarely to 1,000 feet), mainly in the 
Lower Austral Zone, but reaching into the Upper Austral Zone in the 
more northerly localities, and into the Tropical Zone in southern 
Florida. 

General characters. — Similar in general to O. couesi, but pelage 
longer; colors usually darker and duller grayish brown instead of 



22 



NORTH AMERICAN FAUNA. 



[No. 43. 



ochraceous-buffy or ochraceous-tawny; skull differing in various 
details, but dentition about the same. 

Color. — Upperparts in general varying from grizzled grayish brown 
or pale buff, to tawny-olive, clay color, and ochraceous-tawny; the 
face, top of head, and back heavily lined or overlaid with black; 
sides paler, owing to a thinner admixture of blackish hairs; under- 
pays white, varying to buffy white and rarely to pale buff; outer 
sides of ears dusky, the inner sides clothed with grayish or rusty hairs; 
feet whitish ; tail varying from brownish above and whitish below to 
dusky all round. Young (in first pelage): Varying from grayish 

brown to dull tawny- 
olive above, whitish 
below. 

STcull. — (For gen- 
eral outlines see under 
0. palustris group.) 
In general form closely 
resembling that of 0. 
couesi, but sphenopa- 
latine vacuities, ab- 
sent or much reduced 
in size in that species, 
large and widely open; 
antorbital foramen 
with anterior border 
less rounded above, 
less inclined or pro- 
duced forward at the 
base, in many exam- 
ples somewhat excised 
or tending to develop 
a point as in Sigmodon. 
Remarks. — 0. palustris is divisible into four geographic races which 
form a closely intergrading series. The species apparently attains 
its largest size in Florida and along the coast of the Gulf of Mexico east 
of the Mississippi delta. 

ORYZOMYS PALUSTRIS PALUSTRIS (Harlan). 

Swamp Rice Rat. 

[PI. I, figs. 1, la; PI. V, figs, 1, 4; PI. VI, figs. 1, la.] 

Mus palustris Harlan, Silliman's Amer. Journ. Sci., XXXI, p. 38!'), 1837. 

Arvicola oryzivora, Bachman, in Audubon and Bachman, Quadr. North Amer., Ill, 

p. 214, 1854. Type from St. Johns Parish, South Carolina. 
IJ[esperomys] palustris Wagner, in Suppl. Schreber's Saugthiere, III, p. 543, 1843. 
Oryzomys palustris Baird, Mamm. North Amer., p. 459, 1857. 




1. O. p. palustris. 2. O.p.natator. 3. O.p.coloratus. 4. O.p.texensis- 
Fig. 2.— Geographic distribution of subspecies of Oryzomys palustris. 



1918.] ORYZOMYS PALUSTEIS GROUP. 23 

Type locality. — "Fastland/' near Salem, Salem County, New Jersey. 

Type. — Not known to exist. 

Geographic distribution. — Atlantic coastal areas from southern New 
Jersey (not yet known from Delaware or Maryland, but doubtless 
occurs there) south to northeastern Florida, thence westward through 
southern Georgia to the Gulf coast of Alabama and Mississippi, and 
north through Alabama and western Tennessee to southwestern Ken- 
tucky, southern Illinois, and parts of southeastern Missouri. Alti- 
tudinal range from sea level up along streams to about 500 feet (rarely 
to 1,000 feet), mainly in Lower Austral Zone, but reaching into Upper 
Austral Zone in southern New Jersey, southeastern Kentucky, and 
southeastern Missouri (Marble Hill). 

General characters. — Size usually smaller and color more brownish, 
less tawny, than in 0. p. natator and 0. p. coloratus; skull less massive. 
Closely resembling 0. p. texensis, but darker, more brownish than 
topotypes of the latter form; skull broader. 

Color. — Fresh pelage (December): Upperparts grizzled grayish 
brown or pale buff, the brownish or buffy tone most intense on rump, 
darkened on face, top of head, and back by overlying blackish hairs; 
feet whitish; tail brownish above, whitish below, becoming in some 
specimens dark all round near tip. Young (in first pelage) : Grayish 
brown above, dull whitish below. 

Skull. — (For general outlines see under O. palustris group.) Very 
similar to those of 0. p. natator and O. p. coloratus, but narrower; 
braincase decidedly narrower; zygomata less widely spreading; 
frontal region narrower, the supraorbital borders less projecting. 
Compared with that of 0. p. texensis the skull differs mainly in some- 
what larger average size; zygomata usually more widely spreading. 

Measurements. — Average of four adults from Greenwich, N. J. (near 
type locality): Total length, 242 (237-245); tail vertebrae, 112 (109- 
116); hind foot, 31 (30-31.5). An adult from Pope Creek, Va.: 260; 
130; 33. Average of four adults from Georgetown, S. C: 257 (233- 
273); 125 (113-132); 32.5 (31-33). Average of three adults from 
Bon Secour, Ala.: 265 (250-280); 131 (125-138); 30.5 (30-31.5). 
Adult from Bayou La Batre, La.: 233; 116; 30. Adult from Marble 
Hill, Mo.: 252; 115; 29. Adult from Olive Branch, 111.: 255; 113 
29. Adult from Barbourville, Ky.: 270; 133; 30.5. Skull (two 
adults from Greenwich, N. J.: Greatest length, 32.4, 31; zygomatic 
breadth, 17.2, 16.6; interorbital breadth, 5.2, 5.1; width of braincase, 
11.9, 12; nasals, 12.7, 12; anterior palatine foramina, 6.7, 6.7; 
palatal bridge, 6.2, 6.2; upper molar series, 4.5, 4.6. 

Remarks. — 0. p. palustris passes into 0. p. texensis in the Mississippi 
Valley. Specimens from Marble Hill, Mo., and Olive Branch, 111., 
however, seem referable to the typical form. A tendency to develop 



24 NORTH AMERICAN FAUNA. [No. 43. 

the ruddy color and more massive skull of 0. p. natator is exhibited by 
specimens from New Berlin and Burnside Beach, northern Florida, 
and intergradation of the two forms seems evident in that part of the 
State. No rice rats have been recorded from Delaware or Maryland, 
but the favorable character of the country and the narrowness of the 
gap between collecting stations to the north and south point to 
probable continuity of range. Specimens from South Carolina 
apparently representing Arvicola oryzivora of Audubon and Bachman 
are inseparable from typical palustris. 

Specimens examined. — Total number, 233, as follows: 

Alabama: Autaugaville, 1; Bayoi/La Batre, 2; Bon Secour, 3; Elmore, 1; Flor- 
ence, 1; Gallion, 1; Hayneville, 2; Huntsville, 4; Jackson, 2; Mobile, 1; Mont- 
gomery, 1; Mount Weogufka, 1; Reform, 1; Sand Mountain (near Carpenter), 
1; Seale, 1. 

Florida: Burnside Beach, 9; 1 New Berlin, 8. 2 

Georgia: Cumberland Island, 14; 3 > i Hursman Lake, l; 1 Mcintosh County, 1; 
Okefinokee Swamp, l; 2 Ossabaw Island, 12; 1 Riceboro, 7; 5 Saint Marys, 20; 3 > 6 
Savannah, 10; Toccoa, 2. 

Illinois: Olive Branch, 3. 

Kentucky: Barbourville, 3. 

Missouri: Marble Hill, 1. 

Mississippi: Biloxi, 1. 

New Jersey: Cedar Creek, l; 7 Greenwich, 13. 8 

North Carolina: Coinjock, 1; Pea Island, 2; 1 Raleigh, 31. 9 » 10 

South Carolina: Beaufort County, 2; Calhoun Falls, 2; 2 Easley, 1; Frogmore, 
l; 3 Georgetown, 11; Plantersville, 7; Saint Helena Island, 1; Society Hill, 3. 

Tennessee: High Cliff, 1; Lawrenceburg, 2. 

Virginia: Dismal Swamp, 20; Pope Creek (5 miles southeast of Colonial 
Beach), 2; Smith Island, 6; Suffolk, 1; Wallops Island, 3; Warsaw (4 miles 
southwest), 5; Wreck Island, 1. 

ORYZOMYS PALUSTRIS NATATOR Chapman. 

Central Florida Rice Rat. 

Oryzomys palustris natator Chapman, Bull. Amer. Mus. Nat. Hist., V., p. 44, March 
17,' 1893. 

Type locality. — Gainesville, Alachua County, Florida. 

Type. — No. }%%%, <$ adult, American Museum of Natural History; 
collected by F. M. Chapman, January 31, 1889. 

Geographic distribution. — Central Florida, north of Everglades; 
Austro riparian division of Lower Austral Zone. 

1 Collection Mus. Comp. Zool. 7 Collection Acad. Nat. Sci. Philadelphia. 

2 Collection Field Mus. Nat. Hist. 8 Five in collection Acad. Nat. Sci. Philadelphia. 
» Collection Amer. Mus. Nat. Hist. 9 Five in collection Amer. Mus. Nat. Hist. 

< Eleven in collection Mus. Comp. Zool. 10 Two in collection Field Mus. Nat. Hist.; 6 in 

6 Five in collection Field Mus. Nat. Hist. Mus. Comp. Zool. 

6 Three in collection Field Mus. Nat. Hist.; 15 in 
Mus. Comp. Zool. 



1918.] ORYZOMYS PALUSTEIS GROUP. 25 

General characters. — Most like 0. p. coloratus; differing usually in 
less intense tawny suffusion of upperparts, especially cheeks and sides 
of body. Size larger, color more tawny, and skull more massive than 
usual in 0. p. palustris or 0. p. texensis. 

Color. — Fresh pelage (December) : Upperparts varying from grizzled 
grayish brown or pale buff to tawny-olive, clay color, or ochraceous- 
tawny, deepest and richest on lower part of back and rump, becoming 
paler and more buffy on sides, and darkened dorsally by admixture 
of blackish hairs ; face grayish or pale buffy beneath overlying dusky 
hairs; underparts usually white, but in some specimens more or less 
suffused with pale buff; feet white; tail brownish above, whitish be- 
low, becoming in some specimens dark all round toward tip. Young 
(in first pelage) : Brownish or dull tawny-olive mixed with black above, 
whitish below. 

STcull. — Similar to those of 0. p. palustris and 0. p. texensis, but 
broader; braincase decidedly broader; zygomata more widely spread- 
ing; frontal region broader, the supraorbital borders more projecting. 
Comparison with the skull of 0. p. coloratus reveals no appreciable 
difference. 

Measurements. — Type: Total length, 295; tail vertebrae, 143; hind 
foot (dry skin), 33. Average of 10 adults (type and 9 topotypes): 
276.7 (271-300); 142.9 (132-156); 34.3 (32.5-37). SJcull (average of 
6 adults, type and 5 topotypes): Greatest length, 32.4 (31.4-33.7); 
zygomatic breadth, 17.2 (16.4-17.7); interorbital breadth, 5.4 (5.1- 
6.1); width of braincase, 12.4 (12-12.8); nasals,- 12.7 (12-13.5); an- 
terior palatine foramina, 6.9 (6.4-7.2); palatal bridge, 6.1 (5.7-6.5); 
upper molar series, 4.7 (4.5-4.9). 

Remarks. — In northern Florida, not far to the northward of the 
type locality, 0. p. natator passes into 0. p. palustris, as shown by 
specimens from New Berlin and Burnside, which, however, seem more 
properly placed with the latter form. In the vicinity of Lake Oke- 
chobee natator merges with 0. p. coloratus, a richer colored form 
inhabiting the southern part of the State. 

Specimens examined. — Total number, 121, as follows: 

Florida: Anastasia Island, 2; 1 Canaveral, 4; Cape Canaveral, 3; Cartersville, 1; 
Crystal River, l; 1 Enterprise, 26 ; 2 Espanita, 3; 3 Fort Kissimmee, 1; Gaines- 
ville, 19 (type and topotypes); 4 Geneva, 1; Kissimmee, 2; Kissimmee River, 
2; Lake Harney, 11; Lake Kissimmee, 19; Micco, 9; 5 Mullet Lake, 1; Oak 
Lodge, 9; 1 Ocala, 2; Tarpon Springs, l; 6 Titusville, 4. 

i Collection Mus. Comp. Zool. 

2 Fourteen in collection Amer. Mus. Nat. Hist.; 10 in Field Mus. Nat. Hist.; 2 in Mus. Comp. Zool. 

s Collection Field Mus. Nat. Hist. 

* Nine in collection Amer. Mus. Nat. Hist.; 5 in Field Mus. Nat. Hist.; 5 in Mus. Comp. Zool. 

6 Three in collection Amer. Mus. Nat. Hist.; 3 in Field Mus. Nat. Hist. 

« Collection Acad. Nat. Sci. Philadelphia. 



26 NORTH AMERICAN FAUNA. [No. 43. 

ORYZOMYS PALUSTRIS COLORATUS Bangs. 

Everglades Rice Rat. 

(PI. I, figs. 2, 2a.) 

Oryzomys palustris coloratus Bangs, Proc. Boston Soe. Nat. Hist., XXVIII, p. 189, 
March, 1898. 

Oryzomys natator floridanus Merriam, Proc. Washington Acad. Sci., Ill, p. 277, July 
26, 1901. Type from Everglade, Florida, No. 71349, $ ad., U. S. Nat. Mus. (Bio- 
logical Survey collection); collected by J. Alden Loring, March 29, 1895. 

Type locality. — Cape Sable, Monroe County, Florida. 

Type. — No. 4470, <? adult, Museum of Comparative Zoology (col- 
lection of E. A. and O. Bangs) ; collected by C. L. Brownell, April 17, 
1895. 

Geographic distribution. — Tropical southern Florida, north to Lake 
Okechobee. 

General characters. — Closely resembling 0. p. natator; differing in 
more intense tawny suffusion of upperparts; size about the same. 
Size larger than usual in 0. p. palustris and 0. p. texensis, and color 
much more tawny than either. 

Color. — About as in 0. p. natator, but general tone of upperparts 
slightly richer, more tawny or rufescent. 

STcull. — Like that of 0. p. natator. 

Measurements. — Type: Total length, 301; tail vertebrae, 150; hind 
foot, 35. Average of three adult topotypes: 296 (278-305); 144 
(133-152); 33.4 (33.4-33.4). STcull (average of four adults, type 
and three topotypes): Greatest length, 32.2 (31.8-32.7); zygomatic 
breadth, 17.1 (16.8-17.5); interorbital breadth, 5.8 (5.8-5.9); width 
of braincase, 12.4 (12.2-12.5); nasals, 12.4 (12.1-12.6); anterior 
palatine foramina, 6.6 (6.3-7); palatal bridge, 6.1 (5.7-6.6); upper 
molar series, 4.8 (4.7-4.9). 

Remarks. — O. p. coloratus requires close comparison with 0. p. 
natator, from which it apparently differs only in color. The richer 
tone in coloratus is most noticeable when specimens are turned on 
their sides, and the cheeks and flanks contrasted with those of exam- 
ples of natator. As Bangs rightly states, coloratus "occupies only 
the southern, tropical part of the Florida peninsula." 

In describing "Oryzomys natator fioridanus,^ Merriam overlooked 
the name coloratus, which had already been applied to the animal 
of the region; the two are clearly synonymous. 

Specimens examined. — Total number, 50, as follows: 

Florida: Cape Sable, 11 (type and topotypes); 1 Eden, 1; Everglade, 16 (includ- 
ing type of "floridanus"); Flamingo, 13 ; 2 Juno (Lake Worth), 5; Jupiter^; 1 
Miami, 1; Miami River, 1. 

i Collection Mus. Comp. Zool. 2 Twelve in collection Mus. Comp. Zool.; 1 in Amer. Mus. Nat. Hist. 



1918.] ORYZOMYS PALUSTRIS GROUP. 27 

ORYZOMYS PALUSTRIS TEXENSIS Allen. 

Texas Rice Rat. 

Oryzomys palustris texensis Allen, Bull. Amer. Mus. Nat. Hist., VI, p. 177, May 31, 
1894. 

Type locality. — Rockport, Aransas County, Texas. 

Type. — No. ^-ftf, c? ad., American Museum of Natural History; 
collected by H. P. Attwater, November 15, 1893. 

Geographic distribution. — From Corpus Christi Bay north and east 
along the Gulf coast of Texas and Louisiana to the delta of the Mis- 
sissippi, thence north in the Mississippi Valley to southeastern Mis- 
souri; general range reaching southeastern Kansas, probably by way 
of the Arkansas River valley through Oklahoma (not yet known 
from Oklahoma) ; altitudinal range in Austroriparian Zone, mainly 
below 500 feet, but extending up to about 1,000 feet in Kansas. 

General diameters. — Closely resembling 0. p. palustris, but typical 
examples paler; skull usually narrower. Color paler, less rufescent, 
and skull decidedly narrower, less massive, than in O. p. natator and 
O. p. color atus. 

Color. — About as in 0. p. palustris, but averaging slightly paler. 
An ochraceous-tawny suffusion of upperparts and underparts is 
shown in rare examples. 

Skull. — About like that of O. p. palustris, but averaging somewhat 
smaller, with less widely spreading zygomata. Similar to that of 
0. p. natator, but narrower; braincase decidedly narrower; zygomata 
less widely spreading; frontal region narrower, the supraorbital 
borders less projecting. 

Measurements. — Type: Total length, 256 ; tail vertebrae, 139; hind 
foot, 30.5. Average of eight adult topotypes: 242 (226-279); 120 
(108-133); 29 (28.5-30.5). Skull (average of 5 adults— type and 4 
topotypes): Greatest length, 31 (30-32.1); zygomatic breadth, 16 
(15.2-16.8); interorbital breadth, 5.3 (5.2-5.4); width of braincase, 
11.8 (11.1-12.3); nasals, 12.5 (12-12.9); anterior palatine foramina, 
6.1 (5.7-6.5); palatal bridge, 5.8 (5.5-6.1); upper molar series, 4.4 
(4.3-4.7). 

Remarks. — Specimens from the type locality are paler than those 
from other localities in the immediate vicinity, some of which are 
practically indistinguishable from many typical examples of 0. p. 
palustris. Moreover, in cranial characters, especially in width of 
braincase and outward spread of zygomata (characters winch dis- 
tinguish 0. p. texensis from 0. p. natator and 0. p. coloratus), texensis 
very closely approaches palustris. The skull of the type and larger 
topotypes of texensis seem inseparable from some of the smaller skulls 
of comparable age from the region of the type locality of palustris. 
The cranial difference noted, however, affects the majority of indi- 



28 



NORTH AMERICAN FAUNA. 



[No. 43. 



viduals, and together with a tendency toward pallid coloration 
exhibited by animals inhabiting a wide area, seems to entitle texensis 
to recognition as a separate form. The few specimens available from 
Arkansas, western Tennessee, and extreme southeastern Missouri 
seem referable to texensis, but approach palustris so closely that they 
might with nearly equal propriety be assigned to that subspecies. 
Specimens examined. — Total number, 110, as follows: 

Lake City, 1; Wilmot, 1. 



3; Lake Catherine, 



Arkansas: Camden, 2; Delight, 1; 

Kansas: Neosho Falls, 2. 1 

Louisiana: Burbridge, IS; 2 Gibson, 4; 2 Houma, 1; Iowa, 
6; 3 Main Pass, 2; Mermenton, 4; 4 New Orleans, 2. 

Mississippi: Fayette, 1. / 

Missouri: Kennett, 2; Portageville, 1. 

Tennessee: Arlington, 1. 

Texas: Corpus Christi, 7; 5 Matagorda, 7; Matagorda Island, 2; Matagorda Penin- 
sula, 1; Nueces Bay, 5; Padre Island, 4; Port Lavaca, 4; Rockport, 24 (type 
and topotypes); 6 Sabine, 1; Victoria, 1; Virginia Point, 1 ; Wharton County , l. 3 




Fig. 3. — Geographic distribution of Oryzomys couesi and related species. 

ORYZOMYS COUESI (Alston). 

[Synonymy under subspecies.] 

Geographic distribution. — River valleys and marshy areas from 
southern Texas on the east and southern Sinaloa on the west, south 
through central and southern Mexico, Guatemala, Honduras, and 
Nicaragua to northern Costa Rica. Altitudinal range from sea level, 
regardless of latitude, to over 7,000 feet in the Valley of Mexico; 
zonal range mainly Tropical but reaching into Sonoran Zone on the 
tableland of Mexico (fig. 3). 



1 One in Mus. Comp. Zool. 
* Collection Mus. Comp. Zool. 
3 Collection Amer. Mus. Nat. Hist. 
« Collection Field Mus. Nat. Hist. 



' One in collection Amer. Mus. Nat. Hist. 
6 Nineteen in collection Amer. Mus. Nat. Hist.; 2 
in Field Mus. Nat. Hist.; 1 in Mus. Comp. Zool. 



1918.] ORYZOMYS PALUSTRIS GROUP. 29 

General characters. — Similar in general to 0. palustris, but pelage 
shorter; colors usually brighter and richer, ochraceous-buffy or 
ochraceous-tawny instead of grayish brown ; skull differing in various 
details, but dentition about the same. 

Color. — Upperparts varying from light buff or pinkish buff through 
ochraceous-buff to ochraceous-tawny, more or less suffused with 
tawny or russet, the general shade paler on cheeks, shoulders, and 
along sides, becoming deeper, more intense, and darkened on face, 
top of head, and back by admixture of black hairs; underparts 
varying from nearly pure white through light buff to light ochraceous- 
buff, more or less suffused with pale salmon color; outer sides of ears 
blackish, inner sides moderately clothed with short hairs varying 
from grayish to ochraceous-buff or rusty reddish; feet wnite; tail 
dark brownish above, whitish or light brownish below. 

Shutt. — (For general outlines see under 0. palustris group.) In 
general form closely resembting that of 0. palustris, but sphenopala- 
tine vacuities, large in that species, absent or much reduced in size; 
antorbital foramen with anterior border more rounded above, more 
inclined or produced forward at the base, not excised or tending to 
develop the somewhat Sigmodon-Yike point often present in palustris; 
interparietal with a more evident posterior angle. 

Remarks. — Ten geographic races of 0. couesi appear to be recog- 
nizable in the area between the valley of the Bio Grande, Texas, and 
northern Costa Rica. While all the more minute steps of inter- 
gradation are not always shown by the material now available, 
mainland forms throughout this wide interval agree so closely in all 
essential details that they seem safely assignable to a single species- 
Several outlying insular forms (and 0. peninsulse of Lower California) 
exhibit more distinctive characters and are accorded specific rank, 
but they are clearly related to the widely ranging 0. couesi section of 
the 0. palustris group. 

ORYZOMYS COUESI COUESI (Alston). 
Coues Rice Rat. 

(PI. I, figs. 3, 3a.) 

Hesperomys couesi Alston, Proc. Zool. Soc. London, 1876, p. 756. 

Oryzomys couesi Thomas, Ann. Mag. Nat. Hist., ser. 6, XI, p. 403, May 1893 (type 

and locality fixed). 
Oryzomys jalaps Allen and Chapman, Bull. Amer. Mus. Nat. Hist., IX, p. 206, June 

16, 1897. Type from Jalapa, Vera Cruz, Mexico (altitude 4,400 ft.). No. fHiHr, S 

ad., Amer. Mus. Nat. Hist.; collected by F. M. Chapman, April 16, 1897. 
Oryzomys jalapx rufinus Merriam, Proc. Washington Acad. Sci., Ill, p. 285, July 26, 

1901. Type from Catemaco, Vera Cruz, Mexico (altitude 1,000 feet). No. 65499, 

$ ad., U. S. Nat. Mus. (Biological Survey collection); collected by Nelson and 

Goldman, April 27, 1894. 
Oryzomys teapensis Merriam, Proc. Washington Acad. Sci., Ill, p. 286, July 26, 1901. 

Type from Teapa, Tabasco, Mexico. No. 99973, <? subad., U. S. Nat. Mus. 

(Biological Survey collection); collected by Nelson and Goldman, April 4. 1900, 






30 NORTH AMERICAN FAUNA. [No. 43. 

Oryzomys goldmani Merriam. Proc. Washington Acad. Sci., Ill, p. 288, July 26, 1901. 

Type from Coatzacoalcos, Vera Cruz, Mexico (near sea level). No. 78110, £ ad., 

U. S. Xat. Mus. (^Biological Survey collection^; collected by Nelson and Goldman, 

April 11, 1S96. 
Oryzomys jalap- apatdius Elliot. Field Columb. Mus.. publ. 90. zool. ser., Ill, p. 266, 

March 8. 1904. Type from San Carlos. Vera Cruz, Mexico. No. 13107, $ ad., 

Field Mus. Nat. Hist.; collected by X. G. Buxton, March 1, 1903. 
Oryzomys richarc 's>" i Mien. Bull. Arner. Mus. Nat. Hist., XXVIII, p. 99, April 30, 

1910. Type from Pena Blanca. Xicaragua. No. 29800, <? ad., Amcr. Mus. Xat. 

Hist.; collected by Wm. B. Richardson, May 25, 1909. 

Type locality. — Coban. Guatemala. 

Type. — In British Museum: collected by Osbert Salvin. 

Geographic distribution. — From northern Vera Cruz southeastward 
through eastern Puebla, eastern Oaxaca, northern and extreme 
southern Chiapas, Tabasco, Campeche, Yucatan, Quintana Roo, 
Guatemala, Honduras, and Nicaragua, to northwestern Costa Rica; 
altitudinal range from sea level to about 5,000 feet mainly in Humid 
Lower Tropical Zone. 

General characters. — Size about as in 0. c. zygomaticus, O. c. mexi- 
canus and 0. c. richmondi; color slightly darker than in zygomuticus, 
decidedly darker than in mexicanus, and decidedly paler than in 
ri-chmondi; skull about like those of mexicanus and richmondi; 
sphenopalatine vacuities absent or represented by very narrow slits 
as usual in the 0. couesi section of the O. palustris group. Similar hi 
general to O. c. peragrus, but somewhat smaller, with upperparts, 
especially cheeks, shoulders, and sides, more ochraceous-buffy: skull 
less massive. 

Color. — Fresh pelage: Upperparts varying from ochraceous-buffy 
to ochraceous-tawny, deepened in rare examples to light cinnamon- 
brown, lightest on cheeks, shoulders, and along sides, the face, top 
of head, and back much darkened by black hairs; underparts varying 
from light buff to light ochraceous buff (rarely dull white); outer 
sides of ears blackish, the inner sides clothed with short ochraceous- 
buffy hairs; feet white; tail brownish above, dull yellowish below 
proximally, becoming light brownish toward tip. 

STcuU. — About as in 0. c. zygomaticus, 0. c. mexicanus, and O. c. 
richmondi; differing mainly in smaller general size, decidedly nar- 
rower braincase, and smaller molars than those of 0. c. crinitus, O. e. 
albivtnter, and other Mexican tableland forms. Similar to that of 
0. c. peragrus, but braincase narrower. 

Measurements. — Average of 4 adults from Tumbala, Chiapas: 
Total length, 252 (242-265) ; tail vertebra?, 130 (127-135) ; hind foot, 
30.7 (30-31). Average of 10 adults from Yaruca, Honduras: 267.5 
(255-2S0); 138(130-145); 29.1 (2S-32). Average of seven adults 
from Orizaba, Vera Cruz: 263 (248-294); 14S (139-174); 33.1 



1918.] OEYZOMYS PALUSTRIS GROUP. 31 

(32-34.5). Skull (average of 5 adults from Yaruca, Honduras): 
Greatest length, 30.5 (29.9-31.3); zygomatic breadth, 16 (15.5-17.2); 
interorbital breadth, 4.8 (4.5-5.1); width of braincase, 11.4 (11.2- 
11.6); nasals, 11.9 (11.4-12.4); anterior palatine foramina, 6 
(5.5-6.2); palatal bridge, 5.5 (5.3-6.1); upper molar series, 4.7 
(4.5-4.8). 

Remarks. — In the absence of material from the type locality aa 
fixed by Thomas, specimens from Tumbala, Chiapas; and Yaruca, 
Honduras, which agree closely with his description, are assumed to 
represent typical couesi and have been used as a basis for comparison. 
Individual variation in size, color, and cranial details exhibited by 
every large series of 0. c. couesi is very striking, but the form main- 
tains with remarkable constancy its essential characters throughout 
its wide range. Examples from northern Vera Cruz and eastern 
Puebla present the same general variations and are not satisfactorily 
separable from those from Honduras and Nicaragua. This variation 
has resulted in the publication of several names based on characters 
which prove to be inconstant in the large number of specimens 
passed in review. Specimens from various localities indicate direct 
intergradation with 0. c. zygomaticus, 0. c. mexicanus, and 0. c. 
peragrus. 

Specimens examined. — Total number, 199, as follows: 

Campeche: La Tuxpeiia, 1. 

Chiapas: Chicharras, 4; Tumbala, 6. 

Costa Rica: Bahia de Salinas, l. 1 

Guatemala: Jacaltenango, 3. 

Honduras: Yaruca, 35. 2 

Nicaragua: Chontales, 22; 1 Matagalpa, 4; 1 Ocotal, 2; 1 PenaBlanca, 8 (including 
type of "richardsoni") ;* Quilali, l; 1 Rio Coco, 14; 1 Rio Grande, l; 1 Rio San 
JuandelNorte, l; 1 RioTuma, 2; 1 San Juan, l^Tuma, S^Uluce, l; 1 Vijagua^. 1 

Oaxaca: Comaltepec, 1; Guichicovi, 3; Reforma, l 3 ; Santo Domingo (mountain 
near), 8; Tuxtepec, 2. 

Puebla: Huauchinango, 1; Metlaltoyuca, 11. 

Quintana Roo: Santa Lucia, 3. 4 

Tabasco: Teapa, 3 (including type of "teapensis"). 

Vera Cruz: Achotal, 4; 3 Buena Vista, 2; Catemaco, 2 (including type of 
"rufinus"); Jalapa, 4 (including type of "jalapx"); 1 Pasa Nueva, l; 1 Coat- 
zacoalcos, 3 (including type of ' 'goldmani' ' ) ; Jico, 2 ; 3 Mjrador, 1 ; Motzorongo, 
1; San Carlos, 3 (including type of "ajmtclius")-? Orizaba, 16; 5 Rivera (75 
miles south), 1; Papantla, 1- Teocelo, If Tlacotalpam, 7; Ubero, 1. 

Yucatan: Rio Lagartos, 2. 3 



i Collection Amer. Mus. Nat. Hist. 

5 Twenty-six specimens in Mus. Comp. Zool.; 6 in Field Mus. Nat. Hist. 

s Collection Field Mus. Nat. Hist. 

* Collection Mus. Comp. Zool. 

5 Two in Field Mus. Nat. Hist. 



32 NORTH AMERICAN FAUNA. [No. 43. 

ORYZOMYS COUESI RICHMONDI Merriam. 

Richmond Rice Rat. 

Oryzomys richmondi Merriam, Proc. Washington Acad. Sci., Ill, p. 284, July 26, 1901. 

Type locality. — Escondido River (50 miles above Bluefields), Nica- 
ragua. 

Type. — No. |f f^g , c? adult, United States National Museum (Bio- 
logical Survey collection) ; collected by Charles W. Richmond, June 
21, 1892. 

Geographic distribution. — Low river valleys of eastern Nicaragua; 
Humid Lower Tropical Zone. 

General characters. — Size and proportions about as in 0. c. couesi; 
color decidedly darker. 

Color. — Similar to that of 0. c. couesi, but decidedly darker, more 
regularly ochraceous-tawny, the back and upper part of sides more 
heavily darkened by admixture of black hairs; underparts light 
ochraceous-buff. 

Skull. — As in 0. c. couesi. 

Measurements. — Type: Total length, 295; tail vertebrae, 150; hind 
foot, 33.5. Average of 10 adults (type and nine topotypes): 275.8 
(255-295); 137 (124-151); 30.9 (29-33.5). SJcull (average of 5 
adults — type and 4 topotypes): Greatest length, 31.8 (31.2-33.3); 
zygomatic breadth, 16.9 (16.4-17.9); interorbital breadth, 5.1 (5-5.3); 
width of braincase, 11.7 (11-11.9); nasals, 12.3 (11.6-13); anterior 
palatine foramina, 6(5.8-6.2); palatal bridge, 5.8 (5.7-5.9); upper 
molar series, 4.5 (4.3-4.6). 

Remarks. — This form, the darkest of the 0. couesi series, is known 
only from low elevations in eastern Nicaragua, where Richmond 
found it inhabiting banana plantations. Although much darker in 
general tone of upperparts than most examples of 0. c. couesi from 
adjacent territory, close agreement in all other important respects 
points to complete intergradation with the latter. 

Specimens examined. — Total number, 35, as follows: 

Nicaragua: Escondido River (50 miles above Bluefields, 12 2 [type and topotypes] ; 
45 miles above Bluefields, 18; 40 miles above Bluefields, 3; 25 miles above 
Bluefields, 1; 16 miles above Bluefields, 1). 

ORYZOMYS COUESI ZYGOMATICUS Merriam. 

Guatemalan Rice Rat. 

Oryzomys zygomaticus Merriam, Proc. Washington Acad. Sci., Ill, p. 285, July 26, 
1901. 

Type locality. — Nenton, Guatemala (altitude 3,000 feet). 

Type. — No. 76794, s adult United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, December 15, 1895. 



1918.] OEYZOMYS PALUSTEIS GROUP. 33 

Geographic distribution. — Known only from the Chiapas River 
valley in southwestern Guatemala and south-central Chiapas; Arid 
Lower Tropical Zone. 

General characters. — Closely resembling 0. c. couesi and 0. c. mexi- 
canus, but upperparts slightly paler than in the former and slightly 
darker than in the latter. 

Color. — About as in 0. c. couesi but averaging slightly paler, the 
general tone ochraceous-buff as in the palest examples of 0. c. couesi; 
underparts nearly pure white in the type, varying to light buff in 
other examples. 

SJcull. — As in 0. c. couesi. 

Measurements. — Type: Total length, 290; tail vertebrae, 152; hind 
foot, 33. Skull (type): Greatest length, 30.9; zygomatic breadth, 
26.9; interorbital breadth, 4.8; width of braincase, 11.6; nasals, 11.6; 
anterior palatine foramina, 6; palatal bridge, 5.9; upper molar series, 
4.6. 

Remarks. — 0. c. zygomaticus seems to be a slightly differentiated and 
rather localized form intermediate in color and geographic position 
between 0. c. couesi and 0. c. mexicanus. It doubtless intergrades 
directly with couesi, the type of which came from a higher elevation 
about 100 miles to the eastward in central Guatemala. Three speci- 
mens from Jacaltenango, at about 5,500 feet altitude, only a few 
miles to the southeast, are appreciably darker in color and seem to 
represent typical couesi. Near the Pacific Coast in southwestern 
Chiapas, zygomaticus doubtless passes into mexicanus, which differs 
mainly in having slightly paler color. 

Specimens examined. — Total number, 5, as follows: 

Chiapas: Ocuilapa, 1. 

Guatemala: N en ton, 4 (type and topotypes). 

ORYZOMYS COUESI MEXICANUS Allen. 

Mexican Rice Rat. 

Oryzomys mexicanus Allen, Bull. Amer. Mus. Nat. Hist., IX, p. 52, March 15, 1897. 
Oryzomys bulleri Allen, Bull. Amer. Mus. Nat. Hist., IX, p. 53, March 15, 1897. Type 

from Valle de Banderas, Tepic, Mexico, No. ffff, <? Bubad., Amer. Mus. Nat. 

Hist.; collected by Audley C. Buller, February 2, 1893. 
Oryzomys rufus Merriam, Proc. Washington Acad. Sci., Ill, p. 287, July 26, 1901. 

Type from Santiago, Tepic, Mexico (altitude 200 feet). No. 91404, 9 old, U. S. 

Nat. Mus. (Biological Survey collection); collected by E. W. Nelson and 

E. A. Goldman, June 20, 1897. 

Type locality. — Hacienda San Marcos, Tonila, Jalisco, Mexico (alti- 
tude 3,500 feet). 

Type.— No. -ff§-£, c? adult, American Museum of Natural History; 
collected by Audley C. Buller, December 30, 1889. 
14521°— 18 3 



34 NORTH AMERICAN FAUNA. [No. 43. 

Geographic distribution. — Pacific coastal plains and basal mountain 
slopes from southern Sinaloa to southeastern Oaxaca, Mexico; alti- 
tudinal range from sea level to about 1,000 feet (rarely to 3,500 feet), 
mainly in Arid Lower Tropical Zone. 

General characters. — Size and proportions about as in 0. c. couesi; 
color of upperparts rather decidedly paler, more ochraceous-buffy; 
underparts usually white, but varying to light buff, or light ochra- 
ceous-buff, the normal shades in couesi. Slightly paler than 0. c. 
zygomaticus; slightly darker than 0. c. aztecus, with smaller molar 
teeth. Similar in color to 0. c. alhiv enter, but size smaller,, and skull 
differing in detail. 

Color. — Fresh pelage: Upperparts in general varying shades of 
ochraceous-buff, more or less suffused with tawny in old adults, be- 
coming warm buff on cheeks, shoulders, and along lower parts of 
sides; the face, top of head, and back moderate^ darkened by 
blackish hairs; underparts usually nearly pure white, but varying to 
light buff; outer sides of ears dusky, the inner sides clothed with 
grayish or rusty reddish hairs; feet white; tail brownish above, dull 
yellowish below on proximal portion, becoming light brownish toward 
tip. Young (in first pelage) : Upperparts near tawny-olive, the gen- 
eral tone darker and duller than in adults. 

Skull. — About as in O. c. couesi. Differing from those of O. c. 
aztecus, O. c. regillus, and O. c. albiventer most noticeably in smaller 
size of molar teeth, the toothrows being correspondingly shortened. 

Measurements. — Average of 10 adults from Escuinapa, Sinaloa: 
Total length, 251.4' (239-273); tail vertebrae, 137.4 (127-165); hind 
foot, 28.9 (27-35). SkuU (average of same): Greatest length, 31.2 
(29.8-33); zygomatic breadth, 17 (16.1-17.8); interorbital breadth, 
4.8 (4.5-5.3) ; width of braincase, 11.6 (11.3-12.2); nasals, 12.2 (11.1- 
13.4); anterior palatine foramina, 6.1 (5.8-6.7); palatal bridge, 5.5 
(4.8-6); upper molar series, 4.4 (4.2-4.8). 

Remarks. — The narrow distribution area of 0. c. mexicanus along 
the west coast of Mexico somewhat parallels that of 0. c. couesi along 
the east coast; and as in that form, wide range of individual varia- 
tion in size and color has resulted in the publication of names which 
appear to be based on unstable characters. While individuals vary 
notably in size and contour, as shown by every large series, skulls of 
mexicanus and couesi seem indistinguishable, their general characters 
being maintained with remarkable uniformity throughout the com- 
bined ranges of the two forms. Intergradation of mexicanus with 
couesi seems to be indicated by specimens from the Isthmus of 
Tehuantepec, and with 0. c. aztecus by examples from the valley of 
the Balsas River in Guerrero. 



1918.] OEYZOMYS PALUSTKIS GROUP. 35 

Specimens examined. — Total number, 106, as follows: 

Colima: Armeria, 11; Hacienda Magdalena, 2. 

Guerrero: Ometepec, 7. 

Jalisco: Hacienda San Marcos, 1 (type). 1 

Michoacan: La Huacana, 5. 

Oaxaca: Huilotepec, 2; Juchitan, 1; Llano Grande, 5; Pluma, 3; Puerto Angel, 8; 

Reforma, l; 2 Santa Efigenia, 1; Tehuantepec, 1. 
Sinaloa: Escuinapa, 47; 1 Mazatlan, l; 3 Rosario, 2. 4 
Tepic: San Bias, 4; Santiago, 2 (including type of "rufus"); Valle de Banderas, 

2 (including type of "bulleri"). 

ORYZOMYS COITESI AZTECUS Merriam. 

Aztec Rice Rat. 

Oryzomys crinitus aztecus Merriam, Proc. Washington Acad. Sci., Ill, p. 282, July 26, 
1901. 

Type locality. — Yautepec, Morelos, Mexico (altitude 4,000 feet). 

Type. — No. 51173, 6" adult, United States National Museum (Bio- 
logical Survey collection); collected by E. W. Nelson and E. A. Gold- 
man, January 16, 1893. 

Geographic distribution. — Interior river valleys of Morelos, southern 
Puebla, northern Oaxaca, and northeastern Guerrero, Mexico; alti- 
tudinal range from about 3,000 to at least 4,000 feet in Arid Lower 
Tropical Zone. 

General characters. — A pale form with white underparts and rather 
heavy dentition. Closely resembling O. c. mexicanus, but paler in 
color and with heavier dentition than usual in that subspecies. Dif- 
fering from O. c. crinitus in paler upperparts, white instead of buffy 
underparts, and in cranial details. 

Color. — As in the paler examples of O. c. mexicanus, the prevailing 
tone beneath the dark hairs on the back pale ochraceous-buff ; under- 
parts nearly pure white. 

Skull. — About like that of O. c. mexicanus, but molar teeth larger. 
Similar to that of 0. c. crinitus, but braincase narrower and molars 
slightly smaller. 

Measurements. — Type: Total length, 290; tail vertebrae, 154; hind 
foot, 35. Two adults from Puente de Ixtle, Morelos: 318, 313; 160, 
170; 34, 33. Skull (type) : Greatest length, 32.4; zygomatic breadth, 
17; interorbital breadth, 5.2; width of braincase, 12.2; nasals, 12.9; 
anterior palatine foramina, 6.4; palatal bridge, 6.2; upper molar 
series, 5. 

Remarks. — O. c. aztecus is the palest form of the 0. couesi series 
but the light ochraceous tone of the upperparts is very closely ap- 
proached by the paler examples of O. c. mexicanus, O. c. albiventer, 

1 Collection Amer. Mus. Nat. Hist. » Collection Mus. Comp. Zool. 

» Collection Field Mus. Nat. Hist. < One in collection Mus. Comp. Zool. 



36 NORTH AMERICAN FAUNA. [No. 43. 

and 0. c. aquaticus. Specimens from the valley of the Balsas River 
in Guerrero and from northern Puebla approach mexicanus in denti- 
tion, as well as color, and might with nearly equal propriety be re- 
ferred to that subspecies. The characters separating aztecus and 
0. c. crinitus are rather slight, as indicated in the original descrip- 
tions, and it seems best to regard both as forms of 0. couesi. 
Specimens examined. — Total number, 20, as follows: 

Guerrero: Balsas, l; 1 Tlalixtaquilla, 1. 

Morelos: Cuernavaca, 1; Puente de Ixtle, 6; 2 Yautepec, 5 (type and topotypes). 

Oaxaca: Cuicatlan, 2. 

Puebla: Piaxtla, 4. 

/ 
ORYZOMYS COUESI CRINITUS Merriam. 

Valley of Mexico Rice Rat. 

Oryzomys crinitus Merriam, Proc. Washington Acad. Sci., Ill, p. 281, July 26, 1901. 

Type locality. — Tlalpam, Federal District, Mexico (altitude 7,500 
feet). 

Type. — No. 50182, J 1 subadult (molars unworn), United States 
National Museum (Biological Survey collection) ; collected by E. W. 
Nelson and E. A. Goldman, November 30, 1892. 

Geographic distribution. — Known only from type locality, at about 
7,500 feet altitude in the Valley of Mexico; Upper Sonoran Zone. 

General characters. — A large, moderately dark form with light 
buffy underparts and heavy dentition. Similar to 0. c. aztecus and 
O. c. albiventer, but upperparts darker than either and underparts 
buffy instead of white ; differing also in cranial details. 

Color. — Very similar to that of 0. c. mexicanus, but general tone of 
upperparts slightly darker; underparts light buffy as in some exam- 
ples of mexicanus. 

Skull. — Rather large with broad braincase and heavy dentition. 
Very similar to those of 0. c. aztecus, 0. c. albiventer, and O. c. regillus, 
but frontal region usually broader posteriorly; dentition about the 
same; interparietal rather large. 

Measurements. — Type: Total length, 307; tail vertebra?, 161 
hind foot, 37. Adult topotype: 280; 148; 35. Shull (type) 
Greatest length, 32.4; zygomatic breadth, 17; interorbital breadth, 5 
width of braincase, 12.3; nasals, 12.4; anterior palatine foramina 
6.7; palatal bridge, 5.7; upper molar series, 4.9. 

Remarks. — In the vicinity of marshes along the southern border of 
the Valley of Mexico, O. c. crinitus occurs at about 7,500 feet, the 
highest altitude attained by any known member of the 0. couesi 
section of the genus. The exact relationship of this form to O.fulgens 
remains to be determined, since it possesses some of the characters 

i Collection Field Mus. Nat. Hist. » Three in collection Field Mus. Nat. Hist. 



1918.] 0RYZ0MYS PALUSTRIS GROUP. 37 

ascribed to the latter species, the exact habitat of which is unknown. 
It is closely allied to the other forms inhabiting river valleys of the 
plateau region of Mexico — 0. c. aztecus, 0. c. albiventer, and 0. c. 
regillus. 

Specimens examined. — Three, from type locality. 

ORYZOMYS COUESI REGILLUS Goldman. 

Michoacan Rice Rat. 

(PI. I, figs. 4, 4a.) 

Oryzomys couesi regillus Goldman, Proc. Biol. Soc. Washington, XXVIII, p. 129, 
June 29, 1915. 

Type locality. — Los Reyes, Michoacan, Mexico. 

Type. — No. 125945, d adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, February 17, 1903. 

Geographic distribution. — Plateau region of northwestern Michoacan, 
Mexico; altitudinal range from about 3,000 to 4,000 feet, mainly in 
Arid Lower Tropical Zone. 

General characters. — A large form closely allied to O. c. albiventer; 
upperparts darker, more rufescent in color. Similar in general to 
O. c. mexicanus, but larger and richer colored; cranial details also 
distinctive. 

Color. — Much as in O. c. mexicanus and O. c. albiventer but upper- 
parts darker and more rufescent, the general tone rich ochraceous- 
buff, the back and rump strongly suffused with tawny and lined with 
black hairs as usual in the group; underparts varying from nearly 
pure white to light buff. 

STcull. — Like that of O. c. albiventer; similar to those of O. c. 
mexicanus and 0. c. couesi, but larger, with broader braincase and 
heavier dentition. 

Measurements. — Type: Total length, 305; tail vertebrae, 169; 
hind foot, 36. Average of three adult topotypes: 308 (285-320); 
168 (155-180); 35 (34-36). Skull (average of 4 adults— type and 3 
topotypes); Greatest length, 33.1 (32.5-33.5); zygomatic breadth, 
18.1 (17.5-18.5); interorbital breadth, 4.9 (4.6-5.3); width of brain- 
case, 12.5 (12.4-12.5); nasals, 12.9 (12.5-13.3); anterior palatine 
foramina, 6.2 (6-6.5); palatal bridge, 6.5 (6.4-6.6); upper molar 
series, 5.1 (4.9-5.3). 

Remarks. — This handsome rice rat differs only in color from its 
near geographic neighbor, O. c. albiventer, and the two doubtless 
intergrade in northern Michoacan. 

Specimens examined. — Total number, 13, as follows: 
Michoacan: Los Reyes, 11 (type and topotypes); Querendaro, 1; Zamora, 1. 



38 NORTH AMERICAN FAUNA. [No. 48. 

ORYZOMYS COUESI ALBIVENTER Merriam. 

White-Bellied Rice Rat. 

Oryzomys albiventer Merriam, Proc. Washington Acad. Sci., Ill, p. 279, July 26, 1901- 
Oryzomys molestus Elliot, Field Columb. Mus., publ. 71, zool. ser., Ill, p. 145, Feb- 
ruary, 1903. Type from Ocotlan, Jalisco, Mexico (altitude 5,000 feet). No. 
8667, c? old, Field Mus. Nat. Hist.; collected by F. E. Lutz, June, 1901. 

Type locality. — Ameca, Jalisco, Mexico (altitude 4,000 feet). 

Type. — No. 82236, <? adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, February 6, 1897. 

Geographic distribution. — River valleys of the plateau region in 
central Jalisco, Mexico; altitudinal range from about 4,000 to 5,000 
feet mainly in Lower Sonoran Zone. 

General characters. — Color about as in O. c. mexicanus, but size 
usually larger and cranial details distinctive. Similar in size and 
proportions to O. c. regillus and O. c. crinitus, but paler than either, 
the upperparts lacking the rich rufescent tone of the former, and the 
underparts normally white instead of buffy as in the latter. 

Color. — About as in O. c. mexicanus, underparts normally white, but 
varying to pale, creamy buff. 

SJcuU. — About like that of O. c. regillus; similar to that of O. c. 
crinitus, but frontal region usually narrower posteriorly. Compared 
with those of 0. c. couesi and 0. c. mexicanus the skull is decidedly 
broader, with heavier dentition. 

Measurements. — Type: Total length, 288; tail vertebrae, 153; 
hind foot, 37.5. Average of 10 adults (type and 9 topotypes): 294 
(276-314); 161 (151-173); 36.6 (34-40). Skull (average of 10 
adults — type and 9 topotypes): Greatest length, 33.1 (31.5-34.4); 
zygomatic breadth, 17.9 (17.3-18); interorbital breadth, 4.8 (4.1-5.2); 
width of braincase, 12.3 (11.5-12.8); nasals, 13 (12.3-14.2); anterior 
palatine foramina, 6.3 (5.9-6.6); palatal bridge, 6.3 (5.8-6.7); 
upper molar series, 5.3 (5-5.5). 

Remarks. — While O. c. albiventer is usually larger in general size, 
with broader skull and decidedly larger molar teeth than O. c. mexi- 
canus, occasional examples of the two forms are difficult to distin- 
guish and point to probable intergradation in western Jalisco. 
" Oryzomys molestus" of Elliot is based on an unusually large old 
adult which is clearly referable to albiventer, as are four examples 
in the Biological Survey collection from the same locality. 

Specimens examined. — Total number, 18, as follows: 
Jalisco: Ameca, 12 (type and topotypes); La Barca, 1; Ocotlan, 5 (including 
type of ''molestus.") 1 

i Collection Field Mus. Nat. Hist. 



1918.1 ORYZOMYS PALUSTRIS GROUP. 39 

ORYZOMYS COUESI PERAGRUS Merriam. 

Rio Verde Rice Rat. 

Oryzomys mexicanus peragrus Merriam, Proc. Washington Acad. Sci., Ill, p. 283, 
July 26, 1901. 

Type locality. — Rio Verde, San Luis Potosi, Mexico. 

Type. — No. 82119, 6* subadult, United States National Museum 
(Biological Survey collection); collected by E. W. Nelson and E. A. 
Goldman, January 8, 1897. 

Geographic distribution. — Known only from type locality, at about 
3,000 feet altitude on the Rio Verde River, in southern San Luis 
Potosi; Lower Sonoran Zone. 

General characters. — Closely allied to 0. c. couesi but slightly larger, 
with upperparts more pinkish instead of ochraceous-buffy in general 
tone; skull more massive. Similar to 0. c. aquaticus but upperparts 
usually darker, the ground color less ochraceous-buffy and the dorsal 
area more heavily lined with black. 

Color. — Similar to that of 0. c. couesi but upperparts paler, less 
ochraceous-buffy, the cheeks, shoulders, and sides near pinkish buff; 
lips and chin whitish. 

SJcull. — About like that of 0. c. aquaticus; larger and more massive 
than that of 0. c. couesi, with broader braincase and broader inter- 
orbital space. 

Measurements. — Type: Total length, 294; tail vertebra?, 167; hind 
foot, 35. Average of 2 adult topotypes: 274 (265-283); 151 (143- 
160); 33.5 (33-34). STcull (average of 3 adults— type and 2 topo- 
types): Greatest length, 31.5 (31-32.2); zygomatic breadth, 16.6 
(16.1-17.1); interorbital breadth, 5.5 (5.4-6.2); width of braincase, 
12.1 (12-12.3); nasals, 12.1 (11.8-12.5); anterior palatine foramina, 
5.8 (5.7-5.8); palatal bridge, 5.9 (5.7-6.2); upper molar series, 4.9 
(4.7-5). 

Remarks. — As nearly as can be determined by the three known 
specimens, all young adults, 0. c. peragrus is somewhat intermediate 
in characters, as well as geographic position, between O. c. couesi and 
0. c. aquaticus. It differs in color from both, but in cranial details 
approaches the latter form. 

Specimens examined.— Three, from type locality. 

ORYZOMYS COUESI AQUATICUS Allen. 
Rio Grande Rice Rat. 

(PI. I, figs. 5, 5a.) 

Oryzomys aquaticus Allen, Bull. Amer. Mus. Nat. Hist., Ill, p. 289, June 30, 1891. 

Type locality. — Brownsville, Cameron County, Texas. 
Type. — No. %m, e adult, American Museum of Natural History; 
collected by F. B. Armstrong, March 6, 1891. 



40 NORTH AMERICAN FAUNA. [No. 43. 

Geographic distribution. — Rio Grande Valley, from Camargo, 
Tamaulipas, to Gulf coast near Brownsville, Texas; altitudinal range 
from sea level to about 300 feet in lower Sonoran Zone. 

General characters. — Size large; closely resembling 0. c. peragrus 
but upperparts usually 'paler, less pinkish, more ochraceous-buffy; 
underparts buffy, as in peragrus. Similar in general to 0. c. couesi 
but larger; general color above paler ochraceous-buff; skull relatively 
broader and more massive. 

Color. — Fresh pelage: Upperparts near ochraceous-buff, palest on 
cheeks, shoulders, and along sides; the face, top of head, and back 
obscured by dusky hairs; underparts varying from light buff to 
warm buff, becoming in some' specimens more or less distinctly 
whitish on lips, chin, and throat; feet whitish; tail brownish above, 
pale yellowish below, becoming pale brownish toward tip. Young 
(in first pelage): Upperparts near tawny-olive; underparts with a 
pale buffy wash, the plumbeous basal color of the fur showing 
through. 

Skull. — About like that of 0. c. peragrus; decidedly larger and 
heavier throughout than that of 0. c. couesi, with broader braincase 
and more widely spreading zygomata; ascending branches of p re- 
maxillae exhibiting a tendency to exceed nasals in posterior extent 
(nasals usually exceeding premaxillae in forms of 0. couesi). 

Measurements. — Type: Total length, 286; tail vertebrae, 140; hind 
foot, 31.2 (dry skin). Average of 5 adult topotypes: 297 (283-310); 
161 (138-180); 34.5 (32-38). Shall (average of same): Greatest 
length, 33 (32-35.4); zygomatic breadth, 18 (16.7-18.8); interorbital 
breadth, 5 (4.7-5.2); width of braincase, 12 (11.6-12.7); nasals, 12.7 
(11.6-13.8); anterior palatine foramina, 6.3 (6.1-6.6); palatal bridge, 
6 (5.4-6.6); upper molar series, 5.1 (4.8-5.3). 

Remarks. — The Rio Grande Valley, inhabited by O. c. aquaticus, 
marks the extreme northern limit of the general range of the 0. couesi 
series. No rice rats are yet known from the interval of coastal plain 
in Tamaulipas, but close resemblance in all essential respects points 
to intergradation through O. c. peragrus with typical couesi. In 
general size and contour the skull of aquaticus is not widely different 
from those of 0. c. albiventer, 0. c. regillus, and O. c. crinitus of the 
Mexican plateau region, but it maintains the lighter dentition of 
typical couesi. 

Specimens examined. — Total number, 41, as follows: 

Texas: Brownsville, 37 (type and topotypes); 1 Lomita Ranch (Hidalgo 

County), 1. 
Tamaulipas: Camargo, 1; Matamoros, 2. 

i Fourteen in Amer. Mus. Nat. Hist.; 3 in Kansas Univ. Mus.; 2 in Field Mus. Nat. Hist.; 2 in Mus. 
Comp. ZooL; 2 in Acad. Nat. Sci. Philadelphia. 



1918.] ORYZOMYS PALUSTRIS GROUP. 41 

ORYZOMYS FULGENS Thomas. 
Thomas Rice Rat. 
Oryzomys fulgens Thomas, Ann. Mag. Nat. Hist., ser. 6, XI, p. 403, May, 1893. 

Type locality. — "Mexico." Southern Mexico, exact locality un- 
known (probably in or near Valley of Mexico) . 

Type. — 70.6.20.3, <? adult, British Museum; purchased of Geale, 
collected by A. Boucard. 

Geographic distribution. — Range unknown. 

General characters. — From original description: "Size large. Fur 
very thick, coarse and woolly. General colour above bright fulvous, 
brighter than in any other Central- American species; anterior half of 
the body, including the head, rather paler and duller than the pos- 
terior half. Ears decidedly small, broadly rounded, thinly haired, 
their hairs practically the same colour as those of the head in general, 
so that they are not distinguishable by colour at a distance. Lips, 
chin, throat, and inguinal region whitish, belly with a strong suffusion 
of fawn, which reaches a maximum on the breast between the fore 
legs; passage of upper colour into lower quite gradual. Outer sides 
of limbs like back, inner sides whitish; upper surfaces of hands and 
feet thinly clothed with pale silvery-fawn hairs. Tail long, thinly 
haired, the scales not hidden by the hairs; above blackish, below yel- 
lowish, darkening towards the tip." 

Skull. — From original description: "Skull readily distinguishable 
from all allied species by its great breadth, the bold expansion of the 
zygomata, and especially by the evenly incurved outline of the 
supraorbital edges; in all other species these edges form two approxi- 
mately straight lines diverging from the narrowest interorbital 
point, but in 0. fulgens the whole inner wall of the orbit forms one 
even curve, the breadth at the posterior end of the olfactory cham- 
ber being scarcely greater than at the anterior end. Nasals broad 
and flattened. Frontal premaxillary processes very narrow and 
barely attaining to the same level as the back of the nasals. Anterior 
palatine foramina large, widely open, their posterior margin just 
level with the front of ml ." 

Measurements. — Dry skin of type (from original description): 
Head and body, 160; tail, 151; hind foot, 37.5 (c. u.). Skull (type): 
Upper length/ 31.8; zygomatic breadth, 17.8 (c); nasals, 13.2; 
interorbital breadth, 4.8; diastema, 9.1; anterior palatine foramina, 
7.2; upper molar series, 5.2. 

Remarks. — The type of 0. fulgens I have been unable to examine, 
and none of the more recently accumulated material from Mexico 
can at present be assigned with certainty to that species. Thomas's 
full description, above quoted, and comparisons kindly made for me 

1 To back of parietal suture only. 



42 NORTH AMERICAN FAUNA. [No. 43. 

by W. H. Osgood, however, seem to indicate that this is a member of 
the widely dispersed 0. couesi section of the genus. Some of the 
characters given are shared in common by various forms now recog- 
nized. In size and color arrangement it closely approaches 0. c. 
crinitus, of the Valley of Mexico, and may be identical with that 
form, but the intensity of color and details presented by the incom- 
plete skull, especially the form of the interorbital region, appear to 
be distinctive and the exact position of fitlgens remains to be 
determined. 

ORYZOMYS GATUNENSIS Goldman. 

Gatun Rice Rat. 
/ 
(PI. I, figs. 6, 6a.) 

Oryzomys gatunensis Goldman, Smiths. Misc. Coll., LVI, no. 36, p. 7, February 19, 
1912. 

Type locality. — Gatun, Canal Zone, Panama. 

Type. — No. 171034, d young (about two-thirds grown), United 
States National Museum (Biological Survey collection) ; collected by 
E. A. Goldman March 7, 1911. 

Geographic distribution. — Known only from type locality, near sea 
level; Humid Lower Tropical Zone. 

General characters. — A dark-colored form externally similar to 0. c. 
richmondi, but with distinctive cranial characters. 

Color. — Fresh pelage: Upperparts near ochraceous-tawny, palest 
on cheeks, shoulders, and along sides; the face, top of head, and 
back much darker by admixture of black hairs; underparts light 
ochraceous-buffy; outer sides of ears blackish, the inner sides clothed 
with ochraceous-buffy hairs; feet thinly covered with very short 
whitish or grayish hairs; tail light brownish above, somewhat paler 
below. 

Skull. — Similar in general to that of 0. c. richmondi, but frontal 
region broader, the lateral margins more developed as supraorbital 
shelves; interparietal much less extended antero-posteriorly ; nasals 
more prolonged posteriorly be3 r ond premaxillse; dentition about as 
in richmondi. 

Measurements. — Type: Total length, 224; tail vertebrae, 115; hind 
foot, 31.5. Skull (type): Greatest length, 27.7; zygomatic breadth, 
14.5; interorbital breadth, 5.3; width of braincase, 1 1 .4 ; nasals, 10.2; 
anterior palatine foramina, 5.6; palatal bridge, 5.2; upper molar 
series, 5. 

Remarks. — This species requires comparison with only 0. c. rich- 
mondi, with which it is nearly identical in color and general external 
appearance. The skull, however, differs in apparently important 
respects from those of all the forms of 0. couesi. Especially notice- 
able is the lateral development of the supraorbital ridges, a character 



1918.] ORYZOMYS PALUSTRIS GROUP. 43 

which in fully adult examples would doubtless be more pronounced ; 
the reduced antero-posterior extent of the interparietal seems to be 
another distinguishing feature. On the other hand the material 
representing 0. gatunensis is scanty and the range of individual varia- 
tion being undetermined, intergradation with couesi and richmondi 
in Costa Rica or western Panama seems not improbable. 
Specimens examined. — Two, from type locality. 

ORYZOMYS COZUMEL^] Merriam. 
Cozumel Rice Rat. 

(PI. I, figs. 7, 7a.) 
Oryzomys cozumelx Merriam, Proc. Washington Acad. Sci., Ill, p. 280, July 26, 1901. 

Type locality. — Cozumel Island, off east coast of Quintana Roo, 
Mexico. 

Type. — No. 108462, <? adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, April 8, 1901. 

Geographic distribution.— Known only from type locality. 

General characters. — Size large; tail much longer than head and 
body; color dark. A large species allied to 0. c. couesi of the neighbor- 
ing mainland, differing in larger size, darker color, and proportionately 
longer tail. 

Color.— Worn pelage: Upperparts between ochraceous-tawny and 
cinnamon-brownish, palest on cheeks, shoulders, and sides, darkened 
on face, top of head, and back by a brownish admixture; underparts 
between light buff and light ochraceous-buff ; feet whitish; tail dark 
brownish above, dull yellowish below basally, becoming dusky all 
around toward tip. 

Skull. — In general outline closely resembling that of 0. c. couesi 
but larger and more massive; rostrum and anterior roots of zygomata 
relatively slightly heavier; dentition about as in couesi. 

Measurements. — Type: Total length, 332; tail vertebrae, 182; hind 
foot, 35. Average of 6 adult topotypes: 306 (285-327); 172 (163- 
177); 34.3 (33-35.5). STcidl (average of 6 adults — type and 5 topo- 
types): Greatest length, 32 (30.5-34.4); zygomatic breadth, 16.9 
(15.9-17.9); interorbital breadth, 5 (4.7-5.6); width of braincase, 
12.1 (11.5-12.6); nasals, 12.5 (11.7-13.6); anterior palatine foramina, 
6.3 (6.2-6.7); palatal bridge, 5.8 (5.7-6.1); upper molar series, 5 
(4.8-5.3). 

Remarks. — This insular species is clearly an offshoot of O. couesi, 
the widely ranging mainland form. Its divergence from typical 
couesi is mainly in the direction of larger general size and the develop- 
ment of a relatively longer tail. 

Specimens examined. — Nine, from type locality. 



44 NORTH AMERICAN FAUNA. [No. 43. 

ORYZOMYS ANTILLARUM Thomas. 
Jamaican Rice Rat. 

Oryzomys ant'dlarum Thomas, Ann. Mag. Nat. Hist., ser. 7, I, p. 177, February, 1898. 

Type locality. — Jamaica. 

Type— No. 45.10.25.48, British Museum; collected by P. H. 
Gosse. 

Geographic distribution. — Known only from Jamaica. 

General characters. — Allied to 0. couesi; size and proportions about 
as in the typical subspecies; color apparently similar; skull differing 
in rather slight details. 

Color. — From original description of type: "General colour dull 
rufous, rather (though not prominently) richer on the rump and 
greyer on the head; black lining of back not prominently marked. 
Belly dull yellowish, not sharply denned, the hairs slaty grey basally. 
No blackish ring round eyes. Ears small, their visible external sur- 
face blackish and internal yellowish, but in neither case very strongly 
contrasting with the general colour. Hands and feet dull whitish 
above. Tail apparently about as long as head and body, very thinly 
haired, almost naked, pale brownish above, rather lighter below." 
Two specimens in the United States National Museum, collected 
many years ago, are very tawny above, but appear to have been im- 
mersed in alcohol, and the naturally ruddy tone thereby intensified. 

Slcull. — Closely resembling that of 0. c. couesi, but nasals reaching 
farther posteriorly beyond premaxillse (nasals and premaxillse more 
nearly conterminous in couesi); maxillary arm of zygoma heavier; 
anterior palatine foramina shorter than usual in couesi; dentition 
about the same. 

Measurements. — From original description of type (measured in 
skin): "Head and body (apparently stretched) 130 millim. ; tail (im- 
perfect at tip) 130; hind foot without claws (moistened), 28." * * * 
Skull (type): "Basal length (c.) 26, basilar length (c.) 24; greatest 
breadth 17; nasals 12.6x4.1; interorbital breadth 5.2; breadth of 
braincase on squamosals 12.9; interparietal 2.8 x 8.5; palate length 
from henselion 14 ; diastema 8.3 ; palatal foramina 5.7 x 2.1 ; length of 
upper molar series 4.6". An adult from Metcalfe Parish (dry skin): 
Total length, 252; tail vertebras, 122; hind foot (c. u.), 29.2. STcuU 
(of same): Greatest length, 30.5; zygomatic breadth, 16.6; interor- 
bital breadth, 5.1 ; width of braincase, 12.2 ; nasals, 12.6 ; anterior pala- 
tine foramina, 5.7; upper molar series, 4.5. 

Remarks. — The relationship of the Jamaican rice rat to 0. couesi 
was pointed out by Thomas in Ms original account of the species. In 
view of its isolation, the general agreement in all the more important 
respects, even to size, a character usually subject to modification in 
insular forms, is remarkable, and suggests the possibility that 0. aniil- 



1918.] ORYZOMYS PALUSTRIS GROUP. 45 

larum, may have been transported from some point on the coast of the 
North American mainland within a comparatively recent period. 
As suggested by Thomas, however, the fact that no specimens appear 
to have been taken since 1877, while rats of the Old World have 
devastated the island, to be persecuted in their turn by the mon- 
goose, introduced in 1872, renders it probable that the rice rat has 
been exterminated there. 

Specimens examined. — Total number, 2, as follows: 
Jamaica: Metcalfe Parish, 1; Spanishtown, 1. 

ORYZOMYS PENINSULA Thomas. 

Lower California Rice Rat. 

(PI. I, figs. 8, 8a.) 

Oryzomys peninsulse Thomas, Ann. Mag. Nat. Hist., ser. 6, XX, p. 548, December, 
1897. 

Type locality. — Santa Anita, Lower California, Mexico. 

Type. — Male adult, British Museum. 

Geographic distribution. — Known only from very limited marshy 
areas near sea level in extreme southern Lower California; Arid Lower 
Tropical Zone. 

General characters. — General size and proportions about as in 
0. c. mexicanus; color similar, but anterior part of body, especially 
head and shoulders, strongly suffused with gray; skull rather broad, 
with squarely spreading zygomata and large interparietal. 

Color. — Fresh pelage: Upperparts between warm buff and pale 
ochraceous-buff (more or less distinctly tawny in worn pelage of old 
adults) most intense on rump, becoming light buffy grayish on head, 
fore limbs, shoulders, and sides; underparts overlaid with white, 
the basal color of the fur usually plumbeous, except on chin and 
throat, where it is pure white to roots; outer sides of ears brownish, 
inner sides clothed with grayish or ochraceous-buff y hairs; feet white; 
tail light brownish above, yellowish below, becoming more or less 
distinctly brownish toward tip. Young (in first pelage) : Upper- 
parts more fuscous, the head and shoulders lacking the grayish suffu- 
sion so noticeable in adults; underparts dull buffy or soiled white, 
becoming pure white on chin and throat. 

STcull. — Similar to that of 0. c. mexicanus, but broader; zygomata 
more widely and squarely spreading; braincase less flattened, higher, 
more rounded or inflated; interparietal larger; anterior palatine 
foramina much broader, more widely open; lateral wings of parietals 
extending farther below temporal ridges at expense of squamosals; 
ascending branches of premaxillse usually reaching posteriorly beyond 
nasals (nasals usually exceeding premaxillse in posterior extent in 
mexicanus) . 



46 NORTH AMERICAN FAUNA. [No. 43. 

Measurements. — Average of nine adults from San Jose del Cabo, 
Lower California (near type locality) : Total length, 283 (270-305) ; 
tail vertebra?, 143 (138-156); hind foot, 32.2 (31.5-34). STcull 
(average of eight adults from San Jose del Cabo, Lower California) 
Greatest length, 32.5 (31.5-33.9); zygomatic breadth, 17.9 (17.1-19) 
interorbital breadth, 5.1 (4.9-5.3) ; width of braincase, 12 (11.4-12.2) 
nasals, 13 (12.6-13.4); anterior palatine foramina, 6.6 (6.2-7.4) 
palatal bridge, 6.2 (6-6.4); upper molar series, 4.8 (4.7-5.1). 

Remarks. — 0. peninsulas, is allied to 0. c. mexicanus of the adjacent 
mainland coast of Mexico, but important differential cranial charac- 
ters are numerous. The range of the species seems to be centered in 
the marshes near the mouth of the San Jose River at San Jose del 
Cabo, extending a few miles inland along the course of the stream. 
The neighboring coast is extremely arid and unsuitable for habitation 
by an Oryzomys. 

Specimens examined. — Total number, 16, as follows: 

Lower California: San Jose del Cabo, 15; Santa Anita, 1 (topotype). 

ORYZOMYS NELSONI Merriam. 

Nelson Rice Rat. 

(PI. II, figs, l, la.) 

Oryzomys nelsoni Merriam, Proc. Biol. Soc. Washington, XII, p. 15, January 27, 1898. 

Type locality. — Maria Madre Island, Tres Marias Islands, off coast 
of Tepic, western Mexico (altitude 800 feet). 

Type. — No. 89200, o* adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, May 13, 1897. 

Geographic distribution. — Known only from type locality, Maria 
Madre Island, where it inhabits moist places on the upper slopes at 
about 800 feet altitude; Arid Lower Tropical Zone. 

General characters. — Size very large; tail much longer than head and 
body; color of upper parts rich ochraceous-buff ; skull rather long and 
narrow, but massive. Allied to 0. c. mexicanus, but differing widely 
in details of structure. 

Color. — Fresh pelage: Upperparts rich ochraceous-buff, most intense 
on rump, paling to warm buff on head, shoulders, and lower parts of 
sides ; somewhat darkened on face, top of head, and back by dusky 
hairs; underparts white; outer and inner sides of ears thinly clothed 
with grayish hairs ; tail light brownish above and all round near tip, 
becoming yellowish below on basal portion. 

STcull. — Massive, upper outline rising high over anterior roots of 
zygomata, the rostrum very heavy and strongly decurved. Some- 
what similar in general to that of 0. c. mexicanus, but much larger 
and heavier; rostrum much more swollen and decurved; zygomata 



1918.] 



OKYZOMYS MELANOTIS GROUP. 



47 



heavier, but relatively less widely spreading, the sides more nearly 
parallel; supraorbital ridges less divergent posteriorly; interparietal 
larger; anterior palatine foramina relatively shorter; dentition about 
as in mexicanus. 

Measurements. — Type: Total length, 324; tail vertebras, 190; hind 
foot, 38. Two adult topotypes: 344, 320; 191, 185; 39, 37. Skull 
(average of 3 adults — t}^pe and 2 topotypes) : Greatest length, 35.8 
(34.5-37.8); zygomatic breadth, 18.7 (18.2-19); interorbital breadth, 
5.4 (5.2-5.6); width of braincase, 12.9 (12.8-13.1); nasals, 14.5 
(14.2-14.8); anterior palatine foramina, 6.3 (6.1-6.7); palatal bridge, 
6.9 (6.6-7.3); upper molar series, 5.3 (5.1-5.4). 

Remarks. — The isolation of 0. nelsoni on an island 70 miles off the 
mainland coast has resulted in the evolution of a well-marked species, 







i*i/ 



■*"/* 



^T: 



HI DA*- 6 ?. 













C? / HON D. 



1. 0. TO. melanotis. 4. 0. r. megadon. 

2. 0. m. colimensis. 5. O. r. yucatanensis. 

3. O. r. rostratus. 

Fig. 4. — Geographic distribution of the Oryzomys melanotis group. 

but agreement in essential characters with forms of 0. couesi places 
it in that widely distributed section of the genus. It differs strikingly 
from all the mainland forms in the remarkable development of the 
rostrum. 

Specimens examined. — Four, from type locality. 

Oryzomys melanotis Group. 

Geographic distributio7i — Mainly lower elevations near the Pacific 
coast of Mexico from southern Sinaloa to Colima, and along the gulf 
coast from southern Tamaulipas to the Yucatan peninsula; altitudinal 
range from sea level to about 3,000 feet; Arid and Humid Lower 
Tropical Zones (fig. 4). 

General characters. — Size small, medium, or large; form rather 
slender; tail about same length as head and body or somewhat longer, 



48 NORTH AMERICAN FAUNA. [No. 43. 

scantily and indistinctly haired; ears large and conspicuous, thinly 
clothed externally with short, fine dusky hairs and internally with 
similarly short, ruf escent hairs ; general pelage short, rather harsh, 
and lacking the woolly quality of 0. palustris; vibrissas about as long 
as head; toes of hind feet more or less distinctly webbed at base, the 
three longest bearing tufts of silvery bristles which project beyond 
ends of claws; claws short, recurved, compressed, and sharp pointed. 
Color of upperparts varying from ochraceous-buff to rich ochraceous- 
tawny, lined with black; underparts white or buffy whitish. 

Skull. — Size small, medium, or large, with rostrum long and brain- 
case rather narrow ; maxillary /arm of zygoma heavy ; outer wall of 
antorbital foramen with projecting border rounded or sloping for- 
ward, the antorbital fossa deep and conspicuous, but less evenly 
circular than in 0. palustris as viewed from above; nasals and pre- 
maxillse about conterminous posteriorly; frontal region wide or 
narrow, constricted near middle, the lateral margins ridged or smooth; 
temporal ridges moderately developed anteriorly along parieto- 
squamosal sutures, becoming indistinct posteriorly in crossing lateral 
wings of parietals; interparietal large; anterior palatine foramina 
short and broad, much shorter than palatal bridge, not normally 
reaching anterior plane of first molars; palatal pits and sphenopala- 
tine vacuities large; interpterygoid fossa broad; audital bullas small, 
the inner sides extensively overlapped by mastoids; basi-occipital 
broad; angle of mandible broad, the inferior border strongly turned 
inward; coronoid process short and low, owing largely to high, thin 
commissural border extending to condyle ; molars small, approaching 
the 0. palustris type but smaller, second upper molar more evenly 
cleft by inner reentrant angle, and third lower molar more deeply 
incised by outer reentrant angle. The skulls of 0. melanotis and its 
allies differ from those of the 0. palustris group most noticeably in 
the elongation of the rostrum, shortness of anterior palatine foramina 
in relation to palatal bridge, small size of audital bulla?, and the dental 
details noted. 

Remarks. — Unlike 0. palustris and its allies, which live on the 
ground mainly in open marshy places or resort to the vicinity of 
water, members of this group favor more forested areas, where they 
are partially scansorial in habits, and occur on well-drained but moist 
mountain slopes sometimes at points distant from water. 

In external appearance, forms of the 0. melanotis group superficially 
resemble those of 0. couesi, and the two often occur together at the 
same localities. In contrast with 0. couesi, however, the upperparts 
are usually more rufescent, the ears are much larger, less conspicu- 
ously hairy, and tufts of silvery bristles project beyond the points 
of the three longer claws on the hind foot (claws reaching beyond 
bristles in couesi). 



1918.] ORYZOMYS MELANOTTS GROUP. 49 

Key to Species and Subspecies of the O. melanotis Group. 

a 1 . Size larger; hind foot 30 or more. (Atlantic coast of Mexico [0. rostratus].) 
b 1 . Upper molar series 4 or more. 

c 1 . Color darker. (Tabasco; Campeche.) O. r. megadon (p. 54). 

c 2 . Color paler. (Southern Tamaulipas; Vera Cruz; northern Puebla; north- 
eastern Oaxaca.) O. r. rostratus (p. 53). 

b 2 . Upper molar series less than 4. (Yucatan.) O. r. yucatanensis (p. 55). 

a 2 . Size smaller; hind foot less than 30. (Pacific coast of Mexico [0. melanotis}.) 

b 1 . Color darker. (Sinaloa; Tepic; Jalisco.) O. m. melanotis (p. 50). 

b 2 . Color paler. (Colima.) O. m. colimensis (p 51). 

ORYZOMYS MELANOTIS Thomas. 

[Synonymy under subspecies.] 

Geographic distribution. — Pacific coastal plains and basal mountain 
slopes from southern Sinaloa south through western Tepic and Jalisco 
to Colima; altitudinal range from sea level to 3,000 feet; Arid Lower 
Tropical Zone. 

General characters. — Size small or medium; general color above 
varying from rich ochraceous-buffy to pale ochraceous-tawny ; simi- 
lar to small forms of 0. rostratus in general external appearance but 
skull differing in important details. 

Color. — Fresh pelage: Upperparts varying from ochraceous - buff 
to pale ochraceous-tawny (sometimes becoming intense tawny or 
rusty reddish in the worn pelage of old adults), rather sparingly 
lined with black hairs, which becoming less numerous along cheeks 
and sides, leave them a purer, brighter color than the back; under- 
pays pale buff y white, the dark basal color of the fur showing through ; 
outer sides of ears blackish, the inner sides thinly clothed with rusty 
reddish hairs; a more or less conspicuous patch of light-colored fur 
under base of ear; feet (epidermis) dull yellowish, thinly covered 
above with short white hairs; tail (epidermis) brownish above, be- 
coming light brownish or dull yellowish below except near lip, which 
is usually dark all around. Young (in first pelage): Upperparts 
duller and darker, the general tone browner than in adults. 

SlcuU. — Similar in general to that of O. rostratus, but upper outline 
less elevated over anterior part of frontals; braincase relatively 
higher, the parietal region more expanded; rostrum more slender, 
less decurved; frontals broader, with narrow and delicate but rather 
well-developed supraorbital borders; parietals more squarely truncate 
anteriorly, the fronto-parietal sutures more widely divergent; tem- 
poral ridges tending to spread more widely posteriorly, rejoining 
squamosals after crossing slightly developed lateral wings of parietals; 
interparietal larger, with a less evident posterior angle; outer wall 
of antorbital foramen narrower, less extended anteriorly; anterior 
palatine foramina short and wide as in rostratus; dentition about the 
same. 

14521°— X8 4 



50 NORTH AMERICAN FAUNA. [No. 43. 

Remarks. — Two closely allied geographic races of 0. melanotis 
occupy a part of the arid coast region of western Mexico, where they 
appear to be completely isolated from their relatives (subspecies of 
0. rostratus) along the Gulf coast of eastern Mexico. The not very 
distant relationship of 0. melanotis to 0. rostratus is evidenced in 
numerous characters, but the two species appear to be distinct. In 
general external appearance, including color, they are much alike; 
in fresh pelage 0. melanotis may usually be distinguished by the 
whitish subauricular spots; the skulls are easily separable by the 
characters pointed out. 

ORYZOMYS MELAN0TIS MELANOTIS Thomas. 
Jalisco Rice Rat. 

(PI. II, figs. 2, 2a; PI. V, fig. 5; PI. VI, figs. 2, 2a.) 
Oryzomys melanotis Thomas, Ann. Mag. Nat. Hist., ser. 6, XI, p. 404, May, 1893. 

Type locality. — Mineral San Sebastian, Jalisco, Mexico. 

Type. — 93.3.6.25, d old, British Museum; collected by Dr. Audley C. 
Buller, January 25, 1893. 

Geographic distribution. — Coastal plains and basal mountain slopes 
in southern Sinaloa, Tepic, and Jalisco; altitudinal range from sea 
level to about 3,000 feet; Arid Lower Tropical Zone. 

General characters. — Size medium; color of upperparts near pale 
ochraceous-taAvny; closely allied to 0. m. colimensis but larger and 
darker colored; externally similar to pale examples of 0. rostratus; 
skull with long, slender rostrum and high-arched braincase. 

Color. — Fresh pelage: Upperparts pale ochraceous-tawny (becoming 
intense tawny or rusty reddish in the worn pelage of old adults), 
rather sparingly lined with black hairs, which becoming less numerous 
along cheeks and sides leave them a purer, brighter color than on 
the back; underparts white or pale buffy white, the dark basal color 
of the fur showing through; outer sides of ears blackish, inner sides 
thinly clothed with rusty reddish hairs; a more or less conspicuous 
patch of whitish fur under base of ear; feet whitish; tail brownish 
above, becoming light brownish or yellowish below except near tip, 
which is usually dark all around. Young (in first pelage) : Upper- 
parts duller and darker, the general tone browner than in adults. 

STcull. — Similar to that of O. m. colimensis, but larger, with com- 
paratively smaller molar teeth. 

Measurements. — Average of five adult topotypes: Total length, 
235 (228-244); tail vertebrae, 128 (124-134); hind foot, 28.2 (27.5-29). 
STcuU (two adult topotypes): Greatest length, 28.5, 27.9; zygomatic 
breadth, 14.4, 14.5; interorbital breadth, 4.8, 5.2; width of brain- 
case, 10.8, 10.7; nasals, 11.5, 10.3; anterior palatine foramina, 4.3, 
5.2; palatal bridge, 5.4, 5.4; upper molar series, 4, 4.3. 



1918.] ORYZOMYS MELANOTIC GROUP. 51 

Remarks. — Specimens from localities near sea level in Jalisco and 
Tepic are larger than those from the type locality at 3,000 feet alti- 
tude on the slope of the mountains, and may represent a slightly 
different form. In size they contrast strongly with the small form 
0. m. colimensis inhabiting the coast of Colima. 

Specimens examined. — Total number, 13, as follows: 
Jalisco: Ixtapa, 2; San Sebastian, 6 (type and topotypes). 
Sinaloa: Los Limones, l. 1 
Tepic: San Bias, 2; Santiago, 2. 

ORYZOMYS MELANOTIS COLIMENSIS, subsp. nov. 

Colima Rice Rat. 

(PI. II, figs. 3, 3a.) 

Type locality. — Armeria, Colima, Mexico (altitude about 100 feet). 

Type.— No. !ftff> 9 adult > United States National Museum (Bio- 
logical Survey collection); collected by E. W. Nelson, March 2, 1892. 
Original number 1987. 

Geographic distribution. — Forested coastal plains and basal moun- 
tain slopes in the State of Colima, Mexico ; altitudinal range from 
sea level to 1,500 feet; Arid Lower Tropical Zone. 

General characters. — A small form closely allied to O. m. melanotis; 
differing mainly in smaller size and paler color; general tone of upper- 
parts ochraceous-buff instead of ochraceous-tawny as in melanotis. 

Color. — Upperparts in general near ochraceous-buff, the top of 
head and back sparingly mixed with black, giving a lined effect; 
cheeks, shoulders, and sides paler than back, the general tone light 
ochraceous-buff; underparts whitish or pale buffy whitish; ears 
blackish or brownish, thinly clothed on outer sides with short dusky 
hairs, and on inner sides with buffy or rusty reddish hairs ; a patch of 
whitish fur under base of ear as in 0. m. melanotis; feet yellowish, 
thinly covered above with short white hairs; tail brownish above, 
yellowish below to near tip, which is dusky all around. 

Skull. — About like that of O. m. melanotis, but smaller; molar teeth 
actually about the same size as those of melanotis, and therefore rela- 
tively larger. 

Measurements. — Type: Total length, 216; tail vertebrae, 116; 
hind foot, 26. Adult topotype: 220; 118; 28. Skull (average of 
two adults, type and topotype): Greatest length, 27 (26.3-27.7); 
zygomatic breadth, 14 (13.9-14.2); interorbital breadth, 4.7 (4.5- 
4.9); width of braincase, 10.2 (9.9-10.6); nasals, 10.7 (10.3-11.1); 
anterior palatine foramina, 4.4 (4.3-4.5); palatal bridge, 5.4 (5.4- 
5.5); upper molar series, 4 (3.9-4.1). 

i Collection Amor, Mus. Nat. Hist. 



52 NORTH AMERICAN FAUNA. [No. 43. 

Remarks. — This small form is known only from the State of Colima, 
but probably ranges to the southward along the coast of Michoacan. 
An adult example from Hacienda Magdalena is dark in color and in 
this respect approaches 0. m. melanotis, but agrees otherwise with 
the small series of topotypes from near the Pacific coast. 

Specimens examined. — Total number, 4, as follows: 
Colima: Armeria, 3 (type and topotypes); Hacienda Magdalena, 1. 

ORYZOMYS ROSTRATUS Merriam. 

[Synonymy under subspecies.] 

Geographic distribution. — Coastal plains and basal mountain slopes 
from extreme southeastern Tamaulipas through northern Puebla, 
Vera Cruz, northeastern Oaxaca, Tabasco, Campeche, and Yucatan 
to northern Quintana Roo; altitudinal range from sea level to about 
1,500 feet; Arid and Humid Lower Tropical Zones. 

General characters. — A rather large, rufescent species, not very 
unlike 0. melanotis in external appearance, but cranial characters 
distinctive. (For additional characters, excepting specific color, see 
under 0. melanotis group.) 

Color. — Upperparts varying from ochraceous-buff to rich intense 
ochraceous-tawny, purest and brightest along cheeks and sides; the 
face, top of head, and back moderately lined with black hairs, which 
alter the general tone; underparts white or pale buffy white, the 
plumbeous basal color usually showing through; outer sides of ears 
blackish, the inner sides thinly and inconspicuously clothed with pale 
buffy or rusty reddish hairs; feet (epidermis) dull yellowish, thinly 
covered with short white hairs; tail (epidermis) varying from nearly 
uniform brownish throughout to irregularly yellowish on under side. 
Young (in first pelage) : Upperparts darker and less rufescent than in 
adults. 

STcull. — Similar in general to that of 0. melanotis, but upper outline 
more elevated over anterior part of frontals; braincase relatively 
lower, the parietal region less expanded; rostrum heavier, more 
decurved; frontals narrower posteriorly; parietals less squarely 
truncate anteriorly, the fronto-parietal sutures less widely divergent; 
temporal ridges tending to spread less widely posteriorly; interpa- 
rietal smaller, with a more evident posterior angle; outer wall of 
antorbital foramen broader, more extended anteriorly; anterior pala- 
tine foramina short and wide as in melanotis; dentition about the 
same. 

Remarks. — Three geographic races of 0. rostratus are recognizable, 
all of which closely intergrade and differ rather slightly in average 
size, color, or cranial details. Viewed as a whole, the forms exhibit a 
progressive decrease in size from west to east, typical rostratus being 
the largest and 0. r. yucatanensis the smallest of the series. 



1918.] ORYZOMYS MELANOTIS GROUP. 53 

ORYZOMYS ROSTRATUS ROSTRATUS Merriam. 
Metlaltoyuca Rice Rat. 

(PI. II, figs. 4, 4a.) 
Oryzomys rostratus Merriam, Proc. Washington Acad. Sci., Ill, p. 293, July 26, 1901. 

Type locality.— Metlaltoyuca, Puebla, Mexico (altitude 800 feet). 

Type. — No. 93112, <? old, United States National Museum (Bio- 
logical Survey collection); collected by E. W. Nelson and E. A. 
Goldman, February 5, 1898. 

Geographic distribution. — Forested coastal plains and basal moun- 
tain slopes in southeastern Tamaulipas, northern Puebla, Vera 
Cruz, and northeastern Oaxaca; altitudinal range from sea level to 
about 1,500 feet; Arid and Humid Lower Tropical Zones. 

General characters. — Similar to O. r. megadon and 0. r. yucatanensis, 
but averaging larger than either; color rather pale, much as in yuca- 
tanensis, slightly paler, less rufescent than usual in megadon; skull 
broad. 

Color. — Fresh pelage: Upperparts rich ochraceous-tawny, purest 
and brightest along cheeks and sides ; the face, top of head, and back 
moderately lined with black hairs which darken the general tone; 
underparts nearly pure white in some specimens, pale buffy white 
in others, the dark basal color, however, usually showing through; 
outer sides of ears blackish, the inner sides thinly clothed with pale 
buffy or rusty reddish hairs; feet (epidermis) dull yellowish, thinly 
covered with short white hairs; tail varying from nearly uniform 
brownish throughout to irregularly yellowish on under side. 

Skull. — Closely resembling those of O. r. megadon and O. r. yuca- 
tanensis in general form, but larger and relatively broader than 
either, the greater breadth most conspicuous in the braincase; 
dentition about as in megadon, decidedly heavier than in yucatanensis. 
Similar to that of O. talamancx, but narrower; zygomata less squarely 
spreading, the sides more divergent anteriorly; frontal region nar- 
rower, the supraorbital ridges weakly developed; parietals with 
lateral wings less developed below temporal ridges; interparietal 
smaller. Contrasted with that of 0. melanotis, the skull is larger, 
the upper outline more arched over anterior part of frontals; brain- 
case relatively lower and natter; frontals relatively narrower pos- 
teriorly; interparietal smaller. 

Measurements. — Type: Total length, 277; tail vertebrae, 141; hind 
foot, 32.5. Average of seven adult topo types: 255 (240-270); 136 
(125-145); 31.9 (30-33). Slull (average of 7 adults— type and 6 
topotypes): Greatest length, 31.4 (30.8-33.3); zygomatic breadth, 
16.1 (15.4-17.4); interorbital breadth, 5 (4.6-5.2); width of braincase, 
11.5 (11.4-11.7); nasals, 12.8 (11.5-13.8); anterior palatine foramina, 
5.2 (4.4-6); palatal bridge, 6.6 (6.3-7.3); upper molar series, 4.3 
(4.1-4.4). 



54 NORTH AMERICAN FAUNA. [No. 43. 

Remarks. — While 0. r. rostratus is distinguished by somewhat 
larger general size, and the greater breadth of the braincase is an 
especially noticeable cranial feature, the wide range of individual 
variation shown in large series of specimens renders the smaller 
examples difficult to separate from some of those of 0. r. megadon. 
Specimens from Pasa Nueva and Achotal, Vera Cruz, are rather 
small and grade toward megadon. Those from Alta Mira, Tamaulipas, 
marking the extreme northern limit of the known range of rostratus, 
average slightly paler than typical examples. 

Specimens examined. — Total number, 48, as follows: 

Oaxaca: Santo Domingo (mountains near), 5. 

Puebla: Metlaltoyuca 14 (type and topotypes). 

Tamaulipas: Alta Mira, 5. 

Vera Cruz: Achotal, 8; 1 Motzorongo, 1; Pasa Nueva, 6; 2 San Carlos, 9. 1 

ORYZOMYS ROSTRATUS MEGADON Merriam. 
Tabasco Rice Rat. 

(PI. II, figs. 5, 5a.) 

Oryzomys rostratus megadon Merriam, Proc Washington Acad. Sci., III, p. 294, July 
26, 1901. 

Type locality. — Teapa, Tabasco, Mexico. 

Type. — No. 99978, <? old, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, March 24, 1900. 

Geographic distribution. — Heavily forested coastal plains and lower 
mountain slopes in Tabasco and Campeche; altitudinal range from 
sea level to at least 500 feet; Humid Lower Tropical Zone. 

General characters.- — Closely resembling O. r. rostratus and 0. r. 
yucatanensis, but color slightly richer, more tawny than either ; skull 
smaller than that of rostratus and larger than that of yucatanensis. 

Color. — About like that of 0. r. rostratus, but slightly darker and 
richer, more intense ochraceous-tawny. 

Skull. — Similar to that of 0. r. rostratus, but smaller and narrower; 
differing from that of 0. r. yucatanensis mainly in larger size and 
heavier dentition. 

Measurements. — Type: Total length, 272; tail vertebrae, 140; 
hind foot 32.5. Average of 2 adult topotypes: 236.5 (236-237) ; 121 
(120-122); 30 (29-31). Skull (average of 3 adults— type and 2 
topotypes): Greatest length, 30.5 (29.1-32.3); zygomatic breadth, 
15.6 (15-16.5); interorbital breadth, 4.6 (4.4-4.9); - width of brain- 
case, 10.6 (10.5-10.7); nasals, 11.5 (11.3-12); anterior palatine 
foramina, 4.9 (4.6-5.1); palatal bridge, 6.3 (6-6.8); upper molar 
series, 4.3 (4.1-4.6). 

1 Collection Field Mus. Nat. Hist. s Collection Amer. Mus. Nat. Hist. 



1918.] ORYZOMYS MELANOTTS GROUP. 55 

Remarks. — 0. r. megadon appears to be a rather localized form. A 
series of specimens from Apazote, Campeche, agree most closely, as a 
whole, with typical megadon, but some are rather pale in color, and in 
the reduced size of molar teeth also approach 0. r. yucatanensis. 
Specimens examined. — Total number, 18, as follows: 
Campeche: Apazote, 11; Champoton, 1. 
Tabasco: Teapa 6 (type and topotypes). 

ORYZOMYS ROSTRATUS YUCATANENSIS Merriam. 
Yucatan Rice Rat. 

(PI. II, figs. 6, 6a.) 

Oryzomys yucatanensis Merriam, Proc. Washington Acad. Sci., Ill, p. 294, July 26, 
1901. 

Type locality. — Chichen Itza, Yucatan, Mexico. 

Type. — No. 108139, S adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, February 9, 1901. 

Geographic distribution. — Forested lowlands of Yucatan and north- 
ern Quintana Roo ; mainly Arid Lower Tropical Zone. 

General characters. — Similar to 0. r. rostratus and 0. r. megadon but 
somewhat smaller than either; skull small, with very small molar 
teeth. 

Color. — Much as in 0. r. rostratus but decidedly paler, the general 
tone ochraceous-buff instead of rich ochraceous-tawny. 

Skull. — Most closely resembling that of 0. r. megadon, but smaller, 
with short anterior palatine foramina and decidedly smaller molar 
teeth. 

Measurements. — Type: Total length, 235; tail vertebrae, 119; 
hind foot, 32. Adult from Puerto Morelos, Quintana Roo, 255; 136; 
31. Skull (type): Greatest length, 28.7; zygomatic breadth, 15; 
interorbital breadth, 4.6; width of braincase, 10.8; nasals, 11.8; ante- 
rior palatine foramina, 4.2 ; palatal bridge, 5.9 ; upper molar series, 3.7. 

Remarks. — This rather small pale form of 0. rostratus apparently 
passes into 0. r. megadon in southern Campeche. The specimens 
from Apazote are referred to megadon, with which the majority agree 
most closely, but several are indistinguishable in color from 0. f. 
yucatanensis, and, as they present no wide departure in cranial 
details, might be assigned to that form but for the presence of the 
larger or darker examples in the same series. 

Specimens examined. — Total number, 5, as follows: 

Quintana Roo: Puerto Morelos, 1. 

Yucatan: Chichen Itza, 4 (type and topotypes). 



56 



NORTH AMERICAN FAUNA. 



[No. 43. 



Oryzomys alfaroi Group. 

Geographic distribution. — Mountainous districts from northern 
Puebla, Mexico, south through Central America and northwestern 
South America to Ecuador; altitudinal range, from about 1,000 to 
10,000 feet, mainly in Humid Upper and Lower Tropical Zones 
(fig. 5). 

General characters. — Size small; form slender; tail usually longer 
than head and body, thinly and rather indistinctly haired; ears 
large and conspicuous, thinly clothed externally and internally with 
short, fine, blackish hairs; general pelage usually rather short and 
harsh, becoming longest in f orms'ranging at high elevations ; vibrissa? 




1. O. a. alfaroi. 7. O.a. saturatior. 

2. O.a.dariensis. 8. O.a.chapmani. 

3. O. a. angusticeps. 9. O. a. dilutior. 

4. O.a.rhabdops. 10. O.guerrerensis. 

5. O. a. caudatus. 11. O. hylocetes. 

6. O. a. palatinus. 
Fig. 5. — Geographic distribution of the Oryzomys 

alfaroi group. 

about as long as head; toes of hind feet inconspicuously webbed at 
base, the longest three bearing tufts of silvery whitish bristles which 
project beyond the ends of the claws as in the 0. talamancse group; 
claws short and recurved, compressed and sharp pointed. General 
color dark, the upperparts usually near ochraceous-tawny, heavily 
mixed with black; underparts dull whitish or pale buffy. 

Skull. — Small and delicate in structure, with rostrum elongated, 
the braincase moderately broad and rather low; maxillary arm of 
zygoma weakly developed; outer wall of antorbital foramen with 
anterior border rounded, the antorbital notch only moderately deep 
as viewed from above; nasals and premaxillaB ending posteriorly in 
about the same plane ; frontal region broad, the lateral margins very 
slightly elevated to form delicate supraorbital ridges; temporal 
ridges slightly developed anteriorly along parieto-squamosal sutures, 



1918.] ORYZOMYS ALFAKOI GROUP. 57 

usually becoming obsolescent posteriorly in crossing lateral wings of 
parietals; interparietal large, somewhat irregularly pointed-elliptical 
in form; anterior palatine foramina short and broad, shorter than 
palatal bridge and not usually reaching anterior plane of first molars; 
palatal pits rather large and rounded; sphenopalatine vacuities 
present but small; interpterygoid fossa broad; audita! bullae small ; 
angle of mandible short and broad, the inferior border even more 
strongly turned inward than in the 0. talamancse group; coronoid 
process with free portion short owing to high connecting ridge reach- 
ing nearly to summit of condyle ; molars small, the reentrant angles 
shallow and the crown arrangement, especially the enamel islands in 
the second upper molars, much as in the 0. palustris group, but inner 
reentrant angles in first upper molars broader. More reliable dis- 
tinguishing characters are presented by the skull and teeth; the 
molars are less deeply cleft by reentrant angles in the 0. alfaroi 
group, and the enamel island present at the postero-internal base of 
the paracone of the second upper molar (moderately worn) is absent 
in the talamancse group. 

Remarks. — This group, comprised of a series of small, dark-colored, 
closely allied, and somewhat localized forms typified by 0. alfaroi, 
extends in an irregular chain along the backbone of the continent 
from southern Mexico to northwestern South America, one form at 
least ranging as far south as Ecuador. While the 0. palustris and 
0. talamancse groups range mainly at low elevations, or are restricted 
to the vicinity of water at the higher levels, alfaroi and its allies often 
inhabit well-drained but moist mountain slopes, where they com- 
monly ascend to high elevations. 

0. alfaroi and its relatives approach the members of the 0. talamancse 
group in general external characters, and the two partially overlap in 
geographic range. In appearance individuals may be much alike, 
both groups having large ears, appearing nearly naked, and slender 
limbs; forms of the alfaroi group are, however, usually smaller and 
decidedly darker in general color; the ears are smaller, but not mark- 
edly different in color. 

Key to Species and Subspecies of the O. alfaroi Group. 

a 1 . Zygomata broader posteriorly than anteriorly. 

ft 1 . Upperparts darker ochraceous-buff or ochraceous-tawny. 
c 1 . Upperparts less intense ochraceous-buff or ochraceous-tawny. 
d 1 . Median dorsal area not distinctly blackish. 
e l . Size smaller; total length less than 240. 
f 1 . Skull broader; zygomatic breadth 13 or more. 
g 1 . Anterior palatine foramina usually 4 or more. 
ft 1 . Rostrum more massive. (Northern Puebla.) O. a. dilutior (p. 68). 
ft 2 . Rostrum less massive. (Central Vera Cruz.) 

O. a. chapmani (p. 67). 



58 NORTH AMERICAN FAUNA. [No. 43. 

g 2 . Anterior palatine foramina usually less than 4. (Mountains of Hon- 
duras; Nicaragua; Costa Rica; western Panama.) 

O. a. alfaroi (p. 59). 
f 2 . Skull narrower; zygomatic breadth less than 13. (Mountains of south- 
ern Tabasco and northwestern Chiapas.). . . O. a. palatinus (p. 65). 
e 2 . Size larger; total length 240 or more. 
f 1 . Pelage short; tail about 140. (Mountains of northeastern Oaxaca.) 

O. a. caudatus (p. 64). 
f 2 . Pelage long; tail less than 135. (High mountains of southwestern Gua- 
temala and of central and southern Chiapas.) 

O. a. angusticeps (p. 62). 
d?. Median dorsal area distinctly blackish. (Mountains of northern Chiapas.) 

/ O. a. saturatior (p. 66). 

c 2 . Upperparts more intense ochraceous-tawny. (Eastern Panama.) 

O. a. dariensis (p. 61). 
b 2 . Upperparts paler ochraceous-buff or ochraceous-tawny. (Pacific slope of Sierra 

Madre in Guerrero and Oaxaca.) O. guerrerensis (p. 69). 

a 2 . Zygomata broader anteriorly than posteriorly. 

b l . Size larger; total length 225 or more. (Southeastern Guatemala.) 

O. a. rhabdops (p. 63). 
b 2 . Size smaller; total length less than 225. (Southern Chiapas.) 

O. hylocetes (p. 70). 

ORYZOMYS ALFAROI Allen. 

[Synonymy under subspecies.] 

Geographic distribution. — Heavily forested mountain slopes from 
northern Puebla south through southern Mexico and Central America, 
at least to Colombia and Ecuador; altitudinal range from about 1,000 
to 10,000 feet, mainly in Humid Upper and Lower Tropical Zones. 

General characters. — Size small; color dark; skull light and rather 
delicate in structure. Similar in general to 0. guerrerensis, but color 
much darker, and skull differing in detail. (For additional general 
characters see under O. alfaroi group.) 

Color. — Fresh pelage: Upperparts varying from dark ochraceous- 
buff to ochraceous-tawny, tawny, russet, or cinnamon-brown, usually 
heavily mixed with black, this color often predominant over dorsum, 
the lighter element purest and becoming more or less ochraceous- 
buffy on cheeks, shoulders, and sides; underparts dull white or buffy, 
thinly overlying the dark plumbeous basal color; nose blackish; inner 
and outer sides of ears thinly clothed with very short black hairs, 
light ochraceous-buffy subauricular spots present in some forms, 
absent in others; feet dull whitish, the elongated silvery tufts on toes 
of hind feet projecting beyond points of longer claws; feet (epidermis) 
yellowish, thinly clothed above with very short glossy white hairs; tail 
nearly naked, brownish or blackish above, yellowish below basally, 
becoming dark all around toward tip. Young (in first pelage) : Upper- 
parts blackish, finely and inconspicuously mixed with ochraceous- 
tawny; underparts darker than in adults, the dark basal color thinly 



1918.] OKYZOMYS ALFAROI GROUP. 59 

overlaid with white; feet varying from dull whitish to brownish; tail 
blackish. 

Skull. — Similar in general to that of 0. guerrerensis, but more 
elongated; braincase higher, less flattened. In the anteriorly spread- 
ing zygomata, one form of 0. alfaroi (0. a. rhabdops) approaches 
0. hylocetes, but the latter is much smaller, with very small teeth, and 
present material seems to indicate specific distinctness. (For addi- 
tional characters see under 0. alfaroi group.) 

Remarks. — All the North American members of the 0. alfaroi group, 
excepting 0. hylocetes and 0. guerrerensis, and the South American 
forms O.palmirse and 0. gracilis, appear to be assignable subspecifically 
to 0. alfaroi. 1 While complete intergradation may not be shown by 
the material examined, the more essential characters prevail with 
such uniformity throughout the series as to leave little room for doubt 
of its existence. The accession of new material may not improbably 
show that liylocetes and guerrerensis are also geographic races of 
alfaroi. Typical alfaroi presents ' closer resemblance to the geo- 
graphically distant race 0. a. chapmani, of Vera Cruz, than to the 
annectent forms inhabiting the high mountains of Chiapas and 
Guatemala. This resemblance between the more widely removed 
subspecies may be due to the fact that the intermediate races occupy 
more diversified areas, most of them having ascended to high eleva- 
tions where peculiar environmental conditions would conduce to 
differentiation. 

OKYZOMYS ALFAROI ALFAROI (Allen). 

Alfaro Rice Rat. 

(PI. ill, figs. 1, la.) 

Hesperomys (Oryzomys) alfaroi Allen, Bull. Amer. Mus. Nat. Hist., Ill, p. 214, April 

17, 1891. 
Oryzomys alfaroi Allen, Abstr. Proc. Linn. Soc. New York, 1893-94, p. 36, July. 20, 

1894. 
Oryzomys alfaroi incertus Allen, Bull. Amer. Mus. Nat. Hist., XXIV, p. 655, October 

13, 1908. Type from Rio Grande, Nicaragua, No. 28584, S ad., Amer. Mus. Nat. 

Hist.; collected by W. B. Richardson, March 28, 1908. 

Type locality. — San Carlos, Costa Rica. 

Type. — No. ff-jHr, 9 subadult, American Museum of Natural His- 
tory; collected by Anastasio Alfaro, December, 1888. 

Geographic distribution. — Heavily forested mountainous portions 
of Honduras, Nicaragua, Costa Rica, and western Panama; alti- 
tudinal range from about 1,000 to 4,000 feet; mainly in Humid 
Lower Tropical Zone. 

1 The South American forms will therefore stand as follows: 

Oryzomys alfaroi palmirse Allen Miraflores, Colombia. 

Oryzomys alfaroi gracilis Thomas Concordia, Medellin, Colombia. 



60 NORTH AMERICAN FAUNA. [No. 43. 

General characters. — A rather small form with short pelage, resem- 
bling 0. a. dariensis, but coloration duller, lessrufescent; skull differ- 
ing in slight details. Very similar in general to 0. c. chapmani, but 
color usually duller; skull with shorter anterior palatine foramina. 
Smaller than 0. a. angusticeps and 0. a. rhabdops, with shorter pelage 
than either. 

Color. — Upperparts varying from ochraceous-buff to dull ochra- 
ceous-tawny, heavily mixed with black. (Other colors as given 
under 0. alfaroi.) 

STcull. — Size medium for the group, rather narrow and elongated 
with narrowly spreading zygomata and short, wide anterior palatine 
foramina. In general form very similar to that of 0. a. dariensis, 
but braincase and frontal region usually broader; apparently differing 
from those of 0. a. angusticeps and 0. a. chapmani most noticeably 
in shorter anterior palatine foramina. Compared with that of 0. a. 
rhabdops the skull is decidedly narrower, with smaller, less inflated 
braincase, and zygomata much less divergent anteriorly. 

Measurements. — Average of 6 adults from Tuis, Costa Kica: Total 
length, 210 (188-221); tail vertebras, 109 (102-114); hind foot, 26.3 
(26-27). STcull (average of same) : Greatest length, 26.7 (25.6-28.3) ; 
zygomatic breadth, 13.5 (13.2-14.1); interorbital breadth, 5.2 (5.1- 
5.5); width of braincase, 10.5 (10.1-10.8); nasals, 10.9 (10.3-11.5); 
anterior palatine foramina, 3.9 (3.5-4.9) ; palatal bridge, 5.4 (5.3-5.7) ; 
upper molar series, 3.7 (3.6-3.8). 

Remarks. — While a considerable gap separates the known geo- 
graphic ranges of 0. a. alfaroi and 0. a. dariensis, these forms exhibit 
such close approach in size, color, and cranial details that intergrada- 
tion may be safely assumed. Specimens from Yaruca, Honduras, are 
referable to alfaroi, but in the larger size shown by some examples are 
not very unlike O. a. angusticeps and 0. a. rhabdops. Three speci- 
mens from Managua, Nicaragua, recorded by Thomas * as 0. gracilis, 
were probably assignable to alfaroi. 

Allen's "0. a. incertus" was based on rather richly colored speci- 
mens which are well within the range of individual variation exhibited 
by typical alfaroi, as the accession of additional material indicated 
to him. 2 

The general range of alfaroi overlaps that of 0. talamancse, and as it 
may closely resemble that species in general appearance some con- 
fusion of the two has resulted. While many examples may be 
inseparable in color, alfaroi is a smaller animal than talamancse. The 
skull of alfaroi is distinguished by its smaller size and more delicate 
structure, the maxillary arm of the zygoma is more slender and the 
teeth are decidedly smaller; the second upper molar has a large, 

i Thomas, Oldfield, Ann. Mag. Nat. Hist., ser. 6, XVI, p. 57, July, 1895. 
» Allen, J. A., Bull. Amer. Mus. Nat. Hist., XXVIII, p. 99, Apr. 30, 1910. 



1918.] 



ORYZOMYS ALFAROI GROUP. 61 



elongated enamel island at the posterointernal base of the paracone, 
and the internal reentrant angle extends less than halfway across 
the crown of the moderately worn tooth, while in talamancx the 
enamel island mentioned is absent or represented only by a small 
island near the apex of the reentrant angle, which in this species 
reaches halfway across the molar crown. 

Specimens examined. — Total number, 86, as follows : 

Costa Rica: San Carlos, 3 (type and topotypes); l Tuis, 15; 1 exact locality 

unknown, 7. 1 
Honduras: Yaruca, 19. 2 
Nicaragua: Chontales, 6; 1 Jalapa, 2; 1 Jicaro, l; 1 Jinotega, l; 3 Rio Coco, 5; 1 

Rio Grande, 3 (including type of "incertus"); 1 San Juan, 5; 1 Tuma, 2; 1 

Uluce, 2; 1 exact locality unknown, l. 1 
Panama: Boquete, 14.* 

ORYZOMYS ALFAROI DARIENSIS Goldman. 
Darien Rice Rat. 

(PI. Ill, figs. 2, 2a; PI. V, figs. G; PI. VI, figs. 3, 3a.) 

Oryzomys alfaroi dariensis Goldman, Proc. Biol. Soc. Washington, XXVIII, p. 128, 
June 29, 1915. 

Type locality. — Cana, eastern Panama (altitude 2,000 feet). 

Type. — No. 178660, $ adult, United States National Museum 
(Biological Survey collection); collected by E. A. Goldman, March 4, 
1912. 

Geographic distribution. — Heavily forested mountain slopes in 
eastern Panama at 2,000 feet altitude, and probably adjacent por- 
tions of Colombia; Humid Lower Tropical Zone. 

General characters. — A small form closely allied to 0. a. alfaroi; 
color of upperparts richer, more rufescent; skull usually narrower. 
Similar to the South American forms O. a. gracilis and to 0. a. pal- 
mirse, but color more rufescent and skull differing in detail. 

Color. — Fresh pelage: Upperparts between ochraceous-tawny and 
tawny, finely mixed with black; becoming paler and ochraceous- 
buffy on cheeks, shoulders, and lower part of sides; underparts, feet, 
and tail as given under O. alfaroi. 

STcull. — About like that of O. a. alfaroi, but braincase and frontal 
region usually narrower. Closely resembling that of O. a. palmirx, 
of South America, but shorter, with more widely spreading zygomata 
and smaller teeth. Compared with Ecuadorean specimens assumed 
to represent O. a. gracilis, the skull is more massive, with more 
widely spreading zygomata. 

1 Collection Amer. Mus. Nat. Hist. 

! Fourteen specimens in Mus. Comp. Zool.; 2 in Field Mus. Nat. Hist. 

» Collection Field Mus. Nat. Hist. 

* Eleven specimens in Mus. Comp. Zool.; 2 in Field Mus. Nat. Hist.; 1 in Amer. Mus. Nat. Hist. 



62 NORTH AMERICAN FAUNA. [No. 43. 

Measurements. — Type: Total length, 203; tail vertebrae, 107; hind 
foot, 25.5. Average of 5 adult topotypes: 220 (212-226); 113 (107- 
117) ; 24.6 (23-26). SJcull (average of 6 adults, type and 5 topotypes) 
Greatest length, 27.3 (26.5-27.8); zygomatic breadth, 14.3(14-14.7) 
interorbital breadth, 4.8 (4.6-5); width of braincase, 10.3 (10-10.5) 
nasals, 11.1 (10.6-11.4); anterior palatine foramina, 4.3 (4-4.8) 
palatal bridge, 5.6 (5-5.9); upper molar series, 3.7 (3.6-3.9). 

Remarks. — This small, slender rice rat differs from typical 0. a. 
alfaroi, of Costa Rica, mainly in richer, more tawny coloration. It is 
closely allied to the Colombian form described as 0. palmirse and the 
latter is clearly assignable to subspecific rank, if it does not prove 
to be identical with 0. a. gracilis, the type of which came from 
farther north in the Cauca Valley. Comparison with specimens from 
northern Ecuador, assigned to gracilis by Mr. Oldfield Thomas, and 
reference to the original description of that species indicate that the 
two are very nearly related. The description of the color of gracilis, 
however, seems to apply to the Ecuadorean specimens, or to palmirse, 
rather than to the Darien animal. Moreover, the skull 0. a. dariensis 
is distinguished from that of gracilis, as here understood, by the 
greater lateral expansion of the zygomata. 

0. talamancse also occurs at the type locality of dariensis and the 
two are superficially much alike. The smaller size, especially the 
smaller hind foot, usually distinguishes dariensis externally, while the 
skull is smaller, more delicate in structure, the maxillary arm of the 
zygoma more slender and the molar teeth much smaller; the second 
upper molar has a large elongated enamel island at the postero-inter- 
nal base of the paracone, and the internal reentrant angle extends less 
than half way across the crown of the moderately worn tooth, much 
as in 0. palustris. In talamancse the enamel island is absent and the 
reentrant angle reaches halfway across the molar crown. 

Specimens examined. — Eleven, from type locality. 

ORYZOMYS ALFAROI ANGUSTICEPS Merriam. 

Volcan Santa Maria Rice Rat. 

Oryzomys angusticeps Merriam, Proc. Washington Acad. Sci., Ill, p. 292, July 26, 
1901. 

Type locality. — Volcan Santa Maria, Guatemala (altitude, 9,000 feet). 

Type. — No. 76816, c? adult, United States National Museum (Bio- 
logical Survey collection); collected by E. W. Nelson and E. A. Gold- 
man, January 22, 1896. 

Geographic distribution. — Heavily forested slopes of high moun- 
tains in southwestern Guatemala and central and southern Chiapas; 
altitudinal range from about 8,000 to 9,500 feet. 

General characters. — Similar to 0. a. alfaroi but larger, with longer 
pelage; anterior palatine foramina longer. Closely approaching 
0. a. rhabdops in size, but color darker and skull differing in detail. 



1918.] ORYZOMYS ALFAROI GROUP. 63 

Color. — Lighter element in upperparts pale cinnamon brownish, 
heavily mixed with black, the general tone appreciably darkened by 
the dark plumbeous basal color of the long pelage; underparts and 
sides of muzzle light ochraceous-buffy; ears, feet, and tail as given 
under 0. alfaroi. 

Skull. — Similar to that of 0. a. alfaroi, but larger, zygomata more 
squarely spreading anteriorly, the sides more nearly parallel; frontal 
region usually narrower; anterior palatine foramina longer. Ap- 
proaching that of 0. a. rhabdops in size, but zygomata less divergent 
anteriorly. 

Measurements. — Type: Total length, 245; tail vertebrse, 134; hind 
foot, 29. STcull (type): Greatest length, 28.6; zygomatic breadth, 
14; interorbital breadth, 4.6; width of braincase, 10.3; nasals, 10.6; 
anterior palatine foramina, 5; palatal bridge, 5.4; upper molar 
series, 3.6. 

Remarks. — Aside from its geographic neighbor, 0. a. rJiabdops, 
no other North American form of the 0. alfaroi group attains so high 
an altitude; like rhabdops it has developed a longer pelage than 
forms ranging at lower elevations. The skull of the type of 0. a. 
angusticeps seems to be abnormally narrow; the topotypes are all 
young and their skulls of little value for comparative purposes. An 
apparent tendency toward greater breadth shown in the skulls of 
specimens from San Cristobal and Pinabete, Chiapas, is believed to 
be within the probable range of individual variation; on the other 
hand, they do not differ appreciably from some skulls of rliabdops, and 
suggest intergradation. In size and color the Chiapas examples are 
very similar to the topotypes. 

Specimens examined. — Total number, 16, as follows: 

Chiapas: Pinabete, 2; San Cristobal, 10. 

Guatemala: Volcan Santa Maria 4 (type and topotypes). 

ORYZOMYS ALFAROI RHABDOPS Merriam. 

Calel Rice Rat. 
(PI. Ill, figs. 3, 3a.) 
Oryzomys rhabdops Merriam, Proc. Washington Acad. Sci., Ill, p. 291, July 26, 1901. 

Type locality. — Calel, Guatemala (altitude, 10,000 feet). 

Type. — No. 76813, S adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, January 15, 1896. 

Geographic distribution. — -Known only from type locality, at about 
10,000 feet altitude on the upper slope of the high mountains in 
southwestern Guatemala. 

General characters. — Size large; pelage very long for a member of the 
O. alfaroi group, the longer hairs on dorsum reaching about 11 mm.; 



64 NORTH AMERICAN FAUNA. [No. 43. 

color rather pale, upperparts near ochraceous-tawny; skull broad 
with anteriorly divergent zygomata. Similar in general to 0. a. 
angusticeps, but color paler, and skull differing in detail. 

Color. — Fresh pelage: Upperparts near ochraceous-tawny, richest 
and purest on cheeks, shoulders, and sides, the back darkened by 
black hairs, but less copiously than in most forms of 0. alfaroi; sides 
of muzzle and underparts in general light ochraceous-buffy; nose 
deep black, except extreme tip which, like lips, is whitish; ears black, 
with more or less distinct light ochraceous-buffy subauricular spots; 
feet and tail as given under 0. alfaroi. Young (in first pelage) : 
Upperparts blackish, inconspicuously tinged with ochraceous-tawny; 
underparts plumbeous, lightly overlaid with light ochraceous-buff. 

Skull. — Skull broad, with large inflated braincase and anteriorly 
diverging zygomata. Similar to that of 0. a. angusticeps, but 
zygomata wider anteriorly than posteriorly (sides more nearly parallel 
in angusticeps). In general form, especially the anteriorly spreading 
zygomata, similar to that of 0. hylocetes, but much larger, with shorter 
anterior palatine foramina. 

Measurements. — Type: Total length, 255; tail vertebrae, 141; hind 
foot, 29.5. Adult topotype: 255; 138; 28.5. Skull (average of 
two adults — type and topotype) : Greatest length, 28.6 (28.4-28.8) ; 
zygomatic breadth, 14.8 (14.6-15.1); interorbital breadth, 4.8 
(4.S-4.9); width of braincase, 10.9(10.8-11.1); nasals, 11 (10.5-11.5); 
anterior palatine foramina, 4.3 (4.2-4.5) ; palatal bridge, 5.7 (5.5-5.9) ; 
upper molar series, 3.8 (3.7-3.9). 

Remarks. — No other North American form of the genus is known to 
range 10,000 feet above sea level, but a near geographic neighbor, 
0. a. angusticeps, which has also pushed up above the Tropical Zones 
on mountains of similar elevation, approaches and may equal this 
altitude. The two forms are evidently closely allied. Distinguishing 
cranial characters are developed mainly in adults, the skulls of most 
of the younger examples being apparently inseparable. In the 
remarkable anterior expansion of the zygomata the skull of O. a. 
rhabdops resembles that of 0. hylocetes, but the latter appears to be a 
very distinct form. 

Specimens examined. — Fourteen, from type locality. 

ORYZOMYS ALFAROI CAUDATUS Merriam. 
Comaltepec Rice Rat. 

(PI. Ill, figs. 4, 4a.) 
Oryzomys chapmani caudatus Merriam, Proc. Washington Acad. Sci , III, p. 289, July 
26, 1901. 

Type locality. — Comaltepec, Oaxaca, Mexico (altitude 3,500 feet). 

Type. — No. 68641, <? adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, July 31, 1894. 



1918.] OEYZOMYS ALFAROI GROUP. 65 

Geographic distribution. — Mountains of northeastern Oaxaca; alti- 
tudinal range from 3,500 to 6,500 feet; Humid Upper Tropical Zone. 

General characters. — Size largest of the 0. alfaroi group; color 
dark; pelage short. Similar in general to 0. a. chapmani, but 
decidedly larger, with the lighter colored admixture in upperparts of 
ochraceous-buffy instead of ochraceous-tawny; skull larger and more 
massive than usual in the alfaroi group. 

Color. — Fresh pelage: Upperparts ochraceous-buffy, heavily mixed 
with black, the latter color predominating, especially along lower part 
of back; underparts and sides of muzzle light ochraceous-buffy; 
lips and chin whitish; nose black; ears, feet, and tail as given under 
0. alfaroi. 

Skull. — Size very large and structure heavy for 0. alfaroi. Very 
similar in general form to that of 0. a. chapmani, but much larger 
with more swollen rostrum. 

Measurements. — Type: Total length, 257; tail vertebrae, 141; hind 
foot, 30. Skull (type) : Greatest length, 29; zygomatic breadth, 15; 
interorbital breadth, 5; width of braincase, 11.3; nasals, 11.4; 
anterior palatine foramina, 4.8; palatal bridge, 5.2; upper molar 
series, 3.9. 

Remarks. — 0. a. caudatus was based on a single specimen which 
seems to indicate a subspecies still larger than 0. a. rhabdops and 
0. a. angusticeps, but the extent of individual variation remains to be 
determined. Although not very widely differing from angusticeps it 
seems to be most closely allied to 0. a. chapmani, the greater size 
being the chief distinguishing character. An immature example 
from 6,500 feet altitude at Totontepec, Oaxaca, is referable to the 
same form. 

Specimens examined. — Total number, 2, as follows: 
Oaxaca: Comaltepec, 1 (type); Totontepec, 1. 

ORYZOMYS ALFAROI PALATINUS Merriam. 
Teapa Rice Rat. 
(PI. Ill, figs. 5, 5a.) 
Oryzomys palatinus Merriam, Proc. Washington Acad. Sci., Ill, p. 290, July 26, 1901. 

Type locality. — Teapa, Tabasco, Mexico (altitude, 3,000 feet). 
Type.— No. 99977, 9 adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, April 1, 1900. 

Geographic distribution. — Forested mountain slopes in southern 
Tabasco and northwestern Chiapas; known altitudinal range from 
about 3,000 to 3,500 feet; Humid Upper Tropical Zone. 

General characters. — A small form, similar to O. a. chapmani in 
color, but skull narrower, lighter, and more like that of the darker 
subspecies O. a. saturatior. 

14521°— 18 5 



66 NORTH AMERICAN FAUNA. [No. 43. 

Color.— Type in fresh pelage: Upperparts rich ochraceous-tawny, 
finely and uniformly lined with black from nose over top of head 
and back to near base of tail, the tawny element becoming purer and 
paler on cheeks and along lower part of sides; underparts and feet 
dull whitish; tail dark brownish, except basal half of under side, 
which is dull yellowish. Specimens from Ocuilapa, Chiapas, appar- 
ently referable to this form, are indistinguishable in color from 0. a. 
chapmani. 

SJcull. — Size small, general form long and narrow, structure light. 
Scarcely distinguishable from some of the narrower skulls of 0. a. 
saturatior; zygomata very slender as in that form. Similar in 
general to that of 0. a. chapmani, but narrower, maxillary arm of 
zygoma more slender. 

Measurements. — Type: Total length, 209; tail vertebrae, 106; 
hind foot, 25. Adult from Tumbala, Chiapas: 222; 116; 27. STcull 
(type): Greatest length, 26.7; zygomatic breadth, 12.6; interorbital 
breadth, 4.8; width of braincase, 10.2; nasals, 10.2; anterior palatine 
foramina, 3.8; palatal bridge, 5.5; upper molar series, 3.8. 

Remarks. — In 0. a. palatinus the general color of 0. a. chapmani 
seems to be combined with the narrower, lighter skull of the darker- 
colored form 0. a. saturatior. The scanty material available indicates 
that intergradation of palatinus with saturatior is probable, since the 
differential characters are slight and both forms inhabit the northern 
slope of the same mountain range. 

Specimens examined. — Total number, 5, as follows: 

Chiapas: Ocuilapa, 3; Tumbala, 1. 
Tabasco: Teapa, 1 (type). 

ORYZOMYS ALFAROI SATURATIOR Merriam. 

Dusky Rice Rat. 

Oryzomys chapmani saturatior Merriam, Proc. Washington Acad. Sci., Ill, p. 290, 
July 26, 1901. 

Type locality. — Tumbala, Chiapas, Mexico (altitude, 5,000 feet). 

Type. — No. 76183, ? adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, October 23, 1895. 

Geographic distribution. — Forested northern slope of mountains of 
northern Chiapas, at 5,000 feet altitude, limits of range unknown; 
Humid Upper Tropical Zone. 

General characters. — Size rather small — about as in O. a. alfaroi and 
0. a. chapmani; color very dark. Closely allied to O. a. palatinus 
and chapmani, but darker than either, the back blackish and under- 
parts usually distinctly buffy. 

Color. — Upperparts cinnamon brownish, very heavily mixed with 
black, the top of head and median dorsal area blackish ; underparts, 



1918.] ORYZOMYS ALFAROI GROUP. 67 

except chin and throat, varying from light to dark ochraceous-buff ; 
chin and throat more or less distinctly whitish, the fur in some 
examples pure white to roots along a narrow median line; ears black; 
feet brownish to toes, the toes dull yellowish, those of hind feet bear- 
ing the silvery terminal bristles present throughout the group; tail 
usually dark all around, but in some examples yellowish on under 
side at base. Young (in first pelage) : Upperparts nearly black, the 
cinnamon brownish hairs inconspicuous and restricted mainly to 
the sides; underparts dark plumbeous, very thinly overlaid with 
ochraceous-buff; feet and tail blackish. 

Skull. — Size small and structure light. About like that of 0. a. 
palatinus, but usually broader. Similar to that of 0. a. chapmani, 
but averaging slightly smaller, with less swollen rostrum and more 
slender zygomata. 

Measurements. — Type: Total length, 218; tail vertebras, 120; hind 
foot, 25.5. Average of 10 adults (type and 9 topotypes) : 208.4 (195- 
225); 111.1 (105-122); 26.1 (25-28). Skull (average of 3 adults- 
type and 2 topotypes): Greatest length, 25.3 (25.2-25.4); zygomatic 
breadth, 12.8 (12.4-13.5); interorbital breadth, 4.7 (4.5-4.8); width 
of braincase, 10.1 (10-10.3); nasals, 9.8 (9.5-10.2); anterior palatine 
foramina, 4.2 (3.7-4.5); palatal bridge, 5.1 (4.9-5.3); upper molar 
series, 3.6 (3.5-3.6). 

Remarks. — The very dark colors of O. a. saturatior appear to be 
directly due to environmental conditions. The area inhabited by 
this form is heavily forested and excessively humid ; torrential rains 
occur during the wet season and fog enshrouds the mountains nearly 
throughout the year. Two specimens from Tumbala, with narrow 
skulls and slender rostra, approach O. a. palatinus, whose range is 
doubtless contiguous, and point to intergradation with that form. 

Specimens examined. — Seventeen, from type locality. 

ORYZOMYS ALFAROI CHAPMANI Thomas. 
Chapman Rice Rat. 

Oryzomys chapmani Thomas, Ann. Mag. Nat. Hist., ser. 7, I, p. 179, February, 1898. 

Type locality. — Jalapa, Vera Cruz, Mexico (altitude, 4,400 feet). 

Type.— 97.9.9.30, British Museum; collected by Frank M. Chap- 
man, March 31, 1897. 

Geographic distribution. — Forested eastern slopes of the Mexico 
plateau region in central Vera Cruz; known altitudinal range from 
4,400 to 6,000 feet; Humid Upper Tropical Zone. 

General characters. — Closely resembling O. a. alfaroi; size and 
color very similar, but general tone of upperparts averaging slightly 
richer, more tawny; cranial characters distinctive. Size about as in 
O. a. dilutior, but color slightly more tawny; skull much less massive; 



68 NORTH AMERICAN FAUNA. [No. 43. 

differing from 0. a. caudatus in much smaller size and more tawny 
color. 

Color. — Fresh pelage: Upperparts near ochraceous-tawny (becom- 
ing russet in some old adults), finely and abundantly mixed with 
black, the tawny element purer but somewhat paler and grading 
toward dark ochraceous-buff along cheeks and lower part of sides; 
underparts whitish or pale buffy, the plumbeous basal color showing 
through except over a narrow median area on throat and chest, 
where the fur in some specimens is pure white to roots; ears black, 
light subauricular spots usually present; feet and tail as given under 
0. alfaroi. Young (in first pelage) : Upperparts blackish, inconspicu- 
ously mixed with ochraceous-tawny; underparts darker than in 
adults, the dark basal color less heavily overlaid with white; feet 
and tail brownish (becoming paler in adults). 

Skull. — Size and general form much as in 0. a. alfaroi, but tending 
to be shorter and relatively broader, the zygomata more widely or 
squarely spreading anteriorly; frontal region usually narrower; ante- 
rior palatine foramina longer, commonly reaching anterior plane of 
first molars; dentition about the same. Similar to that of 0. a. 
dilutior, but less massive; rostrum less swollen and decurved; maxil- 
lary arm of zygoma more slender. Much like that of 0. a. caudatus, 
but much smaller. 

Measurements. — Average of 10 adult topotypes: Total length, 224 
(202-265); tail vertebras, 117.9 (108-132); hind foot, 25.1 (24-27). 
Skull (average of 7 adult topotypes) : Greatest length, 26.4 (25.4- 
27.3); zygomatic breadth, 13.7 (13.1-14.4); interorbital breadth, 4.7 
(4.4-4.8) ; width of braincase, 10.7 (10.2-11); nasals, 10.3 (9.6-10.9); 
anterior palatine foramina, 4.4 (4-4.7); palatal bridge, 5.3 (5-5.6); 
upper molar series, 3.8 (3.7-3.9). 

Remarks. — The general resemblance of O. a. chapmani to O. a. 
alfaroi is noteworthy, in view of their geographic separation and the 
occurrence of apparently annectent forms that differ considerably 
from both. The annectent forms, however, inhabit diversified areas, 
some having ascended to high elevations, where their differentiation 
has probably been the result of rather local environmental conditions. 

Specimens examined. — Total number, 25, as follows: 
Vera Cruz: Jalapa, 17 (type and topotypes); 1 Jico, 6; Mirador, 1; Teocelo, l. 2 

ORYZOMYS ALFAROI DILUTIOR Mereiam. 
Puebla Rice Rat. 

(PI. Ill, figs. 6, 6a.) 
Oryzomys chapmani dilutior Merriam, Proc. Washington Acad. Sci., Ill, p. 290, July 
26, 1901. 

Type locality. — Huauchinango, Puebla, Mexico (altitude 5,000 
feet). 

1 Thirteen in collection Amer. Mus. Nat. Hist. ' Collection Mus. Comp. Zool. 



1918.] ORYZOMYS ALFAROI GROUP. 69 

Type. — No. 93124, <? adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, January 10, 1898. 

Geographic distribution. — Eastern slope of Mexican plateau region 
at 5,000 feet altitude in northern Puebla; limits of range unknown; 
Humid Upper Tropical Zone. 

General characters. — Similar in size to 0. a. chapmani, but color 
slightly paler, less tawny; skull more massive. 

Color. — About as in 0. a. chapmani, but slightly paler, the general 
tone less tawny. 

SJcull. — Most like that of 0. a. chapmani, but more massive; ros- 
trum more swollen, the upper outline more strongly decurved; maxil- 
lary arm of zygoma heavier. 

Measurements. — Type: Total length, 223; tail vertebrae, 117; hind 
foot, 28. An adult topotype: 225; 125; 27.5. SJcull (type and adult 
topotype): Greatest length, 27.6, 27.4; zygomatic breadth, 15, 14.2; 
interorbital breadth, 4.9, 4.6; width of braincase, 10.8, 10.4; nasals, 
10.3, 10.5.; anterior palatine foramina, 4.6, 4.4; palatal bridge, 5.2, 5; 
upper molar series, 4, 4. 

Remarks. — The range of O. a. diluiior marks the northern limit of 
the O. alfaroi group. The subspecies appears to be a well-marked 
form, requiring close comparison only with 0. a. chapmani. 

Specimens examined. — Three, from type locality. 

ORYZOMYS GUERRERENSIS Goldman. 

Guerrero Rice Rat. 

(PI. Ill, figs. 7, 7a.) 

Oryzomys guerrerensis Goldman, Proc. Biol. Soc. Washington, XXXVIII, p. 127, 
June 29, 1915. 

Type locality. — Omilteme, Guerrero, Mexico (altitude 8,000 feet). 

Type. — No. 127517, <$ adult (molars moderately worn), United 
States National Museum (Biological Survey collection); collected 
by E. W. Nelson and E. A. Goldman, May 20, 1903. 

Geographic distribution. — Forested Pacific slope of Sierra Madre in 
Guerrero and Oaxaca, Mexico; altitudinal range from 3,000 to about 
8,000 feet; Humid Upper Tropical Zone. 

General characters. — Size small; color averaging palest of the North 
American forms of the O. alfaroi group. Similar in general to O. a. 
chapmani, but color paler and skull smaller and natter. 

Color. — Upperparts varying from dark ochraceous-buff to dark 
ochraceous-tawny, purest on cheeks, shoulders, and sides; the face, 
top of head, and back darkened by a moderate admixture of black 
hairs; underparts dull grayish white, the dark basal color of the fur 
everywhere showing through, except in 2 examples out of 8, in which 



70 NORTH AMERICAN FAUNA. [No. 43. 

the fur is pure white to roots along a narrow median line on chin and 
throat; outer and inner sides of ears well clothed with deep glossy- 
black hairs; feet whitish; tail brownish above, irregularly yellowish 
below to near tip, which is dusky all around. Young (in first pelage) : 
As in 0. a. chapmani. 

Skull. — Small and short, the braincase broad and somewhat flat- 
tened. Similar to that of 0. a. chapmani, but usually smaller and 
flatter; zygomata tending to curve evenly outward, the sides less 
nearly parallel; sides of rostrum more tapering anteriorly; ascending 
branches of premaxillse usually broader posteriorly; maxillary arm of 
zygoma more slender; incisors smaller. 

Measurements. — Type: Total length, 220; tail vertebrae, 118; hind 
foot, 27. Adult from Pluma, Oaxaca: 221 ; 116; 26.5. Skull (type) : 
Greatest length, 26.3; zygomatic breadth, 14; interorbital breadth, 
4.9; width of braincase, 10.6; nasals, 10.1; anterior palatine foramina, 
4; palatal bridge, 5.5; upper molar series, 3.8. 

Remarks. — The range of 0. guerrerensis marks the northern limit of 
the distribution of the 0. alfaroi group along the western slope of the 
mountains bordering the Pacific coast of Mexico. As in other south- 
ern groups, the general range of 0. alfaroi and its allies seems to bifur- 
cate north of the Isthmus of Tehuantepec, the forms which pass farther 
to the northward along the eastern and western slopes of the interior 
plateau region being isolated and developing differential characters. 
The ranges of 0. guerrerensis and 0. a. chapmani appear to be com- 
pletely separated, and while these forms differ appreciably in numer- 
ous details, they agree rather closely in the more essential features, 
and may prove to intergrade through the much larger form 0. a. 
caudatus. The latter, however, as at present understood, exhibits so 
wide a departure from guerrerensis that close comparison seems un- 
necessary. 

Specimens examined. — Total number, 8, as follows: 

Guerrero: Omilteme, 6 (type and topotypes). 
Oaxaca: Pluma, 2. 

ORYZOMYS HYLOCETES Merriam. 

Chiapas Rice Rat. 

(PI. Ill, figs. 8, 8a.) 

Oryzomys hylocetes Merriam, Proc. Washington Acad. Sci., Ill, p. 291, July 26, 1901. 

Type locality. — Chicharras, Chiapas, Mexico (altitude 3,500 feet). 

Type. — No. 77605, d 1 old, United States National Museum (Bio- 
logical Survey collection); collected by E. W. Nelson and E. A. 
Goldman, February 14, 1896. 

Geographic distribution. — Heavily forested Pacific slope, at 3,500 
feet, of mountains along continental divide in extreme southern 



1918.] ORYZOMYS TALAMANC^E GEOUP. 71 

Chiapas, and doubtless, adjacent portions of Guatemala; Humid 
Upper Tropical Zone. 

General characters. — Size small; color dark; pelage short; skull short 
and relatively broad across anterior roots of zygomata. Somewhat 
similar to 0. a. angusticeps and 0. a. rhabdops, but much smaller 
than either, and cranial characters distinctive. 

Color. — About as in 0. a. saturatior. 

STcull.— General form short, with posteriorly narrow braincase and 
anteriorly divergent zygomata; anterior palatine foramina rather 
long, but not reaching anterior plane of first molars; molars small. 
In size the skull is perhaps nearest to that of 0. a. saturatior, but it 
differs in the anterior expansion of the zygomata and departs from 
that of its larger and nearer geographic neighbor 0. a. angusticeps in 
the same respect. The skull is similar in general outline, especially 
the form of zygomata, to that of 0. a. rhabdops, but very much 
smaller. 

Measurements. — Type: Total length, 217; tail vertebrae, 118; hind 
foot, 27. SJcull (type): Greatest length, 26.2; zygomatic breadth, 
13.6; interorbital breadth, 4.6; width of braincase, 9.9; nasals, 9.7; 
anterior palatine foramina, 4.5; palatal bridge, 5.3; upper molar 
series, 3.5. 

Remarks. — Two examples only of 0. liylocetes are known, one old 
and in worn pelage, the other very young, and the range of individual 
variation the form may present can not now be determined. The 
species inhabits the lower slope of the mountains, while the upper 
levels of the same range are occupied by 0. a. angusticeps, a much 
larger form, with longer pelage and apparently representing a distinct 
specific type; but only through the accession of new material can 
the exact relationship of the two be made clear. The skull in general 
contour bears a striking resemblance to that of the otherwise different 
Guatemalan form, 0. a. rhabdops. 

Specimens examined. — Two, from type locality. 

Oryzomys talamancse Group. 

Geographic distribution. — Forested districts in eastern Costa Rica 
and Panama, and south in South America at least to southern Brazil; 
altitudinal range from sea level to 3,000 feet in Panama; Arid and 
Humid Lower Tropical Zones (fig. 6). 

General characters. — Size large; form rather slender; tail about equal 
to or somewhat exceeding head and body, scantily and indistinctly 
haired; ears large and conspicuous, thinly clothed externally and 
internally, with very short, fine, almost microscopic hairs, general 
pelage short, rather harsh, and lacking the woolly quality of 0. 
palustris; vibrissas about as long as head; toes of hind feet more or 
less distinctly webbed at base, the longest three bearing conspicuous 



72 NORTH AMERICAN FAUNA. [No. 43. 

tufts of silvery bristles which extend beyond the points of the claws; 
claws short, recurved, compressed, and sharp pointed. Color of up- 
perparts varying from rich ochraceous-tawny to cinnamon brown or 
russet, lined with black; underparts white or buffy whitish. 

SJcull. — Size large, with rostrum long and braincase low and some- 
what flattened; maxillary arm of zygoma heavy; outer wall of ant- 
orbital foramen with projecting border rounded or sloping forward, 
the antorbital notch rather shallow as viewed from above; nasals and 
premaxillse about conterminous posteriorly; frontal region wide, the 
lateral margins trenchant, a more or less conspicuous depression on 
median line close to posterior enpls of nasals; temporal ridges moder- 
ately developed anteriorly along parieto-squamosal sutures, becoming 
indistinct posteriorly in crossing lateral wings of parietals; interpa- 
rietal large; anterior palatine foramina short and broad, much shorter 
than palatal bridge, not reaching anterior plane of first molars; palatal 
pits small; spheno-palatine vacuities small; interpterygoid fossa 

broad; audital bullae small, the 
inner sides largely overlapped 
by mastoids; basi-occipital 
broad; angle of mandible mod- 
erately broad, the inferior bor- 
der strongly turned inward; 
coronoid process rather short 
and low; molars moderately 
heavy; the inner reentrant an- 
gles in upper molars and outer 
reentrant angles in lower mo- 
lars, deeply cutting the crowns, 

Fig. 6. — Geographic distribution of Oryzomys talamancse. , ., -, . ., i • i 

mandibular tooth row decid- 
edly narrower posteriorly than anteriorly; tubercle over root of lower 
incisor weakly developed. Contrasted with those of the 0. palustris 
group, skulls of 0. talamancse and its allies differ especially in greater 
length of rostrum, shorter anterior palatine foramina in relation to 
length of palatal bridge, smaller size of audital bullae, and in dental 
detail; the upper molars are more deeply cleft by inner reentrant 
angles and the lower molars by outer reentrant angles; the second 
upper molar differs especially in the more nearly equal size of the 
protocone and hypocone (protocone larger than hypocone in palus- 
tris), owing to more central position of inner reentrant angle, and in 
the absence of the large, elongated enamel island extending, in the 
moderately worn tooth of palusiris, along the postero-internal base 
of the paracone, or the reduction of this island to a smaller one, ap- 
pearing in talamancse near the mesostyle; the third lower molar is 
more triangular in outline than in palustris, the posterior division 
being decidedlv narrower than the anterior. 




1918.] ORYZOMYS TALAMANCJE GROUP. 73 

Remarks. — The group name used is that of the single North Ameri- 
can species, 0. talamancse of Costa Rica. This species typifies a series 
of wide distribution in South America; 0. mollipilosus and 0. medius 
are closely allied Colombian and Venezuelan forms, and others of 
this unrevised group range far south in Brazil. 

The 0. talamancse group is more nearly related to the 0. bombycinus 
group than to any other North American section of the genus, and 
representatives of the two sometimes share the same local habitat. 
Members of the talamancss group are externally separable by the short- 
ness of their pelage, while the heavier rostrum and the extension 
of the lateral wings of the parietals below the temporal ridges are 
distinguishing cranial characters. In dentition the two groups agree 
very closely. 

ORYZOMYS TALAMANCA Allen. 
Talamanca Rice Rat. 

(PI. IV, figs. 3, 3a; PI. V, fig. 7; PI. VI, figs. 4, 4a.) 

Oryzomys talamancse Allen, Proc. U. S. Nat. Mus., XIV, p. 193, July 24, 1891. 

Oryzomys panamensis Thomas, Ann. Mag. Nat. Hist., ser. 7, VIII, p. 252, September, 
1901. Type from City of Panama, Panama. No. 0.5.1.67, $, British Mus.; col- 
lected by E. Andre\ February 25, 1899. 

Oryzomys carrikeri Allen, Bull. Amer. Mus. Nat. Hist., XXIV, p. 656, October 13, 
1908. Type from Rio Sicsola, Talamanca, Costa Rica. No. 25976, 9 ad., Amer. 
Mus. Nat. Hist. ; collected by M. A. Carriker, jr., August 18, 1904. 

Type locality. — Talamanca, Costa Rica (probably near Sipurio, in 
the valley of the Rio Sicsola). 

Type. — No. ||f|| , c? adult (molars much worn), United States 
National Museum; collected by W. M. Gabb. 

Geographic distribution. — Heavily forested regions from eastern 
Costa Rica eastward through Panama to near Colombian frontier; 
altitudinal range from sea level to 3,000 feet; Arid and Humid Lower 
Tropical Zones. 

General characters. — Externally similar to O. mollipilosus of South 
America; skull narrower. (For additional general characters see 
under 0. talamancx group.) 

Color. — General color of upperparts varying from pale ochraceous- 
tawny to cinnamon brown or russet, becoming lighter and in some 
specimens ochraceous-buffy on cheeks, shoulders, and sides; under- 
pays dull white or buffy whitish, the plumbeous basal color showing 
through; ears brownish, indistinctly clothed externally with very 
short dusky hairs and internally with almost microscopic buffy or 
grayish hairs; feet (epidermis) dull yellowish, thinly covered above 
with short glossy white hairs, the hind feet with tufts of silvery 
bristles projecting beyond claws of longer digits; tail (epidermis) 
dark brownish above, varying from light brownish to dull yellowish. 



74 NORTH AMERICAN FAUNA. TNo. 43. 

Skull. — About like that of 0. mollipilosus, but braincase and frontal 
region decidedly broader. 

Measurements. — Type (dry skin) : Total length, 233 ; tail vertebrae, 
115; hind foot, 30. Average of three adults from Rio Sicsola, Costa 
Rica: 251 (240-265); 124 (118-133); 29.8 (29-30.5). Skull (type): 
Greatest length, 31.5; zygomatic breadth, 15.5; interorbital breadth, 
5.3; width of braincase, 11.2; nasals, 12.8; anterior palatine foramina, 
4.3 ; palatal bridge, 7.1 ; upper molar series, 4.6. 

Remarks. — Specimens from the Canal Zone and as far east as 
extreme eastern Panama seem referable to typical 0. talamancx of 
Costa Rica. Two examples from Gatun, Canal Zone-, have been sub- 
mitted to Mr. Oldfield Thomas for comparison with the type of " 0. 
jianamensis" in the British Museum. The result of his examination 
he has kindly written as follows: "We have only one specimen of 
0. panamensis [the type] and it is both larger and more rufous than 
your specimens. But it is older; the skull agrees in general characters 
and the toothrow is of exactly the same length. As to the colour I 
think the difference is only due to the coming on of the faded fulvous 
stage found in the old specimens of most species of Oryzomys. Per- 
sonally I should certainly refer your specimens to panamensis." 
Since the examples used for comparison are regarded as fairly typical 
of talamancx I conclude that panamensis must be placed in the syn- 
onymy of that species. 

The type and two topotypes of "0. carrikeri" Allen agree essentially 
with the type of talamancx. The exact locality of the latter is not 
definitely known, but Gabb, the collector, worked mainly near Sipurio 
and probably secured the specimen there. The Carriker collection 
came from about halfway between Cuabre and the mouth of the Rio 
Sicsola, the two localities being, on this assumption, not far apart in 
the same river valley. 

The ranges of talamancse and 0. alfaroi overlap, and owing to their 
superficial resemblance the two have sometimes been confused; 
talamancx is a larger animal than alfaroi, with a longer hind foot. The 
skulls, however, present the safest distinguishing characters, that ot 
talamancx being more massive, with heavier maxillary arms of zygo- 
mata, and heavier dentition; the first and second upper molars, 
besides differing in details of crown arrangement, are more deeply cleft 
by inner reentrant angles and all the lower molars by outer reentrant 
angles, while in alfaroi they more nearly approach the condition 
shown in 0. palustris. 

Specimens examined. — Total number, 20, as follows: 

Costa Fvica:Boruca, l; 1 Rio Sicsola, 3 (including type of ' ' carrikeri ") ; 2 Talamanca, 

1 (type). 
Panama: Cana, 7; Cerro Brujo, 1; Divala, 1; J Gatun, 6. 

1 Collection Mus. Comp. Zool. 2 Collection Amer. Mus. Nat. Hist. 



1918.] 



ORYZOMYS BOMBYCINUS GROUP. 



75 




Oryzomys bombycinus Group . 

Geographic distribution. — Forested regions from southern Costa 
Rica south through Panama at least to northern Ecuador ; altitudinal 
range from near sea level to at least 3,000 feet; Humid Lower Tropical 
Zone (fig. 7). 

General characters. — Size large; form rather slender; tail about 
equal to or slightly longer than head and body, scantily and very 
indistinctly haired; pelage of upperparts very long (that of back 
measuring about 12 mm.), underfur not woolly as in 0. palustris; 
supraorbital vibrissa? 50 to 70 
mm. in length (exceeding those 
arising from the muzzle) , reach- 
ing posteriorly to sides of body ; 
ears rather large and naked 
appearing, the short, fine hairs 
scattered over inner and outer 
surfaces almost microscopic ; 
hind feet slender, very scantily 
haired above, the plantar sur- 
face generally smooth, but in 
some examples indistinctly 
granular between interdigital 
tubercles ; toes of hind feet more 
or less distinctly webbed at 
base, the longest three bearing conspicuous tufts of silvery bristles pro- 
jecting beyond ends of claws as in the 0. talamancse group ; claws short, 
recurved, compressed, and sharp pointed. Colors dark; upperparts 
near cinnamon brown or russet; underparts overlaid with dull white. 

STcuM. — Size rather large, with rostrum long and narrow, the nasals 
attenuate and slightly exceeding premaxillaB in posterior extent; 
braincase broad and moderately expanded; zygomata slender; outer 
wall of antorbital foramen with projecting upper anterior border 
well rounded, the antorbital notch broad but not deeply cutting 
zygoma as viewed from above; frontal region moderately broad and 
somewhat flattened, the small depression on median line near ante- 
rior border deep and conspicuous, and more or less involving ends 
of nasals ; lateral margins of f rontals rising in distinct, but narrow, 
compressed ridges; temporal ridges well developed, bounding parie- 
tals laterally as far as lateral extension of supraoccipital, where 
they turn abruptly and are continued downward along the anterior 
border of that segment; parietals without lateral wings extending 
across temporal ridges; interparietal large, reaching transversely 
nearly across posterior parietal border; anterior palatine foramina 
short, not reaching anterior plane of first molars; palatal pits 
small; spheno-palatine vacuities very small or absent; interpterygoid 



1. O. 6. bombycinus. [ 2. O. b. alleni. 

Fig. 7. — Geographic distribution of subspecies of 

Oryzomys bombycinus. 



76 NORTH AMERICAN FAUNA. [No. 43. 

fossa broad ; audital bulla? small ; basi-occipital broad ; angle of mandi- 
ble broad, the inferior border strongly turned inward; coronoid process 
rather large and broad across base; molars moderately heavy, the 
crown arrangement of the 0. talamancse type, the crowns deeply incised 
by reentrant angles and the mandibular toothrowmuch narrower pos- 
teriorly; second upper molar about as in talamancse, the elongated 
enamel island present in 0. palustris, 0. melanotis and 0. alfaroi groups, 
being absent or displaced by a bending backward of the commissure of 
the paracone and protocone ; third lower molar long and narrow, the 
large outer reentrant angle extending more than halfway across the 
crown. 

Remarks. — Forms of this group differing only subspecifically among 
themselves evidently range from Costa Rica to Ecuador and may 
extend much farther; the group, therefore, seems to be mainly South 
American in distribution. Owing to lack of definite knowledge of 
the relationships of the South American forms it seems best to treat 
the North American representatives as specifically distinct until a 
revision of the South American species can be undertaken. 

In many important characters, especially of the skull and dentition, 
the 0. hombycinus group approaches the 0. talamancse, group and the 
two sometimes occur together in the same localities. The hombycinus 
group is easily recognizable externally by the much greater length 
of the pelage, while the skull is sufficiently distinguished by the slender 
rostrum and the absence of the lateral wings of the parietals, which in 
the talamancse group extend across the temporal ridges at the expense 
of the squamosals. Unlike many groups of the genus which show a 
preference for open or partially open situations 0. hombycinus seems 
to be at home in the depth of the forest. 

ORYZOMYS BOMBYCINUS Goldman. 

[Synonymy under subspecies.) 

Geographic distribution. — Heavily forested areas from southern 
Costa Rica to eastern Panama, and probably western Colombia; 
altitudinal range from sea level to at least 3,000 feet; Humid Lower 
Tropical Zone. 

General characters. — (See under 0. hombycinus group). 

Color. — General color of upperparts varying from ochraceous- 
tawny to cinnamon brown or russet finely mixed with black; darkest 
on face, top of head, and back, the lighter element restricted to tips 
of hairs and the dark basal color showing through, becoming fighter 
and approaching ochraceous-buff or pale ochraceous- tawny on cheeks, 
shoulders, and sides; underparts dull whitish; ears black, the outer 
and inner sides thinly clothed with very short dusky hairs; feet 
(epidermis) dull yellowish, the hairs whitish; tail dark brownish above, 
light brownish below. Young (about two-thirds grown) : Upper- 



1918.] ORYZOMYS BOMBYCINUS GROUP. 77 

parts blackish, nearly pure black on face, top of head, and back, 
becoming lighter and more distinctly lined with ochraceous-buffy 
hairs on cheeks, shoulders, and sides. 

Skull. — (See under 0. hombycinus group.) 

Remarks. — Two closely allied forms inhabit Panama and Costa 
Rica, and may not improbably prove to be geographic races of 
0. nitidus of Peru, since Ecuadorean specimens believed to be near 
nitidus show close alliance to the North American forms. 

Key to Subspecies of O. bombycinus. 

a 1 . Braincase less inflated. (Panama.) O. b. bombycinus (p. 77). 

a 2 . Braincase more inflated. (Costa Rica.) O. b. alleni (p. 78). 

ORYZOMYS BOMBYCINUS BOMBYCINUS Goldman. 

Long-Haired Rice Rat. 

(PI. II, figs. 7, 7a; PI. V, fig. 8; PI. VI, figs. 5, 5a.) 

Oryzomys bombycinus Goldman, Smiths. Miec. Coll., LVI, No. 36, p. 6, February 19, 
1912. 

Type locality. — Cerro Azul, near headwaters of Chagres River, 
Panama (altitude, 2,500 feet). 

Type.— No. 171105, 6* adult, United States National Museum 
(Biological Survey collection); collected by E. A. Goldman, March 
26, 1911. 

Geographic distribution. — Mountains of east-central Panama; alti- 
tudinal range from 1,000 to 3,000 feet; Humid Lower Tropical Zone. 

General characters. — Size about as in 0. b. alleni, color identical; 
skull with much natter, less expanded braincase. 

Color. — (See under 0. bombijcinus.) 

Skull. — Similar to that of 0. b. alleni, but braincase lower, flatter, 
much less distended, especially anteriorly; frontal region narrower 
posteriorly. 

Measurements. — Type: Total length, 252 ; tail vertebra?, 127; hind 
foot,. 32. Skull (type and adult topotype): Greatest length, 31.3, 
31.7; zygomatic breadth, 15.4, 16.1; interorbital breadth, 5.5, 5.4; 
width of braincase, 11.2, 11.6; nasals, 12.2, 12.2; anterior palatine 
foramina, 5, 4.5; palatal bridge, 6.2, 6.1 ; upper molar series, 4.7, 4.4. 

Remarks. — The Panama form apparently differs from the Costa 
Rican animal only in cranial characters, and the specimens on which 
it is based closely resemble Ecuadorean examples believed to be 
near 0. nitidus. The four Panama specimens are very slightly paler 
in color than five from Ecuador, with skulls also similar, but the 
zygomata more strongly bowed outward, the sides less nearly parallel. 

Specimens examined. — Total number, 4, as follows: 
Panama: Cerro Azul, 3 (type and topotypes); Cerro Brujo, 1. 



78 NORTH AMERICAN FAUNA. [No. 43. 

ORYZOMYS BOMBYCINUS ALLENI Goldman. 
Allen Rice Rat. 

(PI. II, figs. 8, 8a.) 

Oryzomys nitidus alleni Goldman, Proc. Biol. Soc. Washington, XXVIII, p. 128, 
June 29, 1915. 

Type locality. — Tuis (about 35 miles east of Cartago), Costa Rica. 

Type. — No. -f-ffl, S sub adult (teeth slightly worn), American 
Museum of Natural History; collected by George K. Cherrie, July 
15, 1894. 

Geographic distribution. — Mountainous portions of southern Costa 
Rica; altitudinal range from 8<3o to about 2,000 feet; Humid Lower 
Tropical Zone. 

General characters. — Size about as in 0. b. bombycinus; color iden- 
tical; skull with braincase higher, much more fully expanded. 

Color. — As in 0. b. bombycinus. (See under 0. bombycinus group.) 

Skull. — Similar to that of 0. b. bombycinus, but braincase higher, 
more arched, much more distended, especially anteriorly; frontal 
region broader posteriorly. 

Measurements. — Type tyoung): Total length, 218; tail vertebrae, 
111, hind foot, 29. Skull (type): Greatest length, 28.2; zygomatic 
breadth, 14.3; interorbital breadth, 5.5; width of braincase, 11.6; 
nasals, 10.9; anterior palatine foramina, 3.5; palatal bridge 5.5; 
upper molar series, 4.3. 

Remarks. — Cranial comparisons are necessary in order to distin- 
guish this Costa Rican form from 0. b. bombycinus of Panama. 
While the interorbital breadth is about the same, the greater anterior 
expansion of the braincase has resulted in broadening the posterior 
part of the frontal region. 

Specimens examined. — -Total number, 4, as follows 
Costa Rica: Tuis, 3 (type and topotypes); Guapiles, l. 1 

Oryzomys devius Group. 

Geographic distribution. — High mountains of central Costa Rica to 
Colombia and Venezuela; altitudinal range from 4,000 to at least 
5,000 feet; Humid Upper Tropical Zone (fig. 8). 

General characters. — Size very large, but form rather slender; tail 
much longer than head and body, indistinctly haired; ears rather 
large, blackish, the short hairs, thinly distributed over inner and outer 
sides, of about the same color as the epidermis; pelage long, but 
somewhat rigid; vibrissa long (about reaching shoulders); hind feet 
long and narrow, thinly haired above, the plantar surface between 
the large conspicuous tubercles smooth; toes of hind feet with the 
slight basal webbing usual in the genus, the longest three tufted with 

If i Collection Carnegie Mus. 



1918.] 



ORYZOMYS DEVIUS GROUP. 



79 



silvery bristles as in 0. talamancse and other groups; claws short, 
recurved, compressed and sharp pointed as in other partially scan- 
sorial sections of the genus. Colors dark; upperparts between tawny 
and russet ; underparts varying from ochraceous-tawny to dull white, 
the basal color plumbeous except in some examples, which tend to 
exhibit irregular areas on throat where the fur may be pure white 
to roots. 

STcull. — Size very large, with rostrum long and heavy; nasals 
broad and reaching posteriorly slightly beyond premaxillfe; braincase 
moderately broad and inflated; zygomata heavy; the maxillary arm 
extensively overlapping outer side of jugal, in some examples in con- 
tact with squamosal, the gap usual in the genus being completely 
bridged; outer wall of antorbital foramen with projecting upper 
anterior border rounded, the antorbital notch broad as viewed from 
above, but less deeply cutting 
zygoma, the aperture appear- 
ing less evenly circular than in 
the . palustris group; frontal 
region narrow, especially pos- 
teriorly, the anterior median 
depression rather inconspicu- 
ous ; lateral margins of f rontals 
smoothly rounded, or rising in 
slightly upturned ridges; tem- 
poral ridges moderately devel- 
oped, crossing lateral wings of 
parietals to supraoccipital and 
continuing thence downward in 
prominent crests along occipito- 
squamosal border; interparietal very large, extending transversely 
nearly across posterior parietal border, pointed-elliptical in outline 
owing to convexity of anterior margin ; anterior palatine foramina very 
short, not reaching anterior plane of first molars, narrow anteriorly, 
very broad and gaping widely open posteriorly, palatal pits large and 
irregular in form; spheno-palatine vacuities absent; interpterygoid 
fossa moderately broad and extending well forward, the anterior bor- 
der closely approaching posterior plane of last molars; audital bulla? 
variable but rather small; angle of mandible broad, the inferior border 
strongly incurved ; coronoid process large and strongly upturned, the 
high thin connecting ridge extending to near summit of condyle ; 
molars heavy and approaching the 0. talamancse, type, the crowns of 
the upper series deeply incised by inner reentrant angles, and of lower 
series by outer reentrant angles; as in the talamancse group the 
second upper molar is evenly cleft by inner reentrant angle and lacks 




1. O. devius. 2. O. pirrensis. 

Fig. 8. — Geographic distribution of the Oryzomys devius 

group. 



80 NORTH AMERICAN FAUNA. [No. 43. 

the elongated enamel island extending in the moderately worn crown 
of 0. palustris and other groups along the postero-internal base of the 
paracone; mandibular toothrow very narrow posteriorly, the third 
molar elongated and deeply cleft by outer reentrant angle. 

Remarks. — The 0. devius group, as it may be called, includes two 
northern representatives of a widely distributed South American 
section of the genus which in that country has sometimes been 
denominated the 0. meridensis group, from the name of a Vene- 
zuelan species. 

These northern forms generally lack the irregular but extensive pure 
white pectoral and inguinal areas exhibited by the allied Venezuelan 
and Colombian forms 0. meridensis and 0. maculiventer, although a 
tendency to develop them is shown in 0. devius. 

The North American species are externally easily recognizable 
among their congeners of the general region by the combination of 
large size with dark color and very long tails. Except for the more 
hispid pelage they are superficially much like the species of Pero- 
myscus (subgenus Megadontomys) occurring at the same localities 
with them. 

Key to Species of the O. devius Group. 

a 1 . Supraorbital ridges prominent. (Eastern Panama.) O. pirrensis (p. 81) . 

a 2 . Supraorbital ridges not prominent. (Western Panama.) O. devius (p. 80). 

ORYZOMYS DEVIUS Bangs. 

Chiriqui Rice Rat. 
(PL IV, figs. 1, la.) 

Oryzomys devius Bangs, Bull. Mus. Comp. Zool., XXXIX, p. 34, figs, 13, 14, April, 
1902. 

Type locality. — Boquete, Volcan do Chiriqui, Panama (altitude, 
5,000 feet). 

Type.— No. 10324, ? adult, Museum of Comparative Zoology 
(Bangs collection) ; collected by W. W. Brown, jr., January 29, 1901. 

Geographic distribution. — Forested slopes of high mountains in 
central Costa Rica and western Panama; altitudinal range from 4,000 
to at least 5,000 feet; Humid Upper Tropical Zone. 

General characters. — Size large, about as in 0. pirrensis of eastern 
Panama, but color slightly paler, more tawny; skull more smoothly 
rounded; audital bullae much larger. 

Color. — Upperparts dark tawny, inclining toward russet and rather 
heavily mixed with black along median line of dorsum, becoming light 
tawny, or in the paler examples dark ochraceous-buffy along lower 
part of sides; throat whitish (the fur pure white to roots over a 
^mall area in one example), rest of underparts overlaid with ochra- 
ceous-buffy in three examples and with dull white in the other three 
examined; nose and ears blackish; feet dull yellowish or light brown- 



1918.] ORYZOMYS DEVIUS GROUP. 81 

ish; tail (epidermis) dark brownish above, paler below. Young (in 
first pelage) : Upperparts blackish, the tawny element appearing 
rather inconspicuously along sides; underparts (in single specimen 
examined) thinly overlaid with dull white. 

Slcull. — Similar to that of 0. pirrensis, but more smoothly rounded, 
the supraorbital and temporal ridges weakly developed or absent; 
nasals slightly longer, ending posteriorly in plane of lachrymals; zygo- 
mata less widely spreading;' audital bullae decidedly larger. Compared 
with that of 0. meridensis, the skull is larger, with longer nasals and 
larger audital bulla?. 

Measurements. — Type: Total length, 335; tail vertebra?, ISO; hind 
foot, 33. Two adult topotypes: 345, 360; 185, 195; 36, 35. Slcull 
(average of three adults, type, and two topotypes) : Greatest length, 
36.6 (35.8-37.5); zygomatic breadth, 18.4 (18-19); interorbital 
breadth, 5.7 (5.5-6); width of braincase, 12.6 (12.2-12.9); nasals, 
14.3 (13.8-15); anterior palatine foramina, 5.4 (5.2-5.6); palatal 
bridge, 7.8 (7.8-7.9); upper molar series, 5.7 (5.6-5.8). 

Remarks. — 0. devius is clearly allied to 0. pirrensis of eastern 
Panama, but possesses very distinctive cranial characters, and there 
is no hint of intergradation. Both are inhabitants of high moun- 
tains, and their ranges are apparently separated by the intervening 
lowlands in the vicinity of the Canal Zone. 

Specimens examined. — Total number, 8, as follows: 

Costa Rica: Volcan Irazu, 2. 1 

Panama: Boquete, 6 (type and topotypes). 2 

ORYZOMYS PIRRENSIS Goldman. 

Mount Pirre Rice Rat. 

(PI. IV, figs. 2, 2a; PI. V, fig. 9; PI. VI, figs. 6, 6a.) 

Oryzomys pirrensis Goldman, Smiths. Misc. Coll., LX, No. 22, p. 5, February 28, 1913. 

Type locality. — Head of Rio Limon, Mount Pirre, Panama (altitude 
4,500 feet). 

Type.— No. 178993, <? adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. A. Goldman, April 29, 1912. 

Geographic distribution. — Steep, heavily forested slopes of high 
mountains at 4,500 feet altitude in eastern Panama, and probably 
adjacent portions of Colombia; Humid Upper Tropical Zone. 

General characters. — Size large, about as in 0. devius of western 
Panama, but color slightly darker, more russet; skull more angular; 
audital bulla? decidedly smaller. 

Color. — Fresh pelage: Upperparts between tawny and russet, heav- 
ily mixed with black along the broad median line from top of head to 

1 One specimen in Mus. Comp. Zool. 

2 Four specimens in Mus. Comp. Zool.; 2 in Field Mus. Nat. Hist. 

14521°— 18 6 



82 NORTH AMERICAN FAUNA. [No. 43. 

base of tail, becoming lighter, purer tawi^ on cheeks, shoulders, and 
sides; throat whitish or grayish, rest of underparts usually overlaid 
with ochraceous-tawny, but varying to dull white, the basal color of 
the fur everywhere deep plumbeous; nose and ears blackish; fore feet 
blackish, becoming lighter on toes; hind feet dark brown, thinly 
clothed with short hairs to toes, the toes dull yellowish; epidermis of 
tail dark brown above, usually somewhat paler below. Young (in 
first pelage) : Upperparts darker than in a*dults, the blackish element 
in the pelage predominant; underparts with a thinner ochraceous- 
tawny wash. 

Skull. — Similar to that of 0. devius, but more angular, the supra- 
orbital and temporal ridges well developed; nasals slightly shorter, 
not reaching posteriorly to plane of lachrymals; zygomata more 
widely spreading; audital bulla? decidedly smaller. In small size of 
audital bullas the skull agrees with those of 0. meridensis and 0. macu- 
liventer, but contrasts with both in larger general size and angularity. 

Measurements. — Type: Total length, 340; tail vertebrae, 185; hind 
foot, 38. Average of five adult topotypes: 314 (309-322); 164 (159- 
170) ; 35.7 (34-37). Skull (average of six adults, type, and five topo- 
types) : Greatest length, 36.8 (34.9-38.5); zygomatic breadth, 19.2 
(17.8-20); interorbital breadth, 5.8 (5.5-6); width of braincase, 12.4 
(11.8-12.9); nasals, 13.7 (13-14); anterior palatine foramina, 5.8 (5.4- 
6); palatal bridge, 7.5 (7.3-7.8); upper molar series, 5.7 (5.5-5.9). 

Remarks. — In external appearance 0. pirrensis differs only slightly 
from 0. devius of western Panama. The skull, however, combines 
the large general size of that species with the smaller general dimen- 
sions and small audital bullse of 0. meridensis and 0. maculiventer; it 
differs from both in the development of the supraorbital and temporal 
ridges. In color of upperparts 0. pirrensis is similar to the South 
American species, but the underparts show no trace of the pure white 
pectoral and inguinal areas which are so conspicuous in the latter 
forms. 

Specimens examined. — Eight, from type locality. 

Oryzomys tectus Group. 

Geographic distribution. — Southern Costa Rica and southeastward 
through Panama at least to Colombia and Venezuela; altitudinal 
range from near sea level to about 5,000 feet (fig. 9). 

General characters. — Size large, form rather robust; tail about equal 
to or somewhat exceeding head and body, scantily haired, the epi- 
dermal scales small; ears small, moderately clothed externally and 
internally with comparatively coarse hairs of general body color; 
general pelage long, coarse, and rigid; vibrissas somewhat longer than 
head; hind feet short and relatively broad, well haired above, the 
longest three toes bearing conspicuous tufts of silvery bristles, which 



1918.] 



ORYZOMYS TECTUS GROUP. 



83 



project beyond ends of claws; claws short, strongly curved, com- 
pressed, and sharp-pointed. Color of upperparts between rich tawny 
and ochraceous-tawny, mixed with black, the tawny element pre- 
dominating ; underparts varying from nearly pure white to warm buff. 

Skull. — Size large and angular, with rostrum short and braincase 
low and flattened; outer wall of antorbital foramen rounded above, 
the forward projection moderate; nasals short, ending posteriorly in 
the anterior plane of orbits; premaxillse about conterminous with 
nasals posteriorly, the ends slightly expanded, not beveled externally ; 
frontals very broad, the lateral margins overhanging as supraorbital 
shelves; temporal ridges prominent, extending posteriorly to supra- 
orbital border; mastoid process of squamosal short and stout, owing 
to slight excision of squamosal margin; interparietal large, pointed- 
elliptical, the anterior angle well developed; lachrymal very small; 
anterior palatine foramina 
short, moderately broad anteri- 
orly and posteriorly, reaching 
or nearly reaching anterior 
plane of first molars; palatal 
pits normally small; spheno- 
palatine vacuities absent or 
very small ; audital bullae small ; 
ramus of mandible short, the 
angle broad; coronoid process 
broad and strongly hooked, a 
high trenchant ridge connecting 
with condyle; tubercle over 
root of lower incisor large. 
Molars similar to those of the 
0. talamancse group; approaching the 0. palustris type in general char- 
acters, but second upper molar with inner lobes more nearly equal 
in extent (the anterior slightly the larger in palustris), and central 
enamel island present in palustris usually fused with enamel fold 
separating paracone and parastyle; third lower molar more triangular 
in outline, the posterior lobe narrower and the outer reentrant angle 
more nearly dividing crown. 

Remarks.— -O. tectus is typical of a group including also 0. flavicans, 
0. palmarius, and other extralimital forms distinguished by rich 
coloration, coarsely haired ears, and short stout hind feet. In external 
appearance they are not very unlike some species of Rhipidomys, 
but have shorter, less hairy tails and lack the dark metapodial mark- 
ings usually present in that genus; in general characters they appar- 
ently approach the section assigned to generic rank by Thomas under 
the name (E corny s, 1 but in the more essential respects scarcely exhibit 




1. O. t. tectus. | 2. 0. t. frontalis. 

Fig. 9. — Geographic distribution of subspecies of Oryzo- 

mys tectus. 



I Thomas, Oldfield, Ann. Mag. Nat. Hist., ser. 7, XVIII, p. 444, Dec. 1906. 



84 NORTH AMERICAN FAUNA. [No. 43. 

a wider departure from typical Oryzomys than several groups usually- 
assigned to the genus. 

ORYZOMYS TECTUS Thomas. 

[Synonymy under subspecies.] 

Geographic distribution. — Southern Costa Rica, Panama, and prob- 
ably adjacent portions of Colombia; vertical range from near sea 
level to about 2,000 feet, mainly in Arid Lower Tropical Zone. 

General characters. — Similar to 0. jlavicans of Venezuela, but 
larger; color darker; skull larger with laterally expanded frontal 
region. (For additional generaKcharacters see under 0. tectus group.) 

Color. — Fresh pelage: General color of upperparts varying shades 
of tawny and ochraceous-tawny, rather inconspicuously darkened on 
face, top of head, and back by an admixture of dusky hairs, becom- 
ing paler, more ochraceous-buffy on cheeks, shoulders, and sides; 
underparts varying from near pure white to warm buff; ears clothed 
internally and externally with short tawny hairs; feet whitish; tail 
varying from uniform dark brownish all around to dark brownish 
above, and whitish or yellowish below. 

SJcull. — Size large, with remarkably broad frontal region. Similar 
in general to 0. Jlavicans, but decidedly larger; frontal region broader, 
the lateral margins shelving farther over orbits, not strongly upturned 
as in Jlavicans. (For additional characters see under 0. tectus group.) 

Remarks. — 0. tectus is clearly allied to 0. Jlavicans Jlavicans, O.f. 
illectus, and 0. palmarius, all South American forms in which the 
supraorbital ridges are well developed, but are compressed and not 
widely expanded and Tylomys-like as in the forms of tectus. This 
difference, although rather striking, is a relative instead of absolute 
character, and close agreement in other essential respects even sug- 
gests probable intergradation. 

Key to Subspecies of O. tectus. 

a 1 . Upperparta more distinctly tawny. (Western Panama and southern Costa 

Rica.) O. t. tectus (p. 84). 

a 2 . Upperparts less distinctly tawny. (Eastern Panama.) O. t. frontalis (p. 85). 

ORYZOMYS TECTUS TECTUS Thomas. 
Bugaba Rice Rat. 

Oryzomys tectus Thomas, Ann. Mag. Nat. Hist., ser. 7, VIII, p. 251, September, 1901. 

Type locality. — Bugaba, Chiriqui, Panama (altitude 800 feet). 

Type. — No. 0.7.11.43, $ , British Museum; collected by H. J. 
Watson, September 15, 1898. 

Geographic distribution. — Pacific slope of western Panama and 
southern Costa Rica at about 800 feet altitude, limits of altitudinal 
range unknown ; Arid Lower Tropical Zone. 



1918.] OEYZOMYS TECTUS GROUP. 85 

General characters. — Closely resembling 0. t. frontalis, but upper- 
parts richer tawny; underparts more extensively buffy; lips, chin, 
and throat buffy instead of white as in frontalis; skull differing in 
rather slight details. 

Color. — Upperparts near tawny rather thinly mixed with black, 
general color darkest over dorsum, becoming paler and rich ochra- 
ceous-buffy on cheeks and sides ; underparts, including lips, chin, and 
throat overlaid with warm buff; ears clothed with tawny hairs; feet 
whitish; tail (epidermis and hairs) brownish above, whitish or dull 
yellowish below to near tip, which is dark all around. 

Skull. — Very similar to that of 0. t. frontalis, but frontals more 
extended posteriorly on median line between parietals; interparietal 
smaller. 

Measurements. — From original description of type: "Head and 
body, 140 millim. ; tail, 142; hind foot, s. u. 27, c. u. 29.5; ear, 18. 
Skull: Tip of nasals to back of interparietal, 33; greatest breadth, 17; 
nasals, 11.6 X 4 ; interorbital breadth, 6.5 ; palate length, 13.8; diastema 
8.1; palatal foramina, 5X2.2; length of upper molar series, 4.9." 
A rather young example from Boruca, Costa Rica: Total length, 258; 
tail vertebrae, 140; hind foot, 27. 

Remarks. — The type of 0. t. tectus has not been examined by me, 
but specimens from Boruca, Costa Rica, are believed to be typical. 
They are distinguished from 0. t. frontalis of eastern Panama mainly 
by richer general coloration. No other member of the 0. tectus group 
ranges so far into the Central American Subregion. 

Specimens examined. — Two, as follows: 

Costa Rica: Boruca, 2. 

ORYZOMYS TECTUS FRONTALIS Goldman. 

Corozal Rice Rat. 

(PI. IV, figs. 4, 4a; PI. V, fig. 10; PI. VI, figs. 7, 7a.) 

Oryzomys frontalis Goldman, Smiths. Misc. Coll., LV1, No. 36, p. 6, February 19, 1912. 

Type locality. — Corozal, Canal Zone, Panama (altitude 100 feet). 

Type. — No. 171531, ? adult, United States National Museum 
(Biological Survey collection); collected by E. A. Goldman, June 
20, 1911. 

Geographic distribution. — Forested Pacific slope of Panama from 
the Canal Zone to near Colombian frontier, and probably adjacent 
Colombian territory; altitudinal range from sea level to about 2,000 
feet; Arid and Humid Lower Tropical Zones. 

General characters. — Similar in general to 0. t. tectus, but upperparts 
duller, less distinctly tawny; underparts less extensively buffy; lips, 
chin, and throat white instead of buffy as in tectus; skull differing in 
rather slight details. 



86 NORTH AMERICAN FAUNA. [No. 43. 

Color. — Upperparts between tawny and ochraceous-tawny, rather 
sparingly mixed dorsaUy with black, the general color becoming paler, 
purer, and ochraceous-buffy on cheeks and sides; underparts in 
general white in some examples, thinly overlaid with warm buff 
across the abdomen in others, the lips, chin, and throat in all speci- 
mens examined nearly pure white to roots of hairs; ears and feet as 
in 0. t. tectus; tail usually unicolor, dark brownish, but in some 
examples becoming lighter on under side near base. 

STcull. — Closely resembling that of 0. t. tectus, but frontals less 
extended posteriorly on median line between parietals ; interparietal 
larger. 

Measurements. — Type: Total length, 309; tail vertebrae, 161 ; hind 
foot, 30. Average of five adults from Cana, Panama: 288 (281-293) ; 
156 (152-164); 27.8 (26.5-29). Skull (average of same) : Greatest 
length, 32.8 (31.1-34.1); zygomatic breadth, 17.1 (16.6-17.5) 
interorbital breadth, 6.1 (5.3-6.8); width of braincase, 12.2 (12-12.6) 
nasals, 11.4 (10.5-12.4); anterior palatine foramina, 5.3 (4.9-5.6) 
palatal bridge, 6.4 (5.6-6.9); upper molar series, 5.2 (4.8-5.3). 

Remarks. — Two specimens of 0. t. frontalis from eastern Panama 
have been submitted to Mr. Oldfield Thomas, who has kindly com- 
pared them with the type and topotypes of 0. t. tectus in the British 
Museum. While inclined to regard them as referable to the same 
form he writes that "they are not quite so rich in colour as our 
specimens." Two examples from Boruca, Costa Rica, not far from 
the type locality and in the same general f aunal area, and believed to 
be fairly typical of tectus, exhibit a type of coloration and slight 
cranial details indicating that eastern and western Panama are 
inhabited by closely allied but easily recognizable forms. 

Specimens examined. — Total number, 12, as follows: 

Canal Zone: Corozal, 1 (type). 
Panama: Cana, 11. 

ORYZOMYS VICTUS » Thomas. 

St. Vincent Rice Rat. 

Oryzomys victus Thomas, Ann. Mag. Nat. Hist., ser. 7, I, p. 178, February, 1898. 

Type locality. — St. Vincent, Lesser Antilles. 

Type. — No. 97.12.26.1, 9 adult, British Museum; collected by 
H. H. Smith, presented by F. DuCane Godman. 

Geographic distribution. — Known only from St. Vincent. 

General characters. — From original description of type: "Size and 
proportions about as in the larger members of the 0. longicaudatus 
group. General colour dark rufous, but evidently affected by the 
spirit in which the specimen has been preserved. Under surface 

1 Group association undetermined. 



1918.] SUBGENUS OLIGOEYZOMYS. 87 

buffy white, the bases of the hairs slate-colour. Eyes without darker 
rims. Ears short, the anterior part of their backs brown, not strik- 
ingly contrasting with the general colour of the head. Hands and 
feet thinly clothed with fine silvery hairs. Tail almost naked, brown 
above, slightly paler below. Mammae 2-2=8." 

SlcuU. — From original description of type: "Skull with the general 
shape of the South-American 0. longicaudatus, the braincase being 
similarly lengthened as compared with the broadened braincase of 
the Central-American 0. melanotis and its allies. Compared with a 
Rio Janeiro example it is larger, more rounded, the supraorbital 
edges less sharply square, but the parietal ridges thicker and better 
developed. Molars larger and stouter, palate ending only just 
behind the back of m 3 ." 

Measurements. — From original description of type (measured in 
spirit): "Head and body, 96 millim.; tail, 121; hind foot without 
claws, 25; with claws, 26.7; ear, 14. Skull: Basilar length, 21.4; 
basal length, 23.8; greatest breadth, 15.1; nasals, 11.2x3.4; inter- 
orbital breadth, 4.5; interparietal, 3.2x10; palate length from 
henselion, 12.3; diastema, 7.8; palatal foramina, 5.4x1.8; length 
of upper molar series, 4.1." 

Remarks. — -The type of 0. victus has not been examined by me. It 
was originally compared mainly with a South American species, but 
the true affinities remain to be determined. As in the case of 0. 
antillarum, of Jamaica, this rice rat seems likely to be endangered by 
the presence of the mongoose, if it has not already been exterminated 
since the introduction of that indiscriminately destructive animal. 

Subgenus OLIGORYZOMYS Bangs. 

Oligoryzomys Bangs, Proc. New England Zool. Club., I, p. 94, February 23, 1900 (sub- 
genus). Type Oryzomys navus Bangs. 

GeograpJiic distribution. — Southern Mexico south through Central 
America to undetermined limits in South America. 

Subgeneric characters. — Size very small; hind foot usually less than 
25; form slender and Reithrodontomys-lake; ears rather large and 
coarsely haired; tail much longer than head and body; four longer 
toes of hind feet bearing tufts of silvery bristles projecting beyond 
ends of claws. 

Skull delicate in structure, smoothly rounded; supraorbital and 
temporal ridges absent; interorbital region narrow, the constriction 
about equal to width of rostrum between antorbital foramina ; outer 
wall of antorbital foramen projecting slightly forward, as viewed from 
above ; angle of mandible placed well within vertical plane of condyle ; 
molars with small accessory cusps present as in subgenus Oryzomys, 
but reentrant angles usually broader, the salient angles formed by 
worn crowns of tubercles less evenly rounded; upper molars early 



88 NORTH AMERICAN FAUNA. [No. 43. 

exhibiting small, circular central enamel islands which persist until 
obliterated by wear in extreme old age; second upper molar with 
protocone and hypocone of about equal size, the inner reentrant 
angle central in position; third lower molar with outer reentrant 
angle extending less than halfway across crown; tubercle over root 
of lower incisor large. 

Remarks. — The subgenus Oligoryzomys includes a series of forms 
easily distinguishable among their North American congeners by 
diminutive size and external resemblance to species of the genus 
Reiihrodontomys . Oligoryzomys, however, departs from the subgenus 
Oryzomys, as currently restricted, mainly in a combination of relative 
rather than absolute characters. The molar crowns differ in details 
of enamel arrangement, the second upper especially, in the early 
appearance of a single, persistent, normally circular enamel island in 
the broad central space between the apex of the inner reentrant angle 
and the base of the paracone. In the subgenus Oryzomys this molar 
crown varies in pattern; the more typical forms normally present, in 
early stages of wear, an elongated, crescentic enamel island in the 
central space, but in more divergent forms the enamel island may be 
absent or tend to unite with the long, deep enamel fold between the 
paracone and parastyle. Several groups seem at least as fully 
entitled to subgeneric recognition as Oligoryzomys, but the problem of 
further subgeneric divisions can best be solved when more compre- 
hensive study of the genus is undertaken. 

ORYZOMYS FULVESCENS (Saussure). 
[Synonomy under subspecies.) 

Geographic distribution. — From southern Mexico south through 
Guatemala, Honduras, Nicaragua, and Costa Kica to Panama, and 
probably portions of Colombia ; altitudinal range from near sea level 
to about 5,500 feet; mainly Arid and Humid Lower Tropical Zones 
(fig. 10). 

General characters. — {See subgeneric characters under subgenus 
Oligoryzomys.) 

Color. — Upperparts varying from light ochraceous-buff to ochra- 
ceous-buff or tawny, most intense on rump, palest and purest on 
cheeks, shoulders, and sides; the face, top of head, and back mod- 
erately darkened by blackish hairs; underparts varying from nearly 
pure white in general to light ochraceous-buff on abdomen, inguinal 
region, and inner sides of hind limbs ; the lips, throat, and fore limbs 
white ; outer sides of ears blackish, inner sides clothed with ochra- 
ceous-buffy hairs; vibrissas black (about as long as head); feet white, 
the hind feet with tufts of silvery hairs projecting beyond claws of 
longest four digits; tail dark brownish above, lighter brown or yel- 
lowish below, except toward tip, which is dusky all around. Young 



1918.] 



OEYZOMYS FULVESCENS. 



89 



(in first pelage) : Usually darker than adults, the ochraceous-buff or 
tawny element less developed. 

Skull. — (See subgeneric characters under subgenus Oligoryzomys.) 
Remarks. — The North American forms of the subgenus are all 
referable to a single species whose range probably extends well into 
South America. They agree closely among themselves in essential 
characters, and by small size and slender form are sufficiently distin- 
guished from other sections of the genus inhabiting the same region. 








1. O././ulvcscens. 4. O.f. costaricensis. 

2. O.f.lenis. 5. O./.vegetus. 

3. O.f. mayensis. 

Fig. 10. — Geographic distribution of subspecies of 
Oryzomy s fulvescens. 



Key to Subspecies of O. fulvescens. 
a 1 . Size smaller; hind foot less than 24. 
b 1 . Upperparts darker ochraceous-buff. 
c 1 . Upper molar series shorter. (Southern Tamaulipas; eastern Oaxac a; Chiapas; 

northern Honduras.) O. f. fulvescens (p. 89). 

c 2 . Upper molar series longer. (Southwestern Panama; Costa Rica; Nicaragua.) 

O. f. costaricensis (p. 92). 
b 2 . Upperparts paler ochraceous-buff. 
c l . Skull broader; zygomatic breadth 11.5 or more. (Michoacan; Guerrero; 

southern Oaxaca.) O. f. lenis (p. 91). 

c 2 . Skull narrower; zygomatic breadth less than 11.5. (Yucatan; Campeche.) 

O. f. mayensis (p. 92). 
a 2 . Size larger; hind foot 24 or more. (Mountains of western Panama.) 

O. f. vegetus (p. 93). 

ORYZOMYS FULVESCENS FULVESCENS (Saussure). 

Vera Cruz Pygmy Rice Rat. 

[PI. IV, figs. 5, 5a; PI. V, figs. 2, 11; PI. VI, figs. 8, 8a.] 

Hesperomys fulvescens Saussure, Rev. et Mag. Zool., ser. 2, XII, p. 102, March, 1860. 
Oryzomys fulvescens Allen and Chapman, Bull. Amer. Mus. Nat. Hist., IX, p. 204, 
June 16, 1897. 

Type locality. — Orizaba, 1 Vera Cruz, Mexico. 



1 Type locality fixed by Merriam, Proc. Washington Acad. Sci., Ill, p. 295, July 26. 1901. 



90 NORTH AMERICAN FAUNA. [No. 43. 

Type. — In Geneva Museum of Natural History. 

Geographic distribution. — Southern Tamaulipas, Vera Cruz, eastern 
Oaxaca, Chiapas, and east through central Guatemala to eastern 
Honduras; altitudinal range from near sea level to about 5,500 feet; 
mainly Arid and Humid Lower Tropical Zones. 

General characters. — Size small; molar tooth series short. Similar 
to O.f. lenis but upperparts darker ochraceous-buff, the general tone 
less yellowish; skull narrower and less massive. Size about as in 
O.f. costaricensis , but upperparts usually less tawny; molar series 
shorter. 

Color. — Fresh pelage: Upperparts near ochraceous-buff, most in- 
tense and in some examples suffused with tawny on rump, palest and 
purest on cheeks, shoulders, and sides; the face, top of head, and back 
moderately darkened by blackish hairs; underparts varying from 
nearly pure white throughout in rare examples to warm buff on 
abdomen, inguinal area, and inner sides of hind limbs, the white ap- 
pearing only on lips, chin, and inner sides of hind limbs; outer sides of 
ears blackish, inner sides clothed with ochraceous-buffy hairs; feet 
whitish; tail brownish above, yellowish below, except toward tip, 
which is dusky all around. Young (in first pelage) : Darker than 
adults, the ochraceous-buff of upperparts restricted to narrow tips of 
hairs, thus permitting plumbeous basal color to show through and 
alter general tone. 

Skull. — About like that of O.f. costaricensis, but molar tooth series 
shorter. Contrasted with 0. f. lenis, the skull is narrower, with 
less widely spreading zygomata; maxillary arms of zygomata and 
ascending branches of premaxillae less broad and heavy. 

Measurements. — Average of 10 adults from Orizaba, Vera Cruz: 
Total length, 174.7 (168-205); tail vertebras, 107.1 (96-118); hind 
foot, 22 (21-23). Skull (average of same): Greatest length, 21.9 
(21.1-22.5); zygomatic breadth, 11.5 (11-11.7); interorbital breadth, 
3.5 (3.4-3.9); width of braincase, 9.5 (9.3-9.8); nasals, 7.9 (7.6-8.3); 
anterior palatine foramina, 3.5 (3.3-3.6); palatal bridge, 3.7 (3.5-4); 
upper molar series, 2.9 (2.9-3). 

Remarks. — The differential characters are rather slight, and 0. f. 
fulvescens may safely be assumed to intergrade with O.f. lenis near 
the Pacific coast in the vicinity of the Isthmus of Tehuantepec, with 
0. f. costaricensis in eastern Honduras or western Nicaragua, and 
with the paler form inhabiting the Yucatan Peninsula, in southern 
Campeche. A single specimen from Patuca, Honduras, the only one 
available from that country, has a short molar series and seems refer- 
able to the subspecies fulvescens, the known range of which is thus 
materially extended eastward from western Guatemala. 

Specimens examined. — Total number, 73, as follows: 
Chiapas: Huehuetan, 4; Ocuilapa, 1; Tumbala, 11; Yajalon, 1. 
Guatemala: Jacaltenango, 3; Nenton, 1. 



1918.] OEYZOMYS FULVESCENS. 91 

Honduras: Patuca, 2. 

Oaxaca: Santo Domingo (mountains near), 2; Tuxtepec, 1. 
Tamaulipas: Alta Mira, 1. 

Vera Cruz: Jalapa, 24; 1 Jico, 2; Mirador, 1; Orizaba (type locality by fixation), 
17; Pasa Nueva, l; 2 Santiago Tuxtla, 1. 

ORYZOMYS FULVESCENS LENIS Goldman. 

Los Reyes Pygmy Rice Rat. 

(PI. IV, figs. 6, 6a.) 

Oryzomys fulvescens lenis Goldman, Proc. Biol. Soc. Washington, XXVIII, p. 130, 
June 29, 1915. 

Type locality. — Los Reyes, Michoacan, Mexico. 

Type. — No. 125941, <$ adult, United States National Museum (Bio- 
logical Survey collection) ; collected by E. W. Nelson and E. A. Gold- 
man, February 14, 1903. 

Geographic distribution. — Coastal plains and basal mountain slopes 
in Michoacan, Guerrero, and Oaxaca; altitudinal range from near sea 
level to about 3,000 feet, mainly in Arid Lower Tropical Zone. 

General characters. — Similar to O. f. fulvescens, but upperparts 
paler ochraceous-buff , the general tone more yellowish ; skull broader 
and more massive. 

Color. — Fresh pelage: Upperparts between ochraceous-buff and 
light ochraceous-buff becoming warm buff on cheeks, shoulders, and 
lower part of sides; the face, top of head, and back sparingly lined 
with dark hairs; underparts, ears, feet, and tail as in O.f. fulvescens. 

SJcull. — Broader than that of O. f. fulvescens; zygomata more 
widely spreading; maxillary arms of zygomata and ascending branches 
of premaxillse broader and heavier; dentition rather heavy, but 
equaled in some examples of O. f. fulvescens. 

Measurements. — Type: Total length, 198; tail vertebra?, 115; hind 
foot, 23. Adult topotype: 188; 115; 23. Skull (type and adult 
topotype): Greatest length, 22.6, 21.8; zygomatic breadth, 12.3, 
11.9; interorbital breadth, 3.4, 3.5; width of braincase, 10, 10.3; 
nasals, 8.6, 7.8; anterior palatine foramina, 3.9, 3.5; palatal bridge, 
4.1, 3.7; upper molar series, 3, 3. 

Remarks. — The general distribution area of 0. fulvescens is divided 
north of the Isthmus of Tehuantepec by the high, interior plateau 
region, the western arm representing the range of O.f. lenis. Speci- 
mens from Guerrero and Oaxaca approach subspecies fulvescens in 
characters. 

Specimens examined. — Total number, 4, as follows: 
Guerrero: Ometepec (near), 1. 
Michoacan: Los Reyes, 2 (type and topotypes). 
Oaxaca: Pluma, 1. 

i Sixteen in Amer. Mus. Nat. Hist.; 2 in Mus. Comp. Zool. 2 Collection Amer. Mus. Nat. Hist. 



92 NORTH AMERICAN FAUNA. [No. 43. 

ORYZOMYS FULVESCENS MAYENSIS, subsp. nov. 
Maya Pygmy Rice Rat. 

Type locality. — Apazote (near Yohaltum), Campeche, Mexico 
(altitude 200 feet) . 

Type. — No. 107979, <? adult, United States National Museum 
(Biological Survey collection) ; collected by E. W. Nelson and E. A. 
Goldman, January 5, 1901. Original number 14405. 

Geographic distribution. — Peninsula of Yucatan and Campeche; 
altitudinal range from near sea level to about 300 feet; Arid Lower 
Tropical Zone. 

General characters. — Similar to O. f. fulvescens, but paler colored; 
skull slightly narrower. 

Color. — Fresh pelage: Upperparts near ochraceous-buff shading 
toward light ochraceous-buff, darkest and rather sparingly lined 
with black over dorsum, becoming clearer and paler on cheeks, 
shoulders, and sides; underparts, ears, feet, and tail as in O.f. ful- 
vescens. 

STcull. — About like that of O. f. fulvescens, but zygomata and 
braincase slightly narrower. 

Measurements. — Type: Total length, 177; tail vertebrae, 101, hind 
foot, 22. Average of three adult topotypes: 189 (184-196); 113 
(108-119) ; 23 (22-24). Skull (average of three adults, type and two 
topotypes): Greatest length, 21.1 (20.4-21.8); zygomatic breadth, 
11 (10.8-11.2) ; interorbital breadth, 3.6 (3.5-3.8) ; width of braincase, 
9 (8.9-9.1); nasals, 8.1 (7.4-8.4); anterior palatine foramina, 3.3 
(3.1-3.5); palatal bridge, 4 (3.6-4.4); upper molar series, 2.9 (2.9-3). 

Remarks. — The pale color of this form seems to be due to the same 
environmental conditions that have been operative in the evolution 
of pallid races in other mammalian groups inhabiting Yucatan. In 
this region pale coloration is apparently associated with the arid 
climate and white limestone formation, outcropping or thinly over- 
laid with soil over large areas which have risen less than 300 feet above 
the level of the sea. At Chichen Itza the pygmy rice rat was taken in 
the vicinity of imposing Maya Indian ruins, a circumstance that sug- 
gested the subspecific name. 

Specimens examined.— Total number, 7, as follows: 

Campeche: Apazote, 4 (types and topotypes). 
Yucatan: Chichen Itza, 1; Tunkas, 2. 

ORYZOMYS FULVESCENS COSTARICENSIS Allen 
Costa Rican Pygmy Rice Rat. 

Oryzomys costaricensis Allen, Bull. Amer. Mus. Nat. Hist., V, p. 239, September 22, 

1893. 
Oryzomys (Oligoryzomys) nicaragux Allen, Bull. Amer. Mus. Nat. Hist., XXVIII, p. 

100, April 30, 1910. Type from Vijagua, Nicaragua, No. 29543, c? ad., Amer. 

Mus. Nat. Hist. ; collected by W. B. Richardson, March 24, 1909. 



1918.] ORYZOMYS FULVESCENS. 93 

Type locality. — El General, Costa Rica (altitude, 2,150 feet). 

Type. — No. f§-fl> ? subadult, American Museum of Natural His- 
tory, collected by H. Pittier, February, 1891. 

Geographic distribution. — Southwestern Panama east to Canal Zone, 
Costa Rica, and north to northern Nicaragua; altitudinal range from 
near sea level to at least 2,000 feet. 

General characters. — Similar to 0. f fulvescens, but upperparts 
usually more tawny; molars larger. Also similar to O.f. vegetus, but 
smaller and averaging paler. 

Color. — About as in 0. f. fulvescens, but upperparts usually more 
distinctly tawny. 

Skull. — Closely resembling that of O.f. fulvescens, but molars 
larger, the toothrows decidedly longer. General size smaller than 
that of O.f. vegetus; molars about the same. 

Measurements. — Two adults from Buenos Aires, Costa Rica: 
Total length, 183, 190; tail vertebrae, 100, 108; hind foot, 20, 23. 
Skull (same): Greatest length, 21.1, 21.5; zygomatic breadth, 11.4, 
11.4; interorbital breadth, 3.7, 3.7; width of braincase, 9.9, 9.6; 
nasals, 8.2, 8; anterior palatine foramina, 3.6, 3.7; palatal bridge, 
4, 4.2; upper molar series, 3.2, 3.2. 

Remarks. — The general dimensions of O.f. costaricensis are about 
the same as those of O.f. fulvescens; the color difference is slight but 
the upperparts are usually more distinctly tawny. The most dis- 
tinctive feature seems to be the larger molars and resulting elongation 
of the toothrows, a character shared with the larger and darker form 
O.f vegetus. u O. nicaraguae" was based on scanty material from 
Nicaragua which is not satisfactorily separable from costaricensis. 
The type, apparently full grown but with molars rather slightly worn, 
is not so tawny as most examples of costaricensis, but this color ele- 
ment is usually less distinct in younger individuals. The toothrows 
are long, as in costaricensis. 

Specimens examined. — Total number, 19, as follows: 

Costa Rica: Boruca, 7; 1 Buenos Aires, 2; 2 Cerro de Buena Vista, l; 2 El General, 

3 (type and topotypes); 2 La Carpintera, l. 2 
Nicaragua: San Rafael del Norte, l; 2 Vijagua, 1 (type of "nicaraguse "). 2 
Panama: La Chorrera, l; 2 Old Panama, 2. 2 

ORYZOMYS FULVESCENS VEGETUS Bangs. 

Volcan Chiriqui Pygmy Rice Rat. 

Oryzomys (Oligoryzomys) vegetus Bangs, Bull. Mus. Comp. Zool., XXXIX, text fig. 15, 
p. 35, April, 1902. 

Type locality. — Boquete, Volcan de Chiriqui, Panama (altitude, 
4,000 feet). 

i Six in Araer. Mus. Nat. Hist.; 1 in Field Mus. Nat. Hist. 2 Collection Amer. Mus. Nat. Hist. 



94 NORTH AMERICAN FAUNA. [No. 43. 

Type. — No. 10298, 9 old adult, Museum of Comparative Zoology 
(Bangs collection) ; collected by W. W. Brown, jr., April 16, 1901. 

Geographic distribution. — Known only from the vicinity of type 
locality; Upper Tropical Zone. 

General characters. — Size larger and color usually darker than in 
0. f costaricensis; skull larger, but molar teeth relatively small. 

Color. — About as in . f. fulvescens and O.f. costaricensis, but gen- 
eral tone averaging darker than either, the tawny element in upper- 
parts distinct as in costaricensis; abdomen varying in the darkest 
examples to a shade darker than light ochraceous-buff of Ridgway. 

Slcull. — Decidedly larger than that of O.f. costaricensis, with molar 
teeth actually about the same size, therefore relatively smaller. 

Measurements. — Type: Total length, 210; tail vertebrae, 120; hind 
foot, 25. Average of six adult topotypes: 216 (205-235); 122 (115- 
130) ; 24.2 (24-25) . SJcull (average of 5 adults — type and 4 topotypes) : 
Greatest length, 23.6 (22.7-24.3); zygomatic breadth, 12.3 (11.9- 
12.8); interorbital breadth, 3.8 (3.7-4.2); width of braincase, 10.3 
(10-10.5) ; nasals, 8.7 (8.3-9.3) ; anterior palatine foramina, 3.6 (3.4- 
3.9) ; palatal bridge, 3.9 (3.8-4.3) ; upper molar series, 3.1 (3.1-3.1). 

Remarks. — Larger average size and tendency toward darker colora- 
tion usually distinguish O.f. vegetus from the more northern forms of 
the group, but there seems to be no sharp line of demarcation between 
it and O.f. costaricensis. Both share the heavier dentition as com- 
pared with 0. f. fulvescens. Four specimens in the topotype series 
were referred by Bangs to costaricensis. They are the paler examples 
and in color apparently do not differ from costaricensis, but the larger 
size of the skulls indicates that they belong with the remainder of the 
series of vegetus. 

Specimens examined. — Twenty-two, 1 from type locality. 

Subgenus MELANOMYS Thomas. 

Melanomys Thomas, Ann. Mag. Nat. Hist., ser. 7, X, p. 248, Sept., 1902; Novitatea 
Zool., X, no. 1, p. 41, Apr. 20, 1903 (subgenus). Type Oryzomys phxopus 
Thomas. 

Melanomys Allen, Bull. Amer. Mus. Nat. Hist., XXXII, p. 533, Nov. 17, 1913 (genus). 

Geographic distribution. — Andean region of northwestern South 
America from southern Ecuador to northern Colombia, and north- 
westward through Panama and Costa Rica to northern Nicaragua; 
altitudinal range from near sea level to about 8,000 feet; mainly in 
Upper and Lower Tropical Zones. 

Subgeneric characters. — Color very dark, upperparts and under- 
parts not strongly contrasted; form robust; tail about three-fourths 
length of head and body, black all around; feet (epidermis and hair) 
blackish to base of claws, which are light horn color; hind feet 
broad, stout, the digital bristles not projecting be} T ond ends of claws. 

i Fifteen in Mus. Comp. Zool.; 7 in Field Mus. Nat. Hist. 



1918.] SUBGENUS MELANOMYS. 95 

Skull rotund; rostrum short, nearly straight; braincase large and 
inflated; frontals broad, the lateral margins projecting as supraor- 
bital shelves; zygomata slender, but maxillary root decidedly 
expanded above along frontal and premaxillary sutures; antorbital 
foramen rather narrow above, but little compressed and opening 
widely below, the outer wall projecting slightly forward as viewed 
from above; lachrymal articulating mainly with maxilla; mastoid 
process of squamosal short and broad owing to slight excision of 
squamosal border. Molars slightly hypsodont; parastyle and proto- 
cone of first upper molar with oblique commissures; secondary 
parastyle rudimentary or fused with main element; incisors thin 
and weak, the upper ones descending about perpendicularly from 
premaxillae, the points not decidedly curved backward. 

Remarks. — This group of dark-colored forms, segregated by Thomas 
under the name Melanomys as a subgenus of Oryzomys, was elevated 
to generic rank by Allen hi 1913. In general characters, and espe- 
cially dentition, however, it approaches typical Oryzomys so closely 
that subgeneric recognition seems better to express the relationship. 
The most notable dental differences appear to be shown in the rather 
high molar crowns, comparatively straight, thin upper incisors, and 
the tendency toward suppression of the secondary parastyle normally 
present hi typical Oryzomys. The secondary parastyle is normally 
fused with the parastyle, which is correspondingly enlarged. Among 
the more important cranial details are the expansion of the maxillary 
root of the zygoma over the antorbital foramen and the resulting 
alteration in the position of the lachrymal as compared with typical 
Oryzomys. The lachrymal hi Melanomys articulates almost entirely 
with the maxilla, its position being anterior to the fronto-m axillary 
suture, while in typical Oryzomys it bridges this suture and about 
evenly overlaps the maxilla and frontal. 

The subgenus Melanomys differs so strikingly from the subgenus 
Oligoryzomys in appearance that no close comparison is necessary, 
although most of the characters of the latter are found hi varying 
combinations in the subgenus Oryzomys. 

ORYZOMYS CALIGINOSUS (Tomes). 

Hesperomys caliginosus Tomes, Proc. Zool. Soc. London, 1860, p. 263. Type from 

coast of Ecuador, probably Esmeraldas. 1 Type No. 7.1.1.128, British Museum; 

collected by Louis Fraser. 
Akodon caliginosus Allen, Bull. Amer. Mus. Nat. Hist., XIV, p. 45, Jan. 31, 1901. 
Hesperomys (Melanomys) caliginosus Thomas, Ann. Mag. Nat. Hist. ser. 8, XT, p. 406, 

Apr. 1913. 
Melanomys caliginosus caliginosus [et al. subsp.] Allen, Bull. Amer. Mus. Nat. Hist., 

XXXII, p. 537, Nov. 17, 1913. 

» See Allen (op. cit., pp. 533-554, 1913) for locality and revision of group regarded as generically distinct. 



96 



NORTH AMERICAN FAUNA. 



[No. 43. 



Geographic distribution. — (See under subgenus Melanomys) (fig. 11). 
General characters. — (See subgeneric characters under subgenus 
Melanomys.) 

Color. — Fresh pelage: Upperparts varying from tawny to russet, 
abundantly but finely mixed with black, the black tending to become 
predominant especially on rump; underparts overlaid with varying 
shades from dark ochraceous-tawny to cinnamon brown, the dark 
plumbeous basal color showing through; vibrissse black, scarcely 
reaching posteriorly to ears; ears, feet, and tail thinly clothed with 
short hairs, the hairs and epidermis blackish. Young (in first pelage) : 
Similar to adults, but upperparts usually more distinctly blackish, 

and underparts less heavily 
overlaid with ochraceous- 
tawny or cinnamon brown. 

Skull. — (See subgeneric 
characters under subgenus 
Melanomys.) 

Remarks. — In a revision of 
the" Melanomys" group, Allen * 
assigns the component parts to 
several specific types on the 
basis of characters which seem 
comparatively unimportant; 
the slight departures exhibited 
either in color or cranial de- 
tails, in examples from widely 
separated regions, point to the 
probable inclusion of most, if not all, of the forms in the species 
0. caliginosus. Two subspecies range within the limits of the region 
under review. 

Key to Subspecies of O. caliginosus (Exclusive of South America). 
a 1 . Upperparts tending toward tawny; frontal region narrower. (Eastern Panama.) 

O. c. idoneus (p. 96). 

a 2 . Upperparts tending toward russet; frontal region broader. (Western Panama; 

Costa Rica; Nicaragua.) O. c. chrysomelas (p. 97). 

ORYZOMYS CALIGINOSUS IDONEUS Goldman 

Cerro Azul Dusky Rice Rat. 

( PI. IV, figs. 7, 7a; PI. V, figs. 3, 12; PI. VI, figs. 9, 9a.) 

Oryzomys idoneus Goldman, Smiths. Misc. Coll., LVI, No. 36, p. 5, January 19, 1912. 

Melanomys idoneus Allen, Bull. Amer. Mus. Nat. Hist., XXXII, p. 548, November 

17, 1913. 

Type locality. — Cerro Azul, near headwaters of Chagres River, 
Panama (altitude, 2,500 feet). 

Type. — No. 171106, 9 adult, United States National Museum 
(Biological Survey collection); collected by E. A. Goldman, March 
26, 1911. 




1. O. c. idoneus. \ 2. 0. c. chrysomelas. 

Fig. 11. — Geographic distribution of subspecies of 
Oryzomys caliginosus (exclusive of South American 
forms). 



'See Allen (op.cit., pp. 533-554, 1913) for locality and revision of group regarded as generically distinct. 



1918.] ORYZOMYS CALIGINOSUS. 97 

Geographic distribution. — Heavily forested mountain slopes in 
eastern Panama; known altitudinal range from 1,800 to 2,800 feet; 
Humid Lower Tropical Zone. 

General characters. — Similar to 0. c. caliginosus, of Ecuador, but 
upperparts paler, more tawny, instead of russet in color. Closely 
resembling 0. c. chrysomelas, but color paler; skull narrower between 
orbits. 

Color. — Fresh pelage: Upperparts near tawny (becoming most 
intense in worn pelage), heavily and finely mixed with black, the 
mixture usually uniform, but black predominating on rump in some 
examples; underparts, including inner sides of limbs, dark ochra- 
ceous-tawny; inner sides of ankles black; ears, feet, and tail thinly 
clothed with short hairs, the hairs and epidermis blackish. Young 
(in first pelage) : Upperparts usually more distinctly blackish than 
adults, the tawny element restricted to narrower tips of hairs. 

Skull. — Similar to that of 0. c. caliginosus, but frontal region 
narrower; molars slightly larger, the toothrows longer; anterior 
palatine foramina usually shorter. Closely resembling that of 0. c. 
chrysomelas, but frontal region more constricted, the supraorbital 
borders less projecting laterally. 

Measurements. — Type: Total length, 218; tail vertebrae, 88; 
hind foot, 30. Average of 10 adults from Cana, Panama: 217.9 
(196-240); 93.3 (85-105); 26.7 (26-27.5). STcuU (average of same): 
Greatest length, 30.1 (28.6-30.3); zygomatic breadth, 16.2 (15.1- 
16.7); interorbital breadth, 6.3 (5.9-6.5); width of braincase, 11.8 
(11.2-12.3); nasals, 11.4 (10.7-12.6); anterior palatine foramina, 
4.6 (4.4-5); palatal bridge, 6.1 (5.4-6.8); upper molar series, 4.7 
(4.4-5). 

Remarks. — While 0. c. idoneus differs from typical 0. c. caliginosus 
very appreciably in color, close relationship is evidenced in cranial 
characters, some skulls being scarcely distinguishable. In color it 
closely approaches 0. c. columbianus, of the Santa Marta region of 
Colombia, but is slightly darker; the tail is usually shorter. In 
cranial characters similarity to columbianus is also shown, but the 
ascending branches of the premaxillse are narrower posteriorly, the 
ends externally beveled in outline along the interdigitating fronto- 
premaxillary fine of contact. 

Specimens examined. — Total number, 47, as follows: 

Panama: Cana, 46; Cerro Azul, 1 (type). 

ORYZOMYS CALIGINOSUS CHRYSOMELAS (Allen). 
Costa Rican Dusky Rice Rat. 

(PI. IV, figs. 8, 8a.) 

Hesperomys (Habrothrix) caliginosus Allen, Bull. Amer. Mue. Nat. Hist , III p 210 

Apr. 17, 1891 (Costa Rica). ' *' ' 

Oryzomys chrysomelas Allen, Bull. Amer. Mus. Nat. Hist., IX, p. 37, Mar. 11, 1897. 
Oryzomys (Melanomys) chrysomelas Allen, Bull. Amer. Mus. Nat. Hist., XXIV. p 654 

Oct. 13, 1908 (Nicaragua). e 

Zygodontomys chrysomelas Bangs, Bull. Mus. Comp. Zool., XXXIX, p. 37, April, 1902 

(Panama). 
Melanomys chrysomelas Allen, Bull. Amer. Mus. Nat. Hist., XXXII, p. 547. Nov 17 

1913. " ' 

Zogodontomys (sic) chrysomelas Allen, Bull. Amer. Mus. Nat. Hist., XXXII, p. 547. 

Nov. 17, 1913. * ' 

14521°— 18 7 



98 NORTH AMERICAN FAUNA. [No. 43. 

Type locality. — Suerre, Costa Rica (altitude probably between 3,000 
and 4,000 feet). 

Type. — No. VkW> $ adult, American Museum of Natural History ; 
collected by Anastasio Alfaro, July 16, 1895. 

Geographic distribution. — Western Panama, Costa Rica, and north 
to northern Nicaragua ; altitudinal range from near sea level to about 
3,000 feet, mainly in Humid Lower Tropical Zone. 

General characters. — Most closely allied to 0. c. idoneus, but slightly 
darker in color; skull broader between orbits. 

Color. — About as in 0. c. idoneus, but rufescent element in upper- 
parts darker and inclining toward the russet tone of the paler species 
of 0. c. caliginosus. 

Skull. — Similar to that of 0. c. idoneus, but frontal region less con- 
stricted, the supraorbital borders more projecting laterally. 

Measurements. — Average of five adults from Escondido River (45 
miles from Bluefields), Nicaragua: Total length, 210 (202-217); tail 
vertebrae, 89 (87-90); hind foot, 25.9 (25-27). SJcull (average of 
four adults from same locality): Greatest length, 29.1 (28.2-29.5); 
zygomatic breadth, 15.9 (15.8-16.1); interorbital breadth. 6.6 (6.3- 
6.8); width of braincase, 11.8 (11.4-12.2); nasals, 10.9 (10.1-11.5); 
anterior palatine foramina, 4.9 (4.8-5.1); palatal bridge, 5.6 (5.3- 
5.8); upper molar series, 4.5 (4.3-4.7). 

Remarks. — The range of 0. c. chrysomelas marks the northern limit 
of the species in Central America. The Costa Rican subspecies is 
most closely allied to its near geographic neighbor in eastern Panama, 
although in color the darker specimens approach some of the paler 
examples of 0. c. caliginosus and other South American forms. 

Specimens examined. — Total number, 61, as follows: 

Costa Rica: Pacuare, 5; Port Limon, 1; San Carlos, l; 1 Suerre, 5 (type and topo- 

types); 2 Talamanca (probably near Sipurio), 4. 
Nicaragua: Chontales, 3; 1 Escondido River (45-50 miles from Bluefields), 14; 

Greytown, 1; Savala, 2; 1 Pena Blanca, 3; 1 Rio Coco, 3; 1 Rio Grande, l; 1 

RioTumaS; 1 Tuma, 6: 1 Vijagua, 6. 1 
Panama: Mosca, 1. 

Plate I. 

[All subgenus Oryzomys. Natural size; all in U. S. Nat. Mus., Biological Survey collection, except 

figs. 2, 2a.] 

Figs 1 la. Oryzomys palustris palustris (Harlan). Georgetown, South Carolina, c? 
adult. (No. 71367.) 

2, 2a. Oryzomys palustris coloralus Bangs. Type. Cape Sable, Florida. <? 

adult. (No. 4470, Mus. Comp. Zool.) 

3, 3a. Oryzomys couesi couesi (Alston). Yaruca, Honduras, c? adult. (No. 

131814.) 

4 4a. Oryzomys couesi regillus Goldman. Type. Los Reyes, Michoacan. <? 

adult. (No. 125945.) 

5 5a. Oryzomys couesi aquaticus Allen. Topotype. Brownsville, Texas. <? 

adult. (No. 65066.) 

6 6a. Oryzomys gatunensis Goldman. Type. Gatun, Canal Zone, c? young. 

(No. 17i034.) . 

7 7a. Oryzomys cozumelse Merriam. Type. Cozumel Island, Mexico, c? 

adult. (No. 108462.) . . 

8 8a Oryzomys peninsulse Thomas. San Jose del Cabo, Lower California. ? 

adult. (No. 146618.) ^^ 

i Collection Amer. Mus. Nat. Hist. „.,,,, „ T * ,-,. t 

» Four (including type) in Amer. Mus. Nat. Hist.; 1 in Field Mus. Nat. Hist. 



North American Fauna No. 43, U. S. Dept. Agr. Biological Survey. 



Plate I. 




Skulls of Oryzomys. 



1, la. O. p. palustris. 

2, 2a. O. p. coloratus. 

3, 3a. O. c. couesi. 
4,4a. O.c. regillus. 



5, 5a. O. c. aquatieus. 

6, 6a. O. gatunensis. 

7, 7a. O. eozumelse. 

8, 8a. O. peninsulse. 



North America 


n Fauna No. 43, U. S. Dept. Agr. 1 


iiological Survey. 


Plate II. 


Ir- 


II 


»] 


il 


il 


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Skulls of Oryzomys. 



1, la. O. nelsoni. 

2, 2a. O. m. melanotis. 

3, 3a. O. m. colimensis. 

4, -ia. O. r. rostratus. 



5, 5a. O. r. megadon. 

6, 6a. O. r. yucatanensis. 

7, 7a. O. b. bombycinus. 
8,8a. O. b. alleni. 



Plate II. 

[All subgenus Oryzomys Natural size; all in U. S. Nat. Mus., Biological Survey collection, except 

figs. 8, 8a.] 

Figs. 1, la. Oryzomys nelsoni Merriam. Type. Maria Madre Island, Mexico, cf 
adult. (No. 89200.) 

2, 2a. Oryzomys melanotis melanotis Thomas. Topotype. San Sebastian, Ja- 

lisco, c? adult. (No. 88062.) 

3, 3a. Oryzomys melanotis colimensis, subsp. nov. Type. Armeria, Colima. 

$ adult. (No. mn~) 

4, 4a. Oryzomys rostratus rostratus Merriam. Type. Metlatloyuca, Puebla. 

$ adult. (No. 93112.) 
5, 5a. Oryzomys rostratus megadon Merriam. Type. Teapa, Tabasco. <? 
adult. (No. 99978.) 

6, 6a. Oryzomys rostratiis yucatanensis Merriam. Type. Chichen Itza, Yuca- 

tan. <? adult. (No. 108139.) 

7, la. Oryzomys bombycinus bombycinus Goldman. Topotype. Cerro Azul, 

Panama. $ adult. (No. 171103.) 

8, 8a. Oryzomys bombycinus alleni Goldman. Type. Tuis, Costa Rica. <? 

subadult. (No. ffft, Amer. Mus. Nat. Hist.) 



Plate III. 
[All subgenus Oryzornys. Natural size; all in U. S. Nat. Mus., Biological Survey collection. 

Figs. 1, la. Oryzornys alfaroi alfaroi (Allen). Yaruca, Honduras. $ adult. (No. 
131817.) 

2, 2a. Oryzornys alfaroi dariensis Goldman. Type. Cana, Panama. $ adult. 

(No. 178660.) 

3, 3a. Oryzornys alfaroi rhabdops Merriam. Type. Calel, Guatemala, <? 

adult. (No. 76813.) 
4, 4a. Oryzornys alfaroi caudatus Merriam. Type. Comaltepec, Oaxaca. d 
adult. (No. 68641.) 

5, 5a. Oryzornys alfaroi palatinus Merriam. Type. Teapa, Tabasco. ? adult. 

(No. 99977.) 

6, 6a. Oryzornys alfaroi dilutior Merriam. Type. Huauchinango, Puebla. S 

adult. (No. 93124.) 
7, 7a. Oryzornys guerrerensis Goldman. Type. Omilteme, Guerrero, c? 

adult. (No. 127517.) 
8, 8a. Oryzornys hylocetes Merriam. Type. Chicharras, Chiapas. S adult. 

(No. 77605.) 



North American Fauna No. 43, U. S. Dept. Agr. Biological Survey. 



Plate III. 




Skulls of Oryzomys. 



1, la. O. a. alfaroi. 

2, 2a. O. a. dariensis. 

3, 3a. O. a. rhabdops. 
4,4a. O. a. caudalus. 



5, 5a. O. a. palatums. 
6, 6a. O. a. dilution 

7, 7a. O. guerrerensis. 

8, 8a. O. hylocetes. 



North American Fauna No. 43, U. S. Dept. Agr. Biological Survey. 



Plate IV. 




Skulls of Oryzomys. 



1, la. O. devius. 

2, 2a. O. pirrensis. 

3, 3a. O. talamancae. 

4, 4a. O. t. frontalis. 



5, 5a. O. f. fulvescens. 

6,6a. O.f. lenis. 

7, 7a. O. c. idoneus. 

8, 8a. O. c. chrysomelas. 



Plate IV. 

[Natural size; all in U. S. Nat Mus., Biological Survey collection, except figs. 1, la.J 

Figs. 1, la. Oryzomys (Oryzomys) devius Bangs. Type. Boquete, Panama. 9 adult. 
(No. 10324, Mus. Comp. Zool.) 

2, 2a. Oryzomys (Oryzomys) pirrensis Goldman. Type. Mount Pirre, Panama. 

c? adult. (No. 178993.) 

3, 3a. Oryzomys (Oryzomys) talamancse, Allen. Gatun, Canal Zone, c? adult. 

(No. 170981.) 

4, 4a. Oryzomys (Oryzomys) tectus frontalis Goldman. Type. Corozal, Canal 

Zone. ? adult. (No. 171531.) 

5, 5a. Oryzomys (Oligoryzomys) fulvescens fulvescens (Saussure). Orizaba, Vera 

Cruz, d 1 adult. (No. 63688.) 

6, 6a. Oryzomys (Oligoryzomys) fulvescens lenis Goldman. Type. Los Reyes, 

Michoacan. (No. 125941.) 

7, 7a. Oryzomys (Oligoryzomys) caliginosus idoneus Goldman. Cana, Panama. 

c? adult. (No. 178608.) 

8, 8a. Oryzomys (Oligoryzomys) caliginosus chrysomelas (Allen). Port Limon, 

Costa Rica. <? adult. (No. 76265.) 



PLATfe V. 

[Natural size; all in U. S. Nat. Mus., Biological Survey collection.] 

Fig. 1. Oryzomys (Oryzomys) palustris palustris (Harlan). Georgetown, South Caro- 
lina. <? adult. (No. 71367.) 

2. Oryzomys (Oligoryzomys) fulvescens fulvescens (Saussure). Orizaba, Vera 

Cruz. <? adult. (No. 58248.) 

3. Oryzomys (Melanomys) caliginosus idoneus Goldman. Cana, Panama. <? 

adult. (No. 178663.) 

4. Oryzomys (Oryzomys) palustris palustris (Harlan). Greenwich, New Jersey. 

<? adult. (No. 117384.) 

5. Oryzomys (Oryzomys) melanotis melanotis Thomas. San Sebastian, Jalisco. 

<? adult. (No. 88061.) 

6. Oryzomys (Oryzomys) alfaroi dariensis Goldman. Cana, Panama, d adult. 

(No. 178657.) 

7. Oryzomys (Oryzomys) talamancx Allen. Gatun, Canal Zone. 9 adult. (No. 

170979.) 

8. Oryzomys (Oryzomys) bombycinus bombycinus Goldman. Cerro Azul, Panama 

9 adult. (No. 171103.) 

9. Oryzomys (Oryzomys) pirrensis Goldman. Mount Pirre, Panama. <$ adult. 

(No. 178996.) 

10. Oryzomys (Oryzomys) tectus frontalis Goldman. Cana, Panama. <$ adult. 

(No. 178649.) 

11. Oryzomys (Oligoryzomys) fulvescens fulvescens (Saussure). Orizaba, Vera 

Cruz. <? adult. (No. 58248.) 

12. Oryzomys (Melanomys) caliginosus idoneus Goldman. Cana, Panama. <? 

adult. (No. 178684.) 



North American Fauna No. 43, U. S. Dept. Agr. Biological Survey. 



Plate V. 




VgBjJJ 



s^M 



Skulls and Mandibles of Oryzomys. 



1,4. O. p. palustris. 

2. 11. O. f. fulvescens. 

3. 12. O. c. idoneus. 

5. O. m. melanotis. 

6. O. a. dariensis. 



7. O. talamancse. 

8. O. b. bombycinus. 

9. O. pirrensis. 
10. O. t. frontalis. 



North American Fauna No. 43, U. S. Dept. Agr. Biological Survey. 



Plate VI. 





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1, la. O. p. palustris. 

2, 2a. O. m. melanotis. 

3, 3a. O. a. dariensis. 



Molars of Oryzomys. 

4, 4a. O. talamancse. 

5, 5a. O. b. bombycinus. 

6, 6a. O. pirrensis. 



7, 7a. O. t. frontalis. 

8, Sa. O. f. fulvescens. 

9, 9a. O. c. idoneus. 



Plate VI. 

Slightly retouched and enlarged about eight diameters; all in U. S. Nat. Mus., Biological Survey 

collection.] 

Figs. 1, la. Oryzomys (Oryzomys) palustris palustris (Harlan). Dismal Swamp, Vir- 
ginia. <? subadnlt. (No. 75203.) 

2, 2a. Oryzomys (Oryzomys) melanotis melanotis Thomas. Ixtapa, Jalisco, cf 

subadult. (No. 88068.) 

3, 3a. Oryzomys (Oryzomys) alfaroi dariensis Goldman. Cana, Panama. <? sub- 

adult. (No. 178653.) 

4, 4a. Oryzomys (Oryzomys) talamancn Allen. Cana, Panama. 9 subadult. 

(No. 179601.) 

5, 5a. Oryzomys (Oryzomys) bombycinus bovibyeinus Goldman. Cerro Azul, 

Panama. 9 adult. (No. 171103.) 
6,6a. Oryzomys (Oryzomys) pirrensis Goldman. Mount Pine, Panama. 9 sub- 
adult. (No. 179030.) 

7, 7a. Oryzomys (Oryzomys) tectus frontalis Goldman. Cana, Panama. 9 sub- 

adult, (No. 178646.) 

8, 8a. Oryzomys (Oligoryzomys) fulvescens fuhescens (Saussure). Orizaba, Vera 

Cruz, c? subadult. (No. 58244.) 

9, 9a. Oryzomys (Melanomys) caliginosus idoneus Goldman. Cana, Panama. 

9 subadult. (No. 178668.) 



INDEX. 



[New names in bold-faced type; synonyms in italics; principal references in bold-faced figures.] 



Age, 7. 

Akodon caliginosus, 95. 

albi venter, Oryzomys couesi,30,34,35,36,37,38,40. 

alfaroi, Hesperomys (Oryzomys), 59. 

Oryzomys (species), 58-59, 74. 

Oryzomys alfaroi, 8, 59-61, 61, 62, 63, 66, 67, 68. 
alleni, Oryzomys bombycinus, 77, 78. 
angusticeps, Oryzomys alfaroi, 60, 63-63, 64 , 65, 71 . 
antillarum, Oryzomys, 44-45, 87. 
apatelius, Oryzomys jalapx, 30. 
aquaticus, Oryzomys couesi, 36, 39, 39-40. 
Arvicola, 8. 

oryzivora, 9, 22. 

palustris, 9. 
aztecus, Oryzomys couesi, 34, 35-36, 36, 37. 



bombycinus, Oryzomys (species), 76-77. 

Oryzomys bombycinus, 77, 78. 
bulleri, Oryzomys, 33. 



caliginosus, Akodon, 95. 

Hesperomys, 95 ; 97. 

Melanomys caliginosus, 95. 

Oryzomys (species), 95-96. 

Oryzomys caliginosus, 97, 98. 
carrikeri, Oryzomys, 73, 74. 
caudatus, Oryzomys alfaroi, 64-65, 68, 70. 
chapmani, Oryzomys alfaroi, 59, 60, 65, 66, 67, 67-68, 

69, 70. 
Characters, general, 5-7. 
chrysomelas, Melanomys, 97. 

Oryzomys caliginosus, 97, 97-98. 

Zogodontomys, 97. 

Zygodontomys, 97. 
colimensis, Oryzomys melanotis, 50, 51, 51-52. 
Color, 6, 10. 

coloratus, Oryzomys palustris, 23, 25, 26, 27. 
columbianus, Oryzomys caliginosus, 97. 
costaricensis, Oryzomys fulvescens, 90, 92-93, 94. 
couesi, Hesperomys, 9, 29. 

Oryzomys (species), 28-29, 30, 36, 42, 44, 47, 48. 

Oryzomys couesi, 8, 19, 20, 29-31, 32,33, 34, 35, 
37, 38, 39, 40, 43, 44. 
cozumelse, Oryzomys, 43. 
crinitus, Oryzomys couesi, 30, 35, 36-37, 38, 40, 42. 

D. 

dariensis, Oryzomys alfaroi, 60, 61-62. 
devius, Oryzomvs, 80, 80-81, 81, 82. 
dilutior, Oryzomys alfaroi, 67, 68, 68-69. 



E. 



Economic relations, 4-5. 
Explanations, 10-11. 

F. 

flavicans, Oryzomys flavicans, 83, 84. 
floridanus, Oryzomys natator, 26. 
frontalis, Oryzomys tectus, 85, 85-86. 
fulgens, Oryzomys, 36, 41-42. 
fulvescens, Hesperomys, 9, 89. 

Oryzomys (species), 88-89. 

Oryzomys fulvescens, 89-91, 91, 92, 93, 94. 



G. 

gatunensis, Oryzomys, 42-43. 
Geographic variation, 8. 
goldmani, Oryzomys, 30. 
gracilis, Oryzomys, 59, 60, 61, 62. 
Groups, 14-15. 

Oryzomys alfaroi, 15, 56-71, 76. 

bombycinus, 6, 15, 73, 75-78. 

caliginosus, 95-98. 

devius, 15, 78-82. 

fulvescens, 88-94. 

longicaudatus, 86. 

melanotis, 15, 47-55, 76. 

meridensis, 80. 

palustris, 6,14-15, 18-46, 57, 71, 72, 75, 76, 
79 v 80. 

pirrensis, 6. 

talamancaj, 6, 15, 56, 57, 71-74, 75, 76, 79, 
83 

tectus, 15, 82-86. 
guerrerensis, Oryzomys, 58, 59, 69-70. 



Habits. 4-5. 

Habrothrix caliginosus, 97. 

Hesperomys, 9, 11. 

alfaroi, 59. 

caliginosus, 95, 97. 

couesi, 9, 29. 

fulvescens, 9, 89. 

palustris, 22. 
History, 8-10. 
hylocetes, Oryzomys, 59, 64, 70-71. 

I. 

idoneus, Melanomys, 96. 

Oryzomys caliginosus, 96-97, 98. 
illectus, Oryzomys flavicans, 84. 
incertus, Oryzomys alfaroi, 59, 60. 
Individual variation, 7. 
Introduction, 3-11. 



jalapx, Oryzomys, 29. 



lenis, Oryzomys fulvescens, 90, 91. 
Life zones, 11. 

List of species and subspecies, 16-17. 
longicaudatus, Oryzomys, 87. 

M. 

maculiventer, Oryzomys, 80, 82. 

Material, 8-10. 

mayensis, Oryzomys fulvescens, 92. 

Measurements, 10. 

medius, Oryzomys, 73. 

megadon, Oryzomys rostratus, 53, 54, 54-55, 55. 

Megadontomys, 80. 

Melanomys, genus, 11. 

subgenus, 4, 5, 11, 12, 13, 17, 94-98. 
Melanomys caliginosus, 95. 

chrysomelas, 97. 

idoneus, 96. 
melanotis, Oryzomys (species), 49-50, 52, 53, 87. 

Oryzomys melanotis, 50-51, 51, 52. 
meridensis, Oryzomys, 80, 82. 

99 



100 



NORTH AMERICAN FAUNA. 



[No. 43, 1918.] 



mexicanus, Oryzomys couesi, 8, 30, 31, 33, 33-35, 

35,36,37,38,45,46. 
molestus, Oryzomys, 38. 
mollipilosus, Oryzomys, 73, 74. 
Molt, 6-7. 
Mus palustris, 8, 9, 11, 22. 

N. 

natator, Oryzomys palustris, 23, 24, 24-25, 26, 27. 

navus, Oryzomys, 11. 

Neacomys, 13. 

Nectomys, 13. 

nelsoni, Oryzomys, 46-47. 

Nesoryzomys, 13. 

nicaragux, Oryzomys (Oligoryzomys), 92, 93. 

Nyctomys, 14. 

O. 
03eomys, 83. 

Oligoryzomys, subgenus, 4, 5, 11, 14, 17, 87-94, 95. 
oryzivora, Arvtcola, 9, 22. 
Oryzomys, genus, 3, 11-17. / 

subgenus, 4, 11, 16, 17-87, 87, 88, 95. 
Oryzomys albiventer, 30, 34, 35, 36, 37, 38, 40. 

alfaroi, 8, 59-61, 61, 62, 63, 66, 67, 68. 

alleni, 77, 78. 

angusticeps, 60, 62-63, 64, 65, 71. 

antillaruni, 44-45, 87. 

apatelius, 30. 

aquaticus, 36, 39, 39-40. 

aztecus, 34, 35-36, 36, 37. 

bombycinus, 76-77, 77, 78. 

bulleri, 33. 

caliginosus, 96, 97, 98. 

carrikeri, 73, 74. 

caudatus, 64-65, 68, 70. 

chapmani, 59, 60, 65, 66, 67, 67-68, 69, 70. 

chrysomelas, 97, 97-98. 

colimensis, 50, 51, 51-52. 

coloratus, 23, 25, 26, 27. 

columbianus, 97. 

costaricensis, 90, 92-93, 94. 

couesi, 8, 19, 20, 28-29, 29-31, 32, 33, 34, 35, 36, 
37, 38, 39, 40, 42, 43, 44, 47, 48. 

cozumela;, 43. 

crinitus, 30, 35, 36-37, 38, 40, 42. 

dariensis, 60, 61-62. 

devius, 80. 80-81, 81, 82. 

dilutior, 67, 68, 68-69. • 

flavicans, 83, 84. 

floridanus, 26. 

frontalis, 85, 85-86. 

fulgens, 36, 41-42. 

fulvescens, 88-89, 89-91, 91, 92, 93, 94. 

gatunensis, 42-43. 

goldmani, 30. 

gracilis, 59, 60, 61, 62. 

guerrerensis, 58, 59, 69-70. 

hylocetes, 59, 64, 70-71. 

idoneus, 96-97, 98. 

illectus, 84. 

incertus, 59, 60. 

jalapx, 29. 

lenis, 90, 91. 

longicaudatus, 87. 

maculiventer, 80, 82. 

mayensls, 92. 

medius, 73. 

megadon, 53, 54, 54-55, 55. 

melanotis, 49-50, 50-51, 51, 52, 53, 87. 

meridensis, 80, 82. 

mexicanus, 8, 30, 31, 33, 33-35, 35, 36, 37, 38, 45, 
46. 

molestus, 38. 

mollipilosus, 73, 74. 

natator, 23, 24, 24-25, 26, 27. 

navus, 11. 

nelsoni, 46-47. 

nicaragux, 92, 93. 

palatums, 65-66, 66, 67. 

palmarius, 83, 84. 

palmin?, 59, 61, 62. 

palustris, 3, 4, 5, 21-22, 22-24, 25, 26, 27, 29, 48, 
62, 74. 



Oryzomys panamensis, 73, 74. 
penmsulae, 29, 45-46. 
peragrus, 30, 31, 39, 40. 
pirrensis, 80, 81, 81-82. 
regillus, 34, 36, 37, 37, 38, 40. 
rhabdops, 59, 60, 62, 63, 63-64, 65, 71. 
richardsoni, 30. 
richmondi, 30, 32, 42. 
rostratus, 49, 50, 52, 53-54, 54, 55. 
rufinus, 29. 
rufus, 33. 

saturation 65, 66, 66-67, 71. 
talamancae, 53, 60, 61, 62, 73-74. 
teapensis, 29. 
tectus, 84, 84-85, 85, 86. 
texensis, 23, 25, 26, 27-28. 
vegetus, 93, 93-94. 
victus, 14, 86-87. 
yucatanensis, 52, 53, 54, 55, 55. 
zygomaticus, 30, 31, 32-33, 34. 

P. 

palatums, Oryzomys alfaroi, 65-66, 66, 67. 
palmirag, Oryzomys, 59, 61, 62. 
palmarius, Oryzomys, 83, 84. 
palustris, Arvicola, 9. 

Hespcromys, 22. 

Mus, 8, 9, 11, 22. 

Oryzomys (species), 21-22, 29, 48, 62, 74. 

Oryzomys palustris, 3, 4, 5, 22-24, 25, 26, 27. 
panamensis, Oryzomys, 73, 74. 
Pelage, 6. 

peninsulae, Oryzomys, 29, 45-46. 
peragrus, Oryzomys couesi, 30, 31, 39, 40. 
geromyscus, 14, 80. 
phseopus, Oryzomys, 11, 94. 
pirrensis, Oryzomys, SO, 81, 81-82 

R. 

regillus, Oryzomys couesi, 34, 36, 37, 37, 38, 40. 

Reithrodontomys, 14, 87, 88. 

rhabdops, Oryzomys alfaroi, 59, 60, 62, 63, 63-64, 

65, 71. 
Rhipidomys, 13, 83. 
richardsoni, Oryzomys, 30. 
richmondi, Oryzomys couesi, 30, 32, 42. 
rostratus, Oryzomys (species), 49, 50, 52. 

Oryzomys rostratus, 53-54, 54, 55. 
rufinus, Oryzomys jalapx, 29. 
rufus, Oryzomys, 33. 

S. 

saturatior, Oryzomys alfaroi, 65, 66, 66-67, 71. 
Specimens examined, 9-10. 
Subgenera, 14-15, 17. 

T. 

talamanca?, Oryzomys, 53, 60, 61, 62, 73-74. 
teapensis, Oryzomys, 29. 
tectus, Oryzomys (species), 84. 

Oryzomys tectus, 84-85, 85, 86. 
Teeth, 10, 11 (fig.). 

texensis, Oryzomys palustris, 23, 25, 26, 27-28. 
Tylomys, 84. 

V. 
Variation, 7-8. 

vegetus, Oryzomys fulvescens, 93, 93-94. 
victus, Oryzomys, 14, 86-87. 



yucatanensis, Oryzomys rostratus, 52, 53, 54, 55, 55. 
Z. 

Zogodontomys chrysomelas, 97. 
Zonal divisions, 11. 
Zygodontomys, 14. 
chrysomelas, 97. 
zygomaticus, Oryzomys couesi, 30, 31, 32-33, 34. 



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North American Fauna No. 44, U. S. Dept. Agr. Biological Survey. 



Plate 




Small Eastern Flying Squirrel (Glaucomys volans volans). 
From life; Coram, N. Y., August, 1915; photographed by Francis Harper. 



U. S. DEPARTMENT OF AGRICULTURE 

BUREAU OF BIOLOGICAL SURVEY 

E. W. NELSON, Chief 



NORTH AMERICAN FAUNA 

No. 44- 

[Actual date of publication, June 13, 1918] 




REVISION OF THE AMERICAN FLYING SQUIRRELS 



ARTHUR H. HOWELL 

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



J, S. SUPT. OF DOCUMENT^ 




LETTER OF TRANSMITTAL. 



United States Department or Agriculture, 

Bureau of Biological Survey, 
Washington, D. C, March 9, 1917. 
Sir : I have the honor to transmit for publication as North Ameri- 
can Fauna No. 44 a revision of the American flying squirrels (genus 
Glaucomys) , by Arthur H. Howell, assistant biologist of this bureau. 
The revision is based largely upon material in the collection of the 
Biological Survey. Flying squirrels are found over much of North 
America, and this report on the group gives needed information con- 
cerning the number of forms and their distribution. 

Respectfully, 

E. W. Nelson, 
Chief Biological Survey. 
Hon. David F. Houston, 

Secretary of Agriculture. 



CONTENTS. 



Page. 

Introduction 5 

Habits 5 

Voice 6 

Nests 7 

Breeding 8 

Food 9 

Economic status 10 

Pelage and molt 10 

Explanation of cranial measurements 11 

Material examined 11 

Genus Glaucomys 11 

History and nomenclature 12 

Generic names 12 

Specific names 13 

Generic characters 14 

List of species and subspecies, with type localities 16 

Key to species and subspecies 16 

Descriptions of species and subspecies 18 

Glaucomys volans group 18 

Glaucomys sabrinus group j 29 

Tables of cranial measurements 59 

Explanation of plates 62 

Index 63 

3 



ILLUSTRATIONS. 



[Plate I, frontispiece ; Plates II-VII, following page 62.] 
Plate I. Small eastern flying squirrel (Glaucomys volans volans). 

II. Skulls (dorsal view) of Glaucomys volans saturatus, G. v. volans, G. v. texensis, 
G. v. goldmani, G. v. querceti, G. sabrinus bangsi, G. s. alpinus, G. s. 
canescens, G. s. yukonensis, G. s. macrotis, G. s. sabrinus, and G. s. mak- 
kovikensis. 

III. Skulls (dorsal view) of Glaucomys sabrinus stephensi, G. s. flaviventris, G. s. 

klamatliensis, G. s. fuliginosus, G. s. lascivus, G. s. zaphseus, G. s. colum- 
biensis, G. s. latipes, G. s. californicus, G. s. olympicus, G. s. oregonensis, 
and G. s. bullatus. 

IV. Skulls (ventral view) of Glaucomys volans saturatus, G. v. volans, G. v. 

texensis, G. v. goldmani, G. v. querceti, G. sabrinus bangsi, G. s. alpinus, 
G. s. canescens, G. s. yukonensis, G. s. macrotis, G. s. sabrinus, and G. s. 
makkovikensis . 
V. Skulls (ventral view) of Glaucomys sabrinus stephensi, G. s. flaviventris, G. s. 
klamathensis, G. s. fuliginosus, G. s. lascivus, G. s. zaphseus, G. s. colum- 
biensis, G. s. latipes, G. s. californicus, G. s. olympicus, G. s. oregonensis, 
and G. s. bullatus. 
VI . Skulls (lateral view) of Glaucomys volans volans, G. v. querceti, G. v. saturatus, 
G. v. texensis, G. sabrinus vmcrotis, G. s. sabrinus, G. s. columbiensis, G. s. 
oregonensis, G. s. zaphteus, G. s. bangsi, G. s. canescens, and G. s. alpinus. 
VII. Skulls (lateral view) of Glaucomys sabrinus lascivus, G. s. flaviventris, G. s. 
klamathensis, G. s. bullatus, G. s. californicus, G. s. olympicus, G. «. 
fuliginosus, G. s. latipes, and G. s. stephensi; mandible of G. s. sabrinus; 
and skull (mandible and dorsal, ventral, and lateral views) of Pteromys 
biichneri. 

TEXT FIGURES. 

Page. 
Fig. 1. Map showing distribution of Glaucomys sabrinus and G. volans 6 

2. Map showing distribution of the subspecies of Glaucomys volans 19 

3. Map showing distribution of the subspecies of Glaucomys sabrinus except 

G. s. bullatus 30 

4. Map showing distribution of Glaucomys sabrinus bullatus 51 

4 



No. 44. NORTH AMERICAN FAUNA. June 13, 1918. 

REVISION OF THE AMERICAN FLYING SQUIRRELS. 



By Arthur H. Howell. 



INTRODUCTION. 

The American flying squirrels comprise two distinct groups — the 
small species (Glaucomys volcms) of eastern United States and Mex- 
ico; and the larger species {G. sdbrinus) occupying western United 
States and Canada, with one form ranging into New England and 
the border States along the Great Lakes (fig. 1). 

HABITS. 

Flying squirrels are almost exclusively arboreal, seldom descending 
to the ground and apparently never running for any distance on its 
surface. They are unsuspicious and, being easily tamed, make very 
attractive pets. 1 Possessed of intense activity, their movements are 
the acme of grace and agility. They do not fly, in the usual sense 
of the term, but progress from tree to tree by gliding with out- 
stretched membranes from an elevated position to a point lower 
down, usually near the ground. Just before alighting, the animal 
checks its momentum by sweeping upward in a gentle curve and 
alights on the tree trunk with its head up. Ascending the tree by 
climbing, it is ready for another flight. In these gliding leaps, 
which may extend for a distance of 50 yards or more, the squirrel is 
able to change its course to one side or the other with perfect ease. 

Unlike all other American squirrels, the flying squirrels are strictly 
nocturnal. During the daytime they remain concealed in their 
nests and are never seen abroad unless frightened from their re- 
treats. Usually they may be driven readily from their holes by 
pounding with an ax or club on the base of the tree in which they are 

1 A pair of flying squirrels regularly visit the sixth floor of an apartment house front- 
ing the National Zoological Park, Washington, D. C, and although usually averse to the 
glare of an electric light, they will when hungry eat nuts in apparent unconcern within 
5 or 6 feet of an observer. They run up and down the stuccoed sides of the building 
with perfect ease, doubtless reaching the middle stories by jumping from a near-by tree. 
Their visits usually are between dark and midnight. 

5 



6 



NORTH AMERICAN FAUNA. 



[No. 44. 



sleeping. Woodchoppers frequently scare them out when felling 
timber, and anyone who camps frequently in the woods is likely to 
hear them at night running or jumping about in the trees or drop- 
ping nuts to the ground. 




IGlAUCOMYS SABRINUS 

IGlaucomys VOLANS 



Fio. 1. — Map showing the distribution of Glaucomys saorinus and O. volana, based on 
specimens examined and reports of occurrence. 

VOICE. 

The notes of Glaucomys volans are described by Vernon Bailey as 
a " fine, whistling squeak," and Wood says of the same species : " The 



1918.] INTRODUCTION. 7 

flying squirrel has three quite diverse calls or cries. It utters the 
usual ' chuck-chuck ' of squirrels, the usual quick, sharp squeak when 
scolding, and also, more rarely, a clear musical note, commonly me- 
lodious and pleasant, but occasionally shrill. This resembles the 
chirp of a bird and may be kept up for ten minutes at a time." 1 
Seton, writing of the northern flying squirrel, G. sabrinus, in Mani- 
toba, says : " The cry of this species is said to be like that of volans, 
which is a prolonged squeak not unlike the complaint of a red-eyed 
vireo whose nest is threatened." 2 

NESTS. 

These squirrels are found only in wooded regions, and for homes 
they depend chiefly on hollow trees or stubs and deserted woodpecker 
nests. They often take up their abode, also, in attics, in outbuild- 
ings, or in boxes constructed for martins or other birds. Audubon 
and Bachman mention finding a number of flying squirrels in crevices 
of rock at Red Sulphur Springs, Va. In the cavity chosen for a 
home a nest is constructed of shreds of bark, dry leaves, moss, feath- 
ers, fur, or other soft material. Outside nests are often built, or the 
deserted nests of birds or of other squirrels utilized. Concerning 
this habit (in Glaucomys volans) Mearns says: 

Not infrequently it builds outside nests, and even lives in them during the 
winter. Some resemble the leaf nests of the gray squirrel externally, though 
there is always a warm lining within; other nests are indistinguishable from 
those of red squirrels; and others still are deftly woven of the softest possible 
materials. 3 

Helme, 4 writing from Long Island, N. Y., and Rhoads, 5 writing 
from Pennsylvania, both mention this habit in volans, and both state 
that an evergreen tree usually is selected as a site for the nest. Ever- 
mann and Clark G speak of finding in Indiana a large globular nest 
of fibrous material situated in the crotch of a small oak ; and King 7 
describes a summer nest built of small twigs and oak leaves, and 
lined with grass, situated about 10 feet from the ground in a small 
oak. 

The larger squirrels of the sabrlnus group also occasionally con- 
struct outside nests, although, as is the case with volans, hollow trees 
usually are preferred. J. Ellis McLellan, at Gold Beach, Oreg., once 
found a flying squirrel occupying a small spherical nest made of 
sticks and moss placed in a small fir tree. Vernon Bailey, at Mc- 
Kenzie Bridge, Oreg., found soft nests of the flying squirrel in the 

iWood, F. E., Bull. Illinois State Lab. Nat. Hist., VIII, pp. 535-536, 1910. 

2 Seton, E. T., Life Hist. Northern Anim., I, p. 441, 1909. 

3 Mearns, E. A., Bull. Amer. Mus. Nat. Hist., X, p. 342, 1898. 

* Helme, A. H., Abstr. Proc. Linn. Soc. New York, Nos. 13-14, p. 23, 1902. 
5 Rhoads, S. N., Proc. Acad. Nat. Sci. Philadelphia, p. 389, 1894. 
8 Proc. Wasbington Acad. Sci., XIII, p. 15, 1911. 
7 King, F. H., Amer. Nat., XVII, p. 36, 1883. 



8 NORTH AMERICAN FAUNA. [No. 44. 

masses of tree moss (Ilypnum) which cover the trunks and branches 
of maple and ash trees in the deepest forest. There were runways up 
the trunks under the mossy fleece and others where the moss had been 
worn away by constant use. At Elk River, Minn., in November, 
1904, Charles Bailey found two of the large flying squirrels (G. 
sabrlnus rnaerotis) occupying a nest of sticks and moss in a small 
tamarack. At Florence, Mont., on June 11, 1910, Clarence Birdseye 
found in a pile of cordwood a flying squirrel's nest containing 4 
young with eyes not yet open. 

Flying squirrels apparently do not hibernate, but are known to lay 
up stores of food and to remain for a large part of the winter within 
their nests. During the w T inter'and to some extent, also, at other sea- 
sons they are somewhat gregarious in habit. At Mullan, Idaho, in 
June, Clark P. Streator started 8 adults (G. s. latipes) from a hollow 
tree and secured 6 of them, all of which proved to be males. Seton 
mentions finding 9 adults (G. s. canescens) in a hollow stub at Car- 
berry, Manitoba. 1 The same habit is possessed by G. volans; Prof. 
U. O. Cox, of Farmland, Ind., found, in November, 15 of these squir- 
rels in a small rotten stub a little higher than a man's head, 2 and 
at Dothan, Ala., in March, the writer started 6 from a similar stub 
in a wooded swamp. Audubon and Bachman record finding 20 in a 
martin box having 8 or 10 apartments, placed on top of a large locust 
tree, the box being occupied, also, by bats and screech owls. 3 Dr. 
J. Schneck states that he once ran as many as 50 out of one den. 4 
Audubon and Bachman narrate an interesting experience with flying- 
squirrels near Philadelphia, Pa., where not less than 200 were ob- 
served at once about sunset engaged in sportive gambols in a grove 
of trees. 5 

BREEDING. 

The number of young produced at a birth varies from 3 to 6 — 
usually 4 or 5. The period of gestation is said to be one month, 6 
but no definite evidence on this point is available. In the Northern 
States the young of GTaueomys volans are brought forth in April, 
and a second brood may be produced in August or September. Ever- 
mann and Clark 7 describe finding small naked young ones in Indiana 
on August 19, and Langdon 8 records young with eyes not yet open 
at Madisonville, Ohio, September 7. 

1 Seton, E. T., Life Hist. Northern Anim., I, p. 441, 1909. 

2 Evermann and Clark, Proc. Washington Acad. Sci., XIII, p. 16, 1911. 
a Audubon and Bachman, Quadr. North Auier., I, p. 220, 1840. 

4 Wood, F. E., Bull. Illinois State Lab. Nat. Hist., VIII, p. 534, 1910. 

5 Audubon and Bachman, Quadr. North Amer., I, p. 218, 1840. 

6 D. W. C, On the Habits of the American Flying Squirrels (Pteromys rolucella Cuv.) . 
Mag. Nat. Hist., IX, pp. 569-572, 1836. 

7 Evermann and Clark, Proc. Washington Acad. Sci., XIII, p. 15, 1911. 

8 Langdon, Frank W., Journ. Cincinnati Soc. Nat. Hist., Ill, p. 303, 1881. 



191S.] INTRODUCTION. 9 

The large flying squirrels of the sabrinus group apparently breed 
somewhat later than voluns. Seton records a specimen taken at 
Winnipeg, Manitoba, on April 21, which contained 6 young ready 
for birth ; and a nest of 1 young, with eyes not opened, found May 15 \ 
Merriam found nursing young of G. s. macrotis about a month old 
in northern New York on June 18. In the Bitterroot Mountains, 
Mont., at 4,500 feet altitude, Clarence Birdseye took a female of 
G. s. barigs?iL, May 17, 1910, containing two large embryos; and in 
the Bitterroot Valley, near Florence, Mont., found a nest of small 
nursing young on June 11. 

FOOD. 

The food of flying squirrels consists in large part of nuts, including, 
probably, most of the native species — chestnuts, acorns, beechnuts, hick- 
ory nuts, pecans, and others. Catesby includes pine seeds and 
persimmon berries in their diet, and Prof. D. E. Lantz states that 
they frequently gather and store seeds of the cultivated cherry. 
Stomachs of three specimens of volans taken in Virginia in January 
contained finely chewed chestnut meats, a few pieces of acorn, and 
several beetle larvae — probably secured with the nuts. Several stom- 
achs taken at Red Fork, Okla., in June contained only remains of 
insect larva?. Buds of trees are said to be eaten in winter, and corn 
or other grain sometimes is taken. Beetles and perhaps other in- 
sects constitute a part of the animals' fare. They have a decided taste 
for meat, and are so frequently caught by fur trappers in meat- 
baited traps set for larger game as to constitute a nuisance. 
Rowley states that he has known the small eastern species {volans) 
to gnaw the edges of meat hung from the rafters of an outhouse. 2 
Flying squirrels are suspected by some observers of destroying the 
eggs of wild birds, and Merriam remarks that the eagerness with 
which these animals seize and feast upon a dead bird placed within 
their reach would indicate that they are not strangers to such a 
repast. 3 King states that in confinement they eat birds' eggs with 
great satisfaction, and relates an instance of the instant killing by 
them of a nestling chipping sparrow placed in their cage. A large 
moth introduced alive was also quickly captured and eaten. 4 

A writer in " Forest and Stream," describing the habits of flying 
squirrels in captivity, states that they were fond of parsley, pansy 
flowers, and apple-tree twigs, the last being' devoured with great 
avidity, the leaves and bark stripped clean from the wood. This 
writer states also that the squirrels were fond of moths and grass- 

1 Seton, E. T., Life Hist. Northern Anim., p. 442, 1909. 

2 Rowley, J., Abstr. Proc. Linn. Soc. New York, Nos. 13-14, p. 40, 1902. 
s Merriam, C. Hart., Trans. Linn. Soc. New York, II, p. 100, 1884. 
'King, F. H., Anier. Nat., XVII, pp. 40-41, 1883. 



10 NORTH AMERICAN FAUNA. [No. 44. 

hoppers, but refused crickets, and their preference in nuts was for 
acorns, filberts, and pecans. 1 

Stores of food laid up by flying squirrels for winter use have occa- 
sionally been found, indicating that the hoarding habit is probably 
general. Audubon and Bachman record an instance of a martin 
box occupied by a considerable number of flying squirrels, in which 
was found a quantity of hickory nuts, chestnuts, acorns, and corn. 2 

ECONOMIC STATUS. 

The food habits of the flying squirrels are such that they are almost 
entirely harmless. So far as known they do not damage farm crops, 
and the relatively small number of nuts which they consume does not 
seriously affect the total supply. They have been suspected of de- 
stroying the eggs and young of wild birds, but no definite evidence 
on this point is forthcoming. Occasionally they may take posses- 
sion of bird boxes or enter the lofts of dwelling houses and thus be- 
come objectionable, but ordinarily their gentle and confiding ways 
and their interesting habits make them desirable neighbors. 

PELAGE AND MOLT. 

The pelage in members of this genus is of moderate length, very 
dense, fine, and silky in texture. Apparently only one complete molt 
occurs in a year, in the autumn, from September to November. The 
new pelage usually appears first on the sides and spreads thence over 
the back, the head and shoulders being the last parts to be renewed. 
The effects of wear usually are not noticeable until March, but from 
then till November many worn specimens are to be found, though 
apparently a considerable proportion retain the winter pelage 
throughout the summer with little indication of wear. The head, 
nape, and rump are first subject to wear, though frequently the entire 
upperparts and tail become much worn. In summer there is a gen- 
eral tendency to a reddening of the hairs on the upperparts. 

In Glaucomys volans and in most, if not all, of the races of G. 
sabrinus the middle portion of the soles in summer is naked, the same 
portions being furred in winter pelage in all forms except the south- 
ern races of volans. In one race (G. volans volans) the hind toes 
are conspicuously marked with white in winter, but are brown in 
summer. The exact manner of this change is not apparent, but it 
probably is accomplished by a partial spring molt. 3 

1 " C. C. H.," Forest and Stream, LII, pp. 125-126, 1899. 

a Audubon and Bachman, Quad. North Amer., I, p. 220, 1846. 

3 For a detailed account of the molt in O. volans see, under that species, pp. 20-22. 



1918.] GENUS GLAUCOMYS. 11 

EXPLANATION OF CRANIAL MEASUREMENTS. 

The following measurements of skulls of this genus have been taken 
in millimeters : 

Greatest length. — From anterior border of nasals to posterior 
border of supraoccipital in median line. 

Zygomatic breadth. — Greatest breadth across zygomata. 

Mastoidal breadth. — Greatest breadth across mastoids. 

Inter orbital breadth. — Shortest distance across frontals in front of 
postorbital processes. 

Postorbital breadth. — Shortest distance across frontals behind 
postorbital processes. 

Length of nasals. — Greatest length of nasals, measured along 
median line. 

Maxillary toothrow. — Alveolar length of maxillary molar-premolar 
toothrow. 

MATERIAL EXAMINED. 

For the present revision the writer lias assembled for study 
1,052 specimens. About half of these are contained in the United 
States National Museum, including the Biological Survey and Mer- 
riam collections. The remainder have been borrowed from other 
American museums and from private collections, so that practically 
all the available material in this country has been at the disposal of 
the reviser. 1 

Genus GLAUCOMYS Thomas. 

Mus Linnaeus, Syst. Nat, Ed. 10, p. 63 (part), 1758. 

Sciurus Pallas, Nov. Spec. Glires, p. 350 (part), 1778. Not Sciurus Linnaeus, 1758. 
Pteromys Tiedemann, Zool., I, p. 451 (part), 1808. Not Pteromys G. Cuvier, 1800. 
Sciuropterus Lesson, Manuel de Mamm., p. 242, 1827 (part) ; Desmarest, Diet. 

Sci. Nat., p. 140, 1827 (part). Not Sciuropterus, F. Cuvier, 1825. 
Olaucomys Thomas, Ann. and Mag. Nat. Hist. (ser. 8), I, p. 5, 1908. Type, 

Mus volans Linnaeus. 

1 For the loan of this material the writer desires to extend thanks to the owners and 
custodians as follows : Dr. J. A. Allen, of the American Museum of Natural History ; 
Messrs. Samuel Henshaw and Outram Bangs, of the Museum of Comparative Zoology ; 
Dr. Witmer Stone, of the Academy of Natural Sciences of Philadelphia ; Messrs. Charles 
B. Cory and W. II. Osgood, of the Field Museum of Natural History ; Dr. Joseph Grinnell, 
of the Museum of Vertebrate Zoology, University of California; Mr. P. A. Taverner, of 
the Victoria Memorial Museum ; Mr. F. Kermode, of the Provincial Museum, Victoria, 
British Columbia ; Dr. W. J. Holland and Mr. W. E. Clyde Todd, of the Carnegie Museum ; 
Dr. A. G. Ruthven, of the University of Michigan ; Mr. C. D. Bunker, of the Kansas 
University Museum ; Mr. Myron H. Swenk, of the University of Nebraska ; Mr. J. D. 
Figgins, of the Colorado Museum of Natural History ; Mr. J. O. Snyder, of Leland 
Stanford, Jr., University ; Dr. Barton W. Evermann, of the California Academy of 
Sciences ; Mr. Henry L. Ward, of the Milwaukee Public Museum ; Mr. W. E. Saun- 
ders, of London, Ontario ; Mr. Edward R. Warren, of Colorado Springs, Colo. ; 
Mr. G. L. Kirk, of Rutland, Vt, and Dr. H. V. Ogden, of Milwaukee, Wis. ; and to 
Mr. Francis Harper for the photograph from which the frontispiece was made. 



12 NORTH AMERICAN FAUNA. [No. 44. 

HISTORY AND NOMENCLATURE. 

The small eastern flying squirrel {Glaucomys volans), which is 
very common in Virginia and the other South Atlantic States, at- 
tracted the notice of the earliest colonists and through their descrip- 
tions early became known to naturalists. Captain John Smith, in 
his History of Virginia, published originally in 1624, refers to the 
animal in the following words : 

A Small beast they have they call Assapanick, but we call them flying Squir- 
rels, because spreading their legs, and so stretching the largenesse of their skins, 
that they have beene seene to fly 30 or 40 yards. 1 

This species was described more or less fully also in the seventeenth 
and eighteenth centuries by numerous other writers, including Ray, 2 
Catesby, 3 and Edwards. 4 Both Catesby and Edwards figured the 
animal, and Catesby described its habits in the following words : 

These Squirrels are gregarious, travelling from one Tree to another in com- 
panies of ten, or twelve together. When I first saw them, I took them for dead 
Leaves, blown one Way by the Wind, but was not long so deceived, when I 
perceived many of them to follow one another in one Direction: They will fly 
fourscore Yards from one Tree to another. They can not rise in their Flight, 
nor keep in a horizontal Line, but descend gradually, so that in Proportion to 
the Distance the Tree, they design to fly to, is from them, so much the higher 
they mount on the Tree they fly from * * *. 

The species of eastern Canada was briefly described by Sagard- 
Theodat in 163G, 5 and that of southern Mexico by Hernandez in 1651. 6 

GENERIC NAMES. 

Placed first by Linnaeus in the genus Mus, the American flying 
squirrels were later associated with Sciurus by Gmelin, Pallas, and 
Shaw. In 1800 G. Cuvier proposed a new genus, Pteromys, to include 
the European flying squirrel (Sciurus volans Linnaeus) and the large 
East Indian species (Sciurus yetaurista Gmelin). The American 
forms were referred by later writers to this genus. F. Cuvier, in 
1823, 7 pointed out the characters distinguishing the American and 
European flying squirrels from the large Asiatic species, and. in 1825 
proposed the name Sciuropterus for the European animal, 8 but this 
name was not generally accepted until Allen adopted it for the 
American species in his monograph of the Sciurido? (1877). 9 This 

1 Smith, John, The Generall Historie of Virginia, &c, p. 27, 1624. 

2 Ray, John, Synop. Quad., p. 215, 1693. 

3 Catesby, Mark, Nat. Hist. Carolina, II, rp. 76-77, 1743. 

4 Edwards, George, Nat. Hist. Birds, IV, p. 191, 1751. 

5 Sagard-Theodat, Gabriel, Hist. Canada, III, p. 745, 1636. 

6 Hernandez, F., Hist. Anim. et Min. Novre., Hisp., p. 9, 1651 : In Rerum Med. Novte Hisp. 
Thesaurus. 

7 Cuvier, P., Mem. Mus. Hist. Nat, Paris, pp. 125-127, 1823. 

8 Cuvier, F., Dents des Mamm., p. 255, 1825. 

9 Allen, J. A., Monogr. North Amer. Rodentia : Rept. U. S. Geol. Surv. Terr., XI, p. 653, 
1877. 



1918.] GENUS GLAUCOMYS. 13 

name continued in current use until 1914, when Miller 1 showed that 
according to modern rules for fixing types it must give place to 
Pteromys of earlier date, the type of the latter having been fixed by 
Fleming, 2 in 1822, on Sciurus volans — the European frying squirrel. 
Thomas, 3 in 1908, in a revision of the genera of flying squirrels sepa- 
rated the xVmerican forms subgenerically from the European members 
of the genus under the name Glaucomys, and in 1915 the present 
writer raised the group to generic rank. 4 

SPECIFIC NAMES. 

Linnaeus, in the 10th edition of his Sy sterna Naturae, 5 described the 
smaller American species under the name Mus volans, basing his 
description mainly on the account of the animal as given by Ray. 
Curiously enough, on a later page of the same work he described the 
European flying squirrel as Sciurus volans. The two species nat- 
urally were united in one genus by later authors, and the duplica- 
tion of the name volans necessitated the renaming of one of them. 
Pallas, therefore, in 1778, 6 proposed the name volucclla for the Amer- 
ican animal, which designation remained in general use until 1890, 
when Jordan revived the original name volans. 7 The larger Amer- 
ican species was named Sciurus hudsonius, in 1788, by Gmelin, 8 but 
this name proved to be preoccupied by Erxleben's Sciurus vulgaris 
hudsonicus applied to the American chickaree, so, in 1801, Shaw pro- 
posed in its place the name sabrinus. During the nineteenth century 
both these names were used for the northern flying squirrel — kud- 
sonius by Baird, Allen, and others and sahsrinus b} T Richardson, Au- 
dubon and Bachman, and others. Finally, in 1891, the nomenclatural 
question involved having been settled, Rhoads 10 revived the name 
sabrinus, which has remained current ever since. In addition to 
Pallas's name volucella, the small species Mus volans Linnaeus was 
several times renamed during the early years of the nineteenth 
centur}'. Tiedemann, in 1808, proposed for it the name Pteromys 
virginianus, 11 and Oken, in 1816, called the species Pteromys ameri- 
cana. 12 

Richardson, in 1828, 13 described a new species, alpinus, from mate- 
rial collected by Thomas Drummond in western Canada, but the fol- 

1 Miller, G. S., Proc. Biol. Soc. Washington, XXVII, p. 216, 1914. 

2 Fleming, J., Philos. Zool., II, p. 190, 1822. 

•Thomas, O., Ann. and Mag. Nat. Hist. (ser. 8), I, p. 5, 1908. 
« Howell, A. II., Proc. Biol. Soe. Washington, XXVIII, p. 109, 1915. 
E Linna>us, Syst. Nat, Ed. 10, I, p. G3, 1758. 
•Pallas, Nov. Spec. Glircs, p. 351, 1778. 

7 Jordan, D. S., Man. Vert. U. S., Ed. 5, 321, 1890. 

8 Gmelin, J. F., Syst. Nat., I, p. 153, 1788. 
Shaw, Gen. Zool., II, p. 157, 1801. 

10 Rhoads, S. N., Amer. Nat., XXVIII, p. 525, 1894. 
u Tiedemann, F., Zool., I, p. 451, 1808. 

12 Oken, Lorenz, Lehrb. der Naturg., II, p. 865, 1816. 

13 Richardson, J., Zool. Journ., Ill, p. 519, 1828. 



14 NORTH AMERICAN FAUNA. [No. 44. 

lowing year he reduced the form to a subspecies of sabrinus. 1 Bach- 
man, in 1839, named oregonensis, a new species from the lower Colum- 
bia River, and after this date no new names were proposed for the 
group for a period of more than 50 years. 

Audubon and Bachman in their great work, the Quadrupeds of 
North America, 2 treated the four species already described — volucella, 
sabrinus, alpinus, and oregonensis. Baird, in his Mammals of North 
America, 3 recognized the same four species (using the name hud- 
sonius, however, in place of sabrinus), but was able to add very little 
to the history of the group. Allen, in his Monograph of the 
Sciuridse, 4 united the four forms into one species, volucella, with 
hudsonius as a variety, placing alpinus and oregonensis in synonymy. 
Bangs, in 1896, revised the eastern members of the genus, raising 
sabrinus again to the rank of a species and describing as new, querceti, 
a southern race of volans, and silus, a supposed new species from 
West Virginia. 5 Rhoads, in 1897, revised the western forms of the 
genus, reviving alpinus as a full species with oregonensis as a sub- 
species and describing three additional races — fuliginosus from the 
Cascade Mountains, Washington; bangsi from central Idaho; and 
calif ornicus from the San Bernardino Mountains, California. 6 

During the next eight years 9 new forms were added by various 
authors to the 9 already recognized — one (goldmwii) as a race of 
volans from southern Mexico, two (macrotis and makkovikensis) as 
races of sabrinus, five {klamathensis, olympicus, lascivus, Stephens i, 
and zaphceus) as races of alpinus, and one (yukonensis) as a full 
species. 

In 1915 Swenk proposed as new Pteromys volans nebrascensis, 
and the present writer described two races of volans, four of sabrinus, 
and one full species, bullatus, 7 thus raising the total number of recog- 
nized forms to 26. In the present revision all of these are recognized 
except silus of Bangs and nebrascensis of Swenk, which are referred 
to G. volans volans. 

GENERIC CHARACTERS. 

Form sciurine; fore and hind limbs connected by a broad fold of 
skin extending from wrists to ankles, supported anteriorly by a 

1 Fauna Boreali-Amer., I, p. 195, 1829. 

2 Audubon and Bachman, Quad. North Amer., I, pp. 132, 216, 1846 ; III, pp. 202, 206, 
1854. 

3 Baird, S. F., Mamm. North Amer. : Rept. Expl. and Surv. R. R. Pac, VIII, pp. 286- 
290, 1857. 

* Allen, J. A., Monogr. North Amer. Rodentia : Rept. U. S. Geol. Surv. Terr., XI, 
pp. 653-666, 1877. 

6 Bangs, O., Proc. Biol. Soc. Washington, X pp. 162-166, 1896. 

Rhoads, S. N. f Proc. Acad. Nat. Sci. Philadelphia, pp. 314-327, 1897. 

7 Howell, A. II., Proc. Biol. Soc. Washington, XXVIII, pp. 109-114, 1915 ; in the present 
paper bullatus is treated as a subspecies of sabrinus. 



1918.] GENUS GLAUCOMYS. 15 

slender cartilaginous process growing from the wrist. Tail broad, 
much flattened, densely haired, its sides usually nearly parallel or 
slightly narrowed at each end, the tip evenly rounded. Soles with 
4 tubercles at base of toes ; no metatarsal pad ; palms with 5 tuber- 
cles — 3 at base of fingers, one at base of the rudimentary thumb, and 
one opposite the latter on outside of wrist. Mammae, 8, as follows: 
Pectoral, | ; abdominal, § ; inguinal, ^. Skull with smoothly rounded 
braincase, flattened in some species, subglobular in others; nasals 
abruptly depressed at tip; dorsal outline of skull from nasals to 
postfrontal region nearly straight, then abruptly depressed to 
occiput; frontals long and narrow, the interorbital region decid- 
edly longer than broad; interorbital constriction pronounced, 
usually with a distinct notch at anterior base of postorbital 
processes; postorbital processes broad at base, tapering abruptly to 
a point, slightly depressed at tip; zygomata moderately expanded 
posteriorly, less widely anteriorly; audital bulla) large, subcircular 
in outline, smoothly rounded ; pterygoids slender and rather low, the 
hamular processes usually touching audital bullae; dentition, I,f; 
PM, -f; M, f=22; maxillary toothrows approximately parallel; 
anterior upper premolar very small, terete, its crown simple; trans- 
verse ridges of upper molariform teeth continuous. 

The skull of Glaucomys differs from that of Pteromys 1 in the 
following particulars (see PI. VII, figs. 11-14) : Rostrum with sides 
not parallel, but tapering gradually from tip to point of union with 
the zygomata; nasal branches of premaxillae relatively much wider; 
zygomata narrower anteriorly, not standing out squarely from the 
rostrum ; maxillary arm of zygoma much weaker, the median portion 
of the arch nearly vertical (nearly horizontal in Pteromys) ; frontals 
relatively longer and narrower, the surface nearly flat or with 
a very slight depression; postorbital processes broader at the base 
and less attenuate; braincase more swollen, its outlines smooth and 
rounded; occiput not noticeably depressed; anterior palatine fora- 
mina relatively smaller; mandible relatively longer and more slen- 
derly built, the masseteric ridge less strongly developed, terminating 
at a point slightly posterior to the premolar ; angular process weaker, 
its border forming a regular curve (not abruptly bent outwards, as 
in Pteromys)-, coronoid process larger and longer; enamel pattern 
of molariform teeth simple, the transverse ridges of the upper molars 
complete (in Pteromys PM>, MA, and M^ having the posterior ridge 
cut by a deep reentrant fold). 

1 As restricted to the small palaearctic species P. volans, P. buchneri, and related forms. 



16 NORTH AMERICAN FAUNA. INo. 44. 

List of Species and Subspecies, with Type Localities. 

Glaucomys volans group: 

G lav corny s volans volans (Linnaeus) Virginia. 

volans saturatus Howell Dothan, Alabama. 

volans qucrceti (Bangs) Citronelle, Florida. 

volans texensisHoweW Sour Lake, Texas (7 miles north- 
east ) . 

volans gold mani (Nelson) Teopisca, Chiapas (20 miles 

southeast). 
Glaucomys sabrinus group: 

Glaucomys sabrinus sabrinus (Shaw) Mouth of Severn River, Ontario. 

sabrinus makkovikensis (S^rnborger) -Makkovik, Labrador. 

sabrinus macrotis (Mearns) Hunter Mountain (Catskills), 

Greene County, New York. 

sabrinus cancscens Howell Portage la Prairie, Manitoba. 

sabrinus bangsi (Rhoads) Idaho County, Idaho. 

sabrinus alpinus (Richardson) Jasper House, Alberta. 

sabrinus yukonensis (Osgood) Camp Davidson, Yukon River, 

Yukon. 

sabrinus zaphams (Osgood) Helm Bay, Cleveland Peninsula, 

Alaska. 

sabrinus oregonensis (Bachman) "Pine woods of the Columbia, 

near the sea " — probably near 
the present site of St. Helens, 
Oregon. 

sabrinus columbiensis Howell Okanogan, British Columbia. 

sabrinus fid iginosus (Rhoads) Cascade Mountains, near Martin 

Station, Kittitas County, 
Washington. 

sabrinus latipes Howell Glacier, British Columbia. 

sabrinus olympicus (Elliot) Happy Lake, Olympic Mountains, 

Washington. 

sabrinus bullatus Howell Sawtooth [Alturas] Lake, Idaho. 

sabrinus klamathensis (Merriam) Fort Klamath, Oregon. 

sabrinus flavivcntris Howell Head of Bear Creek, Trinity 

County, California. 

sabrinus lascivus (Bangs) Tallac, Eldorado County, Cali- 
fornia. 

sabrinus calif or nicus (Rhoads) Squirrel Inn, San Bernardino 

Mountains, California. 

sabrinus stephensi (Merriam) Sherwood, Mendocino County, 

California. 

Key to Species and Subspecies. 

a 1 . Hairs on helly white at roots (volans group). 

b\ Colors paler (drab, pinkish cinnamon, or yellowish wood brown). 

&. Size larger; hind toes white in winter pelage volans (p. 19). 

c 2 . Size smaller ; hind toes not white in winter. 
d\ Skull relatively short and broad (greatest length, 32.7-34 mm.) ; bulla? 

smaller texensis (p. 27). 

d*. Skull relatively long and narrow (greatest length, 33.2-35.3) ; bull* 
larger . querceti (p. 26). 



1918.] KEY TO SPECIES AND SUBSPECIES. 17 

6 a . Colors darker (dark sayal brown; hair brown; pale snuff brown). 

c 1 . Sides of face marked with fuscous goldmani (p. 28). 

c 3 . Sides of face not marked with fuscous saturatus (p. 24). 

a 2 . Hairs on belly gray at roots (sabrinus group). 

&\ Bulla? very large bullatus (p. 51). 

b 2 . Bullae medium or small. 
&. Soles yellowish. 

d\ Underparts strongly yellowish flaviventris (p. 54). 

d 2 . Underparts not strongly yellowish. 
e\ Underparts tinged with yellow or cream-buff. 

f. Under side of tail darker (deep pinkish cinnamon), 

columbiensis (p. 45). 
f. Under side of tail paler (light pinkish cinnamon or colonial 

buff) klamathensis (p. 52). 

e*. Underparts tinged with avellaneous lascivus (p. 55). 

c*. Soles not yellowish. 

d 1 . Underparts heavily washed with wood brown or some shade of 
cinnamon or buff. 
e 1 . Upperparts strongly rufescent (mikado brown or pecan brown), 

oregonensis (p. 44). 
e 2 . Upperparts not strongly rufescent. 

f. Tail strongly clouded with fuscous beneath zaphaeus (p. 43). 

f. Tail not strongly clouded with fuscous beneath. 
g\ Size larger (greatest length of skull usually more than 42 mm.), 

latipes (p. 48). 
g*. Size smaller (greatest length of skull usually less than 42 mm.). 
h\ Colors darker (wood brown to snuff brown), 

olympicus (p. 49). 

h 2 . Colors paler (pale sayal brown) fuliginosus (p. 47). 

d 3 . Underparts whitish, or moderately washed with wood brown or some 
shade of cinnamon or buff. 
e\ Upperparts vinaceous or brownish (pinkish cinnamon, vinaceous- 
cinnamon, or sayal brown). 

f. Upperparts very pale (light pinkish cinnamon) canescens (p. 37). 

f. Upperparts darker (vinaceous-cinnamon to sayal brown). 
g 1 . Underparts white or faintly washed with pinkish cinnamon or 
wood brown. 
h\ Size larger (greatest length of skull in adults more than 
39 mm.). 
i\ Under side of tail strongly washed with fuscous or dark 
brown. 
j 1 . Size larger (greatest length of skull usually more than 

40.4 mm.) makkovikensis (p. 34). 

f. Size smaller (greatest length of skull usually less than 

40.4 mm.) sabrinus (p. 31). 

i*. Under side of tail not strongly washed with fuscous (pale 
or deep pinkish cinnamon). 

f. Upperparts vinaceous-cinnamon columbiensis (p. 45). 

j'. Upperparts sayal brown stephensi (p. 57). 

h*. Size smaller (greatest length of skull less than 39 mm.), 

macrotis (p. 35), 

14520°— 18— No. 44 2 



18 NORTH AMERICAN FAUNA. [No. 44. 

g 9 . Underparts moderately washed with pinkish cinnamon or 
avellaneous. 
h\ Size larger (average length of tail about 165 mm.), 

yukonensis (p. 41). 
h'. Size smaller (average length of tail about 142 mm.), 

bangsi ( p. 38 ) . 
er. Upperparts drab, gray, or wood brown. 

f. Size larger; tail heavily marked with fuscous alpinus (p. 40). 

f. Size smaller; tail not heavily marked with fuscous. 

g\ Under side of tail deep pinkish cinnamon bangsi (p. 38). 

g 2 . Under side of tail pinkish buff or pale olive-buff. 
h 1 . Skull with deeper braincase; soles never washed with yellow 

or buff . stephensi (p. 57). 

h 2 . Skull with shallower braincase ; soles often washed with 
yellow or buff. 

i\ Larger and darker lascivus (p. 55). 

i 2 . Smaller and paler californicus (p. 56). 

DESCRIPTIONS OF SPECIES AND SUBSPECIES. 

Glaucomys volans Group. 

GLAUCOMYS VOLANS (Linnaeus). 
[Synonymy under subspecies.] 

General characters. — Size small (hind foot 32 mm. or less) ; hairs 
on underparts white to the roots, except on hind legs and flying mem- 
branes, where they are plumbeous at base. 

Cranial characters. — Skull small (greatest length 35.7 mm. or less) ; 
rostrum short ; audital bullae moderately inflated. 

Color. — Upperparts drab, pinkish cinnamon, sayal brown, pale 
snuff brown, hair-brown, or yellowish wood brown, the bases of the 
hairs deep neutral gray ; sides of face smoke gray, often shaded with 
fuscous or buff; borders of flying membrane above, fuscous, clove 
brown, or blackish brown ; fore feet buffy white, hair-brown, or gray- 
ish buff ; hind feet hair-brown, fuscous, or mouse gray, the toes in one 
race white in winter; tail above, hair-brown, snuff brown, verona 
brown, fuscous, or drab; beneath, pinkish cinnamon, vinaceous- 
cinnamon, or pinkish buff; underparts creamy white, the sides often 
edged with pinkish cinnamon, vinaceous-cinnamon, or pinkish buff. 

Geographic distribution. — Eastern United States and extreme 
southern Canada, from southern New Hampshire, northern New 
York (Lewis County), southern Ontario (London), central Michigan, 
northern Wisconsin (Burnett County), and central Minnesota (Ait- 
kin County), south to the Gulf coast and southern Florida (Fort 
Myers); west to eastern Nebraska (Otoe and Nemaha Counties), 
eastern Kansas (Douglas and Woodson Counties), southwestern 
Oklahoma (Wichita Mountains), and eastern Texas (Parker, Bas- 
trop, and DeWitt Counties) ; also the highlands of Chiapas and 



1918.] 



GLAUCOMYS VOLANS GROUP. 



19 



Guatemala and probably locally throughout eastern Mexico. Zonal 
range from the Transition through Upper and Lower Austral Zones 
(% 2). 

GLAUeOMYS VOLANS VOLANS (Linnaeus). 

Small Eastern Flying Squirrel. 

[PI. I ; PI. II, fig. 2 ; PI. IV, fig. 2 ; PI. VI, fig. 1.] 

[Mus] volans Linnaeus, Syst. Nat., ed. 10, p. 63, 1758 (based chiefly on Ray and 

Edwards). 
Sciurus volucella Pallas, Nov. Spec. Glh-es, p. 351, 177S (apparently a renaming 

of Mus volans Linnaeus, but based primarily on Edwards and Catesby). 
Ptcromys virginianus 

Tiedemann, Zool., 

I, p. 451, 1808 (re- 
naming of Mus vo- 
lans Linnaeus). 
Pteromys americana 

Oken, Lehrb. der 

Naturg., II, p. 865, 

1816 (renaming of 

Sciurus volucella 

Pallas). 
Pteromys volucella Des- 

marest, Nouv. Diet. 

d ' H i s t . Nat., 

XXVII, p. 406, 

1818; Audubon and 

Bachman, Quad. 

North Amer., I. p. 

216, 1846, plate 

XXVIII ; Baird, 

M a m m . North 

Amer. : Kept. Expl. 

and Surv. It. R. 

Pac, VIII, p. 286, 

1857. 
Sciuropterus vol ucella 

Lesson, Manual de 

Mamm., p. 242, 1827. 
Sciuropterus americanus 

Desmarest, Diet. Sci. 

Nat., p. 140, 1827. 

? Pteromys cucullatus Fischer, Synop. Mamm., p. 365, 1829 (type locality, Vir- 
ginia (?) ; based on the "Sciurus, Virginianus, volans" of Seba). 1 
Sciuropterus volucella, var. volucella Allen, Proc. Boston Soc. Nat. Hist., XVI, 

p. 189, 1874 ; Monogr. North Amer. Rodentia : Rept. U. S. Geol. Surv. Terr., 

XI, p. 655, 1S77. 




Fig. 2. — Map showing the distribution of the subspecies of 
Glaucomys volans, based on specimens examined. 



1 Seba's figure does not agree with any known species of flying squirrel ; Thomas (Proc. 
Zool. Soc. London, p. 148, 1911) has identified it as a young Petaurista, but the flattened 
tail strongly suggests Glaucomys ; Linnaeus cites Seba's description in the synonymy of 
Mus volans. 



20 NORTH AMERICAN FAUNA. I No. 44. 

8[ciuropterus] volans Jordan, Man. Vert. U. S., ed. 5, p. 321, 1890. 

Sciuropterus situs Bangs, Proc. Biol. Soc. Washington, X, p. 163, 1896. Type, 
No. 4931, Mus. Comp. Zool. (formerly in collection E. A. & O. Bangs) ; 
from top of Katis Mountain, near White Sulphur Springs, West Virginia; 
September 2, 1895. 

Pteromys volans nebrascensis Swenk, University Studies (Lincoln, Nebraska), 
XV, p. 151, "April " [= Sept. 25], 1915. Type, No. 286, Coll. State Entomolo- 
gist, Univ. Nebraska ; from Nebraska City, Nebr., Nov. 26, 1914. 

Type locality. — Virginia. 1 

Distribution. — Northeastern United States and extreme southern 
Canada, from central Minnesota, Wisconsin, and Michigan, southern 
Ontario, northern New York 7 (Lewis County), and southern New 
Hampshire south to North Carolina (Raleigh), Tennessee (Nash- 
ville), and northern Arkansas and Oklahoma (Boston Mountains) ; 
west to eastern Nebraska (Otoe and Nemaha Counties) and eastern 
Kansas (Douglas and Woodson Counties). 

Characters. — Size large (for the group); colors pale; upperparts 
varying from drab to pinkish cinnamon; toes, in winter, usually 
strongly marked with white and soles haired to the bases of the toes. 

Color. — Winter pelage: Upperparts varying from drab through 
numerous intermediate shades to pinkish cinnamon, shaded along 
sides with pinkish buff ; hairs on back with an indistinct, subterminal 
band of brown (not visible on the surface) , their bases deep neutral 
gray; sides of face smoke gray, often washed with pinkish buff; 
eye-ring fuscous or clove brown; ears hair-brown or drab, scantily 
haired ; upper surface of flying membrane fuscous to blackish brown ; 
under surface edged with light pinkish cinnamon ; underparts white, 
with a creamy tinge, the hairs white to the roots (except on hind 
legs, where their bases are neutral gray) ; tail, above, snuff brown to 
drab, shading on sides to hair-brown; beneath, light pinkish cinna- 
mon to pinkish buff, shaded in some specimens with drab-gray ; front 
feet buffy white or hair-brown, sometimes shaded with gray; hind 
feet hair-brown (rarely mouse gray), the toes and inner borders of 
feet white. Summer pelage: Similar to the winter pelage, but gen- 
eral tone of upperparts usually darker, caused in part by reddening 
of the hairs and in part by wearing away of the tips, thus exposing 
portions of the underfur; toes without white markings; middle por- 
tion of soles naked. 

Molt. — From the large series of skins of this race which have been 
examined it has been possible to assemble a series of 97 specimens 
representing every month in the year and showing fully all the pelage 
gradations from one season to another. The following notes on the 

1 Fixed by Elliot, Synop. Mamm. North Amer. : Field Columb. Mus., Zool. Ser., II, p. 109, 
1901 (see also Thomas, Proc. Zool. Soe. London, p. 148, 1911). 



1618.] GLAUCOMYS VOLANS GROUP. 21 

pelage of these specimens, arranged chronologically, will serve to 
make clear the time and manner of molting : 

December. — Five specimens, all in full unworn pelage ; hind feet white, the 
soles fully haired. 

January. — Seven specimens, all in full unworn pelage excepting one (Virginia, 
January 18), which is slightly worn on rump; hind feet of all, white; soles 
haired. 

February. — Eight specimens, all in full unworn pelage ; hind feet white and 
soles haired. 

March. — Fourteen specimens; ten in full unworn pelage; four adults (Wash- 
ington, D. C, March 1) show considerable wear on head, nape, and rump; one 
immature specimen (White Sulphur, West Virginia, March 1) is slightly worn 
on head ; all have white feet and hairy soles. 

April. — Thirteen specimens; four in full pelage showing little wear; six 
much worn on head and nape ; two considerably worn on entire upperparts and 
one on flanks; all but one have white feet and all but one (Long Island, New 
York, April 15), hairy soles. 

May. — Seven specimens; two young individuals (Maryland, May 13) in much- 
worn pelage have brown feet, the soles hairy ; one young from Wisconsin (June 
10) is similar, but shows little wear; one adult (West Virginia, May 17) is in 
full pelage, but has brown feet and naked soles ; two adults in full unworn 
pelage, except on head, have white feet but naked soles; one adult (Mount 
Graylock, Mass., May 6) is slightly worn on head, having white feet and 
hairy soles. 

June. — Eight specimens ; four moderately worn ; three in full unworn pelage ; 
one worn on head only; one (Washington, D. C, June 20) practically same as 
winter specimens (except feet) ; feet dusky, without white markings (except 
in one from Massachusetts, June 27, in which white on toes is faintly indicated.) 

July. — Eleven specimens; three young nearly full grown (July 23, 27); 
two from Massachusetts (July 3, 26), practically full-grown young, show con- 
siderable wear ; adults show only moderate wear on body, considerable on 
tail; one (July 28, Washington, D. C.) has pelage as full as in winter, but 
redder ; feet dusky ; soles naked. 

August. — Two specimens; one (Lake George, New York, August 30) is some- 
what worn on head, very little on body; another (Massachusetts, August 30) 
shows considerable wear on upperparts and tail ; feet dusky. 

September. — Eight specimens ; four show little wear ; two young are consid- 
erably worn; one adult (September 6, Maryland) is much worn, especially 
on nape and tail ; one (Lake George, New York, September 15) has practically 
completed the body molt, except on head and a small area in middle of hinder 
back ; feet dusky and soles naked. 

October. — Five specimens; one young (October 21) shows little wear; one 
adult (Sing Sing, N. Y., October 17) apparently still retains the old pelage, 
which is little worn except on head and tail ; feet noticeably whitened, 
but soles naked; three adults (Massachusetts, October 21; Virginia, October 
19) are molting all over the upperparts, the feet still dusky and soles naked. 

November. — Nine specimens ; two immnture individuals from Maryland are 
much worn and in process of molting ; one adult from Maryland ( $ , November 
9) still retains old pelage on back, being much worn on head and nape, but 
fresh pelage seems to be coming in on sides of body ; feet drab without white 
markings and soles naked; one adult (Washington, D. C, November 1) has 
completed the body molt except on head and a patch on nape; the toes on hind 



22 NORTH AMERICAN FAUNA. [No. 44. 

feet are slightly whitened, and the hair is encroaching on the soles ; five adults 
from Connecticut (November 26) are in full winter pelage, with white feet 
and hairy soles. 

Skull. — Largest of the group; superior outline straight from 
nasals to postf rontal region ; braincase moderately depressed, squar- 
ish in general outline. 

Measurements. — Average of 1G adults from District of Columbia 
and adjacent parts of Virginia and Maryland: Total length, 232 
(220-240); tail vertebra, 101 (90-110); hind foot, 30.4 (29-32). 
Skull: Average of 12 adults from District of Columbia: Greatest 
length, 34.9 (34.1-35.6) ; zygomatic breadth, 21 (20.4-22.2) ; mas- 
toidal breadth, 17.1 (16.5-17.7) ; least interorbital breadth, 6.9 (6.2- 
7.6) ; least postorbital breadth, 9.1 (8.6-9.6) ; length of nasals, 9.6 
(9.2-10.1) ; maxillary toothrow, 6.4 (6.2-6.7). 

Type specimen. — None known to exist. 

Remarks. — This flying squirrel has an extensive range, and 
although individual variation within the subspecies is considerable, 
there is practically no variation which can be correlated with faunal 
areas. Large series of specimens are available from nearly all parts 
of its range, and careful comparison fails to reveal any appreciable 
differences between those of the New England States on the one 
hand and those of Minnesota, Iowa, Kansas, and Nebraska on the 
other. On the southern border of the range, however, a gradual 
darkening of the color of the upperparts is noticeable, and also a 
tendency toward elimination of white on the toes in winter pelage. 
Specimens from Virginia (the type region), however, do not differ 
appreciably from New England and New York individuals. Speci- 
mens from Ealeigh, N. C, and from Stillwell, Okla., show decided 
approach in characters to saturatus; a single winter specimen from 
Gainesville, Tex., although approaching texensis in skull characters, 
agrees almost exactly with typical volans in color and in having 
white toes. Since this specimen shows no approach to saturatus 
(which occurs at Eed Fork, in the Arkansas Valley), it seems prob- 
able that typical volans ranges from Kansas into western Oklahoma, 
perhaps as far as the Wichita Mountains, and thence into northern 
Texas. A single specimen in summer pelage from Oklahoma City, 
although somewhat indeterminate in characters, seems to bear out 
this theory of distribution. Additional material from that part of 
Oklahoma is needed to settle the question. 

The type of Sciuropterus silus Bangs has been examined and 
proves to be an immature individual of volans, evidentty a runt. A 
considerable series is now available from the type locality of silus, 
and all are perfectly typical of volans. 

The type of Pteromys volans nebraseensis Swenk has been exam- 
ined and found to agree essentially with t} 7 pical specimens of volans 



1918.] GLAUCOMYS VOLANS GROUP. 23 

from the vicinity of Washington, D. C. It is a rather pronounced 
example of the gray phase, and, although taken on November 26, 
the hind toes have not yet acquired the clear white which charac- 
terizes the winter pelage. The hind feet are somewhat darker than 
the average of volans — in this respect agreeing with saturatus. The 
tail measurement of the type (115 mm.) is about 14 mm. more than 
the average, but only 5 mm. more than the extremes of volans. In 
the dried skin, however, the tail does not measure any longer than 
in dozens of typical specimens. The skull and hind feet measure 
exactly as in typical examples. 
Specimens examined. — Total number, 356, as follows: 

Connecticut: East Hartford, 5; 1 Liberty Hill, 8. 2 

District of Columbia: Cleveland Park, 13; Washington, 23. 

Illinois: Belleville, 1; Chicago, 2; Golconda, 1 ; 3 Olive Branch, l; s Olney, 

2 ; Parkersburg, 1 ; Warsaw, 8 ; Willow Springs, 1.* 
Indiana : Denver, 2 ; 2 ' 4 Kankakee, 1 ; 3 La Porte, 1 ; 3 Wheatland, 1. 
Iowa : Hillsboro, 2 ; Iowa City, 1 ; 5 Knoxville, 3 ; Thayer, 2. 6 
Kansas: Douglas County, 5; 6 Lawrence, ll; 5 Topeka, 4; 8 Woodson 

County, 1.' 
Kentucky: Eubanks, 3. 
Maryland: Bacon Hill, 3; 8 Branchville, 1; Capitol View, 1; Garrett Park, 

1 ; Kensington, 2 ; Laurel, 3 ; Marlboro, 1 ; Marshall Hall, 2 ; Oxon 

Hill, 1 ; Plummer Island, 2 ; Prince Georges County, 3 ; Silver Spring, 2. 
Massachusetts: Ipswich, 1; Mount Graylock, 1; Wareham, 4; 2 Waverly, 

2 ; 2 Wilmington, 2 ; Woburn, 1. 
Michigan: Ann Arbor, 4; 4 Brooklyn, 1 ; 4 Dexter, 1;* Iosca, l; 4 Ithaca, 2; 4 

Kavanagh Lake, 3; 4 Lodi Township, Washtenaw County, l; 4 Napoleon, 

l; 4 Portage Lake, 2; 4 Sand Point, 10 ; 4 Ypsilanti, 1* 
Minnesota: Aitkin, 1 ; 3 Bridgman, 2; Elk River, 19; Fort Snelling, 3; 

Steele County, 2. 8 
Missouri : Independence, 1 ; Stotesbury, I. 2 
New Hampshire: Hancock, 2. 2 
New Jersey: Cape May County, 1 ;° Culvers Gap, Sussex County, l; 8 

Ellenville, 1 ; Haddonfield, 5 ; 8 Lake Hopatcong, 4 ; 8 Seaville, 2. 8 
New York: Ardsley, 3; 3,8 Geneva, 1; Hastings, If Jamaica, 1; Lake 

George, 5; Lake Grove, 2; Locust Grove, 5; Miller Place, 5;* Sing 

Sing, 7 ; Suffolk County, l.« 
North Carolina : Apex, 1 ; 3 Raleigh, 20. 10 
Ohio: Fort Ancient, l; 9 Madisonville, 3; Ravenna, 1." 
Oklahoma : Oklahoma City, 1 ; Stilwell, 7. 
Ontario : Kingsville, 1 ; u London, 7. 11 

1 Three in collection Acad. Nat. Scl. Philadelphia. 

2 Collection Mus. Comp. Zool. 
Collection Field Mus. Nat. Hist. 
* Collection Univ. Michigan. 

6 Collection D. Stoner, Iowa City, Iowa. 

a Collsction Kansas Univ. Mus. 

T Collection Colorado Mus. Nat. Hist. 

8 Collection Acad. Nat. Sci. Philadelphia. 

8 Collection Amer. Mus. Nat. Hist. 

10 Nine in collection Amer. Mus. Nat. Hist. 

11 Collection W. E. Saunders, London, Out. 



24 NORTH AMERICAN FAUNA. [No. 44. 

Pennsylvania: Carlisle, 1; Chester County, 2; Drury Run, Clinton 
County, 1 ; Huntingdon Furnace, 1 ;* Leasuresville, 3 ; l Moon Township, 
Allegheny County, 1 ; 2 Pine Grove Furnace, 1." 

Tennessee: Nashville, l. 3 

Texas: Gainesville, 2. 

Vermont: Castleton, l; 4 Rutland, 2. 4 

Virginia: Cherrydale, 1; Dranesville, 1; Dunn Loring, 4; Falls Church, 8; 
Fort Myer, 1 ; Henrico County, 1 ; Maywood, 1 ; Mount Rodgers, Gray- 
son County, 2 ; Mount Vernon, 7 ; Nelson County. 2 ; 3 Suffolk, 1 ; War- 
wick County, 2. 

West Virginia: Travellers Repose, 2; White Sulphur Springs, 13." 

Wisconsin : Beaver Dam, 2 ;"■ ' Cassville, 1 ; 3 Delavan, 1 ; Lake Koshkonong, 
l; 7 Racine, 1. / 

GLAUCOMYS VOLANS SATURATUS Howell. 

Southeastern Flying Squirrel. 

[PI. II, fig. 1 ; PI. IV, fig. 1 ; PI. VI, fig. 3.] 

Olaucomys volans saturatus Howell, Proc. Biol. Soc. Washington, XXVIII, p. 
110, May 27, 1915. 

Type locality. — Dothan, Alabama. 

Distribution. — Southeastern United States (excepting peninsular 
Florida and the coast region of Georgia) from South Carolina and 
western North Carolina west to central Oklahoma and north in the 
Mississippi Valley to southwestern Kentucky. 

Characters. — Similar in size and skull characters to vola?is, but 
upperparts darker at all seasons; toes not conspicuously whitened in 
winter. Compared with querceti: Colors averaging darker, face 
grayer (less buffy), and hind feet grayer (less brownish); soles 
haired in winter. 

Color. — Winter pelage: Upperparts pale snuff brown or dark sayal 
brown, varying to hair-brown ; sides of face smoke gray, rarely with 
a tinge of Duff; upper surface of flying membrane dark clove brown 
or blackish brown ; hind feet hair-brown, the toes washed with buff ; 
fore feet grayish buff ; tail above, snuff brown, shaded with bister or 
dark hair-brown (rarely solid hair-brown) ; beneath, pinkish cinna- 
mon or vinaceous-cinnamon (rarely light hair-brown, washed with 
pinkish cinnamon) ; underparts creamy white, the under surface of 
the membranes edged with light pinkish cinnamon or light vinaceous 
cinnamon. Summer pelage: Essentially as in winter, but hind feet 



1 Collection Acad. Nat. Sci. Philadelphia. 

2 Collection Carnegie Mus. 

3 Collection Mus. Comp. Zool. 

* Collection G. L. Kirk, Rutland, Vt. 

8 Ten in collection Mus. Comp. Zool. ; two in Field Mus. Nat. Hist. 

6 Collection Mus. Vert. Zool., Univ. California. 

'Collection E. R. Warren, Colorado Springs, Colo. 



1918.] GLAUCOMYS VOLANS GROUP. 25 

fuscous. Variation (March specimens from Milton, Florida) : 
Upperparts rich pecan brown. 

Skull. — Closely similar to that of volans, but averaging smaller; 
practically same size as that of querceti, but bullae averaging smaller. 

Measurements. — Average of 12 adults from southern Alabama: 1 
Total length, 226 (211-244) ; tail vertebra?, 100 (84-109) ; hind foot, 
30 (28-33). Skull: Average of 10 adults from southern Alabama: 2 
Greatest length, 34 (32.8-34.9) ; zygomatic breadth, 20.3 (19.9-21.1) ; 
mastoidal breadth, 17 (16.6-17.5) ; least interorbital breadth, 7 (6.7- 
7.8) ; least postorbital breadth, 8.9 (8.3-10) ; length of nasals, 9.4 
(8.8-10) ; maxillary tooth row, 6.4 (6.2-6.8). 

Type specimen. — No. 178366, U. S. Nat. Mus. (Biological Survey 
collection) ; $ ad., skin and skull; collected March 13, 1912, by A. H. 
Howell. 

Remarks. — This race is the darkest form in the volans group, differ- 
ing in this respect both from the Florida form (querceti) and from 
the Texas form (texensis). It is much darker than volans, both in 
winter and in summer. Its range is chiefly in the Lower Austral 
Zone, but extends into the Upper Austral in the mountains of eastern 
Tennessee and western North Carolina. Intergradation with volans 
is shown by a series from Magnetic City, foot of Roan Mountain, 
North Carolina, and with querceti by a series from Pinetucky, Ga. 
The Pinetucky specimens are typical of saturatus in skull characters, 
but average paler in color. By reason of lack of material from 
Louisiana, the relationship of this race to texensis and its western 
limits in that State can not be defined. Specimens from Eed Fork, 
Okla., are typical but in the Boston Mountains intergradation with 
volans occurs. 

Two specimens in full winter pelage from Milton, Fla., differ from 
typical saturatus in having the upperparts pecan brown instead of 
snuff brown. These at first were thought to represent an undescribed 
race, but the recent acquisition of 3 adult specimens from Muscogee, 
Fla., only a few miles from Milton, which do not differ in any way 
from typical saturatus, leads to the conclusion that the Milton speci- 
mens represent a color phase of this race. 

Specimens examined. — Total number, 89, as follows : 

Alabama: Ardell, 4; Autaugaville, 10; Carlton, 2; Dothan, 6; Greensboro, 
2; Mobile, 4; Perdido River, 1; Sand Mountain, near Carpenter, 2; 
York, 1. 

Arkansas: Delight, 2. 

Florida: Milton, 2; Muscogee, 2. 

Georgia : Columbus, 1 ; Okef enokee Swamp, 2 ; Pinetucky, 13 ; Reidsville, 2 ; 
Young Harris, 1. 

Kentucky: Hickman, 1. 

1 Dothan, Autaugaville, Carlton, and York. 

2 Dothan, Autaugaville, Carlton, and Greensboro. 



26 NORTH AMERICAN FAUNA. I No. 44. 

Louisiana: Mer Rouge, 2; Powhatan Plantation, near Gibson, l. 1 

Mississippi: Columbus, 1; Washington, 1. 

North Carolina: Cranberry, 1; Magnetic City, foot of Roan Mountain, 9. 

Oklahoma: Red Fork, 4. 

South Carolina : Calhoun Falls, 4 ; 2 Cleora, 1 ; Greenville, 1 ; Plantersville, 

3 ; Santee, 1. 
Tennessee: Watauga Valley, 2. 

GLAUCOMYS VOLANS QUERCETI (Bangs). 

Florida Flying Squirrel. 

[PI. II, fig. 9 ; PI. JV, fig. 9 ; PI. VI, fig. 2.] 

Sciuropterus volans querceti Bangs, Proc. Biol. Soc, Washington, X, p. 166, 

Dec. 28, 1896. 
[Sciuropterus rolucella] querceti Trouessart, Cat. Mamm., I, p. 402, 1897. 

Type locality. — Citronelle, Florida. 

Distribution. — Peninsular Florida (south at least to Fort Myers) 
and the coast region of Georgia. 

Characters. — Very similar in size and color to volans, but upper- 
parts averaging slightly darker; toes without conspicuous white 
markings in winter; soles partly naked at all seasons; skull with 
larger audital bullse. 

Color. — Winter pelage: Upperparts sayal brown to yellowish 
wood brown, shaded on sides and fore legs with pinkish buff or light 
pinkish cinnamon; upper surface of flying membrane blackish 
brown; ears fuscous; sides of face smoke gray, often clouded with 
fuscous or buff; hind feet pale fuscous, the toes grayish buff; front 
feet whitish buff; tail above, snuff brown or verona brown, shading 
(when worn) to dark hair-brown ; beneath, pinkish buff or pinkish 
cinnamon; underparts creamy white, often irregularly shaded with 
pinkish buff or pinkish cinnamon, particularly along edges of flying 
membranes. Summer pelage: Essentially the same as in winter, but 
averaging slightly browner (less drab) above. 

Skull. — Closely similar to that of volans, but averaging slightly 
smaller, with larger audital bullae. 

Measurements. — Average of 8 adults from Florida : 3 Total length, 
229 (221-253); tail vertebras, 94 (81-115); hind foot, 31 (30-32). 
Skull: Average of 13 adults from Florida: 4 Greatest length, 34.3 
(33.2-35.3) ; zygomatic breadth, 20.5 (19.1-21.3) ; mastoidal breadth, 
17.4 (16.8-18.4); least interorbital breadth, 7 (6.5-7.3); least post- 
orbital breadth, 8.5 (8.2-8.9) ; length of nasals, 9.4 (8.7-10) ; maxil- 
lary toothrow, 6.4 (5.9-7.1). 

1 Collection Mus. Comp. Zool. 

2 Collection Field Mus. Nat. Hist. 

3 Citronelle and Enterprise. 

4 Citronelle, Enterprise, Lake Harney, and New Berlin. 



1918.] GLAUCOMYS VOLANS GROUP. 27 

Type specimen. — No. 2451, Mus. Comp. Zool. (formerly same num- 
ber, collection E. A. and O. Bangs) ; 2 ad., skin and skull ; collected 
September 17, 1894, by F. L. Small. 

Remarks. — The Florida flying squirrel resembles volans very 
closely in color, but is easily distinguished in winter pelage b} T the 
absence of white markings on the toes and by naked soles. Compared 
with saturatus it averages considerabty paler. The large size of the 
audital bullae seems to be only an average character, for while pro- 
nounced in some specimens, others do not differ in this respect from 
typical volans. The type specimen has larger bulla? than have two 
topotypes and, indeed, can scarcely be matched in this character by 
any of the Florida specimens examined. Intergradation with satu- 
ratus takes place in southeastern Georgia. Specimens from St. 
Marys, Ga., agree in color with querceti, but have smaller bullrc; 
some individuals in the series have the hind toes marked with buffy 
white, but less extensively than in volans. Considerably more mate- 
rial from Florida and Georgia is needed to show the average char- 
acters and exact distribution of this race. 

Specimens examined. — Total number, 67, as follows : 

Florida : Citronelle, 3 ;* Enterprise, 10 ; 2 Fort Myers, 1 ;* Gainesville, 2 ; s 
Hernando County, 2; Lake Harney, 2; Nassau County, 2 ; 3 New Berlin, 
13 ;* Ocala, 1 ; Shell Hammock, 2 ; Tarpon Springs, 9." 

Georgia : Mcintosh County, 1 ; x Montgomery, 4 j 1 St. Marys, 15. 1 - 3 - B> ' 

GLAUCOMYS VOLANS TEXENSIS Howell. 

Texas Flying Squirrel. 

[PI. II, fig. 5 ; PI. IV, fig. 5 ; PI. VI, fig. 4.] 

Olaucomys volans texensis Howell, Proc. Biol. Soc. Washington, XXVIII, p. 110, 
May 27, 1915. 

Type locality. — Sour Lake, Texas (7 miles northeast). 

Distribution. — Eastern Texas, west to Aledo, Gurley, Elgin, and 
Cuero ; eastern limits of range unknown. 

Characters. — Similar in color and size to volans; upperparts 
slightly more ochraceous; toes without conspicuous white markings; 
skull decidedly shorter and broader. Compared with querceti: Sim- 
ilar in color, but skull shorter, with smaller bullae. Compared with 
saturatus : Colors much paler ; skull shorter and broader. 

Color. — Winter pelage (March) : Upperparts yellowish wood 
brown to drab, shaded on sides and fore legs with pinkish buff; upper 
surface of flying membrane fuscous-black; ears fuscous; front feet 

1 Collection Mus. Comp. Zool. 

2 Three in collection Anier. Mus. Nat. nis. ; two in Field Mus. Nat. Hist. 
8 Collection Amer. Mus. Nat. Hist. 

4 Four in collection Amer. Mus. Nat. Hist. ; three in Acad. Nat. Scl. Philadelphia. 

6 Collection Field Mus. Nat. Hist. 

6 Collection Mus. Vert. Zool., Univ. California. 



28 X0K1H AMIKkWX FAUNA. tNo. 44. 

buffy white: hind foot hair-brown or drab, the toes washed with 

bully white; tail above, fuscous, shaded with snuff brown: beneath, 
light pinkish cinnamon; underparts creamy white. Summer pelage: 
A specimen from Colmesniel, Tex. (July BO), agrees essentially with 

the grayest April specimen from Sour Lake: a much worn specimen 
from Tarkington (November 23) is slightly more ochraceous than 

spring specimens. 

Skull. — Similar to that of safuratus. but shorter and relatively 
broader: bra incase more rounded, deeper, and more abruptly de- 
pressed posteriorly. Differs from querceti in the same characters 
and also in having smaller audita] bullae. 

M ensure ?n<nte. — Average of topotypes (adult) : Total length. 
229 (219-236) : tail vertebrae, 104 (97-111) : hind foot, 30.8 (30-31). 
Skull: Average of 5 topotypes (adult) : Greatest length. 33.5 (32.7- 
S4) : zygomatic breadth. -JO. 9 (20.6-21.1) : mastoidal breadth, 17.2 
(17.1-17.4) : least interorbital breadth. 7.2 (6.9-7.3) : least postorbi- 
tal breadth. 8.6 (8.2-S.9) : length of nasals. 9.4 (9.2-9.8) : maxillary 
toothrow, 0.3 (0.1-6.4). 

Type specimen. — Xo. 136400, U. S. Xat. Mus. (Biological Survey 
collection) : 3 ad., skin and skull: collected March 15. 1905. by J. H. 
Gaut. 

Remarks. — The Texas flying squirrel closely resembles both volant 
and quereeti in color, but differs from them in skull characters. It is 
much paler than ioturatus, which occupies the eastern Gulf States 
from Louisiana to Georgia. The material at hand is too scanty to 
show clearly the relationships and exact range of this form. Inter- 
gradation with oolans is indicated by an intermediate specimen from 
Gainesville. Tex. (referred to colons), and doubtless intergradation 
takes place also with satiwatus where their ranges meet. 

> eclmens txam'neJ. — Total number. 10. as follows: 

Texas: Colmesniel. 1 : Sour Lake. 6 : Tarkington. 1 : Texarkana. 1 : Troup, 1. 

GLAUCOMYS VOLANS GOLDMANI (Nelson). 

Mexican Flyixg Squirrel, 

[PI. II, fig. 6; PI. IV, fig. 6.] 

Sciuroptcrus vohms goidmani Nelson, Proo. Biol. Soc. Washington, XVII, p. 
14S, Oct. 6, 1901 

Type locality. — Teopisca. Chiapas. Mexico (20 miles southeast). 

Distribution. — Known only from type locality — probably the high- 
lands of Chiapas and Guatemala. 

Characters* — Closely similar in color to saturatus. but top and 
sides of nose whitish, sides of face marked with fuscous, and upper 
surface of flying membrane more brownish: soles naked: skull essen- 
tiallv like that of solans. 



391%.] GLAUCOMVS SABKINUS GROUP. 29 

Color. — Spring pelage (April specimen, worn on nape and tail 
only): Upperparte dark sayal brown: sides of face smoke gray. 

strongly varied with fuscous; eye-ring fnscoa flesh color, 

shaded with fuscous: top and sides of nose creamy white; upper 
surface of flying membrane clove brown: hind feet fuscous, the toes 
faintly shaded with buff; front feet buffy white; tail, above, snuff 
brown; beneath, pinkish cinnamon or pale cinnamon buff; under- 
parts creamy white, irregularly washed with pinkish cinnamon, most 
strongly along edges of membranes. 

Skull. — Essentially the same as that of uolans — slightly larger than 
that of saturatne. 

Measurement*. — Type and topotype : Total length. 237, 238 ; tail 
vertebrae. 112. 107: hind foot. 30. 31.5. SkuU: Greatest length. 35.1, 
34.9: zygomatic breadth. 21.3. 21/.*: mastoidal breadth. 17.8. 17.5: 
least interorbital breadth, 8.7, 9.2; least postorbital breadth, 8.7, 9.2; 
length of nasals, 10.3. 0.0: maxillary toothrow, 6.4, 0.3. 

Type specimen. — No. 132833. U. S. Nat. Bins. I Biological Survey 
collection) : $ ad., skin and skull: collected April 8. 1004, by E. A. 
Goldman. 

Remarks. — This race, although widely separated geographically 
from the United States forms of the group, bears so close a resem- 
blance to saiuratus that a subspecific designation seems best to ex- 
press its relationship. From no point between southern Texas and 
Chiapas are specimens of the genus known, and Nelson and Goldman 
in their travels in Mexico heard of flying squirrels in only a few local- 
ities. The animals were reported to occur in the Pinal de Amoles 
range of mountains in eastern Queretaro, but whether these are refer- 
able to this race is unknown. Tomes, under the name Pteromya 
volucella. records a specimen from Duenas. Guatemala, which un- 
doubtedly is referable to the present form. 1 

Specimen* examined. — Two, from type locality. 

Glaucomys sabrinus Group. 

GLAUCOMYS SABRINUS (Shaw). 

[Synonymy under subspecie.-.] 

General characters . — Size medium to large (hind foot 34 mm. or 
more) ; hairs on underparts plumbeous at roots. 

Cranixd characters. — Skull medium to large (greatest length 36 
mm. or more) ; rostrum usually rather long (except in macrotis) ; 
audital bullae moderately or in some races considerably inflated. 

Color. — TTpperparts cinnamon, pinkish cinnamon, vinaceous-cinna- 
mon, vinaceous-fawn, cinnamon-buff, avellaneous, drab, cinnamon- 
drab, wood brown, sayal brown, snuff brown, mikado brown, or pecan 

1 Tomes, Robert F., Proc. Zool. Soc. London, 1861, p. 281. 



30 



NORTH AMERICAN FAUNA. 



[No. 44. 



brown; sides (and in some races front) of face smoke gray or pale 
gray, often washed with buff, cinnamon, or fuscous; eye-ring fuscous; 
borders of flying membrane above, clove brown, fuscous, or blackish 




Fig. 



3. — Map showing the distribution of the subspecies of Glaucomys sabrinus (except 
G. s. bull at us, see fig. 4, p. 51). based on specimens examined. 



brown ; forefeet hair-brown, wood brown, drab, or mouse gray ; hind 
feet hair-brown, fuscous, clove brown, wood brown, snuff brown, or 
mouse gray, the soles buffy white, yellowish white, creamy white, 



1918.] GLAUCOMYS SABRINUS GROUP. 31 

drab, mouse gray, cinnamon-buff, colonial buff, olive-ochre, olive- 
buff, or oil yellow; tail above, cinnamon, hair-brown, wood brown, 
sayal brown, snuff brown, clove brown, fuscous, or fuscous-black, 
usually darkest at the tip ; tail beneath, pinkish cinnamon, vinaceous- 
cinnamon, pinkish-, ochraceous-, cinnamon-, colonial-, cream-, or 
olive-buff, drab, drab-gray, wood brown, avellaneous, clove brown, or 
fuscous-black, frequently edged with hair-brown or fuscous; under- 
pays white, creamy white, or buffy white, more or less washed or 
clouded with pinkish cinnamon, pinkish-, ochraceous-, cream-, cinna- 
mon-, colonial-, or cartridge-buff, straw yellow, greenish yellow, 
Naples yellow, wood brown, or avellaneous. 

Geographic distribution. — Wooded parts of Alaska, Canada, 
northeastern United States, and mountains of western United States, 
from the Yukon Valley, Cook Inlet, Upper Mackenzie River, Fort 
Anderson, Great Slave Lake, northern Manitoba, northern Quebec 
(Hamilton River), and Labrador (Makkovik), south to northern 
Massachusetts, southern New York (Catskill Mountains), Pennsyl- 
vania (Erie ?), central Michigan (Montcalm County), northern Wis- 
consin, central Minnesota (Elk River), South Dakota (Black Hills), 
Utah (Uinta 1 and Wasatch Mountains 2 ), Idaho (Sawtooth Moun- 
tains), and southern California (Sierra Nevada, San Bernardino, 
and San Jacinto Mountains) (figs. 3 and 4). 

GLAUCOMYS SABRINUS SABRINUS (Shaw). 

Hudson Bay Flying Squirrel. 

[PI. II, fig. 11 ; PI. IV, fig. 11 ; PI. VI, fig. 6 ; PI. VII, fig. 10.] 

Sciurus hudsonius Gmelin, Syst. Nat., I, p. 153, 1788 (not Sciurus vulgaris hud- 

sonicus Erxleben, 1777). 
Sciurus sabrinus Shaw, Gen. Zool., II, p. 157, 1801. 
? Pteromys canadensis Geoffroy, Cat. Mamm. Mus. Hist. Nat, Paris, p. 170, 

1803. Type locality, North America [probably Quebec, Canada. 3 ] 

1 Allen (Monogr. North Amer. Rodentia, p. 664, 1877) records a specimen from Uinta 
Mountains which has since disappeared from the collection ; this specimen was taken 
September 20, 1870, by H. D. Schmidt, near the head of Smiths Fork, Utah (see Prelim. 
Kept. U. S. Geol. Surv. Wyoming, etc., pp. 41-44, 461, 1871). 

2 F. E. Crandall, of Victor, Idaho, states that while living in Emery County, Utah, he 
often saw flying squirrels and knew of their being killed by loggers when felling trees. 
His report indicates that some form of the group ranges much farther south than has 
been supposed. 

3 The description under this name was based on a specimen (No. CCCLI) in the Paris 
Museum, but as I am informed by Dr. A. Menegaux in a recent letter, this specimen is 
not now in the museum. The description is inadequate, but the size of the animal is 
given as 150 mm. This measurement probably refers to the length of head and body, 
and if so agrees well with that of certain specimens of sabrinus from southern Ontario. 
It is too large for any member of the volans group. The specimen doubtless was sent 
from Canada (probably from the vicinity of Quebec), as suggested by the common name, 
" Le Polatouche du Canada," and the name canadensis may thus be considered a syno- 
nym of sabrinus of two years earlier date. 



32 NOETH AMERICAN FAUNA. {No. 44. 

Ptcromys sabrinus Richardson, Zool. Journal, III, p. 519, 1828 ; Fauna Boreali- 

Amer., I, p. 193, 1829; Audubon and Bachman, Quad.' North Amer., Ill, p. 

202, 1853, plate CXLIII. 
Fteromys hudsonius Fischer, Synop. Mannn., p. 365, 1829 ; Baird, Mamm. North 

Amer. : Rept. Expl. and Surv. R. R. Pac, VIII, p. 288, 1857. 
Sciuropterus volucella var. hudsonius Allen, Proc. Boston Soc. Nat. Hisc., 

XVI, p. 289, 1874 ; Allen, Monogr. North Amer. Rodentia : Rept. U. S. Geol. 

Surv. Terr., XI, p. 655, 1S77. 
Sciuropterus volucella sabrinus Rhoads, Amer. Nat., XXVIII, p. 525, 1894. 
Sciuropterus sabrinus Bangs, Proc. Biol. Soc. Washington, X, p. 162, 1896. 

Type locality. — Mouth of Severn River, Ontario, Canada. 

Distribution. — Interior of Canada, from Fort Simpson (possibly 
Fort Anderson), Mackenzie, and lower Churchill River, west side of 
Hudson Bay, south to northern Minnesota, extreme northwestern 
Wisconsin, southern Ontario (vicinity of Lake Nipissing) , and south- 
ern Quebec (Lake Edward). 

Characters. — Size medium (for the group) ; general tone of upper- 
parts vinaceous-cinnamon ; sides of face smoke gray; tail strongly 
shaded with fuscous; underparts whitish, shaded with drab; skull 
of medium size, rather long and narrow, with narrow braincase. 

Color. — Winter pelage: Upperparts deep vinaceous-cinnamon, the 
sides faintly washed with light pinkish cinnamon, the underfur dark 
mouse gray; upper surface of flying membrane fuscous, the color 
extending over legs and becoming paler on feet; toes shaded with 
grayish white; front and sides of face smoke gray or pale smoke 
gray; eye-ring fuscous; underparts soiled whitish, faintly washed 
with pale yellowish and more or less strongly shaded with drab or 
hair-brown, especially along sides, the underfur deep neutral gray; 
tail, above, hair-brown or fuscous, often shaded near the base with 
vinaceous-cinnamon, the tip usually clear fuscous; beneath, drab, 
shaded with pale wood brown and bordered on sides and tip with 
hair-brown or fuscous ; soles buffy white on inner side, drab on outer 
side. /Slimmer pelage : Similar to the winter pelage. 

Skull. — Size medium (larger than macrotis, smaller than alpinus) ; 
braincase rather narrow and elongate; zygomata not widely ex- 
panded, contracted anteriorly; dorsal outline nearly straight from 
nasals to postf rontal region ; braincase moderately depressed ; audital 
bullae of medium size. 

Measurements. — Adult male from Cochrane, Ontario: Total 
length, 315; tail vertebrae, 140; hind foot (dry), 40; ear from notch 
(dry), 18. Adult female from Lake Edward, Quebec: 315; 148; 43; 
20.5. Average of 4 adults from Norway House, Manitoba: Hind 
foot (dry), 39.5 (38.5-41). Skull: Subadult male from Cochrane, 
Ontario: Greatest length, 40; zygomatic breadth, 24; mastoidal 
breadth, 18.1 ; least interorbital breadth, 7.3 ; least postorbital breadth, 
8.9; length of nasals, 11.5; maxillary toothrow, 7.8. Two adults 



1918.] GLAUCOMYS SABRINUS GROUP. 33 

from vicinity of Great Slave Lake, Mackenzie : Greatest length, 39.9, 
40.2; zygomatic breadth, 24.5, 24.4; mastoidal breadth, 18.8, 19.2; 
least interorbital breadth, 8.3, 8.2 ; least postorbital breadth, 9.4, 9.2 ; 
length of nasals, 12, 12.8; maxillary toothrow, 7.8, 8.1. 

Type specimen. — Not noAv known to exist. 1 

Remarks. — The Hudson Bay flying squirrel — the first form of the 
group to receive a name — is still imperfectly known, no specimens 
from the type locality being as yet available. A good series of skins 
from Norway House, Manitoba, doubtless represent the typical form, 
and these have been used in making comparisons, but there are no 
skulls in any collection from nearer the type locality than Great 
Slave Lake on one side and Cochrane, Ontario, on the other. Since 
these skulls, however, agree essentially in characters, they are assumed 
to be typical. Two winter specimens from Cochrane, Ontario, differ 
from Norway House specimens in being slightly paler and less vina- 
ceous above, having the hind feet mouse gray and the tail dark hair- 
brown, with scarcely a trace di vinaceous ; whether these specimens 
represent the typical race or an unrecognized form can not now be 
decided. 

The present race has a very wide range and apparently inter- 
grades with all the surrounding forms — with makhovilcensis in east- 
ern Quebec, with macrotis in northern Minnesota, northern Wisconsin, 
northern Michigan, and southern Ontario ; with canescens in southern 
Manitoba, with alpinus in southern Mackenzie and Alberta, and prob- 
ably with bangsi in southwestern Alberta and northwestern Montana. 
Specimens snowing intergradation with alpinus have been examined 
from Fort Providence and Fort Simpson, Mackenzie ; and from Cas- 
cade Rapid, Grand Rapids, Edmonton, and Red Deer, Alberta. The 
northern specimens differ from typical sabrinus in being paler and 
grayer ; 2 those from Red Deer are nearly typical in color, but have 
skulls closely resembling those of alpinus. Specimens from Trout 
Creek, Ontario; Hinckley and Itasca County, Minn.; and Gordon, 
Wis., are intermediate between sabrinus and macrotis. A specimen 
from Godbput, Quebec, agrees with sabrinus in color, but has a skull 
equaling that of makkovikensis in size. Specimens from Rat Portage, 
Ontario, and Winnipeg, Manitoba, are paler than the typical form, 
thus showing approach to canescens. 

A specimen in worn and faded pelage, without tail or skull, from 
Fort Liard, Mackenzie, is provisionally referred to this race. Better 
material from that region may show the form occurring there to be 
alpinus. 



1 The names hudsonius and sabrinus were based on Forster's account (Philos. Trans., 
LXII, p. 379, 1772) of a specimen sent to the Royal Society from the mouth of Severn 
River. 

2 Some of these intermediates bear a surprisingly close resemblance in color to batigsi. 

14520°— 18— No. 44 3 



34 NORTH AMERICAN FAUNA. [No. 44. 

Specimens examined. — Total number, 72, as follows: 
Alberta: Calgary, 1; Cascade Rapid, Athabaska River, 1; Didsbury, 1; 

Edmonton, 4 ;* Grand Rapids, Athabaska River, 1 ; Gull Lake, 1 ; a Red 

Deer, 2. 2 
Mackenzie: Big Island, Great Slave Lake, 3; Fort Anderson, 1 ; 8 Fort 

Liard, l; 4 Fort Providence, 5; B Fort Resolution, 1 ; e Fort Simpson, 2; 

Hay River, Great Slave Lake, 2. 
Manitoba : " Hudson Bay," 1 ; Norway House, 9 ; Oxford House, 1 ; Red 

River Settlement, 3; Selkirk Settlement, 1; Winnipeg, 2. 
Minnesota: Hinckley, 2; Itasca County, 8. 7 
Ontario: Cochrane, 2; 8 - 9 Hannah Bay (near Moose Factory), 1 ; 2 Matawag- 

amingue [=Ft. Mattagami], 5; Moose Factory, l; 2 Rat Portage, 1; 

Trout Creek, 2. 8 
Quebec : Godbout, 2 ; Lake Edward, 1 ; 10 Tadousac, l. 6 
Saskatchewan: Cumberland House, 1. 
Wisconsin : Gordon, 1 ; 7 Nemakagan River, Burnett County, l. 11 

GLAUCOMYS SABRINUS MAKKOVIKENSIS (Sornbokgek). 

Labrador Flying •Squirrel. 

TPl. II, fig. 12 ; PI. IV, fig. 12.] 

Sciuropterus sabrinus makkovikensis Sornborger, Ottawa Nat., XIV, p. 48, June 
6, 1900. 

Type locality. — Makkovik, Labrador. 

Distribution. — Coast region of Labrador and eastern Quebec; ex- 
act limits unknown. 

Characters. — Similar to sabrinus, but larger and darker, with 
darker face, tail, and feet. 

Color. — Adults: Upperparts dark vinaceous-cinnamon (averaging 
a shade darker than in sabrinus) ; sides of face smoke gray; upper 
surface of flying membrane clove brown ; feet clove brown to fuscous, 
the soles buffy white and toes washed with the same color; tail 
ftiscous-black or clove brown, both above and below, moderately 
mixed (chiefly near the base) with pinkish cinnamon; underparts 
soiled whitish, faintly washed with light pinkish cinnamon or 
pinkish buff. Young : Upperparts wood brown, shaded with vinace- 
ous-cinnamon ; tail nearly uniform fuscous-black. 

Skull. — Similar to that of sabrinus, but slightly larger. 

Measurements. — Average of 5 specimens (adult and subadult) 
from Labrador: Total length, 309 (293-330); tail vertebrae, 138 

I Collection Victoria Mem. Mus. 
- Collection Mus. Comp. Zool. 

3 Fragments of skeleton ; provisionally referred. 

4 Provisionally referred. 

6 Three in collection Victoria Mem. Mus. 

6 Collection Amer. Mus. Nat. Hist. 

7 Collection Dr. II. V. Ogden, Milwaukee, Wis. 
* Collection Carnegie Museum. 

8 Collection W. E. Saunders, London, Ont. 
10 Collection Field. Mus. Nat. Hist. 

II Collection Milwaukee Pub. Mus. 



1918.] GLAUCOMYS SABRINUS GROUP. 35 

(128-146) ; hind foot, 41.8 (40-45). Skull: Average of 6 adults from 
Labrador: Greatest length, 40.7 (40.4-41.3) ; zygomatic breadth, 24.5 
(23.5-25.2) ; mastoidal breadth, 18.9 (18.6-19.1) ; least interorbital 
breadth, 7.8 (7.3-8.2) ; least postorbital breadth, 10 (9.7-10.3) ; length 
of nasals, 11.4 (10.7-12.2) ; maxillary toothrow, 7.8 (7.3-8.1). 

Type specimens. — Cotypes, Nos. 10476, 10477, Mus. Comp. Zool. 
(formerly Nos. 1540, 1541, collection J. D. Sornborger) ; skins and 
skulls, the skulls imperfect ; collected in 1899 by Bev. W. W. Perritt. 

Remarks. — The Labrador flying squirrel, like certain other species 
from this region, is larger and darker than the form from the in- 
terior of Canada. It is very much larger than macrotis, of New 
England. Intergradation with sab?inus apparently takes place in 
the vicinity of Godbout, Quebec. There is no material to show what 
form occupies the interior of eastern Quebec. 

Specimens examined. — Total number, 15, as follows : 

Labrador: Cartwriglit, 1; L'Anse au Loup, l; 1 Makkovik, 9; 1 Paradise, 3. 
Quebec : Northwest River, 1. 

GLAUCOMYS SABRINUS MACROTIS (Mearns). 

Mearns Flying Squirrel. 

[PI. II, fig. 10; PI. IV, fig. 10 ; PI. VI, fig. 5.] 

Sciiiropterus volucella liudsonius Merriam, Trans. Linn. Soc. New York, II, p. 

108, 1S84 (not Sciurus hudsonius Gmelin). 
Sciuropterus sabrinus macrotis Mearns, Proc. U. S. Nat. Mus., XXI, p. 353, Nov. 

4, 1898. 

Type locality. — Hunter Mountain (Catskills), Greene County, New 
York; altitude 3,300 feet. 

Distribution. — Nova Scotia, New Brunswick, Maine, New Hamp- 
shire, Vermont, northern Massachusetts, Boreal portions of New 
York, northern Pennsylvania ( ? ) , southern Ontario, northern part 
of Michigan, and northeastern Wisconsin ; west to Elk Kiver, Minne- 
sota. 

Characters. — Similar to sabrinus, but smaller; ears averaging 
slightly longer; upperparts and hind feet paler and underparts 
whiter. 

Color. — Winter pelage (adult) : Upperparts cinnamon or pinkish 
cinnamon (rarely light pinkish cinnamon), shading on sides to 
pinkish buff ; sides of face pale smoke gray, this color extending back 
on sides of neck beyond the ears ; top of nose frequently tinged with 
gray; upper surface of membranes clove brown; fore feet drab, 
clouded with dull white ; hind feet, above, light hair-brown or mouse 
gray ; beneath, soiled whitish, shaded with drab or buff ; tail, above, 
dull cinnamon, more or less mixed with hair-brown or fuscous, the 
general tone varying from hair-brown to pale snuff brown ; beneath, 

1 Collection Mus. Comp. Zool. 



36 NORTH AMERICAN FAUNA. [No. 44. 

pinkish cinnamon or light pinkish cinnamon; underparts white, 
irregularly shaded with light pinkish cinnamon. Summer pelage: 
Similar to the winter pelage, but upperparts usually slightly 
darker — pale orange-cinnamon. Young (October) : Upperparts be- 
tween wood brown and fawn color, otherwise as in adult. 

Skull. — Similar to that of sabrinus, but decidedly smaller; brain- 
case less elongate, more nearly spherical. Compared with G. volans 
volans: Slightly larger (greatest length 36 mm. or more in macrotis; 
35.6 mm. or less in volans) ; fronto-parietal region more elevated; 
braincase deeper and less flattened. 

Measurements. — Average of 16 adults from New York and New 
England: 1 Total length, 275.6 (263-290) ; tail vertebrse, 126.4 (115- 
135); hind foot, 36 (34-38); ear from notch (dry), 2 18.8 (16-20). 
Skull: Average of 11 adults from same region: Greatest length, 
37.3 (36-38.7) ; zygomatic breadth, 22.6 (21.9-23.5) ; mastoidal 
breadth, 17.6 (17.2-18) ; least interorbital breadth, 6.9 (6.2-7.3) ; 
least postorbital breadth, 9.2 (8.6-9.9) ; length of nasals, 10.7 
(10-11.5) ; maxillary toothrow, 7 (6.4-7.7). 

Type specimen. — No. 83152, U. S. Nat. Mus. ; 9 ad., skin and skull ; 
collected August 31, 1896, by Edgar A. Mearns. 

Remarks. — The Mearns flying squirrel is a strongly marked form 
having a rather extensive range in the Northeastern States and 
southern Canada, from Minnesota to Nova Scotia. It is not known 
from any point south of the Catskills in New York and the vicinity 
of Erie, Pa., 3 but may be expected to occur in the mountainous parts 
of Pennsylvania and possibly farther south in the Alleghenies. 

Intergradation with sabrinus is shown by specimens from southern 
Ontario (Gooderham) and northern Michigan (Porcupine Mountains 
and Vermilion). Two specimens from Elk River, Minn., show ap- 
proach to canescens, one of them being almost as pale as that form. 
The skulls are nearly t} 7 pical of macrotis. Specimens from Maine 
have somewhat shorter ears than those from New York, Massachu- 
setts, and New Hampshire. 

The range of this form overlaps that of G. volans volans for a con- 
siderable distance in southern New England, New York, Michigan, 
Wisconsin, and Minnesota. Although some specimens approach 
volans rather closely in size and resemble it in the shape of skull and 
shortness of rostrum, they are always readily separable (except, per- 
haps, in the young) from the latter species by the plumbeous bases 
of the hairs on the belly. The upperparts, also, are decidedly more 
vinaceous and less drab than in volans. 

1 Hunter Mountain and Peterboro, N. Y. ; Wilmington, Mass. ; Ossipee, N. H. ; Green- 
ville, Bucksport, and Moosehead Lake, Maine. 

2 Seven specimens from New York, Massachusetts, and New Hampshire. 

3 The specimens labeled as from Erie are without further data, but are said to have 
been in the collection of Geo. B. Sennett ; they may not have been collected in the immedi- 
ate vicinity of Erie. There are no other Pennsylvania records. 






19187] GLAUCOMYS SABRINUS GROUP. 37 

Specimens examined. — Total number, 78, as follows: 

Maine: Bucksport, 3; 1 Greenville, 8; 1 Lincoln, 6; 2 Moosehead Lake, If 
Steuben, 1 ; Third Mopang Lake, 2. 2 

Massachusetts: Lunenburg, 1; Wilmington, 3; Winchendon, 1. 

Michigan : Le Roy, 1 ; 3 Montcalm County, 1 ; 3 Palmer, 1 ; Porcupine Moun- 
tains, 3; 3 Vermilion, 3. 3 

Minnesota: Elk River, 2. 

New Brunswick: No specific locality, 1. 

New Hampshire: Ossipee, 4. 

New York : Adirondack Mountains, 1 ; Big Moose Lake, 3 ; Hunter Moun- 
tain, Greene County, 1 ; Locust Grove, 1 ; Peterboro, 1. 

Nova Scotia: Annapolis, 3 ; 3, * Halifax, 3; Kings County, 5. 5 

Ontario : Gooderham, 3 ; 6 Maganetewan, 1 ; 2 New Edinburgh.l ; 5 Woodstock, l. T 

Pennsylvania: Erie [=mountains near?], 2. 8 

Vermont: Rutland, 2; 9 Sherburne, 1.* 

Wisconsin : Clarks Lake, Door County, 1 ; Kelley Brook, 1 ; 10 Lakewood, 2 ; 
Langlade County, 1 ;"■ Mamie Lake, Vilas County, 2. 

GLAUCOMYS SABRINUS CANESCENS Howell. 

Pallid Flying Squikkel. 

[PI. II, fig. 7 ; PI. IV, fig. 7 ; PI. VI, fig. 11.] 

Olaucomys sabrinus canescens Howell, Proc. Biol. Soc. Washington, XXVIII, 
p. Ill, May 27, 1915. 

Type locality. — Portage la Prairie, Manitoba. 

Distribution. — Southern Manitoba; eastern North Dakota; Black 
Hills, S. Dak.; and Bear Lodge Mountains, Wyo. ; exact limits 
unknown. 

Characters. — Similar to microtis, but much paler, with grayer 
head and larger skull. Compared with sabrinus: Size smaller; 
upperparts and feet paler; underparts whiter. 

Color. — Winter pelage: Upperparts light pinkish cinnamon, shad- 
ing to pale pinkish cinnamon on sides; front and sides of face (some- 
times nearly the whole head) pale smoke gray; ears pale fuscous; 
eye-ring and upper side of flying membrane fuscous; feet hair- 
brown, the toes marked with grayish white; tail, above, dark cin- 
namon, shaded with hair-brown; beneath, light pinkish cinnamon, 
edged with hair-brown ; underparts and soles creamy white. 

1 Collection Mus. Coinp. Zool- 

2 Collection Acad. Nat. Sci. Philadelphia. 
s Collection Univ. Michigan. 

4 Collection E. R. Warren, Colorado Springs, Colo. 

6 Collection Victoria Mem. Mus. 

8 Collection Field Mus. Nat. Hist. 

7 Collection W. E. Saunders, London, Ont. 

8 Collection Amer. Mus. Nat. Hist. 
•Collection G. L. Kirk, Rutland, Vt. 

10 Collection Milwaukee Pub. Mus. 

u Collection Dr. H. V. Ogden, Milwaukee, Wis. 



38 NORTH AMERICAN FAUNA. [No. 44. 

Skull. — Similar to that of macrotis, but slightly longer and rela- 
tively narrower, with longer nasals; decidedly smaller than that of 
sdbrinus. 

Measurements. — Average of 3 adults: 1 Total length, 297 (290- 
306) ; tail vertebrae, 138 (133-146) ; hind foot, 37.7 (37-38). Skull: 
Two topotypes (adult): Greatest length, 38.5, 38.8; zygomatic 
breadth, 22.8, 22.9; mastoidal breadth, 17.5, 17.6; least interorbital 
breadth, 9.2, 9.5 ; least postorbital breadth, 9.4, 9.2 ; length of nasals, 
11.5, 11.2 ; maxillary toothrow, 7.7, 7.6. 

Type specimen. — No. 7663, Field Mus. Nat. Hist. ; 5 subad., skin 
and skull ; collected February 2f, 1900, by G. F. Dippie. 

Remarks. — This is the palest of the races of sdbrinus, occupying 
chiefly the thinly timbered parts of southern Manitoba and 
eastern North Dakota. The limits of its range are not known, and 
may extend westward to Saskatchewan. Two specimens from the 
Bear Lodge Mountains, Wyo., are referred to this race, but a larger 
series may show characters to separate the form in that region. 
These two specimens show some approach to bangsi in skull char- 
acters and in a slightly more vinaceous coloring on the upperparts. 
One of them has a wash of pinkish cinnamon on the underparts. 
In a specimen from Portland, N. Dak., the upperparts are a deeper 
shade of pinkish cinnamon and the skull is larger, showing, appar- 
ently, intergradation with sdbrinus. Two specimens in worn pelage 
from Pembina, N. Dak., are also considered intermediate between 
canescens and sabrinus. 

Specimens examined. — Total number, 10, as follows : 

Manitoba : Carberry, 1 ; Portage la Prairie, 2. 2 

Minnesota: Breckenridge, 1. 

North Dakota : Grafton, 1 ; Pembina, 2 ; Portland, 1. 

Wyoming: Bear Lodge Mountains (Middle Fork Hay Creek), 2. 

GLAUCOMYS SABRINUS BANGSI (Rhoads). 

Bangs Flying Squirrel. 

[PI. II, fig. 3; PI. IV, fig. 3; PL VI, fig. 10.] 

^Sciuropterus alpinus bangsi Rhoads, Proc. Acad. Nat. Sci. Philadelphia, June, 
1897, p. 321. 

Type locality. — Idaho County, Idaho. 

Distribution. — Mountains of central Idaho, eastern Oregon, south- 
western Montana, and western Wyoming, north to vicinity of Flat- 
head Lake, Montana ; southern limits unknown. 

Characters. — Similar in size and color to sdbrinus, but upperparts 
averaging more drab (less vinaceous or ochraceous) and underparts 
more clouded with pinkish cinnamon (never yellowish white, as in 

1 Two from type locality ; one from Portland, N. Dak. 

2 Collection Field Mus. Nat. Hist. 



1918.] GLAUCOMYS SABRINUS GROUP. 39 

sabrinus) ; feet grayer (less brownish). Compared with alpinus: 
Upperparts decidedly more vinaceous, tail paler and much less 
clouded with fuscous. 

Color. — Winter pelage: Upperparts pale wood brown or avella- 
neous, shading in some specimens to vinaceous-cinnamon; feet pale 
hair-brown, shading to drab-gray, the toes often grayish white ; tail 
above, wood brown, tinged with cinnamon and more or less shaded 
with hair-brown or fuscous; beneath, light pinkish cinnamon, 
shaded with dark hair-brown; underparts whitish, strongly washed 
with pinkish cinnamon or avellaneous. Variation (May specimen 
from Lake Como, Mont.) : Upperparts pale orange-cinnamon; feet 
pale fuscous. 

Skull. — Very similar to that of sabrinus; slightly smaller than that 
of alpinus, with smaller bulla?. 

Measurements. — Average of 6 (adult and subadult) from Idaho 
and Montana : x Total length, 315 (304-327) ; tail vertebrae, 142 (137- 
148) ; hind foot, 39.5 (37-41). Skull: Average of 7 (adult and sub- 
adult) from same localities: Greatest length, 39.9 (38.2-41.2); 
zygomatic breadth, 24.3 (23.3-25) ; mastoidal breadth, 18.7 (18.3- 
19.1) ; least interorbital breadth, 7.5 (6.9-8) ; least postorbital 
breadth, 9 (8.4-9.7) ; length of nasals, 12.2 (11.1-12.5) ; maxillary 
toothrow, 8.1 (7.6-8.7). 

Type specimen. — No. 6959, Mus. Comp. Zool. (formerly same num- 
ber, collection E. A. and O. Bangs) ; $ subad., skin and skull ; 
collected March 8, 1897, by Harbison and Bargamin, Raymond, 
Idaho. 

Remarks. — This subspecies is the smallest of the Rocky Mountain 
forms of sabrinus. It agrees with the typical race in skull characters, 
but shows some approach in color to alpinus, differing from the latter, 
however, in color of tail and skull characters. Intergradation with 
sabrinus is indicated by a specimen from Paola, Mont. With lalipes, 
whose range meets (and possibly overlaps) that of bangsi, there 
seems to be no intergradation. The specimen from Paola shows no 
approach to latipes, while one from Nyack, a few miles farther 
north, is clearly referable to the latter form. Specimens from west- 
ern Wyoming (Pahaska, Kendall, Pacific Creek) are slightly larger 
than typical specimens, but do not differ appreciably in color. An 
August specimen from Anthony, Oreg., provisionally referred to this 
race, has the upperparts rich orange-cinnamon, much darker than any 
of the Idaho series, and quite different from another August specimen 
from Bourne, Oreg., which is in the normal wood-brown phase. 

Specimens examined. — Total number, 31, as follows: 
Idaho: Idaho County, 2; 2 Ketchum, 2; 2 Sawtooth Lake, 2. 

1 Idaho County (type and topotype), Ketchum, and Sawtooth Lake, Idaho; Florence, 
Mont. 

s Collection Mus. Comp. Zool. 



40 NORTH AMERICAN FAUNA. [No. 44. 

Montana : Florence, 6 ; Paola, 1 ; Rock Creek, near Lake Como, 1. 
Oregon: Anthony, 9; 1 Bourne, 1. 

Wyoming: Kendall (12 miles north, at 7,700 feet altitude), 2; Pacific 
Creek, 1 ; Pahaska, 2 ; Wind River Mountains, near Dubois, 2. 

GLAUCOMYS SABRINUS ALPINUS (Richardson). 

Richardson Flying Squirrel. 

[PI. II, fig. 4 ; PI. IV, fig. 4 ; PI. VI, fig. 12.] 

Pteromys alpinus Richardson, Zool. Journal, III, p. 519, 1828; Audubon and 
Bachman, Quad. North Amer., Ill, p. 206, 1854 ; Baird, Mamm. North Amer. : 
Rept. Expl. and Surv., R. R. Pac,, VIII, p. 289, 1S57. 

Pteromys sabriiius var. /3 alpinus Richardson, Fauna Boreali-Amer., I, p. 195, 
1829. 

[Sciuropterus volucella] alpinus Trouessart, Bull. Soc. Angers, X, p. 67, 1880. 

Sciuropterus alpinus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, p. 319, 1897. 

Type locality. — Jasper House, Alberta. 2 

Distribution. — Rocky Mountain region of Alberta and British 
Columbia, from vicinity of Henry House north at least to Peace 
River and Babine Lake, British Columbia (limits of range unknown) . 

Characters. — Similar to sabrinus, but upperparts much grayer (less 
vinaceous), tail darker, and skull larger, with broader braincase. 

Colo / r. — Winter pelage : Upperparts light drab ; sides of face pale 
smoke gray ; eye-ring fuscous ; ears edged with blackish brown ; upper 
surface of flying membrane blackish brown; hind feet hair-brown; 
fore feet similar but slightly paler; soles and palms soiled whitish; 
tail wood brown, much mixed, both above and below, with fuscous 
or clove brown, strongest on sides and tip ; underparts soiled whitish, 
sometimes irregularly marked (chiefly along median line) with light 
pinkish cinnamon. 

Skull. — Similar to that of sabrinus, but larger, with broader brain- 
case ; postorbital constriction relatively narrow. 

Measurements. — Two adults from Henry House, Alberta: Total 
length, 292, 343; tail vertebra?, 123, 155; hind foot, 41.5, 13; average 
of 3 adults from Stuart Lake, B. C. : 322 ; 149 ; 41.7. Skull: Average 
of 3 adults from Henry House and Jasper House, Alberta : Greatest 
length, 41.5 (41.2^2) ; zygomatic breadth, 25.4 (25.1-25.6) ; mas- 
toidal breadth, 19.4; least interorbital breadth, 8.3 (8.2-8.8); least 
postorbital breadth, 9.2 (9.1-9.6) ; length of nasals, 12.5 (12.4-12.6) ; 
maxillary toothrow, 7.8 (7.5-8.1). 

1 Seven in collection Amer. Mus. Nat. Hist. ; two in Mus. Vert. Zool., Univ. California. 

2 The habitat is given by Richardson in the original description as " Vallies in the 
Rocky Mountains." Later (Fauna Boreali-Amer., I, p. 195, 1829) he speaks of specimens 
from the head of the Elk [=Athabaska] River, and from the souih branch of the Mac- 
kenzie [=Liard, specimens probably from Fort Nelson]. As Richardson speaks of Drum- 
mond as the discoverer of the species, the vicinity of Jasper House, on the headwaters of 
the Athabaska, near which place Drummond made extensive collections, may be considered 
the type locality. 



1918.] GLAUCOMYS SABRINUS GROUP. 41 

Type specimen. — None known to exist. 

Remarks. — This handsome race, recognized as distinct by Richard- 
son nearly a century ago, has until recently been imperfectly known. 
Baird, in 1857, admitted the species but was in doubt as to its real 
characters. Allen, in his Monograph of the Sciuridss (1877) placed 
it in synonymy. Rhoads, in 1897, 1 restored it to specific rank, evi- 
dently using New England specimens of macrotis in comparison and 
considering them typical of sabrinus. Additional material now 
available, both of typical alpinus and of sabrinus from various parts 
of its range, shows conclusively that the two forms are subspecincally 
related. As already shown under sabrinus (p. 33), specimens from 
western Mackenzie and northern Alberta are intermediate in color 
between alpinus and sabrinus and those from southern Alberta have 
skulls nearly typical of alpinus but agree with sabrinus in color. Of 
two specimens from Peace River Canyon, B. C, one is typical of 
alpinus, while the other closely matches sabrinm in color. The pres- 
ent form intergrades, also, with zaphceus and columbiensis in British 
Columbia, and possibly with bangsi in southern Alberta. Several 
specimens from Stuart Lake, B. C, are intermediate in color between 
alpinus and columbiensis, while one specimen is so dark as to suggest 
intergradation with zaphams. 

Specimens examined. — Total number, 23, as follows: 

Alberta: Henry House, 3; Jasper House, 2; no specific locality, I. 2 
British Columbia: Babine Lake, 8: Cariboo, 1; 2 Peace River Canyon (near 
Hudsons Hope), 2; Stuart Lake, 6. s 

GLAUCOMYS SABRINUS YUKONENSIS (Osgood). 
Yukon Flying Squirrel. 
[PI. II, fig. 8; Pi. IV, fig. 8.] 

Sciuropterus yukonensis Osgood, North Amer. Fauna No. 19, p. 25, Oct. 6, 1900. 

Type locality. — Camp Davidson, Yukon River, Yukon (near 
Alaska-Canada boundary). 

Distribution. — Yukon River region, from vicinity of Mayo Lake, 
Yukon (head of Stewart River), to Tanana, Alaska; exact limits 
unknown. 

Characters. — Similar to sabrinus in color, but larger, with longer 
tail, broader hind foot, and larger skull. Compared with alpinus: 
Upperparts more ochraceous (less drab) ; tail longer and more 
vinaceous. 

Color. — Upperparts pinkish cinnamon to vinaceous-cinnamon ; 
sides of face pale smoke gray, sometimes clouded with light pinkish 

1 Proc. Acad. Nat. Sci. Philadelphia.. 1897, p. 319. 

2 Collection Field Mus. Nat. Hist. 

•Two in collection Provincial Mus., Victoria, B. C. ; one in Acad. Nat. Sci. Philadelphia. 



42 NORTH AMERICAN FAUNA. [No. 44. 

cinnamon; eye-ring clove brown or fuscous-black; upper surface of 
flying membrane clove brown or fuscous ; legs similar, partially over- 
laid with the color of the back; feet wood brown, hair-brown, or 
mouse gray ; soles and palms cinnamon-buff or whitish, clouded with 
mouse gray ; tail above, sayal brown, shaded with fuscous, becoming 
pure fuscous at the tip ; beneath, vinaceous-cinnamon or light pinkish 
cinnamon, edged with fuscous or hair-brown; underparts soiled 
whitish, moderately washed with light pinkish cinnamon or avella- 
neous. 

Skull.— Closely similar to /that of alpinus; braincase slightly 
larger; nasals slightly shorter. 

Measurements. — Type : Total length, 365 ; tail vertebrae, 180 ; * hind 
foot (dry), 41; ? adult from Tanana, Alaska: 324; 150; 42. Skull: 
Average of 3 adults from Alaska: 2 Greatest length, 41.3 (41-41.5) ; 
zygomatic breadth, 25.2 (25.1-25.3) ; mastoidal breadth, 19.6 (19.5- 
19.7) ; least interorbital breadth, 8.1 (7.8-8.3) ; least postorbital 
breadth, 9.5 (9.3-10.3) ; length of nasals, 11.9 (11.6-12.1) ; maxillary 
toothrow, 8.3 (7.9-8.7). 

Type specimen. — No.$£$$# , U. S. Nat. Mus. ; 9 ad., skin and skull ; 
collected December 8, 1890, by R. E. Carson. 

Remarks. — This is one of the larger members of the group, about 
equaling latipes and bullatus in external measurements. The 
hind feet, although about the same length as those of alpinus, 
are considerably broader and heavier, and the tail averages longer. 
Although the present form resembles sdbrinus in color, its skull 
is practically identical with that of alpinus. No specimens are 
available from the large area between the known ranges of alpinus 
and yukonensis, but in view of the close relationship of these two 
forms, as shown by the skull characters, it seems probable that they 
will be found to intergrade, and for that reason yukonensis is here 
treated as a subspecies of sdbrinus. Osgood states 3 that flying squir- 
rels have been taken in the Knik district, Cook Inlet, but until speci- 
mens can be secured from that region their subspecific identity must 
remain in doubt. 

Specimens examined. — Total number, 10, as follows: 

Alaska: Big Chena River (65 miles from mouth), 1;* Chicken, 1; Tanana, 

3; Toklat River (head), 1. 
Yukon: Camp Davidson (near Alaska-Canada boundary), 2; Fortymile, 

1 ; 5 Mayo Lake, 1.° 

1 In a topotype, the tail (measured from dry skin) is approximately 160. 

2 Camp Davidson, Tanana, and head of Toklat River. 

3 Osgood, W. H., North Amer. Fauna No. 21, p. 63, 1901. 
* Skull only. 

8 Collection Mus. Vert. Zool., Univ. California. 
Collection Victoria Mem. Mus. 



1918.] GLAUCOMYS SABRINUS GROUP. 43 

GLAUCOMYS SABRINUS ZAPH^EUS (Osgood). 

Axaska Coast Flying Squirrel. 

[PI. Ill, fig. 6 ; PI. V, fig. 6 ; PI. VI, fig. 9.] 

Sciuropterus alpinus zaphcetis Osgood, Proc. Biol. Soc. Washington, XVIII, 
p. 133, Apr. 18, 1905. 

Type locality. — Helm Bay, Cleveland Peninsula, Alaska. 

Distribution. — Coast region of southeastern Alaska and northern 
British Columbia ; limits of range unknown. 

Characters. — Similar to alpinus, but upperparts browner (less 
grayish) and underparts darker. Compared with oregonensis: 
Upperparts paler; underparts grayer (without cinnamon or buff). 

Color. — Winter pelage: Upperparts sayal brown; sides of face 
pale smoke gray, often clouded with fuscous-black, the blackish eye- 
ring often pronounced; upper surface of flying membrane fuscous; 
feet dark hair-brown or fuscous, the soles whitish tinged with hair- 
brown on outer margin ; tail above, fuscous, tinged with sayal brown ; 
beneath, dull vinaceous-cinnamon, much clouded with fuscous on sides 
and tip ; underparts soiled whitish, strongly washed with avellaneous 
or wood brown, strongest along sides. Summer pelage: Upperparts 
yellowish wood brown ; underparts washed with pinkish cinnamon. 

Skull. — Similar to that of alpinus, but braincase narrower, bullae 
slightly smaller, and upper toothrow slightly longer. 

Measurements. — Average of 6 topotypes (adult) : Total length, 
30T (292-311) ; tail vertebras, 144 (133-152) ; hind foot (dry skin), 
40.7 (39-42) ; average of 3 topotypes (adult) i 1 306; 145; 40. Skull: 
Average of 6 topotypes (adult and subadult) : Greatest length, 40.1 
(39.3-11.5) ; zygomatic breadth, 24.7 (23.7-26.2) ; mastoidal breadth, 
18.7 (18.2-19.1) ; least interorbital breadth, 7.8 (7.4-8.3) ; least post- 
orbital breadth, 9 (8.7-9.8) ; length of nasals, 12.5 (12.2-12.9) ; maxil- 
lary toothrow, 7.8 (8-8.6). 

Type specimen. — No. 136137, U. S. Nat. Mus. (Biological Survey 
collection); 5 ad., skin and skull; collected January 21, 1905, by 
Cyrus Catt. 

Remarks. — This subspecies is intermediate in color between alpinus 
and oregonensis, being nearer the latter in the color of the upper- 
parts, but lacking the deep cinnamon-buff on the underparts. In 
skull characters it is close to alpinus and intergrades with it in the 
interior of British Columbia. The northern and southern limits of 
its range are unknown, but very probably it intergrades with ore- 
gonensis in southwestern British Columbia and possibly with 
yukonensis in Yukon. 

1 Measured in flesh by Frank Stephens. 



44 NORTH AMERICAN FAUNA. [No. 44. 

Specimens examined.— Total number, 14, as follows: 

Alaska: Bradflelcl Canal, l; 1 Etolin Island, l; 1 Helm Bay, Cleveland 

Peninsula, 10 ; 2 Tongass, 1. 
British Columbia: Nass River, 1. 

GLAUCOMYS SABRINUS GREGONENSIS (Bachman). 

Bachman Flying Squirrel. 

[PI. Ill, fig. 11 ; PI. V, fig. 11 ; PI. VI, fig. 8.] 

Pteromys oregonensis Bachman, Journ. Acad. Nat. Sci. Philadelphia, VIII, 
p. 101, 1839 ; Audubon and Bachman, Quad. North Araer., I, p. 132, 1846. 

[Sciuropterus volucella] oregonentis [sic] Trouessart, Bull. Soc. Angers, X, 
p. G7, 1S80. 

Sciuropterus alpinus oregonensis Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 
1S97, p. 324. 

Type locality. — " Pine woods of the Columbia near the sea " — 
probably near the present site of St. Helens, Oregon. 

Distribution. — Coast region of Oregon, Washington, and southern 
British Columbia ; northern and southern limits unknown. 

Characters. — Similar to zapluvus, but colors more rufescent, both 
above and below (upperparts rich reddish brown, underparts cinna- 
mon or buff) ; sides of face more buffy (less grayish) ; skull slightly 
smaller. 

Color. — Winter pelage: Upperparts mikado brown or pecan brown ; 
sides of face smoke gray or grayish white, often much mixed with 
buff or cinnamon ; upper surface of flying membrane clove brown or 
fuscous; hind feet snuff brown or pale fuscous, the soles pinkish 
buff shaded with drab; fore feet hair-brown or drab; tail above, 
snuff brown, shaded on sides and tip with fuscous or clove brown; 
beneath, cinnamon-buff; underparts heavity washed with pinkish 
cinnamon or cinnamon-buff. 

Skull. — Similar to that of zapka : us, but averaging smaller, with the 
zygomata less widely expanded. Closely similar to that of fuligi- 
nosus, but averaging slightly smaller. 

Measurements. — Average of 4 adults from Oregon : 3 Total length, 
300 (294-310) ; tail vertebrae, 127 (122-138) ; hind foot, 38.6 (38-40). 
Skull: Average of 8 adults from Oregon : 4 Greatest length, 39.9 (38.5- 
40.7) ; zygomatic breadth, 23.7 (23-24.9) ; mastoidal breadth, 18.3 
(17.6-19) ; least interorbital breadth, 7.9 (7.3-8.8) ; least postorbital 
breadth, 8.6 (8.1-9) ; length of nasals, 12.2 (10.9-12.9) ; maxillary 
toothrow, 8.2 (7.9-8.7). 

1 Collection Mus. Vert. Zool., Univ. California. 

2 Four in collection Mus. Vert. Zool., Univ. California. 

3 Portland, Yaquina Bay, and Gold Beach. 

4 Portland (3), Latourelle Falls, Yaquina Bay, Gold-Beach, Salem, and lower Columbia 
River (type). 



1918.] GLAUCOMYS SABRINUS GROUP. 45 

Type specimen. — No. 235, Acad. Nat. Sci. Philadelphia; adult, 
skin and skull; skin (dismounted and skull removed) in fair con- 
dition, possibly somewhat faded ; skull nearly perfect, but portion of 
occiput cut away ; collected in 1839 by J. K. Townsend. 

Remarks. — The Bachman flying squirrel is easily recognized by 
its rich coloration, rufescent above and buffy below. It occupies the 
humid northwest coast belt, but does not range high in the moun- 
tains. Intergradation with fuliginosus is shown by a specimen from 
Marmot, Oreg. (west slope of Mount Hood), and with columbiensis 
by specimen's from Sumas, Chilliwack, and Stave Lake, B. C. These 
latter specimens agree with oregonensis in the color of the upper- 
parts, but are much paler beneath. A specimen from Sulphur 
Springs, Oreg., is considerably paler above than typical specimens. 
Specimens from Yaquina Bay and Gold Beach, Oreg., have smaller 
skulls than the type and other specimens from the Columbia River. 

Specimens examined. — Total number, 41, as follows: 

British Columbia: Agassiz, l; 1 Chilliwack, 1; Harrison, l; 2 Mission, 1 ; 3 
Stave Lake, 1 ; 4 Sumas, 3. 4 

Oregon: Brownsville, 1 ; B Clackamas River, 1; "Columbia River," 3; 8 Elk 
Head, 1; Gold Beach, 1; Latourelle Falls (18 miles east of Portland), 
1 ;' Marmot, 1 ; Portland, 4 ; Port Orford, 4 ; Salem, 1 ; T Sulphur Springs, 
Benton County (near Albany), 1; Yaquina Bay, 1; no specific local- 
ity, 2. 

Washington : Camp Skagit, 1 ; Fort Bellingham, 1 ; " Lewis River," 1 ; 4 
Olympic Mountains (north Fork Skokomish River), 1; Puget Sound, 6; 
no specific locality, 1. 

GLAUCOMYS SABRINUS COLUMBIENSIS Howell. 

Okanagan Flying Squikeel. 

[PI. Ill, fig. 7 ; PI. V, fig. 7 ; PI. VI, fig. 7.] 

Glaucomys sabrinus columbiensis Howell, Proc. Biol. Soc. Washington, XXVIII, 
p. Ill, May 27, 1915. 

Type locality. — Okanogan, British Columbia. 

Distribution. — Interior valleys and foothills of southern British 
Columbia and northern Washington, from Shuswap Lake and Cran- 
brook, British Columbia, south to Lake Chelan, Washington. 

Characters. — Decidedly paler than oregonensis, both above and 
below; similar to Mamathensis, but upperparts more vinaceous and 
tail much darker; very similar to sabrlnus, but soles of hind feet 
often yellow (as in klamathensis) , and skull larger. Compared with 

1 Collection W. E. Saunders, London, Ont. 

2 Collection Provincial Mus., Victoria, B. C. 

3 Collection Victoria Mem. Mus. 

4 Collection Mus. Comp. Zool. 

6 Collection Leland Stanford, Jr., Univ. 
6 Collection Acad. Nat. Sci. Philadelphia. 
1 Collection Game Dept. Oregon. 



46 NORTH AMERICAN FAUNA. [No. 44. 

fuliginosus : Upperparts more vinaceous (less brownish) ; underparts 
paler; tail more brownish. 

Color. — Winter pelage: Upperparts between vinaceous-cinnamon 
and vinaceous-f awn ; sides of face smoke gray, shaded with fuscous; 
front of face sometimes marked with grayish white; upper surface 
of flying membrane clove brown ; feet hair-brown or pale fuscous, the 
soles deep colonial buff, or sometimes grayish white; tail above, 
snuff brown, shading to fuscous on sides and tip; beneath, pinkish 
cinnamon, more or less clouded (often strongly) with fuscous; under- 
parts white, tinged with straw yellow or light pinkish cinnamon. 

Skull. — Closely similar to that of alpinus, but averaging slightly 
narrower, with flatter, shallower braincase; similar to that of ore- 
gonensis, but slightly larger, with shorter toothrow ; very similar to 
that of fuliginosus, but averaging relatively broader. 

Measurements. — Two topotypes (subadult) : Total length, 313; tail 
vertebrae, 143; hind foot, 42. Skull: Average of 7 (adult and sub- 
adult) from southern British Columbia. 1 Greatest length, 41.1 
(40.3-41.6) ; zygomatic breadth, 24.5 (23.6-25.2) ; mastoidal breadth, 
19.1 (18.8-19.5) ; least interorbital breadth, 7.8 (7.3-8.6) ; least post- 
orbital breadth, 9.1 (8.4-9.7) ; length of nasals, 12.6 (12-13.3) ; maxil- 
lary toothrow, 7.8 (7.6-8.2). 

Type specimen. — No. 94310, U. S. Nat. Mus. (Biological Survey 
collection) ; $ subad., skin and skull ; collected May 9, 1898, by Allan 
Brooks. 

Remarks. — This subspecies, like most of the races inhabiting the 
dry interior valleys of southern British Columbia, is considerably 
paler than the coast form (oregonensis) . Intergradation with the 
latter is shown by a series of specimens from Sumas and Chilliwack, 
B. C.j and with fuliginosus by a series from mouth of Salmon River, 
B. C. Two specimens in the latter series are referable to the pres- 
ent form, although having somewhat darker underparts. Other 
specimens, labeled as from the same locality but possibly taken at a 
higher altitude, agree essentially with fuliginosus, to which race they 
are referred. Intergradation with aljnnus is shown by specimens 
from Stuart Lake, B. C. (referred to alpinus). One specimen from 
Cranbrook, B. C, is considerably grayer above than the typical form, 
thus suggesting intergradation with latipes, with which it agrees in 
all other respects. Another from the same place is typical. 

Specimens examined. — Total number, 11, as follows: 

British Columbia: Cranbrook, 2; Grand Prairie (22 miles northwest of 
Okanogan), 1 ; 2 Okanogan, 2; (Okanogan Falls (Shuttleworth Creek), 
l; 2 Penticton, l; 3 Salmon River (mouth), 2; 3 Vernon, 1. 

Washington: Chelan Mountains, 1.* 

1 Okanogan, Vernon, Penticton, Cranbrook, and mouth of Salmon River. 
1 Collection Provincial Mus., Victoria, B. C. 
s Collection Victoria Mem. Mus. 
♦Collection Amer. Mus. Nat. Hist. 



1918.] GLAUCOMYS SABEINUS GROUP. 47 

GLAUCOMYS SABRINUS FULIGINOSUS (Rhoads). 

Cascade Flying Squibrel. 

[PL III, fig. 4 ; PI. V, fig. 4 ; PL VII, fig. 7.] 

Sciuropterus alpinus fuliginosus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 
June, 1897, p. 321. 

Type locality. — Cascade Mountains, near Martin Station, Kittitas 
County, Washington (altitude about 8,000 feet). 

Distribution. — Cascade Range, from southern British Columbia 
south through Washington and Oregon to the Siskiyou Mountains, 
California. 

Characters. — Similar to columbiensis, but upperparts more brown- 
ish (less vinaceous) ; underparts darker, and tail paler (less brown- 
ish). Compared with oregonensis: Colors much less rufescent, espe- 
cially on upperparts and underside of tail. 

Color. — Winter pelage: Upperparts pale sayal brown (between 
sayal brown and wood brown), the sides sometimes faintly washed 
with pinkish cinnamon; upper surface of flying membrane clove 
brown; sides of face smoke gray or pale smoke gray, sometimes 
washed with light buff or pale fuscous ; eye-ring fuscous ; ears hair- 
brown; hind feet dark hair-brown (approaching fuscous), the toes 
marked with buffy white, the soles buffy white or yellowish white, 
edged with hair-brown ; tail above, dark hair-brown, shading to fus- 
cous, sometimes sparingly mixed with dull cinnamon or pinkish cin- 
namon ; beneath, pale ochraceous-buff or light ochraceous-buff , edged 
on sides with fuscous, the bases of hairs pale smoke gray ; underparts 
pinkish cinnamon or light pinkish cinnamon, the throat often creamy 
white. 

Skull. — Closely similar to that of columbiensis, but averaging rela- 
tively narrower ; similar to that of oregonensis, but slightly larger. 

Measurements. — Average of 3 topotypes (adult) : Total length, 
308 (304-317) ; tail vertebrae, 144 (140-153) ; hind foot, 40.7 (40-42). 
Skull: Average of 10 (adult and subadult) from Easton and Keeche- 
lus Lake, Wash.: Greatest length, 41.4 (40.2-42.5); zygomatic 
breadth, 24.1 (23.5-24.9) ; mastoidal breadth, 18.7 (17.9-19.3) ; least 
interorbital breadth, 8.3 (7.7-9) ; least postorbital breadth, 9.1 (8.4- 
9.6) ; length of nasals, 12.9 (12.3-13.3) ; maxillary toothrow, 8.4 
(8.2-8.4) . 

Type specimen. — No. 8058, Acad. Nat. Sci. Philadelphia (formerly 
No. 1058, collection S. N. Rhoads) ; $ subad., skin and skull ; col- 
lected in March, 1893, by Allan Rupert. 

Remarks. — This subspecies, occupying the Cascade Range in Wash- 
ington and Oregon, is markedly paler than the coast form (oregon- 
ensis). It intergrades with the latter on the western slopes of the 
range, with klamathensis on the eastern slopes in Oregon, with colum- 



48 NORTH AMERICAN FAUNA. [No. 44. 

biensis in southern British Columbia, and with flaviventris in north- 
western California. 

Specimens from Vida and Belknap Springs, Oreg., have browner 
backs and paler bellies than the typical series, thus showing approach 
to oregonensis. A specimen from Crater Lake, Oreg., approaches 
klamathensis in having paler and more yellowish underparts. Speci- 
mens from the Siskiyou Mountains, Cal., are intermediate between 
fuliginosus and flaviventris, agreeing with the former in the color 
of the upperparts and feet and in skull characters, but having the 
underparts paler and more yellowish than in typical specimens. 
Specimens examined. — TotaFnumber, 68, as follows : 
British Columbia: Hope, 7 ;*■ 2 Salmon River (mouth), 6. 3 
California: Preston Peak, 1; Siskiyou Mountains, 5. 

Oregon: Belknap Springs, 2 ; Crater Lake, 1 ; McKenzie Bridge, 2 ; Vida, 3. 4 
Washington: Bumping Lake, Yakima County, 1; Cowlitz River, 1; Easton, 
18 ; Keechelus Lake, 14 ; Martin Station, 3 ; Potato Hill, 15 miles north 
of Goldendale, 1 ; Signal Peak, Yakima Indian Reservation, 3. 

GLAUCOMYS SABRINUS LATIPES Howell. 

Broad-Footed Flying Squirrel. 

[PL III, fig. 8 ; PL V, fig. 8 ; PL VII, fig. 8.] 

Olaucomys sabrinus latipes Howell, Proc. Biol. Soc. Washington, XXVIII, p.112, 
May 27, 1915. 

Type locality. — Glacier, British Columbia. 

Distribution. — Selkirk Range, and other ranges in southeastern 
British Columbia, higher mountains of northern Idaho and north- 
western Montana ; south to Mullan and Orofino, Idaho. 

Characters. — Similar to fuliginosus but larger; upperparts aver- 
aging darker and grayer; feet larger and darker. Compared with 
alpinus and bang si: Size larger; upperparts more brownish (less 
drab) ; underparts darker. 

Color. — Winter pelage: Upperparts drab, more or less mixed with 
sayal brown ; sides of face smoke gray ; ears fuscous ; upper surface of 
flying membrane dark clove brown ; hind feet fuscous ; fore feet dark 
hair-brown, the toes shaded with buffy white; tail above, fuscous 
shaded with sayal brown; beneath, pinkish buff or light ochraceous- 
buff, more or less mixed with fuscous; underparts light ochraceous- 
buff or cinnamon-buff. Summer pelage: Upperparts chiefly sayal 
brown with a drab tinge, otherwise as in winter. 

Skull. — Similar in general to that of fuliginosus, but larger (about 
equaling that of bullatus) ; longer and relatively narrower than that 
of alpinus. 

1 Collection Mus. Coinp. Zool. 

2 Collection E. R. Warren, Colorado Springs, Colo. 

3 Collection Victoria Mem. Mus. 

1 Two in collection Game Dept. Oregon. 



1918.] GLAUCOMYS SABKINUS GROUP. 49 

Measurements. — Average of 10 adults: 1 Total length, 342 (315- 
368) ; tail vertebra?, 153 (145-161) ; hind foot, 41.4 (38-44). Skull: 
Average of 7 adults: 1 Greatest length, 42.9 (41.6-44.2) ; zygomatic 
breadth, 24.5 (24-25.4) ; mastoidal breadth, 19.5 (19-20) ; least in- 
terorbital breadth, 8.3 (8-8.8) ; least postorbital breadth, 8.8 (8-9.1) ; 
length of nasals, 13.3 (12.5-14) ; maxillary toothrow, 8.3 (7.9-9). 

Type specimen. — No. 68753, U. S. Nat. Mus. (Biological Survey 
collection), 5 ad., skin and skull; collected August 13, 1894, by J. 
Alden Loring. 

Remarks. — This subspecies is one of the largest of the American 
flying squirrels, nearly equaling bullatus in external measurements 
and size of skull. Although evidently closely related to fuliginosus 
of the Cascades, intermediate specimens are unknown, but are likely 
to be found when further collecting is done in British Columbia. 
The present form apparently does not intergrade either with alpinus 
of the northern Eockies or with bangsi of the Bitterroot and Saw- 
tooth Ranges in Idaho and Montana. From the latter race, 
the range of which meets that of latipes in northern Montana, 
this form differs in much larger size and darker colors. The 
range of latipes is apparently confined to the mountains, since an- 
other form {columbiensis) occurs in the Okanogan Valley and at 
Cranbrook, B. C. One specimen from the latter locality nearly 
matches specimens of latipes in color of the upperparts, but in other 
characters agrees with columbiensis. 

Specimens examined. — Total number, 23, as follows : 

British Columbia: Glacier, 1; Sehoonover Mountain, near Okanogan (4,000 

feet altitude), 2; 2 Shuswap, 1. 
Idaho : Coolin, 4 ; Mullan, 7 ; Oroflno, 1 ; Priest Lake, 1. 
Montana : Nyack 1 ; Stanton Lake, 5. 

GLAUCOMYS SABRINUS OLYMPICUS (Elliot). 

Olympic Flying Squirrel. 

[PI. Ill, flg. 10 ; PI. V, fig. 10 ; PI. VII, fig. 6.] 

Sciuropterus alpinus olympicus Elliot, Field Columb. Mus., Publ. 30, Zool. Ser., 
I, p. 225, Feb. 1, 1899. 

Type locality. — Happy Lake, Olympic Mountains, Washington. 

Distribution. — Olympic Peninsula, Washington, and south along 
the coast to southern Oregon; occurring in some localities with 
oregonensis. 

1 From Glacier, B. C. ; Coolin, Idaho ; and Stanton Lake and Nyack, Mont. 

2 Collection Provincial Mus., Victoria, B. C. 

14520°— 18— No. 44 4 



50 NORTH AMERICAN FAUNA. [No. 44. 

Characters. — Similar to fuliginosus, but upperparts, face, and feet 
much darker; colors much less rufescent than in oregonensis, both 
above and below ; skull similar to that of fuliginosus, slightly larger 
than that of oregonensis. 

Color. — Worn summer pelage: Upperparts wood brown; sides of 
face smoke gray shaded with cinnamon ; eye-ring and upper surface 
of flying membrane fuscous-black; ears pale fuscous; feet fuscous, 
the soles shaded with buffy white; tail above, fuscous; beneath, dull 
light pinkish cinnamon, edged with fuscous; underparts whitish, 
heavily washed with pinkish cinnamon. Winter pelage (specimens 
from Ocean View, Oreg.): 1 Upperparts snuff brown; tail above, 
snuff brown heavily shaded with fuscous or fuscous-black; feet deep 
mouse gray ; otherwise as in summer. 

Skull. — Essentially the same as in fuliginosus; very similar to 
that of oregonensis, but averaging slightly larger. 

Measurements. — Average of 4 topotypes (subadult) : Total length, 
335 (328-346) ; tail vertebra?, 162 (159-166) ; hind foot, 37.3 (35-39). 
Skull: Greatest length, 41 (40.3-41.6); zygomatic breadth, 23.9 
(23.4-24.7) ; mastoidal breadth, 18.6 (18.4-19) ; least interorbital 
breadth, 8.2 (7.6-8.5) ; least postorbital breadth, 9.1 (8.8-9.4) ; length 
of nasals, 12.4 (11.2-13.2) ; maxillary toothrow, 8.4 (8.3-8.5). 

Type specimen. — No. 5902, Field Mus. Nat. Hist. ; $ subad., skin 
and skull ; collected September 10, 1898, by D. G. Elliot. 

Remarks. — The Olympic flying squirrel is apparently the darkest 
of the American forms. It is most nearly related to oregonensis, 
from which it differs in much less rufescent coloration. The material 
at hand is insufficient to show clearly whether the two races inter- 
grade. The occurrence of typical specimens of both forms in the 
Olympic Mountains and at Gold Beach, Oreg., argues for their spe- 
cific distinctness, but, on the other hand, a series from Ocean View, 
Oreg., strongly suggests intergradation, some of the specimens being 
clearly intermediate in color between olympicus and oregonensis. 
In view of the close cranial relationship which olympicus bears to 
both oregonensis and fuliginosus it seems best to unite it with them 
as a subspecies of sahrinus. To clear up the problem much additional 
material is needed — particularly a good series of winter specimens 
from the type region. At present the only available specimens in 
winter pelage are those from Ocean View, Oreg., which, as already 
explained, may be shading toward oregonensis. 

Specimens examined. — Total number, 15, as follows: 
Oregon : Gold Beach, 1 ; 2 Ocean View, 9. s 
Washington : Olympic Mountains, 4 ; 2 Seattle, 1." 

1 Possibly not typical — see Remarks. 

2 Collection Field Mus. Nat. Hist. 

3 Collection Game Dept. Oregon. 



1918.] 



GLAUCOMYS SABEINUS GROUP. 



51 



GLAUCOMYS SABRINUS BULLATUS Howell. 

Sawtooth Mountains Flying Sqtjierel. 

[PL III, fig. 12 ; PL V, fig. 12 ; PL VII, fig. 4.] 

Glaucomys bicllatus Howell, Proc. Biol. Soc. Washington, XXVIII, p. 113, 
May 27, 1915. 

Type locality. — Sawtooth [Alturas] Lake, Idaho. 

Distribution. — Sawtooth Mountains, Idaho, north to Cranbrook, 
British Columbia, and west to the Blue Mountains, Oregon (fig. 4). 

Characters. — Similar to klamathensis, but larger, with much larger 
bullae; upperparts more pinkish, underparts tinged with avel- 
laneous instead of yellow, tail darker above and more vinaceous be- 
low, and soles of hind feet smoke gray instead of olive-ochre. Similar 
in color to bangsi, but upper- 
parts decidedly more pinkish 
(less vinaceous or drab) ; gray on 
face purer and more extensive; 
size much larger. 

Color. — Winter pelage: Upper- 
parts between pinkish cinnamon 
and cinnamon-buff, shaded in 
some specimens with wood brown, 
becoming paler on the face ; sides 
of face and neck pale smoke gray, 
this color extending back to a 
point slightly behind the ears; 
ears dark hair-brown edged with 
fuscous and often partly clothed 
with smoke gray hairs ; upper sur- 
face of flying membrane between 
hair-brown and fuscous; feet be- 
tween mouse gray and hair-brown, 

the fore feet and toes of hind feet often marked with grayish white, 
the soles pale smoke gray ; tail above, pinkish cinnamon much mixed 
with fuscous, becoming dark mouse gray at tip ; beneath, dull light 
pinkish cinnamon; underparts creamy white, strongly shaded with 
avellaneous or light pinkish cinnamon, becoming pure avellaneous 
on sides. Summer pelage: Hind feet somewhat paler (about mouse 
gray) but otherwise as in winter. 

Skull. — 'Size large (averaging largest of the races of sab Anus) ; 
braincase relatively narrow and very deep, abruptly depressed pos- 
teriorly; interorbital region depressed and fronto-parietal region 
markedly elevated; interorbital region broad, the interorbital notch 
obsolete or much reduced; audital bullae very large; basioccipital 
relatively narrow ; molar teeth massive. 




Fig. 4. — Map showing the distribution 
of Glaucomys sabrinus bullatus, based 
on specimens examined. This form 
occurs in some localities with others 
of the same species (see fig. 3, 
P. 30). 



52 NORTH AMERICAN FAUNA. TNo. 44. 

Measurements. — Type: Total length, 340; tail vertebrae, 150; hind 
foot, 46. Average of 6 adults from Ketchum, Idaho: Total length, 
336 (327-354) ; tail vertebra?, 142 (135-150) ; hind foot, 42.5 (39-45). 
Skull: Average of type and of 5 adults from Ketchum, Idaho : Great- 
est length, 43.3 (42.4-44.2) ; zygomatic breadth, 25 (24.4-25.5) ; mas- 
toidal breadth, 19.3 (19-20) ; least interorbital breadth, 8.9 (8.1-9.6) ; 
least postorbital breadth, 9.4 (9-9.7) ; length of nasals, 13.6 (13.5- 
13.9) ; maxillary toothrow, 9.1 (8.8-9.3). 

Type specimen. — No. fffrii U. S. Nat. Mus. (Biological Survey 
collection) : 9 ad., skin and skull; collected September 28, 1890, by 
Vernon Bailey and B. H. Dutcner. 

Remarks. — This race is the largest and one of the handsomest of 
the American flying squirrels. It was originally believed to be a dis- 
tinct species, since it occurs at the same localities with bangsi in Idaho 
and with latipes in British Columbia, and does not intergrade with 
them, but recently acquired material from eastern Oregon shows in- 
tergradation with klamathensis. An adult from Beech Creek, Oreg., 
agrees well with typical bullatus in color, and its skull is slightly 
larger than the largest of the Idaho series; the braincase, however., 
is relatively broader and flatter. Another adult from Cornucopia, 
Oreg., approaches klamathensis in having the belly and the soles 
tinged with colonial buff. A specimen from Cranbrook, British 
Columbia, differs from the Idaho series in having the underparts a 
slightly darker shade of cinnamon and the tail more extensively 
shaded with fuscous. 

Specimens examined. — Total number, 14, as follows : 

British. Columbia: Cranbrook, 1. 

Idaho : Ketchum, 6 ;* Sawtooth Lake, 2. 

Oregon : Anthony, 1 ; 2 Beech Creek, 1 ; Cornucopia, 3. 

GLAUCOMYS SABRINUS KLAMATHENSIS (Meebiam). 

Klamath Flying Squirrel. 

[PI. Ill, fig. 3 ; PI. V, fig. 3 ; PI. VII, fig. 3.] 

Sciuropterus alpinus klamathensis Merriam, Proc. Biol. Soc. Washington, XI. 
p. 225, July 15, 1897. 

Type locality. — Fort Klamath, Oregon. 

Distribution. — Central Oregon, chiefly east of the Cascades; north- 
ern and eastern limits unknown. 

Characters. — Similar to fuliginosus but upperparts averaging 
slightly paler ; gray on face purer and more extensive (less mixed with 
buff); underparts paler, usually washed with pale j'ellow; soles 
strongly tinted with yellow; audital bulla? larger. Similar to bullatus 

1 Collection Mus. Comp. Zool. 2 Collection Amer. Mus. Nat. Hist 



1918.] GLAUCOMYS SABRINUS GROUP. 53 

but bullae smaller, colors paler, and underparts and soles shaded 
with yellowish. 

Color. — Winter pelage: Upperparts cinnamon-drab or pale sayal 
brown shading in some specimens to light yellowish drab; sides of 
face smoke gray, often considerably washed with pinkish cinnamon ; 
upper surface of flying membrane fuscous; ears drab or pale hair- 
brown; hind feet hair-brown, the soles olive-ochre or deep colonial 
buff ; fore feet light brown mixed with buffy white ; tail above, sayal 
brown or dull cinnamon, much mixed with hair-brown, the tip nearly 
pure hair-brown; beneath, pinkish buff, light pinkish cinnamon, or 
colonial buff, moderately shaded with hair-brown, chiefly on sides; 
underparts whitish, rather heavily washed with colonial buff or 
cream-buff, shading to light pinkish cinnamon along sides. 

Skull. — Similar to that of fuliginosus but with larger bullae ; simi- 
lar to that of hullatus but smaller, with smaller bullae and flatter 
braincase. 

Measurements. — Average of 9 topotypes (adult and subadult) : 
Total length, 319 (300-336) ; tail vertebrae, 144 (135-154) ; hind foot, 
40.4 (39.5-42). Skull: Average of 6 topotypes (adult and subadult) : 
Greatest length, 40.9 (40.2-42.3) ; zygomatic breadth, 23.6 (23.2-25) ; 
mastoidal breadth, 18.9 (18.6-19.6) ; least interorbital breadth, 7.8 
(7-9) ; least postorbital breadth, 9.2 (8.6-10) ; length of nasals, 12.7 
(11.8-13.9) ; maxillary toothrow, 8.6 (8.1-9). 

Type specimen. — No. 87310, U. S. Nat. Mus. (Biological Survey col- 
lection) ; 9 ad., skin and skull; collected January 11, 1897, by B. L. 
Cunningham. 

Remarks. — This race occupies the interior of Oregon, chiefly east 
of the Cascades, but extending into the eastern foothills of that range 
and occasionally as high as Crater Mountain (4 miles south of Crater 
Lake) . It intergrades with fuliginosus in the Cascades and with flavi- 
ventris in northeastern California. A series of specimens from Davis 
Mountain, Crook County, Oreg., exhibit the characters of the sub- 
species in a more pronounced way than the series from the type 
locality. In color they are more drab above and have the underparts 
and feet more intensely yellowish; the skulls average somewhat 
larger. A specimen from Paulina Lake, Oreg., differs from typical 
specimens in having gray instead of yellow feet, and approaches 
fuliginosus also in the color of the back and underparts. 

Specimens examined. — Total number, 27, as follows : 

Oregon: Crater Mountain (4 miles south of Crater Lake), 1; Davis Moun- 
tain, Crook County, 7 ;* Foi*t Klamath, 17 ; Paulina Lake, 1 ; " Upper 
Klamath" [Lake], 1." 

'Collection Game Dept. Oregon. 2 Collection Field Mus. Nat. Hist. 



54 NORTH AMERICAN FAUNA. [No. 44. 

GLAUCOMYS SABRINUS FLAVIVENTRIS Howell. 

Yellow-Bellied Flying Squirrel. 

[PI. Ill, fig. 2 ; PI. V, fig. 2 ; PL VII, fig. 2.] 

Glaucomys sabrinus flaviventris Howell, Proc. Biol. Soc. Wash., XXVIII, p. 112, 
May 27, 1915. 

Type locality. — Head of Bear Creek, Trinity County, California 
(altitude, 6,400 feet). 

Distribution. — Northern California, from the Trinity Mountains in 
Siskiyou and Trinity Counties .east to the Warner Mountains, Modoc 
County. 

Characters. — Similar to klamathensis but smaller, with much 
smaller audital bullae; underparts and feet more strongly suffused 
with yellow or buff ; tail darker beneath. 

Color. — Summer pelage : Upperparts pale sayal brown varying in 
some specimens to drab ; sides of head smoke gray, often tinged with 
buff; ears hair-brown; upper surface of ftying membrane fuscous; 
feet hair-brown, often tinged with pale greenish yellow, the soles oil 
yellow ; tail above, dark buffy brown, shading to pale fuscous at tip ; 
beneath, brownish cream-buff, bordered on sides with pale fuscous or 
hair-brown; underparts whitish, usually heavily washed with pale 
greenish yellow or Naples yellow, shading to wood brown along sides ; 
a small white patch on throat, the hairs white to the roots. 

Skull. — Similar to that of klamathensis, but decidely smaller, with 
much smaller bulla?; much smaller than that of fuliginosus; closely 
similar to that of lascivus. 

Measurements. — Average of 5 topotypes (adult) : Total length, 306 
(286-322); tail vertebra?, 133 (122-140); hind foot, 40.4 (40-42). 
Skull: Greatest length, 39.5 (38.7-40.4) ; zygomatic breadth, 23.5 
(23.2-24.2) ; mastoidal breadth, 17.9 (17.6-18.7) ; least interorbital 
breadth, 7.2 (6.3-8.3) ; least postorbital breadth, 8.9 (8.2-9.8) ; length 
of nasals, 12.3 (11.8-12.6) ; maxillary toothrow, 8.4 (7.7-8.8). 

Type specimen. — No. 13319, Mus. Vert. Zool., Univ. California; 
$ ad., skin and skull; collected August 13, 1911, by Annie M. 
Alexander. 

Remarks. — This race is apparently most nearly related to lascivus 
of the Sierra Nevada, from which it differs widely in color of under- 
parts. It differs so much from oregonensis, both in color and cranial 
characters, that intergradation between them seems very improbable. 
Specimens from the Warner Mountains, Cal., are intermediate be- 
tween this form and klamathensis ; in color they resemble flaviventris, 
but one of the two individuals has a skull equaling that of klamath- 
ensis, though with smaller bulla?. A large series from Rush Creek, 
Siskiyou County, average slightly less yellowish beneath and have 
somewhat larger skulls than tj'pical specimens, some of them equaling 



1918.] GLAUCOMYS SABRINUS GROUP. 55 

skulls of fuliginosus in size. Intergradation with lascivus occurs in 
the region around Mount Lassen. 

Specimens examined. — Total number, 27, as follows : 

California: 1 Bear Creek, Trinity County (head, altitude, 6,400 feet), 5; 
Castle Lake, Siskiyou County, 2; Grizzly Creek, Trinity County (alti- 
tude, 6,000 feet), 1; Jackson Lake, Siskiyou County, 3; Rush Creek, 
Siskiyou County, 14 ; Warner Mountains, 2. 

GLAUCOMYS SABRINUS LASCIVUS (Bangs). 

Sierra Flying Squirrel. 

[PI. Ill, fig. 5 ; PI. V, fig. 5 ; PI. VII, fig. 1.] 

Sciuropterus alpinus lascivus Bangs, Proc. New England Zool. Club, I, p. 69, 
July 31, 1899. 

Type locality. — Tallac, Eldorado County, California. 

Distribution. — Sierra Nevada Range and northward to eastern 
Shasta County, California. 

Characters. — Similar to flaviventris, but slightly smaller, with 
smaller hind foot; underparts without yellow suffusion. Compared 
with klamathensis: Size smaller; color above more drab (less 
brownish) ; underparts and feet less yellowish. Similar in color to 
stephensi, but paler. 

Color. — Winter pelage : Upperparts wood brown shading to drab ; 
sides of face smoke gray, often shaded with pale fuscous; upper 
surface of flying membrane blackish brown ; ears fuscous-black ; hind 
feet hair-brown, the toes shaded with whitish or buff; fore feet mouse 
gray; tail above, fuscous-black mixed with wood brown or pale 
snuff brown; beneath, drab-gray shading to avellaneous; underparts 
grayish white faintly washed with avellaneous. Summer pelage: 
Upperparts more brownish, usually rich wood brown; underparts 
faintly washed with cartridge buff or light pinkish cinnamon ; upper 
surface of membranes clove brown; hind feet pale hair-brown, the 
soles whitish drab or faintly tinged with olive-buff; under side 
of tail pinkish buff. 

Skull. — Essentially like that of flaviventris; much smaller than 
that of klamathensis and of fuliginosus. 

Measurements. — Average of 5 topotypes (adult and subadult) : Total 
length, 303 (295-320) ; tail vertebras, 133 (125-150) ; hind foot, 40.2 
(39-43). Average of 11 from Cisco, Cal. : 292; 122; 37.5. Skull: 
Average of 4 topotypes (adult and subadult) : Greatest length, 40.4 
(39.5^tl.5); zygomatic breadth, 23.5 (23-24.1); mastoidal breadth, 
18.2 (17.7-18.8) ; least interorbital breadth, 7.4 (7.1-7.8) ; least postor- 
bital breadth, 9.2 (9-9.5) ; length of nasals, 13 (12.7-13.2) ; maxil- 
lary toothrow, 8.1. 

1 All in collection Mus. Vert. Zool., Univ. California. 



56 NORTH AMERICAN FAUNA. I No. 44. 

Type specimen. — No. 9186, Mus. Comp. Zool. (formerly in collec- 
tion of E. A. and O. Bangs) ; 5 subad., skin and skull; collected 
August 28, 1898, by W. W. Price and P. O. Simons. 

Remarks. — The Sierra flying squirrel is closely related to flavi- 
ventris of northern California, but differs markedly from it in the 
color of the underparts, tail, and feet. Intergradation between the 
two forms is shown by specimens from Mount Lassen, Dana, Pratt- 
yille, and Castle Lake, Siskiyou County, which approach ftaviventris 
in having the soles yellowish and the underparts tinged with the 
same. One specimen from Echo, close to the type locality, also has 
yellow soles and a buffy tail, but no yellow on the belly. In a large 
series from Cisco, Cal., most of the specimens show indications of 
being stained on the underparts with soot. Intergradation with 
calif omicus is probable, but there is no material available from the 
southern Sierra to show it. This race is very similar in color to 
alpinws of the northern Rocky Mountains, but is of much smaller size 
and has a paler tail. 

Specimens examined. — Total number, 34, as follows: 

California: Blue Canyon, 3; 1 Cisco, IS; 1 Dana, 1; Echo, l; 2 Fort Crook, 
1; Fresno [=inoiantains near ?], 1 (skeleton) ; Kings River Canyon, 1 ; 
Mill Creek, south base Mount Lassen, 1 ; Mount Tallac, 4 ; 2 ' 3 Prattville 
(12 miles northeast), 1; Quincy, 1; Red Point, Placer County, 1.* 

GLAUCOMYS SABRINUS CALIFORNICUS (Rhoads). 

San Bernardino Flying Squirrel. 

[PI. Ill, fig. 9 ; PI. V, fig. 9 ; PI. VII, fig. 5.] 

Sciuropterus alpinus californicus Rhoads, Proc. Acad. Nat. Sci., Philadelphia, 

p. 323, June, 1S97. 
Sciuropterus californicus Grinnell, Univ. California. Publ. Zool., V, pp. 138-139, 

1908. 

Type locality. — Squirrel Inn, San Bernardino Mountains, Cali- 
fornia (altitude, 5,200 feet). 

Distribution. — San Bernardino and San Jacinto Mountains, Cali- 
fornia. 

Characters. — Similar to lascivus, but upperparts paler and more 
grayish ; face between eyes usually washed with gray ; skull smaller. 

Color. — Unworn pelage (April to August) : Upperparts light 
drab, shaded in some specimens with yellowish wood brown; sides 
of face pale smoke gray ; front of face often washed with gray ; ears 
hair-brown; upper surface of flying membrane fuscous; feet hair- 
brown or mouse gray, the toes whitish or buffy white, the soles buffy 
white or olive-buff; tail above, hair-brown, sparingly mixed with 

1 Collection Mus. Vert. Zool., Univ. California. 

2 Collection Mus. Comp. Zool. 

s Collection Acad. Nat. Sci. Philadelphia. 
4 Collection Leland Stanford, Jr., Univ. 



1918.] GLAUCOMYS SABRINUS GROUP. 57 

sayal brown ; beneath, pinkish buff or pale olive-buff, edged on sides 
with hair-brown; underparts soiled whitish, faintly shaded with 
cartridge buff. 

Skull. — Similar to that of lascivus, but averaging smaller, with 
shorter nasals and smaller bulla?. 

Measurements. — Average of 8 adults from San Bernardino Moun- 
tains, Cal. : Total length, 297 (280-312) ; tail vertebrae, 137 (127-149) ; 
hind foot, 37.6 (36-39). Skull: Average of 5 adults from same 
locality: Greatest length, 39 (37.3-40); zygomatic breadth, 22.8 
(22.3-23.5); mastoidal breadth, 17.7 (17.2-18.1); least interorbital 
breadth, 7.8 (7.4-8.2) ; least postorbital breadth, 8.7 (7.9-9.6) ; length 
of nasals, 11.9 (11.3-12.4) ; maxillarv toothrow, 8.1 (7.5-8.6). 

Type specimen. — No. 10487, Acad. Nat. Sci. Philadelphia (for- 
merly No. 3487, collection S. N. Rhoads) ; $ ad., skin and skull ; 
collected June 5, 1896, by R. B. Herron. 

Remarks. — This subspecies is the grayest of the races of sabrinus, 
being considerably paler than aZpinus. It is closely related to 
laschnts of the Sierra, and probably intergrades with it, though no 
intermediate specimens have thus far been collected. 

Specimens examined. — Total number, 9, as follows : 
California : San Bernardino Mountains, 8 ; " San Jacinto Mountains, l. 2 

GLAUCOMYS SABRINUS STEPHENS! (Mekriam). 

California Coast Flying Squirrel. 

[PI. Ill, fig. 1 ; PI. V, fig. 1 ; PI. VII, fig. 9.] 

Sciuropterus oregonensis stephensi Merriam, Proc. Biol. Soc. Washington, XIII, 
p. 151, June 13, 1900. 

Type locality. — Sherwood, Mendocino County, California. 

Distribution. — Coast region of northern California; limits of range 
unknown. 

Characters. — Similar to lascivus, but upperparts more reddish 
(less drab) ; soles whiter (never tinged with yellowish) and toes less 
distinctly marked with buffy or white; skull with very deep brain- 
case. Compared with oregonensis : "Upperparts and tail less reddish; 
underparts whiter. Compared with fuliginosus : Size much smaller ; 
color above in fresh pelage somewhat grayer (less ochraceous) and 
underparts paler. 

Color. — Fresh winter pelage: Upperparts wood brown; sides of 
face pale smoke gray ; upper surface of flying membrane fuscous or 

1 Three in collection Acad. Nat. Sci. Philadelphia ; three in Mus. Vert. Zool., Dniv. 
California ; one in Mus. Comp. Zool. 

a Collection Mus. Vert. Zool., Univ. California. 



58 NORTH AMERICAN FAUNA. [No. 44. 

fuscous-black ; feet hair-brown, the soles buffy white ; tail above, hair- 
brown, shaded with fuscous-black, darkest at tip ; beneath, pale pink- 
ish buff edged with hair-brown; underparts whitish, irregularly 
washed with light pinkish cinnamon. Worn winter pelage (May) : 
Upperparts decidedly more brownish, about sayal brown; tail above, 
somewhat browner; beneath, light pinkish buff, edged with. fuscous. 
Summer pelage (July) : Similar to spring pelage but underside of 
tail darker (pinkish buff). 

Skull. — Similar to that of oregonensis and of lascivus, but averag- 
ing slightly smaller, with relatively shorter nasals and slightly 
smaller bulla? ; braincase deeper and more abruptly depressed. 

Measurements. — Average of 3 adults from Laj^tonville, Cal. : 
Total length, 307 (300-310) ; tail vertebra?, 143 (133-149) ; hind foot, 
39.3 (38-40) Skull: Average of 3 adults from Laytonville, Cal.: 
Greatest length, 40 (39.7-40.5) ; zygomatic breadth, 23.5 (23-2*4) ; 
mastodial breadth, 18.3 (18.3-18.4) ; least interorbital breadth, 7.9 
(7.6-8.4) ; least postorbital breadth, 9.2 (8.6-9.9) ; length of nasals, 
11.7 (11.1-12.6) ; maxillary toothrow, 7.9 (7.3-8). 

Type specimen.— -No. 99830, IT. S. Nat. Mus. (Biological Survey 
collection) ; $ subad., skin and skull; collected May 10, 1894, by F. 
Stephens. 

Remarks. — This race, as indicated by its skull, is closely related to 
oregonensis; in color it resembles lascivus rather closely, and doubt- 
less intergrades with both it and aregonensis. The type (from Sher- 
wood) and one nearly adult specimen from La3^tonville have skulls 
in which the occipital region is very abruptly decurved, but in two 
other adults from the same locality this character is less pronounced. 
A specimen, without skull, from 8 miles southeast of Cecilville — 
within the known range of flaviventris — agrees essentially with the 
type of stephensi, and thus suggests that intergradation does not 
occur between these two races. 

Specimens examined. — Total number, 6, as follows: 
California: Cecilville (8 miles southeast), 1 ;* Eureka, l; a Laytonville, 3; 3 
Sherwood, 1. 

1 Collection Mus. Vert. Zool., Univ. California. 

2 Collection Leland Stanford, Jr., Univ. 

3 Collection California Acad. Sci. 



TABLES OF MEASUBEMENTS. 



59 



TABLES OF CRANIAL MEASUREMENTS. 
Table I. — Glaucomys volans. 



Species and locality. 




•S 


■3 

03 
CD 

X5 






03 


03 


.a 
?, 


3 
S 




eg 

o 

Ph 


Mm. 


Mm. 


7 


9.1 


7.3 


9.3 


7 


9.2 


6.2 


8.6 


6.5 


8.4 


7.5 


8.9 


6.9 


9.1 


7 


9.4 


6.0 


8.2 


6.9 


8.6 


6.8 


8.3 


7.2 


8.7 


6.8 


9.1 


7.2 


8.3 


7.3 


9 


7.3 


8.4 


7 


8.2 


7.3 


8.9 


7.3 


8.9 


6.9 


8.5 


7.2 


8.7 


7.2 


9.2 



Remarks. 



Glaucomys volans volans: 
Washington, D. C 

Do 

Do 

Do 

Lake George, N. Y... 
Wilmington, Mass. . . 
Elk River, Minn 

Do 

Glaucomys v. querceti: 

Citronelle, Fla 

Do 

Lake Harney, Fla 

Enterprise, Fla 

Glaucomys v. saturatus: 
Dothan, Ala 

Do 

Do 

Greensboro, Ala 

Glaucomys v. texensis: 

Sour Lake, Tex 

Do 

Do 

Do 

Glaucomys v. goldmani: 
Teopisca, Chiapas 



Mm. 
9.6 
9.8 

10.1 
9.2 
9.6 
9.9 
9.1 
9.9 



9.6 
9.6 



9.7 
9.5 
10 
9.7 



9.5 
9.2 



9.2 
9.2 



10.3 
9.9 



Mm. 
6.2 
6.3 
6.5 
6 

5.6 
6.6 
6.5 
6.4 



6.3 
6.4 
6.5 
6.4 



6.4 
6.5 
6.7 
6.2 



6.4 
6.3 



6.2 
6.1 



6.4 
6.3 



Adult. 

Subadult. 

Adult. 

Do. 

Do. 

Do. 

Do. 

Do. 



Adult; type. 
Adult. 
Subadult. 
Do. 



Adult. 
Adult; type. 
Adult. 
Do. 



Subadult. 
Old adult; 

type. 

Adult. 

Do. 



Adult; type. 
Adult. 



1 Collection Mus. Comp. Zool. 
Table II. — Glaucomys sabrinus. 



Species and locality. 



Remarks. 



Glaucomys sabrinus sabrinus: 

Cochrane, Ontario 

Lake Edward, Quebec 

Big Island, Mackenzie 

Winnipeg, Manitoba 

Glaucomys s. makkovikensis 

Makkovik, Labrador 

Do 

Glaucomys s. macrotis: 

Hunter Mountain, N. Y 

Peterboro, N. Y 

Ossipee ; N. H 

Greenville, Me 

Glaucomys s. canescens: 

Portage la Prairie, Manitoba. 
Do 



Mm. 
40 
39.4 
40.2 
39.4 

40.8 
40.4 

37.5 

36.7 

36 

38.7 



3 38.5 
9 38.8 



Mm. 

24 

24.2 

24.2 

24.3 

24.7 
24.4 

22.4 
22.4 
22.3 
23.5 

22.8 
22.9 



Mm. 
18.1 
18.5 
19.2 
18 



18.6 

17.5 
17.7 
17.2 
17.9 

17.5 
17.6 



Mm. 
7.3 
8.1 

8.2 

7.8 

7.3 
7.6 

7 

7.1 
6.9 
6.8 

7.7 
7.6 



Mm. 
8.9 
9.7 
9.2 
9.3 

10 

9.7 

9.3 
9.2 
8.6 



9.2 
9.5 



Mm. 
11.5 
11.4 
12.8 
12 

12.2 
11.1 

10.6 
10.5 
10.5 
10.1 

11.5 
11.2 



Mm. 
7.8 
7.3 

8.1 

7.7 

7.4 
7.9 

7.1 
6.4 
7.2 

6.7 

7.7 
7.6 



Subadult. 
Adult. 
Subadult. 
Do. 

Adult. 
Do. 

Adult; type. 
Adult. 

Do. 

Do. 

Do. 
Adult; type. 



1 Collection Carnegie Mus.; not numbered. 
» Collection Field Mus. Nat. Hist. 



1 Collection Mus. Comp. Zool. 



60 



NORTH AMERICAN FAUNA. 
Table II. — Glaucomys sabrinvs — Continued. 



[No. 44. 



No. 



Species and locality. 



16959 



31674 
169169 



174398 
174397 

3.5320 
180338 

136137 
136138 
2S790 

3 253 

« 146 
141952 

69416 

94310 

s 1026 
20G543 



93282 
93144 

68573 
159393 
159394 



6 5905 
6 5904 

31675 
18499 
18495 



193991 
87