HARVARD UNIVERSITY
LIBRARY
OF THE
MUSEUM OF COMPARATIVE ZOOLOGY
r
UNIVERSITY OF KANSAS PUBLICATIONS
^
MUSEUM OF NATURAL HISTORY
MUS. GOMPTzmLl
LIBRARY
JUN -8 19! :
HARVARD
UNIVERSITY
Volume 1
1946-1950
EDITORS
E. Raymond Hall
Donald S. Farner
Donald F. Hoffmeister
H. H. Lane
A. Byron Leonard
Edward H. Taylor
Robert W. Wilson
Museum of Natural History
University of Kansas
Lawrence, Kansas
1950
MUSEUM OF NATURAL HISTORY
UNIVERSITY OF KANSAS
LAWRENCE, KANSAS
PRINTED BY
FERD VOILAND, JR.. STATE PRINTER
TOPEKA, KANSAS
1950 *
23-2413 u« | y
I','
JUN -
CONTENTS
1. The pocket gophers (genus Thomomys) of Utah. By Stephen D. Durrant.
Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces pluvialis Cope, and noteworthy records
of other amphibians and reptiles from Kansas and Oklahoma. By Hobart
M. Smith. Pp. 85-89. August 15, 1946.
3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp. 93-96, 1
figure in text. August 15, 1946.
4. Hybridization between two species of garter snakes. By Hobart M. Smith.
Pp. 97-100. August 15, 1946.
5. Selected records of reptiles and amphibians from Kansas. By John Breu-
kelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.
6. Kyphosis and other variations in soft-shelled turtles. By Hobart M.
Smith. Pp. 117-124, 3 figures. July 7, 1947.
7. Natural history of the prairie vole (Mammalian genus Microtus). By
E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6, 1947.
8. The postnatal development of two broods of great horned owls (Bubo
virginianus) . By Donald F. Hoffmeister and Henry W. Setzer. Pp. 157-
173, 5 figures in text. October 6, 1947.
9. Additions to the list of the birds of Louisiana. By George H. Lowery, Jr
Pp. 177-192. November 7, 1947.
10. A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-216.
November 29, 1947.
11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa R. and E.
Raymond Hall. Pp. 217-236, 2 figures in text. November 29, 1947.
12. A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest and E
Raymond Hall. Pp. 237-244, 6 figures in text. December 10, 1947.
13. Tadarida feniorosacca (Merriam) in Tamaulipas, Mexico. By Walter W.
Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure in text. December
10, 1947.
14. A new pocket gopher (Thomomys) and a new spiny pocket mouse
(Liomys) from Michoacan, Mexico. By E. Raymond Hall and Bernardo
Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
15. A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-
264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Oklahoma. By
Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kansas and
Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure in text. August
16, 1948.
18. A new species of heteromyid rodent from the Middle Oligocene of north-
east Colorado with remarks on the skull. By Edwin C. Galbreath. Pp.
285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (Genus Proechimys, Family Echi-
myidae). By Joao Moojen. Pp. 301-406, 140 figures in text. December
10, 1948.
20. Three new beavers from Utah. By Stephen D. Durrant and Harold S.
Crane. Pp. 407-417, 7 figures in text. December 24, 1948.
21. Two new meadow mice from Michoacan, Mexico. By E. Raymond Hall.
Pp. 423-427, 6 figures in text. December 24, 1948.
22. An annotated check list of the mammals of Michoacan, Mexico. By E.
Raymond Hall and Bernardo Villa-R. Pp. 431-472, 2 plates, 1 figure in
text. December 27, 1949.
23. Subspeciation in the kangaroo rat, Dipodomys ordii. By Henry W. Setzer.
Pp. 423-573, 27 figures in text, 7 tables. December 27, 1949.
(Concluded on back cover)
24. Geographic range of hooded skunk, Mephitis macroura, with description
of a new subspecies from Mexico. By E. Raymond Hall and Walter W.
Dalquest. Pp. 575-580, 1 figure in text. January 20, 1950.
25. Pipistrellus cinnamomeus Miller 1902 referred to the genus Myotis. By
E. Raymond Hall and Walter W. Dalquest. Pp. 581-590, 5 figures in text.
January 20, 1950.
26. A synopsis of the American bats of the genus Pipistrellus. By E. Raymond
Hall and Walter W. Dalquest. Pp. 591-602, 1 figure in text. January 20,
1950.
Index pp. 605-638.
The Pocket Gophers (Genus Thomomys)
of Utah
BY
STEPHEN D. DURRANT
SEP 6 1916
University of Kansas Publications
Museum of Natural History
Volume 1, No. 1, pp. 1-82, 1 figure in text
August 15, 1946
UNIVERSITY OF KANSAS
LAWRENCE
1946
The Pocket Gophers (Genus Thomomys)
of Utah
BY
STEPHEN D. DURRANT
University of Kansas Publications
Museum of Natural History
Volume 1, No. 1, pp. 1-82, 1 figure in text
August 15, 1946
UNIVERSITY OF KANSAS
Lawrence
1946
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Donald S. Farner,
Donald F. Hoffmeister
Volume 1, No. 1, pp. 1-82, 1 figure in text.
Published August 15, 1946
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1946
21-2786
{ WW>»VT »
$0,55-3 \ 5LP 6 1SM6
The Pocket Gophers (Genusxhomomys) of Utah
By
STEPHEN D. DURRANT
Contribution from the Department of Biology, University of Utah, and the Museum of
Natural History, University of Kansas.
INTRODUCTION
The history of pocket gophers of Utah begins with J. A. Allen's
mention in 1874 of mounds of these animals. For them he employed
the name "Thomomys rufescens?" (1874:65). Actual specimens
were reported upon a year later by Elliot Coues (1875:251, 256),
who used the name Thomomys talpoides for specimens from "Utah"
but later in the same paper listed specimens from Provo as
Thomomys talpoides bulbivorus. Even as the great variation in
Utah pocket gophers has been perplexing to modern workers, so it
was also to Coues seventy years ago who left the problem with
the statement that animals from Provo "exhibit among themselves
such variations that their labelling becomes a matter of indiffer-
ence"! In the same year in another report, Coues and Yarrow
(1875:112) used the name Thomomys talpoides umbrinus for
animals from Provo. In 1877, Coues again referred these same
animals to Thomomys talpoides bidbivorus, using the name um-
brinus for the animals of only southern Utah (Coues, 1877:627, 628).
The two names Thomomys bottae and Thomomys talpoides, now
applicable to gophers in Utah, were synonomized under the name
Thomomys talpoides bidbivorus by Coues (1875:256; 1877:627).
After this beginning only three other papers, all by J. A. Allen,
appeared in the next twenty years. They were reports on collections
of mammals made by Walter W. Granger and Charles P. Rowley.
One of these contained the description of Thomomys aureus. Like-
wise, in the ensuing twenty years there were only three papers, one
in 1901 by C. Hart Merriam in which he described Thomomys
uinta, one by Allen (1905:119), and Vernon Bailey's (1915) "Re-
vision of the pocket gophers of the genus Thomomys" in which he
summarized the information then available on these animals within
the state. Barnes (1922 and 1927) reprinted the information sum-
marized by Bailey. Since 1927 approximately twenty-five papers,
mostly taxonomic, have been published in which reference is made
to Utah gophers, and especially since 1930 much information has
been accumulated about the distribution and speciation of this
genus within the state.
Specimens to the number of 1,045 have been available for this
study. Whereas Bailey doc. cit.) listed only four kinds belonging
(3)
University of Kansas Publs., Mus. Nat. Hist.
s
to four different species, thirty-five kinds are now known from Utah.
Seven of these are herein described as new. The thirty-five kinds
are found to belong to only two instead of four full species.
Inasmuch as the literature is scattered and since names have been
applied in different ways at different times, I have attempted to
give a synonomy as complete as possible for each form found
within the state.
The bibliographies of Hayward (1936 and 1941) and Miller's
(1924) "List of North American mammals" have been of great use.
Capitalized color terms in the accounts are after Ridgway, Color
Standards and Color Nomenclature, Washington, D. C, 1912.
In the lists of specimens examined, the localities are listed by
counties from west to east, beginning at the northwestern corner
of the state, and within each county from north to south. When
two localities are on the same latitude, the westernmost is listed
first.
I am deeply indebted to Professor R. V. Chamberlin, of the University of
Utah, for encouragement and support in my investigation. I also acknowledge
critical assistance in the preparation of this paper from Professor E. Raymond
Hall of the University of Kansas. For the loan of specimens I am grateful
to the following: Clinton G. Abbott and Lawrence M. Huey, Natural History
Museum of San Diego, San Diego, California; Harold E. Anthony and J.
Eric Hill, American Museum of Natural History, New York City, New York;
Seth B. Benson, Museum of Vertebrate Zoology, University of California,
Berkeley, California; William H. Burt, Museum of Zoology, University of
Michigan, Ann Arbor, Michigan; J. Kenneth Doutt, Carnegie Museum, Pitts-
burgh, Pennsylvania; Ross Hardy, Dixie Junior College, St. George, Utah;
C. Lynn Hayward and Vasco M. Tanner, Brigham Young University, Provo,
Utah; H. H. T. Jackson and Viola S. Schantz, United States Fish and Wild-
life Service, U. S. National Museum, Washington, D. C; Remington Kellogg
and Alexander Wetmore, U. S. National Museum, Washington, D. C; J. S.
Stanford, Utah State Agricultural College, Logan, Utah.
Unless otherwise indicated, specimens are in the Museum of Zoology,
University of Utah, Salt Lake City, Utah. In lists of specimens examined,
abbreviations are employed as follows:
(A. M. N. H.) American Museum of Natural History.
(N. H. M. S. D.) . . Natural History Museum of San Diego.
(M. V. Z.) Museum of Vertebrate Zoology, University of California.
(U. M.) Museum of Zoology, University of Michigan.
(C. M.) Carnegie Museum.
(R. H.) Collection of Ross Hardy.
(B. Y. U.) Brigham Young University.
(U. S. N. M.) United States National Museum.
(U. S. A. C.) Utah State Agricultural College.
(K. U.) Museum of Natural History, University of Kansas.
Durrant — Pocket Gophers of Utah
Fig. 1. Map showing the distribution of species and subspecies of pocket
gophers in Utah.
Guide to subspecies:
12.
T.
b.
aureiventris
24.
T.
b.
lenis
1. T.
t.
gracilis
13.
T.
b.
robustus
25.
T.
b.
levidensis
2. T.
t.
wasatchensis
14.
T.
b.
minimus
26.
T.
b.
osgoodi
3. T.
t.
oquirrhensis
15.
T.
b.
ncsophihis
27.
T.
b.
howelli
4. T.
t.
uinta
16.
T.
b.
stansburyi
28.
T.
b.
wahwahensis
5. T.
t.
■pygmaeus
17.
T.
b.
albicaudatvs
29.
r.
b.
dissimilis
6. T.
t.
ravus
18.
T.
b.
bonnevilln
30.
T.
b.
aureus
7. T.
t.
ocius
19.
T.
b.
centralis
31.
T.
b.
birdseyei
8. T.
t.
moorei
20.
T.
b.
sevieri
32.
T.
b.
virgineus
9. T.
t.
fossor
21.
T.
b.
convexus
33.
T.
b.
planirostris
10. T.
t.
■parowanensis
22.
T.
b.
tivius
34.
T.
b.
absonus
11. T.
t.
levis
23.
T.
b.
contractus
35.
T.
b.
alexandrae
6 University of Kansas Publs., Mus. Nat. Hist.
Genus Thomomys Wied
All pocket gophers of Utah belong to the genus Tho?nomys. There
are only two species within the state, Thomomys bottae with twenty-
four subspecies and Thomomys talpoides with eleven subspecies.
Due to marked mutational capacities and ready response to en-
vironmental pressures and sedentary habits, pocket gophers differ-
entiate readily into numerous subspecies. It is well known that
Utah by its highly varied topography and climate possesses widely
different types of habitats. The aforementioned plasticity of these
animals and possibly the fact that both species are at the extreme
limits of their ranges in Utah account for the numerous forms found
within the state.
The genus may be characterized as follows: Highly specialized
fossorial rodents, with heavy, thick bodies; all four legs of ap-
proximately equal length, but front legs more muscular for
digging, and feet provided with long claws; external fur-lined
cheek pouches; small eyes, short ears and tail; upper incisors long
and projecting external to lips. Skull: Stout and flattened; zygo-
matic arches well developed and usually widely spreading; all teeth
with permanent pulp cavities; incisors superficially smooth, but fine
median groove present on anterior face of each upper incisor;
dental formula, '• — • c- — • p- — • m- — ; external auditory canal long; sta-
pedial artery small and enclosed within an osseous canal.
Thomomys talpoides (Richardson)
Thomomys talpoides is a northern species that in Utah approaches
the southern limits of its range. The animals of this species inhabit
the mountains and high valleys. In the southward extension of
their range, as in Utah, they are found at higher elevations which
zonally represent lower elevations at more northern latitudes. The
specific characters are: Sphenorbital fissure absent; incisive fora-
mina anterior to infraorbital canal; anterior prism of P4 triangular;
interparietal relatively large; lambdoidal suture concave posteri-
orly in region of interparietal, in Utah specimens.
Thomomys talpoides gracilis Durrant
Thomomys quadrat us gracilis Durrant, Bull. Univ. Utah, 39 (No. 6) :3,
February 28, 1939.
Thomomys talpoides gracilis Durrant, Bull. Univ. Utah, 30 (No. 5) :6,
August 24, 1939; Goldman, Journ. Mamm., 25:414, December 12, 1944.
Thomomys quadratics fishrri Hall, Univ. California Publ. Zool., 37:4,
April 10, 1931.
Durrant — Pocket Gophers of Utah 7
Thomomys uinta Bailey, N. Amer. Fauna, 39:114, November 15, 1915;
Barnes, Bull. Univ. Utah, 12 (No. 15) :83, April, 1922; Bull. Univ. Utah,
17 (No. 12): 104, June, 1927.
Type. — Male adult, skin and skull; No. 44866, Museum of Vertebrate Zo-
ology, University of California; Pine Canyon, 6,600 ft., 17 mi. NW Kelton,
Box Elder County, Utah; July 12. 1930; collected by Annie M. Alexander;
original number 676.
Range. — Mountainous regions of extreme northwestern Utah.
Diagnosis. — Size medium (see measurements). Color: Upper parts Buck-
thorn Brown grading over the sides and flanks to Light Buff on the under-
parts; chin white; nose and postauricular patches grayish black. Claws on
front feet long and slender. Skull: Long and slender; rostrum long and
narrow; zygomatic and mastoidal breadths slight; palatal pits deep; upper
incisors narrow; basioccipital wide.
Comparisons. — Compared with topotypes of Thomomys talpoides
jisheri, gracilis is of approximately the same size. Upper parts
darker and underparts lighter; postauricular patches larger and
darker; claws on front feet longer and slenderer. Skull: Generally
longer and narrower; nasals and rostrum longer; basioccipital wider.
As compared with T. t. uinta, gracilis is of approximately the
same size but differs as follows: Color: Lighter throughout; post-
auricular patches markedly smaller and lighter; inguinal and pec-
toral regions much lighter. One characteristic difference is in
the ear. In uinta the external opening of the ear is much larger;
the pinna of the ear is larger, more rounded at the tip, and lacks
most of the pigmentation on the inner margin. Skull: Generally
narrower and longer; nasals longer; zygomatic arches weaker and
less angular; upper incisors narrower.
This form is easily distinguished from bridgeri by smaller size,
and by the skull being longer, narrower and less angular.
From Thomomys talpoides oquirrhensis to the southeast, T. t.
gracilis ean be distinguished by: Total length and ear shorter.
Color: Generally lighter, except the underparts which are about
fhe same; postauricular patches larger and more deeply pigmented.
Skull: Braincase less inflated; nasals truncated posteriorly as op-
posed to rounded; zygomatic and mastoidal breadths less; rostrum
shorter but narrower; upper incisors narrower and shorter.
For comparisons with ivasatchensis see comparisons under that
form.
In general, this mountain form can be distinguished from all other
talpoides in Utah by lighter color, narrow, slender, "graceful" skull
whence the name gracilis is derived.
8 University of Kansas Publs., Mus. Nat. Hist.
Remarks. — In Utah, gracilis is limited to the extreme north-
western corner of the state. This part of the state is in the Snake
River drainage. The main part of the range of this race lies in
south-central and southwestern Idaho and northeastern Nevada.
The center of its range might be considered to be in the Jarbidge
Mountains area of Nevada. The south slopes of these mountains are
in the Humboldt River drainage, while the north slopes are in the
Snake River drainage, and this subspecies occurs as far north as the
Snake River and south and west almost to central Nevada. No
specimens are available from the area in Utah between the Raft
River Mountains inhabited by gracilis and the Wasatch Mountains
in central Utah inhabited by wasatchensis. Judging from the nature
of the terrain, the range of gracilis does not extend eastward much
beyond the Raft River Mountains. The type locality for a gopher
of a different species, Thomomys bottae aureiventris, is in the first
valley east of these mountains. Furthermore, all valleys to the east
and south, as far as known, are inhabited by gophers of the bottae
group. Also, all mountain ranges in this area, as far east as the
Wasatch Mountains are inhabited by members of the bottae group.
No specimens from Utah indicate intergradation between gracilis
and wasatchensis, the form to the east, but specimens from farther
north at Albion, Cassia County, Idaho, do show intergradation.
Bailey (1915:116), Hall (1931:4), and Durrant (1939:6) have re-
ported on these specimens which at the present time seem best re-
ferred to T. t. gracilis.
Specimens examined. — Total, 24, distributed as follows: Box Elder County: Yost, 4
(U. S. A. C.)i Pine Canyon, 0,600 ft., 17 mi. NW Kelton, 7 (M. V. Z.): Lynn Canyon,
Raft River, 4; Park Valley, 3 (U. S. A. C); Etna, 4 (U. S. A. C); Raft River Mountains,
Clear Creek Camp of Minnedoka National Forest, 1 (R. H.); Raft River Mountains, 1,500
feet above Clear Creek Camp of Minnedoka National Forest, 1 (R. H.).
Thomomys talpoides wasatchensis new subspecies
Thomomys quadratus uinta Hall, Univ. California Publ. Zool., 37:4,
April 10, 1931.
Thomomys talpoides vinta Goldman, Journ. Mamm., 20:234. May 14,
1939.
Thomomys uinta Bailey, N. Amer. Fauna, 39:114, November 15, 1915;
Barnes, Bull. Univ. Utah, 12 (No. 15) :83, April, 1922; Bull. Univ. Utah,
17 (No. 12) :104, June, 1927; Stanford. Journ. Mamm., 12:360, November
11, 1931.
Type. — Male, adult, skin and skull, No. 1604, Museum of Zoology, Uni-
versity of Utah; Midway, 5,500 ft., Wasatch County, Utah; September 1,
1936; collected by S. D. Durrant; original number 1049.
Range. — Wasatch Mountains and neighboring high valleys as far south as
Spanish Fork Canyon, Utah County.
Durrant — Pocket Gophers of Utah 9
Diagnosis. — Size medium (see measurements). Color: Upper parts Snuff
Brown, finely mixed with black; sides and flanks Sayal Brown; underparts
overlaid with Cinnamon Buff, with suffusion of black on underfur; postau-
ricular patches black, extending around ear; ears pointed and covered with
Liack hairs; nose, cheeks, chin and top of head dusky; front feet, hind feet
and distal part of tail white; tail covered proximally with light brown hairs.
Skull : Moderately heavy and ridged ; nasals long, wide posteriorly and not
markedly dilated distally ; posterior ends of nasals emarginate ; zygomatic arches
fairly widely spreading and angular, being nearly straight in adults, but tending
to bow out slightly at posterior ends in young; zygomatic processes of maxillae
heavy; interparietal small and variously shaped, but always wider than long;
interorbital region fairly wide; well marked dorsal depression in frontals post-
erior to ends of nasals; interpterygoid space narrowly V-shaped; tympanic
bullae large; occipital condyles large and widely separated; foramen mag-
num large and higher than wide; basioccipital wide; dentition light.
Comparisons. — From topotypes of Thomomys talpoides moorei,
wasatchensis differs as follows: Size slightly larger; ears longer and
more pointed. Color: Generally darker throughout; postauricular
patches smaller. Skull: Zygomatic arches not as widely spreading;
zygomatic processes of squamosals dip farther ventrally; premax-
illae less extended posterior to nasals; nasals wider posteriorly and
less dilated distally; median dorsal depression of frontals present;
tympanic bullae generally larger, but less inflated ventrally; fora-
men magnum larger especially in dorsoventral dimension; occipital
condyles farther apart; basioccipital wider; alveolar length of
upper molar series less; molariform teeth smaller; upper incisors
wider and shorter.
Topotypes of wasatchensis differ from topotypes and near topo-
types of Thomomys talpoides uinta as follows: Size larger in every
measurement taken. Color: Darker throughout; ears longer and
more pigmented; opening of external ear smaller; postauricular
patches larger. Skull: In females larger throughout, more mas-
sive and angular; nasals longer, wider and not so dilated distally;
rostrum longer but wider; zygomatic arches wider, more angular
and less widely spreading posteriorly; extension of premaxillae
posterior to nasals less; tympanic bullae larger, but less inflated
ventrally; foramen magnum larger and more ovoid; width across
occipital condjdes greater; basioccipital wider; molariform teeth
smaller; upper incisors shorter and wider.
Topotypes of wasatchensis can be distinguished from those of
Thomomys talpoides oquirrhensis as follows: Size larger; tail
longer; ears longer. Color: Slightly darker on sides and under-
parts. Skull: Heavier, more ridged and angular; nasals more di-
10 University of Kansas Publs., Mus. Nat. Hist.
lated distally; posterior ends of nasals more deeply emarginate;
zygomatic arches heavier and more widely spreading, but more
nearly parallel and less divergent posteriorly; zygomatic processes
of maxillae much heavier; braincase and tympanic bullae larger;
pterygoid hamulae shorter; interpterygoid space more narrowly
V-shaped; wider across occipital condyles; foramen magnum larger
and more ovoid.
From topotypes of Thomomys talpoides gracilis, wasatchensis
differs as follows: Size larger; hind foot longer; ears longer and
more pointed. Color: Darker throughout; postauricular patches
relatively smaller. Skull: Larger, heavier and more angular;
nasals emarginate posteriorly as opposed to truncate; rostrum heav-
ier; zygomatic arches heavier and more widely spreading; zygo-
matic processes of maxillae much heavier and more angular; mas-
toid breadth greater; interparietal relatively smaller; extension of
premaxillae posterior to nasals actually as well as relatively less;
palatal pits deeper; tympanic bullae larger; interpterygoid space
more narrowly V-shaped; foramen magnum more ovoid; upper in-
cisors wider.
Topotypes of wasatchensis can be readily distinguished from
those of Thomomys talpoides levis and parowanensis by larger size;
more massive, ridged, angular skulls; larger tympanic bullae; large,
ovoid foramen magnum; and relatively smaller interparietal.
Remarks. — Specimens from Mount Timpanogos and environs are
intergrades between moorei and wasatchensis. They resemble
moorei in the shape and size of the tympanic bullae, and are inter-
mediate in the size and shape of the foramen magnum. In the ma-
jority of characters they resemble wasatchensis to which they are
here referred. The animals from east of Salt Lake City in Salt
Lake County are intergrades between oquiirhensis and wasatchensis
and show some characters of uinta, but are referable to wasatch-
ensis. Animals from Morgan County and western Summit County
are intergrades between wasatchensis and uinta. They resemble
uinta in size, shape of nasals and size of tympanic bullae. The re-
mainder of the cranial details place them with ivasatchensis. Mor-
phologically the animals from Wellsville, Cache County, were the
closest to the topotypes of any obtained and are nearly indistin-
guishable from them. Like the topotypes of wasatchensis this popu-
lation inhabits a high valley. The remaining specimens from Cache
County resemble those from Morgan and Summit counties.
Durrant — Pocket Gophers of Utah 11
SpeciTnens examined. — Total, 119, distributed as follows: Cache County: Logan Canyon,
Beaver Basin, Utah-Idaho Line, 2 (U. S. A. C.)i Logan Canyon, Tony Grove Camp, 6
(U. S. A. C); Logan Canyon, Green Camp, 3 (U. S. A. C); Logan Canyon, 3 (U. S. A. C);
Logan Mountains, 20 mi. E Logan, 3 (U. S. A. C); Logan Peak area, 13 (U. S. A. C);
near Providence Peak, Logan Mountains, 1 (U. S. A. C.) ; Wellsville, 10 (U. S. A. C);
Hardware Ranch, Blacksmith Fork, 1 (U. S. A. C.) ; Avon, 1 (U. S. A. C); 1 mi. E Avon,
1 (U. S. A. C); 7-8 mi. E Avon, 1 (U. S. A. C). Weber County: South Fork, Ogden River,
18 mi. E Ogden, 4 (M. V. Z.). Morgan County: East Canyon, 18 mi. NW Park City, 6,000
ft., 1. Davis County: 8 mi. NE Salt Lake City, 1. Salt Lake County: Mouth of Dry
Canyon, 1 mi. NE Salt Lake City, 1 ; 4 mi. above mouth City Creek Canyon, 5,000 ft., 1 ;
mouth of Emigration Canyon, 1 ; mouth of Millcreek Canyon, 1 ; Lambs Canyon, 13 mi.
SE Salt Lake City, 2 (C. M.); mouth of Big Cottonwood Canyon, 1. Summit County:
Park City, 1 (U. S. N. M.). Wasatch County: Midway, 5,500 ft., 29. Utah County: Mt.
Timpanogos, 1 mi. N Aspen Grove, 7,500 ft., 20; Aspen Grove, Mt. Timpanogos, 5 (1, U. S.
A. C. ; 4, B. Y. U.); Head of Grove Creek, Mt. Timpanogos, 4 (B. Y. U.).
Additional Records: Weber County: Ogden, 6. Salt Lake County: Parleys Canyon,
1 (Bailey, 1915:114).
Thomomys talpoides oquirrhensis Durrant
Thomomys talpoides oquirrhensis Durrant, Bull. Univ. Utah, 30 (No.
5): 3, October 24, 1939.
Type. — Male, adult, skin and skull; No. 2605, Museum of Zoology, Uni-
versity of Utah; Settlement Creek, Oquirrah Mountains, 6,500 ft., Tooele
County, Utah; June 11, 1938; collected by S. D. Durrant; original number 1461.
Range. — Known only from the Oquirrh Mountains, which are in Salt Lake,
Tooele and Utah counties, Utah.
Diagnosis. — Size medium (see measurements); ear long; tail short, claws
of front feet long and slender. Color: Upper parts Buckthorn Brown, mixed
with black, grading over the sides and flanks to Pinkish Buff on the ventral
surface ; feet white ; nose grayish black ; postauricular patches medium in size
and black; chin and throat with varying amounts of white; proximal two-
thirds of tail dark brown, distal third white. Skull : Long and slender, but
relatively wide across mastoidal region; nasals long and rounded posteriorly;
rostrum long and narrow; zygomatic arches weak and not widely spreading,
tending to be slightly bowed out posteriorly, but in the main roughly parallel
to the sides of the skull; outer margin of zygomatic arch slightly concave,
and zygomatic arch dips deeply ventrad; dorsal surface of skull smooth, with
weakly defined parietal crests; parietal crest nearly parallel, but bowed me-
dially, in parietal region, and flaring widely posteriorly to pass lateral to inter-
parietal; tympanic bullae large, truncate anteriorly and markedly inflated
ventrally; upper incisors short and fairly robust.
Comparisons. — From Thomomys talpoides uinta, oquirrhensis
may be differentiated as follows: Color: Darker throughout; post-
auricular patches larger and darker; ears longer and more pointed;
inner margin of pinna heavily pigmented; external opening of ear
smaller. Skull: Nasals rounded posteriorly rather than deeply
emarginate, and less flaring distally; zygomatic arches weaker and
markedly less widely spreading; pterygoid hamulae weaker; basi-
sphenoid narrower; upper incisors shorter and wider.
For comparisons between oquirrhensis and Thomomys talpoides
12 University of Kansas Publs., Mus. Nat. Hist.
gracilis, and oquirrhensis and wasatchensis, see comparisons under
those forms.
Topotypical specimens of oquirrhensis can be distinguished from
those of Thomomys talpoides moorei as follows: Color generally
darker, due to greater admixture of black; terminal bands of hair
actually lighter; postauricular patches larger and darker; ears
longer, more pointed and with more heavily pigmented pinnae; tail
shorter. Skull: About the same size; smoother; zygomatic arches
weaker and less widely spreading; nasals rounded posteriorly as
opposed to emarginate; mastoid breadth less; pterygoid hamulae
weaker; upper incisors wider.
Remarks. — This race is limited to the Oquirrh Mountains, a high
mountain range that lies parallel to, and just west of the Wasatch
Mountains, in Utah, Salt Lake and Tooele counties. These moun-
tains were connected in past times to the Wasatch Mountains by
the Transverse Range, and by a sand and gravel bar deposited by
Pleistocene Lake Bonneville. The Jordan River in its course from
Utah Lake to the Great Salt Lake has cut a channel through the
aforementioned bar. This channel has been cut to the level of the
surrounding valleys as is indicated by the meandering nature of the
stream through this part of its course. As a result the Oquirrh
Mountains are relatively isolated. Although separated from the
Wasatch Mountains by the Jordan River Valley only a few miles
wide, the pocket gophers are distinct on each mountain. A popu-
lation of T. bottae is interposed between the two mountain ranges
as is indicated by specimens from Riverton, six miles north of the
Transverse Range. The populations of bottae are subspecifically
the same on the two sides of the Jordan River.
On the east side of the Oquirrh Mountains, pocket gophers col-
lected from the Jordan Valley up Rose Canyon to about 5,000 feet
elevation were all of the species T. bottae. Between 5,000 and
6,000 feet there is an area in which the ranges of bottae and tal-
poides overlap. When trapping, it is possible to predict what spe-
cies will be taken by the types of burrows and soil. Gophers of
the bottae group have their burrows in the areas of the deepest soil
and heaviest vegetation, whereas the areas of shallow, rocky soil
covered with sparse vegetation are the habitat of talpoides. Above
6,000 feet the only gopher encountered is talpoides. Along Settle-
ment Creek on the west side of the Oquirrh Mountains, which is
the type locality of oquirrhensis, bottae and talpoides have essen-
Durrant — Pocket Gophers of Utah 13
tially the same vertical distribution as in Rose Canyon. On this
mountain the two species appear to be in competition.
The available information, based on collections, indicates that
the Oquirrh Mountains are the only mountains west of the Wasatch
Range upon which talpoides occurs. In Utah, all other mountains
to the west, as far as known, are inhabited by subspecies of
of Thomomys bottae.
Specimens examined. — -Total, 41, as follows: Tooele Covnty: Settlement Creek, Oquirrh
Mountains, 6,500 ft., 14. Salt Lake County: Rose Canyon, Oquirrh Mountains, 5,650 ft., 27.
Thomomys talpoides uinta Merriam
Thomomys uinta Merriam, Proc. Biol. Soc. Washington, 14:112, July
19, 1901; Bailev, N. Amer. Fauna, 39:113, November 15, 1915; Barnes,
Bull. Univ. Utah, 12 (No. 15) :83, April, 1922; Bull. Univ. Utah, 17 (No.
12) :104, June, 1927; Stanford, Journ. Mamm, 12:360; November 11, 1931;
Goldman, Journ. Washington Acad. Sci., 28:333, July 15, 1938; Davis,
The Recent mammals of Idaho, pp. 239, 259, The Caxton Printers, Ltd.,
Caldwell, Idaho, April 5, 1939.
Thomomys talpoides uinta Goldman, Journ. Mamm., 20:234, May 14,
1939.
Thomomys quadratus uinta Hall, Univ. California Publ. Zool., 37:4,
April 10, 1931.
22501
Type. — Male, adult, skin and skull, No. 3005T' U. S. National Museum
(Biological Surveys Collection) ; north base Gilbert Peak, Uinta Mountains,
10,000 ft., Summit County, Utah; June 6, 1890; collected by Vernon Bailey;
original number 1262 (after Merriam, type not seen).
Range. — Uinta Mountains in Duchesne County, eastern Wasatch and Sum-
mit counties, and western Uintah County south to the Roan, Brown and Book
cliffs in Carbon County.
Diagnosis. — Size medium (see measurements). Color: Upper parts Snuff
Brown finely mixed with black, paling over sides and flanks to near Pinkish
Buff on underparts ; postauricular patches relatively small and dusky ; external
opening of ear large; pinnae usually lightly pigmented; hind feet white; front
feet usually white only at base of toes; distal third to half of tail white; tail
usually light below, with proximal dorsal half covered with darker hairs; nose,
chin, cheeks and top of head dusky; usually considerable white on throat.
Skull: Small, slender, and not heavily ridged; nasals short and dilated dis-
tally; posterior margins of nasals emarginate; zygomatic arches moderately
widely spreading, widest posteriorly; interparietal pentagonal or subquadrang-
ular; interpterygoid space V-shaped; tympanic bullae well inflated ventrally;
upper incisors long and narrow.
Comparisons. — For comparisons with other subspecies of Thom-
omys talpoides, see accounts of those forms.
Remarks. — The range formerly ascribed to uinta (Bailey,
1915:114; Barnes, 1922:83, 1927:104) is now known to be inhabited
by animals belonging to three distinct subspecies. The range of
uinta as now understood is restricted to the southern and western
14 University of Kansas Plbls., Mus. Nat. Hist.
parts of the Uinta Mountains and their environs. Three specimens
from the Book Cliffs, Sunnyside, Carbon County, are not typical,
but in a majority of their characters agree with uinta to which they
are here referred.
I have seen only one specimen from the type locality. It is one
of the series on which Merriam (1901:112) based his original de-
scription. In addition, I have studied several large series of near
topotypes. From the material at hand, and from Merriam's descrip-
tion (loc. cit.) , I regard the animals on which the name uinta was
based as intergrades between Thomomys talpoides ravus, the race
to the northeast, on the one hand and the animals of the western
and southern parts of the Uinta Mountains on the other hand. The
affinities of the type series are with the animals from the latter area
which are here all referred to uinta.
Specimens examined. — Total, 41, distributed as follows: Summit County: 2 mi. S junc-
tion Bear River and Haydens Fork, 2 (C. M.): N base, Gilbert Peak. 10,000 ft., 1 (U. S.
N. M.); Smith and Moorehouse Creek, 2; Bald Peak, 25 mi. NE Kamas, 15 (8, M. V. Z. ;
6, C. M.). Duchesne County: Petty Mountain, 15 mi. N Mountain Home, 9,500 ft., 6
(C. M.). Wasatch County: Wolf Creek Pass, 18 mi. NW Hanna, 1 (U. S. A. C); Lost
Lake, Uinta Mountains, 10 (B. Y. U.); Current Creek, Uinta Mountains, 1 (U. S. N. M.).
Carbon County: Forks, Sunnyside, 9,000 ft,, 3.
Additional records. — Summit County: Uinta Mountains, 6 (see Bailey, 1915:114).
Thomomys talpoides pygmaeus Merriam
Thomomys -pygmaeus Merriam. Proc. Biol. Soc. Washington, 14:115.
July 19, 1901.
Thomomys talpoides pygmaeus Davis, The Recent mammals of Idaho,
p. 252, The Caxton Printers, Ltd., Caldwell, Idaho, April 5, 1939.
Type. — Male, adult, skin and skull, No. 55251, U. S. National Museum
(Biological Surveys Collection) ; 10 mi. NE Montpelier, in open sagebrush of
Transition Zone, 6,600 ft., Bear County, Idaho; July 29, 1893; collected by
Vernon Bailey; original number 4150 (after Merriam, type not seen; see. also,
Bailey, 1915:109).
Range. — Limited to Daggett County.
Diagnosis. — Size: Small (see measurements). Color: Upper parts near
Bister slightly mixed with black, grading over sides and flanks to Ochraceous
Buff on underparts; postauricular patches small and dusky; hind feet white;
front feet dusky, being white only at base of claws; chin and nose dusky;
tail brown, lighter below and tipped with white. Skull: Very small, slender
and smooth; nasals short and slender; zygomatic arches weak and not widely
spreading; rostrum narrow; extension of premaxillae posterior to nasals short;
parietal ridges hardly noticeable ; interparietal large ; extension of supra-
occipital posterior to lambdoidal suture long; tympanic bullae actually small,
but relatively large; basioccipital narrow; interpterygoid space narrow and
acutely angled; upper incisors markedly recurved; molariform teeth rela-
tively large.
Durrant — Pocket Gophers of Utah 15
Comparisons. — This small pocket gopher can be distinguished
from all other members of Thomomys talpoides occurring in Utah
by remarkably small size, and slender, weak, small skull with
strongly recurved upper incisors.
Remarks. — The specimens used in this study were those recorded
by Svihla (1931:261). She reports that they were obtained in the
flood-plain banks of the streamsides, and preferred the pine belt.
This shows probably an extension of range with reference to life
zones, as heretofore the main reported localities of capture have
been in sagebrush in the Transition Life-zone.
Insofar as I am aware, Mrs. Svihla's specimens are the only ones
of this subspecies ever obtained in Utah. Additional work is neces-
sary in southwestern Wyoming to outline accurately the geographic
distribution of this subspecies. In comparison with topotypes, the
specimens from Utah are lighter in color and some specimens have
slightly larger skulls, suggesting slight intergradation with Thom-
omys talpoides uinta.
Specimens examined. — Total, 18 (all in Museum of Zoology, University of Michigan),
distributed as follows: Daggett Comity: Sheep Creek, 4; 1 mi. W Summit Springs, 4;
Beaver Creek, 22 mi. S Manila, 9; Granite Park, 24 mi. S Manila, 1.
Thomomys talpoides ravus new subspecies
Type. — Male, adult, skin and skull, No. 13690, Carnegie Museum; Vernal-
Manila Highway, 19 mi. N Vernal, 8,000 ft., Uintah County, Utah; August
22. 1937; collected by J. K. and M. T. Doutt; original number 4718.
Range. — Uinta Mountains in Daggett, northern Uintah and northern Sum-
mit counties.
Diagnosis. — Size large (see measurements); ears relatively narrow; hind
foot relatively small. Color: Upper parts between Drab and Light Drab,
darkest along middorsal line due to mixture of hairs tipped with light brown;
sides and flanks Light Drab; entire underparts creamy white; front and hind
feet, ventral surface of tail and end of tail white; proximal two-thirds of
tail covered dorsally with light brown hairs; nose and cheeks dusky; post-
auricular patches black. Skull: Large, heavy and ridged; rostrum long and
narrow; nasals long, moderately dilated distally and with a distal hump;
posterior ends of nasals emarginate; parietal and lambdoidal crests well de-
veloped; zygomatic arches moderately heavy and widely spreading, widest
posteriorly; zygomatic processes of maxillae moderately heavy and flaring
abruptly from base of rostrum; marked middorsal depression in frontals pres-
ent; interparietal pentagonal; extension of premaxillae posterior to nasals
long; posterior tongues of premaxillae long, slender and rounded proximally;
braincase high, vaulted and relatively narrow; tympanic bullae well inflated
ventrally, and ridged in old animals; pterygoid hamulae long; interptergoid
space narrowly V-shaped; upper incisors long and narrow; molariform teeth
medium.
16 University of Kansas Publs., Mus. Nat. Hist.
Comparisons. — Compared with topotypes of Thomomys talpoides
bridgeri, ravus differs as follows: Size larger; hind foot smaller;
ears narrower. Color: Lighter throughout, grayish as opposed to
brown. Skull: Smaller, narrower, less angular and less massive;
nasals, rostrum, zygomatic processes of maxillae, ascending branches
of premaxillae and posterior tongues of premaxillae all narrower;
extension of premaxillae posterior to nasals longer; interparietal
wider; braincase higher and narrower; tympanic bullae approxi-
mately the same size, but more inflated ventrally; interpterygoid
space more narrowly V-shaped; upper incisors narrower; molari-
form teeth weaker.
Compared with topotypes and near topotypes of Thomomys
talpoides uinta, ravus differs as follows: Size larger in every meas-
urement taken. Color: Lighter throughout, being grayish as op-
posed to brown. Skull: Larger in every measurement taken; ros-
trum and nasals actually as well as relatively longer; extension of
premaxillae posterior to nasals longer; upper incisors longer and
wider; molariform teeth larger.
There is only one other gray subspecies of Thomomys talpoides
in Utah, Thomomys talpoides ocius. Topotypes of ravus differ
from it as follows: Size markedly larger in every measurement
taken. Color: Darker, more brown hairs. Skull: Larger in every
measurement taken; premaxillae extended farther posteriorly to
nasals; extension of supraoccipital posterior to lambdoidal suture
markedly less; tympanic bullae actually as well as relatively
smaller; upper incisors longer and more procumbent.
This new subspecies can be readily distinguished from all other
subspecies of Thomomys talpoides occurring in Utah by markedly
greater size and paler, more grayish color.
Remarks. — The range of this form appears to be limited to the
north slopes of the Uinta Mountains, except in Daggett County
where it occurs also on the south slopes. Intergradation in color
and in cranial details with bridgeri is shown by animals from the
East Fork of Blacks Fork, thirty-one miles SSW Fort Bridger, and
by those from Henrys Fork, 8,300 ft., both in Summit County. Due
to the grayish color and the narrower, weaker skull they are re-
ferred to ravus. Intergradation with uinta is shown by specimens
from the type locality of the latter race. The type series of uinta
consists of intergrades between ravus and the animals to the west
and south (see remarks under uinta) .
Durrant — Pocket Gophers of Utah 17
It is doubtful whether bridgeri occurs in Utah. Material from
Rich County and extreme northern Cache County would settle the
question. Perhaps bridgeri is restricted to the lower valleys in
southwestern Wyoming. Two specimens from northern Cache
County, from Logan Canyon, Beaver Basin, Utah-Idaho Line appear
to be intergrades between bridgeri and wasatchensis, but are refer-
able to the latter race.
Specimens examined. — Total, 38, distributed as follows: Summit County: Henrys Fork,
8,300 ft., 8; E Fork, Blacks Fork, 31 mi. SSW Fort Bridger, 4 (C. M.). Daggett County:
Venial-Manila Road, 4 mi. W Green's Lake, 7,500 ft., 6 (C. M.); Elk Park, Uinta Mountains,
5 (B. Y. U.). Uintah County: Trout Creek, SE Trout Peak, 22 mi. NW Vernal, 9,300 ft.,
5 (C. M.); Vernal-Manila Highway, 19 mi. N Vernal, 8,000 ft., 6 (C. M.); Taylor Peak, 17
mi. N Vernal, 4 (C. M.).
Thomomys talpoides ocius Merriam
Thomomys clusius ocius Merriam, Proc. Biol. Soc. Washington, 14:114,
July 19, 1901.
Thomomys clusius Allen, Bull. Amer. Mus. Nat. Hist., 13:246, No-
vember 25, 1896.
Thomomys ocius Bailey, N. Amer. Fauna, 39:107, November 15, 1915;
Barnes, Bull. Univ. Utah, 12 (No. 15) :83, April, 1922; Bull. Univ. Utah,
17 (No 12): 102, June, 1927.
18852
Type. — Male, adult, skin and skull, No. 25586 ■ U. S. National Museum
(Biological Surveys Collection) ; dry sagebrush mesas at Harveys Ranch,
Smiths Fork, 6 mi. SW Fort Bridger, 6,657 ft., Uinta County, Wyoming; May
24, 1890; collected by Vernon Bailey; original number 1194 (after Bailey,
type not seen).
Diagnosis. — Size small (see measurements). Color: Upper parts Tilleul
Buff overlaid with Avellaneous, grading over sides and flanks to nearly white
on underparts; underparts with faint wash of creamy white; postauricular
patches small and dusky and completely circling the ear; nose and cheeks
dusky; front feet, hind feet, throat, ventral surface of tail and distal half of
tail white. Skull: Small, slender but compact; nasals rounded posteriorly;
extension of premaxillae posterior to nasals very short; zygomatic arches ro-
bust, but not widely spreading, widest posteriorly; interparietal large and pen-
tagonal in shape; extension of supraoccipital posterior to lambdoidal suture
long; tympanic bullae actually as well as relatively large; basioccipital nar-
row; pterygoid hamulae long and ridged; upper incisors short and strongly
recurved.
Comparisons. — Compared with one topotype and seven near topo-
types of Thomomys talpoides pygmaeus, ocius differs as follows:
Size larger in every measurement taken. Color: Lighter through-
out, grayish as opposed to brown; distal half of tail white as op-
posed to only a few white hairs at tip of tail. Skull: Larger in
every measurement taken; skull more compact; zygomatic arches
2—2786
18 University of Kansas Publs., Mus. Nat. Hist.
heavier and more widely spreading posteriorly; tympanic bullae
larger; upper incisors larger, but equally strongly recurved; molar-
iform teeth larger.
Topotypes of ocius can be distinguished from those of Thomomys
talpoides uinta as follows: Color: Lighter throughout, grayish as
opposed to brown. Skull: Nasals rounded posteriorly as opposed
to emarginate; zygomatic arches more robust; interparietal penta-
gonal as opposed to subquadrangular; extension of supraoccipital
posterior to lambdoidal suture markedly greater; tympanic bullae
actually as well as relatively much larger; upper incisors short and
strongly recurved as opposed to long and procumbent.
Specimens of this subspecies can be distinguished from all other
members of the species Thomomys talpoides occurring in Utah by
their grayish color, and by small, compact skulls with very large
tympanic bullae and short strongly recurved upper incisors.
Remarks. — Two specimens from Vernal, Uintah County, are in-
tergrades between ocius and uinta. They resemble uinta in size and
dorsal color, but are slightly lighter tending toward the color of
ocius. Ventrally they are intermediate in color but more like ocius.
The skulls are more like those of ocius in general appearance,
extension of supraoccipital posterior to the lambdoidal suture, shape
and thickness of the zygomatic arches, posterior tongues of pre-
maxillae, size of tympanic bullae and recurved upper incisors. They
more closely resemble uinta in shape of posterior ends of nasals,
basioccipital and shape of the zygomatic processes of the squa-
mosals. In all of the above mentioned characters, they are inter-
mediate between the two named forms, but tend towards one or
the other as listed. The majority of characters are more as in ocius
to which they are here referred.
When Goldman (1939:233, 234) listed the named subspecies of
Thomomys talpoides, he hesitated to include ocius and merely men-
tioned that ocius, pygmaeus and idahoensis might also belong to
talpoides. Davis (1939:240, 241) found intergradation between
idahoensis and fuscus and also between idahoensis and pygmaeus,
and, therefore, arranged the last two mentioned forms as subspecies
of talpoides. This present study reveals intergradation between
ocius and uinta, and also between ocius and fossor (see account of
fossor). Therefore, ocius is properly to be treated as a subspecies
of the series of intergrading forms of which talpoides is the earliest
named.
All specimens of ocius known from Utah are from the extreme
Durrant — Pocket Gophers of Utah 19
eastern part of the northeastern corner of the state. The type lo-
cality of ocius is near Fort Bridger, Wyoming, which is north of
Utah. I have seen one specimen from 12 miles west of Linwood,
Daggett County, Utah, on Henrys Fork in Wyoming. Additional
collecting in northern Utah probably will reveal ocius to inhabit
also parts of northern Utah.
Specimens examined. — Total, 4, distributed as follows: Uintah County: Vernal, 2 (C. M.);
Uncompaligre Indian Reservation, 2 (A. M. N. H.).
Thomomys talpoides moorei Goldman
Thomomys jossor moorei Goldman, Journ. Washington Acad. S'ci.,
28:335, July 15, 1938.
Thomomys talpoides moorei Goldman, Journ. Mamm., 20:234, May
14, 1939.
Type.— Male, adult, skin and skull, No. 248222, U. S. National Museum
(Biological Surveys Collection) ; 1 mi. S Fairview, 6,000 ft., Sanpete County,
Utah; February 19, 1928; collected by A. W. Moore; X-catalogue number
24799 (after Goldman, type not seen).
Range. — Wasatch Plateau in Sanpete, Utah, Carbon and Emery counties,
and in Wasatch Mountains south of Spanish Fork Canyon.
Diagnosis. — Size medium (see measurements). Color: Upper parts be-
tween Cinnamon and Sayal Brown, with mixture of black hairs, grading
through Cinnamon on sides and flanks to Pale Pinkish Buff on underparts,
clearest on inguinal and pectoral regions; nose and cheeks dusky; postauricular
patches medium in size and black; ears black; chin buffy white; front and
hind feet white; tail mostly white with brownish hairs on dorsal surface.
Skull: Large, robust; nasals long and deeply emarginate on posterior ends,
and dilated distally; zygomatic arches robust and widely spreading; zygo-
matic processes of maxillae heavy; interparietal comparatively small, but
always wider than long; extension of premaxillae posterior to nasals short;
tympanic bullae moderate in size, but markedly inflated ventrally; pterygoid
hamulae long; interpterygoid space narrowly V-shaped; upper incisors long
and moderately recurved; molariform teeth light.
Comparisons. — Topotypes of moorei differ from topotypes and
near topotypes of Thomomys talpoides uinta as follows: Size
slightly larger. Color: Upper parts and sides lighter; tail lighter;
postauricular patches larger and darker; ears more pointed, smaller
and darker. Skull: Larger, heavier and more massive; nasals
longer, but deeply emarginate posteriorly as in uinta; rostrum
wider and longer; zygomatic arches heavier and more angular;
zygomatic processes of maxillae heavier; interparietal generally
smaller and shorter; braincase wider; tympanic bullae more inflated
ventrally; interpterygoid space more narrowly V-shaped; upper in-
cisors longer, but not as procumbent; molariform teeth smaller.
Topotypes of moorei can be distinguished from those of Tho-
20 University of Kansas Publs., Mrs. Nat. Hist.
momys talpoides oquirrhensis as follows: Size slightly larger; tail
longer; ears larger, less pointed. Color: Lighter throughout; post-
auricular patches larger. Skull: More ridged and angular; nasals
narrower posteriorly, but more dilated distally; posterior ends of
nasals more deeply emarginate (while shallowly emarginate in
oquirrhensis, they tend to be somewhat rounded) ; rostrum nar-
rower; extension of premaxillae posterior to nasals greater; least
interorbital breadth less ; zygomatic arches more angular and widely
spreading; zygomatic processes of maxillae heavier; interparietal
smaller; tympanic bullae larger and more inflated ventrally; upper
incisors generally longer.
The characters that distinguish moorei from Thomomys talpoides
parowanensis are: Color: Lighter throughout. Skull: Broader,
more angular and more nearly flat; zygomatic arches more widely
spreading; zygomatic processes of maxillae heavier; posterior ends
of nasals emarginate rather than rounded; upper incisors longer.
For comparisons of moorei with Thomomys talpoides levis and
ivasatchensis see accounts of these forms.
Remarks. — Specimens from Colton, show intergradation between
moorei, uinta and wasatchensis, but are referable to moorei in the
majority of characters. Specimens from Mount Nebo, and the
mouth of Reddicks Canyon, in the Wasatch and San Pitch moun-
tains, respectively, are intergrades between moorei and wasatchen-
sis, but are referable to moorei.
That part of the Wasatch Mountains south of Spanish Fork Can-
yon is inhabited by pocket gophers that are intergrades between
moorei and wasatchensis, but the cranial details show them to be
referable to moorei. The range here ascribed to moorei consists of
the Wasatch Plateau to the east of Sanpete Valley, the San Pitch
Mountains and the southern part of the Wasatch Mountains. The
type locality of moorei is situated in the southern end of a high
valley that separates the Wasatch Plateau from the San Pitch and
Wasatch mountains. Topotypical animals are larger and have
more ridged, angular skulls than those from the mountains.
Specimens examined. — Total, 48, distributed as follows: Utah County: Near Pay son
Lake, 1 (R. H.) ; Mt. Nebo, 25 mi. SE Payson, 10,000 ft., 20; Colton, 8 (B. Y. U.). San-
pete County: 1 mi. S Fairview, 6,000 ft., 12 (U. S. N. M.). Juab County: Mouth of Red-
dicks Canyon, Wales Mountain (= San Pitch Mountains), 7,500 ft., 5. Emery County:
Lake Creek, 11 mi. E Mt. Pleasant, 2 (C. M.).
Additional records. — Sanpete County: Ephraim, 5 (see Goldman, 1938:336).
Di rrant — Pocket Gophers of Utah 21
Thomomys talpoides f ossor Allen
Thomomys f ossor Allen, Bull. Amer. Mus. Nat. Hist., 5:51, April 28,
1893; Bailey, N. Amer. Fauna, 39:111, November 15, 1915; Bames, Bull.
Univ. Utah, 12 (No. 15) :85, April, 1922; Bull. Univ. Utah, 17 (No.
12):102, June, 1927; Hall, Univ. California Publ. Zool., 37:4, April 10,
1931.
Thomomys talpoides fossor Goldman, Journ. Mamm., 20:234, May 14,
1939.
Type.— Male, adult, skin and skull, No. |||^-. American Museum of Nat-
ural History; Florida, 7,200 ft., La Plata County, Colorado; June 25, 1892;
collected by Charles P. Rowley (after Allen, tj^pe not seen).
Range. — In the mountains of San Juan and Grand counties, east of the
Colorado and Green rivers.
Diagnosis. — Size medium (see measurements). Color: Upper parts Dresden
Brown, grading over sides to Pale Buff on underparts; chin white; ears small,
pointed, with deeply pigmented pinnae; postauricular patches grayish black;
nose dusky. Skull: Long and narrow; nasals long, rounded proximally and
usually simple distally; rostrum long; interparietal triangular; tympanic bullae
large, and well inflated ventrally; basioccipital narrow; palate narrow; palatal
pits shallow; dentition light.
Comparisons. — Near topotypes of fossor can be distinguished from
topotypes of Thomomys talpoides ocius as follows: Size larger
throughout. Color: Darker throughout, being dark brown as
opposed to grayish. Skull: Longer and narrower; nasals and ros-
trum longer; extension of supraoccipital posterior to lambdoidal
suture markedly less; tympanic bullae markedly smaller; upper
incisors longer and not as strongly recurved.
Among the races of Thomomys talpoides occurring in Utah, fossor
most closely resembles Thomomys talpoides uinta in color and size,
but differs from it as follows : Ears smaller, more pointed and with
more darkly pigmented pinnae. Skull: Longer, narrower and
weaker; rostrum longer; nasals longer, and rounded proximally as
opposed to markedly emarginate; interparietal triangular instead
of roughly pentagonal; tympanic bullae larger and more inflated
ventrally; basioccipital narrower; palate narrower, palatal pits
shallower; dentition lighter.
Remarks. — Bailey (1915:111) remarked that fossor was one form
that held its distinctive characters over a wide range. At that time,
its range was understood to include practically all of the moun-
tainous parts of Colorado, Utah as far west as the central part of
the state, and parts of New Mexico, Arizona and Wyoming. Sub-
sequently three new forms have been named from central Utah,
(Goldman 1938:334-337) thereby showing variation to be much
•J2 University of Kansas Publs., Mus. Nat. Hist.
more prevalent than formerly supposed. The range of jossor in
Utah, as now understood, is limited to the mountainous parts of the
state south and east of the Colorado and Green rivers in Grand
and San Juan counties.
The Utah specimens are not typical. At first glance some dif-
ferences are noted in the premaxillae and nasals. Four specimens
in the collections of the Museum of Natural History, University of
Kansas, three from 3 miles east of Creede, Mineral County, and one
from 10 miles east of Lake City, Hinsdale County, Colorado, both
of which lie north and east of the type locality of fossor show the
same characters as the Utah specimens.
Eight specimens from Oak Spring are intergrades between
fossor and ocius. In size and color they are like fossor, but the
skulls are intermediate. Because the animals are more like fossor
in the majority of characters, they are here referred to that race.
As a result of these studies and due to the paucity of specimens
from Utah, it is advisable, for the present, to refer all these Utah
animals to fossor. Additional specimens may reveal characters that
will merit the separation of the Utah animals from typical fossor;
a desertlike area unfavorable to Thomomys exists between the type
locality and eastern Utah.
Specimens examined. — Total, 21, distributed as follows: Grand County: Oak Spring,
Middle Fork Willow Creek, 15 Mi. N Thompson, 8 (C. M.) ; La Sal Mountains, 1 (U. S.
N. M.); Warner Ranger Station, La Sal Mountains, 3 (B. Y. U.). Son Juan County: Geyser
Pass, 18 mi. SE Moab, La Sal Mountains, 3 (1, B. Y. U. ; 2, C. M.); 5 mi. W Monticello,
1 (C. M.); Cooley Pass, 8 mi. W Monticello, 2 (C. M.); Joshua Flat, Elk Ridge, 8,300 ft.,
3 (M. V. Z.).
Thomomys talpoides parowanensis Goldman
Thomomys fossor parowanensis Goldman, Journ. Washington Acad.
Sci., 28:334, July 15, 1938.
Thomomys talpoides parowanensis Goldman, Journ. Mamm., 20:234,
May 14, 1939; Long, Journ. Mamm., 21:176, May 14, 1940.
Thomomys jossor Bailey, N. Amer. Fauna, 39:112, November 15,
1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922; Bull. Univ.
Utah, 17 (No. 12):102, June, 1927; Hall, Univ. California Publ. Zool.,
37:4, April 10, 1931; Presnall, Zion-Bryce Mus. Bull., 2:14, January,
1938; Tanner, Great Basin Nat., 1:111, 1940.
Type.— Male, adult, skin and skull, No. 158072, U. S. National Museum
(Biological Surveys Collection) ; Brian Head, Parowan Mountains, 11,000 ft.,
Iron County, Utah; September 8, 1908; collected by W. H. Osgood; original
number 3483 (after Goldman, type not seen).
Range. — High mountains of eastern Iron and Beaver counties, and western
Kane and Garfield counties.
Diagnosis. — Size medium (see measurements). Color: Upper parts Sayal
Brown moderately mixed with black, lightest on head; sides lightly washed
Durrant — Pocket Gophers of Utah 23
with Buff; underparts Pinkish Buff, clearest on inguinal and pectoral regions;
nose and cheeks dusky; postauricular patches large and black; front feet,
hind feet and distal half of tail white. Skull: Long and fairly slender; zygo-
matic arches not widely spreading; nasals long; rostrum long and slender;
posterior ends of nasals truncate or moderately emarginate; extension of pre-
maxillae posterior to nasals usually short; tympanic bullae relatively small;
upper incisors long and narrow; molariform teeth large.
Comparisons. — Compared with Thomomys talpoides kaibabensis,
parowanensis differs as follows: Size smaller. Skull: Shorter; na-
sals shorter; zygomatic breadth less; nasals truncate or shallowly
emarginate posteriorly as opposed to rounded; upper incisors nar-
rower.
Topotypes of parowanensis differ from topotypes and near topo-
types of Thomomys talpoides uinta as follows: Size larger. Color:
Usually lighter; postauricular patches larger and darker; ears small
with pinnae deeply pigmented as opposed to large and lightly pig-
mented. Skull: Larger; zygomatic arches more widely spreading;
nasals longer; rostrum longer; posterior ends of nasals truncate or
shallowly emarginate as opposed to deeply emarginate; sides of
zygomatic arches nearly parallel and not so divergent posteriorly;
interparietal larger and less quadrangular; extension of premaxillae
posterior to nasals less; upper incisors less procumbent; molariform
teeth larger.
Among named races of Thomomys talpoides, parowanensis most
closely resembles levis, the race nearest geographically to the east,
but differs from levis as follows: Size larger. Skull: Longer and
wider; rostrum and nasals longer; interparietal quadrangular as
opposed to roughly elliptical; upper incisors longer.
For comparisons with Thomomys talpoides moorei and wasatch-
ensis see accounts of those forms.
Remarks. — The mountains of south central Utah are inhabited by
pocket gophers that have been designated as Thomomys talpoides
parowanensis and T. t. levis by Goldman (1938:334, 336). They
are nearly indistinguishable in color and each is variable in cranial
details. The diagnostic characters of each form occasionally ap-
pear, in varying degrees, throughout the range of the other. The
Sevier River Valley separates the ranges ascribed to these two
forms. This valley is inhabited by pocket gophers that belong to
a different species, Thomomys bottae. The ranges of these two
races of talpoides converge southward at the headwaters of the
Sevier River. Specimens of parowanensis from the northern limits
of its range from the Beaver Mountains in eastern Beaver County
24 University of Kansas Publs., Mus. Nat. Hist.
and those of levis from the northern limits of its range in the Fish
Lake Mountains are readily distinguishable from each other. As
the ranges converge to the southward, there is progressively more
intergradation. The type locality of parowanensis is located in the
southern part of its range, while that of levis is in the extreme
northern part of its range. Therefore, due to the convergence of
the two ranges at the south, the specimens from localities near the
type locality of parowanensis show the greatest amount of inter-
gradation, if we regard specimens of parowanensis from the type
locality as typical of the race. Four specimens from Webster Flat,
sixteen miles east of Cedar City, Iron County, and three from Duck
Creek, Cedar Mountains, Kane County could equally well be as-
signed to either levis or parowanensis.
Specimens examined. — Total, 24, distributed as follows: Beaver County: Britts Meadows,
Beaver Mountains, 8,500 ft., 7 (3, M. V. Z. ; 2, U. S. N. M. ; 2, C. M.); Puffer Lake, Beaver
Mountains, 1 (U. S. N. M.); Kents Lake, Beaver Mountains, 1 (R. H.). Iron County: Lava
Beds, 3% mi. SW Panquitch Lake, 1 (C. M.); Brian Head, Parowan Mountains, 2 (1, U. S.
N. M. ; 1, C. M.); Webster Flat, 16 mi. E Cedar City, 4; Bear Valley, 2 mi. E B. V.
Ranger Station, 1 (R. H.). Garfield County: % mi. W Sunset Point, Bryce National Park,
8,000 ft., 1 (M. V. Z.). Kane County: Navajo Lake, 3 (R. H.); Duck Creek, Cedar Moun-
tains, 9,000 ft., 3 (1, R. H.).
Additional records. — Garfield County: Panquitch Lake, 1 (see Goldman 1938:335). Iron
County: Beaver Mountains, 9 (see Bailey, 1915:112); Buckskin Valley, 1 (see Goldman,
1938:335).
Thomomys talpoides levis Goldman
Thornomys fossor levis Goldman, Journ. Washington Acad. Sci., 28:
336. July 15, 1938.
Thomomys talpoides levis Goldman, Journ. Mamm., 20:234, May 14,
1939.
Thomomys fossor Bailey, N. Amer. Fauna, 39:112, November 15,
1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922; Bull. Univ.
Utah, 17 (No. 12):102, June, 1927.
Type .—Female, adult, skin and skull, No. 158079, U. S. National Museum
(Biological Surveys Collection) ; Seven Mile Flat, 5 mi. N Fish Lake, Fish
Lake Plateau, 10,000 ft., Sevier County, Utah; October 1, 1908; collected by
W. H. Osgood; original number 3616 (after Goldman, type not seen).
Range. — Fish Lake Mountains in Sevier County south into Garfield County,
Utah.
Diagnosis.— Size medium (see measurements). Color: Upper parts near
Sayal Brown, moderately mixed with black, darkest on head and middorsal
region, grading to Cinnamon Buff on sides and flanks; underparts Pinkish
Buff, clearest on inguinal and pectoral regions; chin, cheeks and nose dusky;
postauricular patches large and black; front feet, hind feet and distal half
of tail white; ears small and deeply pigmented. Skull: Slender and weak;
zygomatic arches not widely spreading; posterior ends of nasals rounded;
nasals moderately long and narrow; rostrum long and narrow; extension of
premaxillae posterior to nasals short; interparietal usually much wider than
Durrant — Pocket Gophers of Utah 25
long; pterygoid hamulae ridged; interpterygoid space usually narrowly V-
shaped; upper incisors short.
Comparisons. — Compared with topotypes of Thomomys talpoides
moorei, levis differs as follows: Size smaller; tail shorter. Color:
Darker throughout, especially on dorsal surface due to more black
of the underfur; underparts deeper huff. Skull: Narrower, less
massive ; zygomatic processes of maxillae weaker and not as widely
spreading; interparietal generally wider; extension of premaxillae
posterior to nasals less; posterior ends of nasals rounded rather
than emarginate; upper incisors shorter, less procumbent.
Topotypes of levis differ from near topotypes of Thomomys tal-
poides uinta as follows: Size larger. Color: Upper parts slightly
darker; postauricular patches much darker and larger; ears small
and deeply pigmented as opposed to large and lightly pigmented;
tail darker all around at base, with white part more extensive and
with fewer buff-colored hairs. Skull : More convex dorsally ; zygo-
matic arches more widely spreading and angular; nasals longer;
rostrum longer; interparietal wider and more elliptical; posterior
ends of nasals rounded as opposed to emarginate; extension of pre-
maxillae posterior to nasals less; pterygoid hamulae more ridged;
interpterygoid space more narrowly V-shaped; upper incisors
shorter and less procumbent.
Topotypes of levis can be distinguished from those of Thomomys
talpoides kaibabensis by markedly smaller measurements.
For comparisons with Thomomys talpoides parowanensis and
wasatchensis see acccounts of those forms.
Remarks. — Specimens from the Escalante Mountains and the
Aquarius Plateau are not typical. They are of approximately the
same color as levis, but are larger than lev is and have cranial details
that indicate intergradation with kaibabensis to the south. They
resemble kaibabensis in large size, long nasals and widely spreading
zygomatic arches, but are like levis in shape of the interparietal,
extension of premaxillae posterior to the nasals, rounded posterior
ends of nasals, ridged pterygoid hamulae and relatively short upper
incisors. Additional material from these regions may prove these
animals to merit separation and naming.
Specimens examiricd. — Total, 15, distributed as follows: Sevier County: Seven Mile
Flat, 5 mi. N Fish Lake, Fish Lake Plateau, 10,000 ft., 2 (U. S. N. M.); Fish Lake Experi-
ment Station, 2 (U. S. A. C). Garfield County: Posy Lake, Aquarius Plateau, 2 (B. Y. U.) ;
18 mi. N Escalante, 9,500 ft., 3; Steep Creek, Boulder-Teasdale Road, Boulder Mountain,
4 (B. Y. V.); Summit Birch Creek, Escalante Mountains, 2 (B. Y. U.).
26
University of Kansas Publs., Mus. Nat. Hist.
Measurements op Adult Males of Thomomys
(In millimeters)
H
H
f
73
f
ts
2
M
>
W
f
tri
o
"<
<t
sr
3
3
TO
c <!
t? 5.
a
a>
s. <a
ilar
gth
'a
sr
O
s
O
0
•a 2.
o 2
"9-
p
a.
a
3
o
=1
-
5'
p
"1 "1
O
?3
o
o
-
£
a.
3
s
3
s
[S
cr
o a
Cfl ™
"I
0
H
3
o
o
Sf
p-
p"
3"
n
fD O
? 3
■ p
c
3
2
g
ft
IS
!
Av 204
Min 194
Max 210
Av 209
Min 197
Max 216
Av 221
Min 204
Max 237
Av 199
Min 185
Max 208
Av 216
Min 203
Max 236
Av 215
Min 202
Max 228
Av 248
Min 244
Max 253
53
47
63
58
55
60
67
60
75
51
47
54
65
52
72
T. t. gracilis, 4; topotypes
28 31.5 13.4 21.7 18.3 6.4 7.6 1.3
27 30.3 12.9 21.1 17.8 6.3 7.3 1.0
28 33.5 14.2 22.0 19.0 6.5 7.9 1.7
T. t. oquirrhensis, 4; topotypes
28 32.2 13.9 21.9 19.0 6.9 7.6 0.9
28 31.9 13.7 21.4 18.5 6.7 7.2 0.6
29 32.8 14.3 22.8 19.5 7.1 7.9 1.0
T. t. wasatchensis, 10; topotypes
28 31.3 13.4 21.5 18.9 6.5 7.4 1.1
26 27.4 11.6 19.1 17.2 6.0 6.6 0.9
31 34.5 15.2 23.7 20.4 7.3 8.0 2.0
T. t. uinta, 5; SW slope Bald Peak, Uinta Mts.
27 31.5 13.1 21.7 19.4 6.3 7.6 1.1
26 29.6 12.1 20.3 19.0 5.7 7.3 0.7
28 32.8 13.8 22.2 20.0 6.5 7.8 1.4
T. I. moorei, 7; topotypes
29 32.4 13.9 22.9 19.2
27 31.3 13.0 21.5 18.4
31 34.7 14.5 23.7 20.0
15.4
14.7
16.4
15.8
15.5
16.2
15.1
14.0
16.5
15.2
13.5
15.6
T. t. fossor, 8; Cascade Creek, La Plata Co., Colo.
61 29 31.7 13.2 21.2 18.7
54 27 30.5 12.0 20.5 18.2
70 30 33.0 14.4 23.5 19.9
T. t. ravus, 3; topotypes
73 30 35.2 14.6 24.8 21.4
70 29 34.5 14.3 23.6 20.5
74 30 35.9 15.1 25.7 22.5
T. t. pygmaeus, 1 ; topotype
24.6 10.2 16.3 15.1 5.4
No. 55270 (U.S.N.M.)
165 40 20 24.6 10.2 16.3 15.1 5.4 5.9 0.7
No. 177506 (U.S.N.M.) T. t. ocius, 1; 12 mi. W Linwood, Henrys Fork, Wyo.
Av....
.. 215
59
28
34.3
Min. . .
.. 202
48
27
34.1
Max. . .
.. 228
69
29
34.6
7.2
6.7
7.5
7.7
7.5
7.9
7.4
6.7
8.2
7.4
7.2
7.6
6.5
6.0
7.0
7.7
7.3
8.2
1.5
0.9
2.0
15.9
14.8
16.3
7.3
6.7
7.7
Co.,
Colo.
5.9
5.5
6.3
7.5
7.0
7.9
0.6
0.0
1.1
15.5
14.5
16.9
7.1
6.9
7.4
6.3
6.0
6.7
8.3
8.2
8.4
2.4
2.2
2.7
17.1
16.7
17.5
8.2
8.1
8.5
12.0 5.7
200 62 26 27.5 11.5 19.9 17.8 6.2 6.8 1.0 13.5 7.0
T. t. parowanensis, 2; Britts Meadow, Beaver Mountains
14.5 22.4 18.6 6.0 8.1 1.4 17.3 7.9
14.1 22.0 18.4 5.8 8.0 1.0 17.2 7.6
14.8 22.7 18.9 6.2 8.2 1.7 17.3 8.2
Durrant — Pocket Gophers of Utah
27
Measurements op Adult Females of Thomomys
(In millimeters)
9
TO
r
■r.
a
3
V
TO
§
p
c+-
>
TO
M
TO
O
2
"1
>Q (B
—
p
=r
3
o
■5."
0
a-
•o S.
2 »
9
TO
B
P.
-
P
a
a.
B aT
O 3
£$-
^ sr
$ o
W
•a Si
2 3
e-f- w
?§
32,
en T3
p 1
It
9
TO
M
g
(0
P
—
— o
? B
f
2
3
1.2
1.1
1.4
14.0
14.0
14.0
6.5
6.4
6.6
0.8
0.5
1.0
14.8
14.2
15.5
7.2
6.9
7.5
0.9
0.6
1.2
14.6
13.0
16.2
7.2
6.8
7.5
1.3
1.1
1.5
13.5
13.3
13.6
6.8
6.8
6.8
1.3
1.0
1.6
14.6
14.0
15.6
6.8
6.4
7.0
0.7
0.5
1.0
16.2
15.9
16.3
7.3
7.0
7.5
T. t. gracilis, 2; topotypes
Av 190 58 27 29.7 12.0 19.7 17.3 6.4 7.3
Min 185 54 27 29.5 11.9 19.7 16.9 6.3 7.2
Max 194 61 27 29.9 12.0 19.7 17.6 6.5 7.4
T. t. oquirrhensis, 7; topotypes
Av 203 56 27 30.2 12.9 20.4 18.2 6.8 7.5
Min 193 52 25 28.5 12.2 19.5 17.5 6.6 6.7
Max 215 59 28 31.5 13.3 21.0 19.1 7.2 8.0
T. t. wasatchensis, 19; topotypes
Av 205 62 27 31.5 12.7 20.5 18.0 6.5 7.4
Min 180 52 23 28.1 11.2 19.3 17.2 6.2 6.0
Max 222 70 30 32.5 14.5 22.0 19.9 6.7 8.1
T. t. uinta, 2; SW slope Bald Peak, Uinta Mts.
Av 181 49 25 28.4 11.6 19.8 17.3 6.6 7.2
Min 177 47 25 28.3 11.6 19.8 17.2 6.4 7.0
Max 185 50 25 28.4 11.6 19.8 17.4 6.7 7.3
T. t. moorei, 5; topotypes
Av 206 62 26 29.9 12.8 21.5 18.4 6.6 7.3
Min 198 55 24 29.0 12.3 21.0 18.0 6.4 7.0
Max 213 69 28 31.2 14.1 22.5 19.1 6.8 7.5
T. t. fossor, 4; Cascade Creek, La Plata Co., Colo.
Av 215 57 29 32.6 14.2 22.0 19.0 6.0 7.5
Min 204 51 28 31.3 13.6 21.5 18.0 5.7 7.1
Max 223 63 30 34.0 14.8 22.9 19.6 6.3 7.8
No. 13684 (CM.) T. t. ravus, 1; topotype
241 71 28 35.7 14.5 24.4 21.5 6.2 7.8
No. 178868 (U.S.N.M.) T. t. pygmaeus, 1 ; Fossil, Wyo.
167 52 20 24.0 10.2 16.5 14.8 5.2 5.6
T. t. ocius, 3; topotypes
Av 201 60 25 30.0 13.5 20.5 17.9 6.2 7.2
Min 196 57 25 29.9 13.0 19.9 17.5 6.1 7.1
Max 205 64 25 30.1 14.0 21.5 18.6 6.3 7.3
T. t. paroioanensis, 4; Britts Meadow, Beaver Mountains
Av 221 58 29 33.2 14.5 22.8 19.0 6.0 7.8
Min 207 50 28 30.5 12.8 22.7 18.6 5.8 7.4
Max 240 66 30 34.8 15.5 23.0 19.6 6.2 8.1
T. t. levis, 2; topotypes
Av 203 65 27 28.1 11.1 19.2 17.7 6.1 6.9
Min 199 61 26 28.0 10.6 18.9 17.5 5.8 6.6
Max 206 70 27 28.2 11.6 19.5 17.9 6.4 7.2
2.7 17.1 8.1
0.7 11.1 5.8
0.8
0.5
1.0
0.9
0.5
1.5
0.8
0.6
1.0
15.0
14.7
15.3
15.4
14.7
17.8
13.0
12.8
13.2
7.4
7.3
7.5
7.3
7.0
7.7
6.8
6.6
7.0
28 University of Kansas Publs., Mtjs. Nat. Hist.
Thomomys bottae (Eydoux and Gervais)
Thomomys bottae is a southern species that, within the Great
Basin, reaches the most northern limits of its distribution in Utah.
The animals of this species inhabit the lower valleys, and with the
exception of the Oquirrh Mountains, inhabit also the mountains in
that part of the state west of the central mountain ranges. The
specific characters are: Sphenorbital fissure present; incisive fora-
mina posterior to infraorbital canal; anterior prism of P4 rounded;
interparietal relatively small; lambdoidal suture straight in region
of interparietal, in Utah specimens.
Thomomys bottae aureiventris Hall
Thomomys perpallidus aureiventris Hall, Univ. California Publ. Zool..
32:444, July 8, 1930; Univ. California Publ. Zool., 37:3, April 10, 1931.
Thomomys bottae aureiventris Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935.
Type. — Male, adult, skin and skull, No. 43980, Museum of Vertebrate Zo-
ology, University of California; Fehlman Ranch, 3 mi. N Kelton, 4,225 ft.,
Box Elder County, Utah; September 27, 1929; collected by Louise Kellogg;
original number 451.
Range. — Northwestern Utah, and extreme western Utah as far south as
the southern end of the Deep Creek Mountains.
Diagnosis. — Size medium (see measurements) ; claws on front feet small.
Color: Near Cinnamon on dorsal and ventral surfaces; inguinal region, front
and hind feet and distal third to half of tail white; nose, cheeks and post-
auricular patches grayish black. Skull: Moderately angular and ridged; zyg-
omatic arches nearly parallel with sides of skull; jugals vertical; marked
thickening at union of jugal and zygomatic process of maxilla; greatest zygo-
matic breadth at anterior part of arches; interpterygoid space lyre-shaped;
ventral margin of jugal concave dorsally; nasals long and denticulate distally;
parietal ridges bowed in at two places, at coronal suture and at middle of
interparietal; paroccipital processes extremely well developed; dorsal fronto-
maxillary suture usually straight.
Comparisons. — From near topotypes of Thomomys bottae cen-
tralis, aureiventris differs as follows: Size larger; tail shorter; hind
foot longer; claws on front feet shorter. Color: Slightly darker
on upper parts, but with greater extension of white on ventral sur-
face. Skull: Zygomatic breadth greater; greatest width across
zygomatic arches at anterior rather than posterior region; zygo-
matic arches thicker at union of jugals and zygomatic processes of
maxillae; dorsal frontornaxillary suture less convex medially; mas-
toid breadth greater; extension of premaxillae posterior to nasals
less; interpterygoid space lyre-shaped rather than V-shaped.
From topotypes of Thomomys bottae albicaudatus, aureiventris
Durrant — Pocket Gophers of Utah 29
can be distinguished by: Size larger; hind foot longer. Color: Mark-
edly lighter throughout, Cinnamon as opposed to near (13 " " n)
Black. Skull: Larger in all but three measurements taken; exten-
sion of premaxillae posterior to nasals less ; alveolar length of upper
molar series shorter; zygomatic arches widest anteriorly rather than
posteriorly; thickening at union of jugal and zygomatic process of
maxilla markedly greater; interpterygoid space lyre-shaped as op-
posed to V-shaped; lacrimal processes more globose at tips.
Thomomys bottae aureiventris can be readily distinguished from
T. b. bonnevillei, sevieri, wahwahensis, and convexus by larger size
in all measurements taken and darker coloration. The same dif-
ferences obtain in comparison with T. b. tivius and stansburyi ex-
cept that aureiventris is much lighter colored. See comparisons
under those forms.
Remarks. — T. b. aureiventris has one of the most extensive ranges
of any race of T. bottae occurring in Utah. The range extends from
the valleys of the northwest corner of the state south along the
extreme western margin of the state approximately to the southern
end of the Deep Creek Mountains. This ascribed range practically
bounds the northwest and western margins of the great salt desert
in Box Elder and Tooele counties. As far as known, this great
waste area harbors no members of the Geomyidae. Pocket gophers
were available from four localities in addition to the type locality.
In these four localities all of the animals were intergrades. The
three specimens from Queen of Sheba Canyon, Deep Creek Moun-
tains, although smaller than aureiventris in every measurement
taken, resemble it in color and general configuration of the skull.
The animals from Trout Creek and Ibapah at the southern end of
the range, although referred to aureiventris, are intermediate between
it and centralis. In color and measurements they more closely re-
semble centralis, but the skulls closely resemble those of aureiventris.
The skulls show some slight characteristics of bonnevillei, the form
to the east, which indicate an early relationship between the two.
Specimens from the east side of Tecoma Range, adjacent to Pilot
Peak, although referred to aureiventris are intergrades between it
and centralis. Although this locality is nearer the type locality of
aureiventris than any of the other record stations, the animals show
the maximum departure from topotypes in morphological features.
In color they approach centralis, and agree with it in one-half of
the measured characters. The general configuration of the skull
and a majority of the critical diagnostic characters, for example,
30 University of Kansas Publs., Mus. Nat. Hist.
jugal thickening, are more nearly as in aureiventris. From the
above remarks it is readily understood that this subspecies is ex-
tremely variable.
Specimens examined. — Total, 55, distributed as follows: Box Elder County: Fehlman
Ranch, 3 mi. N Kelton, 4,255 ft., 8 (7, M. V. Z.); Utah-Nevada Boundary, E Side Tecoma
Range, 4,300 ft., 12. Tooele County: Ibapah, 5,000 ft., 21. Juab County: Queen of Sheba
Canyon, W side Deep Creek Mountains, 5,600 ft., 11.
Thomomys bottae robustus new subspecies
Type.— Male, adult, skin and skull, No. 2726, Museum of Zoology, Uni-
versity of Utah; Orr's Ranch, Skull Valley, 4,300 ft., Tooele County, Utah;
June 19, 1938; collected by S. D. Durrant; original number 1583.
Range. — Skull Valley, Tooele County, Utah.
Diagnosis. — Size medium (see measurements); tail short; hind foot short.
Color: In a series of 24 animals, upper parts vary from Pale Smoke Gray (4
specimens) through Cinnamon Buff (19 specimens) to Dark Mouse Gray (1
specimen). The Cinnamon Buff color is considered to be typical. Color
grading to lighter on underparts; postauricular patches small and grayish
black; front and hind feet and distal part of tail white. Skull: Small, flat
and heavily ridged; nasals short; zygomatic arches heavy and widely spread-
ing, widest posteriorly at union of jugal and squamosal; union of jugal and
zygomatic process of maxilla thickened, with a ventrally directed spinous
process in sixty percent of the specimens; occasionally there is a second
process, also directed ventrally at union of jugal and zygomatic process of
squamosal; zygomatic arches convex dorsally; deep dorsal depression present
in frontal bones in mature specimens; lacrimal processes prominent, project-
ing well above the arch at the anteromedial angle of the orbit ; interpterygoid
spaces V-shaped; tympanic bullae well inflated ventrally; upper incisors
short, and pale; when placed on a flat plane the dorsal surface of the skull
is nearly parallel to the substratum; space enclosed within the zygomatic
arches nearly quadrangular.
Comparisons. — From topotypes of Thomomys bottae aureiventris,
robushis can be distiguished as follows: Size smaller; tail and hind
foot shorter. Color: Lighter throughout. Skull: Smaller, more
heavily ridged and more nearly flat; nasals shorter; rostrum rela-
tively wider and shorter; zygomatic arches shorter and relatively
more widely spreading with greatest width posteriorly as opposed
to anteriorly; junction of jugal and zygomatic process of maxilla
not as prominent; aureiventris shows no spinous process at this
junction; lacrimal processes larger and projecting farther dorsally;
enclosed space within zygomatic arches roughly quadrangular as
opposed to triangular; mastoidal part of tympanic bullae less ex-
posed; sphenorbital fissure smaller; interpterygoid space V-shaped
rather than lyre-shaped; palatal pits smaller and shallower; tym-
panic bullae smaller, but more inflated ventrally; basioccipital
Durrant — Pocket Gophers of Utah 31
averaging relatively wider; molars smaller; upper incisors shorter,
smaller and cadmium yellow as opposed to orange yellow.
Comparisons of robustus with topotypes of Thomomys bottae al-
bicaudatus show the following: Size smaller. Color: Lighter
throughout; postauricular patches smaller and lighter. Skull:
Smaller, more compact and more nearly flat; rostrum shorter and
more nearly straight; lacrimal processes larger, projecting higher
above the anteromedial angle of the orbit; parietal ridges uniformly
heavier; mastoid width actually as well as relatively wider; zygo-
matic arches heavier and relatively much wider (males 76.2 per-
cent of basilar length, females 73.8 percent as opposed to males
73.8 percent and females 73.5 percent) ; union of jugal and zygo-
matic process of maxilla uniformly more thickened; spinous process
at jugal-maxillary suture present; zygomatic arches much more
concave on ventral surface; uniform deep depression present in
mature adults, between frontal processes of premaxillae, and an-
terior interorbital region of frontals; extension of premaxillae
posterior to nasals less; sphenorbital fissure more constricted; tym-
panic bullae more inflated ventrally, extending well ventrad of
basioceipital; palatal pits shallower and smaller; molars smaller;
upper incisors shorter, narrower and paler (see comparison of aure-
iventris).
From near topotypes of Thomomys bottae centralis from 1 mile
east of Garrison, Millard County, Utah, robustus differs in: Size
smaller; tail and hind foot shorter. Color: Lighter, terminal bands
of hair cinnamon, but because more black in underfur the animals
appear darker; postauricular patches smaller and lighter. Skull:
Shorter, more nearly flat and much more heavily ridged; nasals
shorter; rostrum shorter and wider; lacrimal processes larger and
projecting higher above anteromedial angle of orbit; zygomatic
arches heavier, shorter, more angular and actually as well as rela-
tively wider; jugals thicker; angle between maxillary plate and
rostrum less obtuse; spinous process at jugal-maxillary suture pres-
ent; extension of premaxillae posterior to nasals less; parietal ridges
much more pronounced; looked at from above, space enclosed
within zygomatic arches more quadrangular in shape as opposed to
roughly triangular; tympanic bullae more inflated ventrally; molars
smaller; upper incisors shorter, narrower and paler.
The characters that distinguish robustus from topotypes of Tho-
momys bottae ivuhwahensis are: Size slightly smaller. Color:
Darker throughout. Skull: Rostrum longer and narrower; nasals
32 University of Kansas Publs., Mus. Nat. Hist.
longer; zygomatic arches wider and longer; lacrimal processes
larger and projecting higher above anteromedial angle of the orbit;
parietal ridges more roughened; tympanic bullae much larger and
more inflated ventrally; supraoccipital higher; middorsal depression
in frontals present. For comparisons with Thomomys bottae bon-
nevillei see account of that form.
The remaining forms from the Bonneville Basin, namely, Tho-
momys bottae sevieri, convexus, twins and stansburyi are all easily
distinguished from robustvs. Specimens of sevieri are paler, smaller
in every measurement taken, and the skulls are weaker and less
angular. All specimens of convexus are paler, the skulls are more
convex dorsally and narrower, with less ridging and angularity.
Both tivius and stansburyi are small dark forms, with weak, smooth,
small .skulls as compared with robustus which is light colored and
has compact, ridged and angular skulls.
Remarks. — Twenty-three specimens were obtained at a small
isolated spring. Critical study of animals taken only a few miles
to the east prove them to be so different as to be referable to an-
other subspecies, albicaudatus. T. b. robustus is an endemic form
in this desert valley. The variable color is noteworthy but difficult
to explain in an isolated population as small as this one. All five
of the gray animals are females of which four are lactating adults.
The affinities of this subspecies are with albicaudatus to the east,
but enough time has elapsed since isolation to enable them to dif-
ferentiate.
Specimens examined. — Total, 23, from the type locality.
Thomomys bottae minimus Durrant
Thomomys bottae minimus Durrant, Proc. Biol. Soc. Washington,
52:161, October 11, 1939; Marshall, Joum. Mamm, 21:154, May 14, 1940.
Type. — Male, adult, skin and skull, No. 263942, U. S. National Museum
(Biological Surveys Collection) ; Stansbury Island, Great Salt Lake, Tooele
County, Utah; June 25, 1938; collected by William H. Marshall; original
number 141.
Range. — Known only from the type locality.
Diagnosis. — Size small (see measurements) ; tail relatively long. Color :
Upper parts Pinkish Buff, darker on head; underparts Pale Pinkish Buff;
front and hind feet white; nose, chin and postauricular patches black. Skull:
Long, slender and nearly devoid of ridges; braincase moderately inflated; in-
terparietal quadrangular; zygomatic arches weak, widest in temporal region,
but neither widely spreading nor angular; nasals straight and truncate pos-
teriorly; extension of premaxillae posterior to nasals relatively great; tym-
panic bullae moderately inflated; palatal pits deep; rostrum short but narrow;
Durrant — Pocket Gophers of Utah 33
interpterygoid space moderately lyre-shaped; upper incisors narrow; molars
light.
Comparisons. — Compared with topotypes of Thomomys bottae
albicaudatus, minimus differs as follows: Size markedly smaller;
claws on front feet shorter and weaker. Color: Markedly lighter
throughout, being Pinkish Buff as contrasted with near (13 " " n)
Black. Skull: Smaller in every measurement taken; slender,
smooth, weak and nonangular as opposed to ridged, robust, wide
and angular; zygomatic arches much weaker and not so widely
spreading posteriorly ; ascending processes of premaxillae much nar-
rower; extension of premaxillae posterior to nasals less; interptery-
goid space moderately lyre-shaped as opposed to V-shaped; denti-
tion lighter.
Topotypes of minimus differ from those of Thomomys bottae
aureiventris as follows: Size markedly smaller. Color: Lighter
dorsally and no "gold color" on underparts. Skull: Markedly
smaller in every measurement taken; weak, smooth and slen-
der as opposed to ridged, angular and robust; zygomatic arches
weak and widest posteriorly rather than heavy and widest anteri-
orly; no great thickening at region of union of jugal and zygomatic
process of the maxilla; jugals more nearly straight rather than
concave laterally; interpterygoid space not so markedly lyre-
shaped; dentition lighter.
The races nearest geographically to minimus are Thomomys
bottae nesophilus and T. b. stansburyi. For comparisons see ac-
counts of those forms.
Remarks. — This subspecies is the smallest of all the races of
Thomomys bottae occurring in Utah. As far as known it is en-
demic to Stansbury Island, and since the Pleistocene Lake Bonne-
ville attained its highest level has remained on that part of Stans-
bury Island that was above this high level. (See comments under
nesophilus.) The sandy nature of the soil and the desert condi-
tions of the area that has since been exposed at lower levels ap-
parently do not constitute a favorable environment. Unlike neso-
philus from Antelope Island, this form does not have its affinities
with albicaudatus, the valley form of the adjacent mainland, but
does show affinities with stansburyi, the nearest mountain form on
the mainland. This is easily understood when one realizes that
Stansbury Island is only an isolated part of Stansbury Mountain
that projects northward as a peninsula into Great Salt Lake. The
3—2786
34 University of Kansas Publs., Mus. Nat. Hist.
history of Stansbury Island with reference to isolation of minimus
parallels that of nesophilus on Antelope Island. See discussion
under nesophilus.
Specimens examined. — Total, !>, as follows: Tooele County: Stansbury Island, Great Salt
Lake, 5 (U. S. N. M.).
Thomomys bottae nesophilus Durrant
Thomomys bottae nesophilus Durrant, Bull. Univ. Utah, 27 (No. 2) :2,
October, 1936; Marshall, Journ. Mamm, 21:156, May 14, 1940.
Type. — Male, adult, skin and skull. No. 1136, Museum of Zoology, Uni-
versity of Utah; Antelope Island, Great Salt Lake, Davis County, Utah; April
20, 1935; collected by S. D. Durrant; original number 761.
Range. — Known only from the type locality.
Diagnosis. — Size medium (see measurements) ; claws on front feet long.
Color: Upper parts Cinnamon Buff; lighter below; sides Pinkish Buff inter-
spersed with gray; pectoral and inguinal regions Cinnamon; nose grayish
black; postauricular patches black. Skull: Interparietal wedge-shaped;
tympanic bullae small; dorsal surface of lambdoidal prominence 3 mm. wide
rather than developed as a crest; jugals nearly straight; zygomatic arches
strongly rectangular.
Comparisons. — Compared with topotypes of Thomomys bottae
albicaudatus, nesophilus is of approximately the same size, but
differs as follows: Claws on front feet longer. Color: Lighter
throughout; tail white terminally, but much darker at base; post-
auricular patches smaller. Skull: Interparietal wedge-shaped as
opposed to roughly quadrangular; lambdoidal eminence more of a
crest than a ridge; tympanic bullae smaller; jugals more nearly
straight; zygomatic arches more nearly rectangular.
From topotypes of Thomomys bottae aureiventris, nesophilus
differs in: Size smaller; claws on front feet longer. Color: Darker
throughout; postauricular patches larger. Skull: Heavier, more
massive; zygomatic arches more robust and convex laterally rather
than concave; interparietal wedge-shaped rather than roughly
quadrangular; braincase more nearly flat; tympanic bullae mark-
edly smaller; upper molariform series longer; molariform teeth
wider and heavier; interpterygoid space V-shaped rather than lyre-
shaped.
The race nearest geographically to nesophilus is T. b. minimus
from Stansbury Island, Great Salt Lake. It can easily be distin-
guished from minimus by the following features: Size much larger;
claws on front feet longer and thicker. Color: Darker throughout;
postauricular patches larger and with more admixture of buff col-
Dirrant — Pocket Gophers of Utah 35
ored hairs. Skull: Larger in every measurement taken; wide and
robust as opposed to narrow and slender; zygomatic arches more
widely spreading and angular; braincase more nearly flat; tympanic
bullae actually larger, but relatively smaller; lambdoidal eminence
flat-topped rather than a crest; interparietal wedge-shaped as op-
posed to quadrangular; teeth larger.
Remarks. — The affinities of nesophilus of Antelope Island are
unquestionably with albicaudatus of the eastern and southern
mainland. At the time of this writing (1945), Antelope Island is
not truly an island, but only the tip of a broad peninsula projecting
westward into Great Salt Lake. Nevertheless, the area of occur-
rence of nesophilus is effectively isolated by the exposed, sandy
lake bottom that is unsuited to occupancy by pocket gophers.
Fluctuations in the level of the Great Salt Lake have broken and
reestablished this connection with the mainland many times. Each
of the several other kinds of mammals which are known from both
the island and the mainland show no differentiation on the island.
These are kinds (see Marshall, 1940:156), which more freely cross
the exposed, sandy lake bottom. I, myself, have noted tracks of
coyotes going to and from the island. The pocket gopher, neso-
philus, so far as known is the only mammal which has developed a
subspecies endemic to the island. The beach levels of Pleistocene
Lake Bonneville are well marked on both Antelope Island and
Stansbury Island, which is fifteen miles west of Antelope Island.
On the eastern side of Antelope Island the lower beach levels of
this prehistoric lake are farmed. Although sought for elsewhere
on this island, pocket gophers were found only in the farmed
land. On Stansbury Island there has been no farming, and the
endemic pocket gophers, minimus, although sought for elsewhere on
that island were found only above the highest beach levels of the
ancient lake. Evidently these pocket gophers still occupy only that
part of Stansbury Island that projected above water during the
greatest height of Lake Bonneville. Farming on Antelope Island
may have developed a more favorable environment for pocket
gophers, thus causing them to move down to the lower levels from
that part of the island that was above water during Pleistocene
times.
Specimens rxamined. — Total, 5, from the type locality.
36 University of Kansas Publs., Mus. Nat. Hist.
Thomomys bottae stansburyi new subspecies
Type. — Female, adult, skin and skull, No. 2045, Museum of Zoology, Uni-
versity of Utah; South Willow Creek, Stansbuiy Mountains, 7,500 ft., Tooele
County, Utah; July 2, 1937; collected by O. S. Walsh and S. D. Durrant;
original number 1257 of Durrant.
Range. — Stansbury Mountains, Tooele County, Utah.
Diagnosis. — Size small (see measurements). Color: Upper parts Saccardo's
Umber, darker on head; sides and underparts Pinkish Buff; nose, chin and
postauricular patches black; front and hind feet and distal third to half of
tail white. Skull: Small, slender, weak and smooth; zygomatic arches light
and not widely spreading; zygomatic arches actually as well as relatively
short; interparietal generally quadrangular; nasals relatively long and slender;
interpterygoid space narrowly V-shaped; basioccipital fairly wide; tympanic
bullae moderately inflated ventrally; dentition light.
Comparisons. — Topotypical specimens of stansburyi can be read-
ily distinguished from those of Thomomys bottae centralis, aurei-
ventris and albicaudatus by being smaller in every measurement
taken, particularly those of the skull; the skull is weaker and
smoother. In color stansburyi is like albicaudatus but is much
darker throughout than aureiventris and centralis.
Comparisons of topotypes of stansburyi with those of Thomomys
bottae sevieri show them to be of approximately the same size, but
to differ as follows: Color: Darker throughout. Skull: Zygomatic
arches shorter; tympanic bullae less inflated ventrally; zygomatic
breadth less; mastoid breadth greater; width across alveolar pro-
cesses of maxillae greater; alveolar length of upper molar series
greater; molariform teeth larger.
Compared with topotypes of Thomomys bottae minimus, stans-
buryi is seen to be of larger size and darker color throughout, with
a skull that is larger in most every measurement taken, although
of the same slender, smooth, nonangular type.
Among named races of Thomomys bottae, stansburyi most closely
resembles tivius, a small, dark, mountain form from central Utah.
Size and color are almost the same but stansburyi differs in: Tail
shorter; hind foot averaging slightly longer. Skull: Generally
larger in every measurement taken; zygomatic arches shorter;
width across alveolar processes of maxillae greater; zygomatic
arches more widely spreading, and widest in extreme posterior re-
gion rather than in region of jugal-squamosal suture.
Remarks. — The Stansbury Mountains are separated from the
Oquirrh Mountains by the Stockton Bar, and from the Onaqui
Mountains, which are in reality a continuation of the Stansbury
Durrant — Pocket Gophers of Utah 37
Mountains, by only a low pass. Pocket gophers from Clover Creek,
Onaqui Mountains and Little Valley, Sheeprock Mountains, al-
though intergrades between robustus and albicaudatus are dark in
color like stansburyi. These intergrades are large, dark colored,
and have heavy, ridged, angular skulls. It appears that stans-
buryi is a mountain subspecies derived from albicaudatus of the
valley. It would be instructive to artificially transplant gophers
from mountains to valleys, and vice versa, so as to reveal what ef-
fects if any on the animals' morphology the environment might
have in one or a few generations. Gophers are well known to be
very plastic, and such an experiment as suggested might call for
modification of the view, held here, that the differential features of
gophers from South Willow Creek and, say, Bauer, are hereditary.
Specimens examined. — Total, 11, from the type locality.
Thomomys bottae albicaudatus Hall
Thomomys perpallidus albicaudatus Hall, Univ. California Publ. Zool.,
32:444, July 8, 1930; Univ. California Publ. Zool., 37:3, April 10, 1931.
Thomomys bottae albicaudatus Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935; Durrant. Bull. Univ. Utah, 28 (No. 4) :5, August
18, 1937.
Thomomys -perpallidus aureiventris Hall, Univ. California Publ. Zool.,
37:3, April 10, 1931.
Type. — Male, adult, skin and skull, No. 43971, Museum of Vertebrate
Zoology, University of California; Provo, 4,510 ft., Utah County, Utah; Octo-
ber 17, 1929; collected by Annie M. Alexander; original number 506.
Range. — From the area between the Great Salt Lake and the Wasatch
Mountains south along the western margin of the central mountains of the
state to the Sevier River, in Juab County, west into Tooele County to the
Onaqui and Sheeprock mountains.
Diagnosis. — Size medium (see measurements) ; claws on front feet medium.
Color: Upper parts near (13""w) Black, grading over sides and flanks to
Pinkish Cinnamon on underparts; chin, nose, top of head and postauricular
patches black; front feet, hind feet and distal third to half of tail white.
Skull: Angular and ridged; zygomatic arches moderately wide spreading,
widest posteriorly; paroccipital processes weak; zygomatic processes of
maxillae convex anteriorly; lacrimal processes small and peglike; jugals con-
vex dorsally on ventral surface; nasals short, rounded distally and truncate
proximally; parietal crests bowed in, in two places; interpterygoid space
broadly V-shaped.
Comparisons. — For comparisons of albicaudatus with Thomomys
bottae aureiventris and centralis see accounts of those forms.
Topotypes of albicaudatus are dark colored and can be distin-
guished from those of Thomomys bottae birdseyei, tivius, stans-
buryi and contractus which are also dark forms, by larger size and
larger, more robust skulls (see accounts of those forms). It can be
38 University of Kansas Publs., Mus. Nat. Hist.
distinguished from the remainder of the known subspecies of Thom-
oiyiys bottae in Utah by darker color and by cranial details (see ac-
counts of those forms).
Remarks. — The range of albicaudatus is larger than that of any
other race of Thomoinys bottae limited to Utah. Specimens are
available from thirty localities which represent widely varied habi-
tats and environments. This subspecies consists of many highly
variable local populations, and the marginal populations intergrade
freely with adjacent races. In many populations, it is really difficult
to recognize the relationships on account of the great variation, and
one is frequently tempted to name some of them as distinct. Care-
ful study of the large number of specimens has enabled me to recog-
nize diagnostic characters common to all of these variable popula-
tions. The animals range from large and dark at the north to small
and light at the south.
The Jordan River bisects Salt Lake County from north to south.
Pocket gophers were taken at nine places east of the river, and at
three places west of it.
Gophers from Salt Lake City and environs (east of the river)
vaiy in color from almost black to dark cinnamon. Specimens from
Draper, which locality is likewise east of the river, are uniformly
lighter, but also vary in color. The skulls of animals from both
localities are indistinguishable from each other and closely resemble
those of topotypes. Specimens from the west side of the river,
from Riverton, two miles west of Murray and Rose Canyon,
Oquirrh Mountains, all are lighter in color than topotypes. The
color varies from darkest at the north at Murray to lightest at the
south at Riverton. This is exactly the reverse of what would be
expected since Riverton is the locality geographically nearest to the
type locality, Provo. The skulls are quite uniform and are all re-
ferable to albicaudatus. The Jordan River may be one factor which
causes this lack of uniformity between the animals from the two
sides of the river. Davis (1939:56-57) states that rivers are not
barriers to movement of pocket gophers where the river completely
freezes over and has the ice covered with thick snow. Although the
Jordan River does occasionally freeze over, it is never frozen for
more than a few days at a time, and snow in this area does not last
for long periods. The material at hand indicates that the gophers
from both sides of the river are referable to the same subspecies
albicaudatus. The animals from the east side of the river are in
the aggregate of characters the most typical of albicaudatus of any
Durrant — Pocket Gophers of Utah 39
in the entire range. Those from the west side of the river, although
definitely referable to albicaudatus do show some intergradation
with Thomomys bottae robustus, the subspecies to the west.
The specimens from Bauer, Tooele County, are relatively uniform
in color, and are considerably lighter than topotypes of albicauda-
tus. Their upper parts vary from Sepia to Saccardo's Umber as
compared with near (13 ' ' ' ' n) Black of the topotypes. The sides
and underparts are lighter, due primarily to much less black in the
underfur. They average slightly longer in total length, but shorter
in hind foot. All cranial measurements are slightly smaller than
in topotypes of albicaudatus. The shape of the skull closely re-
sembles that of albicaudatus, although the rostrum, nasals, upper
incisors and posterior tongues of the premaxillae tend to be nar-
rower. This narrowness indicates intergradation with Thomomys
bottae stansburyi, the race nearest to the west. These animals are
in the majority of characters referable to albicaudatus.
Bauer is situated in extreme western Tooele Valley at the foot
of Stockton Bar, a low pass between the Stansbury and the Oquirrh
mountains. This valley lies to the west of the aforementioned Jor-
dan River. Although these gophers are definitely referable to albi-
caudatus they are more unlike topotypes than are the animals from
Riverton.
The specimens from Settlement Canyon, Oquirrh Mountains,
Tooele County, show the same characteristics as those from Bauer.
In a large series of animals from St. John, in Rush Valley, Tooele
County, the upper parts vary from black, even darker than topo-
types of albicaudatus, to Tawny Olive, and the underparts vary
from black through Cinnamon Buff to Pinkish Buff. Most of the
animals are Cinnamon Buff. Although variable they approach albi-
caudatus in color. The total length, tail and hind foot of males
are longer than in topotypes of albicaudatus; females differ in the
same direction but only slightly. In both sexes the zygomatic
breadth is less, but the mastoid breadth is greater than in albi-
caudatus. In size and shape of the lacrimal processes, and the
great thickening of the jugal at the maxillo-jugal suture they ap-
proach robustus. They are much larger, however, and in the ma-
jority of characters are referable to albicaudatus.
What has just been said relative to the animals from St. John
applies also to those from Clover Creek in the Onaqui Mountains
of Tooele County. At the latter locality the tendencies towards
robusttis are accentuated. This is to be expected, since this locality
40 University of Kansas Publs., Mus. Nat. Hist.
is midway between St. John and the type locality of robustus. All
characters considered, these animals are all referable to albicaudatus.
The animals from Little Valley, Sheeprock Mountains, Tooele
County, resemble albicaudatus in color. They vary on the upper
parts from near (1) Sepia to Clay Color, and ventrally from nearly
black to Pinkish Buff. They are markedly smaller in every meas-
urement taken, except zygomatic and mastoidal breadths, and ex-
tension of premaxillae posterior to nasals. This relatively greater
breadth indicates intergradation with robustus to the west. These
gophers are smaller in most measurements than any other popula-
tion referred to albicaudatus. This is understandable because
gophers from mountains usually are smaller and have weaker,
smoother skulls than animals from low lands. Although approach-
ing robustus in size and in some aforementioned cranial details,
the aggregate of characters including color, make these animals re-
ferable to albicaudatus.
The animals from Fairfield, Utah County, are closer geograph-
ically to the type locality of albicaudatus than any other series, but
morphologically are the least like topotypes. At first glance one
is struck with the differences. They are uniformly Clay Color
above, with Cinnamon Buff sides and flanks and Pinkish Buff un-
derpays. Their color closely approaches that of robustus to the
west which has Cinnamon Buff on the upper parts. Examination
of eleven measurements of males and the same number for females,
shows that the animals are nearest to robustus in two measurements,
to albicaudatus in 12, distinct in 7 and intermediate in one. The
general appearance of the skull is intermediate between that of the
two above mentioned forms. The differences from albicaudatus in
size and color may be correlated with the differences in soil at Fair-
field and Provo. At Fairfield the soil is light-colored clay, but at
Provo it is sandy and darker. Although they are intergrades be-
tween robustus and albicaudatus, the animals are referred to the
latter race. Utah Lake and its outlet, the Jordan River, make a
partial barrier between populations at Fairfield and at the type
locality at Provo. During Pleistocene times, when Lake Bonne-
ville was present it formed a complete barrier. Enough time has
evidently elapsed since the disappearance of this lake to allow
albicaudatus, the mainland form, to expand its range to the west.
Intergradation has taken place, with the result that the animals
from Fairfield, although unstable, agree with the mainland form,
albicaudatus, in a majority of their characters.
Durrant — Pocket Gophers of Utah 41
Pocket gophers were taken at four localities from north to south
in eastern Juab County. They range in color from Ochraceous
Tawny on the upper parts and Cinnamon Buff on the underparts to
shades that are slightly lighter. All are much lighter than topotypes
of albicaudatus. The general configuration of the skull is the same
as that of albicaudatus, and this is especially true in the females.
In the narrower rostrum and weaker dentition they approach con-
tractus, but are distinctly lighter colored. Hall (1931:3) referred
one specimen from Nephi, Juab County, to Thomomys bottae aurei-
ventris. Since that time Thomomys bottae lenis which has some
affinities with aureiventris has been described (see account of con-
tractus). The large series now available from Nephi and nearby
localities do show some intergradation with lenis, in that four char-
acters are more as in lenis and contractus and seven characters are
more as in albicaudatus. Although differing markedly in many re-
spects from topotypes of albicaudatus they fit the aforementioned
concept of this subspecies, and are being treated as a variable local
population of it.
Provo is the locality listed for specimens which were available to
naturalists from 1875-1877. To these specimens the following names
were applied: Thomomys talpoides bulbivorus Coues (1875:256;
1877:627) and' Thomomys talpoides umbrinus Coues and Yarrow
(1875:112). Possibly these names were applied to the animals cur-
rently known as Thomomys bottae albicaudatus which does occur
at Provo. Without the opportunity to examine the actual specimens,
which so far as I know are no longer in existence, I cannot exclude
the possibility that the locality designation ''Provo" was used in a
general sense to include pocket gophers taken a few miles to the
eastward of Provo, where it is known that pocket gophers of only
the species Thomomys talpoides (current terminology) occur.
Specimens examined. — Total, 239, distributed as follows: Davis County: Bountiful,
4,500 ft., 1. Salt Lake County: Salt Lake City and environs, 4,300 ft., 51; 2 mi. W
Murray, 4,300 ft., 6; Riverton, 4,300 ft., 11; Draper, 4,500 ft., 7; Rose Canyon, Oquirrh
Mountains, 5,650 ft., 4. Tooele County: Bauer, 4,500 ft., 30; Settlement Creek, Oquirrh
Mountains, 6,500 ft., 1; St. John, 4,300 ft,, 28; Clover Creek, Onaqui Mountains, 5,500 ft.,
15; Vernon, 4,300 ft., 2 (U. S. A. C); Little Valley, Sheeprock Mountains, 5,500 ft., 20.
Utah County: Fairfield, 4,800 ft., 24; Provo, 4,400 ft., 20 (8, B. Y. U. ; 12, M. V. Z.).
Juab Caunty: Neff Farm, 4 mi. N Nephi, 5,000 ft., 2 (1, R. H.); Nephi, 5,000 ft., 1
(M. V. Z.); 2 mi. S Nephi, 4,700 ft., 14; 7 mi. SW Nephi, 6,000 ft., 2.
Thomomys bottae bonnevillei new subspecies
Type. — Male, adult, skin and skull, No. 3576, Museum of Zoology, Uni-
versity of Utah; Fish Springs, 4,400 ft., Juab County, Utah; June 8, 1940;
collected by S. D. Durrant; original number 1955.
Range. — Known only from the type locality.
42 University of Kansas Publs., Mus. Nat. Hist.
Diagnosis. — Size medium (see measurements); claws on front feet small.
Color: Entire dorsal surface Warm Buff; sides near (e) Cinnamon Buff, un-
derpays near (16") Pale Pinkish Buff; inguinal region, front and hind feet
and distal part of tail white; top of head, nose and cheeks grayish black;
postauricular patches small and grayish black; ears small, pointed and with
heavily pigmented pinnae. Skull: Angular, short and wide; nasals of me-
dium length, narrow proximally but widely flared distally; interparietal small;
lambdoidal suture concave towards the interparietal; zygomatic arches uni-
formly widely spreading; interpterygoid space widely V-shaped; extension of
premaxillae posterior to nasals long; lambdoidal crest well developed.
Comparisons. — From topotypes of Thomomys bottae aureiventris,
bonnevillei differs as follows: Size smaller, hind foot shorter. Color:
Upper parts and sides lighter; underparts pale buff rather than
"gold." Skull: Shorter and relatively wider; rostrum wider and
heavier; zygomatic arches relatively wider and more massive, with
greatest width posteriorly instead of anteriorly; interpterygoid
space widely V-shaped rather than lyre-shaped ; thickening at union
of jugal and zygomatic process of maxilla less developed; anterior
palatine foramina larger; nasals shorter and more markedly flared
distally; zygomatic breadth relatively, and mastoidal breadth actu-
ally, wider; extension of premaxillae posterior to nasals greater;
tympanic bullae more inflated ventrally; upper incisors wider.
From near topotypes of Thomomys bottae centralis, from 1 mile
east of Garrison, Millard County, Utah, bonnevillei differs as fol-
lows: Size smaller; hind foot and tail shorter. Color: Generally
darker above and lighter below; top of head darker; postauricular
patches smaller and lighter. Skull: Shorter and wider (zygomatic
breadth expressed in percent of basilar length being, in males, 74.5
in bonnevillei and 71.5 in centralis) ; interpterygoid space more
widely V-shaped; interparietal smaller, and more triangular; nasals
shorter and much more dilated distally, as well as more constricted
proximally; lacrimal processes smaller and less globuse at tips; tem-
poral fossae larger; braincase and entire dorsal surface of skull more
nearly flat; lambdoidal suture convex posteriorly as opposed to
nearly straight; tympanic bullae more inflated ventrally.
Comparisons of bonnevillei with the type and type series of
Thomomys bottae wahwahensis show them to be of approximately
the same size, but to differ as follows: Color: Slightly darker above
and lighter below; postauricular patches smaller and lighter. Skull:
Larger in every measurement taken, except breadth of rostrum
which is smaller; skull not as flat; tympanic bullae more inflated
ventrally; nasals and rostrum longer; extension of premaxillae pos-
Durrant — Pocket Gophers of Utah 4:J
terior to nasals greater; interparietal smaller and more triangular;
zygomatic arches more bowed out laterally; jugals heavier; inter-
pterygoid space more widely V-shaped; upper incisors less massive.
The characters that distinguish bonnevillei from Thomomys bot-
tae albicaudatus are: Size smaller. Color: Markedly lighter
throughout. Skull: Shorter and wider; mastoid and zygomatic
breadths greater; rostrum narrower but shorter; angle between ros-
trum and zygomatic processes of maxillae less; interparietal smaller
and more triangular; extension of premaxillae posterior to nasals
greater; upper incisors shorter, narrower and more recurved.
T. b. bonnevillei is indistinguishable in color from Thomomys
bottae convexus, but differs from it in the following features: Size
larger in nearly every measurement taken. Skull: Flattened dor-
sally as opposed to convex; zygomatic arches longer and weaker;
jugals more nearly perpendicular; tympanic bullae larger; upper
incisors longer; alveolar length of upper molar series the same, but
molars narrower; rostrum longer but nasals shorter; extension of
premaxillae posterior to nasals greater.
Topotypes of bonnevillei can be distinguished from those of both
Thomomys bottae tivius and stansburyi by being larger in every
measurement taken, by markedly lighter color throughout, and by
ridged, massive, angular skulls rather than smooth, weak, non-
angular skulls.
The races closest geographically to bonnevillei are Thomomys
bottae robustus and T. b. sevieri. Compared with topotypes of ro-
bustus, bonnevillei differs in: Size larger. Color: Lighter through-
out. Skull: Larger, although not as compact; zygomatic arches
mure widely spreading; jugals lighter; lacrimal processes not as
prominent; zygomatic processes of maxillae not as robust; nasals
more flared distally; extension of premaxillae posterior to nasals
greater; alveolar length of upper molar series longer; molars larger;
upper incisors longer, wider and darker in color; when placed
ventral side down on a surface, the dorsal face of a skull of robustus
is approximately parallel to the surface, whereas one of bonnevillei
dips down in the occipital region.
T. b. sevieri can be easily distinguished from bonnevillei by being
smaller in every measurement taken, darker in color, and by small,
weak, smooth skulls as opposed to large, robust, ridged skulls.
Remarks. — Fish Springs, where bonnevillei occurs is a marshy
area south of the barren, salt-desert country of western Utah. The
source of water is springs at the base of the north end of the Fish
44 University of Kansas Publs., Mus. Nat. Hist.
Springs Mountains. Only the moist area supports pocket gophers.
Specimens from Trout Creek, Juab County, twenty-five miles to
the southwest are intergrades between bonnevillei and aureiventris,
and are referred to the latter subspecies. The country between Fish
Springs and Trout Creek in 1937 and 1940 lacked pocket gophers;
it was of the playa and sand type. Probably T. b. bonnevillei was
derived from T. b. aureiventris, a western mainland form of Pleis-
tocene Lake Bonneville, through isolation and subsequent differen-
tiation morphologically. The moist soils at Cane Springs, seven
miles south of Fish Springs, had no pocket gophers when visited in
1940.
Specimens examined. — Total, 11, from the type locality.
Thomomys bottae centralis Hall
Thomomys perpallidus centralis Hall, Univ. California Publ. Zool..
32:445, July 8, 1930.
Thomomys bottae centralis Goldman, Proc. Biol. S'oc. Washington,
48:156, October 31, 1935; Hall and Johnson, Proc. Utah Acad. Sci. Arts
and Letters, 15:121, 1938.
Type. — Male, adult, skin and skull, No. 41688, Museum of Vertebrate
Zoology, University of California; 2l/-> mi. E Baker (1*4 mi. W Nevada-Utah
boundary on 39th parallel), 5,700 ft.. White Pine County, Nevada; May 30,
1929; collected by E. Raymond Hall; original number 2683.
Range. — Extreme western Utah, in Millard, Beaver and Iron counties.
Diagnosis. — Size medium (see measurements); tail long; claws on front feet
long. Color: Near Cinnamon Buff on upper parts, darker in middorsal
region, grading to Pinkish Buff on underparts, more accentuated in pectoral
and inguinal regions; nose, cheeks and postauricular patches grayish black;
front and hind feet and distal half of tail white. Skull: Robust and moder-
ately ridged; zygomatic breadth about the same for entire length of arches;
jugals vertical posterior to middle; moderate thickening present at region of
maxillo-jugal suture; interpterygoid space narrowly V-shaped; dorsal fronto-
maxillary sutures convex medially; lacrimal processes globose and well de-
veloped; nasals long and with distal denticulations; paroccipital processes well
developed.
Comparisons. — Compared with topotypes of Thomomys bottae
albicaudatus, centralis differs as follows: Size larger; tail longer;
claws on front feet longer. Color: Lighter throughout, Cinnamon
Buff as opposed to near (13 " " n) Black. Skull: Basilar length
and length of nasals greater; zygomatic breadth less; zygomatic
arches thicker at region of maxillo-jugal sutures; interpterygoid
space more broadly V-shaped; dorsal frontomaxillary sutures convex
medially as opposed to straight; paroccipital processes more de-
veloped; zygomatic arches approximately the same width through-
out as opposed to widest posteriorly.
Dukrant — Pocket Gophers of Utah 45
For comparisons with Thomomys bottae aureiventris see account
of that form.
T. b. centralis can be distinguished from Thomomys bottae bon-
nevillei, robustus, sevieri and convexus by larger size throughout
and generally darker color (see accounts of those forms). From
Thomomys bottae stansburyi and tivius, centralis differs in larger
size throughout and lighter color (see accounts of those forms) .
Remarks. — Thomomys bottae centralis has one of the most ex-
tensive ranges of any of the known races of T. bottae. The eastern
limits extend into extreme western Utah. Specimens from Utah for
the most part are intergrades between centralis and aureiventris, the
race to the north. Some minor intergradation is also noted between
centralis and sevieri and bonnevillei, the races to the east. Inter-
gradation is the expected condition because the animals belonging
to centralis are at the extremes of their range in this area. The
greater affinities of these animals with aureiventris is to be expected
because both aureiventris and centralis are forms of the western
mainland of the Pleistocene Lake Bonneville; while the races to the
east, although closest geographically, were isolated from the gophers
of the western mainland during prehistoric times by this lake. They
are still isolated and enough time has elapsed so that only vestiges
of morphological intergradation exist between centralis and these
eastern forms. Two specimens from Cedar City, Iron County, are
intergrades between Thomomys bottae wahwahensis, centralis and
planirostris. Their skulls are slightly convex as in planirostris, and
the rostrum is short and wide as in wahwahensis. In shape of the
zygomatic arches, length of the nasals, and color, they resemble
centralis to which they are here referred.
Specimens examined. — Total, 49, distributed as follows: Millard County: 1 mi. SE Gandy,
5,000 ft., 15 (M. V. Z.); White Valley (Tule Spring), 60 mi. W Delta, 4, (3 in R. W.
Fautin Vertebrate Collection) ; Robison Ranch, 5,300 ft., (on Hendry Creek) Simonsons
Ranch, 4,596 ft., 2 (M. V. Z.); 1 mi. E Garrison, 5,000 ft., 21; 5 mi. S Garrison, 5,400 ft.,
5 (M. V. Z.). Iron County: Cedar City, 2 (M. V. Z.).
Thomomys bottae sevieri new subspecies
Type.— Female, adult, skin and skull, No. 2530, Museum of Zoology, Uni-
versity of Utah; Swasey Spring, House Mountains, 6,500 ft., Millard County,
Utah; May 16, 1938; collected by S. D. Durrant; original number 1380.
Range. — Known only from the type locality.
Diagnosis. — Size medium (see measurements) ; claws on front feet short
and weak; ears short; tail relatively long. Color: Upper parts Pinkish Buff,
grading over sides to Pale Pinkish Buff on underparts; nose, top of head,
chin and cheeks grayish black; postauricular patches small and grayish black;
front and hind feet and distal two-thirds of tail white. Skull: Small, weak
46 University of Kansas Publs., Mtjs. Nat. Hist.
and smooth; rostrum narrow ; nasals narrow, not markedly flared distally;
zygomatic arches weak, not angular, and of "graceful" contour; lacrimal pro-
cesses small; characteristic dorsal depression present in region of sa^itto-
coronal suture; mastoid and zygomatic breadths narrow; occiput narrow and
high; braincase well inflated; paroccipital processes small and smooth; inter-
pterygoid space narrowly V-shaped; tympanic bullae small, but well inflated
ventrally; alveolar length of upper molar series short; molars small; upper
incisors short, but narrow.
Comparisons. — From topotypes of Thomomys bottae aureiven-
tris, sevieri differs as follows: Size smaller. Color: Lighter
throughout, no "gold" on underparts. Skull: Much smaller in
every measurement taken, less massive and not angular; zygomatic
arches weaker and widest posteriorly rather than anteriorly; union
of jugal and zygomatic process of maxilla not greatly thickened;
interpterygoid space narrowly V-shaped rather than lyre-shaped;
pterygoid hamulae shorter and weaker; tympanic bullae smaller,
but markedly more inflated ventrally; dentition smaller and
weaker.
From near topotypes of Thomomys bottae centralis, sevieri can
be distinguished by the following features: Size markedly smaller.
Color: Lighter throughout. Skull: Markedly smaller in every
measurement taken, weaker and smoother; zygomatic arches
weaker, less angular and more "graceful"; rostrum shorter, but
narrower; lacrimal processes smaller; tympanic bullae smaller, but
more inflated ventrally, being triangular in shape as opposed to
ovate and with anteromedial margin decidedly pointed; pterygoid
hamulae smaller and weaker; dentition smaller and weaker.
T. b. sevieri can readily be distinguished from Thomomys bottae
albicaudatus by the following features: Size smaller in every mea-
surement taken. Color: Markedly lighter throughout. Skull:
Smaller, and weaker; rostrum shorter and narrower; ascending
processes of premaxillae narrower; extension of premaxillae pos-
terior to nasals shorter; posterior tongues of premaxillae narrower;
dentition much lighter.
Comparisons of sevieri with topotypes of Thomomys bottae wah-
wahensis show them to be of approximately the same size, but to
differ as follows: Hind foot longer; ear shorter. Color: Slightly
darker. Skull: Smaller, weaker, less ridged; zygomatic breadth
less; zygomatic arches markedly less angular; mastoid breadth less;
rostrum much longer and narrower, not as blunt nor flattened;
tympanic bullae much larger and more inflated ventrally; braincase
vaulted as opposed to flattened.
Durrant — Pocket Gophers of Utah 47
From topotypes of Thomomys bottae bonnevillei, sevieri differs
in: Size smaller throughout. Skull: Smaller in every measurement
taken, weaker, smoother and less angular; dentition smaller and
weaker.
Topotypes of sevieri are easily distinguished from those of Tho-
mcmys bottae robustiis by smaller size, and smaller, markedly
weaker skull which is less angular and ridged.
Among named races of Thomomys bottae, sevieri is closest geo-
graphically to convexus, but differs from it as follows: Size larger;
hind foot longer. Skull: Smaller in every measurement taken; na-
sals shorter and not so flaring distally; rostrum weaker, narrower
:md not so depressed; zygomatic arches markedly weaker and less
angular; lacrimal processes smaller; supraoccipital narrower and
higher; paroccipital processes weaker; tympanic bullae smaller;
dentition markedly weaker.
Topotypical specimens of sevieri can be readily distinguished
from those of Thomomys bottae tivius by Pinkish Buff instead of
Mummy Brown on upper parts. Tympanic bullae larger and mark-
edly more inflated; nasals longer; zygomatic and mastoidal breadths
greater; rostrum longer and more depressed; upper incisors longer
and wider; molariform teeth smaller. The skulls of sevieri re-
semble those of tivius more closely than those of any other sub-
species.
Remarks. — The House Mountains in western Millard County are
surrounded by desertlike terrain that is seemingly unsuited to pocket
gophers. In these mountains, gophers were sought in vain at sev-
eral localities, including Antelope Springs which superficially ap-
peared suitable for the animals. Pocket gophers were found only
at the type locality, Swasey Spring, which is well above the high-
est level of the Pleistocene Lake Bonneville. T. b. sevieri, like T .
b. minimus on Stansbury Island, Great Salt Lake, appears to re-
main only on land that was an island when Lake Bonneville was
at its highest level.
Specimens examined. — Total, 10, from the type locality.
Thomomys bottae convexus Durrant
Thomomys bottae convexus Durrant, Proc. Biol. Soc. Washington,
52:159, October 11, 1939.
Type. — Male, adult, skin and skull, No. 2482, Museum of Zoology, University
of Utah; E side Clear Lake, 4,600 ft, Millard County, Utah; May 20, 1938;
collected by S. D. Durrant; original number 1401.
48 University of Kansas Publs., Mus. Nat. Hist.
Range. — Westcentral Utah in Delta Valley.
Diagnosis. — Size medium (see measurements). Color: Upper parts and
sides Pinkish Buff, purest on sides; underparts Pale Pinkish Cinnamon; in-
guinal and pectoral regions Pale Pinkish Buff; nearly all specimens have
white on perineal region; nose grayish black; front feet, hind feet and distal
third to half of tail white; postauricular patches black. Skull: Braincase
moderately convex on dorsal surface; rostrum strongly depressed, giving the
entire dorsal surface of the skull a "rocker-shape"; zygomatic arches heavy,
short and widely spreading, widest posteriorly; upper incisors recurved, short
and wide; molariform teeth large; alveolar length of upper molar series long;
palatal pits deep; tympanic bullae moderately inflated ventrally; mastoidal
breadth actually as well as relatively wide.
Comparisons. — Compared with topotypes of Thomomys bottae
wahwahensis, conv exits is of approximately the same color, but
differs as follows: Size smaller; tail, hind foot, and ear shorter.
Skull: Rostrum longer, narrower and more depressed; skull convex
rather than flat; nasals longer, and convex rather than flat; tym-
panic bullae larger; zygomatic arches shorter and more massive;
molariform teeth larger.
From topotypes of Thomomys bottae centralis, convexus differs
in: Size smaller; tail and hind foot shorter. Color: Uniformly
lighter, more white in perineal region. Skull: Smaller, more con-
vex; rostrum shorter, wider and more depressed; zygomatic arches
shorter and heavier; mastoidal breadth actually, as well as rela-
tively wider; tympanic bullae more inflated ventrally; upper in-
cisors shorter and wider.
Comparatively, topotypes of convexus can be distinguished from
those of Thomomys bottae aureiventris by: Size smaller; tail and
hind foot shorter. Color: Darker on upper parts; no "gold" on
underparts. Skull: Smaller and more nearly flat; rostrum shorter
and more depressed; zygomatic arches shorter, heavier and widest
posteriorly rather than anteriorly; interpterygoid space V-shaped
as opposed to lyre-shaped; upper incisors shorter, narrower and
more recurved.
Topotypical specimens of convexus differ from those of Thom-
omys bottae nesophilus as follows: Size smaller; tail and hind foot
shorter. Color: Uniformly lighter throughout, Cinnamon Buff as
opposed to Pinkish Buff. Skull: Smaller; rostrum heavier, shorter
and more depressed; zygomatic arches shorter, heavier and not so
widely spreading; no widening of supraoccipital as in nesophilus',
upper incisors shorter and more recurved.
When compared with topotypes of Thomomys bottae albicauda-
tus, convexus shows the following differences: Size smaller; tail and
Durrant — Pocket Gophers of Utah 49
hind foot shorter. Color: Markedly lighter throughout, Skull:
Smaller, more convex and compact; rostrum shorter, heavier, more
depressed and compact; zygomatic arches shorter and more robust;
upper incisors shorter and more recurved.
Thomomys bottae tivius is the race closest geographically to con-
vexus. From it, convexus can be readily distinguished by: Size
larger; tail shorter; hind foot longer. Color: Markedly lighter
throughout. Skull: Much heavier and more compact, weights of
skulls of males and females of the two subspecies being 2.4 grs., 1.6;
1.6, 1.2, respectively; rostrum heavier, wider and more depressed;
zygomatic arches shorter, and more massive ; upper incisors shorter,
wider and more recurved; molariform teeth larger.
For comparisons with Thomomys bottae lenis, contractus, sevieri,
bonnevillei, and robustus see accounts of those forms.
Remarks. — T. b. convexus is limited to the area around Clear
Lake in Millard County. This lake is surrounded by areas of loose,
shifting sand and flat areas of barren alkali. The lake is fed by
springs which flow from lava outcroppings on its eastern side. As
far as discernible, the only area populated by pocket gophers (1938)
was that adjacent to the lake where vegetation had trapped the
sand. The factor which limits the extension of range of this sub-
species probably is plant food. Also, the soil is mechanically poor
for burrowing, since it caves in easily and burrows were found only
in the sand where salt grass (Distichlis stricta) had trapped and
stabilized it. Burrows were found from the edge of the water back
as far as this grass persisted.
Specimens examined. — Total, 17, from the type locality.
Thomomys bottae tivius Durrant
Thomomys bottae tivius Durrant, Bull. Univ. Utah, 28 (No. 4) :5,
August 18, 1937.
Type.— Female, adult, skin and skull, No. 1827, Museum of Zoology, Uni-
versity of Utah; Oak Creek Canyon, 6 mi. E Oak City, 6,000 ft., Millard
County, Utah; September 14, 1936; collected by S. D. Durrant; original num-
ber 1100.
Range. — Limited to the Canon Mountains, Millard County.
Diagnosis. — Size small (see measurements). Color: Upper parts Mummy
Brown, grading through Cinnamon on the sides to Pale Cinnamon on the
underparts; cheeks Cinnamon; postauricular patches black; distal third to
half of tail white. Skull: Small, weak; zygomatic arches weak, not widely
spreading, widest posteriorly; tympanic bullae large; interpterygoid space
V-shaped; nasals short, usually simple distally, but with some denticulations
4—2786
50 University of Kansas Publs., Mrs. Nat. Hist.
in some specimens; palatal pits deep; palate narrow; paroccipital processes
small; incisors, both upper and lower, narrow; molariform teeth small.
Comparisons. — Topotypes of tivius differ from those of Thomo-
mys bottae albicaudatus as follows: Size markedly smaller in every
measurement taken. Color: Lighter, Mummy Brown as opposed
to near (13 " " n) Black. Skull: Smaller, slenderer and weaker;
zygomatic arches weak and not widely spreading as opposed to
massive and wide spreading; nasals and rostrum narrower and
shorter; extension of premaxillae posterior to nasals shorter; tym-
panic bullae smaller; molariform teeth smaller.
For comparisons with Thomomys bottae stansburyi and T. b.
contractus see accounts of those forms.
The four subspecies tivius, albicaudatus, stansburyi, and contrac-
tus are the darkest in color of all the Thomomys bottae occurring
within the state.
Remarks. — This small, dark subspecies is limited to the Canon
Mountains in eastern Millard County. Apparently it is a mountain
derivative of Thomomys bottae contractus which occurs in the val-
leys to the east and west of these mountains. Intergradation is
noted with animals from the valleys on either side. For further
comments on distributional problems of this type see remarks under
Thomomys bottae stansburyi.
Specimens examined. — Total, 12, from the type locality.
Thomomys bottae contractus new subspecies
Thomomys perpallidus albicaudatus Hall, Univ. California Publ. Zool.,
37:3, April 10, 1931.
Thomomys bottae albicaudatus Dun-ant. Bull. Univ. Utah, 28 (No.
4) A, August 18, 1937.
Type. — Male, adult, skin and skull, No. 1851, Museum of Zoology, Uni-
versity of Utah; Scipio, 5,315 ft,, Millard County, Utah; September 17, 1936;
collected by S. D. Durrant; original number 1125.
Range. — -Extreme eastern Millard and Beaver counties, Utah.
Diagnosis. — Size medium (see measurements). Color: Upper parts Cinna-
mon Buff, mixed with black giving a color of Dresden Brown; sides between
Cinnamon Buff and Pinkish Buff; underparts Pinkish Buff, purest on inguinal
and pectoral regions; postauricular patches medium in size and black; ears
covered with black hairs; nose, chin, cheeks and top of head dusky; front
feet, hind feet and distal third to half of tail white; proximal part of tail
covered all around with buff-colored hairs. Skull: Long, slender, moderately
ridged and convex transversally at proximal ends of nasals; nasals long; ros-
trum long and narrow; posterior ends of nasals truncate or shallowly emar-
ginate; ascending processes of premaxillae slender; extension of premaxillae
posterior to nasals long; zygomatic arches neither robust nor widely spread-
. Ditrrant — Pocket Gophers of Utah 5]
ing; interparietal subquadrangular ; supraoccipital extending horizontally well
behind lambdoidal suture instead of dropping off abruptly to the foramen
magnum ; interpterygoid space moderately V-shaped in some specimens, but
somewhat lyre-shaped in others; tympanic bullae large and truncate anteri-
orly; upper incisors long and narrow; molariform teeth small and light.
Comparisons. — Compared with topotypes of Thomomys bottae
albicaudatus, contractus differs as follows: Tail longer. Color:
Lighter throughout. Skull: Slenderer, less ridged and angular;
rostrum narrower; zygomatic and mastoidal breadths less; ascend-
ing processes of premaxillae narrower; posterior tongues of pre-
maxillae narrower; posterior ends of nasals less truncate; zygo-
matic arches weaker, less angular, and less widely spreading pos-
teriorly; interparietal larger; paroccipital processes weaker; inter-
pterygoid space not as widely V-shaped; upper incisors longer and
narrower; molariform teeth smaller.
Topotypes of contractus can be distinguished from those of Tho-
momys bottae convexus by the following: Size larger, tail longer;
hind foot larger. Color: Darker throughout. Skull: Longer, nar-
rower, and not as massive; top of skull moderately, as opposed to
strongly, convex; nasals arched rather than straight; zygomatic
arches neither as widely spreading, angular nor massive ; space en-
closed within zygomatic arches longer; interparietal larger; inter-
pterygoid space more narrowly V-shaped ; upper incisors longer and
narrower; molariform teeth much lighter.
Comparisons of topotypes of contractus with near topotypes of
Thomomys bottae centralis show them to be approximately the
same size, but to differ as follows: Color: Darker throughout,
Skull: Shorter and slenderer; rostrum narrower; region between
posterior tongues of premaxillae narrower and more convex trans-
versally; nasals more truncate; zygomatic breadth less, but arches
relatively more widely spreading posteriorly; interparietal larger;
interpterygoid space generally narrower; upper incisors longer and
narrower; molariform teeth smaller.
Topotypes of contractus differ from those of Thomomys bottae
aureiventris as follows: Size smaller; tail longer; hind foot shorter.
Color: Darker throughout. Skull: Shorter but slenderer; rostrum
narrower; nasals shorter but slenderer, and more truncate pos-
teriorly; extension of premaxillae posterior to nasals longer; zygo-
matic arches weaker and less angular; zygomatic processes of max-
illae weaker and with no marked thickenings at union of maxilla
and jugals; interparietal larger; interpterygoid space more gener-
52 University of Kansas Publs., Mus. Nat. Hist.
ally V-shaped; upper incisors longer and narrower; molariform
teeth smaller.
Compared with topotypes of Thomomys bottae planirostris,
contractus differs in: Size smaller throughout. Color: Darker,
more black and less Cinnamon in pelage. Skull: Smaller in every
measurement taken; rostrum narrower; nasals arched instead of
flat; zygomatic arches neither angular, massive nor widely spread-
ing; upper incisors narrower; molariform teeth markedly smaller
and weaker.
Topotypes of contractus differ from those of Thomomys bottae
levidensis in larger size, darker color and longer, slenderer skulls.
Among named races of T. bottae, contractus is closest morpho-
logically to tivius. It differs from it as follows : Size larger through-
out. Color: Lighter throughout. Skull: The same general shape
and proportions, but larger in every measurement taken; rostrum
longer and narrower; extension of premaxillae posterior to nasals
longer; posterior tongues of premaxillae narrower.
Remarks. — Fifteen animals from Oak City are intergrades be-
tween contractus and tivius. Intergradation with lenis is also
shown in some specimens by the widely spreading zygomatic
arches. In the majority of characters including the diagnostic long,
slender, narrow rostrum they are more like contractus to which they
are here referred.
Nine animals from Beaver were considered by Hall (1931:3) and
Durrant (1937:4) to be intergrades between Thomomys bottae
albicaudatus and Thomomys bottae centralis. Restudy of these
specimens in the light of additional material now shows them to be
intergrades between T. b. centralis, T. b. planirostris and T. b. con-
tractus. The majority of these animals are intermediate in color
between centralis and contractus, but a few have the reddish cast of
planirostris. The shape of the nasals is characteristic of planiros-
tris, while the zygomatic arches are as in centralis. In the re-
mainder of the diagnostic characters they are like contractus to
which they are here referred.
Strong affinities exist between albicaudatus, tivius and contractus.
All three of these races probably stemmed from a dark form which
formerly inhabited the eastern mainland of the Pleistocene Lake
Bonneville. At present, tivius is isolated on the Canon Mountains
in eastern Millard County, while the range of albicaudatus and con-
tractus have been separated by that of lenis. T. b. lenis has the
majority of its affinities with aureiventris which is an inhabitant of
Durrant — Pocket Gophers of Utah 53
the western mainland of this ancient lake. An understanding of
the history of the Sevier River Valley will probably clarify this
distribution of pocket gophers.
Specimens examined. — Total, 39, distributed as follows: Millard County: Oak City, 6,000
ft., 15; Scipio, 5,315 ft., 15. Beaver County: Beaver, 6,000 ft., 9 (M. V. Z.).
Thomomys bottae lenis Goldman
Thomomys townsendii lenis Goldman, Proc. Biol. Soc. Washington,
55:75, June 25, 1942.
Thomomys perpallidus aureus Moore, Journ. Mamm., 10:259; No-
vember 11, 1931.
Type.—MsAe, adult, skin and skull, No. 264805, U. S. National Museum
(Biological Surveys Collection); Richfield, 5,308 ft., Sevier County, Utah;
March 11, 1928; collected by A. W. Moore; X-catalogue number 28835
(after Goldman, type not seen).
Range. — Sevier River Valley from Piute County north to southwestern Juab
and northeastern Millard counties, Utah.
Diagnosis. — Size large (see measurements). Color: Upper parts Cinnamon
Buff mixed with black in middorsal region; sides, flanks, forearms, thighs and
underparts Pinkish Buff; inguinal region, front feet, hind feet, underpart of
tail and end of tail white; postauricular patches small and dusky; chin,
cheeks, nose and top of head dusky. Skull : Largest of Utah gophers, massive
and angular; nasals long and denticulate distally; rostrum long and relatively
narrow; zygomatic arches widely spreading and heavy throughout; jugals
nearly vertical; zygomatic processes of maxillae heavy and flaring out
abruptly from base of rostrum; union of zygomatic process of maxilla and
jugal greatly thickened; extension of premaxillae posterior to nasals long;
posterior tongues of premaxillae relatively narrow; lacrimal processes small;
pterygoid hamulae long; interpterygoid space moderately V-shaped, tending
to be somewhat lyre-shaped in some specimens; tympanic bullae somewhat
flattened, only moderately inflated ventrally; upper incisors long and narrow;
molariform teeth actually large, but relatively small.
Comparisons. — Topotypes of lenis can be distinguished from
those of Thomomys bottae tivius, convexus, contractus, albicau-
datus, levidensis, centralis and aureiventris by the following mark-
edly greater average measurements of males: Total length, 250
mm.; length of nasals, 15.5; zygomatic breadth, 28.3; mastoid
breadth, 22.5; and length of rostrum, 18.3. Other distinguishing
characters are: Zygomatic arches more widely spreading; length
of zygomatic processes of maxillae greater; and relatively longer,
narrower rostrum.
Remarks. — Twenty-one animals obtained from Lynndyl, Millard
County, are all intergrades between lenis and aureiventris. They are
like aureiventris in the shape of the zygomatic arches, and in the
bowing of the parietal crests. Slight intergradation with centralis
is indicated by color and the shape of the nasals. The transverse
54 University of Kansas Publs., Mus. Nat. Hist.
arching of the posterior part of the rostrum is indicative of some
relationship with contractus. In six other characters studied they
most closely approach lenis to which they are here referred.
Large size is the distinctive feature of Thomomys bottae lenis.
The skulls are the largest of any species or subspecies of Thomomys
found in Utah. In total length, however, these animals are no
longer than the extremes found in other named races. When Gold-
man (1942:75) described this race as new, he referred it to the
species Thomomys townsendii, but remarked that the animal from
Richfield was different enough from any other form then named to
merit probably full specific status. I know of no character other
than size to separate Thomomys townsendii from Thomomys bottae,
and since intergradation has been shown to exist between these al-
leged townsendii from Richfield and animals from extreme western
Utah known to belong to the species bottae, lenis is here arranged
as a subspecies of Thomomys bottae which name has priority over
Geomys townsendii.
The range here ascribed to this race is the Sevier River Valley
from Piute County as far downstream as the town of Lynndyl which
is near the eastern mainland of Pleistocene Lake Bonneville. The
Sevier River continues farther out into Delta Valley ultimately to
empty into Sevier Lake, which at present is adjacent to the area
that formerly constituted the western mainland of the aforemen-
tioned ancient lake. This watercourse may have provided a mi-
gration route in ancient times, during the fluctuations of Lake Bon-
neville, whereby the animals formerly of the western mainland
were able to come far eastward. The animals from Lynndyl which
are intergrades between lenis, an eastern mainland form, and cen-
tralis and aureiventris which are western mainland forms of Lake
Bonneville lend support to this hpyothesis.
Specimens examined. — Total, 26, distributed as follows: Millard County: Lynndyl, 4,796
ft., 21. Juab County: U. B. (= Yuba) Darn, 5,000 ft., 1. Sevier County: Salina, 4,575
ft., 1; Richfield, 5,308 ft., 3 (U. S. N. M.).
Thomomys bottae levidensis Goldman
Thomomys bottae levidensis Goldman, Proc. Biol. Soc. Washington,
55:76, June 25, 1942.
Thomomys perpallidits aureus Bailey, N. Amer. Fauna, 39:75, No-
vember 15, 1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12):100, June, 1927.
Type.— Male, adult, skin and skull, No. 191962, U. S. National Museum
(Merriam Collection); Manti, 5,500 ft., Sanpete County, Utah; December 6,
1888; collected by Vernon Bailey; original number 427 (after Goldman, type
not seen).
Durrant — Pocket Gophers of Utah 55
Range. — San Pitch River Valley, Sanpete County, Utah.
Diagnosis. — Size small (see measurements). Color: Upper parts and sides
Cinnamon Buff, finely mixed with black along median line of back; under-
parts Pinkish Buff; nose, cheeks and chin grayish black; postauricular patches
fairly large and grayish black; front and hind feet white (examples from type
series badly stained) ; tail light buff but apparently white distally (the color
of these specimens has apparently changed with age). Skull: Small, fairly
robust; basilar length short; zygomatic arches weak, but widely spreading;
tympanic bullae small; nasals short and simple distally; ventral margin of
jugals convex dorsally; extension of premaxillae posterior to nasals relatively
a? well as actually long; posterior tongues of premaxillae relatively wide.
Comparisons. — Topotypes of levidensis differ from those of Tho-
momys bottae absonus as follows: Size smaller. Color: Lighter
throughout. Skull: Shorter, weaker and less ridged and angular,
but relatively wider.
Compared with topotypes of Thomomys bottae albicaudatus, lev-
idensis differs as follows: Size smaller in every measurement taken.
Color: Markedly lighter throughout. Skull: Smaller in every
measurement taken; width relatively greater; skull smooth, weak
and nonangular as opposed to ridged, robust and angular.
For comparisons with Thomomys bottae lenis and contractus see
accounts of those forms.
Remarks. — The range here ascribed to levidensis is the San Pitch
River Valley, which gradually merges southward into the Sevier
River Valley. The latter valley in this area is inhabited by pocket
gophers that belong to another subspecies, lenis. Nephi Valley to
the west of San Pitch River Valley is inhabited by animals belong-
ing to the subspecies albicaudatus. No known specimens show in-
tergradation between lenis and levidensis, but intergradation be-
tween lenis and albicaudatus is noted in the Nephi Valley animals
(see account of albicaudatus). Superficially levidensis resembles
absonus in size and color, but the skulls closely resemble those of
albicaudatus, except for size in which they are smaller in all mea-
surements. T. b. albicaudatus is the most variable subspecies of
T. bottae occurring in Utah, and additional material from the Se-
vier River Valley between San Pitch River Valley and Nephi Val-
ley may show levidensis to be only a local variant of the highly
variable subspecies, albicaudatus.
Specimens examined. — Total, 6, from the type locality.
56 University of Kansas Publs., Mus. Nat. Hist.
Thomomys bottae osgoodi Goldman
Thomomys perpallidus osgoodi Goldman, Joum. Washington Acad.
Sci.. 21:424, October 19, 1931.
Thomomys bottae osgoodi Goldman, Proc. Biol. Soc. Washington,
48:156; October 31, 1935.
Thomomys perpallidus aureus Bailey, N. Amer. Fauna, 39:75, No-
vember 15, 1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April., 1922;
Bull. Univ. Utah, 17 (No. 12):100, June, 1927.
Type.— Male, adult, skin and skull, No. 158530, U. S. National Museum
(Biological Surveys Collection); Hanksville, Wayne County, Utah; October
20, 1908; collected by W. H. Osgood; original number 3701 (after Goldman,
type not seen).
Range. — Eastern Utah in the valleys of the drainage of the San Rafael,
Dirty Devil and Price rivers.
Diagnosis. — Size medium (see measurements). Color: Upper parts near
(e) Pale Ochraceous Buff, definitely yellow in appearance; sides Pale Ochra-
ceous Buff; entire underparts white, with a wash of Light Buff in the pectoral
and inguinal regions; top of head, nose, cheeks, and chin dusky; postauricular
patches grayish black; front feet, hind feet and distal part of tail white.
Skull: Fairly robust but narrow; zygomatic arches concave medially in mid-
jugal region; skull moderately convex dorsally, due to swelling in region of
base of rostrum; lambdoidal suture situated well ahead of posterior margin
of skull, with supraoccipital forming a side shelf at posterior part of skull ;
interpterygoid space narrowly V-shaped; tympanic bullae well inflated ven-
trally; basioccipital short; nasals rounded posteriorly; molariform teeth large.
Comparisons. — Topotypes of osgoodi differ from those of Thomo-
mys bottae absonus as follows: Size generally smaller. Color:
Lighter throughout, more yellowish in appearance as opposed to
buffy. Skull: Smaller in all measurements, except length of nasals,
mastoid breadth, and alveolar length of upper molar series which
are larger; rostrum shorter but relatively wider; zygomatic arches
more robust and concave medially; palate wider; supraoccipital
more bulging posteriorly; tympanic bullae more inflated ventrally;
molariform teeth larger.
For comparisons with Thomomys bottae aureus and T. b. dis-
similis see accounts of those forms.
Remarks. — The animals here referred to osgoodi are remarkably
uniform in color, but vary widely in cranial details. Specimens
from Carbon County are not typical and when more material be-
comes available it may prove that these animals from the northern
part of the range of osgoodi will merit separation and naming. The
specimens from Emery County are not typical but resemble osgoodi
more than do the animals from Carbon County.
The range here ascribed to osgoodi is in that part of the eastern
Utah desert that is bounded on the east by the Green and Colorado
Durrant — Pocket Gophers of Utah 57
rivers, on the west by the high mountains of central Utah, on the
north by the Book Cliffs and on the south by the Dirty Devil River.
This area is an uninviting wasteland in which there are relatively
few roads and little water. In addition, it is greatly cut up by
washes and gullies which contain water only during a few weeks of
the year. The continuation of this area of wasteland southward
beyond the Dirty Devil River is inhabited by pocket gophers be-
longing to the subspecies absonus. If specimens were available they
would undoubtedly show intergradation to exist between osgoodi
and absonus.
Specimens examined. — Total, 14, distributed as follows: Carbon County: y2 rni. N
Spring Glen, 6,150 ft., 2; Spring Glen, 6,200 ft., 2; 2 mi. E Spring Glen, 6,200 ft., 1.
Emery County: Price River, 2 mi. SE Woodside, 4,600 ft., 2 (C. M.); Green River, 4,080
ft., 5 (M. V. Z.). Wayne County: Hanksville, 2 (U. S. N. M.).
Thomomys bottae howelli Goldman
Thomomys bottae howelli Goldman, Journ. Washington Acad. S'ci.,
26:116, March 15, 1936.
Type. — Female, adult, skin and skull, No. 25684, U. S. National Museum
(Biological Surveys Collection) ; Grand Junction, 4,600 ft., Mesa County, Colo-
rado; November 7, 1895; collected by A. H. Howell; original number 493
(after Goldman, type not seen).
Range. — In the valleys of eastern Utah, east of the Green River and north
of the Colorado River.
Diagnosis and Comparisons. — Inasmuch as there is but one specimen, the
holotype known, and as it was impossible to study it, the following diagnoses
and comparisons are from Goldman, (1936:116).
"General characters. — A rather large, pallid subspecies with a broad, flat-
tened cranium. Similar to the palest specimens of Thomomys bottae aureus
of the San Juan River Valley, southeastern Utah, in color, but under parts
more thinly overlaid with buffy white, and cranial characters, especially the
broad, flat braincase, distinctive. Approaching Thomomys bottae osgoodi of
the Fremont River Valley, Utah, in color, but much larger and skull widely
different.
''Color. — Type (winter pelage) : Upper parts in general between tilleul
buff and pale olive buff (Ridgway 1912), somewhat darkened on head by a
mixture of cinnamon buff and brown; a few inconspicuous dusky-tipped hairs
along median line of back; muzzle dusky; ears and postauricular spots deep,
contrasting black; underparts thinly overlaid with buffy white, the hairs be-
coming pure white to roots on inguinal region; thighs pure white to roots all
around; feet white; tail buffy whitish, slightly paler below than above.
"Skull. — Similar in general to that of T. b. aureus, but braincase conspicu-
ously broader and flatter; zygomata more widely spreading; nasals shorter;
premaxillae more attenuate posteriorly; interparietal larger; audital bullae
more rounded and fully inflated anteriorly; incisors short, as in aureus, but
less strongly recurved. Compared with that of T. b. osgoodi the skull is much
larger, with flatter braincase, shorter nasals, and posteriorly narrower pre-
maxillae."
58 University of Kansas Publs., Mtjs. Nat. Hist.
Remarks. — Six specimens, in the Carnegie Museum from 10 miles
north of Moab, Grand County, Utah, were available for this study.
They are not typical of howelli as it is diagnosed by Goldman (loc.
tit.). They appear to be intergrades between howelli and osgoodi
in cranial characters, but more closely resemble howelli, particularly
in the flat, widened, low braincase. In color, some specimens seem
to intergrade toward aureus.
The range ascribed to this form in Utah appears to be one of
the most natural ones within the state since it is bounded by the
Green and Colorado rivers which have formed deep rocky gorges
in this region.
Specimens examined. — Total, 6, as follows: Grand County: 10 mi. N Moab, 6 (C. M.).
Thomomys bottae wahwahensis Durrant
Thomomys bottae. wahvxihensis Durrant, Bull. Univ. Utah, 28 (No.
4):4, August 18, 1937.
Type. — Male, adult, skin and skull, No. 1750, Museum of Zoology, Uni-
versity of Utah, Wah Wah Springs, 30 mi. W Milford, 6,500 ft., Beaver County,
Utah; July 22, 1936; collected by S. D. Durrant; original number 989.
Range. — Westcentral Utah, in Wah Wah Mountains, and Pine Valley to
the west of these mountains.
Diagnosis. — Size medium (see measurements). Color: Upper parts Pink-
ish Buff; underparts Pale Pinkish Buff with considerable admixture of gray;
inguinal and pectoral regions Pale Pinkish Buff; nose and cheeks grayish
black; postauricular patches small and black; front feet, hind feet and distal
one-third to one-half of tail white. Skull: Flat dorsoventrally ; rostrum
short and wide; premaxillae broad and heavy; nasals short and straight, with
no arching as viewed laterally; tympanic bullae small; space enclosed within
zygomatic arches short antero-posteriorly; alveolar length of upper molar
series short; molariform teeth small.
Comparisons. — From topotypes of Thomomys bottae centralis,
wahwahensis differs as follows: Size smaller in every measure-
ment taken. Color: Lighter, Pinkish Buff as opposed to Cinnamon
Buff. Skull: Rostrum wider, shorter and more nearly flat; nasals
straight as opposed to moderately convex; tympanic bullae smaller
and less inflated ventrally; zygomatic arches more widely spread-
ing and angular; molariform teeth smaller; extension of premaxillae
posterior to nasals less.
From topotypes of Thomomys bottae albicaudatus, wahwahensis
differs as follows: Hind foot shorter. Color: Lighter throughout,
Pinkish Buff as opposed to (13 " " n) Black. Skull: Smaller and
more nearly flat; rostrum shorter, wider and more nearly flat;
nasals straight as opposed to convex; zygomatic breadth less but
mastoid breadth greater; tympanic bullae smaller, and less inflated
Durrant — Pocket Gophers of Utah 59
ventrally; extension of premaxillae posterior to nasals less; molari-
form teeth smaller.
From topotypes of Thomomys bottae aureiventris, wahwahensis
differs in the following features: Size smaller; hind foot shorter.
Color: Lighter throughout, no "gold" on underparts. Skull:
Smaller in nearly every measurement taken; rostrum shorter and
relatively wider; zygomatic arches more angular and relatively
more widely spreading; nasals shorter and more nearly flat; thick-
ening at union of jugal and zygomatic process of maxilla less; inter-
pterygoid space V-shaped as opposed to lyre-shaped; tympanic
bullae much smaller, and less inflated ventrally; molariform teeth
much smaller.
Topotypes of wahwahensis can be easily distinguished from those
of Thomomys bottae tivius by their markedly larger size in every
measurement taken, lighter color, and larger, more robust and more
nearly flat skull.
For comparisons of wahwahensis with Thomomys bottae sevieri,
robustus, bonnevillei and convexus see comparisons under those
forms.
Among the named races of Thomomys bottae, wahwahensis defi-
nitely has its affinities with planirostris from Zion National Park.
Both possess flat skulls with wide, short rostra. It differs from the
latter in: Size smaller in every measurement taken. Color: Lighter
throughout. Skulls: Nasals and rostrum shorter and more nearly
flat; tympanic bullae markedly smaller; alveolar length of upper
molar series shorter; molariform teeth markedly smaller and
weaker.
Remarks. — Wah Wah Springs, the type locality of wahwahensis,
are on the summit of a low pass in the Wah Wah Mountains in the
desert of west central Utah. The surrounding valleys, for many
miles, as far as my investigations show, are not inhabited by pocket
gophers, except the Desert Range Experiment Station of the United
States Forest Service in Pine Valley to the west of these mountains.
There, pocket gophers were obtained which are intergrades between
centralis and wahwahensis. In five out of seven characters investi-
gated these gophers resemble wahwahensis, to which they are here
referred. Study of the topography reveals the probable means by
which the animals reached this valley. The long axis of the Wah
Wah Mountains is north and south, but a westward arm forms the
northern boundary of Pine Valley. Around springs in this west-
ward projecting arm workings of pocket gophers were found. With
60 University of Kansas Publs., Mus. Nat. Hist.
the development of water at the Desert Range Experiment Station,
and subsequent improvement of forage, these animals probably
came down into the valley from the springs to the north.
The terrain between the Desert Range Experiment Station in
Pine Valley and Snake Creek (where centralis occurs) to the west
is not inhabited by pocket gophers at present. This area, however,
forms part of the southwest mainland of Pleistocene Lake Bonne-
ville, which mainland in times past was probably suitable for
pocket gophers. Since the close of the Pleistocene, aridity has ren-
dered most of it unfit for pocket gophers, and they remain only in
isolated areas where suitable environments still persist.
Specimens examined. — Total, 18, distributed as follows: Millard County: Desert Range
Experiment Station, United States Forest Service, Sec. 9, T. 25 S, R. 17 W, Salt Lake Base
Meridian, 6. Beaver County: Wah Wah Springs, Wah Wah Mountains, 30 mi. W Milford,
6,500 ft., 12 (2, M. V. Z.).
Thomomys bottae dissimilis Goldman
Thomomys perpallidus dissimilis Goldman, Journ. Washington Acad.
Sci., 21:425, October 19, 1931.
Thomomys bottae dissimilis Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935.
Thomomys perpallidus aureus Bailey, N. Amer. Fauna 39:75, Novem-
ber 15, 1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12): 100, June, 1927.
Type.— Female, adult, skin and skull, No. 158526, U. S. National Museum
(Biological Surveys Collection) ; E slope Mount Ellen, Henry Mountains,
8,000 ft., Garfield County, Utah; October 15, 1908; collected by W. H. Osgood;
original number 3677 (after Goldman, type not seen).
Range. — Known only from the type locality.
Diagnosis.— Size small (see measurements) . Color : Upper parts Light Buff,
grading over sides to nearly white on underparts; underparts lightly washed
with Pale Buff, more marked in inguinal and pectoral regions; postauricular
patches grayish black; nose, chin, cheeks and top of head dusky; front feet,
hind feet and distal half of tail white. Skull: Small and weak; zygomatic
arches long, but lying close to skull, giving it a slender appearance; supra-
occipital markedly projecting posteriorly from lambdoidal suture; rostrum
relatively long and narrow; nasals long; tympanic bullae well inflated ven-
trally, with a median ventral ridge; pterygoid hamulae weak; interpterygoid
space narrowly V-shaped; upper incisors short and light in color; molariform
teeth relatively large.
Comparisons.— Comparison of one topotype of dissimilis with
topotypes of Thomomys bottae aureus shows it to differ as follows:
Size smaller throughout. Color: Lighter dorsally and on sides, pale
buff as contrasted with rich ochraceous; underparts more buffy.
Skull: Smaller in every measurement taken; zygomatic arches
markedly less widely spreading; braincase narrower and more
Durrant — Pocket Gophers of Utah 61
vaulted; tympanic bullae with a median ventral ridge as opposed
to smooth; pterygoid hamulae slenderer; interpterygoid space nar-
rowly V-shaped as opposed to U-shaped; upper incisors smaller and
lighter in color.
Compared with topotypes of Thomomys bottae absonus, dis-
similis differs in the following features : Size smaller in every meas-
urement taken. Color: Lighter throughout. Skull: Smaller in
every measurement taken, except alveolar length of upper molar
series which is greater; skull narrower and weaker; zygomatic arches
weaker and less widely spreading; tympanic bullae more ridged on
ventral surface and shorter (more rounded) in antero-posterior
measurement; upper incisors shorter and narrower; molariform teeth
larger.
Thomomys bottae dissimilis resembles T. b. osgoodi more than
any other subspecies but differs in : Size smaller throughout. Color :
Slightly darker dorsally. Skull: Smaller in every measurement
taken, and slenderer; rostrum relatively longer; zygomatic arches
weaker, and less widely spreading, more converging anteriorly;
tympanic bullae less rounded, more ridged medioventrally ; upper
incisors shorter but narrower; molariform teeth smaller.
Remarks. — The Henry Mountains, in eastern Garfield County,
are in the Colorado River drainage. The surrounding country is
desertlike and cut by gullies and washes with sheer escarpments
and precipitous draws. The type locality of dissimilis is possibly
in an isolated area. Only three specimens were available to Gold-
man when he named dissimilis. He commented on the close re-
semblance to osgoodi which inhabits the country to the north. I
have examined only one of the three specimens available to Gold-
man. Although I can see the characters that he mentioned, I am
not fully convinced that dissimilis is separable from osgoodi. Two
specimens from Escalante, Garfield County, are referred to absonus,
but they show intergradation with dissimilis.
Specimens examined. — One (U. S. N. M.) from E slope Mount Ellen, Henry Mountains,
8,000 ft., Garfield County.
Thomomys bottae aureus Allen
Thomomys aureus Allen, Bull. Amer. Mus. Nat. Hist., 5:49, April 28,
1893.
Thomomys bottae aureus Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935; Benson, Univ. California Publ. Zobl, 40:450,
December 31, 1935.
Thomomys fidvus aureus Goldman, Journ. Washington Acad. Sci.,
21:417, October 19, 1931; Joum. Washington Acad. Sci., 23:464, October
15, 1933.
62 University of Kansas Publs., Mus. Nat. Hist.
Thomomys perpallidus aureus Bailey, N. Amer. Fauua, 39:74, No-
vember 15, 1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12) :100, June, 1927.
Type.— No. _|||.. American Museum of Natural History; Bluff City, San
Juan County, Utah; May 12, 1892; collected by Charles P. Rowley (after
Allen, type not seen).
Range. — All of San Juan County (except extreme southwestern part) and
Grand County east of the Colorado River.
Diagnosis. — Size large (see measurements). Color: Upper parts Cinnamon
Buff, lighter on sides; underparts generally white, or if colored at all with
only a faint wash of Light Buff; nose and chin blackish gray; top of head
blackish due to admixture of black hairs; postauricular patches small and
dusky; front feet and hind feet white. Skull: Long, narrow but massive;
zygomatic arches not widely spreading, but heavy; jugals thick, union of
jugals and zygomatic processes of maxillae thickened; rostrum long but wide;
top of rostrum convex in lateral view; ascending processes of premaxillae wide
and heavy; nasals thin proximally; braincase long and narrow; tympanic
bullae well inflated ventrally; alveolar length of upper molar series long;
molars large; pterygoid hamulae heavy; interpterygoid space U-shaped; palate
arched; upper incisors long and wide.
Comparisons. — Compared with topotypes of Thomomys bottae
osgoodi, aureus differs as follows: Size larger in every measure-
ment taken, except tail which is shorter. Color: Darker throughout
except on ventral surface which is lighter. Skull: Larger, longer
and wider; nasals longer; rostrum wider and longer; zygomatic
arches more nearly straight and heavier; ascending processes of
premaxillae wider; basioccipital longer; interpterygoid space U-
shaped as opposed to V-shaped; tympanic bullae larger; upper in-
cisors longer, wider; molars larger.
Topotypical specimens of aureus can be distinguished from those
of Thomomys bottae dissimilis by: Size larger throughout. Color:
A trifle darker on dorsal surface. Skull: Larger in every meas-
urement taken; zygomatic arches heavier and more nearly straight;
tympanic bullae larger and more inflated ventrally; interpterygoid
space U-shaped as opposed to V-shaped; alveolar length of upper
molar series longer; molars larger; upper incisors longer and wider.
Topotypes of aureus differ from those of Thomomys bottae ab-
sonus as follows: Size larger in every measurement taken. Color:
Darker dorsally, Light Ochraceous as opposed to Cinnamon Buff;
due to admixture of gray, absonus has more of a grayish cast.
Skull: Larger in every measurement taken, longer, narrower and
more compact; zygomatic arches heavier; ascending processes of
premaxillae wider; jugals heavier; tympanic bullae larger; inter-
Durrant — Pocket Gophers of Utah 63
pterygoid space U-shaped rather than V-shaped; upper incisors
longer and wider; molars larger.
From topotypes of Thomomys bottae planirostris, aureus can be
distinguished as follows: Size larger; tail shorter. Color: Lighter
throughout. Skull: Larger in every measurement taken except
zygomatic breadth, extension of premaxillae posterior to nasals, and
length of upper molariform series which are less; rostrum longer,
wider and more convex; nasals slightly arched rather than straight;
depression absent rather than present in posterior region of nasals ;
zygomatic arches not so widely spreading, but equally heavy.
For comparisons with Thomomys bottae alexandrae, see accounts
under that form.
Remarks. — Topotypes of aureus are among the largest pocket
gophers in the state. They are exceeded in total length only by T. b.
lenis and are approached by T. b. aureiventris and T. b. pla?iiros-
tris. On the average they have the longest hind foot, body and ear.
The length of the skull is second only to that of lenis as also is the
length and breadth of the rostrum relative to the basilar length.
From the time of the original description of aureus in 1893 until
1930, all light colored gophers from Utah were referred to that
form. Barnes (1927:100) gives the range of aureus as extending
completely across southern Utah and on the west and east sides as
far north as central Utah. Since 1930, forms named by E. R. Hall,
W. H. Burt, E. A. Goldman and the writer have restricted the range
of aureus in Utah to that part of the state east of the Colorado
River.
Specimens examined. — Total, 22, as follows: San Juan Count)/: Bluff, 3,300 ft., 22 (15,
M. V. Z.).
Thomomys bottae birdseyei Goldman
Thmaomys bottae birdseyei Goldman. Proc. Biol. Soc. Washington,
50:134, September 10, 1937.
Thomomys perpallidus aureus Bailey, N. Amer. Fauna, 39:75, No-
vember 15. 1915; Barnes. Bull. Univ. Utah. 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12):100, June, 1927.
Type.— Male, adult skin and skull, No. 1G1G54, U. S. National Museum
(Biological Surveys Collection) ; Pine Valley Mountains, 5 mi. E Pine Valley,
8,300 ft., Washington County, Utah; April 10. 1909; collected by Clarence
Birdseye; original number 861 (after Goldman, type not seen).
Range. — High mountains and plateaus of southwestern Utah.
Diagnosis. — Size medium (see measurements). Color: Upper parts between
Cinnamon and Sayal Brown, finely mixed with black in median dorsal region,
grading over sides and flanks to Cinnamon on underparts; front feet, hind
feet, and distal part of tail white; postauricular patches, chin, cheeks and top
64 University of Kansas Publs., Mus. Nat. Hist.
of head grayish black. Skull: Depressed along median line of frontals and
posterior ends of nasals; region of nasofrontal suture concave ventrally;
zygomatic arches heavy and widely spreading, widest posteriorly; posterior
ends of nasals straight, tending to be somewhat rounded in some specimens;
extension of premaxillae posterior to nasals moderate; tympanic bullae mod-
erately inflated ventrally; basioccipital wide; interpterygoid space widely
V-shaped.
Comparisons. — Topotypes of birdseyei differ from near topotypes
of Thomomys bottae virgineus, from Beaverdam Wash as follows:
Size larger; tail and hind foot longer. Color: Darker throughout,
between Cinnamon and Sayal Brown as opposed to Cinnamon Buff.
Skull: Larger in every measurement taken except extension of
premaxillae posterior to nasals, and length and width of rostrum
which are less; skull more depressed in nasofrontal region; zygo-
matic arches more widely spreading; zygomatic processes of squa-
mosals shorter; pterygoid hamulae longer; tympanic bullae smaller
and less inflated ventrally.
Among named races of T. bottae, birdseyei most closely resem-
bles trumbullensis in size, but differs as follows: Hind foot and
tail longer. Color: Lighter throughout; postauricular patches
smaller and lighter. Skull: Larger; mastoid breadth less; zygo-
matic arches wider and more widely spreading posteriorly; median
frontal depression more marked; extension of premaxillae posterior
to nasals greater; tympanic bullae less inflated ventrally; molari-
form teeth larger.
For comparisons with Thomomys bottae planirostris see account
of that form.
Remarks. — T. b. birdseyei is apparently endemic to the moun-
tainous area of southwestern Utah in Washington and Iron coun-
ties. It intergracles with virgineus and with planirostris as de-
scribed in the account of the latter.
Specimens examined. — Total, 8, distributed as follows: Washington County: Pine Valley,
1 (TJ. S. N. M.); Pine Valley Mountains, 5 mi. E Pine Valley, 8,300 ft., 3 (U. S. N. M.);
Pine Valley campground, 6,800 ft., 1 (R. H.) ; % mi. E town of Pine Valley, 6,500 ft., 3
(R. H.).
Additional records. — Washington County: Hebron, 1; Mountain Meadows, 2 (Bailey
1915:75).
Thomomys bottae virgineus Goldman
Thomomys bottae virgineus Goldman, Proc. Biol. Soc. Washington.
50:133, September 10, 1937.
Type.— Male, adult, skin and skull, No. 262016, U. S. National Museum
(Biological Surveys Collection) ; Beaverdam Creek, near confluence with Vir-
gin River, Littlefield, 1,500 ft., Mohave County, Arizona; October 16, 1936;
Durrant — Pocket Gophers of Utah 65
collected by Luther C. Goldman; original number 67 (after Goldman, type
not seen).
Range. — Extreme southwestern Utah, in Beaverdam Wash, Washington
County, Utah.
Diagnosis. — Size medium (see measurements). Color: Upper parts Cin-
namon Buff, finely mixed with black; sides and flanks Pinkish Buff; under-
parts Pale Pinkish Buff; front feet, hind feet, and distal part of tail white;
nose, cheeks, chin and top of head grayish black. Skull: Robust, with mod-
erately wide zygomatic arches; z3'gomatic processes of maxillae wide; zygo-
matic processes of squamosals long; jugals concave laterally, giving the
zygomatic arches the appearance of double bowing; nasals long; extension
of premaxillae posterior to nasals long; tympanic bullae well inflated ven-
trally; pterygoid hamulae heavy; interpterygoid space widely V-shaped;
molariform teeth large.
Comparisons. — For comparisons of virgineus with Thomomys
bottae planirostris and T. b. birdseyei see accounts under those
forms.
Topotypical specimens of virgineus can be distinguished from
those of Thomomys bottae trumbullensis as follows: Size smaller.
Color: Lighter throughout. Skull: Zygomatic arches less widely
spreading; jugals more bowed medially; zygomatic processes of
squamosals longer; extension of premaxillae posterior to nasals
greater; tympanic bullae larger and more inflated ventrally; molari-
form teeth larger.
Compared with topotypes of Thomomys bottae centralis, vir-
gineus differs in: Size smaller; tail shorter; hind foot smaller.
Color: Deeper Cinnamon Buff, thus darker in overall appearance.
Skull: Smaller, but relatively wider; zygomatic processes of maxil-
lae heavier; region of maxillo-jugal sutures thicker; jugals more
concave laterally; tympanic bullae more inflated ventrally; molari-
form teeth larger.
Remarks. — This pocket gopher occupies practically the same
range in Utah as the large kangaroo rat Dipodomys deserti deserti
Stephens. Both are found in the Beaverdam Wash. The type lo-
cality of virgineus is but a short distance down the Beaverdam
Creek at Littlefield, Arizona. It intergrades with birdseyei, the
mountain form to the north and east (see remarks under birdseyei).
There are evidences of intergradation with planirostris of the Vir-
gin River Valley above the narrows of the Virgin River where it cuts
through the Beaverdam Mountains (see the discussion under
planirostris). There are intergradational tendencies exhibited to-
5—2786
66 University of Kansas Publs., Mus. Nat. Hist.
wards centralis in some specimens. Some of the animals are prac-
tically indistinguishable in color and there are intergrading cranial
characters in the nasals, zygomatic arches and tympanic bullae.
Specimens examined. — Total, 20, distributed as follows: Washington County: Beaverdam
Wash, 8 mi. N Utah-Arizona border, 7; Beaverdam Wash, 5 mi. N Utah-Arizona border,
2,600 ft., 13.
Thomomys bottae planirostris Burt
Thomomys perpallidus planirostris Burt, Proc. Biol. Soc. Washington,
44:38, May 8, 1931.
Thomomys bottae planirostris Hall and Davis, Proc. Biol. Soc. Wash-
ington, 47:52, February 9, 1934; Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935; Presnall, Zion-Bryce Mus. Bull., 2:14, Janu-
ary, 1938; Long, Journ. Mamm, 21:176, May 14, 1940.
Thomomys perpallidus aureus Bailey, N. Amer. Fauna, 39:75, No-
vember 15, 1915; Barnes. Bull. Univ. Utah, 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12): 100, June, 1927; Woodbury, Ecological
Monographs, 3:193, April, 1933.
Thomomys bottae centralis Hall and Davis, Proc. Biol. Soc. Washing-
ton, 47:52/ February 9, 1934; Presnall, Zion-Bryce Mus. Bull., 2:14,
January, 1938.
Thomomys perpallidus centralis Hall, Univ. California Publ. Zool.,
23:445, July 8, 1930.
Thomomys bottae nicholi Goldman, Journ. Washington Acad. Sci.,
28:337, July 15, 1938, type from Shivwits Plateau, 20 mi. S Wolf Hole
(road to Parashonts), 5,000 ft., Mohave County, Arizona; Hardy, Eco-
logical Monographs, 15:98, January, 1945.
Thomomys bottae trumbidlensis Hall and Davis, Proc. Biol. Soc.
Washington, 47:52, February 9, 1934.
Type.— Male, adult, skin and skull, No. 8395, Collection of Donald R.
Dickey; Zion National Park, Washington County, Utah; May 4, 1920; col-
lected by A. Brazier Howell; original number 2184 (after Burt, type not seen).
Range. — Valley of the Virgin River from Zion National Park west to the
Beaverdam Mountains.
Diagnosis. — Size large (see measurements); tail long. Color: Upper parts
Sayal Brown ; underparts between Vinaceous Cinnamon and Cinnamon, grading
to Pinkish Cinnamon in some specimens; nose, chin, cheeks, postauricular
patches, and top of head grayish black; front feet and hind feet white; tail
Pinkish Buff, with distal third white. Skull: Massive and ridged; nasals
straight and flat, simple distally; dorsal surface of rostrum slightly concave
at proximal end of nasals; zygomatic arches widely spreading, widest posteri-
orly; zygomatic processes of maxillae heavy; premaxillae broad and extend-
ing far beyond posterior end of nasals; rostrum wide and heavy; palate slightly
arched; pterygoid hamulae heavy; interpterygoid space V-shaped; tympanic
bullae moderately inflated ventrally, somewhat compressed laterally; upper
incisors long and heavy; molariform teeth large.
Comparisons. — Compared with topotypes of Thomomys bottae
birdseyei, planirostris differs as follows: Size larger, except total
length which averages slightly less in females. Color: Lighter
throughout. Skull: Larger in every measurement taken; more
Durrant — Pocket Gophers of Utah 67
massive; rostrum wider, longer and more nearly flat; nasals straight
and not inflated dorsally on distal end; premaxillae wider at pos-
terior ends; extension of premaxillae posterior to nasals greater;
zygomatic arches heavier, especially the zygomatic processes of
the maxillae; posterior ends of nasals more nearly truncate as op-
posed to generally rounded; tympanic bullae more nearly flat and
relatively smaller; upper incisors longer and heavier; interptery-
goid space more narrowly V-shaped; molariform teeth much heavier.
Topotypes of planirostris differ from near topotypes of Thomo-
mys bottae virgineus as follows: Size larger; tail and hind foot
longer. Color: Slightly darker dorsally, but markedly darker ven-
trally; postauricular patches smaller and lighter. Skull: Larger
in every measurement taken ; skull more massive ; nasals flat, neither
arched nor swollen distally; rostrum wider; nasofrontal region flat-
tened or concave as opposed to convex; premaxillae relatively nar-
rower; zygomatic arches heavier, especially in the processes of the
maxillae; tympanic bullae smaller and less inflated ventrally; in-
terpterygoid space generally more narrowly V-shaped; upper in-
cisors longer and heavier; molariform teeth larger.
From topotypes of Thomomys bottae trumbullensis, planirostris
differs in: Size larger throughout; tail longer. Color: Much lighter
throughout. Skull: More convex dorsally; rostrum wider and more
depressed distally; extension of premaxillae posterior to nasals
greater; zygomatic arches shorter, and not as widely spreading pos-
teriorly; interpterygoid space more narrowly V-shaped; tympanic
bullae smaller; upper incisors wider and longer; molariform teeth
larger.
Topotypes of planirostris can be easily distinguished from those
of Thomomys bottae absonus by darker color throughout and mark-
edly larger size.
Remarks. — From the synonomy at the beginning of this account
one may note that the animals here ascribed to this subspecies have
had nearly as many subspecific names applied to them as there have
been investigators who have written about them. Although each
of the previous writers had but a small amount of material upon
which to base his opinion, the diversity of opinion as to subspe-
cific status bespeaks the instability of these animals. The present
study is based upon eighty animals including additional compara-
tive material.
All animals from Zion National Park have the characters pointed
out by Burt (1931:38) in his description of this form. Farther
68 University of Kansas Publs., Mus. Nat. Hist.
down the Virgin River Valley towards St. George, however, some
very perplexing problems of intergradation are encountered. St.
George and environs may correctly be thought of as a "melting pot."
Each of the fifty-seven animals studied from this region is an in-
tergrade; some specimens combine the characters of three sub-
species.
As may be seen on the distribution map, three different subspecies
of Thomomys bottae occur in Washington County. Down the river,
below St. George, the race virgineus inhabits the Virgin River
Valley below the narrows of the Beaverdam Mountains. Because
these narrows are filled with water from wall to wall during periods
of high runoff, they form an effective barrier at present to migra-
tion of pocket gophers. The mountains to the north of St. George
are inhabited by the dark form, birdseyei. The type locality of
planirostris is on the middle reaches of the Virgin River, in Zion
National Park. In addition Mount Trumbull to the south, in north-
ern Arizona, is the locality of another subspecies, trumbullensis.
Unquestionably the easiest route of migration into the St. George
area is down the Virgin River from Zion National Park ; no barrier
to gophers occurs between the Park and St. George. Although the
animals from St. George are all intergrades, the majority of their
affinities as would be expected are with planirostris from Zion Na-
tional Park. The river itself is not an impassable barrier for
gophers to the north and south of it, since this stream frequently
changes its course, and often nearly dries up. The Virgin River
Valley in Zion National Park is in the bottom of a relatively deep,
narrow canyon which has sheer rock escarpments. The upper
reaches of the river are inhabited by pocket gophers of another
species, Thomomys talpoides.
Two specimens from St. George, north of the Virgin River, were
identified as centralis by Hall and Davis (1934:52), but were stated
to be intergrades between centralis, trumbullensis and planirostris.
Goldman (1938:338) referred twelve specimens from St. George
to nicholi, but stated that they intergraded with planirostris.
Twenty-six other specimens from three miles southwest of St.
George on the west side of Santa Clara Creek, about one-half mile
above its confluence with the Virgin River and on its north side,
like the topotypes of planirostris were taken in May and have com-
plete, fresh summer pelage. With the exception of two specimens
which show the ventral color of virgineus, these animals are indis-
tinguishable in color from the topotypes of planirostris. A study
Durrant — Pocket Gophers of Utah 69
of eleven measurements of the males of this series yield the follow-
ing data: Like planirostris in four measurements, birdseyei in one,
virgineus in one; intergrade between planirostris and birdseyei in
two, planirostris and virgineus in two and birdseyei and virgineus
in one. Corresponding measurements of the females show the ani-
mals to be: Like planirostris in four measurements, birdseyei in
one, virgineus in two; intergrade between planirostris and birds-
eyei in two, planirostris and virgineus in one and birdseyei and
virgineus in one. In eight of eleven measurements the males either
are like planirostris or intergrade towards it, and the females are
similarly allied to planirostris in seven out of eleven measurements.
In none of the measurements was either sex referable to trumbullen-
sis.
Intergradation was noted in still other cranial details. In the
heavy, relatively straight zygomatic arches, a majority of the
skulls resemble those of planirostris, although some show the elon-
gated zygomatic processes of the squamosals that are characteristic
of virgineus. Some skulls show a tendency toward birdseyei in the
widely spreading posterior regions of the zygomatic arches. The
nasals for the most part are as in planirostris. Intergradation
between all three subspecies is shown in the extension of the pre-
maxillae posterior to the nasals. Some skulls show the lateral con-
cavity of the jugals which is characteristic of virgineus. The tym-
panic bullae are variable but on the average are intermediate be-
tween those of planirostris and birdseyei, but more as in the latter.
The size of the pterygoid hamulae is like that of planirostris, but
the shape of the interpterygoid space is more like that of birds-
eyei. The size of the molariform teeth is as in birdseyei. The in-
cisors are intermediate between those of planirostris and birdseyei,
but more like those of birdseyei.
Eighteen specimens from St. George and its environs, on the
north side of the Virgin River, agree with the twenty-six specimens
just described, except that they show more evidence of intergrada-
tion with birdseyei in slightly darker color, length of hind foot,
length of nasals and alveolar length of the upper molar series.
One specimen from three miles south, two from two miles south-
west, another from four miles southeast of St. George, and four
immature animals from Short Creek Road south of the town of
Virgin, all on the south side of the Virgin River, are darker than
topotypes of planirostris and show intergradation with trumbul-
lensis to the south. In size they are likewise closer to the latter
70 University of Kansas Publs., Mus. Nat. Hist.
race. They intergrade with trumbullensis in the size and shape of
the zygomatic arches and tympanic bullae. In the majority of
cranial details, however, they are like planirostris to which they are
here referred.
One specimen, a skin only, from Danish Ranch, 5 miles north-
west of Leeds, north of the Virgin River is an intergrade in size and
color between birdseyei and planirostris, but referable to the latter.
Three specimens from the East Entrance, and three from near the
east entrance to Zion National Park are much darker than topo-
types of planirostris. All of these animals are in worn pelage, thus
allowing a great amount of the black underfur to show, which gives
a markedly darker color. The unworn hair is only slightly darker
than that of the topotypes. The cranial details prove these animals
to be intergrades between planirostris and trumbullensis. They re-
semble trumbullensis in size of tympanic bullae, extension of the
premaxillae posterior to the nasals and shape of the nasals. The
majority of the cranial details are as in planirostris to which they
are here referred.
When Goldman (1938:337) named Thomomys bottae nicholi
from northern Arizona he referred twelve specimens from St.
George, Washington County, Utah, to his newly named race. He
noted that the animals from this region intergrade with planirostris.
I have had occasion to study one-fourth of the material available
to Goldman for his original description of nicholi. For his speci-
mens listed as from St. George, the exact locality of capture, which
is so essential in this distributional study, was not given. All of
the specimens that I have seen from the Biological Surveys Collec-
tion are from the south side of the Virgin River, while St. George
itself is on the north side. As noted earlier in this account there are
differences between the gophers from the two sides of the Virgin
River in this area. Those from the north side are intergrades be-
tween birdseyei, planirostris and virgineus, while those from the
south side are intergrades between planirostris and trumbullensis.
Goldman (loc. cit.) mentioned several times that the skulls of
nicholi were nearly indistinguishable from, or closely resembled
those of, trumbullensis. Color was the only truly diagnostic char-
acter mentioned by Goldman. My study reveals the same differ-
ences and likenesses found by Goldman, but I consider color alone
insufficient basis in this instance for establishing a new subspecies,
and regard Thomomys bottae nicholi as a synonym of the earlier
proposed name, Thomomys bottae trumbullensis.
Durrant — Pocket Gophers of Utah 71
The animals from the south side of the Virgin River, labelled as
from St. George, Washington County, heretofore referred by Gold-
man to nicholi, are intergrades between trumbullensis and plani-
rostris and along with other specimens from the same place are
referable to the latter race.
Specimens examined. — Total, 68, distributed as follows: Washington County: Danish
Ranch, 5 mi. NW Leeds, 1; Zion National Park, 2 (M. V. Z.); Grotto Camp, Zion National
Park, 4,300 ft., 6 (N. H. M. S. D.); Springdale, 3,400 ft., 4 (K. U.); near Short Creek Road,
S town of Virgin, 4 (R. H.); St. George, N Virgin River, 2,950 ft., 21 (4, M. V. Z. ; 8, R.
H. ; 9, N. H. M. S. D.); Santa Clara Creek, 3 mi. SW St. George, 2,800 ft., 26; St. George, S
Virgin River, 5 (2, M. V. Z. ; 3, U. S. N. M.) ; 2 mi. SE St. George, 2,950 ft., 2 (N. H. M.
S. D.); 3 mi. S St. George, 1 (C. M.); 4 mi. SE St. George, S Virgin River, 1 (R. H.); 6 mi.
S St. George, 2,700 ft., 6 (K. U.). Kane County: East Entrance Zion National Park, 5,725
ft., 3 (N. H. M. S. D.); near East Entrance Zion National Park, 5,500 ft., 3 (N. H. M. S.
D.).
Additional records. — Washington County: Zion National Park, 22; Washington, 7 (Burt,
1931:39); St. George, 5; Santa Clara, 2 (Bailey, 1915:75).
Thomomys bottae absonus Goldman
Thomomys perpallidus absonus Goldman, Journ. Washington Acad.
Sci., 21:425, October 19, 1931.
Thomomys bottae absonus Hall and Davis, Proc. Biol. Soc. Washing-
ton, 47:52, February 9, 1934; Goldman, Proc. Biol. Soc. Washington,
48:156, October 31, 1935.
Thomomys perpallidus aureus Bailey, N. Amer. Fauna, 39:75, No-
vember 15, 1915; Barnes, Bull. Univ. Utah, 12 (No. 15) :85, April, 1922;
Bull. Univ. Utah, 17 (No. 12):100, June, 1927.
Type.— Male, adult, skin and skull, No. 250016, U. S. National Museum
(Biological Surveys Collection) ; Jacobs Pools, Houserock Valley, 4,000 ft.,
Coconino County, Arizona; June 7, 1931; collected by E. A. Goldman; orig-
inal number 23569 (after Goldman, type not seen).
Range. — Southern Utah in Kane and Garfield counties, in the drainages of
Kanab Creek, Johnson Creek, Paria River and Escalante River.
Diagnosis. — Size medium (see measurements). Color: Upper parts Och-
raceous Buff mixed with dusky; sides and underparts Light Ochraceous Buff;
chin, nose, cheeks and top of head grayish black; postauricular patches black
mixed with buff; front feet, hind feet, inguinal region and distal third of tail
white. Skull: Nasals relatively long; rostrum narrow; ascending processes of
premaxillae narrow; extension of premaxillae posterior to nasals short; lamb-
doidal and sagittal crests poorly developed; zygomatic arches light; jugals
nearly straight; palate narrow; molariform teeth small.
Comparisons. — Compared with topotypes of Thomomys bottae
trumbullensis, absonus differs in: Size smaller. Color: Markedly
lighter throughout. Skull: Smoother, less angular; zygomatic arches
weak as opposed to robust; nasals more convex as viewed later-
ally; extension of premaxillae posterior to nasals less; ascending
processes of premaxillae narrower; palate narrower; palatal pits
shallower; rostrum narrower; molariform teeth smaller.
72 University of Kansas Publs., Mus. Nat. Hist.
For comparisons of absonus with Thomomys bottae aureus see
account under that form.
Among named races of Thomomys bottae, absonus most closely
resembles planirostris, but can be distinguished from the topotypes
as follows: Size markedly smaller. Color: Lighter, more buffy
throughout. Skull: Smaller, less ridged and more nearly flat; nasals
convex as opposed to flat; extension of premaxillae posterior to
nasals less; width of ascending processes of premaxillae less; zygo-
matic arches weaker; palate narrower; alveolar length of upper
molar series shorter; tympanic bullae more inflated ventrally; mo-
lariform teeth smaller and lighter.
Remarks. — One specimen from Kanab is an intergrade between
trumbullensis and absonus. The majority of its characters are with
absonus to which it is referred (see Hall and Davis, 1934:52).
Two specimens from Escalante are intergrades between absonus
and dissimilis, but are referable to absonus.
Specimens examined. — Total, 3, distributed as follows: Garfield County: Escalante, 5,258
ft., 2 (B. Y. U.). Kane County: Kanab, 4,925 ft., 1 (M. V. Z.).
Thomomys bottae alexandrae Goldman
Thomomys alexandrae Goldman, Journ. Washington Acad. Sci.,
23:464, October 15, 1933.
Thomomys bottae alexandrae Benson, Univ. California Publ. Zool.,
40:449, December 31, 1935.
Type. — Male, adult, skin and skull, No. 250969, U. S. National Museum
(Biological Surveys Collection) ; 5 mi. SE Rainbow Lodge, near Navajo Moun-
tain, Coconino County, Arizona; June 16, 1933; collected by E. A. Goldman;
original number 23613 (after Goldman, type not seen).
Range. — In extreme southwestern San Juan County, Utah. Known only from
Navajo Mountain, probably limited to the area enclosed on the north by the
Colorado and San Juan rivers, on the east and west by Navajo and Piute
canyons, respectively.
Diagnosis. — Size small (see measurements). Color: Upper parts Cinnamon
Buff, grading over the sides to Pinkish Buff on underparts; nose and top of
head grayish black; hind feet and tail white; postauricular patches large and
dark. Skull: Small and not heavily ridged; zygomatic arches widely spread-
ing but weak; zygomatic arches nearly parallel; tympanic bullae moderately
inflated ventrally; palate not arched; interpterygoid space U-shaped; denti-
tion light.
Comparisons. — Compared to topotypes of Thomomys bottae ab-
sonus, alexandrae differs as follows: Size smaller in every measure-
ment taken. Color: Upper parts Cinnamon Buff as contrasted
with Light Ochraceous Buff. Skull: Smaller in every measurement
Durrant — Pocket Gophers of Utah 73
taken except interorbital breadth and alveolar length of upper molar
series which are larger; molariform teeth larger.
Among named races of Thomomys bottae occurring in Utah,
alexandrae most resembles T. b. aureus to the northeast. It can be
distinguished from topotypes of the latter by: Size smaller in every
measurement taken. Color: Darker throughout. Skull: Smaller,
slenderer and more nearly flat; palate nearly flat as opposed to
arched; zygomatic arches weaker and not so widely spreading; in-
terparietal narrower; tympanic bullae smaller; dentition weaker.
Remarks. — Goldman (1933:464) accorded alexandrae full specific
status, because he found no intergradation with other races, from
which he thought alexandrae had been isolated perhaps for thou-
sands of years by the barriers of the surrounding terrain. Benson
(1935:450) noted resemblances between alexandrae and specimens
of latirostris from Keams Canyon, Zuni Well, and Winslow in
Navajo County, Arizona (— aureus), and also between alexandrae
and absonus from Houserock Valley, Arizona. He thought that
alexandrae is no more differentiated or isolated than each of several
other kinds of desert pocket gophers, and, therefore, accorded alex-
andrae only subspecific status, as I, also, am inclined to do.
Specimens examined. — One (M. V. Z.) from Soldier Spring, Navajo Mountain, 8,600 ft.,
San Juan County. Fourteen topotypes from Arizona also were examined.
74
University of Kansas Publs., Mus. Nat. Hist.
Measurements of Adult Males of Thomomys
(In millimeters)
H
— i
t-1
T)
f
N
—
>
H
t-1
*
1
"D
3
2
»
!5
3
'<
rt
c 5
S
2
ft
s.
5*
->
5"
5
B"
o
C
3
£8
5"
g
a.'
—
O
•a 2.
0 f»
c-*- co
5'
?3
B
B-
O
a
B"
O
3*
;_
1 '
p.
1
0
8
3
—
-I
O 3
IT'S-
s
o
CO
o
CO
a.
—
=r
0.
?§ o
3. Hi
?B
i
l
o
(S
«■
;
;
CO
Av 243
Min 232
Max 253
Av 237
Min 215
Max 250
Av 228
Mia 223
Max 235
Av 222
Min 214
Max 236
Av 206
Min 198
Max 215
Av 230
Min 220
Max 242
Av 184
Min 179
Max 189
Av 251
Min 248
Max 255
Av 229
Min. 209
Max 255
T. b. aureiventris, 4; topotypes (Hall, 1930:446)
67 32 36.4 14.7 26.5 21.5 6.6 7.9
59 31 35.3 14.0 25.5 20.9 6.1 7.8
72 33 37.1 15.3 27.3 22.3 6.9 8.0
T. b. centralis, 9; topotypes (Hall, 1930:446)
75 30 36.3 14.6 25.2 20.7 6.6 8.0
61 29 34.5 13.9 24.6 19.7 5.8 7.5
83 32 38.0 15.9 26.1 21.9 7.2 8.7
T. b. albicaudatus, 7; topotypes (Hall, 1930:446)
65 31 35.4 14.0 26.1 20.5 6.6 8.1
59 29 34.9 13.4 24.9 19.8 6.4 7.8
72 32 36.1 15.1 27.8 21.1 6.9 8.4
T. b. robustus, 9; topotypes
65 29 34.1 13.6 26.0 20.8 6.4 7.8
59 28 32.6 13.0 25.2 20.0 6.1 7.3
70 31 35.7 14.4 26.7 21.5 6.7 8.2
T. b. stansburyi, 5; topotypes
60 28 32.3 12.4 22.4 19.1 6.3 7.6
58 26 30.6 12.0 21.5 18.2 6.2 7.0
68 31 33.4 13.0 23.1 20.1 6.5 8.0
T. b. nesophilus, 4; topotypes
69 32 35.3 14.4 25.5 20.4 6.8 8.4
60 30 33.6 14.1 24.9 19.8 6.5 8.2
75 33 36.5 14.8 26.2 21.1 7.1 8.7
T. b. minimus, 2; topotypes
60 25 30.7 11.3 21.3 18.7 6.4 7.4
55 24 28.7 10.2 20.2 17.8 6.3 7.3
64 26 32.8 12.5 22.4 19.6 6.4 7.6
T. b. lenis, 2; topotypes
80 32 39.7 16.0 28.6 22.6 6.8 8.3
74 31 39.4 15.8 28.4 22.4 6.6 8.2
86 32 29.9 16.2 28.7 22.7 6.9 8.5
T. b. contractus, 8; topotypes
74 31 33.3 12.5 23.7 19.1 6.6 7.6
63 28 30.0 10.9 21.4 17.7 6.3 7.2
85 33 37.4 14.5 26.4 20.9 6.9 8.0
2.4
1.8
3.4
3.2
2.2
4.5
3.2
3.0
3.8
2.7
2.0
3.0
2.8
2.5
3.0
2.5
2.1
2.9
2.5
2.5
2.5
3.4
3.0
3.7
3.0
2.4
3.5
15.7 8.4
14.7 8.1
17.0 8.8
14.7
14.1
15.4
13.9
12.9
15.0
18.4
17.9
18.8
7.5
7.1
7.8
17.1 8.2
16.4 7.6
18.4 8.6
7.5
7.0
7.9
8.8
8.6
8.9
15.4 7.3
13.5 6.5
18.2 8.0
Durrant — Pocket Gophers of Utah
75
Measurements of Adult Males of Thomomys — Continued
*3
Q
n
0
9
to
t->
M
f
s
CO
D
1
ED
"1
9.
B*
5'
c
5
I
o
tr
CO
O
5.'
cr
I
CO
a.
Q.
c <
•o 2.
S3
B 5*
O 3
»9
-> a-
W
O ST
CO 9
e-»- Co
S'
*° ^,
S~
■-»
•— i*d
" 3
CO
■A
9-
o
3-
ts
u
o
8.
No. 191959 (U. S. N. M.)
222 65 28
T. b. levidensis, 1 ; topotype
33.3 12.7 24.5 19.0 6.5
7.6 3.3 15.1 8.0
T. b. convexus, 6; topotypes
Av 213 59 28 33.1 14.3 24.9 21.7 6.6 8.0 2.6
Min 206 57 27 31.3 13.9 23.8 21.0 6.5 7.7 2.1
Max 233 68 29 35.0 14.6 26.7 22.3 6.8 8.1 2.8
T. b. tivius, 7; topotypes
Av 208 69 27 31.5 12.2 22.4 18.4 6.4 7.2 2.4
Min 199 67 25 29.3 11.9 20.6 17.1 6.0 7.0 2.1
Max 227 70 30 34.1 12.8 25.0 19.8 6.6 7.6 3.0
T. b. bonnevillei, 3; topotypes
Av 228 70 30 35.4 13.9 26.4 21.8 6.6 8.1 3.7
Min 221 62 30 33.6 13.2 25.4 20.5 6.5 8.1 3.4
Max 236 79 30 37.4 14.9 28.0 22.5 6.7 8.1 4.3
T. b. sevieri, 3; topotypes
Av 216 67 '30 32.7 12.9 22.9 18.7 6.4 7.2 2.5
Min 210 66 29 31.7 11.8 22.2 18.0 6.2 7.0 1.8
Max 222 68 31 33.5 13.5 23.4 19.3 6.7 7.2 3.0
T. b. wahwahensis, 4; topotypes
Av 228 66 29 34.7 13.5 25.5 20.7 6.6 7.3 2.3
Min 210 60 26 33.0 13.1 24.6 20.1 6.5 7.0 2.2
Max 250 78 30 37.6 14.6 27.0 21.4 6.8 8.0 2.5
T. b. planirostris, 8; topotypes (Burt, 1931:39)
Av 238 76 32 35.6 13.8 25.9 20.4 6.6 8.5 3.7
Min 222 66 31 33.3 12.5 24.4 19.8 6.2 8.2 3.0
Max 261 83 34 38.7 15.3 27.6 21.3 7.2 8.9 4.5
T. b. birdseyei, 3; topotypes
Av 227 64 31 34.9 13.8 26.2 20.9 6.2 8.4 2.6
Min 214 52 30 34.5 13.1 26.0 20.1 6.0 8.1 2.2
Max 243 81 32 35.2 14.1 27.4 21.5 6.5 8.8 2.8
T. b. virgineus, 5; Beaverdam Wash, 5 mi. N Utah-Arizona Line
Av 226 68 29 34.6 13.5 25.6 20.7 6.3 8.0 3.0
Min 216 62 27 33.5 12.8 25.0 20.0 6.1 7.6 2.4
Max 235 70 30 34.9 14.4 26.0 21.1 6.6 8.4 3.5
T. b. aureus, 3; topotypes
Av 242 68 34 36.6 14.3 25.3 21.4 6.6 8.3 2.4
Min 233 65 32 35.3 13.8 24.6 20.6 6.4 7.7 2.0
Max 251 70 36 37.8 14.9 25.8 22.0 6.8 8.7 2.5
16.2
15.2
17.2
8.2
8.0
8.6
14.0
13.2
15.0
7.1
6.5
7.9
17.6
16.1
18.1
8.5
8.2
8.7
15.3
14.5
16.4
7.6
7.5
7.7
15.7
14.9
17.1
8.7
8.5
9.0
8.8
8.3
9.4
16.3
16.0
16.9
8.3
8.2
8.4
16.5
15.3
17.4
8.5
8.3
8.7
17.2
16.7
17.9
8.7
8.3
9.0
78 University of Kansas Publs., Mus. Nat. Hist.
Measurements of Adult Females of Thomomys — Continued
H
■H
f
T)
t"1
ts
g
1— 1
13
>
M
f
a
o
■o
n>
I
A
*<
w X
2
S-
3
3
3
TO
C <
•O r*
a
(0
2. <g.
— 3-
TO
13*
0
TO
B"
0
3
0
0
cr
■a q
•a 2.
o 2
t» 3
"5-
3-
to
Q.
o
Q
i -i
o
JO -i
D*
1
3
g
3"
0
0
o
•*
3" «
B"
s
3
3-
3*
0
Hi
a
£0
cn
&
CD
C3
3"
-1
m
P
E
H
m
'J
0 3
1 v
O
-♦»
o
a
O
o
o
o
'/i
—
ct> o
? 3
:
f
T. 6. convexus, 11; topotypes
Av 197 57 27 29.9 12.5 21.7 19.3 6.6 7.7 2.6
Min 182 43 26 27.9 11.2 21.0 18.8 6.2 7.1 2.1
Max 204 63 28 30.9 13.4 22.3 19.8 7.1 7.9 3.1
T. b. tivius, 5; topotypes
Av 203 68 27 29.5 11.1 21.1 17.8 6.5 7.2 2.4
Min 192 63 26 28.0 10.5 20.1 17.3 6.3 7.1 2.0
Max 215 74 30 31.3 11.4 22.9 19.0 6.7 7.5 3.0
T. b. bonnevillei, 7; topotypes
Av 199 57 28 31.7 11.8 22.2 19.3 6.6 7.7 3.2
Min 184 50 24 29.4 10.1 20.3 18.1 6.4 7.1 2.6
Max 216 66 29 34.3 13.6 24.3 20.3 7.0 8.5 4.1
T. b. sevieri, 7; topotypes
Av ... 205 62 28 30.2 11.8 21.6 18.0 6.4 7.0 2.7
Min 199 54 28 29.4 11.3 20.6 17.7 6.1 6.6 2.1
Max 212 70 29 30.7 12.6 22.1 18.6 6.8 7.4 3.0
T. b. wahwahensis, 8; topotypes
Av 185 56 27 28.7 11.3 20.6 17.6 6.3 7.1 2.1
Min 180 50 26 26.3 10.2 19.0 16.5 5.8 6.9 1.1
Max 197 62 29 30.7 12.6 22.0 19.0 6.7 7.8 2.9
T. b. planirostris, 8; topotypes (Burt, 1931:39)
Av 215 71 31 32.2 12.4 23.2 18.7 6.5 8.1 3.6
Min . 205 61 30 31.5 11.8 22.3 18.1 6.4 7.5 2.8
Max 228 78 33 33.0 12.9 24.1 19.5 6.7 8.6 4.5
T. b. birdseyei, 3; topotypes
Av 220 71 29 31.6 11.8 22.7 18.6 6.1 7.4 2.4
Min . 217 68 28 31.4 11.0 22.4 18.3 6.0 7.3 1.6
Max 223 75 30 32.0 12.8 23.0 19.1 6.2 7.4 3.0
T. b. virgineus, 4; Beaverdam Wash, 5 mi. N Utah- Arizona Line
Av 211 64 29 31.6 12.2 22.6 19.4 5.9 7.5 3.1
Min 202 60 27 31.3 11.3 22.4 18.8 5.8 7.3 2.4
Max 218 68 30 32.1 12.8 22.7 20.0 6.1 7.8 3.7
T. b. aureus, 3; topotypes
Av 226 57 31 33.2 13.3 23.8 19.8 6.7 8.2 1.9
Min 217 54 30 32.8 12.5 23.3 19.6 6.4 8.0 1.6
Max'..... 233 64 31 34.0 14.2 24.4 19.8 6.9 8.4 2.0
No. 20300 (C. M.) T. b. howelli, 1; 10 mi. N Moab
202 59 28 32.4 12.3 21.1 19.2 6.4 7.9 2.4
14.7
13.3
15.2
7.4
7.1
7.7
13.5
12.7
14.2
6.8
6.4
7.2
14.9
13.5
16.6
7.3
6.9
7.7
14.2
13.9
14.7
7.1
6.6
7.6
12.6
10.8
14.0
7.1
6.4
7.6
....
7.9
7.5
8.1
14.7
13.3
15.3
7.5
7.4
7.5
15.1
14.4
15.5
7.3
7.2
7.6
15.3
14.5
16.4
8.2
8.2
8.3
15.8 8.3
Durrant — Pocket Gophers of Utah
79
Measurements of Adult Females of Thomomys — Concluded
H
o
3
•a
9
3
J}
3
cd
EC
3
T5
3
3
»
a,
3-
O
5-'
o
c <
•a ™
•a 2.
2 »
B m-
O 3
IT1*
a> o
d r-t-
GO 3
o"
BS.
3
3
B*
a
a>
B)
a.
**■
B*
O
o
No. 158524 (U. S. N. M.)
188 61 27
No. 158528 (U. S. N. M.)
203 61 27
A.v
.. 205
63
28
.. 195
57
27
Max . . .
. . 215
70
29
T. b. dissimilis, 1; topotype
28.2 10.1 19.0 16.7 6.
T. b. osgoodi, 1; topotype
29.6 11.5 .... 18.3
T. b. alexandrae, 3; topotypes
30.9 11.8 20.8 17.9 6.4
28.7 11.5 20.5 17.2 6.3
31.5 12.1 22.2 18.6 6.5
7.4 2.1 12.8 6.5
6.9 7.4 2.0 14.0 7.3
7.6
1.8
14.1
7.5
7.5
1.5
13.6
7.2
7.7
2.0
14.7
7.7
80 University of Kansas Publs., Mus. Nat. Hist.
LITERATURE CITED
Allen, J. A.
1874. Notes on the mammals of portions of Kansas, Colorado, Wyoming
and Utah, Part IV. On the mammals of the Great Salt Lake Valley,
Utah. Bull. Essex Inst., 6:61-66, 1874.
1893. Descriptions of four new species of Thomomys with remarks on
other species of the genus. Bull. Amer. Mus. Nat. Hist., 5:47-68,
April 28, 1893.
1893. List of mammals collected by Mr. Charles P. Rowley in the San
Juan region of Colorado, New Mexico and Utah, with descriptions
of new species. Bull. Amer. Mus. Nat. Hist., 5:69-84, April 28, 1893.
1896. List of mammals collected by Mr. Walter W. Granger in New Mexico,
Utah, Wyoming and Nebraska, 1895-1896, with field notes by the
collector. Bull. Amer. Mus. Nat. Hist., 8:241-258, November 25, 1896.
1905. Mammals from Beaver County, Utah, collected by the Museum ex-
pedition of 1904. Brooklyn Inst. Mus. Sci. Bull., 1:117-122, March
31, 1905.
Bailey, Vernon.
1915. Revision of the pocket gophers of the genus Thomomys. N. Amer.
Fauna, 39:1-136, pis. 8. 10 figs., November 15, 1915.
Barnes, Claude T.
1922. Mammals of Utah. Bull. Univ. Utah. 12 (No. 15): 1-176, 30 figs.,
April, 1922.
1927. Utah mammals. Bull. Univ. Utah. 17 (No. 12): 1-183, 32 figs., June,
1927.
Benson, Seth B.
1935. A biological reconnaissance of Navajo Mountain, Utah. Univ. Cali-
fornia Publ. Zool., 40:439-455, December 31, 1935.
Burt, William H.
1931. A new pocket gopher of the genus Thomomys from Utah. Proc.
Biol. Soc. Washington, 44:37-40, May 8, 1931.
Coues, E.
1875. Abstract of results of a study of the genera Geomys and Thomomys.
Part III. Zoology, in explorations of the Colorado River of the
West and its tributaries, explored in 1869, 1870, 1871 and 1872 under
the direction of the Smithsonian Institution, reported by J. W.
Powell, Gov't Printing Office. Washington, D. C, 1875.
1877. Monographs of North American Rodents, No. X, Geomyidae, pp.
601-629, U. S. Geol. Surv. of the territories, Gov't Printing Office,
Washington, D. C, 1877.
Coues, E., and Yarrow, H. C.
1875. Report upon the collection of mammals made in portions of Ne-
vada, Utah, California, New Mexico and Arizona during the years
1871-74. Wheeler's Rept. Expl. W of 100th Mer. vol. 5, pp. 35-129,
1875.
Dirrant — Pocket Gophers of Utah 81
Davis, William B.
1939. The Recent mammals of Idaho. The Caxton Printers, Ltd., Caldwell,
Idaho, pp. 1-400, pis. 2. 33 figs, April 5, 1939.
Durrant, Stephen D.
1937. Two new gophers from Utah. Bull. Univ. Utah. 28 (No. 4):l-7,
August 18, 1937.
1939. A new pocket gopher of the Thomomys quadratm group from the
northern Great Basin region. Bull. Univ. Utah, 39 (No. 6):l-6,
February 28, 1939.
Goldman, E. A.
1933. New mammals from Arizona. New Mexico and Colorado. Journ.
Washington Acad. Sci, 23:463-473. October 15, 1933.
1936. New pocket gophers of the genus Thomomys. Journ. Washington
Acad. Sci., 26:111-120, March 15. 1936.
1938. New pocket gophers of the genus Thomomys from Arizona and
Utah. Journ. Washington Acad. Sci., 28:333-343, July 15. 1938.
1939. Remarks on pocket gophers, with special reference to Thomomys
talpoides. Journ. Mamm.. 20:231-244. May 14, 1939.
1942. Three new rodents from southern Utah. Proc. Biol. Soc. Washing-
ton, 55:75-78, July 25. 1942.
Hall, E. Raymond.
1931. Critical comments on mammals from Utah, with descriptions of new
forms from Utah, Nevada and Washington. Univ. California Publ.
Zool.v 37:1-13, April 10, 1931.
Hall, E. Raymond, and Davis, Wtilljam B.
1934. Notes on Arizona rodents. Proc. Biol. Soc. Washington, 47:51-56,
February 9, 1934.
Haywakd, C. Lynn.
1936. A bibliography of Utah mammalogy; including references to names
and type localities applied to Utah mammals. Utah Acad. Sci. Arts
and Letters, 13:122-146, 1936.
1941. A bibliography of Utah mammalogy; including references to names
and type localities (first supplement). Great Basin Nat., 2:125-136,
December 31. 1941.
Marshall, William H.
1940. A survey of the mammals of the islands in Great Salt Lake, Utah.
Journ. Mamm, 21:149-159. 2 pis, 1 map, May 14, 1940.
Merriam, C. Hart.
1901. Descriptions of twenty-three new pocket gophers of the genus Tho-
momys. Proc. Biol. Soc. Washington, 14:107-117. July 19, 1901.
6-2786
82 University of Kansas Publs., Mus. Nat. Hist.
Miller, Gerritt S., Jr.
1924. List of North American Recent mammals, 1923. U. S. Nat. Mus.
Bull., 128, pp. I-XVI, + 1-673, Govt. Printing Office, Washington,
D. C., March 18, 1924.
Svihla, Ruth Dowell.
1931. Mammals of the Uinta Mountains region. Journ. Mamm., 12:256-
266, pis. 1, 1 fig., August 24, 1931.
□
21-2786
The Systematic Status of Eumeces pluvialis
Cope, and Noteworthy Records of Other
Amphibians and Reptiles From
Kansas and Oklahoma
BY
HOBART M. SMITH
University of Kansas Publications
Museum of Natural History
Volume 1, No. 2, pp. 85-89
August 15, 1946
UNIVERSITY OF KANSAS
LAWRENCE
1946
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Donald S. Farner, Donald F. Hoffmeister
Volume 1, No. 2, pp. 85-89
Published August 15, 1946
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR. . STATE PRINTER
TOPEKA. KANSAS
1946
21-2765
gO,SS3
The Systematic Status of Eumeces pluvialis Cope, and
Noteworthy Records of Other Amphibians and
Reptiles from Kansas and Oklahoma
By
HOBART M. SMITH
A number of noteworthy items have come to attention in the
course of a survey of material for a handbook on the herpetology of
Kansas. Some of the items, which follow, can be recorded here more
appropriately than in the handbook.
Eumeces anthracinus pluvialis Cope
Recent material in addition to information presented in Taylor's
monograph of Eumeces (Univ. Kansas Sci. Bull, 23, 1935) re-
veals that Eumeces anthracinus is composed of three geographically
distinct populations : One occurs from western New York to northern
Georgia, and west to Kentucky, in the Appalachian uplands or
northward of them; a second centers about the Ozark uplands but
extends into northwestern Louisiana, eastern Texas, central Okla-
homa, eastern Kansas, and nearly as far east as the Mississippi
river in northern Arkansas and southern Missouri ; the third popula-
tion occurs in extreme southern Alabama and Mississippi.
These populations differ in at least the color of the young. Speci-
mens from the eastern area are marked at birth like the adults ; those
from the western area are black at birth and develop stripes as they
grow older; unfortunately young specimens from the southern area
are not known.
Obviously at least two races are involved, the eastern and the
western. Whether the southern population belongs to one of these
races or is distinct is unknown. Until this point is settled the name
for the western race will remain in doubt. The eastern race is the
typical one, Eumeces a. anthracinus (Baird) (Journ. Acad. Nat. Sci.
Phila., 1 (ser. 2) :294, 1850; type locality North Mountain, Carlisle.
Pennsylvania). The southern population has been named pluvialis
by Cope (Ann. Rept. U. S. Nat. Mus., 74:663-664, 1900; type lo-
cality Mobile, Alabama). Unfortunately no name is available for
the western population. It may either be called Eumeces anthra-
cinus pluvialis, or be given a new name, according to the ultimate
(87)
88 University of Kansas Publs., Mus. Nat. Hist.
decision on its consubspecificity with the southern population. I
suggest retention of the name pluvialis at least until a more careful
study indicates the necessity of further change.
Eurycea lucifuga (Rafinesque)
On October 21, 1945, E. W. Jameson, Jr., discovered a specimen of
this species in a small cave situated in a park l1/^ miles south of
Galena, Cherokee County, Kansas, on the north side of Shoal Creek,
NW !/4 of Sec. 35, T. 35 S., R. 25 E. Later the same day Claude
W. Hibbard and I returned to the same cave, and with the help of
Jameson found two more specimens. All were found under stones
in the twilight zone. Exploration of deeper recesses of the cave was
impossible because the larger entrances to them had been closed off
with cement to prevent children from entering. No water was run-
ning from the cave at the time we were there, although there was
visible evidence of a previous heavy flow of water, probably in times
of heavy and prolonged rains. The only other salamanders found
in the limited area available for exploration belonged to Eurycea
longicauda melanopleura (Cope), a form considerably more abun-
dant in the cave than E. lucifuga.
This constitutes the first published record of the occurrence of
E. lucifuga in Kansas. Previous records from Arkansas, Missouri
and Oklahoma, as well as a sight record by Taylor (Smith, Amer.
Midi. Nat., 15:382-383, 1934) have indicated its probable occur-
rence in Kansas.
The largest specimen obtained is an adult male measuring 166 mm.
in total length; it exceeds by 2 mm. the maximum previously known.
The pattern and other characters of all specimens appear typical.
The specimens are in the Museum of Natural History of the Uni-
versity of Kansas.
Hyla crucifer crucifer Wied
In 1943 Bragg (Great Basin Nat,, 4:67, 1943) stated that Hyla
crucifer crucifer has been recorded with certainty from only one
county in Oklahoma, McCurtain County in the extreme southeastern
part of the state. Reports of their call being heard in Le Flore
County, immediately north of McCurtain County had also been
transmitted to him.
In Kansas the species is still known only from the northern half
of the extreme eastern part of the state (Smith, Amer. Midi. Nat.,
Smith — Eumeces pluvialis 89
15:472, 1934). Between this area and southeastern Oklahoma no
record of occurrence of the species has been available.
An adult specimen taken by Dr. Joseph Tihen in the extreme
southeastern corner of Delaware County, Oklahoma (Mus. Nat.
Hist., Univ. Kans., No. 20827), thus provides a second definite lo-
cality for the species in Oklahoma and suggests the probability that
it ranges along the entire eastern border of both Kansas and Okla-
homa. The specimen is in poor condition but enough of the pattern
and some other features can be discerned to permit reliable identi-
fication.
□
21-2765
The Tadpoles of Bufo cognatus Say
BY
HOBART M. SMITH
University of Kansas Publications
Museum of Natural History
Volume 1, No. 3, pp. 93-96, 1 figure in text
August 15, 1946
a^ Zoology VN
UNIVERSITY OF KANSAS
LAWRENCE
1046
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Donald S. Farner, Donald F. Hoffmeister
Volume 1, No. 3, pp. 93-96
Published August 15, 1946
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1946
21-2764
The Tadpoles of Bufo cognatus Say
By
HOBART M. SMITH
The tadpoles of this species have been described by Bragg (Copeia,
1936: 14-20, figs. 1-13; Amer. Midi. Nat., 18:273-284, figs. 1-5,
1937). The drawings and descriptions of the mouthparts, how-
ever, appear to have been taken from dried, or immature, or trans-
forming individuals, for they do not agree among themselves nor do
they agree with larvae obtained in the field and now in the Museum
of Natural History of the University of Kansas.
At hand are two series of tadpoles of this species; one series was
collected July 2, 1938, 1.5 miles east of Meade County State Park,
Fig. 1. — Mouthparts of a tadpole of Bufo cognatus. Disk widely spread.
Approximately X 45.
Kansas, and the other lacks data. The second lot contains numerous
sizes of tadpoles from 14 mm. to 31 mm., and several transforming
specimens which clearly possess the pattern so typical of this species.
Mouthparts in both series (consisting all told of about 200 speci-
mens) are fairly constant except in the transforming and extremely
young specimens. The accompanying figure shows them as seen
with the mouth disk widely spread. The indentations at the corners
of the mouth in the papillary fringe are more prominent when the
mouthparts are less extended. The outer row of teeth of the lower
labium is sometimes a little shorter or longer than the figure shows,
but the average is about as indicated. The extent of the medial
edge of the papillae on the lower labium varies somewhat; in some,
(95)
9G University of Kansas Publs., Mus. Nat. Hist.
the papillae barely reach the level of the ends of the outer row of
teeth, while in others they overlap the ends slightly.
Measurements agree with those given by Bragg, except that ap-
pearance of the hind legs occurs at about 15 mm.; the fore legs appear
at about 28 mm. A pattern recognizably similar to that of the adult
is evident at about 20 mm.
These tadpoles show such a striking similarity to those referred by
Wright, to Bafo compactilis Wiegmann (Proc. U. S. Nat. Mus., 74:4,
pi. 5, fig. 6, 1929) that their conspecificity is suggested. If on the
other hand, the specimen figured by Wright is properly identified,
then the two species must in reality be very closely related. A direct
comparison of positively identified tadpoles of each species is much
to be desired.
□
21-2764
Hybridization Between Two Species of
Garter Snakes
BY
HOBART M. SMITH
University of Kansas Publications
Museum of Natural History
Volume 1, No. 4, pp. 97-100
August 15, 1946
UNIVERSITY OF KANSAS
LAWRENCE
1946
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Donald S. Farner, Donald F. Hoffmeistn
Volume 1, No. 4, pp. 97-100
Published August 15, 1946
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR. . STATE PRINTER
TOPEKA. KANSAS
1946
21-2763
M o ^> ^
Hybridization Between Two Species of Garter Snakes
By
HOBART M. SMITH
The chief characters distinguishing Thamnophis radix (Baird and
Girard) and T. marciana (Baird and Girard) in southern Kansas
are:
marcxana
lateral light line involving only the
3d scale row anteriorly,
dorsal light line without distinct
edges, varying in width from less
than 1 to nearly 3 scale rows, at
various places on body,
several anterior lateral spots fused
across lateral light stripes.
2 posterior upper labials not light-
centered, unlike others.
A well -developed, white, black-
edged crescent behind angle of jaws
(postrictal crescent).
radix
1. lateral light line involving rows 3
and 4 anteriorly.
2. dorsal light line with straight, even
edges, IY2 scale rows wide.
usually no anterior lateral spots
fused across lateral light stripes.
2 posterior labials light-centered,
like others.
typically no well-developed post-
rictal crescent.
Typical specimens of radix are available from several localities in
Morton County of southwestern Kansas (Spring Creek; twelve
miles and eighteen miles north of Elkhart; Elkhart) ; from the State
Lake and Meade in Meade County; from Hunters, Harper County;
Coolidge, Hamilton County; and Ingalls, Gray County.
Typical marciana is available from Spring Creek, Morton County ;
Liberal, Seward County; and Clark County (no locality). An over-
lap of range with radix is evident, and from Spring Creek in Morton
County typical specimens of both species are available. Accordingly,
at present, I conclude that the two forms are correctly regarded as
distinct species.
Yet there is a rather marked tendency of radix to approach the
characters of marciana in southwestern Kansas. Two specimens
(one from Morton County, one from Gray County) have the dorsal
stripe slightly broken up by infiltration of the ground color onto the
edges of the scales. All southwestern radix develop the distinct
postrictal crescent so characteristic of marciana, and occasional
specimens fail to have light centers in the last two labials. Finally,
one specimen from Meade, Meade County (No. 5434), appears to be
(99)
100 University of Kansas Publs., Mus. Nat. Hist.
actually intermediate, and may be regarded as a hybrid. The mid-
dorsal stripe is not sharp-edged; the lips are barred exactly as in
marciana, the postrictal crescent is well defined, and the lateral
light stripe extends onto the fourth scale row only very slightly. I
refer the specimen to T. radix on the basis of the middorsal light
stripe which still is not as irregular as in marciana, upon the nature
of the lateral dark spots (not fused), and upon the slight extension
of the lateral light stripe onto the fourth scale row. Yet the speci-
men is definitely atypical of radix; no other of the 135 specimens
examined deviates so strongly from the normal condition.
Because the two kinds of garter snakes in question maintain their
distinctness at other places where they occur on common ground,
it seems best to interpret specimen No. 5434 as a hybrid rather than
an intergrade.
□
21-2763
Selected Records of Reptiles and Amphibians
from Kansas
BY
JOHN BREUKELMAN AND HOBART M. SMITH
University of Kansas Publications
Museum of Natural History
Volume 1, No. 5, pp. 101-112
August 15. 1946
UNIVERSITY OF KANSAS
LAWRENCE
1946
University or Kansas Publications, Museum of Natural, History
Editors: E. Raymond Hall, Chairman, Donald S. Farner, Donald F. Hoffmeister
Volume 1, No. 5, pp. 101-112
Published August 15, 1946
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR. . STATE PRINTE..
TOPEKA. KANSAS
1 946
21-2762
Selected Records of Reptiles and Amphibians
from Kansas
By
JOHN BKEUKELMAN AND HOBART M. SMITH
Preparation of a handbook of reptiles and amphibians by the
junior author has led to a survey of the collections of these animals
at Kansas State Teachers College in Emporia. Numerous locality
records of interest and importance have been accumulated there
through the efforts of the senior author and a number of his stu-
dents, particularly Mr. Allen Downs. The more important records,
including the first record for Kansas of Rana sylvatica, are reported
here. We have not mentioned specimens that are from counties from
which the University of Kansas Museum of Natural History al-
ready has specimens.
Specimens examined by Smith are indicated by an asterisk; those
identified by the late Dr. F. N. Blanchard are indicated by an en-
circled period (none of these specimens are now available). All
other specimens here recorded have been examined either by the
senior author or by Mr. Allen Downs, or by both. Specimen num-
bers, unless otherwise indicated, are those of the Kansas State
Teachers College collection.
Triturus viridescens louisianensis (Wolterstorff), Newt.
Cherokee Co.: 1 mile north and 4 miles east of Crestline (No. 164).*
This is a terrestrial adult, and provides the second known locality
for the species in the state.
Ambystoma texanum (Matthes), Narrow-mouthed Salaman-
der.— Lyon Co.: Emporia.
Ambystoma tigrinum mavortium (Baird), Tiger Salamander.
— Lyon Co.: (No. 292) ; 2 miles east of Americus. Ness Co.: Ness
City (No. 591).
Scaphiopus bombifrons Cope, Plains Spadefoot. — Ness Co.: 4
miles west, 1.5 miles north of Ness City (No. 592).
Bufo americanus americanus (Holbrook), American Toad. —
Chase Co.: 10 miles southwest of Saffordville. Cherokee Co.: 4
miles southeast of Columbus. Lyon Co.: 6 miles south of Plymouth
(No. 290)*; Emporia (Nos. 442, 443).* The records from Chase
and Lyon counties represent the westernmost localities for the species
in Kansas.
(103)
104 University of Kansas Publs.. Mus. Nat. Hist.
Bufo cognatus Say, Great Plains Toad. — Ness Co.: 4 miles
west and 1.5 miles north of Ness City (No. 594) .
Bufo woodhousii woodhousii (Girard), Rocky Mountain Toad.
—Clark Co.: 11 miles south of Bucklin (No. 401).* Decatur Co.:
Sappa Creek near Oberlin (2 spec).* Ford Co.: 5 miles southwest
of Dodge City (1 spec.).* Lyon Co.: Emporia (No. 352).* Ness
Co.: Ness City (Nos. 502-504, 595, 596)*; 4 miles west, 1.5 miles
north of Ness City (No. 593).* Sheridan Co.: Sheridan County
State Park (Nos. 565-568).
Acris crepitans Baird. Northern Cricket Frog. — Ness Co.: 4
miles west and 1.5 miles north of Ness City (Nos. 506, 507, 597-
606).*
Pseudacris nigrita triseriata (Wied), Striped Chorus Frog. —
Lyon Co.: 10 miles south of Plymouth; 3 miles north of Emporia
(No. 300) ; 7 miles west of Olpe; 2 miles northeast of Emporia (Nos.
434-441).* Neosho Co.: 3 miles west of Erie.
Hyla versicolor versicolor (LeConte), Common Tree Toad. —
Chautauqua Co.: Elk City (No. 621).
liana catesbeiana Shaw, Bullfrog. — Ness Co.: 4 miles west and
1.5 miles north of Ness City (No. 607).* Wallace Co.: 3 miles east
of Sharon Springs (1 spec.).*
Rana pipiens brachycephala Cope, Leopard Frog. — Clark Co.:
11 miles south of Bucklin (Nos. 398-400).* Ness Co.: 4 miles west
and 1.5 miles north of Ness City (Nos. 505, 508, 509, 608).*
Rana sylvatica cantabrigensis Baird, Wood Frog. — Lyon Co.:
extreme southwestern corner, 3 miles east of Chase County line, be-
tween the Verdigris River and the corner of the county (1 specimen,
now Mus. Nat, Hist., Univ. Kans., No. 23149).* This specimen pro-
vides for the first time a basis for inclusion of the species in the fauna
of Kansas. It measures 50 mm. snout to vent; hind leg from vent 80
mm.; tibia 23 mm. The ratio of hind leg to snout-vent measure-
ment is 0.625, and that of the tibia to snout-vent measurement is
2.17. Both figures are too high for Rana s. sylvatica, in which the
former ratio varies between 0.53 and 0.62, the latter ratio between 1.6
and 1.88. The ratios agree well with those of R. s. cantabrigensis, in
which the former ratio varies from 0.62 to 0.75, the latter from 1.93
to 2.3. Direct comparison of the specimen with typical examples of
both subspecies substantiates its allocation to R. s. cantabrigensis.
In the vicinity of Kansas, specimens of this species are known
from Missouri (St. Louis and Stone Counties only) and northwestern
Arkansas (Washington County: Winslow and Prairie Grove, Mus.
Breukelman and Smith — Reptiles and Amphibians 105
Nat. Hist., Univ. Kans., Nos. 16526, 18820, 18823). Reexamination of
these specimens confirms their identity as Rana sylvatica sylvatica
to which the Missouri specimens from Stone County undoubtedly
also belong. Accordingly this race is still to be anticipated in ex-
treme southeastern Kansas.
Reference of the (Specimen from Lyon County to Rana s. canta-
brii/ensis presents a problem in distribution, for the race is not known
from nearer Kansas than North Dakota, Minnesota, Wisconsin and
southern Illinois, except for a record given by Cope (Bull. U. S. Nat.
Mus., No. 34:437, 1889) from "western Missouri." Hurter (Trans.
St. Louis Acad. Sci., 20:123, 1911) restricts this record to Cooper
County, and presumably verifies Cope's identification. Hurter, too,
recognized the other form, R. syhratica, in Missouri (Marble Cave,
Stone County). Cope distinguished between the two races (as they
are now recognized) and recorded typical R. sylvatica from St. Louis.
Accordingly the specimen from Cooper County may be considered
properly identified racially. It apparently is from the locality
nearest to Kansas at which the race has been taken.
It seems highly probable that the Kansas occurrence, and pos-
sibly those in Arkansas and Missouri also, is a relict one. It is
highly improbable that the species has a continuous distribution in
either state. A wider or more southern distribution in the past
seems evident. The group to which it belongs certainly has had a
more southern range, as indicated by Taylor's discovery in Meade
County, Kansas (Univ. Kans. Sci. Bull., 28:217, 1942), of a fossil
species of Rana (parvissinia) , from the Upper Pliocene, presumably
related to sylvatica. It may or may not have been a direct ancestor
of the living species.
Microhyla olivacea (Hallowell), Northern Narrow-mouthed
Toad. — Lyon Co.: 6 miles southwest of Emporia. Wilson Co.: 7
miles northeast of Fall River.
Crotaphytus collaris collaris (Say), Collared Lizard. — Geary
Co.: 4 miles south of Fort Riley. Wabaunsee Co.: 2 miles north-
east of Alma.
Holbrookia maculata maculata (Girard), Earless Lizard. —
Chase Co.: 7 miles south of Saffordville (No. 350)*; 6 miles south-
west of Saffordville; 1 mile south of Saffordville (No. 338)*; 10
miles southwest of Olpe. Hodgeman Co.: Jetmore. Lyon Co.: 5
miles south of Plymouth; 6 miles southeast of Emporia; 9 miles
southwest of Emporia. Ness Co.: 4 miles west and 1,5 miles north
106 University of Kansas Publs., Mus. Nat. Hist.
of Ness City (Nos. 480, 481, 484-497, 609-611)*, 6 miles west and 0.5
miles south of Ness City (Nos. 482, 483, 498).*
Sceloporus undulatus garmani Boulenger, Northern Plains
Lizard. — Ellsworth Co.: Carneiro; 10 miles south of Ellsworth.
McPherson Co.: 4 miles west of Roxbury (No. 133). Ness Co.: 4
miles west and 1.5 miles north of Ness City (No. 479, 612).*
Phrynosoma cornutum (Harlan), Texas Horned Lizard. — Ells-
worth Co.: 10 miles south of Ellsworth. Lyon Co.: 1 mile south of
Emporia; 8 miles southwest of Emporia. Saline Co.: Coronado
Heights; 3 miles northwest of Lindsborg.
Ophisaurus ventralis (Linnaeus), Glass-snake Lizard. — Lyon
Co.: Emporia; 1 mile southwest of Emporia (No. 288).* Rooks Co.:
5 miles southwest of Stockton (No. 407).*
Cnemidophorus sexlineatus (Linnaeus), Six-lined Racerunner.
— Ellsworth Co.: Carneiro. Lyon Co.: 1.5 miles northwest of
Reading. Shawnee Co.: 5 miles east of Topeka (No. 14).*
Leiolopisma laterale (Say), Brown Skink. — Labette Co.: 7
miles northwest of Mound Valley (No. 301).* Lyon Co.: 1.5 miles
northwest of Reading. Wilson Co.: 4 miles southwest of Coyville
(No. 281).*
Eumeces fasciatus (Linnaeus). Common Five-lined Skink. —
Bourbon Co.: 1 mile north of Fulton. Chase Co.: 7 miles south-
west of Saffordville ; 6 miles south of Clements; 2 miles south of
Saffordville. Franklin Co.: 8 miles east of Ottawa; 2 miles south
of Ottawa; 2 miles southwest of Lane; 4 miles east of Ottawa; 5
miles southwest of Ottawa. Labette Co.: 2 miles southwest of
Dennis; 7 miles northwest of Mound Valley. Lyon Co.: 1.5 miles
northwest of Reading. Miami Co.: 2.5 miles south of Fontana.
Montgomery Co.: 5 miles west of Independence. Neosho Co.: 4
miles northwest of Erie (No. 318).*
Eumeces obsoletus (Baird and Girard), Sonoran Skink. — Cof-
fey Co.: 4 miles south of Gridley (No. 467).* Ellsworth Co.: 10
miles south of Ellsworth. Franklin Co.: 2 miles south of Lane.
Linn Co.: 0.5 miles north of Trading Post. Lyon Co.: 1.5 miles
northwest of Reading; 10 miles south of Plymouth; 2.5 miles north-
east of Dunlap; 4 miles southwest of Bushong; Emporia (No. 433)*;
Dunlap (No. 444).* McPherson Co.: 4 miles west of Lindsborg.
Morris Co.: 5 miles east of Skiddy ; 1 mile east of Skiddy. Neosho
Co.: 15 miles north of Parsons. Wilson Co.: 3 miles east of Buffalo.
Eumeces septentrionalis septentrionalis (Baird), Northern
Breukelman and Smith — Reptiles and Amphibians 107
Prairie Skink. — Chase Co.: 6 miles south of Clements; 1 mile south
of Saff ordville ; 11 miles southwest of Olpe (No. 348).
Diadophis punctatus arnyi (Kennicott), Prairie Ring-necked
Snake. — Bourbon Co.: 1 mile north of Fulton. Chase Co.: 5 mile*
southwest of Saffordville (No. 334)*; Elmdale (No. 146)*; 3 miles
west of Bazaar. Franklin Co.: 2.5 miles southeast of Peoria; 2
miles south of Lane. Linn Co.: 0.5 miles north of Trading Post.
Lyon Co.: 1.5 miles northwest of Reading (Nos. 6, 372)* ; Emporia.
Morris Co.: 5 miles south of Council Grove (Nos. 469-472). Neosho
Co.: 4 miles northwest of Erie (No. 316).* Osage Co.: 8 miles
southwest of Auburn. Shawnee Co.: 5 miles east of Topeka. Wa-
baunsee Co.: 2 miles northeast of Alma. Wilson Co.: 3 miles east
of Buffalo.
Carphophis amoena vermis (Kennicott), Western Worm Snake.
— Bourbon Co.: 6 miles northwest of Fort Scott. Chase Co.: 6 miles
southwest of Cottonwood Falls (No. 365).* Geary Co.: 5 miles
southwest of Wreford. Greenwood Co.: 4 miles northwest of La-
mont (Nos. 516, 517).* Johnson Co.: 3 miles east of De Soto.
Labette Co.: 9 miles northeast of Parsons (No. 313).* Linn Co.:
3.5 miles south of Pleasanton. Lyon Co.: 2 miles northeast of
Reading; 5 miles northwest of Emporia. Neosho Co.: 4 miles north-
west of Erie (No. 314).* Shaumee Co.: Wakarusa. Wilson Co.: 2
miles northwest of Neodesha (No. 322).*
Heterodon contortrix contortrix (Linnaeus), Common Hog-
nosed Snake. — Saline Co.: Coronado Heights; 3 miles northwest of
Lindsborg.
Heterodon nasicus nasicus Baird and Girard, Western Hog-
nosed Snake. — Chautauqua Co.: Peru. Ness Co.: 6 miles west
and 0.25 miles south of Ness City (No. 501)*; 5 miles northwest of
Ness City (Nos. 619,620).* Rooks Co.: Stockton (No. 418). Scott
Co.: Near Scott City (Nos. 511-513. 515).*
Coluber constrictor flaviventris (Say), Blue Racer. — Butler
Co.: 3 miles south of El Dorado. Chase Co.: 5 miles south of
Saffordville (Nos. 4, 110, 122-129, 656, 657).* Chautauqua Co.: 1
mile south of Chautauqua (No. 375).* Geary Co.: 5 miles south-
west of Wreford. Labette Co.: 7 miles northwest of Mound Valley
(No. 356).* Lyon Co.: 5 miles northwest of Reading (No. 226)*;
2 miles west of Olpe (No. 341)* ; 5 miles northwest of Emporia (No.
397)* ; 17 miles southwest of Emporia (No. 655).* McPherson Co.:
4 miles west of Roxbury. Morris Co.: 4 miles west of Delavan.
Neosho Co.: 4 miles northwest of Erie; 8 miles southeast of Chanute.
108 University of Kansas Publs.* Mus. Nat. Hist.
Ness Co.: 5 miles northwest of Ness City (No. 617).* Wilson Co.:
3 miles east of Buffalo; 2 miles northwest of Neodesha; 7 miles
northeast of Fall River.
Masticophis flagellum flagellum (Shaw), Eastern Coachwhip.
— Wilson Co.: 2 miles northwest of Neodesha (No. 302).* Elk Co.:
5: miles west of Grenola (No. 3).*
Masticophis flagellum testaceous (Say), Western Coachwhip.
— Ness Co.: 5 miles northwest of Ness City (No. 616).* Rooks
Co.: Stockton (Nos. 411, 412).*
Elaphe laeta laeta (Baird and Girard), Emory Rat Snake. —
Chase Co.: 5 miles southwest of Saffordville (Nos. 117-120, 130, 326,
354)*; Wolf Creek; 2 miles northeast of Strong City (No. 366).*
Coffey Co.: 7 miles east of Lebo. McPherson Co.: Lindsborg.
Morris Co.: 10 miles south of Council Grove (No. 230).* Saline
Co.: Salemsborg. Wilson Co.: 3 miles east of Buffalo (No. 161).*
Elaphe obsoleta obsoleta (Say), Pilot Black Snake. — Atchison
Co.: Atchison (No. 15).* Labette Co.: 4 miles north of Oswego
(No. 320).* Lyon Co.: Emporia (Nos. 12, 374, 514)*; 5 miles
northwest of Emporia (No. 337) ; 1.5 miles northwest of Reading
(No. 634).* Morris Co.: 0.5 miles north of Wilsey. Neosho Co.: 4
miles northwest of Erie (Nos. 321, 359).* Wabaunsee Co.: 4 miles
southwest of Alma. Wilson Co.: 7 miles northeast of Fall River.
Pituophis catenifer sayi (Schlegel), Common Bull Snake. —
Atchison Co.: Atchison. Chase Co.: 4 miles east of Elmdale; Toledo;
13 miles west of Emporia; Saffordville (No. 212).* Cherokee Co.:
4 miles southeast of Columbus. Coffey Co.: 6 miles west of Waverly.
Ford Co.: Bucklin (No. 405).* Franklin Co.: 2 miles southwest of
Lane. Hodgeman Co.: Jetmore. Jefferson Co.: 3 miles south of
Norton ville. McPherson Co.: Lindsborg. Morris Co.: 3 miles
southeast of Diamond Springs; 6 miles west of Council Grove; 4
miles west of Dwight; 3 miles north of Burdick; 3 miles east of
Delavan. Ness Co.: 4 miles west and 1.5 miles north of Ness City
(Nos-. 499, 500, 615).* Rooks Co.: 5 miles southwest of Stockton
(Nos. 409, 410).*
Lampropeltis calligaster calligaster (Harlan), Yellow-bellied
King Snake. — Butler Co.: U. S. Highway 54 near Greenwood
County line. Coffey Co.: 13 miles east of Emporia. Franklin Co.:
5 miles southwest of Ottawa (No. 207).* Lyon Co.: 8 miles east of
Emporia (No. 2)*; 3 miles east of Emporia; 3 miles southeast of
Olpe; southwest of Emporia (No. 216) ; 6 miles south of Plymouth
(No.- 22)*; 1.5 miles northwest of Reading (No. 633).* McPherson
Breukelman and Smith — Reptiles and Amphibians 109
Co.: Western edge of Lindsborg. Osage Co.: 4 miles northeast of
Overbrook.
Lampropeltis getulus holbrooki (Stejneger), Speckled King
Snake. — Chase Co.: 5 miles southwest of Saffordville (No. 109); 2
miles southwest of Elmdale (No. 363).* Hodgeman Co.: Jet-more.
Lyon Co.: 5 miles east of Emporia; 4 miles southwest of Bushong
(No. 200) .* Marion Co.: 4 miles east of Antelope (No. 10) .* Mor-
ris Co.: 1 mile east of Skiddy. Woodson Co.: Lake Fegan (No.
626).* Wilson Co.: 3 miles east of Buffalo (No. 162).*
Lampropeltis triangulum gentilis (Baird and Girard), West-
ern Milk Snake. — Chase Co.: 5 miles southwest of Saffordville (Nos.
121, 131, 406). 0 Gove Co.: Fair Grounds (No. 18). Greenwood
Co.: 4 miles southwest of Lamont (No. 376)°; 6 miles south of
Wilbur. Scott Co.: near Scott City (No. 510).*
Lampropeltis triangulum syspila (Cope), Red Milk Snake. —
Cherokee Co.: 3 miles east of Crestline (No. 559). Franklin Co.:
2 miles southwest of Lane (No. 174) ®
Sonora episcopa (Kennicott), Great Plains Ground Snake. —
Wilson Co.: 2 miles northwest of Neodesha (Nos. 303-305, 323-
325).*
Natrix erythrogaster transversa (Hallowell), Yellow-bellied
Water Snake. — Chase Co.: 6 miles south of Clements; 6 miles south-
west of Saffordville; 3 miles east of Cottonwood Falls; 10 miles east
of Matfield Green; 7 miles south of Plymouth (No. 287) ; Elmdale
Hill, 0.5 miles east of Elmdale (No. 291)*; 10 miles southwest of
Olpe (No. 343).* Lyon Co.: 9 miles south of Plymouth (No. 25) ;
Emporia (No. 30)*; 5 miles northwest of Emporia (No. 67) ; 1 mile
north of Hartford (No. 108)*; 7 miles southeast of Saffordville (No.
283).
Natrix grahami (Baird and Girard I, Graham Water Snake. —
Lyon Co.: Admire; 5 miles south of Plymouth (No. 19)*; 6 miles
east of Emporia (No. 40)*; 0.5 miles north of Hartford (No. 85)*;
2 miles east of Emporia (No. 208)*; Emporia (No. 588).*
Natrix rhombifera (Hallowell), Diamond-backed Water Snake.
— Lyon Co.: 1 mile south of Emporia (Nos. 218-225)*; 8 miles
northwest of Emporia (Nos. 28, 29, 240, 261)*; 2 miles southeast of
Emporia (Nos. 32-35)*; 5 miles northwest of Reading.
Natrix sipedon sipedon (Linnaeus), Common Water Snake. —
Barber Co.: 8 miles west of Medicine Lodge. Bourbon Co.: 1 mile
north of Fulton (No. 184).* Lyon Co.: 5 miles northeast of Em-
poria (No. 5)*; 9 miles south of Plymouth (No. 23)*; 1 mile west
110 University of Kansas Publs., Mus. Nat. Hist.
of Neosho Rapids; 2 miles southeast of Emporia (No. 142, 211)*; 9
miles northeast of Emporia (No. 41) ; 3 miles northwest of Emporia
(No. 66) ; 8 miles northwest of Emporia (Nos. 75, 78, 241, 254, 272)*;
5 miles south of Hartford (No. 86) ; 1 mile north of Hartford (Nos.
91, 100) ; 7 miles southwest of Emporia (No. 116) ; Emporia (No.
239). Morris Co.: 3 miles southwest of Council Grove. Shawnee
Co.: 4 miles east of Topeka (No. 31).*
Haldea striatula (Linnaeus), Southern Ground Snake. — Chero-
kee Co.: 3 miles east of Crestline (No. 317)*; 2 miles north of
Baxter Springs; 1 mile north and 4 miles east of Crestline.
Thamnophis radix radix (Baird and Girard), Plains Garter
Snake. — Chase Co.: 5 miles southwest of Saffordville. Lyon Co.:
Emporia (Nos. 209, 210)*; 1.5 miles northwest of Reading. Ness
Co.: 5 miles northwest of Ness City (No. 618).*
Thamnophis sauritus proximus (Say), Western Ribbon Snake.
—Chase Co.: 1 mile south of Saffordville (No. 340).* Lyon Co.:
2 miles southeast of Emporia (No. 38)*; 5 miles northwest of Em-
poria (Nos. 68-70)*; 12 miles southeast of Emporia (No. 215)*; 5
miles northwest of Reading (No. 229).* Wilson Co.: 3 miles east
of Buffalo.
Thamnophis sirtalis parietalis (Say). Red-sided Garter Snake.
— Barber Co.: 8 miles north of Medicine Lodge. Dickinson Co.:
1.5 miles northwest of Herington. Lyon Co.: 2.5 miles southeast
of Emporia (No. 39)*; 1 mile northeast of Emporia (Nos. 43-48)*;
5 miles northwest of Emporia (No. 71)*; 8 miles northwest of Em-
poria (No. 84).* Wabaunsee Co.: 2 miles northeast of Alma.
Tropidoclonion lineatum (Hallowell), Lined Snake. — Chase
Co.: Saffordville; 3 miles northeast of Bazaar. Labette Co.: 1 mile
north of Montana (No. 362).* Lyon Co.: Emporia; 9 miles south
and 5 miles west of Emporia. Marion Co.: 4 miles east of Antelope
(No. 11).* Morris Co.: 3 miles east of Woodbine (Nos. 518-520).*
Rooks Co.: 5 miles northwest of Stockton (Nos. 414, 415).*
Tantilla gracilis Baird and Girard, Slender Tantilla. — Cherokee
Co.: 3 miles east of Crestline (Nos. 540-5441. Geary Co.: 4 miles
south of Fort Riley. Wilson Co.: 3 miles east of Buffalo; 7 miles
northeast of Fall River; 2 miles northwest of Neodesha.
Tantilla nigriceps nigriceps Kennicott, Great Plains Black-
headed Snake. — Rooks Co.: 5 miles northwest of Stockton (No.
416) ; Stockton (No. 417). This is the northernmost known record
for the species.
Breukelman and Smith — Reptiles and Amphibians 111
Agkistrodon mokeson mokeson (Daudin), Southern Copper-
head.—Atchison Co.: Atchison (Nos. 201, 202, 573, 578)*; 5 miles
north of Atchison (No. 653).* Bourbon Co.: 6 miles northwest of
Fort Scott (No. 294).* Cherokee Co.: 1 mile north and 4 miles
east of Crestline (Nos. 165-170)*; 2 miles east of Riverton (No.
293).* Coffey Co.: 4 miles northeast of Burlington. Franklin Co.:
2 miles southwest of Lane (Nos. 187-192, 194).* Lyon Co.: 1.5
miles northwest of Reading (No. 7).* Wabaunsee Co.: 2 miles
northeast of Alma (No. 195).* Woodson Co.: Lake Fegan (Nos.
627,628,630-632,649).*
Sistrurus catenatus tergeminus (Say), Western Massasauga.
—Chase Co.: 5 miles southwest of Saffordville (Nos. 8, 26, 112, 113,
295)*; 3 miles southwest of Elko (No. 145)*; 11 miles northeast of
Matfield Green (No. 231)*; 8 miles south of Clements; 2 miles
southwest of Elmdale (No. 333) ; 10 miles southwest of Olpe (No.
344).* Lyon Co.: 10 miles south of Plymouth (Nos. 20, 121)*;
8 miles southwest of Emporia (No. 114)*; 5 miles northwest of
Bushong (No. 353)*; 11 miles northeast of Emporia (No. 474).
Wabaunsee Co.: Kansas State Highway 99 just north of Lyon
County (No. 641).*
Crotalus horridus horridus (Linnaeus). Timber Rattlesnake. —
Atchison Co.: Atchison (Nos. 204-206)*; 5 miles north of Atchison
(Nos. 642-652).*
Crotalus viridis viridis (Rafinesque) , Prairie Rattlesnake. —
Hodgeman Co.: Jetmore.
Sternotherus odoratus (Latreille), Common Musk Turtle. —
Cherokee Co.: 1 mile north and 4 miles east of Crestline (No. 171).
Kinosternon flavescens flavescens (Agassiz), Yellow Mud
Turtle. — Ford Co: Rattlesnake Creek 2 miles south of Bucklin (1
spec.).* Pratt Co.: 5 miles southeast of Pratt. Sheridan Co.:
Sheridan County State Park (No. 569).
Chelydra serpentina serpentina (Linnaeus). Common Snap-
ping Turtle. — Chase Co.: 10 miles southwest of Olpe (No. 345) ; 3
miles east of Cottonwood Falls; 5 miles northeast of Strong City.
Greenwood Co.: (1 spec.).* Lyon Co.: 1.5 miles northwest of
Reading (No. 336); 5 miles south of Plymouth; 10 miles north of
Emporia; Admire; 4 miles northwest of Olpe; Emporia. Sheridan
Co.: State Lake; 7 miles northeast of Quint er.
Terrapene ornata (Agassiz), Ornate Box Turtle. — Chase Co.:
14 miles southwest of Olpe ; 6 miles south of Clements ; 5 miles south-
west of Saffordville. Coffey Co.: 4 miles south of Gridley (No.
112 University .of Kansas Publs., Mus. Nat. Hist.
468) * ; 1 mile west of Agricola (No. 638).* Ellsworth Co.: 10 miles
south of Ellsworth. Greenwood Co.: (1 spec.).* Hodgeman Co.:
Jetmore. Lyon Co.: 6 mile* south of Plymouth; 8 miles southwest
of Emporia; 7 miles west of Olpe. Morris Co.: 5 miles northwest
of Council Grove; 1 mile east of Skiddy; 5 miles south of Council
Grove. Rice Co.: Sterling. Rooks Co.: Solomon River near Stock-
ton (No. 408).*
Terrapene triunguis (Agassiz), Carolina Box Turtle. — Coffey
Co.: 1 mile west of Agricola (No. 637).*
Chrysemys picta bellii (Gray), Painted Turtle. — Chase Co.:
Kahola Creek, near Morris County line. Dickinson Co.: 1.5 miles
north of Herington. Ford Co.: Rattlesnake Creek; 2 miles south of
Bucklin (1 spec.).* Lyon Co.: 3 miles north of Emporia; 6 miles
south of Plymouth. Ness Co.: 4 miles west and 1.5 miles north of
Ness City (Nos. 613, 614).* Sheridan Co.: Sheridan County State
Park (No. 570). Wilson Co.: 4 miles southeast of Buffalo. Wood-
son Co.: Owl Creek north of Yates Center (1 spec.).*
Pseudemys floridana hoyi (Agassiz), Toothed Turtle. — Green-
wood Co.: Holmer Creek south of Hamilton on Kansas State High-
way 99 (Mus. Nat. Hist.. Univ. Kans., No. 23148) .* This is the sec-
ond published locality for the species in Kansas; it has previously
been reported from a locality 5.5 miles northeast of Coyville, Wood-
son County (Burt and Hoyle, Trans. Kans. Acad. Sci., 37:198, 1934).
Pseudemys scripta elegans (Wied), Scribe Turtle. — Chase Co.:
7 miles southwest of Saffordville. Lyon Co.: 10 miles northwest of
Emporia; 7 miles south of Plymouth.
Amyda mutica (Le Sueur), Spineless Soft-shelled Turtle. — Mc-
Pherson Co.: Lindsborg.
Amyda spinifera spinifera (Le Sueur), Spiny Soft-shelled Tur-
tle.— Chase Co.: 10 miles southwest of Olpe; 7 miles southwest of
Saffordville (No. 351).* Lyon Co.: 5 miles east of Emporia. Ness
Co.: 5.5 miles northwest of Ness. Sheridan Co.: State Lake; 7 miles
northeast of Quinter.
□
21-2762
5 - N A - u
Kyphosis and other Variations in
Soft-shelled Turtles
BY
HOBART M. SMITH
MUS. C08SP. ZOOL
LIBRARY
flAR -8 !•
i:
University of ^Kansas Publications
Museum of Natural History
Volume 1, No. 6, pp. 117-124
July 7, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Dcnald S. Farner.
Donald F. Hoffmeister
Volume 1, No. 6, pp. 117-124
July 7. 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1947
21-6301
MUS. COL?. ZQOL
LIBRARY
MAR -8 I960
Kyphosis and other Variations in Soft-shelled Turtles
I
HQBAKT M. SMITH
Kyphotic (hump-backed) soft-shelled turtles have been known
for many years in Asia and America. Gressitt (Peking Natural
History Bulletin, 2 (pt. 4): 413-415, figs. 1-5, 1937)' has reviewed
accounts of such turtles, and recorded the anomaly in Amyda sinen-
sis (Wiegmann) and .4. steindachneri (Siebenrock) of Asia and in
unidentified species in the United States. Records of kyphosis in
American species apparently are few.
Three skeletons in the University of Kansas Museum of Natural
History demonstrate occurrence of the condition in at least 3 Amer-
ican species: Amyda emoryi (Agassiz), A. mutioa (Le Sueur) and
.4. spinifera (Le Sueur). The specimen of A. emoryi (Catalog No.
2219) was taken at Phoenix, Maricopa Co., Arizona, by Victor H.
Householder, on May 1. 1926. The second specimen, called to my
attention by C. W. Hibbard, was taken in 1936 from the Kansas
River at Lawrence, Douglas Co., Kansas, by Max Wheatley, to
whom I am indebted for the accompanying photographs and per-
mission to describe the specimen which he has added to the Mu-
seum's collections (No. 23230). The identity of No. 23230 is es-
tablished as A. mutica by the absence of spines (see fig. 3) and by
a number of cranial characters. The specimen of A. spinifera (No.
23026) is without locality data; its identification is verified by the
presence of spines on the front of the carapace.
In the specimen of A. mutica (see figures) the hump forms a
smooth, high curve, closely resembling the condition in Gressitt's
specimens of A. steindachneri (op. cit.: fig. 1). In the other two
the hump is lower and its apex forms a relatively sharp angle; in
the specimen of .4. spinifera the posterior face of the hump is more
nearly vertical than the anterior face. In A. emoryi the rear edge
of the apex is sharply inclined (at an angle of about 45°), whereas
the remainder of the surface slants at an angle of about 35°.
In the accompanying table of measurements of specimens in the
University of Kansas Museum of Natural History the height is
measured from the end of the rib opposite the highest elevation to
the crest of the elevation, by projected lines. The length is mea-
sured from the anterior border of the nuchal plate to the posterior
edge of the last costal plate. The width is measured from tip to
(119)
120 University of Kansas Publs., Mtjs. Nat. Hist.
tip of the longest ribs. Catalogue numbers of the specimens, with
indication of the localities of capture are as follows: Nos. 2215-9,
2803, 2824, 2837, Phoenix, Maricopa Co.. Arizona; Nos. 19459-60,
Ozark. Franklin Co., Arkansas; Nos. 2225-9, Lewisville, Lafayette
Co., Arkansas; Nos. 1867-70, 1874-6, 1879, 1881, 1930-1, 2666, 2761-
2, 2826, 2838-42, Devalls Bluff, Prairie Co., Arkansas; No. 16528,
Orange Co., Florida; Nos. 1872, 1878, 1943, 1964, Doniphan Lake,
Doniphan Co., Kansas; No. 2220, Douglas Co., Kansas; No. 23230,
Kansas River, Douglas Co., Kansas; No. 18159. Harper Co., Kan-
sas; No. 2757, Smoky Hill River, Trego Co., Kansas; No. 23026,
no data.
The three abnormal specimens vary in width/height ratio from
1.83 to 3.14. In the 37 normal turtles measured, the corresponding
ratio is 4.64 to 7.85. The ratio of 4.64 is possibly subject to correc-
tion since the shell tends to warp in some specimens, especially in
those retaining the skin about the periphery of the shell. The
warping does not produce' a marked convexity in transverse section,
but does so in longitudinal section. Accordingly the height as here
measured is little effected, and the comparison with width rather
than length of shell provides for the lesser error from warping.
There appears to be no close correlation of proportions with either
size or sex.
It is of interest that Arnyda ferox is the most distinctive in pro-
portions of the carapace. Its carapace is longer in relation to its
width than that of any of the other species. The average relative
length of the carapace of A. emoryi is intermediate between that of
A. ferox and the averages of ,4. spinijera and A. mutica, but the
overlap in range with the latter two is complete.
The cause for kyphotic anomalies is unknown. That it is ac-
companied by a greater degree of growth in the vertebral column
than in the periphery of the costal plates is obvious. There seems
to be no well-established accommodation for the difference in
growth, since the hump produced by it varies considerably in form.
There is no trend from small to large specimens in size of the
hump; large and small humps occur in both large and small speci-
mens. Accordingly it seems that the humped condition is developed
in the late embryo or early post-embryonic life, and does not later
change.
An apparently reasonable hypothesis is that the costal plates an-
kylose distally with the ribs early enough in embryonic life so that
anv differential in growth rate between them and the vertebral
Smith — Soft-shelled Turtles 121
column is translated into abnormal contortions of the body. Agas-
siz and others have shown that the costal plates normally do not
fuse with the ribs by the time of hatching; the fusion then does not
normally occur in the embryonic stage. Presumably, once fused,
the costal plates and vertebral column normally have equal growth
rates, since the height/width ratio does not change significantly
with increased size. It is well known that fusion takes place in
young specimens soon after hatching; in all skeletons examined of
this genus, from the smallest (62 mm. in length) to the largest (295
mm.), the fusion has occurred. Therefore, the normal time of fu-
sion must be approximately at the time of hatching.
Although costal plates and the vertebral column grow in direct
proportion to each other throughout life from a period shortly
after hatching, the vertebral column apparently grows more rapidly
than the costals shortly before and possibly also shortly after hatch-
ing, at least in kyphotic and probably also in normal specimens.
An exceptionally early date of fusion of costal plates and ribs would
thus result in a kyphotic condition, and it may well be assumed
that the earlier the fusion, the greater the hypertrophy would be.
Whether or not this hypothesis correctly accounts for kyphosis in
turtles can be ascertained only by further study.
Stejneger (Bull. Mus. Comp. Zool., 94: 12, 1944) regards the
presence of 8 neurals as opposed to 7 as an important peculiarity
of .4. mutica. The 42 specimens for which the number of neurals
is recorded reveals, however, that there is greater variation than
previously supposed: in 16 .4. mutica more than half (9) have 7
neurals and the remainder (7) have 8. Eight neurals were recorded
also in 2 of 18 spinifera, and in 1 of 7 A. emoryi. Seven neurals
are present in the single specimen of A. ferox examined.
It is of interest also that the number of costals, which has been
reported to be consistently 7 in New World species and 8 in Old
World species, varies markedly. In New World specimens, one
.4. mutica has 7 on one side, 8 on the other, and 8 occur on both
sides of one other (of a total of 16 examined). One of twenty A.
spinifera, and one of eight A. emoryi have 8; the single A. ferox
(Schneider) has 7. Accordingly the suggestion by H. M. Smith
(Field Mus. Nat. Hist., Zool. Ser., 23:19, 1939) that Platypeltis
Baur be resurrected for the American soft-shelled turtles on the
basis of the occurrence of only 7 costals, is untenable.
The generic allocation of American soft-shelled turtles has varied
considerablv in recent years: Smith (loc. cit.) uses Platypeltis;
122 University of Kansas Publs., Mrs. Nat. Hist.
Pope (Turtles of the United States and Canada, p. 343, 1939) uses
Trionyx Geoffroy; and Stejneger (op. cit., p. 8) uses Amyda Gebf-
froy. As stated above, use of Platypeltis at the present time is un-
warranted, since no constant difference has been discovered that
would support generic separation of Asiatic and American members
of this group. New World turtles should be placed either in Tri-
onyx or in Amyda, depending upon the interpretation of type desig-
nation for the latter name. Malcolm Smith (Bull. Raffles Mus. 3:2,
1930) and others have considered that, as a part of the original de-
scription, Geoffroy (Ann. Mus. Hist. Nat, Paris, 14:20, 1809) des-
ignated the type species of his new generic name Trionyx as aegyp-
ticus E. Geoffroy (= triunguis Forskal a well-recognized species).
Stejneger argues that Geoffroy did not adequately designate a type
from among the many species he treated in his genus Trionyx, and
that it remained for Fitzinger (Syst. Rept., p. 30, 1843) to select
one of these as a type; he chose coromandelicus Geoffroy, which is a
synonym of granosa Schoppff, a species belonging to a different
genus (as now recognized) from that to which triunguis belongs,
although Geoffroy had made the mistake of considering both groups
as members of his genus Trionyx. Now if Fitzinger's type designa-
tion is accepted, the name Trionyx is to be applied to that group
containing granosa (only one other form is known in the genus,
and both forms occur only in India and Burma), whereas the name
Amyda of Geoffroy {op. cit., p. 1) is applied to the genus (as now
recognized) which includes triunguis and some 20 other species of
Asia and North America. The type of Amyda is a typical member
from Asia (cartilagineus Boddaert). On the other hand, if Geof-
froy's type designation is accepted, the American forms (and the
others of that genus) would take the generic name Trionyx, of
which Amyda would be a synonym, and the genus to which granosa
belongs would take the name Lissemys Malcolm Smith (Fauna Brit.
India, Rept. Amph., 1:154, 1931).
Stejneger discussed the various aspects of this problem (op. cit.,
pp. 6, 7), and I can add nothing to his discussion. His arguments
for the acceptance of Fitzinger's type designation rather than that
of Geoffroy are well founded upon the statement of the Interna-
tional Rules of Zoological Nomenclature, while those of Smith arc
not. In weighing these two alternatives, the practical value of
maintenance of the "status quo" is not here important, for the
whole system of nomenclature in this field is completely upset; any
conclusive decision would be of great practical value and one al-
S.m ith — Soft-shelled Turtles
123
ternative holds no special, practical advantage over the other. Ac-
cordingly, it seems reasonable to consider the matter closed with
Stejneger's analysis, retaining Amy da for the American and related
species of soft-shelled turtles. That this assemblage contains nat-
ural subgroups that may warrant subdivision into other genera is
obvious, but to none of these will the name Trionyx be applicable.
Table of Data on Amyda
Species
Number
.Sex
Width
(mm.)
Length
(mm.)
Ratio,
width-
Height
(mm.)
Ratio,
width-
Neurals
Costals
length
height
cmoryi
2219*
2215
2216
c?
81
62
1.30
34
2.38
7
8
7
7
11
104
88
1.18
18
5.77
7
**
2217
2218
7
7
8
• t
106
93
1.14
21
5.04
7
• i
2803
9
150
132
1.13
28
5.35
7
7
1 (
2824
9
204
198
1.03
32
6.37
7
7
*'
2837
19460
9
7
8
7
"
97
77
1.26
14
6.93
7
ferox
16528
9
282
295
0.99
53
5.32
7
7
tnutica
2841
9
99
75
1.32
16
6.18
7
7
*4
23230*
101
78
1.29
55
1.83
7
7
ti
2838
9
106
79
1.34
17
6.23
7
7
"
1964
&
110
95
1.15
18
6.11
7
7
i i
2839
9
115
77
1.49
18
6.39
7
7
"
2840
9
115
85
1.35
17
6.76
8
7-8
• t
19459
131
106
1.23
20
6.55
7
7
1 1
2220
9
144
116
1.24
22
6.54
7
7
"
1874
162
137
1.18
32
5.06
7
7
■ <
1930
9
180
138
1.30
33
5.45
8
7
• i
1875 •
181
164
1.10
39
4.64
8
8
1 1
1881
1868
9
8
7
7
"
185
167
1.10
39
4.74
7
"
1876
9
190
177
1.07
33
5.75
8
7
< i
1870
9
194
166
1.27
35
5.54
8
7
' *
1943
98
76
1.29
18
5.44
?
7
spin if era
1872
129
101
1.27
17
7.59
7
7
• t
1931
&
148
102
1.45
26
5.69
7
7
"
18159
9
151
129
1.17
26
5.80
?
7
i *
1878
9
163
132
1.23
25
6.52
8
7
■ *
2225
9
165
131
1.17
21
7.85
7
7
. "
23026*
9
170
133
1.27
54
3.14
7
7
..
2227
9
191
175
1.09
39
4.89
7
7
• •
2228
1867
9
L96
207
167
164
1.17
1.26
7
7
7
n
26
7.58
7
"
2757
213
196
1.08
30
7.10
7
8
* '
2229
215
178
1.20
28
6.78
7
7
' '
2762
9
219
184
1.19
40
5.47
7
7
1 1
1879
223
187
1.19
38
5.87
7
7
■ i
2761
9
233
182
1.28
43
5.41
7
7
1 4
2666
234
208
1.12
42
5. .57
8
7
1 1
2226
9
239
215
1.11
38
6.29
7
7
"
1869
245
211
1.16
44
5.55
7
7
"
2842
245
219
1.12
45
5.44
7
7
41
2826
9
245
237
1.03
45
5.44
7
7
* Kyphotic
University of Kansas, Museum
Kansas.
of Natural History, Lawrence,
124
University of Kansas Publs., Mrs. Nat. Hist.
Figs. 1-3. Amyda mutica, Univ. Kans., Mus. Nat. Hist., No. 23230, Law-
rence, Kansas. All views approximately half natural size. 1, Frontal view.
2, Lateral view. 3, Dorsal view.
□
*C -5 19
NATURAL HISTORY OF THE
PRAIRIE VOLE
(Mammalian Genus Microtus)
BY
E. W. JAMESON, Jr.
University of Kansas Publications
Museum of Natural History
Volume 1, No. 7, pp. 125-151
October 6, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; Donald S. Farner, H. H. Lane,
Edward H. Taylor
Volume 1, No. 7, pp. 125-151
October 6, 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR. STATE PRINTER
TOPEKA. KANSAS
1947
21-6957
/j^* Zoolo,:.
vpEC -5 I
Natural History of the Prairie Vole
(Mammalian Genus Microtus)
By
E. W. JAMESON, JR.
CONTENTS
PAGE
Introduction 128
Methods 130
Molt 131
Food and Habitat 132
Types of cover 132
Cuttings 133
Food caches 134
Plants used as food and as cover 135
Associates 137
Nest and Burrows 137
External Parasites 138
Fleas (Siphonaptera) 139
Lice (Anoplura) 141
Mites (Acari except Ixodoidea) 142
Ticks (Ixodoidea) 143
Reproduction 144
Age classes 144
Fecundity 144
Size of litters 146
The breeding season 147
Summary 149
Literature Cited 150
(127)
INTRODUCTION
The prairie vole (Microtus ochrogaster) at Lawrence, Kansas, is
approximately 5^ inches in length, of which the tail comprises l1/^
inches, and weighs approximately l1/^ ounces. The color on the
dorsum is dark gray with a grizzled appearance from the mixture of
black and fulvous on the long hairs; the venter is paler, sometimes
pale fulvous or cinnamon. The animal is compactly built much as
are the other microtine rodents. The short legs and short tail, small
eyes and partly hidden ears, and heavy and flattened head all sug-
gest its semifossorial mode of life. The prairie vole spends most
of its time in an elaborate system of tunnels (some entirely below
the ground) and in almost hidden galleries in the dense grass.
Microtus ochrogaster can be separated from other voles in its
geographic range by a combination of several characters. The plan-
tar tubercles usually number five, although a few individuals with six
tubercles were found at Lawrence, Kansas. Microtus pennsylvanicus,
normally with six plantar tubercles, as Bole and Moulthrop (1942:
156) pointed out, sometimes has only five. Therefore, the number
of plantar tubercles alone is not a certain means for separating
pennsylvanicus from ochrogaster. The color of the venter of ochro-
gaster is usually fulvous or cinnamon instead of grayish as in penn-
sylvanicus, but there is variation in this respect too; some prairie
voles also have a grayish venter. The shorter tail of ochrogaster
will assist in establishing its identity where it occurs with pennsyl-
vanicus. The third upper molar has two closed triangles in ochro-
gaster and usually three in pennsylvanicus. The pelage of ochro-
gaster is coarse whereas pennsylvanicus has fine fur. Prairie voles
may be separated from pine mice (Pitymys nemoralis and P. pine-
torum) with which they are sometimes found, by the larger eyes,
less rusty color, and longer tail. The Cooper lemming mouse (Syn-
aptomys cooperi) differs from the prairie vole in having the upper
incisors grooved, and in possessing a shorter tail which approximates
the hind foot in length.
Of Microtus ochrogaster from Lawrence, Douglas County, Kansas,
average measurements of twenty-five adult males are: total length,
143 (121-167) mm.; tail, 32 (25-42) mm.; hind feet, 20 (17-22)
mm.; weight, 43 (38-55) grams. Twenty-five adult females from
the same place average: total length, 150 (131-170) mm.; tail, 33
(31-41) mm.; hind foot, 19 (17-21) mm.; weight, 45 (38-58) grams.
(128)
Jameson — Natural History of Prairie Vole
129
The prairie vole is found in suitable habitats in the central part of
North America. It has been recorded from Edmonton, Alberta, in
the northwest (Bailey, 1900:76), southeastward to Chesapeake, Ohio
(Bole and Moulthrop, op. cit. : 156), and in the southwest as far as
Ft. Reno, Oklahoma (Bailey, op. ci£.:74). See figure 1 showing the
known range of Microtus ochrogaster. Microtus ludovicianus, a close
relative of ochrogaster, has been taken along the southern part of
the boundary between Texas and Louisiana (Lowery, 1943:247).
The activities of voles, especially those of the genus Microtus, at-
tracted the attention of naturalists even in early times. Aristotle
(translated by Thompson, 1910) wrote: "The rate of propagation
of field mice in country places, and the destruction that they cause,
Figure 1. Range of the Prairie Vole (Microtus ochrogaster).
are all beyond telling. In many places their number is so incalcul-
able that but very little of the corn-crop is left to the farmer; and
so rapid is their mode of proceeding that sometimes a small farmer
will one day observe that it is time for reaping, and on the follow-
ing morning, when he takes his reapers afield, he finds his entire
crop devoured. Their disappearance is unaccountable: in a few
days not a mouse will be there to be seen."
Several early naturalists in this country commented on the fluctu-
ations in numbers of individuals, and on the breeding and feeding
habits of voles. Kennicott (1857) in an agricultural report on the
mammals of Illinois wrote about the breeding of the prairie vole.
He described its stores of plants and commented on the behavior
of some captives. Quick and Butler (1885) discussed the habits of
130 University of Kansas Publs., Mus. Nat. Hist.
Microtus ochrogaster as well as those of M . pennsylv aniens, Pitymys
pinetorum, and Synaptomys cooperi in Indiana, and described the
feeding and breeding habits of these species. Criddle (1926) gave an
account of the feeding and breeding habits of Microtus ochrogaster
in Manitoba, and Fisher (1945) published a short description of the
food and reproduction of the same species as he observed it in Mis-
souri. Stone investigated the fauna in the nests of this vole in the
same state, but has not yet, as of March, 1946, published his findings.
METHODS
The information in the present account was obtained by observing
animals in the field, and by examining trapped animals that were
brought into the laboratory. Five hundred individuals were caught
in snap-traps, and forty additional voles that were marked were cap-
tured a total of 157 times. More than 90 per cent of the specimens
were trapped at Lawrence, Douglas County, Kansas, but voles were
examined also in Ellsworth, Atchison, and Jefferson counties, Kan-
sas, and in Douglas County, Illinois. My data pertain to Microtus
ochrogaster in the above named areas from October, 1945, until
August, 1946. The findings may not be typical of this species in
other areas and in other years.
The museum special traps were used both with and without bait.
The bait consisted of a piece of walnut meat on the treadle. By
placing the trap crosswise in the runway, voles were captured
whether or not the treadle was baited. Immediately upon removal
from the trap, each vole was placed in a white flannel sack, one sack
sufficing for several voles when necessary. In this way the loss of
ectoparasites was kept to a minimum. The fleas were counted, and
the numbers of lice and mites were estimated; some specimens of
ectoparasites were saved for identification.
The voles taken in live traps were marked and released. The
marking was done by cutting off one or more toes in such a manner
that the vole could later be identified. From left to right, the toes
were assigned numbers from one to five on the left hind foot, and by
tens from ten to fifty on the right hind foot. Number 33, therefore,
was assigned to the one vole of which the middle toe of each hind
foot had been cut off. Each time an animal was captured alive, it
was weighed, specimens of fleas, lice and mites were preserved, and
the external appearance of the reproductive organs was noted. The
extent of the molt line, if the vole was molting, was recorded. Cor-
responding data were kept for each dead vole caught in a snap trap.
Jameson — Natural History of Prairie Vole 131
Assistance is acknowledged from Professors E. Raymond Hall, A.
Byron Leonard, Worthie H. Horr, and Donald F. Hoffmeister; and I
have had also much helpful advice from Professors W. J. Hamilton,
Jr., and P. C. Stone.
MOLT
The skins of 44 molting prairie voles were pinned out flat. The
flesh sides clearly show the areas of molt. Various stages in the molt
process were observed also in animals caught in live traps. The
molt begins when the animal is three or four weeks old; at this time
the juvenal pelage is replaced by the subadult pelage. The second
molt occurs when the prairie vole is between eight and twelve weeks
old, and is the means by which the adult pelage replaces the sub-
adult pelage. These same two molts were found by Hatfield (1935)
to occur in captive Microtus californicus. Molting voles of the
species ochrogaster were trapped in each month of the year.
The molting processes of juveniles and subadults follow the same
pattern. The first area of molting is in the pectoral region. The
molt patch extends caudad toward the tail and cephalad toward the
chin. New pelage separates this area of active molt into two strips
on the fourth or fifth day. By this time each strip has spread laterad
to the legs and sides, and is 10 to 20 mm. wide. Ultimately each
strip unites with its opposite, usually at the center of the dorsum.
This area of molt then spreads cephalad and caudad. Fourteen to
fifteen days after the beginning of the molt, the entire dorsum is in
process of being covered with new pelage. Shortly before the com-
pletion of the molt, the dorsal area of molt divides into two patches,
one on the rump and one on the nape. The areas last to be covered
with new pelage are the crown and that between the ears and the
eyes. A slight variation in the above process occurred in some
specimens in which the lateral strips joined immediately cephalad
of the tail instead of at the center of the dorsum. The entire process
takes approximately three weeks.
Large voles (45 grams or more) grow hair in irregular patches
that measured 5 to 15 mm. In these large voles the molt is accom-
plished slowly and does not cover large areas of the body at any one
time. The small areas of molt are visible for 7 to 10 days, and were
found on approximately three quarters of the large voles examined.
132 University of Kansas Publs., Mrs. Nat. Hist
FOOD AND HABITAT
The diet of the prairie vole reflects both its environment and its
choice of food. The plants eaten are usually green and succulent,
but some dry, hard seeds and small stems of woody plants are also
eaten. The vegetation, which supplies the food for the vole, is im-
portant as cover or nesting material. For this reason food and
habitat are discussed together.
Types of Cover
Prairie voles inhabit areas where the dominant plants in summer
are clover or grasses or both. The lawn on the campus at the Uni-
versity of Kansas consists mostly of several kinds of grasses, but in
some places alfalfa (Medicago sativa) replaces clover {Trifolium
sp.), and in other places sedges (Scirpus spp.) are found in addition
to the grasses. The grass is short; it is mowed to a length of 4 to 6
inches. Bluegrass (Poa pratensis) and crabgrass (Digitaria ischae-
mum) form most of the sod. Bluejoint (Andropogon furcatus) is
common in a sparsely wooded part of the campus, an area which has
many voles. Foxtail (Setaria lutescens and S. viridis) and prairie
threeawn (Aristida oligantha) are also common on the lawn, but
these become dry in late summer, and at that time supply neither
food nor cover for the voles. The voles make well-beaten depres-
sions in the sod, and the grass arches over them to form canopies.
In the winter, when the snow flattened the grass on the campus
so that there were no longer protective canopies of blades over the
runways of the voles, they migrated into areas of Japanese honey-
suckle (Lonicera japonica). At this season the honeysuckle was
their main food. In areas where this vine was not available, the
voles abandoned their surface runways and remained below the
ground, coming to the surface only under the protection of a blanket
of snow. The voles returned to the grass and clover habitat in
March and April in 1946.
One pure stand of Ladino clover in Jefferson County, Kansas, was
studied in late November and early December of 1945. The clover
was 2 to 4 inches high, and although it was the sole food of the
voles, it furnishes but little cover. They were common here; 300
traps yielded 111 voles in two nights.
Jameson — Natural History of Prairie Vole 133
Cuttings
The voles seek particularly the tender heads of grasses and the
terminal leaves of sweet clover (Melilotus alba). To obtain these
parts, the voles begin by cutting through the base of the plant. The
surrounding plants are often near enough to support the freshly cut
piece in an upright position. The vole makes successive cuttings,
40 or 50 millimeters from the ground, until the desired parts of the
plant are within reach. The cuttings that have accumulated at the
base of the plant may be eaten, but frequently they remain as evi-
dence of the vole's feeding activity.
On May 12, 1946, an analysis was made of the cuttings found in
an area of alfalfa, grasses, and weeds. From table 1 it may be seen
that quackgrass, alfalfa, wild lettuce, and cleavers were common.
In three nights 70 traps caught 8 prairie voles and 3 deer mice; no
pine mice or cotton rats were caught on the area. The stomachs of
the voles and the deer mice were examined, and only the stomachs
of the voles contained green material. Analysis of the cuttings (see
table 2) indicates that alfalfa was eaten in greater quantity than
any other plant; it made up almost three quarters of the cuttings al-
though but one quarter of the cover. All other plants occurred less
commonly in the piles of cuttings than they did in the estimated
composition of the cover. Grasses and wild lettuce were next to
alfalfa in importance.
Table 1. — The Relative Abundance of Plants in an Area of Alfalfa, Grasses,
and Weeds *
Percentage by number
Species of plants
Quackgrass (Agropyron repens) 30
Speargrass (Poa annua) 1
California brome (Bromus carinatus) 1
Smooth brome (Bromus inermis) 1
Alfalfa (Medicago sativa) 25
Peppergrass (Lepidium densiflorum) 2
Cleavers (Galium aparinc) 15
Wild lettuce (Lactuca- scariola) 25
Table 2. — Composition of Ten Piles of Cuttings
Species
Agropyron repens 1
Poa annua 0
Bromus carinatus 0
Bromus inermis 0
Medicago sativa 40
Lepidium densiflorum ... 0
Galium aparine 0
Lactuca scariola, 6
* Analysis made on May 12, 1946, on an area 20 x 80 yards, at Lawrence, Kansas.
* Each of the first ten vertical columns gives the composition of one pile of cuttings. The
last column gives the percentage of occurrence in the piles of cuttings of each species of plant
in the area. Place and date for data in table 2 same as for table 1.
Frequency of
Ten pili
?s of cuttings
occurrence
0
0
2
0
0
6
19
0
4
13
0
0
0
0
0
0
0
0
0
00
0
0
0
0
10
0
0
0
0
04
0
0
0
0
0
0
0
0
0
00
14
30
30
31
5
0
0
21
4
73
0
0
0
0
0
0
0
0
0
00
0
0
0
1
0
1
0
0
0
01
2
1
2
5
0
0
0
2
4
09
134 University of Kansas Publs., Mrs. Nat. Hist.
Approximately one out of every ten voles caught in snap traps had
a piece of plant in its mouth. Occasionally a vole took a piece of
food into a live trap. Evidently the food is not always eaten where
it is procured. Grasses of the genus Poa are the kinds most fre-
quently found in the mouths of dead voles. Bromus carinatus, B.
inermis and sweet clover (Melilotus alba) were found in the runways.
The pulpy fruit of the horse nettle (Solarium carolinense) was found
partly eaten, especially near the entrances to underground passages.
Food Caches
Caches of seeds and underground parts of plants are stored in sub-
terranean chambers. One lot of food was found stored on the surface
of the ground. Four times, piles of seeds in runways indicated the
species of plants which the voles were storing.
One underground cache was found on May 27, 1946, on the Uni-
versity campus, by John Evans, Richard Edgar, and the writer.
This cache was in a large chamber in a tunnel system of the prairie
vole, on an east-facing hillside of walnut trees, catalpas, and Ken-
tucky coffee trees. The oval chamber was 250 mm. wide, 400 mm.
long, and 200 mm. high. The roof, at its highest point, was 30 mm.
below the surface of the ground. There were two entrances to the
cavity, both on the downhill side. The cache consisted of eight
quarts of seeds (approximately 2,800) of the Kentucky coffee tree
(Gymnocladm dioica). The seeds were packed with earth and all
were well preserved. The site of this cache was in an area which
was shaded by a small coffee tree. A seed of this tree is spheroidal,
measures 17 mm. in width, and weighs 2 grams.
Several times in the fall of 1945, in the above-mentioned grove,
the writer found pods of the coffee tree lying in the runs of the
voles. These pods were sometimes entire, but more often they had
been gnawed; frequently only part of a pod remained, indicating
that the voles were storing or feeding upon the seeds, although the
possibility that the mice were storing food did not occur to the writer
at the time. Three times, seeds of other plants were found piled at
the entrances of the burrows of voles. Twice these piles consisted
of from 50 to 70 seeds of the common dandelion [Taraxacum
officinale). The third pile was composed of 20 seeds of the giant
ragweed (Ambrosia trifida) .
A pasture of Canadian bluegrass (Poa compressa) , wild millet
(Echinochloa crusgalli) , sedges (Scirpus spp.), and clover (Tri-
folium sp.) in Atchison County, Kansas, was examined in Novem-
Jameson — Natural History of Prairie Vole 135
ber, 1945. This area was the home of a dense population of prairie
voles. Wherever a path of the voles crossed a deep imprint of a
horse's hoof, there was a collection of cuttings from the horizontal
stems of the clover which bordered the runways. Some of the cut-
tings may have been made by lemming mice (Synaptomys cooperi)
which were also common in the area.
Several kinds of voles store food. Bailey (1920) wrote of the
caches of Microtus pennsylv aniens in North Dakota, where, in one
locality, this vole was known as the bean mouse. He stated that the
Indians dug up beans (Falcata comosa) and the tubers of the
Jerusalem artichoke (Helianthus tuberosus) which the voles had
stored. Lantz (1907:17) found a cache of the roots of wild morn-
ing glory (Convolvulus septum) laid away by Microtus pennsyl-
vanicus. Nelson (1893:140) wrote that, as winter approached,
Microtus operarius gathered small bulbous roots, sometimes storing
a peck or more in a single cavity. Fisher (1945) in Missouri found
a gallon of the fruits of the horse nettle (Solarium carolinense)
stored in a hollow stump by the prairie vole. Kennicott (1857:99)
found five or six quarts of roots of two species of spike-flower
(Liatrus), Helianthus, and various grasses among the winter provi-
sions of the prairie vole in Illinois.
Plants Used as Food and as Cover
Table 3 lists, according to their families, the species of plants
which the prairie vole was observed to use for food. The same
species are sometimes used as cover. The majority of the plants are
in three families: the grass family (Graminae), the pulse family
(Leguminosae), and the composite family (Compositae).
The grasses that supply the voles' food and cover are mostly Poa
(the bluegrasses) and Bromus (bromegrass, chess, or cheat). Poa
pratensis is a common lawn and pasture grass, P. annua is a weed
species. The bluegrasses begin to grow in late winter about Law-
rence, Kansas, and they remain green until late in the fall. During
this time, the voles eat the blades and heads of bluegrass, and make
their runways under the culms. The prairie voles utilize several
species of Bromus. Bromus inermis and B. carinatus are important
range and pasture grasses, but japonicus is a weed of little or no
economic value. These are soft, tender grasses, but, in contrast to
the bluegrasses, they become dry in midsummer, and are then un-
suitable as food. However, they continue to form a protection over
the runways of the voles.
136 University of Kansas Publs., Mus. Nat. Hist.
The legumes, which appeared to be most important to the prairie
vole, are clover {Trifolium spp. and Melilotus alba) and alfalfa
{Medicago sativa) . These plants are common in both cultivated and
feral states. They form a different type of cover from that made
by grasses. Voles, living in clover and alfalfa, do not make run-
ways as distinct as they do in grasslands. The clover and alfalfa
plants are branched and of a spreading growth form, whereas the
grasses have leaves which are appressed to the main stem. The in-
dividual grass plants grow close together, and a vole cannot run
through grass without trampling some of it. As voles use the same
paths repeatedly, the grass in their runs becomes flattened and dies.
There is sufficient room between the stems of the clover and alfalfa
plants to allow the voles to pass through without treading on the
stems. In such a habitat, vole runways are poorly developed, and
are difficult to find. Voles in grasslands feed in runways, as at-
tested by the piles of cuttings found in the runways and the nibbled
grass which borders them. Voles in clover or alfalfa feed at the
bases of the plants wherever the plants may grow. In the latter
type of cover the cuttings are rather evenly distributed.
Compositae formed a minor part of the cover in most of the habi-
tats studied. Many grasslands have a stand of dandelions; sow
thistle, wild lettuce, and ragweed were also common in some grass-
lands. The voles ate the leaves and sometimes the seeds and under-
ground parts of these plants.
Table 3. Plants Used for Food by the Prairie Vole
Graminae Solanaceae
Poa annua Solanum carolinense
P. compressa Boraginaceae
P. pratensis ~ ,.
Bromus inermis Gahum aPanne
B. carinatus Caprifoliaceae
B. japonicus Lonicera japonica
Andropogon. furcatus
Agropyron repens Compositae
Setaria lutescens Loctuca scariola
S. viridis Sonchus arvensis
T • Ambrosia trifida
Leguminosae A. artemsiifolia
Mehlotus alba Taraxacum officinale
Medicago sativa
Trifolium spp.
Gymnocladus dioica
Jameson — Natural History of Prairie Vole 137
ASSOCIATES
In the mixed areas of grassland and clover that were described
above, the cotton rat (Sigmodon hispidus) , the deer mouse {Pero-
myscus maniculatus) , and the little short-tailed shrew (Cryptotis
parva) were commonly caught in the runways of the prairie vole.
Less frequently trapped were the common mole (Scalopus aquati-
cus), the large short-tailed shrew (Blarina brevicauda) , the Cooper
lemming mouse [Synaptomys cooperi) , the pine mouse (Pitymys
nemoralis), and the harvest mouse (Reithrodontomys megalotis).
In the dense growth of Japanese honeysuckle, the prairie vole shared
runways with the white-footed mouse {Peromyscus leucopus), the
large short-tailed shrew, and the pine mouse.
NEST AND BURROWS
The prairie vole makes a tortuous network of paths through the
grass and honeycombs the topsoil with its tunnels. The underground
passages lead to nests or to chambers where food is sometimes stored.
The runways through the grass are 40 to 50 mm. wide, and usually
lie slightly below the surface of the ground. By using the same
path repeatedly, the voles create little ruts in which they run. The
bottom of the runways are bare soil or are covered with only a thin
layer of trampled grass. Cotton rats, on the other hand, apparently
do not use their runs over long periods, for they are not well-beaten
runways, but are made merely by parting the grass and not by
trampling it down or cutting it off. Voles were trapped in runways
of the cotton rats, but no cotton rat was caught in a typical runway
of a vole.
The burrows of the prairie vole are 40 to 50 mm. in diameter, and
the shallowest part is usually 50 to 100 mm. below the surface of the
ground. Burrows leading to nests or food chambers may descend
deeper than the others. Some prairie voles were trapped in tunnels
of the common mole (Scalopus aquaticus). The voles make their
own burrows, and are especially active at this task when a hard rain
has loosened the previously hard, dry soil. The rain in the first two
weeks of October, 1945, made the soil much more friable than it had
been at the beginning of the month, and the voles took advantage of
the favorable opportunity to construct many new burrows. In Oc-
tober, particles of soil were packed beneath the toenails of many
specimens.
In this time fifteen nests were found. They were 6 to 18 inches
below the surface of the ground, and two tunnels led from each nest
138 University of Kansas Publs., Mus. Nat. Hist.
to the surface runway. The nest cavities were spheroidal, and
measured 150 to 200 mm. horizontally, and 80 to 100 mm. vertically.
The floors were slightly concave and were covered with loose dirt
and a mixture of dried grass and one or two leaves. The remainder
of the cavity was filled with the dry grass of which the nest was
composed. Criddle (1926) stated that at Treesbank, Manitoba, this
vole makes its nests in the burrow systems of the pocket gopher
{Thomomys talpoid.es) ; and Kennicott (1857:98) found nests of
the prairie vole in old ant hills.
Each of two nests that had been recently occupied was placed in a
Berlese funnel, and in this way the arthropod fauna of the nests
was collected. The most common arthropods in the nests were mites
(parasitic, predaceous, and free-living) and springtails. Sowbugs,
centipedes, spiders, and fleas were also present.. Of these arthropods,
the laelaptid mites, one kind of tick, and one kind of flea have a
direct relationship with the vole. These parasites are the same
species which are found on the vole itself. The mites were Eulaelaps
stabularis (Koch) and Atricholaelaps glasgoivi (Ewing). One adult
tick, Ixodes sculptus Newman, was in one nest. The fleas, about a
dozen in each nest, were Ctenophthalmus pscudagyrtes Baker, the
flea most frequently found on the prairie vole.
EXTERNAL PARASITES
The pelage of prairie voles, pine mice, deer mice, and shrews forms
a habitat for many kinds of parasitic arthropods. The fleas, lice,
and mites from the prairie vole were collected, counted, and identi-
fied. The ectoparasites from the other small mammals living in the
same habitat as the prairie vole were also considered. Some ecto-
parasites begin to leave the host when it dies, and any counts of
ectoparasites made from snap-trapped voles may fall short of the
number which was on the animal when it was alive. The average
number of fleas recorded from live voles exceeds that found on snap-
trapped voles (see table 4). The numbers of lice and mites were
estimated, but selected voles were examined to obtain absolute num-
bers of these kinds of ectoparasites.
The fleas, lice, and mites were mounted on one inch by three inch
glass slides; the ticks were preserved in 70 per cent alcohol. Dr. E.
W. Baker identified the mites; Dr. R. A. Cooley and Dr. Glen M.
Kohls, the ticks; Dr. G. W. Wharton, the chiggers; and Dr. Gordon
F. Ferris, the lice. To each of these gentlemen I am grateful. The
fleas were identified by myself.
Jameson — Natural History of Prairif. Vole 139
Fleas (Siphonaptera)
The information on the average numbers of fleas on voles was ob-
tained from live-trapped and some snap-trappd voles. Fleas were
counted only on voles which were removed from the traps within
twenty-four hours after the traps had been last examined. The
average numbers of fleas found on prairie voles in this study are
given in table 4.
Table 4. Average Numbers, of Fleas on Prairie Voles*
Subadults Adults
Live-trapped voles 1.9 (73) 3.4 (29)
Snap-trapped voles 1.1 (26) 1.3 (27)
* The fleas on the live-trapped voles are all Cten&phthalmius pseudagyrtes Baker, and
those on snap-trapped voles represent several species (see table 2). The numbers in paren-
theses are the numbers of voles examined.
Table 5 shows the average degree of infestation for ten months of
an eleven month period. The monthly averages for the most part
show no variations. The latter half of February provides an ex-
ception in that a series of 22 snap-trapped voles and 11 live-trapped
voles taken at that time had on the average, 9.7 and 5.3 fleas re-
spectively. Pine mice (Pity my s nemoralis) occurred in small num-
bers in the area where Microtus ochrogaster was live-trapped, and
Ctenophthalmus pseudagyrtes was the flea found to be common on
both of these voles.
Table 5. — Monthly Averages of Fleas on Prairie Voles
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
.6 5.1 5* .... 3 1.8 1.4 1.7 .... 1.1 2 2
(6) (11) (6) .... (6) (88) (26) (6) .... (8) (14) (2)
* This figure is high because one vole had the high number of 19 fleas. The numbers in
parentheses show the number of live voles examined for each month. All fleas were Cteno-
phthalmus pseudagyrtes Baker.
Some fleas have a habitat preference as well as a host specificity.
As voles from different areas were examined, different kinds of fleas
were encountered. A population of free-living voles under observa-
tion on the Campus at Lawrence was parasitized only by Ctenoph-
thalmus pseudagyrtes. From 90 prairie voles collected in a field
of clover 4 miles northwest of Lawrence, the only species of flea
recovered was Orchopeas leucopus. In both places the prairie vole
was the most common mammal, but in the field of clover three deer
mice (P. maniculatus) also were trapped. In a third field, one mile
west of Lawrence, the prairie vole was host to both the above men-
tioned fleas. Here both the prairie vole and the cotton rate [Sigmo-
don hispidus) were common.
140 University of Kansas Publs., Mus. Nat. Hist.
The host distribution of fleas on seven small mammals which lived
in the same habitats as the prairie vole is given in table 6.
Table 6. — Frequency of Occurrence of Fleas on Seven Species of
Small Mammals*
v.
s
-2 '■ b
a ~ £
5 c p. a a -a
ft | § g .5 § I
S * I I 5 §
i •§ s S § g g
§i £ ? p s & ?»
s>
et
o eq Qn a, &s < a,
Orchopeas leucopus (Baker) 0 0 53 31 37 6 10
Orchopeas houmrdii (Baker) =
O. wickhami (Baker) 0 0 0 0 0 1 0
Nosopsyllus fascia tus (Bosc) 0 0 0 0 0 1 0
Epitedia wenmanni (Rothschild) 0 0 0 9 0 2 0
Rectofrontia fraterna (Baker) 0 0 0 0 0 1 0
Corrodopsylla hamiltoni (Traub) 47 8 0 0 0 0 0
Ctenophthalmus pseudagyrtes Baker, 0 38 0 0 4 25 53
Peromyscopsylla scotti I. Fox 0 0 0 6 0 0 0
Total number examined 34 13 34 35 57 414 21
* The numbers represent the percentage of each species which was parasitized by fleas. The
mammals were collected at Lawrence, Douglas County, Kansas, between October, 1945, and
June, 1946. These data are entirely from snap-trapped animals with the exception of those
from Microtias and Pitymys which are from both snap-trapped and live-trapped animals.
It is seen that some fleas are rather specific in their choice of hosts,
and that others are commonly found on two or more small mammals
in the same habitat. In each of these groups there are fleas which
have a habitat preference, that is to say, the flea lives on the host
when the host lives in a given habitat, but is absent when the host-
lives in another habitat.
Group 1: Fleas with a Host Preference
Epitedia wenmanni was found on the white-footed mouse {Pcro-
myscus leucopus) and only rarely on the prairie vole. Corrodop-
sylla hamiltoni was taken only from the two kinds of shrews
(Blarina brevicauda and Cryptotis parva). Fleas on shrews may
have a well-developed host preference. At any rate, Elton, Baker,
Ford, and Gardner (1931) found that Doratopsylla dasyenemus
rarely strayed from its normal host (Sorex araneus) to other small
mammals. Peromyscopsylla scotti was taken from the white-footed
mouse (Peromyscus leucopus), and had a habitat preference also.
It was found only on those white-footed mice which were trapped in
the woodlands at various places in Douglas County; white-footed
mice which were trapped in areas of brush were free of this parasite.
Jameson — Natural History of Prairie Vole 141
Group 2: Fleas Commonly Found on Two or More Kinds of
Small Mammals
Orchopeas leucopus was an outstanding example of this group. It
was the most common flea on the deer mouse, the white-footed
mouse, and the cotton rat. In certain areas it was common on the
two voles (Pitymys nemoralis and Microtus ochrogaster) . Cteno-
phthalmus pseudagyrtes is the most abundant flea on the two kinds
of voles and on the large shrew (Blarina brevicauda) , and was found
sparingly on the cotton rat.
Several kinds of fleas do not belong in either of the above groups.
Some fleas were accidental strays from mammals not included in
table 6; and one flea {Rectofrontia fraterna) may prove to be a com-
mon nest parasite. Orchopeas howardii is common on tree squirrels
(Sciurus niger and S. carolinensis) . Nosopsyllus fasciatus is a cos-
mopolitan flea on Rattus norvegicus. Rectofrontia fratema was
taken once from a prairie vole. Since the only specimens in the Uni-
versity of Kansas Entomological Collections are from "mouse nests,"
this flea may be found to be a nest inhabiting parasite.
Some fleas are possible bridges by which a blood parasite could be
transmitted from one kind of a mammal to another. If Ctenoph-
thalmus pseudagyrtes acted as the intermediate host of a disease-
causing organism, an epizootic from Microtus ochrogaster might be
transmitted to Pitymys nemoralis or to Sigmodon hispidus or Blarina
brevicauda. There are several other such potential bridges for blood
parasites. Although table 6 does not prove that individual fleas
wander from one host to another, the frequency with which the sev-
eral kinds of fleas are removed from live mice suggests that the fleas
occasionally do so.
Lice (Anoplura)
Lice collected from the prairie vole were all of one species, Hop-
lopleura acanthopus (Burmeister) . Of 59 voles examined for the
presence of lice, 33 were found to be parasitized; the 59 voles had
an average of 3.4 lice each. Other mice which used the same run-
ways as the prairie vole had their own species of Anoplura. The
cotton rat was host to Hoplopleura hirsuta Ferris, and the two
species of Peromyscus were parasitized by Hoplopleura hespero-
mydis (Osborn).
The writer collected Hoplopleura acanthopus from Microtus cali-
jornicus at Calaveras Dam, Almeda County, California, and from
M. pennsylvanicus at Ithaca, Tompkins County, New York. Elton,
142 University of Kansas Publs., Mxis. Nat. Hist.
Ford, Baker, and Gardner (1931) recorded this same species from
M. argestis in England.
Lice on the prairie vole are the same species as those found on
other species of Microtus in other areas, but since Anoplura of the
prairie vole do not parasitize the cotton rat, the white-footed mouse,
and the deer mouse, this host specificity of lice makes it unlikely
that lice would carry blood parasites from the prairie vole to any
of the latter named rodents.
Mites (Acarina except Ixodoidea)
Many of the small mammals examined in this study had mites,
some of which were collected and identified. Mites were collected
from other species of voles in several localities in the United States
and in one locality in Canada ; as voles in widely separated regions
are sometimes hosts to the same species of mites, these records will
be presented here.
The frequency of some kinds of mites in the identified material
suggests that they are more abundant than other kinds. The occur-
rence of mites on small mammals from Lawrence, Kansas, is pre-
sented in table 7.
The following comments can be made concerning the specificity
and geographic ranges of several species of mites:
Liponyssus occidentalis Ewing was found only on Cryptotis parva.
Eulaelaps stabularis (Koch) was one of the more common kinds
found on the prairie vole. This mite is rather large (about 1 mm.
long) and is frequently (with the following species) seen running
through the pelage of its host. In addition to the records for this
species in table 1, it was found to be a common parasite on Pitymys
pinetorum at Point Abino, Welland County, Ontario. Elton, Ford,
Baker and Gardner (1931) found this same mite on Apodemus
sylvaticus and Clethrionomys glareolus in England.
Atricholaclaps glasgowi, like the preceding species, was one of the
commoner mites on the prairie vole. It was found also on Pitymys
pinetorum at Point Abino, Welland County, Ontario; on Microtus
pennsylv aniens at Ithaca, Tompkins County, New York; and on
M. californicus at Calaveras Dam, Almeda County, California.
Atricholaclaps sigmodoni occurred only on the cotton rat.
Laelaps kochi was less commonly found than Eulaelaps stabu-
laris and Atricholaclaps glasgowi. In Kansas the prairie vole and
the cotton rat were hosts to Laelaps kochi, and it occurred on
Jameson — Natural History of Prairie Vole 143
Microtus pennsylvanicus at Ithaca, New York, and on M. cali-
jornicus at Berkeley, California.
Trombiculidae are commonly known by their larvae which are
called chiggers or harvest mites. The white-footed mouse, the cot-
ton rat, and the prairie vole were parasitized at Lawrence. In the
winter these mites live in the ears of these small mammals, but in the
summer they were found both in the ears and on the rump. Those
obtained in winter were Ascoschongastia brevipes (Ewing) ; other
species may be involved.
Listrophoridae was represented on the prairie vole by a species of
Myocoptes and a species of Listrophorus. These mites cling to the
hairs of their host, and do not occur on the skin of the voles.
No evidence was seen that mites had any ill effect on the health of
their hosts. No voles had scabs on the skin; and the ears were not
swollen and disfigured as they sometimes are by chiggers. Al-
though the identity of a specimen of mite could not be determined
until it was mounted, a person could tell whether or not it was one
of the larger, very active Laelaptidae, one of the hair-clinging
Listrophoridae, or one of the tiny, orange Trombiculidae.
On July 12, 1946, three prairie voles were examined to determine
the number of mites they supported. The voles were freshly caught,
no one of them having been dead for more than five minutes before
they were examined. These three voles had an average of 25
Laelaptidae, 22 Listrophoridae, and 53 Trombiculidae.
Six species of mites (Ixodoidea excepted) were found on the
prairie vole. Four of these were collected also from other small
mammals living in the same habitat as this vole. Two species of
mites were found to occur on voles in New York, Kansas, and Cali-
fornia.
Ticks (Ixodoidea)
Two kinds of ticks were found. One adult specimen of Ixodes
sculptus Neumann was clinging to the head of a vole, just in front
of its eye. This species of tick was taken also from the thirteen-
lined ground squirrel (Citelhis tridecimlineaus) at Lawrence. One
nymph of Dermacentor variabilis (Say) was found attached to the
scapular region of a prairie vole. Both of these specimens were
taken in June.
144 University of Kansas Publs., Mus. Nat. Hist.
Table 7. Host Distribution of Mites on Seven Small Mammals*
c
ec
3
s
ec
■S
c
c
^
-*
rs
O
ft
s
V.
j
ft
•5
v.
•c
^
<~
c
s
cr
>
•a
P.
5
s
c
c
CO
t
a;
ft,
c
tz,
o
Ascoschongastia brevipes (Ewing) . . .
Liponyssus occident-alis Ewing
Eulaelaps stabularis (Koch)
Atricholaelaps glasgowi (Ewing) ....
Atricholaelaps sigmodoni Strandtmann
Laelaps kochi Oudemans
Myocoptes sp
Listrophorus sp
X
X
X
X X
X
X
X
X
X
X
X
X
X
X
* These data are from material collected at Lawrence, Douglas County, Kansas.
REPRODUCTION
Age Classes
Each prairie vole was assigned to one of three age classes (juvenile,
subadult, or adult) principally on the basis of weight, but partly on
the quality and color of the pelage. The three age classes are char-
acterized in table 8.
Table 8. Characters of Juvenile, Subadult, and Adult Prairie Voles
Juvenile
Less than 21 grams
Weight usually less
than 20 grams
Entire pelage dull
Dorsal color black
Subadult
21-38 grams
Average weight
30-32 grams
Pelage of rump
dull; rest of
pelage glossy
Dorsal color
grizzled except
on rump
Adult
38 grams or more
Average weight
40-45 grams
Pelage usually
entirely glossy
(rump sometimes dull)
Entire dorsal color
grizzled except
sometimes on rump
Fecundity
Hamilton (1941:4) found for Microtus pennsylv aniens that mac-
roscopic tubules of the cauda epididymis were an indication of
fecundity. By noting the size of the tubules (whether macroscopic
or not) and by making smears from them in approximately every
25th male caught, I found that the presence of sperm was positively
correlated with large-sized tubules of the cauda epididymis in Micro-
tus ochrogaster. Inferentially, males with sperm were fecund.
There is a relationship almost positive between the size of the
tubules of the cauda epididymis and the length of the testes. Testes
Jameson — Natural History of Prairie Vole
145
longer than 7 mm. have macroscopic tubules in the cauda, and in
testes shorter than 7 mm. these tubules cannot be seen with the
naked eye. Hamilton (1937b) found that in M. pennsylvanicus
>
100
90
80
70
60
50
o 40
B
o>
o
£ 30
o
a
m
°- 20
10
\
^uL
""•?
Oct
Nov.
Dec Jan
1945 1946
Feb
Mac
Apr
May
June
July
Aug
Figure 2. Fecundity of Prairie Voles by Months. Adults and Subadults are
Considered Together.
Z 7
«
E
= 6
E
.£ 5
- 4
it
* 3
_L
_L
Oct.
Nov
Dec
1945
Jan
1946
Feb
Mar
Apr.
May
June
July
Aug.
Figure 3. Seasonal Changes in the Length of Testes.
146 University of Kansas Publs., Mus. Nat. Hist.
testes smaller than 8x4 mm. did not contain sperm. The testes of
the prairie vole descend into the scrotum in the breeding season. In
the two winter months, when the voles did not bring forth young, the
testes decreased in size (see figure 3) and were withdrawn into the
body cavity. The presence of the testes in the body cavity does
not mean that a vole is not in breeding condition, for many speci-
mens with abdominal testes were fecund.
The females were considered to be fecund if they were gravid, or
if there were placental scars in the horns of the uteri.
Size of Litters
The number of mammae characteristic of a species of vole may
be a rough guide to the average size of a litter for that species.
The prairie vole has fewer mammae (three pairs) than some other
voles in North America, and might, therefore, be expected to have
smaller litters. Fifty-eight gravid females of Microtus ochrogaster
examined by me had an average of 3.4 embryos each; the number
of embryos ranged from one to seven. Hamilton (1936a) gave 5.07
as the average number of young per litter in M. pennsylvanicus.
Hatfield (1935) stated that M. californicus has an average of 5.7
young in a litter. Both pennsylvanicus and californicus normally
have four pairs of mammae. The expectation as to the size of the
litter seems to be realized. In the prairie vole one pair of mammae
is pectoral and two pairs are abdominal. Usually a lactating vole
showed evidence of only the abdominal mammae having been in use.
The size of litters was found to vary with the season of the year
(see table 9). Gravid females were collected in every breeding
month except September.
Table 9. Average Size of Litters of Microtus ochrogaster by Months*
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
0 2.8 3.9 3.2 3.4 3.1 2.8 3.0 ... 3.2 2.6 0
.. (4) (10) (6) (8) (9) (5) (2) ... (5) (5) ..
* These months are from October, 1945, until August, li)46. The numbers in parentheses
indicate the number of gravid females collected each month.
Table 9 shows that the prairie vole produced the largest litters in
March. A comparison of table 2 with figure 2 shows that the largest
litters were produced at the height of the breeding season. Baker
and Ransom (1933), studying Microtus agrestis, also found that
larger litters were characteristic of the height of the breeding season;
and that at the beginning and at the end of the breeding season the
litters averaged smaller.
Jameson — Natural History of Prairie Vole 147
The size of litters varied also with the age of the female. To place
a gravid female in its proper age class, the weight of the embryos
was subtracted from the total weight, and the remaining weight was
used as the body weight. The average size of the litters of 14 sub-
adults was 2.9, and in 35 adults it was 3.4. Hatfield (op. cit.)
found that the younger females of M. californicas gave birth to
smaller litters than did the adults.
Not included in either of the above analyses are nine gravid
females collected in November in a pasture watered by an artesian
spring in Atchison County, Kansas. In this pasture there was a
high concentration of prairie voles, and the percentage of fecundity
was much higher than in Douglas County at the same time. In
November only 29 per cent of the female prairie voles in Douglas
County were fecund, as against 59 per cent in Atchison County. The
average number of embryos of these nine voles was 4.1. Data from
Atchison County are not included in table 9.
The Breeding Season
In October, 1945, when this study was begun, the prairie vole was
bringing forth young. In the winter of 1945-'46 at Lawrence, Kan-
sas, there was a cessation of reproduction. The reproductive activity
was measured in terms of the fecundity of the subadults and the
adults of both sexes. Figure 2 suggests that the decline was most
marked in December and January ; no gravid females were collected
in these two months, although two females trapped in the first week
of December were lactating. In October, November, and December,
85 per cent of the breeding females were adults. In October, 85 per
cent of the adult females were fecund, and in November, this figure
was 80 per cent. Reproduction at this season, in the females, it
appears, was largely a function of the adults. The proportion of
adults to the rest of the population was calculated for each month ;
and the monthly changes in relative numbers of adults is shown in
figure 4. In November, December, and January there was a scarcity
of adult voles in the population. The autumnal decline in repro-
duction occurred simultaneously with the disappearance of these
adults, and is thought to have been largely a result of it.
Reproductive activity began in February; and in this month one-
third of the females contained embryos, and 90 per cent of the males
were fecund. Reproduction reached its height in March when fecun-
dity for the females and males was 77 per cent and 100 per cent
respectively. In April both sexes showed signs of being less pro-
148 University of Kansas Publs., Mus. Nat. Hist.
ductive, and still later in the spring the percentage of fecundity re-
mained at slightly over 65 for both sexes, this figure being higher
for the males than for the females for any one month. From Jan-
uary to February there was a 30 per cent increase in the percentage
of adults in the population; and for this period, there was a 33 per
cent increase in the fecundity of both males and females. In Feb-
ruary, 80 per cent of the fecund females were adults. The breeding
in the late winter, as in the fall, is thought to depend upon the per-
centage of adults in the population. Hamilton (1937b) noted a
100
90
80
70
<B
S 60
3
O
O
w
U.
50
40
30
20
10
Oct
Nov.
Dec
1945
Jan
1946
Feb
Mar
Apr
May
June
July
Auo,
Figure 4. Seasonal Changes in the Numbers of Adults in Relation to the Total Population
of Prairie Voles.
similar correlation between winter breeding and dominance of adults
in Microtus pennsylv aniens in New York. Fisher (1945) found that
the prairie vole continued to breed throughout the winter of 1943-'44
in Missouri; in such a case, one would expect to find a large pro-
portion of adults in the population.
Throughout the winter of 1945-'46, at Lawrence, the majority of
males were fecund ; but fecundity in the females was much less, and
in January, no females showed signs of reproductive activity. From
this it appears that the females, not the males, limit the breeding
season of this species.
Jameson — Natural History of Prairie Vole 149
SUMMARY
In the eleven month period, October, 1945, until August, 1946, in
northeastern Kansas, more than five hundred specimens of the
prairie vole (Microtus ochrogaster) were examined in the flesh ; and
forty free-living voles were examined 157 times — an average of
slightly less than four times each.
There is a complete molt from juvenal to subadult pelage, and
one from subadult to adult pelage. These molts require three weeks
each. Subsequent molts are irregular and extend over longer periods
of time.
This vole, in summer, inhabits areas of grass, clover, and alfalfa.
In winter, habitats with some woody growth may be sought.
Twenty-two kinds of plants were found to be used for food. Al-
though most of these were succulent plants, seeds and small woody
stems were sometimes eaten. The prairie vole, like some other
species of Microtus, lays away stores of food, usually underground;
the maximum quantity found in one cache was two gallons.
Nine other species of small mammals occur in the same habitat
with the prairie vole, and frequently use its runways. The vole
makes a network of paths through the grass, and constructs its own
burrows which lead to its neste and food stores. Each of fifteen
nests found were underground. Most, if not all, of the underground
tunnels are dug when the soil is moist, not when the soil is dry.
The commonest flea on the prairie vole is Ctenophthalmus pseu-
dagyrtes; it averages 1.9 (for subadult voles) to 3.4 (for adult voles)
per individual vole. Other fleas on this vole are Orchopeas leucopns,
Orchopeas howardii, Nosopsyllus fasciatus, Epitedia wenmanni, and
Rectofrontia fraterna. The two species of fleas which were actually
common on the vole (C. pseudagyrtes and 0. leucopus) , parasitized
also some other small mammals which lived in the same habitat as
the vole. One species of sucking louse (Hoplopleura acanthopus)
and two kinds of mites {Laelaps kochi and Atricholaelaps glas-
gowi) which occur on the prairie vole in Kansas, occur also on
Microtus calif ornicus in California and on M. pennsylvanicus in
New York. Only three ticks (1 Dermacenter variabilis and 2
Ixodes sculptus) were found on the prairie vole.
Fifty-eight gravid females had an average of 3.4 embryos. Litters
at the height of the breeding season are larger than those at the be-
ginning and at the end of the breeding season. Reproduction in
Microtus ochrogaster ceased in December, 1945, in northeastern
Kansas, and the first evidence of reproduction in 1946 was observed
in February.
150 University of Kansas Publs., Mus. Nat. Hist.
LITERAURE CITED
Bailey, V.
1900. Revision of the American voles of the genus Microtias. N. Amer.
Fauna, 17:1-88, June 6, 1900.
1920. Identity of the bean mouse of Lewis and Clark. Jour. Mamm.,
1:70-72, November 28, 1919.
Baker, J. R., and Ransom, R. M.
1933. Factors affecting the breeding of the field mouse (Microtus agrestris).
Part 11. Temperature and food. Royal Soc. London Proc, (Ser. B)
112:39-46, November 1, 1932.
Bole, B. P., Jr., and Moulthrop, P. N.
1942. The Ohio Recent mammal collection in the Cleveland Museum of
Natural History. Sci. Pub. Cleveland Mus. Nat. Hist., 6-83-181, Sep-
tember 11, 1942.
C riddle, S.
1926. Habits of Microtus minor in Manitoba. Jour. Mamm., 7:193-200,
August 9, 1926.
Elton, C. S., E. B. Ford, J. R. Baker, and A. D. Gardner.
1931. The health and parasites of a wild mouse population. Proc. Zool.
Soc. London. 101:657-721, September 30, 1931.
Fisher, H. J.
1945. Notes on voles in central Missouri. Jour. Mamm., 26:435-437, No-
vember, 1945.
Hatfield, D. M.
1935. A natural history study of Microtus californicus. Jour. Mamm.,
16:261-271, November 15, 1935.
Hamilton, W. J., Jr.
1937a. The biology of microtine cycles. Jour. Agr. Res., 54:779-790, May
15, 1937.
1937b. Growth and life span of the field mouse. American Nat., 71:500,
September-October, 1937.
1941. The reproduction of the field mouse, Microtus peunsylvanicus (Ord).
Cornell Univ. Agr. Exp. Sta, Memoir 237, pp. 1-23, May, 1941.
Kennicott, R.
1856. The quadrupeds of Illinois. Part I, Rep. Commiss. Patents: Agricul-
ture, pp. 52-110, 1857.
Lantz, D. E.
1907. An economic study of field mice (genus Microtus). IT. S.D. A. Bull.
Biol. Surv., 31:1-64, October 28, 1907.
Lowery, G. H, Jr.
1943. Check-list of the mammals of Louisiana and adjacent waters. Occas.
Papers Mus. Zool., Louisiana State Univ., 13:213-257. November 22,
1943.
Jameson — Natural History of Prairie Vole 151
Nelson, E. W.
1893. Description of a new species of Armcola, of the Mynomes group, from
Alaska. Proc. Biol. Soc. Washington, 8:140-142, December 28, 1893.
Quick, E. W., and A. W. Butler.
1885. The habits of some Arvicolinae. American Nat., 19:113-118, Feb-
ruary. 1885.
Transmitted August IS, 1946.
□
21-6957
6
>*n\
nAR -4
THE POSTNATAL DEVELOPMENT OF
TWO BROODS OF GREAT
HORNED OWLS
(Bubo virginianus)
KY
DONALD F. HOFFMEISTER AND HENRY W. SETZER
University of Kansas Publications
Museum of Natural History
Volume 1, No. 8, pp. 157-173
October 6, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Donald S. Farner, H. H. Lane,
Edward H. Taylor
Volume 1, No. 8, pp. 157-173
October 6, 1947
University of Kansas
Lawrence, Kansas
PRINTED Bf
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1947
1-6958
IAR -4 ■
The Postnatal Development of Two Broods of Great
Horned Owls
(Bubo virginianus)
By
DONALD F. HOFFMEISTER AND HENRY W. SETZER
Opportunity regularly to observe at the nest the development of
young Great Horned Owls, Bvbo virginianus (Gmelin), under
favorable conditions, was afforded when a pair nested and reared
their three offspring in 1945 and one offspring in 1946 on the vine-
covered north wall of the Museum of Natural History at the Uni-
versity of Kansas. The observations here reported are based pri-
marily on the three young raised in 1945 when daily observations
were made. These have been supplemented by other observations
made of the one nestling in 1946. Unless otherwise stated, obser-
vations pertain to the nest and three young in 1945.
NEST SITE
In 1945 the nest was situated on a metal-covered cement ledge,
two feet wide and 48 feet above the ground, at the northeast corner
of the Museum Building. The nest was protected on the east by a
stone abutment of the building and on the south by the north wall
of the building itself. Here the nest could be observed at will
through a laboratory window without disturbing the birds. The
taking of notes was begun at the time of egg-laying and extended
to the time at which the young left the nest, February 3 through
April 26, 1945. In 1946 the owls nested farther down the north
side of the building, behind two cement pillars, approximately 25
feet above the ground. To examine the nest in 1946 it was neces-
sary to lower an observer down the side of the building by means
of a rope. Observations of this nest were never made more fre-
quently than every other day. The adult owls were first seen at
the nest on February 3, 1946; careful examination of the nest be-
gan when the one egg hatched on March 7 and continued until
April 25, shortly before the young owl left the nest.
One large cottonwood tree, used by the parent-owls as a landing
place whenever they were forced from the nest, was situated ap-
proximately 110 feet to the north and a five-story building was
located 80 feet farther to the north. Numerous smaller trees line
(159)
160 University of Kansas Publs., Mus. Nat. Hist.
the street to the east and there are some on the lawns around the
Museum. Also, there are about two acres of trees 225 feet west
of the nest-site where the parent-owls took refuge when forced from
the cottonwood tree.
The nest, if it can be called a nest, was no more than a few bare
branches of the Virginia creeper, which covers the side of the build-
ing, together with some excrement which the owls tended to push
to the periphery of the nest. For most of the time the three eggs
in 1945 lay directly on the metal which covered the ledge, because
there was no definite floor to the nest. The single egg in 1946 lay
on the cement shelf between the pillars and the wall of the building.
This laxity in nest building by Great Horned Owls apparently is
not uncommon (see Bent, 1938:300).
PERIOD OF INCUBATION
Incubation of the eggs probably began, in 1945, on February 5,
the day the first egg was laid. It has usually been assumed that,
in birds of prey, incubation begins when the first egg is laid. The
last of the three eggs was laid February 7. In 1946, the single egg
was being incubated on February 4. Since another egg had been
laid two or three days before this — a broken egg was found beneath
the nest and there were remnants of the egg in the nest — incuba-
tion may have started as early as February 1 or February 2. In
comparing these dates of initial incubation with other recorded
dates of nesting, only those from places at, or near, the latitude
of Lawrence, Kansas, in the central United States, should be ex-
pected to be approximately the same since the times of egg-laying
and incubation are progressively later in the year as approach is
made toward the polar region. Baumgartner (1938:279) has pre-
viously pointed this out.
The incubation period for the Great Horned Owl in the central
United States has usually been regarded as 28 to 29 days. In the
nest under observation in 1945, two eggs hatched on March 12 and
are assumed to be the first two eggs laid, with an incubation period
for each of 35 and 34 days, respectively, and the third egg hatched
on March 14 with an incubation period of 35 days. In 1946, the
single egg hatched on the 33rd day, assuming that incubation began
on February 2, for the egg hatched March 7. In the period of egg-
laying and also in incubation, the parent bird in 1945 was fre-
quently disturbed by persons who peered at it through the window.
Curious observers handled the eggs at least once and vigorous
HOFFMEISTER AND SETZER — GREAT HORNED OwLS 161
pounding by carpenters in the room adjacent to the nest frequently
flushed the adult bird but did not cause desertion of the nest. It
may be that such disturbances prolonged the incubation period.
However, in 1946, the brooding birds were undisturbed, yet the in-
cubation period was nearly as long. If an observer near the nest
exposed himself in the daytime to the incubating bird, the adult
flew, but exposure at 50 feet or more from the nest only caused the
incubating bird to remain alert on the nest. When flushed, the
parent usually returned to the nest within 15 minutes or less after
the observer withdrew. On the thirty-second and thirty-third days
of incubation in 1945, the crew of carpenters demolished partitions
within the building on which the owl was nesting, and within 15
feet of the nest itself. At first the adult would fly from the nest
at each outburst of hammering and, at one time, remained away
from the nest for more than two hours. After a few hours of inter-
mittent hammering, however, the parent bird remained on the nest
despite all the noise produced. These observations bear out, rather
than refute, Baumgartner's statement (1938:281) that "the horned
owl incubates very closely," for a strong stimulus was necessary to
keep the owl from covering the eggs.
The egg hatched on March 14, 1945, and approximately two days
later than the. other two, is judged to be the one laid last. This
owl, III, was always 5 to 21 per cent lighter in weight than the older
birds when weights for corresponding ages were compared. Whether
this difference was the result of a lack of food because of dominance
of the two older birds, or because of a sexual difference, we do not
know. The owl that hatched in 1946 was likewise markedly lighter
than the first two birds hatched in 1945 (figure 1). A series of
adults from Meade County, southwestern Kansas, shows a pro-
nounced secondary sexual difference in weight. In this sample the
mean weight of 17 males, 1,208 grams, was 21 per cent less than that
of 25 females, 1,531 grams.
GROWTH OF JUVENILES
The principal measurement of growth taken by us was the weight
of the owls. In 1945 each of the three owls was weighed daily, with
two or three exceptions when a 48-hour period was interposed be-
tween weighings. The young were removed from the nest to a
nearby balance, weighed, and examined. The owl last hatched
(owl III) was weighed on the first day of life and on most subse-
quent days. The other two owls (designated as owls I and II)
162 University of Kansas Publs., Mus. Nat. Hist.
were first weighed when they were between 53 and 60 hours old.
On some days the birds were weighed twice, once in the morning
and once in the late afternoon; on most days, they were weighed
only in the late afternoon. The owl hatched in 1946 was weighed
when seven days old and at irregular, but usually two day, intervals
thereafter. It was weighed always slightly before midday.
The growth of the four owls is well shown by the changes in
weight recorded in figure 1. For the period during which the
young owls remained at the nest, growth can be divided into two
,omO 5 10 6 20 25 30 35 40 45 ,0,.
<cOO- I i i i I i i i i I i i i i i i i i i i i I i i i i i i i t i i i i i i i i i !?00
Owl
Owi n
Owl ffl
Owl EZ } 1946
AGE IN DAYS
JI50
.1100
.1050
JOOO
L950
1900
.850
.800
t.750
.700
1650
:600
L550
1.503
-450
1400
350
300
1250
1200
1150
JOO
150
1 1 1 1 1 1 1 1 1 1 i 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
0 5 10 15 20 25 30 36 40
1 1 1
45
Fig. 1. Growth of four Great Horned Owls as shown by changes in weight from near the
time of hatching until the time of leaving the nest.
HOFFMEISTER AND SETZER — GREAT HORNED OWLS 163
phases: (1) a rapid increase in weight during the first 3% or 4
weeks while the parent birds are supplying the young with ample
food; and (2) a subsequent period of slower growth, marked by
fluctuations (actual losses as well as gains) in weight resulting from
the failure of the parent birds to provide an ample supply of food.
If there is an initial period of about one week in postnatal develop-
ment in which there is a rather slow gain in weight, as suggested
by Sumner (1933:284), it was poorly marked in this instance. Owl
IV remained at the nest until the 50th day of age, and on the 47th
and 49th days (not shown on chart, fig. 1) weighed 1,011.4 grams
and 971.4 grams, respectively. By this age, the growth curve had
definitely flattened out. The fact that owl IV was consistently
heavier than owl III might be accounted for, in part, by the fact
that owl IV was always weighed in the morning when it was gorged
with food. However, Riddle, Charles, and Cauthen (1932) have
pointed out that when there were two or more pigeons in a nest,
each grew less rapidly than if there was only one present.
Within about 12 hours after hatching, the smallest of the three
owlets (III) weighed 48.7 grams. During the first four weeks of
postnatal growth, each owl gained in weight, daily, an average of
SSYs grams or an increase of 11.1 per cent. Owl I gained an average
of 36.1 grams each day, or a daily increase of 10.7 per cent; owl II,
37.8 grams, or 11.2 per cent; and owl III, 26.1 grams, or 11.4 per
cent. From the beginning of the fifth week until the time the young
left the nest, the three owls gained on the average only 12.7 grams or
approximately 1.6 per cent in weight daily. Individually, the daily
mean increases were as follows: I, 9.6 grams or 0.93 per cent; II,
12.7 grams or 1.86 per cent; III, 15.8 grams or 1.97 per cent. Prior
to the twenty-sixth to twenty-eighth day of age, there seldom was
any loss in weight from day to day, whereas after this period, ap-
proximately one weight in four was less than on the previous day.
These data support the earlier statement that during the first 3%
or 4 weeks, there is a relatively uniform and rapid increase in weight
whereas after this period weight fluctuates.
Growth as measured by changes in weight in these young Great
Horned Owls parallels growth in some other young birds. For ex-
ample, nestling Red-tailed Hawks, as reported upon by Fitch, Swen-
son, and Tillotson (1946:215), increase in weight rapidly for about
the first three weeks and then more gradually. Sumner's (1929b)
graphs indicate the same pattern of growth in the Barn and Great
Horned owls and Red-tailed and Cooper hawks. Pigeons, judging
164 University of Kansas Publs., Mrs. Nat. Hist.
from the growth curves for bodily weight as given by Riddle,
Charles, and Cauthen (1932), increase in weight rapidly until some-
where between the twenty-fourth and thirty-second day of post-
natal development. However, in the Golden Eagle, the early part
of postnatal development is not one of rapid growth, judging from
Sumner's diagram (1929a:164), but after the fourth week there is
a rapid increase in weight. Graphs that Sumner (1929b) gives for
Sparrow Hawks, Long-eared Owls, and Screech Owls, indicate that
in these instances also the increase in weight during the first few
days of postnatal development was not so rapid as it was after the
end of the first week. Stoner (1935) indicates that in the young
Barn Swallow, increase in weight was most rapid between the fourth
and tenth days, with the young remaining at the nest until the
twentieth day. Much the same pattern of weight increase was noted
by Stoner (1945) in the Cliff Swallow. Huggins' (1940:228) sig-
moid curve for increase in weight in the House Wren indicates that
the period of rapid growth in this species does not begin until the
second day. Sumner (1934:284) cites other studies which he be-
lieves, for altricial birds, indicate three periods of growth, an initial
period of rather slow gain, a period of maximum increase in weight,
and a final period of fluctuations. As previously indicated, for the
Great Horned Owls under observation, and in some other species
as indicated by published growth curves, the initial period of slow
gain is lacking.
The period of a decelerated rate of growth in the young Great
Horned Owls is correlated with the occasional lack of food. The
parent birds, during this latter period, remain off the nest more of
the time during the day, and their failure to provide the young with
food may represent an attempt to force the young to become pro-
ficient in flight or to force them away from the nest site, which
amounts to the same thing. When only slightly more than a month
old, the young began to test their wings, springing into the air, and,
in general, becoming more active and alert. Sumner (1929b: 110)
has suggested some other possible reasons for the period of de-
celerated rate of growth.
Although there was a daily increase in weight in the early stages
of growth, there was a decided fluctuation in any twenty-four hour
period. On any given day, the young always were heavier in the
morning than in the afternoon (see figure 2) ; presumably they were
gorged with food early in the morning.
HOFFMEISTER AND SETZER — GREAT HORNED 0\VL.S 165
80CL
750.
700.
650
600.
550.
_i i i i , i .,750.
AM PM AM PM AM PM AM PM AM PM AM PM
28
15001
AM PM AM PM AM PM AM PM AM PM AM PM
29 30
31
1 2
28
29 30
31
1 2
MARCH
APRIL
MARCH
APRIL
Fig. 2.
Morning and afternoon weights of two Great Horned Owls. Note that in the
morning the owls weigh more than in the afternoon.
When the young left the nest, they were approximately three-
fourths grown. When owl I on the 44th day and owl II on the 45th
day left the nest, they weighed 1,120 and 1,139 grams, respectively, or
73 and 74 per cent, respectively, of the average weight of 25 females
(1,530.9 grains). Owl III weighed 943.3 grams on the 43rd day
and owl IV weighed 971.4 grams on the 49th day, or 78 and 80 per
cent, respectively, of the average weight of 17 mature males (1,207.7
grams) . Owl I left the nest 18 hours before owl II did. Owl III
attempted to leave when 43 days old, but for it coordinated flight
was impossible and the bird landed on the lawn after a 150-foot
glide. When attempting to take owl IV from the nest on the 49th
day, it sprang into the air and by gliding, aided by an occasional
flap, sailed more than 300 feet before alighting on the ground.
After we returned the owl to the nest, it immediately sailed forth
for even a longer distance. When attempt was made to pick up
owls III and IV after they had flown down to the ground, they
rolled over on their backs and used both claws and beaks de-
fensively. Such a reaction never was noted at the nest; there our
hands were inspected, and sometimes bitten by the owls as possible
sources of food, but the claws were rarely used offensively or de-
fensively.
Slightly elevated remiges and rectrices, still in the sheath, were
visible on the ninth day. Some remiges first ruptured the feather
sheath on the 14th day; nearly all of the primaries ruptured the
sheaths by the 19th day and the secondaries by the 20th day. The
166 University of Kansas Publs., Mus. Nat. Hist.
Table 1. — Changes with age in certain parts of a young Great Horned Owl
hatched in 1946.
(Measurements are in millimeters)
Age in days
19
21
26
33
37
39
49
Length of erupted portion
of "average" primary. . . .
6.0
10.0
26.0
93.0
87.5
99.2
Length of erupted portion
of "average" secondary. .
5.0
25.0
60.0
78.0
95.0
Length of erupted portion
of "average" rectrix
17.0
16.0
28.0
78.0
Length of exposed culmen
without cere
19.6
22.5
24.0
23.7
Length of total culmen
30.4
36.0
38.5
40.0
Length of femur.
69.0
87.5
89.5
amount of eruption from the sheath for primaries, secondaries, and
rectrices, as given in table 1, was determined by measuring the one
feather of, say, the secondaries, judged to be near the mean in de-
gree of eruption. The feathers of the wing at 21 and 47 days of age
are shown in figure 5. On the eighth day, or slightly before, the
white nestling down of the newly hatched bird was replaced by a
downy immature plumage, which was more yellowish than the pre-
ceding plumage. The development of the plumage in the birds
under observation was much the same as that recorded by Sumner
(1933) in Bubo virginianus pacificus Cassin.
Traces of the egg-tooth were retained until the ninth day in two
owls and until the 11th day in another. In owl IV, the egg-tooth
was lost sometime between the 9th and 14th day. Changes in the
length of the culmen are indicated in Table 1. The length of total
culmen of owl IV when 47 days old is slightly greater than the
average for three adult male owls from Lawrence, Kansas (40.0
mm. as contrasted with 39.2 mm. in the adults) . The length of ex-
posed culmen, without cere, in the same bird when 47 days old is
less than the average of this measurement in the three adults from
Lawrence (23.7 mm. as contrasted with 26.5 mm. in the adults).
The femur was measured on three occasions as accurately as pos-
HOFFMEISTER AND SETZER — GREAT HORNED OWLS 167
sible through the skin and flesh. The precise boundaries of the
femur could not be determined and the thickness of the skin and
certain muscle is included. These measurements are not given to
indicate actual length of the femur, but to indicate the relative
changes in length of this bone.
Remnants of the yolk stalk were clearly evident at seven days
of age (see fig. 5) in the owl hatched in 1946 and were still present
when the owl was last examined (49 days of age) just before the
young left the nest. The scablike remnant was not noted in the
three young hatched in 1945, but close inspection was not made to
see if it was present.
The instinctive reactions of young horned owls shortly after
hatching have been fully described by Sumner (1934). By the
third day our owls could raise their heads, but when a bird was
undisturbed its head lay on the nest floor and the wings were
slightly spread. The eyes of owls I and II opened at about 7%
days, those of owl III on the 6th day, and those of owl IV some-
time between the 7th and 9th days. After the eyes opened, the
head was held erect more of the time. The young responded with
''cries" when disturbed by handling, when stimulated by certain
movements of the parent, or by movements of our hands near their
heads, which suggested to them the possibility of being fed. Cries
were evident but weak in the unhatched, pipped, egg, but soon after
hatching increased in intensity, and beginning at six days of age
were replaced, in part, by the characteristic "bill-snapping" of more
mature birds. These cries of the young may serve, among other
things, for recognition, inasmuch as they were given when the parent
was inspecting the young. When the parent returned to the nest
and covered the young, after having been flushed, it sometimes
uttered a special note, "hut, hut, hut," much like the "cluck" note
of the hen of the domestic chicken. The young responded to these
notes with faint cries, in contrast to the loud cries signifying alarm
and possibly hunger, which they gave when the parents were absent
from the nest.
The first definite evidence that the young were attempting to feed
themselves was obtained when they were 23 days old. Frequently
thereafter, fresh blood was found on their beaks and claws, but as
late as the 34th day a parent was seen feeding them. That day,
after being flushed, a parent returned to the nest at 7 p. m., and be-
gan tearing away parts of a cottontail which had previously been
brought to the nest. Bones in the hind leg of the rabbit broke
168
University of Kansas Publs., Mrs. Nat. Hist.
readily under pressure of the parent's bill, and the three young
crowded in close, opening their bills widely and placing them around
that of the parent. Of cottontails, the only parts consistently un-
eaten were the upper cheek teeth and the supporting maxillae and
connecting palatal bridge.
FOOD BROUGHT TO THE NEST
In the 45 days that the young remained at the nest site in 1945,
ninety-one individuals of 16 different species of animals were
brought by the parent owls (table 2). Probably a few smaller
animals, of which we saw no traces, were caught and eaten at night.
In 1946, two additional kinds of birds were brought to the nest:
1 Baldpate (Mareca americana) and 1 Pied-billed Grebe (Podilym-
Tabijq 2. Number of food items brought to the nest by the Great Horned
Owls in 1945
Birds
Rock Dove (Columba livia) .... 32
Robin (Turdus migratorius) 6
Starling (Sturnus vulgaris) 4
Mourning Dove (Zenaidura
macroura) 10
Meadowlark (Sturntlla sp.) 3
Red-wing (Agelaius phoeniceus) . . 1
Bronzed Grackle (Quiscalus
versicolor) 1
Mockingbird (Mimus
polyglottos) 1
Birds
Brown Thrasher (Toxostoma
rujum) 1
Grasshopper Sparrow (Ammo-
dramus savannarum) 1
Coot (Fulica americana) 3
Sora (Porzana Carolina) 1
Blue-winged Teal (Anas discors). 1
Mammals
Sylvilagus floridanus 19
Rattus norvegicus 6
Microtus ochrogaster 1
bus podiceps) . The large number of Rock Doves in the list can be
explained by their abundance on the buildings on the University
campus, including the Museum building where some were nesting as
close as 100 feet to the owl nest. When the owls were less than a
week old, only small birds and mammals were brought (young Rock
Doves, Robins, Starlings, Grasshopper Sparrow, meadow mouse,
and Norway rat) . The first rabbit was brought when the owls were
eight days old.
After the 28th day, only 18 food items, or slightly less than 20
per cent of the total number, were brought to the nest. These last
18 food items brought after the owls were 4 weeks old were no
larger or bulkier than those brought in the previous 20 days. The
beginning of this period of reduced amount of food corresponds to
HOFFMEISTER AND SETZER — GREAT HORNED OWLS 169
the beginning of the second phase of growth characterized by marked
fluctuations in weight.
Fox squirrels (Sciurus niger) are abundant on the University
campus, yet there were no remains of this mammal at the nest.
This may be explained by the fact that fox squirrels are principally
diurnal and Great Horned Owls feed principally at night. Yet in
early February, 1946, when the owls were preparing the nest, they
frequently flew on and off the nest in the early twilight of evening
while one or two fox squirrels fed in the periphery of trees not more
than 25 feet away. Yet the owls flew off to the west and left this
source of food unmolested.
Whether both owls regularly attended the young we do not know,
for the adults were not distinctively marked. On March 17, 1945,
when weighing the young, one parent bird started to return to the
nest but was frightened away by the observer who at the same time
noted the other parent perched in an adjacent tree. This was the
first time two adults were seen at the same time near the nest. In
1946, two adult owls (presumably both were parents) were within
sight at one time when the young owl first sailed forth and landed
in a wooded area some 100 yards away.
SUMMARY
Great Horned Owls {Bubo virginianus virginlanus) have em-
ployed as nest sites the protruding shelves of the stone wall of the
Museum of Natural History at the University of Kansas for several
years. In 1945, daily observations were made on one such nest and
its three young, and in 1946 irregular observations were made on
another such nest and the one young owl. The incubation time for
the three owls, hatched in 1945, was 35 days for two of the young
and 34 days for the third; for the one owl hatched in 1946, the in-
cubation time was at least 33 days. Two owls were consistently
smaller; when these smaller two left the nest they were, respectively,
21 and 17 per cent lighter than the other two. The smaller two
were judged to be males because adult males in Kansas average
smaller by 21 per cent than adult females.
Growth of the nestling young is divisible into (1) a period of
rapid increase in weight during the first 25 to 28 days, and (2) a
subsequent period marked by gains and losses in weight. The fluc-
tuations in this latter period are correlated with a reduction in food
brought to the nest by the parent birds and with the development
of habits of flight, This second period may be considered to be a
170 University of Kansas Publs., Mus. Nat. Hist.
period of "weaning." By the forty-fifth day, the young owls are
able to fly short distances and thus are able to leave the site of the
nest permanently. At this time they are about three-fourths grown.
Ninety-one individuals of 16 species of birds and mammals made
up the food items brought to the nest in 1945. Two factors seem to
be concerned in the acquisition of prey: (1) its availability and (2)
appropriate size of the prey.
LITERATURE CITED
Baumgartner, F. M.
1938. Courtship and nesting of the Great Horned Owls. Wilson Bull.,
50:274-285.
Bent, A. C.
1938. Life histories of North American birds of prey (Part 2), Orders Fal-
coniform.es and Strigiformes. U. S. Nat, Mus., Bull. 170, viii + 482 pp.
Fitch, H. S., Swenson, F., and Tillotson, D. F.
1946. Behavior and food habits of the Red-tailed Hawk. Condor, 48:205-
237.
Huggins, S. E.
1940. Relative growth in the House Wren. Growth, 4:225-236.
Riddle, O., Charles, D. R., and Cauthen, G. E.
1932. Relative growth rates in large and small races of pigeons. Proc. Soc.
Exp. Biol, and Med., 29:1216-1220.
Stoner, D.
1935. Temperature and growth studies on the Barn Swallow. Auk, 52:400-
407.
1945. Temperature and growth studies of the Northern Cliff Swallow. Auk,
62:207-216.
Sumner, E. L., Jr.
1929a. Notes on the growth and behavior of young Golden Eagles. Auk.
46:161-169.
1929b. Comparative studies in the growth of young raptores. Condor,
31:85-111.
1933. The growtli of some young raptorial birds. Univ. California Publ.
Zool, 40:277-307.
1934. The behavior of some young raptorial birds. Univ. California Publ.
Zool, 40:331-361.
HOFFMEISTER AND SETZER — GREAT HORNED OWLS
171
Fig. 3. Young Great Horned Owls in nest. Two owls are 7, 12, 18, 32, and 3f> days of
age, respectively ; the third owl is about 2 days younger in each instance.
Fig. 4. Young Great Homed Owl hatched in 1946. The two lower pictures show the developing
facial mask. Photographs by Joao Moojen.
HOFFMELSTER AND SETZER — GREAT HORNED OWLS
173
21 days
21 days
Fig. 5. Young Great Horned Owl hatched in 1946. Upper row: Ventral views showing scar of
volk sac and ventral side of wing. Middle row: Ventral (left) and dorsal view of wing at 21
days. Bottom row: Ventral (left) and dorsal view of wing at 47 days. Photographs by
Joao Moojen.
21-6958
S^ N A - L
Additions to the List of the Birds
of Louisiana
BY
GEORGE H. LOWERY, JR.
LIBRARY
1AR -8 i9! !
HARVARB
University of Kansas Publications
Museum of Natural History
Volume 1, No. 9, pp. 177-192
November 7, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, H. H. Lane
Edward H. Taylor
Volume 1, No. 9, pp. 177-192
Published November 7, 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA. KANSAS
1947
21-6959
1AR -8 !«• i
Additions to the List of the Birds of Louisiana
By
GEORGE H. LOWERY, JR.
Oberholser's "Bird Life of Louisiana" (La. Dept. Conserv. Bull.
28, 1938), was a notable contribution to the ornithology of the Gulf
Coast region and the lower Mississippi Valley, for it gave not only
a complete distributional synopsis of every species and subspecies
of bird then known to occur in Louisiana but also nearly every rec-
ord of a Louisiana bird up to 1938. However, at the time of the
appearance of this publication, one of the most active periods in
Louisiana ornithology was just then beginning. The bird collection
in the Louisiana State University Museum of Zoology had been
started only the year before, and the first comprehensive field work
since the time of Beyer, Kohn, Kopman, and Allison, two decades
before, was still in its initial stage. Since 1938 the Museum of Zo-
ology has acquired more specimens of birds from Louisiana than
were collected there in all of the years prior to that time. Many parts
of the state have been studied where no previous work at all had
been done. Also in the last eight years some capable ornithologists
have visited the state as students at Louisiana State University,
and each has contributed greatly to the mass of new data now avail-
able. Despite the excellence of Oberholser's compilation of records,
it is, therefore, not surprising that even at this early date twenty-
four additions can be made to the list of birds known from Louisi-
ana. Furthermore, this recently acquired information permits the
emendation of the recorded status of scores of species, each pre-
viously ascribed to the state on the basis of comparatively meager
data.
The plan is to publish eventually a revision of the birds of Lou-
isiana which will incorporate all of the new information, but the
projected scope of this work is such that many years may elapse
before it is finished. The present paper is intended to record only
the more pertinent additions, particularly records that may be sig-
nificant in connection with the preparation of the fifth edition of
the American Ornithologists' Union's "Check-list of North American
Birds." There are numerous species for which Oberholser cited
only a few records, but of which we now have many records and
(179)
180 University of Kansas Publs., Mus. Nat. Hist.
large series of specimens. If, in such instances, the treatment given
in the fourth edition of the American Ornithologists' Union's
Check-list would not be materially affected, I have omitted men-
tion of the new material in this paper.
I am indebted to a number of ornithologists who have presented
their notes on Louisiana birds to the Museum of Zoology and who
have done much to supplement its collections. Outstanding among
these are Thomas R. Howell, Robert J. Newman, Sam M. Ray,
Robert E. Tucker, Harold E. Wallace, and the late Austin W. Bur-
dick. Their efforts in behalf of the Museum have been untiring. I am
grateful also to Thomas D. Burleigh and Jas. Hy. Bruns, both of
whom have played an integral part in our field activities in recent
years and without whose help much less would have been accom-
plished. John S. Campbell, Ambrose Daigre, James Nelson Gowan-
loch, Sara Elizabeth Hewes, E. A. Mcllhenny, Edouard Morgan,
and George L. Tiebout, Jr., have generously contributed notes and
specimens which are duly attributed in the following text. For as-
sistance in taxonomic problems, or for the loan of comparative ma-
terial, I wish to thank John W. Aldrich, Herbert Friedmann, How-
ard K. Gloyd, Alden H. Miller, Harry C. Oberholser, James L.
Peters, Karl P. Schmidt, George M. Sutton, J. Van Tyne, and Al-
exander Wetmore.
Sula sula sula (Linnaeus), Red-footed Booby
An immature individual of this species came aboard a boat of the Louisiana
Department of Conservation near the mouth of Bayou Scofield, 7 miles below
Buras, Plaquemines Parish, on November 1, 1940. It was captured by J. N.
McConnell, who delivered it to James Nelson Gowanloch of the Department
of Wildlife and Fisheries. The bird was then turned over to me in the flesh for
preparation and deposit in the Louisiana State University Museum of Zool-
ogy. It has since been examined by James L. Peters and Alexander Wetmore,
who confirmed the identification. This is the first specimen of the species
obtained in the United States. The only other record of its occurrence in
this country is that of individuals observed near Micco, Brevard County,
Florida, on February 12, 1895 (Bangs, Auk, 19, 1902: 395-396). To eliminate
possible confusion in the literature, attention is called here to the fact that
the above-listed specimen was erroneously recorded by an anonymous writer
(La. Conserv. Rev., 10, Fall Issue, 1940: 12) as a Gannet, Moms bassanus (Lin-
naeus).
Butorides virescens virescens (Linnaeus), Eastern Green Heron
No winter records for the occurrence of this species were available to Ober-
holser in 1938, the latest date cited by him being October 27. Recently, how-
ever, it has been noted several times in winter on the coast of Louisiana.
Kilby and Croker (Aud. Mag., 42, 1940: 117) observed it at the mouth of
Lowery — Birds of Louisiana 181
the Mississippi River, near Pilot Town, on December 25, 1939, and Burleigh
and I each obtained a specimen at Cameron on December 13, 1940. Another
was shot by me at the same place on February 2, 1946. The species is there-
fore of casual occurrence in the state in winter.
Dichromanassa rufescens (Gmelin), Reddish Egret
Although previously reported only as a casual summer visitor along the
coast, the Reddish Egret is known now to occur regularly in small numbers
during the winter. Since Oberholser (op. cit., 56) cited only one specific
record of occurrence in the state, all additional records are listed here. On
East Timbalier Island, one to three were seen daily, August 16-19, 1940, and
two to five were seen daily, November 15-17, 1940. In Cameron Parish, the
species has been noted as follows (Lowery, et al.) : two on December 14, 1940;
one on January 3, 1943; three on September 3 and two on November 4, 1944;
one on April 29, 1945. Several specimens were collected.
Plegadis falcinellus falcinellus (Linnaeus), Eastern Glossy Ibis
Plegadis mexicana (Gmelin), White-faced Glossy Ibis
Considerable confusion exists concerning the specific identity of the glossy
ibises inhabiting Louisiana. The fourth edition of the A. O. U. Check-list
(1931: 33) stated that falcinellus "breeds rarely and locally in central Florida
and probably in Louisiana." In 1932, Holt visited the marshes of Cameron
Parish in southwestern Louisiana where he studied the ibises nesting in a
large rookery. Later he definitely stated (Auk, 50, 1933: 351-352) that the
birds seen by him were Eastern Glossy Ibises (Plegadis falcinellus). It was
doubtless Holt's identification that influenced Oberholser to list falcinellus as
a fairly common local resident in the state (op. cit., 78). This, however, is
contrary to the evidence at my disposal. My associates and I have studied
thousands of glossy ibises in the marshes of southwestern Louisiana in the
past ten years. These observations include numerous field trips into the re-
gion where ibises are plentiful throughout the year, especially during the
breeding season. I have also visited a large nesting rookery in Cameron
Parish, the only one in the state known to me, and the one which I have
every reason to believe is the same colony visited by Holt in 1932. Although
Holt identified as falcinellus the birds seen by him at a nesting rookery in
Cameron Parish, I have never seen that species anywhere in Louisiana except
at Grand Isle, 150 miles east of Cameron, as henceforth noted.
In winter when the White-faced Glossy Ibis lacks the white on its face,
some difficulty might be encountered in differentiating that species from the
Eastern Glossy Ibis. The perplexing thing, however, is that Holt made his
observations in the nesting season when no possible confusion should exist;
also he was in the middle of a nesting rookery with birds close at hand on
all sides. This fact notwithstanding, the ibis nesting in the Cameron Parish
rookery (known locally as "The Burn") on May 28, 1942, was the white-
faced species (Plegadis mexicana), as evidenced by moving pictures taken by
J. Harvey Roberts and by specimens of varying ages collected at the same
time by me. In all, the Louisiana State University Museum of Zoology has
19 specimens of mexicana taken in Cameron Parish in April, May, November,
December, and January. Field records are available also for the months of
February, March, July, and September.
182 University of Kansas Publs., Mus. Nat. Hist.
Aside from Holt's statement, Oberholser had only five other records for
falcinellus in Louisiana, one being a market specimen with incomplete data
and therefore of questionable scientific value. The remaining four specimens
were taken by E. R. Pike near the mouth of the Mississippi River on Novem-
ber 13 and 17, 1930, and are now on deposit in the Chicago Academy of
Sciences. Recently I borrowed these specimens for reexamination with the
following results. The three taken on November 17, 1930, are mexicana and
not falcinellus as labeled and so reported by Oberholser. The single speci-
men taken on November 13 is, however, correctly identified as falcinellus.
Alexander Wetmore kindly examined the material for me and confirmed my
identifications. The occurrence of falcinellus in Louisiana thus hinged on
Holt's statement and one preserved specimen. However, on July 23, 1944, in
the marshes on Grand Isle, Jefferson Parish, Louisiana, I encountered a flock
of 12 immature ibises that impressed me by their blackness in contrast to the
color of glossy ibises with which I was familiar in Cameron Parish. Two
specimens were collected and both proved to be falcinellus.
Holt's published observations cannot be positively refuted, for we cannot
be sure that a colony of falcinellus did not exist in Cameron Parish in 1932,
nor that the portion of the rookery under his observation did not consist of
a segregated population of that species. However, ten years of field observations
by other ornithologists have failed to disclose the species which Holt considered a
common nesting bird in an area where we now know that only the White-faced
Glossy Ibis occurs. The fact that Holt specifically stated that he failed to
find the white-faced bird at any time in his stay in Cameron Parish is difficult
to explain, but this much is certain — the present known status of falcinellus
in Louisiana is that of only a rare and casual visitor.
Branta canadensis hutchinsii (Richardson), Hutchins Goose
Oberholser (op. tit., 89) cited only one Louisiana record for this goose.
The bird in question was shot but apparently not preserved. Consequently,
the status of the race on the Louisiana list was subject to question. Recently,
however, two typical specimens of hutchinsii were obtained in the state, one
by Edouard Morgan, near Lake Catherine, on November 7, 1942, and the
other by Herman Deutsch, four miles above the mouth of the Mermentau
River, on November 2, 1944. The former is displayed in the Louisiana Wild-
life and Fisheries Exhibit in the Louisiana State Museum, and the latter is
now in the Louisiana State University Museum of Zoology.
Oxyura dominica (Linnaeus), Masked Duck
A mounted specimen of this species was found by T. D. Burleigh and myself
in a sporting goods store in Lake Charles, Louisiana. Through the kindness of
Mr. Jack Gunn, owner, it was donated to the Louisiana State University
Museum Collection. The bird was shot approximately 25 miles southeast of
Lake Charles at Sweet Lake, Cameron Parish, on December 23, 1933, by R.
T. Newton. This is the first recorded occurrence of the species in Louisiana,
as well as one of the very few instances of its appearance anywhere in the United
States.
Lowery — Birds of Louisiana 183
Buteo lineatus texanus Bishop, Texas Red-shouldered Hawk
Although this race has been recorded previously only from Texas and north-
eastern Mexico, it appears to be of regular occurrence in southern Louisiana
in the fall and winter. The six specimens in the Louisiana State University
Collection, identified by Herbert Friedmann as texanus, are as follows: West-
over, November 25, 1937; Baton Rouge, October 20, 1936, November 1, 1938,
and September 3, 1940; University, November 14, 1942; Hoo-shoo-too, October
12, 1941 (Lowery, Tiebout, and Wallace). Another specimen, taken at Baton
Rouge on September 17, 1940 (Ray), was acquired by Louis B. Bishop, who
identified it as texanus.
Numenius americanus americanus Bechstein, Long-billed Curlew
Numenius americanus parvus Bishop, Northern Long-billed Curlew
Thirteen specimens of this species in the Louisiana State University Museum
have been identified subspecifically (in part by J. Van Tyne) as follows:
JV. a. americanus — 4 9 , Cameron, November 21 and 22, 1940, and December
5, 1942. N. a. parvus— 4 $ , 1 2 , Cameron, November 21 and 23, 1940, and
April 11 and October 31, 1942; 1 2, East Timbalier Island, August 18, 1940.
Three are intermediate in size and therefore not identifiable with certainty.
Contrary to published accounts, the Long-billed Curlew is a fairly common
migrant in certain parts of southern Louisiana. About seventy-five were
counted on the beach near Cameron on November 1, 1941, and twenty-five
were noted at the same place on December 6, 1942. Almost invariably a few
are present there during every month of the year.
Charadrius alexandrinus nivosus (Cassin), Western Snowy Plover
Charadrius alexandrinus tenuirostris (Lawrence), Cuban Snowy Plover
Oberholser (op. cit., 216-217) listed the Cuban Snowy Plover as a rare
transient in Louisiana, and cited only four definite records based on three
specimens. Our recent studies, however, have yielded twelve additional
specimens and a number of sight records, all of which indicate that the species
is a regular and sometimes common migrant in spring and fall. Eleven speci-
mens in the series are identifiable with certainty as examples of nivosus and
therefore constitute an addition to the state list. They were taken at East
Timbalier Island on November 15 and 16, 1940 (Burleigh, Lowery, and Ray),
at Grand Isle on March 27, 1943 (Burleigh), and near Cameron on November
20 and 21, 1941, April 3 and October 17, 1942, and September 3, 1944 (Burdick,
Howell, and Lowery). On April 29, 1945, Tucker saw twenty on the beach
near Cameron, but he did not obtain a specimen. A single adult male in our
series, taken on East Timbalier Island, on November 15, 1940 (Ray), is re-
ferable to tenuirostris.
Charadrius hiaticula semipalmatus Bonaparte, Semipalmated Plover
Oberholser (on. cit., 218) made special mention of the absence of definite
winter records for this species, but, in recent years, it has been noted on
numerous occasions in Louisiana in that season. For example, ten were seen
at Cameron on December 13, 1940, and the same number was noted there on
January 22 and 23, 1941 (Lowery, et al.). A specimen was shot at Cameron
on December 5, 1942 (Lowery) .
184 University of Kansas Ptjbls., Mus. Nat. Hist.
Charadrius wilsonia wilsonia Ord, Wilson Plover
Oberholser's single winter record for this species (op. tit., 220) has now been
supplemented by two others — fifteen birds seen and three collected at Cam-
eron on January 22, 1941 (Burleigh, Wallace, and Ray) ; one taken at the
same place on December 5, 1942 (Burdick).
Pluvialis dominica dominica (Muller), American Golden Plover
The presence of the Golden Plover on the northern Gulf coast in winter
already has been reported by Burleigh ("Bird Life of the Gulf Coast Region
of Mississippi," Occas. Papers Mus. Zool. La. State Univ., 20, 1944: 367), but
since there are no published instances of its occurrence in Louisiana at that
season, the following four specimens are noteworthy: two collected near
Creole by Lowery and Ray on November 21, 1940; two others shot at the
same place by Burdick and Tucker on December 6, 1942; and one seen, but
not taken, near Cameron on November 22, 1941 (Lowery, et al.).
Erolia bairdii (Coues), Baird Sandpiper
Since there is only one previous definite record of the occurrence of this
species in the state, the following records are significant. A male was ob-
tained by Burdick at University, 3 miles south, on October 25, 1942. I saw
three at the same place on October 29 and shot a male there on November
9. The only spring record is that of a bird seen by me at University, 1 mile
south, on May 16, 1945.
Steganopus tricolor Vieillot, Wilson Phalarope
Apparently the first definite record of this species in the state is that of an
adult female, in breeding plumage, shot by E. A. Mcllhenny at Avery Island,
Louisiana, on May 10, 1939, and later sent to the Louisiana State University
Museum of Zoology. A second specimen, a male in winter plumage, was
taken by Burdick 5 miles south of the University on September 12, 1943.
Limosa fedoa (Linnaeus), Marbled Godwit
This species was listed by Oberholser (op. cit., 271) as a very rare winter
resident along the Gulf coast region of southern Louisiana and he cited only
two records of occurrence in the state. The following additional records
should clarify its present-day status. In 1940 two were seen on East Timbalier
Island on August 19, eight on November 15, and seventy-five on both Novem-
ber 16 and 17. Three were seen near Cameron on November 21, 1941. In
1942, two were seen near Cameron on April 4, five on April 5, three on April
11, two on April 22, and one on April 23. Another was noted near Cameron
on October 7, 1943 (Lowery, et al.). A small series of specimens was taken
from the birds mentioned above. In connection with this species, it may be
of interest to note that the Hudsonian Godwit (Limosa haemastica) has not
been observed in Louisiana by me or my associates.
Geococcyx calif ornian us (Lesson), Road-runner
The Road-runner inhabits the northwestern part of the state where it has
been reported for many years by local residents. However, since confirma-
tion of its occurrence was lacking, previous publications on the birds of the
Lowery — Birds of Louisiana 185
state have not listed it. The first definite record is that of a bird killed near
Shreveport, on May 1, 1938, by an unspecified collector. Another was shot
four miles north of Keatchie, De Soto Parish, on July 9, 1943, by Delmer B.
Johnson, at that time field biologist with the Louisiana Department of Wild-
life and Fisheries. Both specimens are in the Louisiana State University
Museum. Johnson states that he has seen the species on a number of oc-
casions, specific records being in April and May, 1943, twelve miles east of
Mansfield, and two miles east of Logansport. Various reports of nests have
been received, but as yet no completely satisfactory breeding record for the
state has been obtained.
Columbigallina passerina pallescens (Baird), Mexican Ground Dove
The Louisiana State University Museum of Zoology now has a series of
21 specimens of Columbigallina passerina collected in Louisiana since the pub-
lication of Oberholser's book, in which only a few records for C. p. passerina
alone are cited. Examination of the new material reveals that eleven speci-
mens are clearly referable to pallescens, providing, therefore, an addition to
the avifauna of the state. As might be expected, pallescens prevails in the
western part of the state, although, at least occasionally, it migrates farther
east. The specimens identifiable as pallescens are as follows: 7$, 1$, Cam-
eron, April 3, 1938 (Lowery) ; December 15, 1940 (Wallace) ; November 1 and
20, 1941 (Burdick and Lowery) ; October 31, 1942 (Burdick and Tucker).
Two females were taken at White Castle on January 18, 1938 (Hewes), and
another was shot at Carville on January 15, 1941 (Lowery). No Louisiana
breeding record for the species is yet available, but in 1939 I saw a pair in
the last week of May at Baton Rouge, another near Plaquemine on May 17,
1946, and George M. Sutton and I noted a pair almost daily at Cameron be-
tween April 22 and 30, 1942. If the bird breeds in Cameron Parish, the nest-
ing race may prove to be pallescens, since a bird taken there on April 3, as
listed above, belongs to that subspecies.
Chordeiles minor minor (Forster), Eastern Nighthawk
Since the one previous record (Oberholser, op. cit., 348) of the occurrence
of this subspecies in the state now proves to be an example of C. m. howelli,
the following specimens, all taken after the publication of Oberholser's book,
constitute the only Louisiana records: 45, 19, University, October 3, 5, 12,
23, 1941 (Burdick, Howell, Ray, and Lowery) ; 4 5, 12, University, May 15,
18, 22, 30, 1942 (Burdick and Lowery) ; 1 $ , Creole, September 2, 1944 (Bur-
dick).
Chordeiles minor howelli Oberholser, Howell Nighthawk
The only state records known, all previously unpublished, are as follows:
19, Colfax, May 15, 1937 (Lowery); 2$, 12, University, May 23 and 24 and
October 3, 1941 (Ray and Lowery); 3$, University, May 22 and 25, 1942
(Burdick); 1$, Chloe, 10 miles south, April 28, 1945; 1$, Creole, 2 miles
west, April 30, 1945 (Tucker).
186 University of Kansas Publs., Mus. Nat. Hist.
Chordeiles minor aserriensis Cherrie, Cherrie Nighthawk
Three specimens, one male and two females, taken from flocks of migrating
nighthawks at University on September 29 and October 3 and 9, 1941 (Ray
and Lowery), are the only records of the occurrence of this race in the state.
Chordeiles minor sennetti Coues, Sennett Nighthawk
A female taken at University on September 29, 1941 (Burdick), and a
male shot at the same place on May 22, 1942 (Lowery), constitute the basis
for the addition of this subspecies to the Louisiana list.
Chordeiles acutipennis texensis Lawrence, Texas Nighthawk
At dusk on April 10, 1942, in company with Burdick and Ray, I en-
countered a small flock of nighthawks feeding over the marsh near the beach
a few miles from Cameron. Darkness came before more than two could be
collected, but both of these proved to be the Texas Nighthawk, a species not
heretofore recorded from Louisiana. On the following day a nighthawk was
found perched in a tree near the marsh where the birds had been seen the
previous evening. It was collected and likewise proved to be texensis.
Muscivora forficata (Gmelin), Scissor-tailed Flycatcher
The nesting of this species in northwestern Louisiana has been indicated
for some time, especially after Wallace noted it at Lucas, in Caddo Parish,
on June 16 and July 21, 1942. However, the first authentic breeding record for
the state was furnished by a freshly built nest found by Edgar W. Fullilove
and myself several miles below Bossier, on July 3, 1945. At least two pairs
were found there in a large cotton field in which an occasional pecan tree had
been left standing. The nest was in one of these trees, about 25 feet from the
ground and far out on the end of a limb. Fullilove informed me that to his
knowledge the species had nested in this field for at least ten years and that
on numerous previous occasions he had seen both nests and young.
Myiarchus cinerascens cinerascens (Lawrence), Ash-throated Flycatcher
The first record of the occurrence of this species in Louisiana is that of a
male collected by Howell at University, on March 20, 1943. On December 23,
1945, I shot a second specimen, a female, on the bank of False River opposite
New Roads. When found, both birds were actively pursuing insects and on
being skinned, both were found to be very fat.
Empidonax flaviventris (Baird and Baird), Yellow-bellied Flycatcher
Oberholser (op. cit., 394) listed this species as a rare autumn transient,
citing one definite Louisiana record for that season. On the contrary, the species
is quite regular in fall. Six specimens have been collected at University, one
each on September 12, 17, 18, and 28, 1940, October 22, 1942, and September
26, 1943 (Lowery and Wallace). Two others have been taken at Cameron,
on October 7, 1943 (Burleigh), and September 2, 1944 (Lowery). There are
numerous sight records, but since the species cannot be distinguished with
certainty in the field from extremely yellow-plumaged Acadian Flycatchers,
none of these is recorded.
Lowery — Birds of Louisiana 187
Empidonax traillii traillii (Audubon), Alder Flycatcher
This species long has been regarded as an uncommon transient in Louisiana
in both spring and fall. However, recent field work has shown the bird to
occur regularly and sometimes abundantly in autumnal migration. Forty-one
specimens have been collected at University on dates ranging from August
17 to October 5 (Lowery, et al.). Specimens taken by Burleigh at New Orleans
on September 27, 1941, and August 23, 1943, are in the Louisiana State Univer-
sity Museum.
Empidonax minimus (Baird and Baird), Least Flycatcher
Oberholser (op. tit., 397) listed this species as an uncommon transient
since he had only a few sight records at hand. Since field identification of all
eastern empidonaces in fall is open to question, our recent data, based on
collected material, are significant. Six specimens have been taken at Univer-
sity on dates ranging from September 15 to October 5, and five at Cameron
between July 25 and October 17 inclusive (Lowery, et al.). Another specimen
in the collection is that of a bird taken by Burleigh at New Orleans on October
1, 1942. There is, as yet, no unquestionable spring record for Louisiana.
Pyrocephalus rubinus mexicanus Sclater, Vermilion Flycatcher
Oberholser (op. cit., 401) listed only one record for this species, a male ob-
served by H. E. Wallace at University, on February 6, 1938, and shot the next
day by me. Since 1938, however, it has been found regularly and frequently at
numerous localities in southern Louisiana in winter. At Baton Rouge, for ex-
ample, an adult male was noted almost daily between October 19, 1941, and Jan-
uary 7, 1942, at a small pond on the University campus. An immature male
was seen there also on November 25, 1941, but not thereafter. In the following
autumn another adult male appeared at the same place on October 23, and was
observed regularly until January 15, 1943. Again, an adult male returned to the
same area on November 10, 1943, and remained until the middle of January,
1944. W. C. Abbott informs me that for several years one or two individuals
have spent the winter at a small willow-bordered pond at his home near Hope-
villa, Iberville Parish. Like the individuals noted at Baton Rouge, Abbott's
birds arrived in October or November and remained until the following January
or February. H. B. Chase, Jr., noted two individuals at City Park Lake in New
Orleans in the winter of 1944-45, and three at the same place in the winter of
1945-46. I have seen the species frequently in Cameron Parish, in south-
western Louisiana, where six specimens have been collected on dates ranging
from November 4 to January 22. Atwood (Auk, 60, 1943: 453) has also re-
corded its presence near the Laccasine Refuge in Cameron Parish. An im-
mature male was obtained at False River, near Lakeland, in Pointe Coupee
Parish, on November 8, 1942 (Burdick). E. A. Mcllhenny writes me that he
has seen the species many times at Avery Island and recently he sent me a skin
of an adult female which he collected there on October 25, 1945 (also cj.
Mcllhenny, Auk, 52, 1935: 187). From these data it is evident that the
Vermilion Flycatcher is now a regular winter visitor to southern Louisiana.
188 University of Kansas Publs., Mus. Nat. Hist.
Troglodytes troglodytes pullus (Burleigh), Southern Winter Wren
A rather large series of Winter Wrens, all taken later than the date of pub-
lication of Oberholser's book, includes three specimens of this race and provides
an addition to the state list. Two of the specimens are males collected at
Baton Rouge on November 23 and December 21, 1943 (Burleigh), and the
other is a male shot at the same place on January 23, 1944 (Burdick). Several
additional specimens in the series are noticeably darker than the average
hiemalis and may have migrated from a zone of intergradation.
Turdus migratorius nigrideus Aldrich and Nutt, Newfoundland Robin
The only two records for the occurrence of this race in Louisiana are those
of specimens taken at Baton Rouge on February 1, 1937, and February 9, 1946
(Lowery).
Hylocichla ustulata swainsoni (Tschudi), Eastern Olive-backed Thrush
Hylocichla ustulata almae Oberholser, Alma Olive-backed Thrush
Only four Louisiana specimens of the Olive-backed Thrush were available
to Oberholser in 1938. He identified two as swainsoni and two as almae. We
have since collected twenty-five specimens in the state, seven of which are
definitely almae. Of the remaining, all are clearly swainsoni with the excep-
tion of a few that appear intermediate in color. The specimens of almae
were collected at Cameron, Baton Rouge, and Baines on dates ranging from
April 26 to May 16 and from September 29 to October 6. The specimens of
swainsoni were taken at New Orleans, Port Hudson, Baton Rouge, and Baines
between April 20 and May 16 and between September 12 and October 28.
Hylocichla fuscescens salicicola Ridgway, Willow Thrush
Oberholser (op. cit., 474) recorded this race as a rare spring transient on
the basis of two records. However, eleven out of twenty-three recently taken
specimens are referable to salicicola, indicating that salicicola and fuscescens
possibly occur in approximately equal numbers, in both spring and fall. The
dates on which salicicola have been collected range from April 22 to May 16,
and from September 14 to 27. They were taken at Cameron, Port Hudson,
Baton Rouge, University, and Baines.
Anthus spinoletta pacificus Todd, Western Pipit
The only Louisiana record for this far western race is that of a female taken
by me at Jennings, on January 3, 1943. The specimen was sent to Alden H.
Miller, who compared it with material in the Museum of Vertebrate Zoology
and verified the identification. As a rule, I scrutinize closely with binoculars
all flocks of pipits, and as a result, on several occasions have detected pale
individuals that stood out from the remainder of the flock. However, the
above-mentioned specimen is the only individual so detected that I succeeded
in shooting.
Vireo solitarius alticola Brewster, Mountain Vireo
Four specimens out of a series of twenty-eight Blue-headed Vireos taken
in Louisiana since 1938 are referable to this race. It has not been recorded
previously from the state. The specimens consist of a male and a female col-
Lower y — Birds of Louisiana 189
lected at Bogalusa on February 9, 1939, a male taken at Tunica on March 30,
1939, and a female at Erwinville on March 11, 1941 (Lowery).
Helmitheros vermivorus (Gmelin), Worm-eating Warbler
Although there are no published nesting records of this species in Louisiana,
it is now known to be a common summer resident in the beech-magnolia
forests of the Bayou Sara-Tunica Hills section north of St. Francisville. Jas.
Hy. Bruns has supplied me with copious data on the birds seen in the nesting
season at Baines, and the two of us have spent a great deal of time searching
for a nest, without success. However, Bruns obtained a juvenile female, just
out of a nest, on June 28, 1942.
Seiurus aurocapillus furvior Batchelder, Newfoundland Oven-bird
Seiurus aurocapillus cinereus A. H. Miller, Gray Oven-bird
Four specimens in our series of Oven-birds are identifiable without question
as examples of furvior. Two were collected by me at University on Septem-
ber 15 and 25, 1940, and Tucker shot one there on September 27, 1942, and
another at Cameron on April 29, 1945. There are also two specimens in the
series referable to cinereus, as well as several that are intermediate between
cinereus and S. a. aurocapillus. Burdick shot one of the typical examples of cin-
ereus at University on September 24, 1942, and I shot the other at the same
place on May 16, 1945.
Seiurus noveboracensis noveboracensis (Gmelin), Northern Water-thrush
Seiurus noveboracensis limnaeus McCabe and Miller, British Columbia
Water-thrush
A. H. Miller has recently examined our large series of migrant Water-
thrushes and identified three as good examples of limnaeus, and six as nove-
boracensis, neither one of which has been recorded previously from the state.
The specimens of limnaeus were taken at or near University on October 2,
1942 (Howell), October 12, 1943, and May 11, 1945 (Burleigh). The specimens
of noveboracensis were collected at University on September 14, 1941 (Low-
ery) ; at Baines on September 4, 1943, August 20, 1944, and May 6, 1945
(Bruns) ; at New Orleans on October 20, 1941 (Burleigh) ; and at Cameron
on April 26, 1942 (Lowery).
Geothlypis trichas occidentalis Brewster, Western Yellow-throat
I have found it impracticable to determine subspecifically every specimen
in our series of 104 Yellow-throats from Louisiana. However, two female
specimens taken by me, one at Cameron on December 4, 1938, and the other
on False River at Lakeland on February 11, 1941, are without doubt repre-
sentatives of the race now known as occidentalis, a subspecies not previously
recorded from this state. Several additional specimens in the series are prob-
ably also of that race, but I am deferring, for the time, recording them as Buch.
Icteria virens virens (Linnaeus), Yellow-breasted Chat
The only winter record for Louisiana is that of a female taken by me at
Hackberry on January 24, 1941.
190 University of Kansas Publs., Mus. Nat. Hist.
Wilsonia pusilla pusilla (Wilson), Wilson Warbler
The only winter record for the state is that of a female shot by T. D. Bur-
leigh on December 20, 1944, in a thicket along the Mississippi River at Uni-
versity. He first found the bird at this place in November, and he saw it
several times in December before he succeeded in obtaining it. Since Ober-
holser cited so few Louisiana records, it might be well to mention in this con-
nection that the species is after all a fairly common fall migrant in southern
Louisiana. At Baton Rouge it occurs regularly between September 11 and
October 24, and at Cameron it has been noted between October 17 and No-
vember 21. There are still no spring records for southern Louisiana.
Sturnella neglecta Audubon, Western Meadowlark
In 1938 Oberholser cited only two Louisiana records, both from the north-
western part of the state. However, recently the species has been found in
the south-central region. Two were collected at Churchill on February 11,
1941 (Lowery and Wallace), and another was shot at University on December
9, 1942 (Burdick). There are in addition several sight records, all of birds in
song.
Cassidix mexicanus prosopidicola Lowery, Mesquite Great-tailed Grackle
I am indebted to E. A. Mcllhenny for material that now permits the defi-
nite recording of this subspecies from Louisiana. On occasions during the
winters of 1938, 1939, and 1940, Mcllhenny sent me specimens of grackles in
the flesh which he had removed from his bird-banding traps at Avery Island.
Selection was based primarily on eye-color; individuals with clear yellow
irises proved invariably to be examples of prosopidicola, whereas those with
brown or yellow-brown irises were always major. The final basis for sub-
specific identification was, however, size and plumage color. The series pro-
vided by Mcllhenny consists of six females taken on November 24 and De-
cember 20, 1938, December 18, 1939, January 22 and March 5, 1940. Since the
range in Texas of typical prosopidicola extends eastward to within thirty miles
of the Louisiana line, it is not surprising that occasional individuals or flocks
wander into Louisiana in winter.
Passerculus sandwichensis mediogriseus Aldrich, Southeastern Savannah
Sparrow
Passerculus sandwichensis labradorius Howe, Labrador Savannah Sparrow
Passerculus sandwichensis nevadensis Grinnell, Nevada Savannah Sparrow
Our series of 107 Savannah Sparrows, collected in Louisiana almost en-
tirely since the publication of Oberholser's book, includes representatives of
five geographical races, as follows: 37 savanna, 24 oblitus, 12 mediogriseus, 8
labradorius, and 7 nevadensis. The remaining 19 specimens show various
combinations of characters and appear to be intergrades, and so have not
been assigned definitely to any one race. I am indebted to James L. Peters
for the identification of most of our specimens. Since mediogriseus and la-
bradorius have not been reported previously from Louisiana, and since there
is only one Louisiana record of nevadensis (Miles, Auk, 60, 1943: 606-607),
actual dates and localities of occurrence for these races are listed here. P. s.
Lowery — Birds of Louisiana 191
medio griseus (specimens by Burdick, Howell, Lowery, Ray, Tucker, and Wal-
lace)— University, January 31, 1939; February 11 and 29, April 29, November
28, and December 16, 1940; December 6 and 7, 1941; October 10 and 25, 1942;
April 14, 1943. Erwinville, March 11, 1941. P. s. labradorius (specimens by
Burleigh, Lowery, Mcllhenny, Ray and Wallace) — University, February 15 and
November 8, 1940; January 1, 1941; December 11, 1943. 2 mi. NE Baton
Rouge, January 1, 1941. Burtville, December 8, 1939. Avery Island, May 3,
1939. Lake Charles, November 20, 1940. P. s. nevadensis (specimens by Bur-
dick, Lowery, and Wallace) — Iowa Station, January 23 and 24, 1940. Univer-
sity, February 10 and March 10, 1940. University, December 7, 1941, and No-
vember 15, 1942. Cameron, December 6, 1942. There are at present no bona
fide records of P. s. anthinus in Louisiana, since the one recorded example of
that race (Oberholser, op. cit., 647) appears, on reexamination, to be referable
to savanna (fide J. L. Peters).
Ammodramus savannarum pratensis Vieillot, Eastern Grasshopper Sparrow
Eight specimens of the Grasshopper Sparrow taken recently in Louisiana
are without exception referable to pratensis. Our one remaining specimen, a
male collected at Pride on December 19, 1937, is an example of perpallidus as
recorded by Oberholser (op. cit., 648). Although the present series is inade-
quate for determining the prevailing form in the state in the winter, it would
appear that pratensis is more common, rather than perpallidus as indicated by
Oberholser.
Chondestes grammacus strigatus Swainson, Western Lark Sparrow
Oberholser cited only one Louisiana record for this race. The following
additional records are now available : a specimen was taken by Howell at
Cameron on October 31, 1942, and one was obtained by me at University on
April 13, 1945. The species is a transient in both localities. A supplementary
winter record for the Lark Sparrow in Louisiana is that of an individual seen
at Port Hudson on December 23, 1945, by Howell and Newman. The bird
was shot, but unfortunately, it was not retrieved.
Junco hyemalis cismontanus Dwight, Cassiar Junco
The only specimen in our series of Slate-colored Juncos that is a clear-cut
example of this race is a male taken by Ambrose Daigre at Catahoula Lake
on November 29, 1939. A. H. Miller has confirmed the identification.
Calcarius lapponicus alascensis Ridgway, Alaska Longspur
Oberholser listed this species as a casual winter visitor in northern Louisi-
ana, which was possibly no more than was indicated by records then available
to him. Since 1938, however, the species has been observed in large flocks at
various localities in the southern part of the state, notably in January, 1941,
when the whole state was blanketed with snow. Nevertheless, snow is ap-
parently not prerequisite to the appearance of the species this far south, for
on January 1 and 3, 1943, a flock of approximately a thousand individuals
was seen a few miles north of Jennings. Again, on February 14, 1943, about
half of what may have been the original flock was observed there. In neither
instance was there snow anywhere in Louisiana. Of the thirty specimens in
192 University of Kansas Publs., Mus. Nat. Hist.
the Louisiana State University Collection, eleven have been identified by
Alexander Wetmore as somewhat intermediate between alascensis and lapponi-
cus, but closer to the former. Only lapponicus has been previously recorded
from Louisiana. The specimens of alascensis were taken at Baton Rouge on
January 25 and 28, 1940; Cornor, January 27, 1940; Lottie, January 27, 1940;
and 10 miles north of Jennings, January 1 and February 14, 1943 (Burdick,
Campbell, Hewes, Lowery, and Wallace).
Transmitted February 1, 1947.
21-6959
A Check-List of the Birds of Idaho
BY
M. DALE ARVEY
LIBRARY
1AR
-8 19.50
;
HARVARD
UNIVERSITY 0F_KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
Volume 1, No. 10, pp. 193-216
November 29, 1947
University of Kansas
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, H. H. Lane, Edward H. Taylor
Volume 1, No. 10, pp. 193-216
Published November 29, 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
I 947
•1\ -69 111
<P. 2001 J
LIBRARY
HAR -8 I960
A Check-list of the Birds of Idaho
By
M. DALE ARVEY
There is comparatively little literature dealing with the avifauna
of Idaho, mostly because relatively few persons have done field work
in the state. In the ornithological literature, there is nothing even
comparable to a "state list," so that when birds supposedly unre-
ported previously from Idaho are found, it is difficult to know
whether or not they should be recorded as "new" to the state. The
present paper has been prepared in the hope that it will stimulate
additions to, and corrections of, the list. It is, admittedly, a be-
ginning.
Material for the present article was obtained from personal col-
lecting in the five years and ten months in which I resided in the state
(October, 1938-September, 1944). Also, the published reports that
could be found have been drawn upon ; these publications are listed
in the appended bibliography. Taxonomic problems, of which many
are unsolved, are not here considered, since this is merely a list in-
dicating whether or not the species or subspecies, as now understood,
is known to be present, whether it is common, and where it might be
found.
The nomenclature is that of the Fourth Edition of the American
Ornithologists' Union Check-list and its supplements, except where
a revision has been made that is seemingly valid but which has not
yet been acted upon by the A. 0. U. Committee. For each species
or subspecies the objective is to give at least one reference to occur-
rence, as to date and place, as accurately as possible.
Reference is made to southern, central, and northern Idaho. These
references denote the Snake River Plains, characterized by sage-
brush desert; the wooded regions immediately to the north of this
and in the foothills, extending to Idaho County in the west; and the
so-called Panhandle, respectively. In all, 292 kinds of birds are
recorded in the following list.
LIST OF SPECIES
Gavia immer elasson Bishop. Lesser Loon. Uncommon resident in the lakes
of northern Idaho, and generally distributed. Merrill (1897:350) states that
the species is common and resident at Fort Sherman.
Gavia stellata (Pontopiddan). Red-throated Loon. Davis (1935b :234)
records specimens taken in migration in Minidoka County at the Minidoka
(195)
196 University of Kansas Ptjbls., Mus. Nat. Hist.
Irrigation Project, and Rust (1915:121) states that this species is rare in
Kootenai County.
Colymbus grisegena holbollii (Reinhardt). Holboell Grebe. Merrill (1897:
349) records this species as common in migration at Fort Sherman.
Colymbus auritus Linnaeus. Horned Grebe. Uncommon resident. Davis
(1935b :234) records the bird as a summer visitant at the Minidoka Project.
Colymbus nigricollis californicas (Heermann). Eared Grebe. Fairly com-
mon resident along rivers and in lakes. Rust (1915:121) records one specimen
taken on Lake Coeur d'Alene in October, 1912.
Aechmophorus occidentalis (Lawrence). Western Grebe. Uncommon resi-
dent. Merrill (1897:349) records one specimen from Fort Sherman.
Podilymbus podiceps podiceps (Linnaeus). Pied-billed Grebe. Common
resident. Merrill (1S97:350) states that it is common at Fort Sherman in the
spring and autumn.
Pelecanus erythrorhynchos Gmelin. White Pelican. Resident along the
Snake River; large nesting colonies are to be found in Bear Lake County.
See Davis (1935b :234) for nesting dates.
Phalacrocorax auritus albociliatus Ridgway. Farallon Cormorant. Davis
( 1935b :234) records this bird in the Minidoka Project as a regular migrant and
gives dates of occurrence. The resident population at the Bear Lake Refuge
has been reported as subspecies auritus by Behle (1944:68), but probably is
albociliatus.
Ardea herodias treganzai Court. Treganza Great Blue Heron. Common
resident in suitable localities. (Dale Arvey 1505, 7 mi. NE Moscow, Latah
County, Idaho, February 19, 1940.)
Leucophoyx thula brewsteri (Thayer and Bangs). Brewster Egret. Davis
(1935b :234) records one specimen from the Minidoka Project, taken on Sep-
tember 16, 1919, and Hayward (1934:39) reports the species as breeding at
Bear Lake Valley in Bear Lake County.
Nycticorax nycticorax hoactli (Gmelin). Black-crowned Night Heron.
Common locally. Hayward (1934:39) reports the bird as resident in Bear
Lake Valley.
Botaurus lentiginosus (Montagu). American Bittern. Fairly common res-
ident in suitable localities. Merrill (1897:351) records the American Bittern
as rather common at Fort Sherman.
Plegadis mexicana (Gmelin). White-faced Glossy Ibis. Vagrant. Re-
Corded as common at the Minidoka Project by Kenagy (1914:122).
Cygnus columbianus (Ord). Whistling Swan. Resident in the winter in
the larger lakes, and transient along the Snake River. (D. A. 1783, 1 mi. S
Hagerman, Gooding County, February 1, 1940.)
Cygnus buccinator Richardson. Trumpeter Swan. Merriam (1891:91)
states that Bendire found this swan breeding on Henry Lake in 1877. and
that two were collected in August of that year. Rust (1915:123) records the
species as a rare fall migrant on Lake Coeur dAlene. There are no recent
records.
Branta canadensis (Linnaeus). Canada Goose. Fairly common resident.
See Aldrich (1946b) for records of each subspecies.
a. moffitti Aldrich. Great Basin Canada Goose. This is the resident race.
b. occidentalis (Baird). White-cheeked Goose. Migrant.
Arvey — Birds of Idaho 197
c. leucopareia (Brandt). Lesser Canada Goose. Migrant.
Branta hutchinsii hutchinsii (Richardson). Hutchins Cackling Goose. Mi-
grant. See Aldrich (1946b) for the srtatus of this goose.
Branta bernicla nigricans (Lawrence). Black Brant. Davis ( 1935b :234)
records this species as a regular migrant in Minidoka County, and indicates
that some remain all winter.
Anser albijrons albijrons (Scopoli). White-fronted Goose. Uncommon
migrant. Jones (1943:120) records a specimen from "about 10 mi. north
Pocatello, Bingham County."
Chen hyperborea hyperborea (Pallas). Lesser Snow Goose. Fairly com-
mon transient along the Snake River. Two specimens are in the State Game
Department's mounted collection from the Snake River, probably from near
Payette, Payette County.
Chen rossi (Cassin). Ross Goose. Transient along the Snake River. The
Game Department collection has two mounted skins from "along the Snake
River."
Anas platyryhnchos platyryhnchos Linnaeus. Mallard. Very common resi-
dent. (D. A. 1753, Boise River, 1 mi. S Middleton, Canyon County, November
24, 1940.)
Anas acuta tzitzihoa (Vieillot). American Pintail. Resident and common
during migration. (D. A. 1752, Snake River, 1 mi. S Hammett, Elmore County,
November 16, 1940.)
Ayias carolinensis Gmelin. Green-winged Teal. Common resident. (D. A.
1261, Thorn Creek, 7 mi. S Moscow, Latah County, October 30, 1938.)
Anas discors Linnaeus. Blue-winged Teal. Rare resident. Merriam (1891:
90) records two shot on Saw Tooth Lake (=Alturas Lake, Blaine County),
about October 1.
Anas cyanoptera Vieillot. Cinnamon Teal. Uncommon resident. I ob-
served a female with four young in Bellevue, Blaine County, in July, 1942, and
Merrill (1897:350) records a female with young on June 11 at Fort Sherman.
Anas strepera Linnaeus. Gadwall. Resident locally; fairly common in
migration. (D. A. 1310, Havenor's, 7 mi. NW Pocatello, Power County,
January 2, 1939.)
Mareca americana (Gmelin). Baldpate. Common during migration, and
resident along the Snake River. (D. A. 1747, 1 mi. W Bowman Ranch on Boise
River, Canyon County, October 26, 1940.)
Spatula clypeata (Linnaeus). Shoveller. Common in migration, and breeds
locally. (D. A. 1492, Wallace, Shoshone County, October 22, 1939.)
Aix sponsa (Linnaeus). Wood Duck. Fairly common in migration, and resi-
dent locally. Merrill (1897:350) records it as a summer resident at Fort
Sherman.
Ay thy a americana (Eyton). Redhead. Fairly common migrant. Recorded
by Merrill (1897:350) at Fort Sherman.
Aythya collaris (Donovan). Ring-necked Duck. Uncommon transient.
Merrill (1897:350) records it at Fort Sherman.
Aythya valisineria (Wilson). Canvas-back. Fairly common in migration,
and recorded by Low and Nelson (1945:131) as breeding in Bonneville and
Caribou counties.
198 University of Kansas Publs., Mus. Nat. Hist.
Ay thy a marila (Linnaeus). Greater Scaup Duck. Fairly common migrant.
Davis (1935b :236) records one bird from the Minidoka Project taken on March
28, 1920.
Aythya affinis (Eyton). Lesser Scaup Duck. Common during migration.
Davis (1935b :235) lists this bird as a regular winter visitant in Minidoka
County from October 30 to May 31.
Glaucionetta clangula americana (Bonaparte). American Golden-eye. Com-
mon resident. (D. A. 1476, Bellevue, Blaine County, June 28, 1939.)
Glaucionetta islandica (Gmelin). Barrow Golden-eye. Uncommon tran-
sient. Davis (1935b :234) records one specimen taken at the Minidoka Project.
Glaucionetta albeola (Linnaeus). Buffle-head. Common migrant. (D. A.
1852, Snake River, 1 mi. S Hammett, Elmore County, November 15, 1941.)
Histrionicus histrionicus pacificus Brooks. Western Harlequin Duck. Un-
common. Rust (1915:122) records one specimen taken on the marshes of the
St. Joseph River in Kootenai County, and Merrill (1897:350) states that it is
occasionally taken on the St. Joseph and Coeur d'Alene rivers.
Melanitta jusca subsp. ?. White-winged Scoter. Rust (1915:122) re-
cords this bird as common on Lake Coeur d'Alene in the winter of 1913.
Melanitta perspicillata (Linnaeus). Surf Scoter. Rust (1915:122) states
that this is a rare fall migrant in Kootenai County.
Oxyura jamaicensis rubida (Wilson). Ruddy Duck. Common migrant
on the Snake River. Merrill (1897:350) records this duck as "not uncommon
in the spring and autumn" at Fort Sherman.
Lophodytes cucullatus (Linnaeus). Hooded Merganser. Common resident
in suitable localities. (D. A. 1389, Lewiston, Nezperce County, April 2, 1939.)
Mergus merganser americanus Cassin. American Merganser. Common
resident. Merrill (1897:350) states that the bird is common in fall and winter
at Fort Sherman.
Mergus serrator Linnaeus. Red-breasted Merganser. Uncommon. Merrill
(1897:350) records one specimen taken "near Fort Sherman."
Cathartes aura teter Friedmann. Western Turkey Vulture. Common resi-
dent in southern Idaho, and transient elsewhere. Merrill (1897:352) records it
as a summer resident at Fort Sherman.
Accipiter gentilis striatulus (Ridgway). Western Goshawk. Fairly com-
mon migrant, and possibly resident. Hand (1933b :36) reports it as resident in
northern Idaho. (D. A. 1317, 1318, Nezperce, Lewis County, January 9 and
12, 1939.)
Accipiter striatus velox (Wilson). Sharp-shinned Hawk. Common resident.
(D. A. 1296, 4y2 mi. NE Genessee, Latah County, November 27, 1938.)
Accipiter cooperii (Bonaparte). Cooper Hawk. Common resident in the
forests. (D. A. 1450, Sandpoint, Bonner County, May 24, 1939.)
Buteo jamaicensis calurus Cassin. Western Red-tailed Hawk. Common resi-
dent. (D. A. 1352, Moscow, Latah County, March 18, 1939.)
Buteo platypterus platypterus (Vieillot). Broad-winged Hawk. Davis
(1936:86) records one specimen of this hawk taken on May 23, 1935, at Castle
Creek, 8 mi. S Oreana, Owyhee County.
Buteo swainsoni Bonaparte. Swainson Hawk. Common resident. (D. A.
1451, Moscow, Latah County, May 21, 1939.)
Arvey — Birds of Idaho 199
Buteo lagopus s. johannis (Cmelin). American Rough-legged Hawk. Com-
mon migrant and possibly resident. (D. A. 1301, 11 mi. SE Gcncssee, Nezperce
County, November 27, 1938.)
Buteo regalis (Gray). Ferruginous Rough-leg. Uncommon migrant. (D. A.
1326. 4 mi. N Minidoka Power Plant, Minidoka County. January 27, 1939.)
Aquila chrysaetos canadensis (Linnaeus). Golden Eagle. Uncommon resi-
dent. Merrill (1897:353) stated that the species occurred "sparingly" at Fort
Sherman.
Haliaeetus leucocephalus washingtoniensis (Audubon). Northern Bald
Eagle. Uncommon resident in northern Idaho. Merrill (1897:353) stated that
a few pairs bred about Lake Coeur dAlene.
Cirais cyaneus hudsonius (Linnaeus). Marsh Hawk. Very common resi-
dent. (D. A. 1371, Havenor's, 7 mi. NW Pocatello, Power County, April 1,
1939.)
Pandian haliaetus earolinensis (Gmelin). Osprey. Uncommon resident.
Merrill (1897:353) reported the bird as frequent in the summer at Fort
Sherman.
Falco mexicanus Schlegel. Prairie Falcon. Fairly common resident. (D. A.
1319, American Falls, Bingham County, January 16, 1939.)
Falco percgrinus anatum Bonaparte. Duck Hawk. Uncommon resident.
Bond (1946:104) lists this bird as a rare breeder in Idaho.
Falco columbarius bendirei Swann. Western Pigeon Hawk. Rust (1915:124)
records one specimen from Coeur d'Alene as subspecies columbarius ; although
the skin has not been checked by me, it would seem to be more likely of
subspecies bendirei, corresponding to others taken in northern Idaho.
Falco sparverius sparverius Linnaeus. Eastern Sparrow Hawk. Common
resident. (D. A. 1267, Little Bear Ridge, 5 mi. SW Troy, Latah County,
November 2, 1939.)
Dendragapus obscurus (Say). Blue Grouse. Common resident.
a. obscurus (Say). Dusky Grouse. Specimens from southeastern Idaho
are referable to this race.
b. richardsonii (Douglas). Richardson Grouse. This is the resident
race of southwestern Idaho north to Idaho County, where intergrada-
tion occurs with the next form. (D. A. 1431, 1432, 10 mi. SW Rig-
gins, Idaho County, May 14, 1939.)
c. pallidus Swarth. Oregon Dusky Grouse. Birds in the northern por-
tion of the state are of this race.
Canachites jranklinii (Douglas). Franklin Grouse. Uncommon resident.
I have observed the birds in the Selway National Forest, in Idaho County,
and specimens have been taken in Bonner County. (D. A. 1336, 1337, 6 mi.
S Coolin, Bonner County, February 19, 1939.)
Bonasa umbellus (Linnaeus). Ruffed Grouse. Common resident. See
Aldrich and Friedman (1943) for ranges of the following races.
a. phaia Aldrich and Friedmann. Idaho Ruffed Grouse. This is the
race resident in southwestern Idaho, and it intergrades with the two
following forms.
b. umbelloides (Douglas). Gray Ruffed Grouse. Resident in northern
Idaho.
200 University of Kansas Publs., Mus. Nat. Hist.
c. incanics Aldrich and Friedmann. Hoary Ruffed Grouse. Resident in
southeastern Idaho.
Lagopus leucurus altipetens Osgood. Southern White-tailed Ptarmigan.
Several specimens of this bird are mounted in a collection in Idaho City,
having been collected "in the vicinity."
Pcdioecetes phasianellus columbianus (Ord). Columbian Sharp-tailed
Grouse. One specimen was sent me from Bonner County, where the species
was said to be fairly abundant. (D. A. 1513. 15 mi. N Priest River, Bonner
County, April 1, 1940.)
Centrocercus urophasianus (Bonaparte). Sage Grouse. Common locally.
Previously numerous, and now recovering from a severe decline in numbers.
Merriam (1891:93) speaks of using these birds for fresh meat during much
of his trip.
Perdix perdix perdix (Linnaeus). European Partridge. Common since its
introduction.
Colinus virginianus texanus (Lawrence). Texas Bob-white. Common res-
ident in southern Idaho. Merriam (1891:92) states that the birds were first
introduced at Boise, Ada County.
Lophortyx californica brunnescens Ridgway. California Quail. Intro-
duced into southern Idaho; not numerous but establishing itself in the foot-
hills.
Oreortyx picta picta (Douglas). Plumed Quail. Common resident. Wy-
man (1912c :538) states that this species was not present in Idaho prior to
about 1900, having at that time extended its range from Oregon.
Phasianus colchicus Linnaeus. Ring-necked Pheasant. Common resident
since its introduction; there is considerable admixture of races in the stock.
Grus canadensis tabida (Peters). Sandhill Crane. Uncommon resident.
Merriam (1891:91) reports the bird breeding near Fort Lapwai, Nezperce
County, in June 1871, and Davis (1935b :234) states that it is a regular mi-
grant at the Minidoka Project.
Rallus limicola limicola Vieillot. Virginia Rail. Davis (1923) states that
this rail is uncommon at the Minidoka Project, but that it was abundant in
earlier years.
Porzana Carolina (Linnaeus). Sora. Uncommon resident. Merriam (1891:
91) recorded this species from Big Lost River, "about 8 mi. above Arco,"
Butte County, on July 26.
Fulica americana Gmelin. American Coot. Common resident. (D. A.
1745, Notus, Canyon County, October 20, 1940.)
Charadrius voriferus vocijerus Linnaeus. Killdeer. Common resident
in the Transition Life-zone. Rust (1915:123) records the earliest arrival date
for the bird in Kootenai County as March 9, 1913, and says that it leaves by
September 1.
Pluvialis dominica julva (Gmelin). Pacific Golden Plover. Sloanaker
(1925:73) records one specimen of this bird, shot from a flock of four near
Coeur d'Alene on Lake Chactolet on October 1, 1923.
Squatarola squatarola (Linnaeus). Black-bellied Plover. Rust (1915:123)
records one specimen of this bird taken on the St. Joseph marshes, Kootenai
County.
Arvey — Birds of Idaho 201
Capella gallinago delicata (Ord). Wilson Snipe. Fairly common resident.
(D. A. 1739, Boise River, 3 mi. W Boise, Ada County, October 17, 1940.)
Numenius americanus Bechstein. Long-billed Curlew. Uncommon resi-
dent. See Oberholser (1918) for ranges of the following subspecies.
a. americanus Bechstein. Long-billed Curlew. Resident in southern
Idaho.
b. parvus Bishop. Northern Curlew. The resident population in north-
ern Idaho is referable to this subspecies.
Actitis macularia (Linnaeus). Spotted Sandpiper. Common resident in the
Canadian Life-zone. (D. A. 1807, junction of Simmon's Cr. and Boise River,
Boise County, July 5, 1941.)
Tringa solitaria cinnamomea (Brewster). Western Solitary Sandpiper.
Davis (1935b :236) took one specimen on April 9, 1920 at the Minidoka Proj-
ect, and records the bird as erratic in occurrence.
Catoptrophorus semipalmatus inornatus (Brewster). Western Willet. Da-
vis (1935b :235) records this bird as a summer visitant at the Minidoka Proj-
ect, and gives dates of its occurrence there.
Totanus melanoleucus (Gmelin). Greater Yellow-legs. Davis (1935b :234)
records this bird at the Minidoka Project in migration.
Totanus flavipes (Gmelin). Lesser Yellow-legs. Fairly common in migra-
tion. (D. A. 1742, Notus, Canyon County, October 20, 1940.)
Erolia melanotos (Vieillot). Pectoral Sandpiper. Merrill (1897:351) re-
cords this bird as common in 1896 from August to October at Fort Sherman,
and a number of specimens were taken.
Erolia minutilla (Vieillot). Least Sandpiper. Fairly common migrant.
Davis (1935b :234) gives dates of migration of this bird at the Minidoka Proj-
ect.
Limnodromus griseus scolopaceus (Say). Long-billed Dowitcher. Merrill
(1897:351) collected five specimens on September 12 on the St. Joseph
marshes.
Micropalarna himantopus (Bonaparte). Stilt Sandpiper. Davis (1935b:
234) collected one bird at the Minidoka Project on May 13, 1919, and stated
that the species was erratic in occurrence.
Ereunetes mauri Cabanis. Western Sandpiper. Rust (1917:32) recorded
this bird on August 27 near Spencer, Fremont County, and also at Henry Lake.
Limosa fedoa (Linnaeus). Marbled Godwit. Davis (1935b :236) records
one specimen taken on August 1, 1920, at the Minidoka Project.
Limosa haemastica (Linnaeus). Hudsonian Godwit. Davis (1935b :236)
records one bird taken at the Minidoka Project on July 7, 1919.
Crocethia alba (Pallas). Sanderling. Davis (1935b :236) records this bird
from the Minidoka Project in migration, and he took one specimen on May
19, 1921.
Recurvirostra americana Gmelin. Avocet. Uncommon resident in southern
Idaho. (D. A. 1631, Snake River at Hagerman, Gooding County, June 16,
1940.)
Himantopus mexicanus (Miiller). Black-necked Stilt. Davis (1935b :235)
records this bird from Minidoka as a summer visitant, and gives dates of its
occurrence.
202 University of Kansas Publs., Mus. Nat. Hist.
Phalaropus fulicarius (Linnaeus). Red Phalarope. Hand (1935:180) re-
ports one bird of this species in October on the St. Joseph River at St. Maries,
Benewah County.
Steganopus tricolor Vieillot. Wilson Phalarope. Uncommon. Davis (1935b:
236) took one specimen at the Minidoka Project on May 13, 1919.
Lobipes lobatus (Linnaeus). Northern Phalarope. Uncommon resident.
Davis (1935b :236) reports the species as erratic at the Minidoka Project,
where he took one specimen on May 13, 1919.
Stercorarius pomarinus (Temminck). Pomarine Jaeger. Davis (1935b :236)
took one bird "on the Snake River," on September 4, 1919.
Larus argentatus thayeri Brooks. Thayer Gull. Merrill (1897:350) records
several birds of this species taken in the fall and winter on Lake Coeur
d'Alene.
Larus californicus Lawrence. California Gull. Common in the winter, and
possibly breeds along the Snake River. Davis (1935b :235) records this bird
as a common summer visitant at the Minidoka Project.
Larus delawarensis (3rd. Ring-billed Gull. Uncommon straggler. Merrill
(1897:350) records it in the winter at Fort Sherman.
Larus pipixcan Wagler. Franklin Gull. Late winter and spring straggler.
See Slipp (1942).
Larus Philadelphia (Ord). Bonaparte Gull. This gull is recorded by Mer-
rill (1897:350) as taken at Fort Sherman in November.
Sterna jorsteri Nuttall. Forster Tern. Davis (1935b :235) lists this bird as
a summer visitant in Minidoka County, and gives dates of its occurrence
there.
Sterna hirundo hirundo Linnaeus. Common Tern. Rust (1915:121) states
that this tern is rare in Kootenai County.
Hydroprogne caspia (Pallas). Caspian Tern. Common during migration.
Davis (1935b :234) records the species as common in migration at the Mini-
doka Project, and gives dates of its occurrence.
Chlidonias nigra surinamensis (Gmelin). Black Tern. Fairly common on
lakes; evidently resident. Rust (1915:121) records this bird as common in
June, 1914, on the St. Joseph Marshes.
Columba jasciata jasciata Say. Band-tailed Pigeon. Rare at present.
Merrill (1897:349) states that Cooper listed this bird in what is now Idaho.
Zenaidura macroura marginella (Woodhouse). Western Mourning Dove.
Common summer resident, frequently remaining in winter. Rust (1915:123)
lists the bird as a fairly common summer resident in Kootenai County.
Ectopistes migratorius (Linnaeus). Passenger Pigeon. Extinct. Merrill
(1897:349) states that Cooper listed this species from Montana and from what
is now Idaho.
Coccyzus americanus occidentalis Ridgway. California Cuckoo. This bird
was reported by Davis (1935b :236), as taken May 16, 1918 at the Minidoka
Project, and he says that nests have been taken near Rupert by Kenagy.
Coccyzus erythroplhalmus (Wilson). Black-billed Cuckoo. One breeding
bird of this species was reported by Arvey (1941 :291), taken at Slide Gulch on
the Boise River, Boise County, on July 10, 1941. Since this time I have ob-
served the bird twice in Boise, Ada County, in the summer.
Arvey — Birds of Idaho 203
Tyto alba pratincola (Bonaparte). Barn Owl. Uncommon resident. One
specimen in the University of Idaho collection of mounted birds was taken
near Moscow, Latah County.
Otus asio (Linnaeus). Screech Owl. Common resident.
a. macjarlanei (Brewster). MacFarlane Screech Owl. Resident in south-
ern Idaho. (D. A. 1861, Boise, Ada County, April 11, 1942.)
b. brcwsteri Ridgway. Brewster Screech Owl. Resident in northern
Idaho. (D. A. 1312, Lapwai, Nezperce County, December 25, 1938.)
Otus flammeolus flammeolus (Kaup). Flammulated Screech Owl. Rare
resident. Specimens have been taken in two localities. Merriam (1891 :96) took
one. specimen on the west side of Big Wood River, "only a few miles north of
Ketchum, September 22," 1890. The record from Blaine County and the one
of Rust (1915:125), near Fernan Lake, September 28, 1914, are the only two
positive records of this species to my knowledge.
Bubo virginianus (Gmelin). Great Horned Owl. Common resident. See
A.O.U. Check-list (1931).
a. wapacuthu (Gmelin). Arctic Horned Owl. Migrant.
b. occidentalis Stone. Montana Horned Owl. Resident in central and
south-eastern Idaho.
c. lagophonus (Oberholser). Northwestern Horned Owl. Resident in
western and northern Idaho. (D. A. 1486, 10 mi. SW Riggins, Idaho
County, September 15, 1939.)
Nyctea scandiaca (Linnaeus). Snowy Owl. Casual migrant. Merrill (1897:
352) stated that there was an invasion of owls of this species in the winter of
1896-'97, and many were observed during that time at Fort Sherman.
Surnia idula caparoch (Miiller). American Hawk Owl. Uncommon. Hand
(1933a :32) reports one specimen of this owl taken at Stanley Butte, 10 mi. S
Lochsa River, Idaho County, on November 3, 1925, and mentions one other
observed in the summer. He suggests that the bird breeds in northern Idaho.
Glaucidium gnoma calijornicum Sclater. California Pygmy Owl. Fairly
common resident in the Canadian Life-zone. Specimens seem referable to sub-
species pinicola, recently synonymized by the A. 0. U. Committee. (D. A. 1311,
Priest River, Bonner County, January 3, 1939.)
Speotyto cunicularia hypugaea (Bonaparte). Western Burrowing Owl.
Fairly common local resident. (D. A. 1388, 10 mi. W Boise, Ada County,
April 2, 1939.)
Sirix nebulosa nebulosa Forster. Great Gray Owl. Vagrant. A specimen,
D. A. 1303, taken on December 8, 1938, was sent me from 9 mi. NE Grangeville,
Idaho County, December 8, 1938.
Asio otus wilsonianus (Lesson). Long-eared Owl. Fairly common resident.
(D. A. 1532, 5 mi. SW Moscow, Latah County, April 29, 1940.)
Asio flammeus flammeus (Pontoppidan). Short-eared Owl. Very common
resident in the Transition Life-zone. (D. A. 1346, 2 mi. S Moscow, Latah
County, March 7, 1939.)
Aegolius funereus richardsoni (Bonaparte). Richardson Owl. Rust
(1915:125) records this bird as a rare winter visitor in Kootenai County, and
Merrill (1897:353) lists two specimens taken "early in the spring of 1894 . . .
about seven miles from the fort."
204 University of Kansas Publs., Mus. Nat. Hist.
Aegolius acadicus acadicus (Gmelin). Saw-whet Owl. Rare. Davis
(1935b :235) says that this is a regular winter visitor at the Minidoka Project,
and Merrill (1897:353) lists one specimen taken at Fort Sherman, on January 19.
Phalaenoptilus nuttallii nuttallii (Audubon). Nuttall Poorwill. Uncom-
mon resident. Merriam (1891:98) records this species from "the lava beds
west of Blackfoot" on July 17, 1872.
Chordciles minor hesperis Grinnell. Pacific Nighthawk. Common resident
in the Transition Life-zone. (D. A. 1468, 2 mi. S Hailey, on Wood River,
Blaine County, June 25, 1939.)
Chaetura vauxi vauxi (Townsend). Vaux Swift. Merrill (1897:354) re-
ports this bird as resident at Fort Sherman, as does Burleigh (1923:658) at
Clark's Fork, Bonner County.
Aeronautes saxatalis saxatalis (Woodhouse). White-throated Swift. Fairly
common resident in suitable localities. The Museum of Vertebrate Zoology
has one specimen of this bird taken on Salmon Creek, 8 mi. W Rogerson, Twin
Falls County.
Archilochus alexandri (Boucier and Mulsant). Black-chinned Hummingbird.
Rust (1915:125) records this species as resident in Kootenai County.
Selasphorus platycercus platycercus (Swainson). Broad-tailed Humming-
bird. Common resident in southern Idaho. Davis (1935b :236) states that the
bird is of erratic occurrence at the Minidoka Project.
Selasphorus rujus (Gmelin). Rufous Hummingbird. Fairly common resi-
dent. Merrill (1897:355) states that this species is common in spring at Fort
Sherman.
Slellula calliope (Gould). Calliope Hummingbird. Common resident.
(D. A. 1541, 10 mi. NE Moscow, Latah County, May 10, 1940.)
Megaceryle alcyon caurina (Grinnell). Western Belted Kingfisher. Com-
mon resident in suitable localities. (D. A. 1518, 7 mi. NE Moscow, Latah
County, April 19, 1940.)
Colaptes cajer (Gmelin). Red-shafted Flicker. Common resident.
a. collaris Vigors. Red-shafted Flicker. Resident in southwestern and
northern Idaho. Many specimens show yellow remiges and roctricos.
and are perhaps hybrids with the species auratus. (D.A. 1731, Owl
Creek, in Blaine County, September 8, 1940.)
b. canescens Brodkorb. Red-shafted Flicker. Resident in southeastern
Idaho. See Brodkorb (1935a :1).
Hylatomus ' pileatus picinus (Bangs). Western Pileated Woodpecker.
Fairly common resident in the Transition Life-zone. (D.A. 1498, 10 mi. NE
Moscow, Latah County, November 18, 1939.)
Asyndesmus lewis Gray. Lewis Woodpecker. Common resident. Merrill
(1897:354) records this bird as common "around Fort Sherman."
Sphyrapicus varius rmchalis Baird. Red-naped Sapsucker. Fairly common
resident. (D. A. 1485, 10 mi. SW Riggins, Idaho County, September 15, 1939.)
Sphyrapicus thyroideus thyroideus (Cassin). Williamson Sapsucker. Un-
common resident. The Museum of Vertebrate Zoology has one specimen
taken on the W rim Copenhagen Basin, 8400 ft., Wasatch Mountains, Bear
Lake County.
Dendrocopos villosus monticola Anthony. Rocky Mountain Hairy Wood-
Arvey — Birds of Idaho 205
pecker. Common resident. (D. A. 1662, 4 mi. NW Pollock, Idaho County,
July 1, 1940.) .
Dcndrocopos pubescens leucurus (Hartlaub). Batchelder Woodpecker. Com-
mon resident. (D. A. 1495, Potlatch, Latah County, November 3, 1939.)
Dcndrocopos albolarvatus albolarvatus (Cassin). Northern White-headed
Woodpecker. Uncommon resident. (D. A. 1434, 10 mi. SW Ri^gins, Idaho
County, May 14, 1939.)
Pico'ides arcticus (Swainson). Arctic Three-toed Woodpecker. Uncommon
resident in northern Idaho. Merrill (1897:354) reports these birds as resident
at Fort Sherman.
Pico'ides tridactylus (Linnaeus). Uncommon resident.
a. dorsalis Baird. Alpine Three-toed Woodpecker. Resident in south-
ern Idaho ; the Museum of Vertebrate Zoology has specimens taken
at W rim Copenhagen Basin, 8400 ft., Wasatch Mountains, Bear Lake
County.
b. fasciatus Baird. Alaska Three-toed Woodpecker. Resident in north-
ern Idaho. There are specimens in the Museum of Vertebrate Zool-
ogy taken at Coolin, Priest Lake, Kootenai County.
Tyrannic tyrannies (Linnaeus). Eastern Kingbird. Common resident in
northern Idaho; casual in southern portion. (Univ. Idaho, No. 39, Moscow,
Latah County, May 19, 1937.)
Tyrannus verticalis Say. Arkansas Kingbird. Common resident in south-
ern Idaho. (D. A. 1794, Arrowrock Reservoir, Boise County, June 15, 1941.)
Myiarchus cinerascens cinerascens (Lawrence). Ash-throated Flycatcher.
Uncommon resident in southern Idaho. (D. A. 1S37, Head Taylor Creek.
Boise National .Forest, Boise County, August 7, 1941.)
Sayornis saya saya (Bonaparte). Say Phoebe. Fairly common resident in
southern Idaho. (D.A. 1720, 4 mi. NW Pollock, Idaho County.)
Empidonax traillii brewsteri Oberholser. Little Flycatcher. Fairly com-
mon resident in the Transition Life-zone. (Univ. Idaho No. 121, Moscow
Mountain, Latah County, June 15, 1938.)
Empidonax hammondii (Xantus). Hammond Flycatcher. Uncommon res-
ident in the Transition Life-zone. (Univ. Idaho No. 62, Avery, Latah County,
July 10, 1937.)
Empidonax wrightii Baird. Wright Flycatcher. Common resident in the
Transition Life-zone. (D.A. 1560, Robinson's Lake, 10 mi. E Moscow, Latah
County, May 16, 1940.)
Empidonax griseus Brewster. Gray Flycatcher. Davis (1934) records one
specimen of this species taken June 3, 1934, at Riddle, Owyhee County.
Contopus richardsonii richardsonii (Swainson). Western Wood Pewee.
Common resident. (D.A. 1617, 9 mi. ESE Moscow, Latah County, June 5,
1940.)
Nuttallornis boreali-s (Swainson). Olive-sided Flycatcher. Uncommon resi-
dent. (D.A. 1786, Idaho City, Boise County, May 23, 1941.)
Ercmophila alpestris (Linnaeus). Horned Lark. Common resident. See
Behle (1942) for ranges of the following races.
a. lamprochroma Oberholser. Oregon Horned Lark. Southwestern Idaho,
and intergrading with the next two races.
206 University of Kansas Publs., Mus, Nat. Hist. _
b. utahensis Behle. Great Salt Lake Horned Lark. Resident in central
and southeastern Idaho.
c. merrilli Dwight. Dusky Horned Lark. Northern Idaho.
Tachycineta thalassina lepida Mearns. Violet-green Swallow. Common
resident. (D. A. 1654, 4 mi. NW Pollock, Idaho County, June 27, 1940.)
Iridoprocne bicolor (Vieillot). Tree Swallow. Fairly common resident.
Burleigh (1923:655) records the birds at Clark's Fork, Bonner County.
Riparia riparia riparia (Linnaeus). Bank Swallow. Fairlj' common resi-
dent in suitable localities. (D. A. 1453, 4.Vz mi. SW Moscow, Latah County,
May 26, 1939.)
Stelgidopteryx ruficollis serripennis (Audubon). Rough-winged Swallow.
Low (1945:132) records a colony of these birds and Bank Swallows nesting to-
gether at Gray's Lake, in Caribou County.
Hirundo ruslica erythrogaster Boddaert. Barn Swallow. Common resident.
(D. A. 1420, Troy, Latah County, May 6, 1939.)
Petrochelidon pyrronola albijrons (Rafinesque). Northern Cliff Swallow.
Common resident. (D. A. 1415, Troy, Latah County, May 6, 1939.)
Perisoreus canadensis bicolor A. H. Miller. Idaho Jay. Common resident
in central and northern Idaho. (D. A. 1344, Blue Creek, 8 mi. NE Priest
Lake, Bonner County, March 5, 1939.)
Cyanocitta stelleri annectens (Baird). Black-headed Jay. Common resi-
dent. (D.A. 1257, Moscow Mountain, Latah County, October 25, 1938.)
Aphclocoma coerulescens woodhousei (Baird). Woodhouse Jay. Uncom-
mon resident in southern Idaho. The A. O. U. Check-list records this species
from southern Idaho; it is resident in the pihon-juniper association.
Pica pica hudsonia (Sabine). American Magpie. Common resident. (D.
A. 1782, Star, Canyon County, May 1, 1940.)
Corvus corax sinualus Wagler. American Raven. Common resident in
southern Idaho. Davis (1935b :235) lists the bird as a regular winter visitant
at the Minidoka Project.
Corvus brachyrynchos hesperis Ridgway. Western Crow. Common resi-
dent. Davis (1935b :235) lists the bird as a winter visitant at the Minidoka
Project.
Gymnorhinus cyanocephalus Wied. Pihon Jay. Resident locally in
piiion-juniper association. Davis (1935b :235) states that this is a regular
winter visitant in Minidoka County.
Nucijraga Columbiana (Wilson). Clark Nutcracker. Common resident of
forested areas of central and northern Idaho. See Burleigh (1923:655).
Par us atricapillus Linnaeus. Black-capped Chickadee. Very common resi-
dent. See Duvall (1945) for ranges of the following races.
a. septentrionalis Harris. Long-tailed Chickadee. Resident in eastern
Idaho; intergrades with the next two races.
b. nevadensis (Linsdale). Pallid Black-capped Chickadee. Resident in
southwestern and south-central Idaho.
c. fortuilus (Davison and Bowles). Columbian Black-capped Chicka-
dee. Resident in northern and central Idaho.
Par us gambcli Ridgway. Mountain Chickadee. Common resident in the
Transition Life-zone.
Arvey — Birds of Idaho 207
a. grinnelli (van Rossem). Grinnell Chickadee. Resident in central
and northern Idaho. (D. A. 1508, 10 mi. ESE Moscow, Latah County,
March 18, 1940.)
b. inyoensis (Grinnell). Inyo Chickadee. Resident in southeastern
Idaho. (D.A. 1361. Havenor's, 7 mi. NW Pocatello, Power County,
April 1, 1939.)
Pants rufesci ns rufescens Townsend. Chestnut-backed Chickadee. Resi-
dent in central and northern Idaho. Rust (1915:129) records the bird from
Fernan Lake, Kootenai County.
Parus inornatus griseus (Ridgway). Gray Titmouse. Fairly common resi-
dent in southeastern Idaho in the pinon-juniper association. (D. A. 1366, Po-
catello Creek, 3 mi. E Pocatello, Bannock County, April 2, 1939.)
Psallripants minimus plumbeus (Baird). Lead-colored Bush-tit. Uncom-
mon resident in the pihon-juniper association of southern Idaho. The Mu-
seum of Vertebrate Zoology has specimens collected by me at S Fork Owyhee
River, 12 mi. N Nevada line, Owyhee County.
Sit la carolinensis tenuissima Grinnell. Inyo Nuthatch. Fairly common
resident in the Transition Life-zone. (D.A. 1286, 3 mi. NE Princeton, Latah
County, November 20, 1938.)
Sitta canadensis Linnaeus. Red-breasted Nuthatch. Common resident in
the Transition Life-zone. (D.A. 1905, 11 mi. SSW Idaho City, Boise County,
October 20, 1946.)
Sitta pygmaea melanotis van Rossem. Black-eared Nuthatch. Fairly com-
mon resident in the Transition Life-zone. (D.A. 1552, 10 mi. NE Moscow,
Latah County, May 11, 1940.)
Certhia famiKaris caurina Aldrich. Northwestern Creeper. Common resi-
dent in the Transition Life-zone. (D. A. 1304, Paradise Ridge, 3 mi. S Mos-
cow, Latah County, December 10, 1938.)
Cinclus mexicanus unicolor Bonaparte. Dipper. Common resident. Rust
(1915:128) reports that this bird is regularly seen along mountain streams in
Kootenai County.
Troglodytes a'edon parkmanii Audubon. Western House Wren. Common
resident. (Univ. Idaho No. 50, Moscow, Latah County, May 25, 1937.)
Troglodytes troglodytes pacificus Baird. Western Winter Wren. Uncom-
mon resident in the Canadian Life-zone of central and northern Idaho. (D.
A. 1269, Lochsa River, at Van Camp, Idaho County, November 5, 1939.)
Telmatodytes palustris pulverius Aldrich. Northwestern Long-billed
Marsh Wren. Common resident in suitable localities. (D. A. 1769, 2 mi. SW
Notus, Canyon County, February 20, 1941.)
Catherpes mexicanus griseus Aldrich. Northern Canyon Wren. Uncom-
mon resident in southern Idaho, extending north at least to Idaho County.
(D. A. 1702, 4 mi. NW Pollock, Idaho County, July 15, 1940.)
Salpinctes obsoletus obsoletus (Say). Common Rock Wren. Resident in
southern Idaho. (D.A. 1799, Boise, Ada County, June 24, 1941.)
Dumetella carolinensis ruficrissa Aldrich. Western Catbird. Common resi-
dent in northern Idaho, and possibly in the southern portion of the state.
(D. A. 1467, 2 mi. NE Moscow, Latah County, June 2, 1939.)
208 University of Kansas Publs., Mus. Nat. Hist.
Orcoscoptes montanus (Townsend). Sage Thrasher. Resident in the sage-
brush area from Idaho County south. (D. A. 1645, 4 mi. NW Pollock, Idaho
County, June 25, 1940.)
T urdus migratorius Linnaeus. Robin. Common resident in the Transition
Life-zone.
a. caurinus (Grinnell). Northwestern Robin. Common migrant. (Univ.
Idaho No. 216, Moscow, Latah County, August 25, 1937.)
b. propinquus Ridgway. Western Robin. Resident. (D. A. 1893, Boise,
Ada County, May 1, 1944.)
Ixoreus naevius meruloides (Swainson). Northern Varied Thrush. Un-
common resident in the Transition Life-zone. (D. A. 1231, Moscow, Latah
County, October 7, 1938.)
Hylocichla guttata (Pallas). Hermit Thrush. Fairly common resident.
a. guttata (Pallas). Alaska Hermit Thrush. The A. O. U. Check-list
(1931) states that these birds migrate through Idaho.
b. auduboni (Baird). Audubon Hermit Thrush. Resident. (D. A. 1230,
Moscow, Latah County, October 1, 1938.)
Hylocichla ustulata almae Oberholser. Western Olive-backed Thrush.
Fairly common resident. (D. A. 1616, 9 mi. ESE Moscow, Latah County,
June 5, 1940.)
Hylocichla fuscescens salicicola Ridgway. Willow Thrush. Fairly common
resident. The Museum of Vertebrate Zoology has specimens of this species,
taken at Castle Creek Ranger Station, Idaho County, 7 mi. SE Murphy,
Owyhee County, and 3 mi. W Swan Valley, Bonneville County.
Sialia mexicanus occidentalis Townsend. Western Bluebird. Resident in
northern Idaho. Rust (1915:129) states that the species is fairly common at
Coeur d'Alene Lake.
Sialia currucoides (Bechstein). Mountain Bluebird. Very common resi-
dent. (D. A. 1789, Black Creek, 12 mi. SE Boise, Ada County, March 7, 1941.)
Myadestes townsendi (Audubon). Townsend Solitaire. Uncommon resi-
dent in the boreal zones. (D. A. 1294, 7 mi. E Genessee, Latah County, No-
vember 27, 1938.)
Polioptila caendca amoenissima Grinnell. Western Gnatcatcher. Brodkorb
(1935b :312) records one specimen of this bird taken at 6,000 ft. "about eight
miles southwest of Raymond, Bear Lake County," on October 7, 1932.
Regidus satrapa olivaceus Baird. Western Golden-crowned Kinglet. Resi-
dent; fairly common in winter. (D.A. 1229, Moscow, Latah County, Octo-
ber 1, 1938.)
Regulus calendula cineraceus Grinnell. Western Ruby-crowned Kinglet.
Resident; one of the most common winter birds. (D.A. 1902, Cottonwood
Creek, 5 mi. NNE Boise, Ada County, October 5, 1946.)
Anthus spinoletta pacificus Todd. Western Pipit. Common migrant.
(D.A. 1849, Black Creek Reservoir, 12 mi. SE Boise, Ada County, October
11, 1941.)
Bombycilla garmlus pallidiceps Reichenow. Bohemian Waxwing. Common
sporadically in winter. Taylor (1918:226) reported this bird breeding near
Sandpoint, Bonner County.
Bombycilla cedrorum Vieillot. Cedar Waxwing. Very common in winter,
Arvey — Birds of Idaho 209
often with the preceding species; resident in Kootenai and Bonner counties,
:tnd probably elsewhere in the State. Rust (1915:128) records a nest with
three fresh eggs on June 28 at Fernan Creek, Kootenai County.
Lanius excubitor invictus Grinnell. Northwestern Shrike. Casual migrant.
(D. A. 1875, Boise, Ada County, February 3, 1943.)
Lanius ludovicianus gambeli Ridgway. California Shrike. Miller (1931:79)
states that the resident population of this species is referred to this race.
Common resident in the Sonoran zones.
Sturnus vulgaris Linnaeus. Starling. These birds have been reported for
several years; specimens were first reported by Jones (1946:142) from Ban-
nock County.
Vireo huttoni huttoni Cassin. Hutton Vireo. Very common resident in the
Transition Life-zone. (D. A. 1413, Troy, Latah County, May 6, 1939.)
Vireo solitarius cassinii Xantus. Cassin Vireo. Common resident in the
Transition Life-zone. The Museum of Vertebrate Zoology has a specimen
taken 3 mi. W Payette Lake, Adams County.
Vireo olivaceus (Linnaeus). Red-eyed Vireo. Common resident. The
Museum of Vertebrate Zoology has a specimen of this vireo taken 4 mi. W
Meadow Creek, Idaho County.
Vireo gilvus swainsonii Baird. Western Warbling Vireo. Very common
resident. (Univ. Idaho No. 119, Moscow, Latah County, June 14, 1938.)
Vermivora celata orestera Oberholser. Rocky Mountain Orange-crowned
Warbler. Common resident. (Univ. Idaho No. 204, Moscow, Latah County,
August 16, 1938.)
Vermivora ruficapilla ridgwayi van Rossem. Calaveras Warbler. Burleigh
(1923:662) states that this warbler is fairly common at Clark's Fork, Bonner
County, in July and August.
Dendroica petechia morcomi Coale. Rocky Mountain Yellow Warbler.
Very common resident. (Univ. Idaho No. 175, Moscow Mountain, Latah
County, July 29, 1938.)
Dendroica auduboni auduboni (Townsend). Audubon Warbler. Common
resident. (D. A. 1555, 10 mi. NE Moscow, Latah County, May 11, 1940.)
Dendroica nigrescens (Townsend). Black-throated Gray Warbler. Fairly
common in migration, and probably resident. The Museum of Vertebrate
Zoology has a specimen taken at Indian Creek, 12 mi. SE Riddle, Owyhee
County.
Dendroica toumsendi (Townsend). Townsend Warbler. Fairly common in
migration. Burleigh (1923:663) states that the bird is resident at Clark's
Fork, Bonner County.
Seiurus noveboracensis notabilis Ridgway. Grinnell Water-thrush. Merrill
(1897:349) records this bird from the State.
Oporornis tolmiei (Townsend). Macgillivray Warbler. Common resident.
(D. A. 1421, Troy, Latah County, May 6, 1939.)
Geothlypis trichas occidentalis Brewster. Western Yellow-throat. Com-
mon resident in suitable localities. (D. A. 1863, 2 mi. W Boise, Ada County,
May 8, 1942.)
Icteria virens auricollis (Lichtenstein). Long-tailed Chat. Common res-
ident. (D.A. 1800, Cinch Creek, Arrowrock Reservoir, Boise County, June
28, 1941.)
210 University of Kansas Publs., Mus. Nat. Hist.
Wilsonia pusilla pileolata (Pallas). Northern Pilcolated Warbler. Bur-
leigh (1923:663) records this bird as a common resident at Clark's Fork, Bon-
ner County; uncommon in southern Idaho.
Setophaga ruticilla (Linnaeus). American Redstart. There are some rec-
ords of casual visitants in southern Idaho, and Burleigh (1923:663) states that
it is a summer resident at Clark's Fork, Bonner County.
Passer domesticus (Linnaeus). English Sparrow. This cosmopolitan bird
can be found wherever there is a human habitation.
Dolichonyx oryzivorus (Linnaeus). Bobolink. Resident in northern Idaho.
Burleigh (1923:655) states that the bird is resident at Clark's Fork, Bonner
County.
Sturnella neglecta Audubon. Western Meadowlark. Common resident.
(D. A. 1876, Boise, Ada County, May 12, 1943.)
Xanthocephalus xanthocephalus (Bonaparte). Yellow-headed Blackbird.
Common resident along the Snake River in southern Idaho. (D. A. 1628,
Hagerman, on Snake River, Gooding County, June 16, 1940.)
Agelaius phoeniceus (Linnaeus). Red-wing. Common resident.
a. jortis Ridgway. Thick-billed Red-wing Resident in southeastern
Idaho. (D. A. 1624, Hagerman on Snake River, Gooding County,
June 16, 1940.)
b. nevadcnsis Grinnell. Nevada Red-wing. Resident in southwestern
and northern Idaho. (D. A. 1765, Star, Canyon County, May 1, 1941.)
Icterus bullockii bullockii (Swainson). Bullock Oriole. Common resident.
(D. A. 1655, 4 mi. NW Pollock, Idaho County, June 27, 1940.)
Euphagus cyanocephalus (Wagler). Brewer Blackbird. Common resident.
(D. A. 1894, nest and four eggs, Boise, Ada County, May 10, 1944.)
Molothrus ater artemisiae Grinnell. Nevada Cowbird. Fairly common
bird in the Upper Sonoran Life-zone. (D. A. 1460, 4Mj mi. SW Moscow, Latah
County, May 26, 1939.)
Piranga ludoviciana (Wilson). Western Tanager. Very common resident
in the Transition Life-zone. (D.A. 1570, 10 mi. ESE Moscow, Latah County,
May 19, 1940.)
Pheucticus melanocephalus melanocephalus (Swainson). Rocky Mountain
Grosbeak. Resident in the Transition Life-zone. (Univ. Idaho No. 51 Mos-
cow Mountain, Latah County, May 30, 1937.)
Passerina amoena (Say). Lazuli Bunting. Very common resident in the
Upper Sonoran Life-zone. (D. A. 1802, Cinch Creek, Arrowrock Reservoir,'
Boise County, June 28, 1941.)
Hesperiphona vespertina brooksi Grinnell. Western Evening Grosbeak.
Resident in the Transition Life-zone ; large flocks of these birds are commonly
observed in winter. (D. A. 1527, 10 mi. ESE Moscow, Latah County, April
20, 1940.)
Carpodacus cassinii Baird. Cassin Purple Finch. Common resident in the
Transition Life-zone. (D. A. 1822, Head Crooked River, Sawtooth Range,
Boise County, August 6, 1941.)
Carpodacus mexicanus solitudinis Moore. Desert House Finch. Common
resident, (D.A. 1889, Boise, Ada County, April 24, 1944.)
Pinicola enucleator montana Ridgway. Rocky Mountain Pine Grosbeak.
Arvey — Birds of Idaho 211
Resident on the boreal summits of the mountains. (D. A. 1321, Moscow
Mountain, Latah County, January 26, 1939.)
Leucosticte tephrocotis Swainson. Rosy Finch. Resident in the boreal
zones; observed casually in winter. Various races of this species are present
in the State, but only the following two are here listed until there is further
clarification of the status of the other races of the species.
a. lit (oralis Baird. Hepburn Rosy Finch. Winter visitant. (D. A. 1347,
2 mi. N Moscow, Latah County, March 18, 1939.)
b. tephrocotis (Swainson). Gray-crowned Rosy Finch. According to
the 1931 A.O.U. Check-list, this subspecies breeds in the State.
Leucosticte atrata Ridgway. Black Rosy Finch. Resident in the Salmon
Mountains. See A.O.U. Check-list (1931) for the range of this species.
Acanthis flarnmea flammea (Linnaeus). Common Redpoll. Rust (1915:
127) lists this bird as a winter visitant in Kootenai County, and one specimen
was obtained in Bonner County. (D. A. 1334, 6 mi. S Coolin, Bonner County,
February 19, 1939.)
Spinus pinus vagrans Aldrich. Western Pine Siskin. Common resident in
the Transition Life-zone. (D. A. 1857, Horseshoe Bend, Boise County, De-
cember 10, 1941.)
Spinus tristis pallidas Mearns. Pale Goldfinch. Common resident. (D. A.
1622, 4 mi, ESE Boise, Ada County, March 14, 1941.)
Loxia curvirostra Linnaeus. Red Crossbill. Uncommon resident in the'
Canadian Life-zone.
a. bendirei Ridgway. Bendire Crossbill. Resident. (D. A. 1525, 10 mi.
ESE Moscow, Latah County, April 20, 1940.)
b. benti Griscom. Bent Crossbill. Winter visitant. (Univ. Idaho No.
94, Moscow, Latah County, December 5, 1937.)
Loxia leucoptera leucoptera Gmelin. White-winged Crossbill. Davis
(1935b :236) records this bird from the Minidoka Project on December 18,
1919, and Jewett (1912b :193) took one specimen in the Sawtooth Mountains.
Chlorura chlorura (Audubon). Green-tailed Towhee. Breeding individuals
of this species have been taken at the Minidoka Project by Davis (1930:136).
Pipilo macidatus Swainson. Spotted Towhee. Common resident in the
Transition Life-zone.
a. arcticus (Swainson). Arctic Towhee. Resident in northern Idaho.
(Univ. Idaho No. 163, Coeur d'Alene, Kootenai County, July 20, 1938.)
b. curtatus Grinnell. Nevada Towhee. Resident in southern Idaho.
(D. A. 1804, Dutch Creek and Boise River, Boise County, July 4,
1941.)
Calamospiza melanocorys Stejneger. Lark Bunting. Davis (1935b :236) re-
cords this species as erratic at the Minidoka Project, where he took a specimen
on May 29, 1921.
Passerculus sandwichensis nevadensis Grinnell. Nevada Savannah Sparrow.
Common resident. (Univ. Idaho No. 57, Moscow, Latah County, September
25, 1937.)
Pooecetes gramineus confinis Baird. Western Vesper Sparrow. Common
resident. (D. A. 1391, Moscow, Latah County, April 16, 1939.)
Chondestes grammacus strigatus Swainson. Western Lark Sparrow. Com-
212 University of Kansas Publs., Mus. Nat. Hist.
mon resident. (D.A. 1579, 3 mi. SW Moscow, Latah County, May 21, 1940.)
Amphispiza belli nevadensis (Ridgway). Northern Sage Sparrow. Resident
in southern Idaho. Davis (1935b :236) took one specimen in Minidoka on
May 19, 1921.
Junco hyemalis cismontanus Dwight. Slate-colored Junco. Fairly common
winter visitant with other juncos. See Miller (1941:329) for records of these
birds.
Junco oreganus Townsend. Oregon Junco. Common resident. See Miller
(1941:238) for ranges of the following subspecies.
a. mearnsi Ridgway. Pink-sided Junco. Resident in Custer and Fre-
mont counties.
b. montanus Ridgway. Montana Junco. Resident in northern and
western Idaho.
Junco caniceps caniceps (Woodhouse). Gray-headed Junco. Miller (1941:
180) states that some hybridization occurs between this species and oreganus
in Bannock and Cassia counties. It is resident in southeastern Idaho.
Spizella arborea ochracea Brewster. Western Tree Sparrow. Fairly com-
mon resident in central and northern Idaho. (D.A. 1516, nest and eggs,
Moscow, Latah County, April 6, 1940.)
Spizella passerina arizonae Coues. Western Chipping Sparrow. Very com-
mon resident in the Transition Life-zone. (D. A. 1805, junction of Dutch
Creek and Boise River, Boise County, July 4, 1941.)
Spizella breweri breweri Cassin. Brewer Sparrow. Resident in southern
Idaho. Davis (1935b :235) records the bird as a summer resident at the Mini-
doka Project.
Zonotrichia querula Nuttall. Harris Sparrow. Wyman (1911a :267) records
this bird from Nampa, Valley County, in winter.
Zonotrichia leucophrys (Forster). White-crowned Sparrow. Common res-
ident.
a. gambeli (Nuttall). Gambel Sparrow. Migrant. (Univ. Idaho No. 6.
Moscow, Latah County, September 26, 1936.)
b. leucophrys (Forster). White-crowned Sparrow. Resident in the
Hudsonian and Canadian zones. See A.O.U. Check-list (1931) for
range of this subspecies.
Zonotrichia albicollis (Gmelin). White-throated Sparrow. Wyman (1912b:
247) reported this bird from Nampa, Valley County, in winter.
Passerella iliaca schistacea Baird. Slate-colored Fox Sparrow. Uncommon
resident in the Transition Life-zone, and fairly common in migration. (D. A.
1365, Pocatello Creek, 3 mi. E Pocatello, Bannock County, April 2, 1939.)
Melospiza lincolnii alticola (Miller and McCabe). Montane Lincoln Spar-
row. Resident in the boreal zones, and fairly common in migration. See
Miller and McCabe (1935:149) for range of this subspecies.
Melospiza melodia (Wilson). Song Sparrow. Common resident.
a. fallax (Baird). Mountain Song Sparrow. Resident in southern Idaho.
(D.A. 1839, Head Taylor Creek, Boise County, August 7, 1941.)
b. merrilli Brewster. Merrill Song Sparrow. Resident in central and
northern Idaho. (Univ. Idaho No. 103, Moscow, Latah County, Feb-
ruary 22, 1938.)
Calcarius lapponicus alascensis Ridgway. Alaska Longspur. Uncommon
Arvey — Birds of Idaho 213
migrant. Merrill (1S98:15) records one specimen of this species taken at
Fort Sherman on November 13, 1896.
Plcctrophenax nivalis nivalis (Linnaeus). Eastern Snow Bunting. Un-
common migrant. Rust (1915:127) records the bird as rare in migration in
Kootenai County, and Merrill (1898:15) states that it is irregular in winter
at Fort Sherman.
BIBLIOGRAPHY
Aldrich, J. W.
1944. Notes on the races of the white-breasted nuthatch. Auk, 61:592-604.
1946a. New subspecies of birds from western North America. Proc. Biol.
Soc. Washington, 59:129-136.
1946b. Speciation in the white-cheeked geese. Wilson Bull.. 58:94-103.
Aldrich, J. W. and Friedmann, H.
1943. A revision of the ruffed grouse. Condor, 45:85-103.
American Ornithologists' Union Committee.
1931. Check-list of North American birds. Lancaster Press.
1944. Nineteenth supplement to the American Ornithologists' Union check-
list of North American birds. Auk, 61:441-464.
1945. Twentieth supplement to the American Ornithologists' Union check-
list of North American birds. Auk, 62:436-449.
1946. Twenty-first supplement to the American Ornithologists' Union
check-list of North American birds. Auk, 63:428-432.
1947. Twenty-second supplement to the American Ornithologists' Union
check-list of North American birds. Auk, 64:445-452.
Arvey, M. D.
1941. Black-billed cuckoo in Idaho. Condor, 43:291.
1944. Eastern blue-jay in Idaho. Condor, 46:205.
Behle, W. H.
1942. Distribution and variation of the horned larks (Otocoris alpestris) of
western North America. Univ. California Publ. Zool., 46:205-316.
1944. Check-list of the birds of Utah. Condor, 46:67-87.
Bendire, C. E.
1877. Birds of southeastern Oregon. Proc. Boston Soc. Nat. Hist., 19:109-
149.
Bond, R. M.
1946. The peregrine population of western North America. Condor, 48:
101-116.
Brewster, W.
1896. Description of a new warbler and a new song sparrow. Auk, 13:44-47.
Brodkorb, P.
1935a. Two new subspecies of the red-shafted flicker. Occ. Pap. Mus. Zool.,
Univ. Michigan, 314:1-3.
1935b. A new bird for Idaho. Auk, 52:312.
Burleigh, T. D.
1923. Notes on the breeding birds of Clark's Fork, Bonner County, Idaho,
Auk, 40:653-665.
214 University of Kansas Publs., Mus. Nat. Hist.
Coole, H. K.
1915. The present status of the trumpeter swan (Olor buccinator). Auk,
32:82-90.
Coues, E.
1892. Original description of Lewis's woodpecker. Auk, 9:394.
Davis, W. B.
1923. On the avifauna of Minidoka County, and adjacent territory. Mur-
relet, 4:3-4.
1930. Meet Oreospiza chlorura. Oologist, 47:136.
1934. Bird notes from Owyhee County, Idaho. Murrelet, 15:69-72.
1935a. Noon-day feeding of the Pacific nighthawk. Condor, 37:176.
1935b. An analysis of the bird population in the vicinity of Rupert, Idaho.
Condor, 37:233-238.
1936. Broad-winged hawk in Idaho. Condor, 38:86.
Davis, W. B. and Stevenson, J.
1934. The type localities of three birds collected by Lewis and Clark in
1806. Condor, 36:161-163.
DlJVALL, H. J.
1945. Distribution and taxonomy of the black-capped chickadees of North
America. Auk, 62:49-69.
Evendon, F. G., Jr., and Evendon, J. R.
1944. A house finch census at Mountain Home, Idaho. Condor, 46:209.
Grin nell, J.
1904. The origin and distribution of the chestnut-backed chickadee. Auk.
21:364-382.
Hand, R. L.
1933a. The hawk-owl in northern Idaho. Condor, 35:32.
1933b. Summer occurrence of the goshawk in Idaho. Condor, 35:36.
1935. A sight record of the red phalarope (P. Julicans) in northern Idaho.
Auk, 52:180-181.
1938. Notes on some birds nesting in northern Idaho. Condor, 41:84.
Hay ward, C. L.
1934. Important heron rookeries in southeastern Idaho. Auk, 51:39-41
Hxtrley, J. B.
1926. Birds observed in Idaho, Washington, and Oregon. Murrelet. 7:
35-36.
Jewett, S. G.
1912a. Western records of the catbird. Auk, 29:106.
1912b. Some birds of the Sawtooth Mountains, Idaho. Condor, 14:191-194.
Jones, V. E.
1943. White-fronted goose in Idaho. Condor, 45:120.
1946. The starling in Idaho. Condor, 48:142-143.
Kenagy, F.
1914. A change in fauna. Condor, 16:120-123.
Low, J. B.
1945. Clay bank has multiple use for wildlife. Condor, 47:132-133.
Arvey — Birds of Idaho 215
Low, J. B., and Nelson, M.
1945. Recent records of breeding waterfowl in Utah and southern Idaho.
Condor, 47:131-132.
Marshall, W. H.
1940. An "Eagle Guard" developed in Idaho. Condor, 52:166.
McCabe, T. T., and McCabe, E. B.
1933. Hermit thrushes of the northwestern states. Condor, 35:122-123.
Merriam, C. H.
1891. Results of a biological reconnaisance of south-central Idaho. X.
Amer. Fauna, 5:1-108.
1892. The dwarf screech owl (Megascops fiammeolus idahoensis Merriam).
Auk, 9:169-171.
Merrill, J. C.
1897. Notes on the birds of Fort Sherman, Idaho. Auk. 14:347-357.
1898. Notes on the birds of Fort Sherman, Idaho. Auk, 15:14-22.
Miller, A. H.
1931. Systematic revision and natural history of the American shrikes
(Lanius). Univ. California Publ. Zool., 38:11-242.
1933. The Canada jays of northern Idaho. Trans. San Diego Soc. Nat.
Hist., '7:287-296.
1941. Speciation in the avian genus Junco. Univ. California Publ. Zool.,
44:173-434.
Miller, A. H. and McCabe, T. T.
1935. Racial differentiation in Passerella (Melospiza) lincolnii. Condor.
37:144-160.
Moore, R. T.
1939. A review of the house finches of the subgenus Burrica. Condor, 41:
177-205.
Oberholser, H. C.
1918. Notes on the subspecies of Numenius americanus Bechstein. Auk,
35:188-195.
Olson, A. C, Jr!
1943. Starling in northern Idaho.' Condor, 45:197.
Palmer, R. H.
1928. Relative abundance of bird species in southern Idaho, Fresno County,
California, and King County, Washington. Murrelet, 9:28-38.
RlDGWAY, R.
1901-1918. The birds of North and Middle America. U. S. Nat. Mus. Bull.
50, pts. 1-8.
Rust, H. J.
1913. Birds new to the vicinity of Lake Coeur d'Alene, Kootenai County,
Idaho. Condor, 15:41.
1914. Some notes on the nesting of the sharp-shinned hawk. Condor, 16:
14-24.
1915. An annotated list of the birds of Kootenai County, Idaho. Condor,
17:118-129.
216 University of Kansas Publs., Mus. Nat. Hist.
1916. Additional notes on the birds of Kootenai County, Idaho. Condor,
18:81-82.
1917. An annotated list of the birds of Fremont County, Idaho, as observed
during the summer of 1916. Condor, 19:29-43.
1919. A favorite nesting haunt of the Merrill song sparrow. Condor, 21 :
145-153.
1920. The home life of the western warbling vireo. Condor, 22:85-94.
Slipp, J. W.
1942. Franklin's gull in Idaho. Condor, 44:226-227.
Sumnaker, J. L.
1925. Notes from Spokane. Condor, 27:73-74.
Snyder, J. O.
1900. Notes on a few species of Idaho and Washington birds. Auk, 17:
242-245.
Stone, W.
1915. Type locality of Lewis's woodpecker and Clarke's nutcracker. Auk,
32:371-372.
SUGDEN, J. W.
1937. The status of the sandhill crane in Utah and southern Idaho. Con-
dor, 40:18-22.
Tayerner, P. A.
1914. A new subspecies of Dendragapus (Dendragapus obscurus fiemmingi)
from southern Yukon Territory. Auk, 31 :385-388.
Taylor, W. P.
1918. Bohemian waxwing (Bombycilla garrula) breeding within the United
States. Auk, 35:226-227.
Tracy, H. C.
1910. The bobolink in Idaho. Condor, 12:80.
VAN ROSSEM, A. J.
1929. A northern race of the mountain chickadee. Auk, 45:104-105.
Wyman, L. E.
1911a. Harris's sparrow (Zonotrichia querula) in southern Idaho. Auk, 28:
267-268.
1911b. The bobolink again in Idaho. Condor, 13:75.
1911c. The catbird in southern Idaho. Condor, 13:108.
1912a. Bobolink again in Idaho. Condor. 14:41.
1912b. White-throated sparrow in Idaho. Auk, 29:247.
1912c. Oreortyx in Idaho. Auk, 29:538-539.
Transmitted February 12, 1947.
21-6960
S-/VA-L
Subspeciation in Pocket Gophers of Kansas
By
BERNARDO VILLA-R. and E. RAYMOND HALL
ZOOL
LIBRARY
MAR -8 1950
HARYARB
UNIVE 5ITY <
»---■ J
University of Kansas Publications
Museum of Natural History
Volume 1, No. 11, pp. 217-236
November 29, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, H. H. Lane, and Edward H. Taylor
Volume 1, No. 11. pp. 217-236
Published November 29. 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR., STATE PRINTER
TOPEKA. KANSAS
1947
21-8188
MUS. CO^P. ZOOL
LIBRARY
1AR -8 19! 1
Subspeciation in Pocket Goph
ers of Kansas
T By
BERNARDO Vl!fcfcA-R-r-AN©-fc-RA-¥AHOND HALL
Several full species of the genus Geomys have been recorded from
Kansas. The purpose of the study now reported upon was to de-
termine the present taxonomic status of these animals and the dis-
tribution of each within the boundaries of Kansas. No pocket
gopher of any kind has been reported from the southeastern part of
the state; in all other parts Geomys is locally common.
HISTORY
The first published reference that we have found to pocket go-
phers of Kansas is Prof. Spencer F. Baird's (1857:377, 380) mention
of two specimens from Fort Riley. One he identified as Geomys
bursarius (p. 377) and the other (p. 380) he doubtfully referred to
Geomys breviceps. Both specimens were obtained by Dr. W. A.
Hammond. J. A. Allen (1874:49) reported pocket gophers from
Kansas under the generic name "Geomys?". Professor M. V. B.
Knox (1875:21) published a list of Kansas mammals in which he
used the names Geomys bursarius Shaw and Geomys breviceps
Baird, the last one for the specimen taken by Dr. Hammond, at
Fort Riley. Baker (1889:57) employed the name Geomys bursarius
Rich, for the gopher "found along the hundredth meridian, between
N latitude 38° 30' and 39° 307' He reported this animal as com-
mon in western Kansas. Merriam (1895:129) recorded G. bursarius
and G. lutescens from Kansas. Allen (1895:265) recorded five
specimens of Geomys lutescens collected between September 16 and
October 13 at Long Island, Phillips County, Kansas, by W. W.
Granger. Since that time several papers, some of them dealing
mostly with habits of pocket gophers, have been published in which
reference is made to Geomys in Kansas. Hibbard (1933:240) rec-
ognized three species: G. bursarius, G. lutescens, and G. breviceps
llanensis. In 1944 (74-75) he recorded Cratogeomys from Meade
County, on the basis of two skulls dug out of the ground, and he
recognized the same three full species of the genus Geomys that he
did in 1933, along with two additional subspecies.
Specimens to the total number of 335 from Kansas have been
available for the present study of the five subspecies recognized.
The reason for arranging all of the named kinds as subspecies of a
single species is that intergradation has been found to occur be-
(219)
220 University of Kansas Publs., Mus. Nat. Hist.
tween every pair of kinds having contiguous geographic ranges. The
characters previously thought by some writers constantly to differ-
entiate, say, Geomys lutescens of western Kansas from Geomys bur-
sarius of eastern Kansas, prove not to do so; instead, in areas geo-
graphically intermediate between the geographic ranges of the two
kinds, the pocket gophers are intermediate in morphological charac-
ters and therefore are regarded as intergrades. Intergradation of
this kind here is accepted as the criterion of subspecies, and lack
of such intergradation as the criterion of species. Search for struc-
tural characters, distinctive of the different kinds, additional to
those characters noted by other writers, has resulted in the finding
of a few such characters but they too are subject to intergradation.
Therefore the several kinds are arranged as subspecies of a single
species which takes the name Geomys bursarius because it is the
oldest available name. Detailed comment on specimens showing
intergradation are to be found in the accounts of G. b. bursarius
and G. b. major.
METHODS AND ACKNOWLEDGEMENTS
The series with the largest number of individuals from one re-
stricted locality was selected for initial study. These individuals
were segregated by sex, and specimens of each sex were arranged
from oldest to youngest. Each series was divided into age-groups,
and within a given age-group of one sex from one locality of what
was considered as one species, estimation was made of the amount
of individual variation. Thus, it was possible when comparing dif-
ferent kinds of pocket gophers to use only one age class of one sea-
son of one sex.
Age was estimated to some extent by size of animal and nature
of its pelage. The immature pelage is grayer and the hair is more
crinkled than in adults. A more certain guide to age, however, is
furnished by the skull. With increasing age some sutures disappear,
the rostrum increases in length and the ridges marking the limits
of the temporal muscles come to fuse and eventually, in males,
form a high sagittal crest.
Cranial measurements were taken as follows:
Basilar length. — From the anteriormost inferior border of the foramen mag-
num to a line connecting the posteriormost margins of the alveoli of the first
upper incisors.
Length of the nasals. — The greatest length of the nasals.
Zygomatic breadth. — The greatest distance across the zygomatic arches.
Mastoid breadth. — The greatest distance across the mastoids.
Villa-R. — Pocket Gophers of Kansas
221
Breadth of rostrum. — Width, perpendicular to long axis of the skull.
Interorbital constriction. — The least distance between the orbits.
Maxillary tooth row. — The greatest length of the upper molariform tooth
row at the alveolar border.
Extension of premaxillae posterior to nasals. — From the posteriormost bor-
der of the nasals to the posterior end of the extension of a premaxilla.
Depth of skull. — From the median suture of the frontals, on the dorsal
surface of the skull to the median suture of the palatines at the level of the
first molar (not premolar).
Length of rostrum. — From the anterior border of the nasal to the maxilla
at the lateral end of the hamulus of the lacrimal.
In the list of specimens examined, localities are arranged by counties from
west to east, beginning at the northwestern corner of the state; specimens in
each county are arranged from north to south. If several localities are in the
same latitude, the westernmost is listed first. Capitalized color terms are af-
ter Ridgway, Color Standards and Color Nomenclature, Washington, D. C,
1912.
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10 o to *o Miles
lil I ■
• SPECIMEN EXAMINED
<§)lYPE LOCALITY
G.b. lutescens
3
4
G.b. majusculus 5
■■, :.:,;
G.b. jugossicularis
G. b. industrius
G.b. major
Fig. 1. Map showing the geographic distribution of the five subspecies of
the Mississippi Valley pocket gopher, Geomys bursarius, in Kansas, with insert
showing range of the species.
222 University of Kansas Publs., Mtjs. Nat. Hist.
In connection with this study each of the authors acknowledges assistance
from the John Simon Guggenheim Memorial Foundation and one of us (Villa)
is grateful for assistance also to Drs. Isaac Ochoterena and Roberto Llamas
of the Biological Institute of Mexico. For the loan of specimens we are grate-
ful to Dr. William B. Davis, of the Agricultural and Mechanical College of
Texas; Dr. G. C. Rinker, of Hamilton, Kansas; and Mr. A. J. Kirn, of Somer-
set, Texas. Unless otherwise indicated, specimens are in the University of
Kansas Museum of Natural History.
ACCOUNTS OF SUBSPECIES
Geomys bursarius lutescens Merriam
Geomys bursarius lutescens Merriam. North Amer. Fauna, 4:51, Octo-
ber 8, 1890; Scheffer, Technical Bull., U. S. Dept. Agric, 224:6, January,
1931.
Geomys lutescens Merriam, North Amer. Fauna, 8:127-29, January 31.
1895; Lantz, Trans. Kansas Acad. Sci., 19:175, 1905; Lantz, Kansas State
Agric. College Bull., 129:335, April, 1905; Hibbard, Trans. Kansas Acad.
Sci., 36:240, 1933; Black, 30th Bienn. Rept. Kansas State Board Agric,
35:182, 1937; Swenk, Missouri Valley Fauna, 2:1, February 1, 1940; Allen,
Kansas State Teachers College, Emporia, Bull. Inf. in Educ, 20 (no. 5)
:15, May, 1940; Hooper, Occas. Papers Mus. Zool., Univ. Michigan, 420:3,
June 28, 1940.
Geomys lutescens lutescens, Hibbard, Trans. Kansas Acad. Sci., 47:74,
1944.
•
Type locality. — Sandhills on Birdwood Creek, Lincoln County, western Ne-
braska.
Distribution in Kansas. — Northwestern Kansas, eastward certainly to Ellis
County, southward certainly to Scott County.
Description. — Animals with total length averaging no more than 272 mm.;
length of vertebrae of tail averaging no more than 92; hind foot averaging no
more than 35. Color: In autumn pelage, upper parts Light Ochraceous-Buff
becoming Buckthorn Brown in middorsal region and there forming a faint
longitudinal band; sides Pale Yellow Orange. In summer, Buckthorn Brown
on upper parts with a dorsal band, especially distinct on specimens from Ellis
and Trego counties; specimens from farther west lack the distinct dorsal band.
Underparts Gray Drab and sometimes whitish, usually whitish in young speci-
mens; basal color of pelage Deep Neutral Gray; fore and hind feet whitish.
Skull: Zygomatic arch broadly and squarely spreading anteriorly; temporal
impressions uniting to form a low sagittal crest in adult males, but in adult
females and in young males the impressions usually remain apart; shape of
interparietal varying from subquadrate in young specimens to subtriangular
or triangular in adults; in some young specimens the interparietal is reduced
to a minute, ovoid bone.
Comparisons. — See comparisons in the accounts of other sub-
species occurring in Kansas.
Remarks. — In his monographic revision of the pocket gophers,
Merriam (1895:129) recorded 3 "typical or nearly typical" speci-
mens from Trego County, and 18 "non typical" specimens as fol-
lows: Garden Plain, Sedgwick County, 4; Belle Plain, Sumner
Villa-R. — Pocket Gophers of Kansas 223
County, 5; Cairo, Pratt County, 6; Kiowa, Bather County, 2; and
Ellis, Ellis County, 1. A detailed discussion of Merriam's account
of the distribution of Geomys lutescens in Kansas is given by Swenk
(1940:11-12).
Judging by specimens in the University of Kansas Museum of
Natural History, G. bursarius lutescens in Kansas is restricted to
the northwestern part of the state, reaching southward certainly to
Scott County and eastward certainly to Ellis County; precise limits
of distribution of this subspecies are unknown. Additional collecting
is necessary to determine where the range of lutescens meets the
ranges of the other subspecies. The specimens studied are remark-
ably uniform. One specimen obtained in October, in Trego County,
is slightly lighter colored than any other from Kansas. In other
characteristics it agrees with specimens from northwestern Kansas
and from the type locality.
Specimens examined. — Total number 32, as follows: Cheyenne County: 23 mi. (by road)
NW St. Francis, 3. Rawlins County: 2 mi. NE Ludell, 10. Logan County: 5 mi. W El-
kader, 3; no locality more precise than county, 1. Trego County: Wakeeney, 4; 12 mi. S
Collyer, Perrington Ranch, 3; no locality more precise than county, 5. Scott County: 4 mi.
S Scott City, 2. Ellis County: Hays State College Campus, Hays, 1.
Geomys bursarius majusculus Swenk
Geomys bursarius majusculus Swenk, Missouri Valley Fauna, 1:6, De-
cember 5, 1939; Hibbard, Trans. Kansas Acad. Sci., 47:74, 1944.
Geomys bursarius, Baird, Expls. and surveys for a railroad route from
the Mississippi River to the Pacific Ocean, pt. 1, Mammals, 377, 1857;
Merriam, North Amer. Fauna, 8:120, January, 1895; Lantz, Trans. Kansas
Acad. Sci., 19:175, 1905; Lantz, Kansas State Agric. College Bull., 129:335,
April, 1905; Scheffer, Kansas State Agric. College Ento. and Zool. Dept.
Bull., 172:199, September, 1910; Hibbard, Trans. Kansas Acad. Sci., 36:
240, 1933; Allen, Kansas State Teachers College Emporia Bull. Inf. Stud,
in Educ, 20 (no. 5): 15, May, 1940.
. Geomys bursarius bursarius, Black, 30th Bienn. Rept. Kansas State
Board Agric, 35:181, 1937.
Geomys breviceps, Baird, Expls. and surveys for a railroad route from
the Mississippi River to the Pacific Ocean, pt. 1, Mammals, 380, 1857.
Type locality. — Lincoln, Lancaster County, Nebraska.
Distribution in Kansas. — Northeastern Kansas, westward certainly to Clay
and Marion counties and southward certainly to Greenwood County.
Description. — Color: Upper parts Mummy Brown in fresh appearing pelage
of February but in more worn pelage of March more reddish being near (16')
Prout's Brown; top of head and sometimes back darker than rest of upper
parts; underparts usually with some whitish anteriorly; fore and hind feet
and approximately distal half of tail white. Size: Large, total length aver-
aging more than 280 mm. in males and 257 in females; hind foot averaging
35 mm. or more in males. Skull: Large; rostrum averaging more than twice
as long as wide; sagittal crest high in males and barely present in females;
occiput vertical when skull is laid top down ; least width of braincase less than
224 University of Kansas Publs., Mus. Nat. Hist.
distance from alveolus of upper incisor to middle of lateral border of P4 at
alveolar border.
Comparisons. — From Geomys bursarius lutescens, majusculus
differs as follows: Color darker, Mummy Brown to Prout's Brown
instead of Buckthorn Brown. In both sexes: head and body a fifth
to a sixth longer; hind foot 5 to 6 per cent longer; skull averaging
larger in all parts measured except that premaxillae (in each sub-
species) extend equally far posteriorly to nasals; diastema longer in
relation to basilar length; rostrum longer relative to its width; sagit-
tal crest higher; rostrum often more depressed distally; angle of
suture between maxilla and jugal more obtuse.
From G. b. bursarius, according to Swenk (1939:6), majusculus
differs in larger size.
From G. b. illinoensis, majusculus. according to Komarek and
Spencer (1931:405), differs in brownish instead of slate-gray colora-
tion and in two cranial characters as follows: Nasals straight-sided
instead of shaped like an hour-glass, and superficial canals on pala-
tine extending anteriorly beyond first molar, and from there ante-
riorly more or less separated. The first of these characters does not
always hold; occasional individuals of majusculus, for example
some from Douglas County, have the nasals shaped like an hour-
glass.
From G. breviceps dutcheri, majusculus differs in larger size
(hind foot more than 33 mm. in males, and 29 in females; basilar
length more than 42 mm. in males and 36 in females) ; dorsal ex-
posure of jugal longer than width of rostrum measured between
ventral margins of infraorbital foramina.
From G. bursarius major of southcentral Kansas (for example
Harvey County), majusculus differs in slightly darker color, being
Mummy Brown instead of Prout's Brown; size larger (in males
total length more than 284 mm., hind foot 35 or more, basilar length
of skull more than 42, and in females total length 265 or more, hind
foot averaging 33 or more, and basilar length 40 or more).
Skull : Averaging larger, in all parts measured, except that pre-
maxillae do not extend so far posteriorly to nasals in either males
or females; interorbital constriction slightly narrower in adult fe-
males; temporal ridges forming a more prominent sagittal crest in
adult males (sagittal crest barely present in some adult males of
major from Harper County).
Remarks. — In employing the subspecific name majusculus we are
following Swenk (1939:6) who on the basis of larger size differen-
Villa-R. — Pocket Gophers of Kansas 225
tinted the animals from southeastern South Dakota, the eastern
parts of Nebraska and Kansas, and the western and southern parts
of Iowa, from G. bwsarius bursarius to which he assigned a more
northern geographic range. In the absence of comparative mate-
rials of the northern subspecies we cannot make an independent de-
cision on the validity of majusculus and recognize that if it is in-
separable from G. b. bursarius the latter name will apply to
specimens from northeastern Kansas. We are the more uncertain
about applying the name majusculus to specimens from eastern
Kansas because they average smaller than topotypes. Only at the
northeasternmost locality in Kansas (3 mi. N Cummings, Atchison
County) do specimens average as large as topotypes of majusculus.
Farther southward they become progressively smaller in eastern
Kansas, and we interpret this as intergradation with the still smaller
subspecies major, to the southwest. The average external measure-
ments of two adult males from Atchison County are: 321-99-35.
Thirty-six miles farther south, in Douglas County, 16 adult males
average 289-80-36. From Hamilton, Greenwood County, 80 miles
farther southwest, nine adult males average 284-83-35. The maxi-
mum total length recorded at these three localities is: Atchison
County, 342 (1 of 2 specimens), Douglas County, 308 (1 of 16
specimens) , Greenwood County, 357 (in coll. of Dr. Glenn C. Rinker
and 1 of 15 males of all ages involved). It will be seen, therefore,
that although there is a trend to smaller average size toward the
southward, the maximum of 357 millimeters total length at Hamil-
ton exceeds the maximum of 352 millimeters recorded by Swenk
(1939:3) among 86 males at Lincoln where the recorded average is
largest.
Four specimens from Salina (Debold Farm) are intermediate
structurally, as they are also geographically, between G. b. majus-
culus on the one hand and Geomys bursarius lutescens and Geomys
bursarius major on the other hand. In color they agree with majus-
culus, as they do also in width of nasals posteriorly, in more obtuse
angle of the rostrum and maxillary arm of the zygomatic arch. They
agree with G. b. lutescens in having the occiput inclined anterodor-
sally, and are intermediate between majusculus and lutescens, but-
nearer the latter in size of skull and in length of the rostrum relative
to its width.
Specimens examined. — Total number, 148, as follows: Clay County: 6 mi. SW Clay
Center, 3. Jackson County: 10% mi. WSW Holton, 1; no locality more precise than county,
1. Atchison County: 3 mi. N Cummings, 2. Jefferson County: Oskaloosa, 1. I^eavenworth
County: Fort Leavenworth (Government Hill, 2; Engineer Hill, 1), 6; no locality more pre-
226 University of Kansas Publs., Mus. Nat. Hist.
cise than county, 19. Saline County: Salina, Debold Farm, 4 (coll. of A. J. Kirn). Morris
County: \y2 mi. N Council Grove, 3. Douglas Comity: 1 mi. NW Midland, 2;1 mi. N
Lawrence, 1; iy2 mi. W Lawrence, 2; 1 mi. W K. U. Campus, 2; 1 mi. W Lawrence, 2; y2
mi. W Lawrence, 2; "W K. U. Campus," 2; K. U. Campus, 4; Lawrence, 23; South Law-
rence, 1 ; y2 mi. SW K. U. Campus, 2 ; Southwest K. U. Campus, 1 ; Haskell Institute, 1 ;
iV2 mi. S Lawrence, 1; 7 mi. SW Lawrence, 6; 7V2 mi. SW Lawrence, 1; 8 mi. SW Law-
rence, 1; 10 mi. S Lawrence, 1; 11 mi. SW Lawrence, 3; no locality more precise than
county, 15. Marion County: \y2 mi. NE Lincolnville, 6; 4 mi. SE Lincolnville, 1; 6 mi.
S Lincolnville, 1. Greenwood County: Hamilton, 1; y2 mi. S Hamilton, 4; 1 mi. S Hamil-
ton, 4; 4 mi. S and 14 mi. W Hamilton, 6; 8 mi. SW Toronto, 1; sy2 mi. SW Toronto, 5;
no locality more precise than county, 6.
Geomys bursarius jugossicularis Hooper
Geomys lutescens jugossicularis Hooper, Occas. Papers Mus. Zool., Univ.
Michigan, no. 420: 1, June 28, 1940; Hibbard, Trans. Kansas Acad. Sci.,
vol. 47, p. 75, 1944.
Type locality. — Lamar, Prowers County, Colorado.
Distribution in Kansas. — Extreme southwestern part of state, northward cer-
tainly to Hamilton County and south certainly to Morton and Seward counties.
Description. — A yellowish-cinnamon colored animal, with body of medium
size, zj'gomatic plate of maxilla deep and mastoid process small.
Comparisons. — Differs from Geomys bursarius industrius in
slightly lighter color; occiput not strongly inclined anterodorsally.
From G. b. lutescens, jugossicularis differs in less buffy coloration
and deeper zygomatic plate of maxilla.
Remarks. — G. bursarius jugossicularis and G. bursarius industrius
intergrade in the southern part of Meade County. Some specimens
from this area show a coloration resembling that of G. b. jugossicu-
laris; nevertheless, one specimen from Morton County has the occi-
put anterodorsally inclined as in G. b. industrius.
Specimens examined from Hamilton County correspond closely to
G. b. jugossicularis; they agree with it both in color and in cranial
characters.
Specimens examined. — Total number, 20, distributed as follows: Hamilton County: 1 mi.
E Coolidge, Conard Farm, 4. Morton County: 12 mi. NE Elkhart, 2; Cimarron River, 12
mi. N Elkhart, 4; no locality more precise than county, 6. Seward County: 1 mi. E
Arkalon, 4.
Geomys bursarius industrius, new subspecies
Geomys lutescens Merriam, North Amer. Fauna, 8:127, January 31, 1895.
Geomys breviceps llanensis, Hibbard, Trans. Kansas Acad. Sci., 36:240.
1933; Black. 30th Bienn. Rept. Kansas State Board Agric, 35:181. 1937.
Geomys lutescens jugossicularis Hooper, Occas. Papers Mus. Zocil.,
Univ. Michigan, 420:1. June 28, 1940.
Type.— Male, adult, skin and skull, no. 14083 Museum of Natural History,
University of Kansas; from 1V> miles north of Fowler, Meade County, Kan-
sas; obtained December 30, 1941, by H. H. Hildebrand, original number 16.
Distribution in Kansas. — Southwestern Kansas from Meade County east-
Villa-R. — Pocket Gophers of Kansas
227
ward certainly to Pratt and Clark counties; from Pawnee County southward
probably to the Oklahoma boundary.
Diagnosis. — Size of body medium; color of upper parts Cinnamon Brown;
skull with occiput .strongly inclined anterodorsally in males.
Description. — Color: Upper parts Cinnamon Brown, slightly reddish, but
in some specimens collected in September, in Pawnee County, near (15' i)
Ochraceous-Tawny; underparts usually Wood Brown, somewhat whitish an-
Fi<i. 2. Three views of the skull of the type specimen of Geomys bursarius industrius
A. Lateral view; B. Dorsal view; C. Ventral view. All natural size.
teriorly; forefeet white; hind feet and approximately distal half of tail whit-
ish. Size: Medium (see measurements), total length averaging not more
than 271 mm. in males and 254 in females; hind foot averaging not more
than 35 mm. in males and less than 32 in females. Skull: In males, least
width of braincase equal to distance from alveolus of incisor to anterior bonier
of alveolus of first upper molar, occiput strongly inclined anterodorsally. tem-
poral impressions usually united in a low sagittal crest, zygomatic arch heavy
and curved at level of jugal bone. In adult females least width of braincase
approximately equal to distance from alveolus of incisor to anterior bonier
of alveolus of first upper molar (not premolar) ; occiput less inclined antero-
228 University of Kansas Publs., Mus. Nat. Hist.
dorsally than in males; temporal ridges not forming a sagittal crest. In young
females the width of the braincase is more than the distance between the
alveoli of the incisor and first molar.
Comparisons. — G. lutescens industrius differs from G. lutescens
lutescens in: Color darker; least width of braincase not equal to
(usually more than) the distance from the alveolus of incisor to
the anterior border of the alveolus of the first upper molar.
G. lutescens industrius differs from G. lutescens jugossicularis in:
Color slightly darker, the former being Cinnamon Brown instead of
Vinaceous Cinnamon, with hairs basally Deep Neutral Gray in up-
per parts and underparts. Skull: Jugular part of zygomatic arch
more curved (convex dorsally) and occiput far more inclined antero-
dorsally ; lower part of mastoidal ridge more prominent.
For comparison with G. I. major, see account of that subspecies.
Remarks. — Judging from the known specimens of this subspecies,
it has the smallest geographic range of any of the subspecies in
Kansas, but additional collecting in Hodgeman County and coun-
ties to the north and west of it may extend the known range in those
directions; collecting in Comanche County and in adjoining parts
of Oklahoma may extend the known range to the southward.
The anterodorsal inclination of the occiput in males is the one
cranial character in which industrius differs from all of the sub-
species with adjoining geographic ranges. The existence of this
unique (among adjoining subspecies) cranial character is the prin-
cipal reason for according subspecific status to this animal. Al-
though it has other characters which are fairly uniform over a
considerable geographic area, these other characters, namely, Cin-
namon Brown color of the upper parts and medium size of the body,
after all, are conditions intermediate between those in jugossicu-
laris to the west and those in the darker and larger animals assigned
to major to the eastward. Considering the intermediate geographic
position of industrius, the color and size are approximately what a
person would predict by study of only the animals to the west and
those to the east. Therefore, the color and size probably are in-
dicative of intergradation between jugossicularis and major. Still,
there is the anterodorsally inclined occiput in males— a character
of a unique sort — and this influences us to give subspecific status to
this animal with full recognition of the fact that it is a "weak" sub-
species as compared with any one of the adjoining subspecies.
Hooper (1940:2) in naming as new Geomys lutescens jugossicu-
laris referred to his new subspecies a skin-only from Meade County
Yilla-R. — Pocket Gophers of Kansas 229
State Park. Our more abundant material from there shows the
cranial conformation to be that of industrius to which we accord-
ingly assign the specimens. However, with only a skin available,
we, too, would have used the name jugossicularis because the color
is paler than in other specimens of industrius and this paleness in-
dicates intergradation between the two named subspecies. Speci-
mens from Pratt County are slightly darker than industrius thereby
indicating intergradation between industrius and major.
Specimens examined. — Total number, 58, distributed as follows: 'Pawnee Count;/: Jet.
Pawnee and Arkansas rivers, .Larned, 6; 1 mi. S and 1 mi. E Larned, 7. Edwards County:
1 mi. W and 3% mi. S Kinsley, 1. Kiowa County: Rezeau Ranch, 5 mi. N Belvidere, 2.
Pratt County: Pratt, 14; no locality more precise than county, 1. Meade County: 3% mi.
NE Fowler, 2; 2 mi. N Fowler, 2; iy2 mi. N Fowler, 2; 1% mi. N and % mi. E Fowler,
2; 7 mi. N Meade, Cudahy Ash Pit, 2; 13 mi. SW Meade, 9; State Lake, 2; State Park, 4.
Clark County: 7 mi. SW Kingsdown, E. A. Stephenson Ranch, 1; .6 mi. S Kingsdown, 1.
Geomys bursarius major Davis
Geomys hdescens major Davis, Texas Agric. Exp. St., Bull. no. 590:32,
August, 1940; Hibbard, Trans. Kansas Acad. Sci., 47:75, 1944.
Geomys lutescens Merriam, N. Amer. Fauna, 8:129, January 31, 1895.
Geomys breviceps llancnsis, Lantz, Trans. Kansas Acad. Sci., 20 (pt. 2) :
215, 1907; Hibbard, Trans. Kansas Acad. Sci., 36:240, 1933; Black, 30th
Bienn. Rept. Kansas State Board Agric, 35:182, 1937; Swenk, Missouri
Valley Fauna, 2:12, February 1, 1940.
Type locality. — Eight miles west of Clarendon, Donley County, Texas.
Distribution in Kansas. — Southcentral Kansas, northward certainly to Ells-
worth County, westward certainly to Stafford and Barber counties and east-
ward to Cowley County.
Description. — Color: Upper parts varying from Brussels Brown in some
specimens to nearly Prout's Brown, especially in specimens from central part
of state. Top of head, and sometimes back, darker than rest of upper parts,
but no well denned black stripe; underparts varying from whitish to nearly
Buffy Brown; fore and hind feet and approximately distal half of tail white.
Size: Large (see measurements). Skull: Sagittal crest absent in females and
barely present in males; least width of braincase more than distance from al-
veolus of incisor to middle of lateral border of P4 at alveolar border. Length
of auditory bulla (from anteroventral edge of paroccipital process of exoccipi-
tal to hamulus of peterygoid), in each sex, more than 8 mm.; occiput usually
vertical when skull is laid top down; zygomatic arch broadly and squarely
spreading, divergent anteriorly; rostrum averaging less than <wice as long as
wide.
Comparisons. — From G. bursarius lutescens, major differs in color
darker, premaxillae extending slightly farther posteriorly ; temporal
impressions usually forming a more well-marked sagittal crest in
males; ventral side of zygomatic arch, at level of jugal bone, more
curved.
From G. bursarius majiisculus} major differs in slightly lighter
230 University of Kansas Publs., Mus. Nat. Hist.
color, smaller size of body; in males, total length less than 284 mm.;
hind foot 34 or less ; basilar length of skull less than 42 ; in females
total length less than 264, hind foot no more than 33, and basilar
length less than 39.
From G. bursarius industrius, major differs in color, being Prout's
Brown, instead of Cinnamon Brown (less Fuscous) ; body averaging
10 per cent longer; total length in males from 9 to 9.7 per cent
longer, hind foot 9.7 per cent longer on the average ; skull averaging
larger in all parts measured. Occiput less inclined anterodorsally ;
top nearly fiat, less arched than that of G. b. industrius; auditory
bulla averaging slightly larger and less inflated.
Remarks. — Specimens of this subspecies from Norman, Cleveland
County, Oklahoma, and Canton, Dewey County, Oklahoma, and
most of those from Kansas, are more Fuscous than topotypes and
tend toward G. bursarius majusculus. Specimens from McPherson
County have a darker dorsal stripe resembling that of G. bursarius
majusculus. One adult from Little Salt Marsh, Stafford County, is
pale, closely resembling topotypes.
Most of the cranial characters, nevertheless, are constant in all
available specimens, except that in specimens of each sex from the
type locality the basilar length averages 4 to 5 per cent shorter. In
the constancy of size of the relatively large auditory bullae and in
the nearly flat dorsal profile of the cranial part of the skull, the
specimens from Kansas agree with the specimens from the type
locality.
Specimens from Harper County have the occiput slightly inclined
anterodorsally and thus are reminiscent of industrius which has an
even greater inclination of the occiput. Probably the appearance
in dilute fashion of this character in Harper County is properly to
be interpreted as intergradation with industrius. If so, the actual
intergradation may be to the northwest via Pratt County since
specimens from Barber County, immediately west of Harper and
lying between Harper County and the range of industrius, do not
have the occiput so inclined.
Of a pair of adults from eight miles west of Rosalia, Butler
County, the female is indistinguishable in color from adults of G. b.
industrius from northern Meade County and from two specimens
from eleven miles west of Clarendon, Donley County, Texas, near
the type' locality of G. b. major. The male from eight miles west of
Rosalia is darker as compared either with G. b. industrius or G. b.
major and the coloration of the upper parts resembles those in G. b.
Villa-R. — Pocket Gophers of Kansas 231
majusculus; the underparts are only slightly paler than the upper
parts as in majusculus. Measurements of the skulls are intermedi-
ate between the averages for G. b. majusculus and those for G. b.
major. These specimens from eight miles west of Rosalia are inter-
mediate structurally, and since they are intermediate geograph-
ically between G. b. majusculus and G. b. major, they suggest inter-
gradation of the two subspecies. The specimens in question are re-
ferred to major because the size is nearer that of major. It is
mainly the intermediate nature of these two specimens from Butler
County, and the intermediate nature of the specimens from Mc-
Pherson County, Kansas, that have caused us to treat G. b. majus-
culus as only subspecifically distinct from the more western sub-
species, major.
Specimens examined. — Total number, 77, as follows: Ellsworth County: 2 mi. S Ells-
worth, 1. McPherson County: Smoky Hill River, 1 mi. S and V2 mi. W Lindsborg, 5 ; Ms
mi. E McPherson, 1. Stafford County: Little Salt Marsh, 12; no locality more precise than
county, 3. Reno County: 8 mi. N and 1 mi. E Haven, 2. Harvey County: 1 mi. E and V2
mi. N Halstead, 1; Halstead, 3. Butler County: 8 mi. W Rosalia, 2. Barber County:
near South Bridge, Sun City, 1; 2 mi. S Sun City, 1; Wells Ranch, Aetna, 5; "1 mi. W
Aetna," 3; near South Bridge, Aetna, 1; near Bridge, 1 mi. S Aetna, 2. Harper County:
414 mi. NE Danville, 8; 1 mi. N Harper, 11; 3 mi. S Harper, 1. Cowley County :f 3 mi.
SW Arkansas City, 4; 3 mi. SE Arkansas City, 9; 3 mi. S Arkansas City, 1.
232 University of Kansas Publs., Mus. Nat. Hist.
Measurements of Adult Males of Geomys
(In millimeters)
~^> 3
•a ai a
C M 3
— a ~
u
O > 3
p m a
E g-3
3 3 o
ex
C
O
H
o
c
•a
c
j5
g
M
a
<u
-a
o
03
09
s
s
o
w
c
bx
03
01
>>
ffl
J
S3
-a
03
O
■a
J3 o
.3
JZ c
03
11
9 H
O O
P. aj
afi+» t*-.
2 « G
S 03
-S3
3
3
5 ave.
min.
max.
16 ave.
min.
max.
4 ave.
min.
max.
8 ave.
min.
max.
11153
11152
12870
12892
266 82.0
257 76.0
276 91.0
12088 272 92.0
289 79.8
273 70.0
308 95.0
265 82 . 0
250 68.0
285 92 . 0
265 82.0
247 70.0
280 90.0
11724 256
240
240
G. b. lutescens; topotypes
34.2 40.0 17.7 30.5 26.8 11.5 6.7
33.0 38.3 16.0 29.1 26.1 11.2 6.3
36.0 42.4 20.3 31.7 27.5 11.9 6.9
2 mi. NE Ludell, Rawlins Co., Kansas
35.0 43.2 19.1 32.3 27.7 11.3 6.6
8.6
8.1
9.2
G. b. majusculw; Douglas Co., Kansas
36.3 47.1 21.0 *34.1 30.4 12.1 6.8 9.3
32.0 44.7 18.9 30.5 27.5 11.1 6.5 8.2
55.0 49.9 23.2 38.0 34.5 13.5 7.6 10.3
G. b. jugossicularis ; Morton Co., Kansas
34.2 40.7 16.9 30.0 27.9 10.7 6.0 8.6
30.0 38.5 16.1 29.0 27.5 10.5 5.5 8.2
37.0 42.4 17.4 31.1 28.4 11.0 6.2 9.2
G. b. i7idustrius; Meade Co., Kansas
35.0 40.9 18.1 30.0 28.0 11.0 6.2 8.8
33.0 37.9 15.5 28.2 26.5 9.9 5.7 8.0
36.0 43.4 21.0 32.4 29.5 11.6 7.0 9.1
3.9
3.5
4.2
17.1 20.8
16.2 19.1
17.7 23.6
8.4 2.8 18.0 22.1
3.7 18.5 24.9
2.9 17.3 22.9
5.7 20.0 28.1
4.7
5.5
4.3
2.9
G. b. major; Wells Ranch, Aetna, Barber Co., Kansas
66.0 34.0 41.0 18.3 31.6 28.2 10.6 6.1 9.0 4.0
1 mi. W Aetna, Barber Co., Kansas
75.0 32.0 36.7 15.7 26.9 24.6 9.9 5.9 8.8 4.0
65.0 32.0 36.0 14.2 26.1 25.4 10.9 5.6 8.5 5.0
17.3 21.2
16.4 20.2
17.9 22.0
17.7 21.8
16.8 19.5
19.1 24.2
17.0 21.3
15.0
15.5
19.5
18.5
3 mi. SE Arkansas City, Cowley Co., Kansas
246 76.0 32.0 42.1 J16.0 33.7 29.7 11.5 6.3 9.4 4.5 17.6 $21 3
3 mi. SW Arkansas City, Cowley Co., Kansas
282 84.0 33.0 41.7 17.3 .... 27.7 10.8 6.4 8.9 4.2 17.2 21.5
Villa-R. — Pocket Gophers of Kansas
233
Measurements of Adult Females of Geomys
(In millimeters)
Number of individ-
uals averaged or
catalogue number
J3
+^>
&
c
CD
"a
o
'3
o
t
a
►J
Length of hind foot
Basilar length
Length of nasals
Zygomatic breadth
Mastoid breadth
Breadth of rostrum
Interorbital constric-
tion
1
.a $
c ^
£ C
a
J3V.
o a
l a
<
1 fe
eu o
a fc
■ i~ «
°*
c a
■S3
S i
V. a
to nasals
Depth of skull
1
O
o
x;
-&
a
CD
G. b. lutescens; topotypes
6 ave.
min.
max.
233
215
254
72.3
63.0
76.0
31.1 35.3 15.0 25.9 23.7 10.4 6.1
30.0 33.5 13.9 24.6 21.8 10.1 5.6
32.0 37.0 16.8 26.7 24.8 10.7 6.6
2 mi. NE Ludell, Rawlins Co., Kansas
8.3
8.1
8.5
3.7
2.9
4.5
15.4
14.8
16.2
18.4
17.3
19.8
11733
12155
230
245
63.0
70.0
31.0 35.3 15.1 26.5 24.1 9.3 6.1
30.0 35.6 14.6 25.2 24.1 10.6 6.4
G. b. nmjusculus ; Douglas Co., Kansas
7.5
7.5
2.4
3.1
15.0
14.9
18.2
18.2
17 ave.
min.
max.
265
222
304
78.6
59.0
92.0
32.8 °40.6 °17.2 *28.6 26.4 10.9 6.5
30.0 37.1 15.9 26.7 24.9 10.0 5.9
35.0 47.0 20.1 33.4 29.1 12.3 7.3
G. b. jugossicularis ; Morton Co., Kansas
9.1
8.5
10.0
3.6
2.0
5.9.
16.6
15.2
19.1
21.0
18.8
24.1
5012
5395
244
230
72.0
72.0
30.0 36.2 16.4 25.4 25.0 10.0 5.9
30.0 34.6 13.9 24.7 24.8 9.8 5.8
8.0
8.0
4.2
4.5
16.0* 19.3
15. 2{ 17.5
G. b. industrius; Meade Co., Kansas
7 ave.
min.
max.
238 §73.0
231 65.0
256 75.0
31.3 t36.4 14.9 26.3 f24.8 10.0 6.0
30.0 35.4 14.0 25.8 24.5 9.5 5.6
32.0 37.8 16.1 27.8 25.9 10.3 6.5
8.4
8.1
8.7
4.1
3.6
4.7
16.2
15.5
17.6
18.6
17.5
19.9
G. b. major; 1 mi. S Aetna, Barber Co., Kansas
10069
257
95.0
32.0 37.0 16.4 26.4 25.5 10.8 6.2
Aetna, Barber Co., Kansas
9.0
3.4
16.4
19.4
10070
242
83.0
30.0 36.8 15.7 26.2 25.0 10.1 65
Wells Ranch, Aetna, Barber Co., Kansas
9.1
3.3
15.8
19.1
12238
239
65.0
31.0 34.2 14.5 24.6 23.7 9.6 6.0
1 mi. S Sun City, Barber Co., Kansas
8.0
3.6
15. 2_
17.7
11075
232
66.0
28.0 34.2 14.4 25.0 23.6 9.9 5.9
3 mi. SW Arkansas City, Cowley Co., Kansas
8.0
3.4
15.0
17.0
12872
242
66.0
30.0 38.1 15.0 28.0 26.2 10.3 6.3
7.8
4.5
16-U.19.1
3 mi. SE Arkansas City, Cowley Co., Kansas
12894
12893
230
246
82.0
83.0
30.0 38.5 15.5 28.0 25.6 10.0 6.7
32.0 36.5 14.2 25.6 24.8 9.6 6.6
8.7
8.7
4.0
4.6
16.6
15.4
19.5
18.1
* 15 averaged.
° 16 averaged.
| 6 averaged,
t 5 averaged.
t aproximate.
234 University of Kansas Publs., Mus. Nat. Hist.
SUBSPECIES OF THE SPECIES GEOMYS BURSARIUS
If Geomys lutes cens major Davis is correctly judged to intergrade
with Geomys busarius majusculus Swenk, the name for the full
species will be Geomys bursarius because bursarius is the oldest
name among those available. Some new combinations of names
are required. According to our present understanding, the eleven
kinds of pocket gophers named below are properly to be arranged
as subspecies of the species Geomys bursarius :
Geomys bursarius bursarius (Shaw). Type from unknown locality in Up-
per Mississippi Valley.
Geomys bursarius majusculus Swenk. Type from Lincoln. Lancaster
County, Nebraska. -
Geomys bursarius hylaeus Blossom. Type from 10 mi. S Chadron, Dawes
County, Nebraska.
Geomys bursarius levisagittalis Swenk. Type from Spencer, Boyd County,
Nebraska.
Geomys bursarius vinaceus Swenk. Type from Scottsbluff. Scotts Bluff
County, Nebraska.
Geomys bursarius lutescens Merriam. Type from Sandhills on Birdwood
Creek, Lincoln County, Nebraska.
Geomys bursarius illinoensis Komarek and Spencer. Type from 1 mi. S
Momence, Kankakee County. Illinois.
Geomys bursarius jugossicularis Hooper. Type from Lamar, Prowers
County, Colorado.
Geomys bursarius industrius new subspecies. Type from l1/? mi. N Fowler,
Meade County, Kansas.
Geomys bursarius major Davis. Type from 8 mi. W Clarendon, Donley
County, Texas.
Geomys bursarius llanensis Bailey. Type from Llano, Llano County, Texas.
Villa-R. — Pocket Gophers of Kansas 235
LITERATURE CITED
Allen, J. A.
1874. Notes on the mammals of portions of Kansas, Colorado, Wyoming
and Utah. Part I. On the mammals of middle and western Kansas.
Bull. Essex Inst., 6 (no. 2) :43-52. February, 1874.
1895. List of mammals collected in the Black Hills region of South Dakota
and in western Kansas by Mr. Walter W. Granger with field notes
by the collector. Bull. Amer. Mus. Nat. Hist, 7:259-274. August 21,
1895.
Allen, P.
1940. Kansas mammals. Kansas State Teachers College, Emporia, Bull.
Inf. Stud, in Educ, Number 20 (no. 5) : 1-62. May, 1940.
Baker, A. B.
1889. Mammals of western Kansas. Trans. Kansas Acad. Sci, 11:56-58
(for 1887-88).
Baird, S. F.
. 1857. Explorations and surveys for a railroad route from the Mississippi
River to the Pacific Ocean. War Department. Mammals, Part I,
xxxii + 757, pis. 17-60, 35 figs, in text, 1857.
Black, J. D.
1937. Mammals of Kansas. Thirtieth Bienn. Rept. Kansas State Board of
Agric, 35:116-217.
Davis, W. B.
1940. Distribution and variation of pocket gophers (Genus Geomys) in the
southwestern United States. Texas Agric. Exp. Station, Bull., 590:
1-38, 6 figs, in text. October 23, 1940.
Hibbard, C. W.
1933. A revised check list of Kansas mammals. Trans. Kansas Acad. Sci.,
36:230-249.
1944. A checklist of Kansas mammals, 1943. Trans. Kansas Acad. Sci.,
47:61-88.
Hooper, E. T.
1940. A new race of pocket gopher of the species Geomys lutescens from
Colorado. Occas. Papers, Mus. Zool., Univ. Michigan, 420:1-3. June
28, 1940.
Knox, M. V. B.
1875. Kansas Mammalia. Trans. Kansas Acad. Sci., 4:18-22.
Komarek, E. V, and Spencer, D. A.
1931. A new pocket gopher from Illinois and Indiana. Journ. Marnm., 12:
404-408, 1 pi, 1 fig. in text. November 11, 1931.
Lantz, D. E.
1905. Kansas mammals in their relations to agriculture. Kansas State
Agric. College Bull, 129:331-404. April, 1905.
1905. A list of Kansas mammals. Trans. Kansas Acad. Sci, 19:171-178.
1907. Additions and corrections to the list of Kansas mammals. Trans.
Kansas Acad. Sci, 20 (pt. 2) :214-217.
236 University of Kansas Publs., Mus. Nat. Hist.
Merriam, C. H.
1890. Descriptions of twenty-six new species of North American mammals.
N. Amer. Fauna, 4: v + 60, 3 pis., 3 figs, in text. October 8, 1890.
1895. Monographic revision of the pocket gopher Family Geomyidae . . . .
N. Amer. Fauna, 8:1-258, 19 pis. and frontispiece, 71 figs, in text, 4
maps. January 31, 1895.
Scheffer, T. H.
1910. The pocket gopher. Kansas State Agric. Coll. Ent. and Zool. Dopt.,
Bull., 172:197-233, illustrated. September, 1910.
1931. Habits and economic status of the pocket gophers. TJ. S. Dept. Agric,
Tech. Bull., 224:1-27, 8 pis., 2 figs, in text. January, 1931.
Swenk, M. H.
1939. A study of local size variations in the prairie pocket gopher (Geomys
bursarius), with description of a new subspecies from Nebraska. Mis-
souri Valley Fauna, 1:1-8. December 5, 1939.
1940. A study of subspecific variation in the yellow pocket gopher (Geomys
lutescens) in Nebraska, and the geographical and ecological distribu-
tion of the variants. Missouri Valley Fauna, 2:1-12. February 1,
1940.
Transmitted May 80, 1947.
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TOPEKA. KANSAS
1947
21-8188
S -NA-L.
EL L_ &- UJ"v*<^wC£ ) Kari,^
A New Bat (Genus Myotis) From Mexico
BY
WALTER W. DALQUEST and E. RAYMOND HALL
. ZOOL
LIBRARY
MAR -8 19'
University of jtansas Publications
Museum of Natural History
Volume 1, No. 12, pp. 237-244
December 10, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, H. H. Lane, Edward H, Taylor
Volume 1, No. 12, pp. 237-244
December 10, 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1947
22-1402
MUS. GO
LIB
zeoL
MAR -
8
50
A New Bat (Qenus Myotis) From Mexico*
I ■ By — '
WALTER W. DALQUEST AND E. RAYMOND HALL
While one of us (Dalquest) was in a dugout canoe that was being
paddled up a small unnamed tributary of the Rio Coatzacoalcos,
through dense jungle, he grasped a decayed and termite damaged
tree-trunk projecting approximately three feet above the surface of
the water to steady the canoe. At that instant two bats were de-
tected in one of the many small holes in the trunk, which was eight
to nine inches in diameter. It was a simple matter to enlarge the
hole and extract the animals. Superficially they resembled silvery-
haired bats (Lasionycteris) but their naked interfemoral membranes
and other features suggested that they belonged to the genus Myotis.
Subsequently, study in the laboratory showed this to be the fact and
revealed also that they are of an heretofore unnamed species which
may be known as :
Myotis argentatus, new species
Type. — Male, adult, skin with skull, No. 19228, Mus. Nat. Hist., Univ. Kan-
sas; 14 kilometers southwest of Coatzocoalcos, 100 feet elevation, Veracruz,
Mexico; 2 February 1947; obtained by Walter W. Dalquest; original No. 7052.
Range. — Known only from the type locality.
Diagnosis. — Size medium for the genus (see measurements), tail short; foot
long; ears and membranes black; pelage long (maximum length on middle of
back 9 mm.) and black; upper parts with overhairs tipped with whitish es-
pecially on rump; underparts from posterior part of thorax posteriorly with
all of the hairs tipped with this same whitish color; skull with preorbital part
small in relation to brain case; teeth small in relation to total area of palate;
brain case much inflated; ventral margin of foramen magnum evenly rounded.
Comparison. — From Myotis albescens (E. Geffroy) known to us by speci-
mens in the United States National Museum from Paraguay (Tacural), Pan-
ama (Tabernilla), and Nicaragua (Prinzapolca R. and Escondido R.), argen-
tatus differs in: Body and foot longer; tail relatively shorter (57 and 58% of
length of head and body versus 76 (62-83)% in albescens); tibia shorter;
pelage longer, and black instead of brown; silver tipping of fur on hinder
back markedly more conspicuous; precranial part of skull, when viewed from
above, larger in relation to brain case; postorbital constriction less abrupt,
that is to say, skull "longer-waisted" ; occlusal surfaces of teeth of equal area
and therefore occupying a relatively smaller percentage of total area of palatal
* Assistance with field work is acknowledged from the University of Kansas Endowment
Association.
(239)
240
University of Kansas Publs., Mus. Nat. Hist.
surface; ventral margin of foramen magnum less deeply indented; ventrally
prominent part of basioccipital twice as wide.
Remarks. — The relatively slight wear on the teeth of the female
of M. argentatus and the large ends on the bones of the wings indi-
cate that it is immature. Its measurements, recorded below, aver-
age smaller than those of the adult holotype, a male, and the silvery
tipping on the upper parts is almost lacking from the pelage which
is shorter than in the holotype.
Myotis.
From left to right, dorsal, lateral and ventral views.
All X 2. *
Figs. 1-3. Myotis argentatus, no. 19228, Univ. Kan. Mus. Nat. Hist., type.
Figs. 4-6. Myotis albescens, no. 105664, $ , U. S. Nat. Mus., from Tacuaral,
Paraguay; obtained on November 13, 1900, by Wm. T. Foster, orig. no. 128.
Among at least American kinds of Myotis, argentatus is extreme
in small area of occlusal surface of the upper molariform teeth in
relation to the total area of the palatal surface of the skull. M . al-
bescens previously was regarded as extreme in this feature. The
distance across the third upper molars, from the outside of one tooth
Dalquest and Hall — New Bat from Mexico 241
to the outside of the other, is 5.5 mm. in the holotype of argentatus
and 5.4 mm. in a specimen of corresponding age and sex of albescens.
The distance between the third upper molars, from the lingual side
of one tooth to the lingual side of the other, is 2.9 mm. in argentatus
and 2.8 mm. in albescens.
In each of our two specimens there is no sagittal crest but instead
a low ridge one millimeter wide which marks the space between the
margins of the two temporal muscles.
Allusion already has been made to the resemblance of the newly
named Myotis argentatus to the silvery-haired bat, Lasionycteris
noctivagans (LeConte). The whitish tips of the hairs are slightly
more yellowish in argentatus but the difference is so slight as to be
detected by only the most careful comparison. The remainder of
the pelage in argentatus is black as in the darkest individuals of
Lasionycteris.
Among named kinds of the genus Myotis, the species argentatus
most closely resembles Myotis albescens which, up to now has been
recorded from as far south as Argentina, in South America, and as
far north as Nicaragua, in Central America (Miller and Allen, Bull.
U. S. Nat. Mus., 144:202, 203, 1928). The differences detected be-
tween the two species are indicated above in the paragraph of com-
parisons and some other differences can be detected by comparing
measurements given below with those of M. albescens as recorded
by Miller and Allen (op. cit.: 204-205). In initial comparisons with
albescens, only Paraguayan specimens were employed. It was felt
that specimens of albescens from the northernmost localities of oc-
currence might more closely resemble argentatus. Accordingly, we
appealed a second time to Dr. A. R. Kellogg for comparative ma-
terial and he lent us the specimens (alcoholics with skulls separate)
in the U. S. National Museum from Central America. These also
differ from our newly named bat in the same fashion as do the
South American specimens. Further, the number and magnitude of
the differences between albescens and argentatus greatly exceed any
that can be pointed to between the American subspecies of any other
one full species of the genus Myotis. Full specific, rather than mere
subspecific, status, therefore, is suggested for the bat here named
Myotis argentatus.
Measurements. — The adult, male type, and the immature female specimen
measure, respectively, as follows: Head and body, 55, 51 mm.; tail, 32,
29; tibia, 13.7, 135; foot, 8, 9; forearm, 33.0, 34.5; thumb, 5.8, 5.7; third meta-
242 University of Kansas Publs., Mus. Nat. Hist.
carpal, 32.2, 30.5; fifth metacarpal, 31.5, 30.3; greatest length of skull, 14.5,
14.0; condylobasal length, 13.8, 13.0; zygomatic breadth, 9.1, 9.0; interorbital
constriction, 4.3, 4.0; breadth of brain case, 7.5, 7.4; occipital depth 5.7, 5.7;
mandible, 10.5, 10.0; maxillary tooth row, 5.3, 5.0; maxillary breadth at M3,
5.5, 5.7; mandibular tooth row, 5.6, 5.3.
Specimens examined. — Two, from the type locality.
Transmitted October 20, 1947.
D
«=7 -/vA-L
Tadarida femorosacca (Merriam)
in Tamaulipas, Mexico
BY
WALTER W. DALQUEST and E. RAYMOND HALL
'. ZOQL
LIBRARY
HAR -8 !■
University of Kansas Publications
Museum of Natural History
Volume 1, No. 13, pp. 245-248
December 10, 1947
UNIVERSITY OF KANSAS
LAWRENCE
1947
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, H. H. Lane, Edward H. Taylor
Volume 1, No. 13, pp. 245-248
December 10, 1947
University of Kansas
Lawrence, Kansas
PRINTED BY
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1947
22-1401
MUS. CO
LIBRARY
1AR -3 1950
Tadarida femorosacca (Merriam) In Tamaulipas,
Mexico
- ' Dj>' ■'
WALTER W. DALQUEST AND E. RAYMOND HALL
On January 23, 1946, two pocketed free-tailed bats {Tadarida
femorosacca, Catalogue nos. 17852 and 17853) were obtained in a
large cave 10 kilometers north-northeast of the village of Antiguo
Morelos, in the state of Tamaulipas, Mexico. This extends the
known range of this species to the Atlantic Slope and more than
300 miles to the northeast of Zacoalco, Jalisco, the only locality in
Fig. 1. Map showing localities of known occurrence of the pocketed free-tailed
bat (Tadarida femorosacca) .
central Mexico from which the species was previously known (see
Shamel, H. H., Proc. U. S. Nat. Mus., vol. 78, art. 19, p. 13, 1931).
The total length of the skull (18 mm.) and the basal length (15.0,
(247)
248 University of Kansas Publs., Mus. Nat. Hist.
15.2) are less than recorded by Shamel (op. cit.) for any one of the
eight specimens studied by him. Otherwise our two specimens an-
swer the description of feynorosacca. They were found lying on the
floor of the cave. One was dead and the other alive but incapable
of flight. Shooting into the cracks of the roof of the cave more
than a hundred feet high failed to dislodge other bats but stimu-
lated a volume of squeaking of bats which indicated that thousands
of individuals, possibly of this species, were ensconsed there. The
cave had long been used by bats as attested by the large deposit of
guano, much of which had been removed for fertilizer.
Transmitted October 20, 1947.
□
A New Pocket Gopher (Thomomys) and A New
Spiny Pocket Mouse (Liomys) from
Michoacan, Mexico
BY
E. RAYMOND HALL AND BERNARDO VILLA R.
«. . ZOOL
MAR -8 !•■ I
y _J
University of Kansas Publications
Museum of Natural History
Volume 1, No. 14, pp. 249-256, 6 figs, in text
July 26, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; A. Byron Leonard,
Edward H. Taylor
Volume 1, No. 14, pp. 249-256, 6 figs, in text
July 26, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3338
fMAR -8 1950
A New Pocket Gopher (Thomomys) and a New Spiny
Pocket Mouse (Liomys) from Michoacan, Mexico
E. RAYMOND HALL and BERNARDO VILLA R.
A series of 17 pocket gophers of the species Thomomys umbrinus
obtained in 1943 from points 3, 4 and 5 miles south of Patzcuaro
proves upon comparison to be an hitherto unrecognized subspecies
which is described and named as follows:
Thomomys umbrinus pullus, new subspecies
Tfype.— -Male, adult, skin and skull; No. 100151, Univ. California Mus. Vert.
Zool.; 5 mi. S Patzcuaro, 7800 ft., Michoacan, Mexico; March 10, 1943; ob-
tained by Hubert H. Hall, original No. 117.
Range. — Known only from 3 to 5 miles south of Patzcuaro, Michoacan.
Diagnosis. — Size small (see measurements) ; color black or between Cinna-
mon-Brown and Snuff Brown; distal half of tail whitish and all of tail whitish
in one specimen; lambdoidal crests perpendicular to sagittal plane of skull;
posteroventral face of tympanic bulla rugose; jugal vertical (flat surface not
oblique) ; interpteiygoid space truncate at apex with sides curved outward
(see figure).
Comparison. — From topotypes of Thomomys umbrinus supcrnus Nelson and
Goldman, pullus differs as follows: More individuals wholly black (except
distal half of tail); underparts lacking white; rostrum broader; braincase an-
Figs. 1-3. Three views of the skull of the type specimen of Thomomys
umbrinus pullus. X 1-
teriorly slightly more expanded dorsally; lambdoidal crests perpendicular to
sagittal plane rather than inclined posteromediad; interparietal broader, $ 5.7
(5.0-7.0) versus 4.5, and in $ 6.5 (5.6-7.1) rather than 4.8 (4.4-5.1); flattened
middle part of jugal vertical rather than oblique; in side view, mastoid and
paroccipital processes farther apart thus exposing larger surface of mastoidal
bulla; incisors, in both upper and lower jaws, slightly narrower; molariform
teeth smaller, interpteiygoid space truncate, at apex, with sides convex mediad,
rather than V-shaped ; ventral face of tympanic bullae rugose in posterior half
rather than smooth.
(251)
252 University of Kansas Publs., Mus. Nat. Hist.
Remarks. — Among named subspecies of Thomomys umbrinus,
T. u. pullus most closely resembles T. u. supernus, the subspecies
next adjacent to the northward. Therefore, the results of compari-
sons with only that subspecies are here reported upon. T. u. tolucae
to the eastward is for one thing a much larger animal and has
slightly less procumbent upper incisors. So far as we know, Tho-
momys umbrinus has not heretofore been reported from Michoacan.
Of our seventeen skins, eight are brown, six are black and two are
intermediate in color.
Most of these pocket gophers lived where there was a good growth
of pine trees in the same areas where large pocket gophers of the
species Cratogeomys gymnurus occurred. The field notes of the col-
lector of the type of T. u. pullus record that when he was making a
shallow excavation to reveal the gopher burrow in which he trapped
the holotype, he found the burrow approximately five inches below
the surface of the ground and that in digging deeper than was nec-
essary he accidentally broke into the burrow of a Cratogeomys.
Another member of our field party (E. R. Hall) when removing
from its burrow a trapped Thomomys that was caught only by the
hind leg, dug around the animal whose burrow was approximately
six inches underground and in doing so he also broke through the
roof of a burrow of Cratogeomys. The burrow of Cratogeomys was
approximately sixteen inches below the ground. Nowhere else, ex-
cept 3 to 5 miles south of Patzcuaro, have the authors found two
kinds of pocket gophers living together. The two-story arrangement
south of Patzcuaro was possible because of the different levels at
which the two kinds of animals made their burrows and the two-
story arrangement was accidental and exceptional rather than the
rule.
Measurements. — Average and extreme measurements of five adults of each
sex, are as follows: Total length, male 184 (178-198), female 185 (174-194);
length of tail, 54 (48-60), 53 (47-57); length of hind foot, 26.8 (25-29), 27.6
(26-29); weight, 86.1 (78.7-96.9), 74.3 (70.2-84.8) grams; basilar length, 30.2
(28.8-31.3), 28.6 (27.8-29.1); zygomatic breadth, 23.2 (22.3-24.6), 21.3 (20.8-21.8);
least interorbital breadth, 5.9 (5.8-6.1), 6.4 (6.0-6.8); mastoid breadth, 17.8
(17.1-18.7), 17.2 (16.6-17.5); length of nasals, 12.4 (11.8-13.0), 11.5 (11.0-12.5);
breadth of rostrum, 7.5 (6.9-8.2), 7.1 (6.9-7.3); length of rostrum, 14.1 (13.4-14.5),
13.3 (12.7-13.5); alveolar length of maxillary tooth-row, 7.0 (6.7-7.5), 6.9 (6.8-
7.0); palato-frontal depth, 13.2 (13.0-13.4), 12.9 (12.3-13.5).
Specimens examined. — Total, 17, all from 7800 ft., Michoacan, as follows:
3 mi: S Patzcuaro, 1; 4 mi. S Patzcuaro, 10; 5 mi. S Patzcuaro, 6.
Hall and Villa: A New Pocket Gopiiku
253
In 1943 a series of fifteen spiny pocket mice, Liomys irroratus,
was obtained within a radius of five miles of Patzcuaro and, mostly
on geographic considerations, the animals were assigned to Liomys
irroratus alleni (Coues). In fact, in his "Revision of the Spiny
Pocket Mice," Goldman (N. Amer. Fauna, 34:57, 1911) had thus
identified the one specimen available to him from Patzcuaro. Crit-
ical examination of the series, however, revealed cranial features not
described in the named kinds from adjoining geographic areas, and
comparisons showed that the animal from Patzcuaro differs sub-
specifically from any named kind. The new subspecies may be
known as:
Liomys irroratus acutus, new subspecies
Type.— Female, adult, skin and skull; No. 100171, Univ. California Mus.
Vert. Zool.; 2 mi. W. Patzcuaro, 7700 ft., Michoacan, Mexico; March 10, 1943;
obtained by E. R. Hall and J. R. Alcorn, original No. 3837 of Alcorn.
Range. — Known only from the vicinity of Patzcuaro, Michoacan.
Diagnosis. — Size large (see measurements) ; upper parts dark brown ; pos-
terior border of nasals V-shaped with apex directed anteriorly ; frontomaxillary
suture medially concave or rarely straight; interparietal subcircular; basisphe-
noid wide; tympanic bullae large.
Comparisons. — From Liomys irroratus alleni, acutus differs as follows: Color
slightly darker brown on upper parts; size slightly less; posterior border of
nasals V-shaped rather than truncate; frontomaxillary suture medially concave
Figs. 4-6. Three views of the skull of the type specimen of Liomys irroratus
acutus. X 1-
or straight instead of convex; interparietal subcircular (anterior border) rather
than triangular; basisphenoid broader; tympanic bullae larger and more in-
flated. From Liomys irroratus jaliscensis (topotypes), acutus differs as follows:
Color slightly darker brown on upper parts; size larger, without overlap, in
external measurements and in basilar length, length of nasals and mastoid
breadth; posterior border of nasals V-shaped rather than almost truncate;
frontomaxillary suture medially concave or straight rather than convex; inter-
parietal subcircular rather than quadrilateral; basisphenoid wider; tympanic
bullae larger. From Liomys irroratus pullus, acutus differs in longer body,
254 University of Kansas Publs., Mus. Nat. Hist.
shorter tail, slightly longer hind foot; all of upper parts, and especially upper
side of tail, more brownish and less blackish; posterior border of nasals and
frontomaxillary suture differing in same way as from alleni; interorbital region
narrower in relation to length of skull; over-all length of skull greater; inter-
parietal anteroposteriorly longer; tympanic bullae more inflated.
Remarks. — This relatively large, dark-colored, spiny pocket mouse
of east-central Michoacan differs from its geographic neighbors in
V-shape of posterior border of nasals, semicircular shape of inter-
parietal, medially concave maxillofrontal suture, wide basisphcnoid
and larger tympanic bullae. The latter character is not constant.
Intergradation with L. i. alleni is shown by specimens from Queren-
daro in which the shape of the interparietal is exactly intermediate
between those of topotypes of the two subspecies and also in that
the basisphenoid is wider than in acutus but narrower than in alleni.
Intergradation with L. i. jaliscensis is shown, by specimen No.
120275 (U. S. N. M.) from Zamora, in shape of posterior end of
nasals, direction of maxillofrontal suture, and shape of interparietal.
In each of these features the specimen from Zamora is almost ex-
actly intermediate between acutus and jaliscensis. In large size of
tympanic bullae and wider basisphenoid the specimen agrees with
acutus, but otherwise is nearly as small as jaliscensis to which it is
here referred. Actually the specimen could, with almost equal pro-
priety, be referred to either subspecies.
Measurements. — The measurements of two males, Nos. 100184, 100182, and
average and extreme measurements of five females, are, respectively, as fol-
lows: Total length, 257, 267, 244 (230-251); length of tail, 130, 128, 122 (105-
129); length of hind foot, 32, 31, 31 (30-33); length of ear from notch, 16, 17,
15.3 (13.0-19); weight in grams, 71.5, 65.1, 50.8 (44.8-61.8); greatest length of
skull, 35.2, 34.9, 33.6 (32.7-34.2); zygomatic breadth, 17.7, 17.5, 16.5 (16.1-17.1);
interorbital breadth, 8.4, 8.1, 7.8 (7.5-8.0); length of nasals, 15.1, 14.9, 14.0
(13.3-14.5); width of braincase, 15.9, 15.1, 15.0 (14.7-15.1); alveolar length of
upper molariform tooth-row, 6.0, 6.0, 5.6 (5.5-5.9). The measurements were'
taken according to the method of Goldman (N. Amer. Fauna, 34:10, 1911).
Each of the specimens of which measurements are given above is adult; the
transverse enamel fold has been obliterated in Ml, is represented by only an
isolated lake in M2 (except in one female where all trace of the fold has worn
away) and is present in M3.
Specimens examined. — Total, 16, all from Michoacan, Mexico, and unless
otherwise indicated in the University of California Museum of Vertebrate
Zoology, as follows: 3 mi. NW Patzcuaro, 6700 ft., 1; 2 mi. W Patzcuaro,
7700 ft., 5; 2 mi. W Patzcuaro, 6700 ft,, 2; Patzcuaro, 1 (U. S. Nat. Mus.);
5 mi. S Patzcuaro, 7800 ft., 7.
For the loan of comparative materials we are grateful to Dr.
Harold E. Anthony of the American Museum of Natural History,
Hall and Villa: A New Pocket Gopher 255
Mr. Stanley P. Young and Dr. Hartley H. T. Jackson of the Bio-
logical Surveys Collection in the United States National Museum,
Dr. Charles P. Lyman of the Museum of Comparative Zoology, and
for assistance with the field work to the John Simon Guggenheim
Memorial Foundation and to Miss Annie M. Alexander.
Transmitted April 1, 1948.
□
22-3338
S-A/A-L
A New Hylid Frog from Eastern Mexico
BY
EDWARD H. TAYLOR
MUS.
ZOOL
!AR -8 19! )
University of Kansas Publications
Museum of Natural History
Volume 1, No. 15, pp. 257-264, 1 fig. in text
August 16, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor
Volume 1, No. 15, pp. 257-264, 1 fig. in text
August 16, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3339
P. IML
1AR -8 1950
A New Hyiid Frog frorm Eastern Mexico
By
EDWARD H. TAYLOR
A small collection of Mexican reptiles and amphibians recently-
acquired by the University of Kansas Natural History Museum
contains five specimens of a species of the genus Hyla {sensu lato)
which is here described as new.
Hyla proboscidea sp. nov.
Type. — University of Kansas Museum of Natural History, No. 23626, col-
lected 2 km. west of Jico, Veracruz, Mexico, at an elevation of 4,200 ft., Oct.
28, 1946, by Walter W. Dalquest.
Paratypes.— Nos. 23624, 23625, 23627, 23628, collected with the type.
Diagnosis. — A medium sized member of the genus with known maximum
length of male, 57 mm. Canthus rostralis well defined; tip of snout with a
bulbous projection; fingers more than one-third webbed, foot nearly com-
pletely webbed; tympanum distinct; skin smooth above, granular below; very
prominent inner metatarsal tubercle, small outer tubercle; tibiotarsal articu-
lation reaches to nostril; a well-defined outer tarsal fold; anal opening ventral,
covered by a free triangular flap; pupil of eye horizontal.
Description of the type. — Head longer than broad, the distance between the
eye and nostril slightly greater than distance between nostril and tip of snout;
canthus rostralis sharply defined, continued to above nostril; upper part of
loreal region sloping abruptly, lower part sloping more gently to edge of lip;
area in front of nostril somewhat swollen, the nostril large, directed strongly
backward; tip of snout forming a short rounded proboscis; upper jaw rather
strongly overhanging lower jaw.
Width of an upper eyelid contained in interorbital distance about 1% times;
horizontal diameter of eye about equal to distance between eye and nostril,
about 1% times diameter of tympanum; tympanum distinct, its distance from
orbit equal to its diameter, overhung by a glandular fold running back from
eye.
Choanae large; vomerine teeth in two elevated patches which lie between,
and reach the posterior level, of choanae, the patches closer to each other than
to choanae ; tongue rather small, subcircular, not or but very indistinctly
notched behind, not at all free behind; opening to vocal sacs behind level of
tongue, the openings a short slit directed backwards. (Vocal sacs not evident
externally in type or paratypes.)
Skin of dorsal surfaces generally smooth (under magnification surface mi-
nutely corrugated and wrinkled) ; ventral surface of abdomen, the thighs, and
lower part of lateral surface of body strongly granulate, the granules unequal in
size and elevation ; breast, chin, and under side of arm with sparse granules or
tubercles.
Anal opening ventral, covered by a small, free, triangular flap; a small
thickened fold, slightly free, on each side of anus partly covered by triangular
flap.
(259)
260
University of Kansas Publs., Mus. Nat. Hist.
Arms rather short, upper arm slender, forearm much thickened; a small
axillary web present; disks on three outer fingers distinctly larger than tym-
panum, of first finger equal to or somewhat smaller than tympanum; outer
fingers, between one-third and one-half webbed; on inner fingers webbing less
than one-third; first finger more or less opposed to other three, its base widened,
Measurements (in mm.),
width, 1S.6.
-Snout to vent, 58; leg, 86; head length, 20; head
and the upper surface covered by a large patch of minute dark, horn-colored
nuptial asperities, that extend to near the terminal disk; subarticular tubercles
strongly elevated with numerous supernumerary tubercles on palm; a some-
what enlarged elevated palmar tubercle; under surface of forearm with a row
of distinct tubercles; other smaller scattered granules present. Toes more
than four-fifths webbed, the membrane reaching the base of the terminal disks,
on one side at least, of all toes save fourth; subarticular tubercles strongly
Taylor: A New Hylid Frog
261
elevated, with numerous supernumerary tubercles on sole; a large elevated
inner metatarsal tubercle; a small outer tubercle; a continuous, well-defined,
tarsal fold extending entire length of tarsus. Tibiotarsal articulation reaches
nostril when leg is brought forward.
Color and marking. — General color of type (preserved in formalin, then
transferred to alcohol) dull grayish purple, darker anteriorly and somewhat
lighter and more mottled posteriorly; color very much lighter on sides with
a few cream, dark-edged spots in groin and on sides of abdomen; front and
back surfaces of thigh and shank with some darker and lighter flecking that is
continued more or less on the foot. When submerged in water, very dim dark
spots or bars visible on limbs; ventral surface dirty brownish flesh, without
markings.
Variation. — The series consists of five adult male specimens. It is presumed
that the female is considerably larger, and may lack the nasal proboscis which
I suspect is a secondary sexual character.
There are some differences in the shade of coloring in the preserved speci-
mens, some being darker, some lighter than the type. In two the lateral dark-
edged, cream spots extend to the axilla, and the light and dark markings on
the front and back surfaces of the leg are much more distinct in most of the
specimens than in the type. When the specimens are submerged in water, the
black bars on the limbs are evident in all specimens. The tympanum is
sometimes darker, sometimes lighter than its surroundings.
In the field notes of Mr. Dalquest I find the statement that the color in
life is bright yellow, which presumably applies to all of the specimens. No
trace of this color remains at the present time.
The ventral granules of some of the paratypes are very unequal in elevation,
some being elongated, nipplelike.
The following table gives the variation in measurements of the type and
paratypes :
Measurements of Hyla proboscidea in mm.
Snout
Head
Head
Foot and
Eye
to
vent
length
width
Arm
Leg
longest
toe
diam-
eter
23626
58
20
18.6
29
86
39
5
56
19
18
29
81
37
5
23627
53
18.5
17
30
81
35
5
23624
50
16.8
15.5
26
70
29.8
5
23628
50
17
16
28
71
30
5
Relationship. — It would appear that the relationship of this species
is with Hyla bistincta, a widespread Mexican species likewise oc-
curring in the same general area but at a much higher elevation. It
also has an elongated flap which carries the anal opening nearly to
the level of venter but the terminal part lacks the triangular flap.
262 University of Kansas Publs., Mus. Nat. Hist.
There is no prolongation of the snout tip. There are also numerous
other differences, so that it would be difficult to confuse the two
forms.
Remarks. — This adds another very distinctive species to the Vera-
crucian fauna. Despite the fact that this state has probably been
explored at greater length than any other Mexican state it still is a
likely place for the discovery of novelties.
Transmitted April 8, 1948.
□
22-3339
A New Extinct Emydid Turtle from the Lower
Pliocene of Oklahoma
BY
EDWIN C. GALBREATH
University of Kansas Publications
Museum of Natural History
Volume 1, No. 16, pp. 265-280, plate 1
August 16, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor
Volume 1, No. 16, pp. 265-280, plate 1
August 16, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3340
J '* ':-^ * '-^i/i
f
J5^^ i
Plate 1. Chrysemys limnodytes. Univ. Kans. Mus. Nat. Hist., Vert. Paleo.
Coll. No. 7676. Fig. A, dorsal view of carapace; fig. B, ventral view of plastron.
Both views approximately X -4.
riAR -8 \r" ]
A New Extinct Emydid Turtle from the Lower Pliocene
of Oklahoma
EDWIN C. GALBREATH
In the summer of 1946 a party from the University of Kansas
Museum of Natural History visited exposures of the Laverne forma-
tion in Beaver County, Oklahoma, at the invitation of Dr. Stuart
Schoff of the United States Geological Survey. When examining
the marl beds an Emydid turtle was discovered which appears to
be an unnamed species of the genus Chrysemys. A description of
the new species follows.
Chrysemys limnodytes, new species
Holotype. — University of Kansas Museum of Natural History No. 7676,
vertebrate paleontological collection, a turtle consisting of a fragmental an-
terior portion of a carapace, left part of the plastron, and several marginals
collected by the 1946 paleontological field party of the University of Kansas
Museum of Natural History.
Geological Age and locality. — Marl beds of the Laverne formation, early
Pliocene age, in SW % Sec. 15, T. 4 N., R. 25 ECM, Beaver County, Okla-
homa. The specimen was removed from the marl immediately below the fossil
leaf zone (see Chaney and Eiias, 1936; Frye and Hibbard, 1940).
Diagnosis. — Size large (see measurements) and differing from other species
of Chrysemys in having: The anterior end of the carapace broadly concave,
the posterolateral marginals not greatly flared, the posterior end of the plastron
broadly indented, the carapace more sculptured and relatively wider.
Description of type. — The specimen had been badly damaged before preser-
vation, and had suffered further damage from exposure before discovery. The
anterior and posterior lobes of the plastron had been folded over the bridge,
forming a three-ply thickness of bone. Of the carapace, only the following
parts are known: Fragment of the nuchal; right 1st, 7th, 8th, and 9th mar-
ginals, left 1st, 2d, 7th, 8th, 9th, and 11th marginals; costals 1-5 on the right
side; costals 1-4 on the left side; and 1st, 2d, 3d, and 4th neurals. The left
half of the plastron is relatively complete, lacking only the epiplastron and
entoplastron. The left 7th, 8th, and 9th marginals are joined to the plastron
at the inguinal buttress, and the right 7th, 8th, and 9th marginals are attached
to the fifth costal. The carapace has smooth contours with no keel present,
but on the lower half of the costals there are seven or eight ridges, and the
remaining surface of the costals and neurals are rugose. The marginals lack
ridges, and the posterolateral marginals are not serrated. The anterior end of
the carapace has a broad shallow notch. The first neural is rounded, and the
2d, 3d, and 4th are hexagonal, with the broad ends forward. Anterior margins
of the 2d and 3d neurals are concave, and the anterior margin of the 4th neural
is straight.
(269)
270 University of Kansas Publs., Mus. Nat. Hist.
The sulci bounding the scutes are moderately impressed. The width of the
first vertebral scute, anteriorly and posteriorly, is less than the width of the
second vertebral scute. The costal scutes join the marginal scutes on the
marginal plates.
The plastron, broadly indented at the posterior end, does not have the
posterior lobe flared laterally as it is in Recent species of Chrysemys, and lacks
any pronounced notch at the femoro-anal suture. The humero-pectoral sulcus
crosses the plastron behind the entoplastron in a straight line, and reaches the
border anterior to the axillary notch. The pectoro-abdominal suture is an-
teriorly convex at the sides and concave at the midline.
A comparison of this carapace and plastron with a series of specimens of
Recent Chrysemys picta and Pseudemys scripta of approximately the same
size reveals characters indicated in the following chart:
Galbreath: A New Emydid Turtle
271
RECENT Chrysemys
Chrysemys limnodytes
RECENT Pseudemys
Not serrated.
Posterolateral marginals
not serrated.
Serrated.
Not notched.
Carapace with broad shal-
low notch at anterior
end.
Notched.
Occasional faint notch
at femoro-anal suture.
Plastron does not have a
pronounced notch at
femoro-anal suture.
Distinct notch at femoro-
anal suture.
Posterior lobe of plas-
tron flares laterally.
Posterior lobe of plastron
does not flare laterally.
Posterior lobe of plastron
does not flare laterally.
Carapace smooth.
Carapace has smooth
contours.
Carapace has depressions
and elevations.
Old specimens occasion-
ally have five or six
ridges near border of
costals.
Seven or eight ridges on
lower half of costals.
Remaining surface of cos-
tals and neurals rugose.
Ridges cover costals.
No ridges on marginals.
No ridges on marginals.
May or may not have
ridges on marginals.
Nuchal smooth.
Nuchal smooth.
Nuchal has ridges.
Carapace not greatly
arched.
Probably arched less than
in Psexidemys, but more
than in any Recent
Chrysemys.
Greatly arched.
Keel often present at
birth, but soon lost.
No keel present.
Keel often present.
Anterior and posterior
widths of first verte-
bral scute approxi-
mately same as width
of second vertebral
scute.
Anterior and posterior
widths of first vertebral
scute less than width
of second vertebral
scute.
Anterior width of first
vertebral scute less than
posterior width, or both
dimensions less than
width of second verte-
bral scute.
Ribs do not tend to be
prominent on costals.
Ribs not prominent on
costals.
Ribs tend to be prominent
on costals.
272 University of Kansas Publs., Mus. Nat. Hist.
Hay attached considerable taxonomic importance to the characters of the
nuchal and I find its characters to be fairly constant in the specimens of
Emydidae examined. Although the nuchal of Chrysemys limnodytes is in-
complete, it can be distinguished from the nuchals described by Hay as types
of his several fossil Emydids. Differences in the nuchal, together with those
in the carapace and plastron, serve to distinguish the species from other genera
of the Emydidae.
When the specimen is compared with Chrysemys timida Hay, of the Ne-
braska Pleistocene, many similarities, mostly of generic rank, are seen.
Chrysemys limnodytes is broader in relation to length than is either C. timida
or any Recent specimen examined of the same size. The greatest allowance
possible in estimating the length of C. limnodytes fails to bring the ratio of
its breadth to length within the range of Recent specimens of similar size.
Data from 96 specimens of Recent Chrysemys picta show that the ratio of
length to width is not affected by sex, but that the ratio does vary with the
age of the specimen. In young animals the length and width are approximately
equal, but with further growth the length becomes relatively greater. Speci-
mens in the length group of 135 to 144 mm. have the widths ranging from 71
to 81 per cent of the lengths. In all specimens larger than this, the ratio is in
the low seventies, and the largest specimen, 177 mm. in length, has the width
of the carapace amounting to only 74 per cent of the length. The fossil species,
C. timida, with a length of 160 mm., has the width amounting to 75 per cent of
the length, and C. limnodytes, with an estimated length of 180 mm., has the
width amounting to 80.5 per cent of the length. C. timida is widest anteriorly,
whereas C. limnodytes and the other species of the genus are widest posteriorly.
Less obvious differences between the two fossils are the narrower anterior mar-
gin of the nuchal, the concave anterior end of the carapace, the sculptured
surface of the carapace, and the relatively wider neurals and longer vertebrals
of C. limnodytes.
DIMENSIONS OF THE TYPE SPECIMEN
(In millimeters)
Total length of carapace, 180 (estimated) ; greatest width of carapace, 145
(estimated) ; height of carapace, more than 50.
Length of plastron, 165 (estimated) ; width of plastron, 130 (estimated) ;
length of anterior lobe, 45 (estimated) ; width of anterior lobe, 75 (estimated) ;
length of posterior lobe, 62; width of posterior lobe, 82; length of bridge from
axillary to inguinal notch, 60.
Plates of the Carapace and Plastron
Nuchal: Width of anterior margin, 12 (estimated); greatest width, 37 (esti-
mated); length at midline, 35 (estimated).
First neural: Greatest width, 13; length at midline, 17. Second neural:
Greatest width, 16; length at midline, 14. Third neural: Greatest width, 18;
length at midline, 16. Fourth neural: Greatest width, 18; length at mid-
line, 16.
Costals: Thickness at proximal end, 3-5; thickness at distal end, 2. First
costal: Length of margin bordering nuchal, 23 (this and the following measure-
ments of the costal and marginal plates are of plates from the right side of the
animal except those indicated by an "L") ; length of margin bordering neurals,
17; length of margin bordering marginals, 38; length of margin bordering 2d
costal, 51. Second costal: Length of margin bordering 1st costal, 53; length
Galbreath: A New Em void Turtle 273
of margin bordering neurals, 16; length of margin bordering marginals, 25;
length of margin bordering 3d costal, 56. Third costal : Length of margin
bordering 2d costal, 55 (L), 56; length of margin bordering neurals, 19; length
of margin bordering marginals, 18; length of margin bordering 4th costal, 58.
Fourth costal: Length of margin bordering 3d costal, 58; length of margin
bordering neurals, 16; length of margin bordering marginals, 22; length of
margin bordering 5th costal, 55. Fifth costal: Length of margin bordering
4th costal, 52; length of margin bordering neurals, 16; length of margin bor-
dering marginals, 20; length of margin bordering 6th costal, 41.
First marginal: Length of margin bordering nuchal, 23 (L), 21; length of
outer margin, 23 (L), 23; length of inner margin, 12 (L), 12; length of margin
bordering 2d marginal, 22 (L), 21. Second marginal: Length of margin bor-
dering 1st marginal, 22 (L) ; length of outer margin, 22 (L) ; length of inner
margin, 15 (L) ; length of margin bordering 3d marginal, 15 (L). Seventh
marginal: Length of margin bordering 6th marginal, 18 (L), 17 (estimated);
length of outer margin, 25 (L), 23 (estimated) ; length of inner margin, 18 (L),
18; length of margin bordering 8th marginal, 22 (L), 22. Eighth marginal:
Length of margin bordering 7th marginal, 22 (L), 23; length of outer margin,
22 (estimate of L), 22; length of inner margin, 18 (L), 18; length of margin
bordering 9th marginal, 24 (estimate of L), 23. Ninth marginal: Length of
margin bordering 8th marginal, 24 (L) ; length of outer margin, 20 (L), 19;
length of inner margin, 19 (L) ; length of margin bordering 10th marginal,
23 (L), 23. Eleventh marginal: Length of margin bordering 10th marginal,
22 (L) ; length of outer margin, 16 (L) ; length of inner margin, 12 (L) ; length
of margin bordering pygal, 18 (L).
Entoplastron : Width, 24 (estimated).
Hyoplastron: Length of margin bordering epiplastron, 25; length of margin
on midline, 37; length from junction of epiplastronal border and outer border
to point on posterior border equidistant from midline, 53; width from midline
to axillary notch, 39; distance between axillary notch and posterior border, 31.
Hypoplastron : Length of margin bordering midline, 42; length of posterior
(xiphiplastronal) margin, 40; distance from junction of xiphiplastronal margin
and outer margin to point on anterior border equidistant from midline, 49
(estimated) ; distance between inguinal notch and anterior border, 29.
Xiphiplastron : Length of anterior (hypoplastronal) margin, 38; length of
margin along midline, 43; distance from extreme posterior extension of xiphi-
plastron to midline, 14.
Scutes of Carapace and Plastron
First marginal scute: Length of margin bordering 2d marginal, 15 (L), 14;
length of anterior margin, 15 (L) ; length of posterior margin, 14 (L); length
of inner margin. 13 (L) ; length of outer margin, 23 (L). Third marginal
scute: Length of anterior margin, 14 (L). Eighth marginal scute: Length of
anterior margin, 15 (L), 15; length of posterior margin, 16 (estimate of L),
16; length of inner margin, 20 (L), 20; length of outer margin, 25 (estimate
of L), 25. Ninth marginal scute: Length of anterior margin, 17 (L), 16;
length of posterior margin, 17 (L), 17; length of inner margin, 18 (L), 20;
length of outer margin, 21 (L), 21. Tenth marginal scute: Length of anterior
margin, 17 (L), 17. Eleventh marginal scute: Length of posterior margin,
14 (L).
First costal scute : Length of margin bordering vertebrals, 45. Second costal
scute: Length of margin bordering vertebrals, 35 (L), 35; length of margin
bordering 3d costal scute, 52.
First vertebral scute: Length of anterior margin, 24 (estimated); greatest
width, 32 (estimated); length at midline, 35 (estimated). Second vertebral
scute: Length of anterior margin, 27; greatest width, 42; length at midline, 29.
Third vertebral scute: Length of anterior margin, 33; greatest width, 42;
length at midline, 40 (estimated).
Pectoral scute : Length of humero-pectoral sulcus from midline to outer
border, 38; length of margin of pectoral scute on midline, 18; distance between
274 University of Kansas Publs., Mus. Nat. Hist.
junction of humero-pectoral sulcus and outer border and point on pectoro-
abdominal sulcus equidistant from midline, 19; distance from axillary notch to
point on pectoro-abdominal sulcus equidistant from midline, 17.
Abdominal scute: Length of margin of scute on midline, 43; width of
posterior border of abdominal scute from midline to inguinal notch, 41; dis-
tance from inguinal notch to a point on pectoro-abdominal sulcus equidistant
from midline, 44.
Femoral scute: Length of border of scute on midline, 24; width of anterior
border of scute from midline to inguinal notch, 41 ; width of posterior border
of scute from midline to outer border (along sulcus), 40; length of outer margin
of scute from inguinal notch to femoro-anal sulcus, 46.
Anal scute: Length of margin at midline, 36; length of femoro-anal
sulcus, 40.
Remarks. — Noteworthy is the intermediate nature of C. limno-
dytes when compared with species of the genera Chrysemys and
Pseudemys. However, any resemblance to Pseudemys is not to be
considered as evidence that C. limnodyt.es is in any way ancestral to
the genus Pseudemys. The fossil specimens of Pseudemys from the
Pliocene are too poorly known to allow the student certainly to
place them in their correct systematic positions. The fossil Emydids
from Western Europe, listed as species of Chrysemys, differ very
much from this species, or belong to other genera of the family.
Only a few turtles are known from the Laverne formation. Hesse
(Chaney and Elias, 1938) reported a small Testudo from the La-
verne of Beaver County, Oklahoma, but neglected to state whether
it was among the material borrowed by him from the University of
Kansas Museum of Natural History. The Museum has an incom-
plete carapace and plastron (No. 3101) of a small Testudo from
that locality and formation. In Harper County, Oklahoma, the
field party from the University recovered a large number of frag-
ments of a large Testudo. Although this specimen is as yet unpre-
pared, enough fragments have been pieced together to reveal that
the tibia is 127 mm. long. This dimension and those of some of the
fragments indicate that the animal may have been four to five feet
long.
Mrs. Bernita Mansfield of the Geology Department, University
of Kansas, prepared the plate.
Galbreath: A New Emydid Turtle 275
LITERATURE CITED
Bergouxioux, Frederic-Marie.
1935. Contribution a l'etude paleontoligique des cheloniens: Cheloniens
fossiles du Bassin d'Aquitaine. Memoires de la Societe geologique de
France, vol. 11, Mem. 25, pp. 1-215, 44 figs., 16 pis.
1937. Relations fauniques entre des chelonien fossiles de l'Espagne et de la
France. Comptes Rendus Acad. Sci. Paris, vol. 204, pp. 793-795.
1938. Cheloniens fossiles d'Espagne. Bulletin de la Societe d'histoire
Naturelle de Toulouse, vol. 72, pp. 257-288, 7 figs.
Chaney, R. W., and Elias, M. K.
1936. Late Tertiary Floras from the High Plains, with a Chapter on the
Lower Pliocene Vertebrate Fossils from the Ogallala Formation
(Lavern Zone) of Beaver County, Oklahoma, by Curtis J. Hesse.
Publ. Carnegie Inst. Wash., No. 476, pp. 1-72, 11 figs., 7 pis.
Frye, J. C, and Hibbard, C. W.
1941. Pliocene and Pleistocene Stratigraphy and Paleontology of the Meade
Basin, Southwestern Kansas. University of Kansas Publications, State
Geological Survey of Kansas, Bulletin 38, pp. 389-424, 3 figs., 4 pis.
Hay, 0. P.
190S. The Fossil Turtles of North America. Publ. Carnegie Inst. Wash., No.
75, pp. i-iv, 1-568, 704 figs., 113 pis.
Transmitted March 1, 1948.
□
22-3340
S-WA-L
t L*.
LO^v-4*>
Pliocene and Pleistocene Records of Fossil
Turtles from Western Kansas and
Oklahoma
BY
EDWIN C. GALBREATH
m. ' zool
LIBRARY
frlAR -3 13! 3
University of Kansas Publications
Museum of Natural History
Volume 1, No. 17, pp. 281-284
August 16, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor
Volume 1, No. 17, pp. 281-284
August 16, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3341
ZOCL
i v
HAR -8 1-1
Pliocene and Pleisjtocene Records] of Fossil Turtles from
Western Kansas and! Oklahoma
By
EDWIN C. GALBREATH
In the vertebrate paleontological collection at the University of
Kansas Museum of Natural History there are many fragments of
turtles that have been collected, generally in connection with the
excavation or recovery of other fossils. The generic identification
of this material is possible in many instances, and such identifica-
tions give new and important geological and distributional records
for genera in existence today.
All catalogue numbers refer to the vertebrate paleontological col-
lection in the University of Kansas Museum of Natural History.
Family KINOSTERNIDAE
Kinosternon sp. No. 7729 consists of fragments of marginals,
costals, and plastronal elements collected from Edson Quarry, Sher-
man County, Kansas. The age is middle Pliocene (Hemphillian).
No. 7679, consisting of a nuchal, and fragments of marginals, costals,
and neurals is from Nye Sink, XI Ranch, Meade County, Kansas,
and is of Pleistocene age.
Any fossil record of Kinosternon is a welcome find, and these two
specimens give new data both as to age and distribution. However,
it should be emphasized that these identifications are based on frag-
ments, and are tentative.
Family EMYDIDAE
Pseudemys sp. No. 5613, Sherman County; 6784, Seward
County; and 4728, Meade County, are three of many fragments,
mainly elements of the nuchal plate and plastron, that were collected
from Edson Quarry, Sherman County, Kansas, and from middle and
late Pliocene beds in Seward and Meade counties, Kansas, respec-
tively. The species represented cannot be differentiated from species
of Pseudemys living today. Species of Pseudemys are common also
in most of the Pleistocene deposits of western Kansas.
Family TESTUDINIDAE
Testudo sp. In general, two recognizable lines of the genus
Testudo existed in western Kansas during early Pliocene to mid-
Pleistocene time— a line of large testudinates with a carapace three
(283)
284 University of Kansas Publs., Mus. Nat. Hist.
to four feet long, and one line of smaller tortoises with a rugose
carapace approximately six to nine inches in diameter. Hibbard's
Testudo riggsi (Hibbard, 1944) is the best known of these smaller
turtles.
Gopherus sp. No. 5935 is from the Pleistocene of Harper County,
Kansas, and No. 7677 is from the early Pleistocene of Beaver
County, Oklahoma. Each is composed of costals and neurals which
may be referred to this genus. Their size indicates an animal con-
siderably larger than any on record, and probably these specimens
represent a new species.
Family CHELYDRIDAE
Chelydra sp. No. 6821 is the anterior part of a plastron from
the Edson Quarry, Sherman County, Kansas. The age is middle
Pliocene. This fragment has been listed three times in print, once
merely as Chelonia (having reference to the order) (Adams and
Martin, 1931), and twice by Hibbard (1934 and 1939) as Chelonia
sp. Hibbard's listing was obviously a lapsus calami for Chelydra
since he placed the genus in the family Chelydridae.
No. 6479 is part of a costal and marginals from Meade County,
Kansas. This specimen was associated with Aenocyon dims, and
is Pleistocene in age.
Family TRIONYCHIDAE
Amyda sp. No. 6800 is part of a costal collected in Seward
County, Kansas. It is possibly of early Pleistocene age. No. 7568
is part of a carapace from Meade County, Kansas, probably of the
same age.
LITERATURE CITED
Adams, L. A., and Martin, H. T.
1931. An addition to the Urodele Fauna of Kansas from the lower Pliocene.
Univ. Kansas Sci. Bull., 19:2S9-297, pis. 30-32.
Hibbard, C. W.
1934. Two new genera of Felidae from the middle Pliocene of Kansas.
Trans. Kansas Acad. Sci., 37:239-255, pis. 4-6.
1939. Notes on additional fauna of Edson Quarry of the middle Pliocene of
Kansas. Trans. Kansas Acad. Sci., 42:457-462, 6 figs.
1944. A new land Tortoise, Testudo riggsi, from the middle Pliocene of
Seward County, Kansas. Univ. Kansas Sci. Bull., 30:71-76, 2 figs.
Transmitted, March 8, 194S.
□
22-3341
A New Species of Heteromyid Rodent from the
Middle Oligocene of Northeast Colorado
with Remarks on the Skull
BY
EDWIN C. GALBREATH
m%. CBKP. ZOGL
LIBRARY
MAR -8 19
HARVAR8
University of Kansas Publications
Museum of Natural History
Volume 1, No. 18, pp. 285-300, 2 plates
August 16, 1948
University of Kansas
LAWBENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman; H. H. Lane, Edward H. Taylor
Volume 1, No. 18, pp. 285-300, 2 plates
August 16, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3312
Plate 2. Heliscomys tenuiceps. Univ. Kans. Mus. Nat. Hist., Vert. Paleo.
Coll. No. 7702. A., dorsal view; B, lateral view; C, ventral view. All views
approximately X 5.
SRal ^%
N MAX
A
. .--*>
Plate 3. Heliscomys tenuiceps. Univ. Kans. Mus. Nat. Hist.. Vert. Paleo.
Coll. Xo. 7702. A, lateral view of ri^ht side of skull showing structures in
orbital area. Al.S. alisphenoid. FR, frontal. MAX. maxillary. OS., orbito-
sphenoid. PAL, palatine. PC, presphenoid canal. SF. sphenoidal fissure.
SFr, sphenofrontal foramen. SPal, sphenopalatine foramen. Approximately X
9.3; B, occlusal view of P4-M3. Approximately X 23.4.
30L
HAR -8 19: )
A New Species of Jleteromyid Rcxlonl from the Middle
Oligocene of Northeast Ottrorido with Remarks
on the Skull
By
EDWIN C. GALBREATH
Heretofore our knowledge of the osteology of Heliscomys Cope
has been extremely limited; this genus previously was known by its
teeth, fragmental maxillaries, incomplete palatine bone and mandi-
ble, and part of one forelimb. In the summer of 1946 the writer, as
a member of the University of Kansas Museum of Natural History
field party, discovered the anterior part of a skull of Heliscomys in
the middle Oligocene deposits of Logan County, Colorado. This
specimen, representing a new species, yields a welcome, and greatly
desired addition to our fund of information about the genus.
The writer is indebted to Dr. Robert W. Wilson for advice and
helpful criticism in the course of this study, and to Mr. Bryan Pat-
terson of the Chicago Natural History Museum for the loan of com-
parative material. Mrs. Bernita Mansfield of the Geology Depart-
ment, University of Kansas, prepared the plates.
Family HETEROMYIDAE
Heliscomys tenuiceps, new species
Ho lo type.— Anterior part of a skull with left P4-M3, No. 7702, Vertebrate
Paleontological Collection, Museum of Natural History, University of Kansas.
Geological Age and locality. — Silts of Orellan age in the Cedar Creek facies
of the Brule formation in "Chimney Canyon," Sec. 3, T. 11 N, R. 54 W, Logan
County, Colorado.
Diagnosis. — Size larger than any known species; P4 with posteroexternal
cusp (metacone) anterior to central (hypocone) and lingual (entostyle) cusps,
which are connected by a cingulum; internal cingula of molars undivided, and
as high as paracone and metacone; style of each cingulum opposite the straight
median valley; rostrum deep and laterally compressed.
Description. — The type consists of the preorbital and interorbital parts of a
skull. Its size is comparable to that of the Recent heteromyid, Liomys pictus
Merriam. L. pictus is the species referred to in the comparisons below when
only the generic name Liomys is mentioned. Both incisors have been broken
off. The right tooth-row is missing, but the left row is complete, and its
orientation indicates that the tooth rows were parallel. The zygomata are
broken off close to the rostrum, which is relatively narrow in comparison with
its length and depth. In this narrowness, the specimen resembles Florentiamys
Wood more than it does such Recent heteromyids as Liomys or Heleromys,
where the rostrum is much wider at the dorsal surface than at the ventral
(289)
290 University of Kansas Publs., Mus. Nat. Hist.
surface (correlating with the wide interorbital dimension). In No. 7702 the
rostrum is not appreciably expanded on the dorsal surface. The wide inter-
orbital dimension also gives a tapering appearance to the rostrum of the Re-
cent heteromyids, when viewed dorsally, which is not seen in the fossil speci-
men. Like those of most heteromyids, the nasals and premaxillaries project
forward beyond the incisors.
H. tenuiceps has a distinctly heteromyidlike appearance, and it is obvious
that the features of the anterior part of the skull, which characterize the
heteromyids, had been established by middle Oligocene time.
The nasal bone extends caudad as far as does the premaxillary ; they termi-
nate at the anterior border of the orbit. The nasal is widest anteriorly where
it curves downward on the side to meet the anterior projection of the pre-
maxillary bone beyond the incisor. Posteriorly, the two nasals have practically
parallel lateral borders much as in Liomys.
The frontal bone dorsally is relatively narrower than in any Recent hetero-
myid, and closely resembles that of the geomyids. There is a slight depression
in the midline of the skull where the two frontals unite, but no evidence of a
ridge for the attachment of the temporal muscle. In lateral view, the ledge
seen in Liomys at the dorsal surface is absent, nor is this surface rounded as
in Geomys. Preservation around the nasolacrimal canal is poor, but traces of
sutures indicate that the frontal bone is not involved in the posteromedial
wall of that canal. The orbital plate is broad, comparatively flat, and extends
farther ventrad than in Liomys, and enters into the composition of the spheno-
palatine foramina. Ventrally the frontal bone meets the orbital processes of
the palatine and maxillary bones, and posterolaterally meets the orbito-
sphenoid.
In the anterodorsal angle of the rim of the orbit the lacrimal bone rests
against the frontal and maxillary bones, where the body of the lacrimal con-
tributes to the formation of the posteromedial wall of the nasolacrimal canal.
Only a slight part of the maxillary process of the lacrimal remains on each side.
The premaxillary bone, which constitutes most of the anterior part of the
rostrum, is typically heteromyid in shape. The frontal process is long and
slender. On the side of the rostrum the premaxillary forms the anterointernal
border of the infraorbital foramen. The ventrolateral border of the bone is
expanded slightly and aids in the formation of the tuberosity made by the
maxillary bone at the ventroposterior border of the foramen. Ventrally the
premaxillary makes up the anterior two-thirds of the lateral wall of the in-
cisive (anterior palatine) foramen. It is not possible to establish what part
of the median septum between the foramina is made up of premaxillary bones.
The incisor arches through the premaxillary in a manner similar to that in
Liomys, with the upper wall of the root canal being formed by the upper
surface of the bone. Due to the narrowness of the rostrum, the root of the
incisor is prominently outlined on the side of the rostrum, both in the pre-
maxillary and maxillary bones. With this modeling of the side of the rostrum
because of the incisor root canal, and the flaring of the posterior and ventral
edges of the infraorbital foramen, the side wall of the premaxillary appears
as a depressed area. Anterior to the incisor root the tip of the premaxillary
projects forward, and parallels its opposite, laterally, instead of turning inward
Galbreath: A New Heteromyid Rodent 291
as in Liomys. This condition, together with the prominence of the root canal,
makes the anterior tip project as a flange. The premaxillary extends down-
ward as a plate of bone, and embraces the posterior and lateral sides of the
incisor as in Recent heteromyids. The interpremaxillary foramen, if present,
is obscure. However, there appears to be a foramen posterior to the incisor,
which possibly has taken over the function of the interpremaxillary foramen.
Both maxillary bones are incomplete, and lack the zygomatic processes.
The rostral part of the maxillary is compressed laterally, as is the premaxillary.
The anterior border of the maxillary contributes to the formation of the border
of the anterior opening of the infraorbital canal where, at the posteroventral
border of the opening, the bone is produced into a prominent tuberosity which
projects laterally approximately one millimeter on each side. The infraorbital
foramen (anterior opening of the infraorbital canal) lies about midway between
the anterior end of the skull and the root of the zygoma. High on each side
of the rostrum, and beneath the dorsal edge of the masseteric plate, is an area
containing small foramina. The zygomasseteric plate is inclined forward at
the dorsal end, and extends anteriorly almost to the highest part of the arch
of the canal for the root of the incisor. The posterior end of the infraorbital
canal lies on the median side of the zygomatic root as it does in H. hatcheri
Wood. Ventrally the zygomatic root rises above the fourth premolar as in
H. gregoryi Wood, H. hatcheri, and in Recent heteromyids. The ventral part
of the orbit, containing the sphenopalatine foramen, presphenoid foramen, and
the sphenoidal fissure, is not constricted as in Liomys, but is open like that
of the squirrels. This condition is emphasized by the narrowness of the inter-
orbital part of the skull and the more vertical position of the orbital plate.
The alisphenoid bone is large and forms part of the posteromedial wall of
the orbit. The sphenofrontal foramen lies in the suture between the extreme
anterior margin of this bone and the frontal bone.
The orbitosphenoid bone makes up little of the orbital wall. It occupies
the posterior area of the' orbit between the alisphenoid and palatine, and is
in contact with these bones and the frontal. The presphenoid canal between
the orbits is large, and the entrance at each end is well separated from the
sphenoidal fissure. Damage to the sphenoidal fissure, which occurred prior to
preservation, obscures its relationship to the optic foramen. No bar was
found that would indicate that the two openings were widely separate. Antero-
ventrally the sphenoidal fissure is bounded by the orbitosphenoid bone, and
dorsolateral^ by the alisphenoid bone. Between the presphenoid foramen and
the orbitosphenoid-frontal suture there is a distinct ridge, and the suture be-
tween the two bones lies in an elongate pit or trough formed by the anterior
sloping side of the ridge and the impressed lateral wall of the frontal bone.
The palatine bone is represented by fragments joined to other bone9 of the
skull. The maxillary process of the left palatine bone is united to the maxil-
lary by a highly sinuous suture. The union of the palatines to the maxillaries
make a suture in the shape of a "V" with the base forward and somewhat blunt.
The canal for the palatine artery and nerve has a multiple opening on the
palate. One major foramen opens on each side of the palatomaxillary suture,
and two or possibly three smaller foramina open posteriorly on the palatine
bone. Prominent on the palatine bone, posteromedial to the third molar, is
292 University of Kansas Publs., Mus. Nat. Hist.
the foramen (palatine pit) for the palatine vein. Collectively, this complex
of foramina is often known as the posterior palatine foramina. Wood (1933)
states that H. gregoryi has two posterior palatine foramina as in Recent
genera, the anterior one opening opposite the posterior end of Ml, and the
posterior one opposite the median part of M3. The orbital process of the left
palatine bone lies inside (medial to) the palatine process of the maxillary.
Anteriorly this orbital process meets the orbital process of the maxillary bone,
and the sphenopalatine foramen is found in the suture between these two
bones and the frontal.
As previously mentioned, the preserved dentition of this specimen consists
of the complete left row of cheek teeth and roots of the incisors.
The incisor is compressed laterally, more so than in any Recent heteromyid.
The anterior face is rounded, asulcate, and covered with a heavy band of
enamel, whereas the posterior side, due to lateral compression, is drawn out
into a thin blade. The root of the incisor is at the lateral border of the pre-
maxillary, so it is obvious that the two incisors converged on each other at the
midline to form a cutting surface. The writer has not examined the asulcate,
laterally compressed incisors of //. halcheri, and cannot say how they compare
with this specimen.
The most significant features of the cheek teeth are their size, and the un-
divided internal cingulum. The molars are well worn, but the pattern, as a
whole, is easily discernable.
P4 has an anterior cusp and three posterior cusps as in other members of
the genus. However, the buccal cusp (metacone) of the metaloph is consider-
ably anterior to the central (hypocone) and lingual (entostyle) cusps, and
the three cusps do not form a curve as in other species. In size the central
«usp is largest, the buccal cusp is practically as large, and the lingual cusp is
small. A cingulum connects the lingual and central cusps at the posterior
margin of the tooth. In the Pipestone Springs specimen of Heliscomys re-
ported by McGrew (1941) the central and buccal cusps were connected by a
cingulum, and some H. hatcheri specimens have all three cusps connected in
a similar manner. A low arm or ridge extends from the lingual cusp forward
to the lingual side of the base of the anterior cusp. The valleys between the
posterior cusps are shallow. There is no sign of the small cuspule on the
anteroexternal base of the anterior cusp seen in H. gregoryi, H. hatcheri, and
the Pipestone Springs specimen. However, when one sees the variability of
the cuspules on P4 of H . hatcheri, the presence of a minor cuspule does not
seem to be of taxonomic importance.
Ml deviates from the pattern typical of Heliscomys- move than do any of
the other molar teeth. However, it must be kept in mind that some of the
differences may be due to wear. For example, the protocone and paracone,
and the hypocone and metacone are united to form protoloph and metaloph
respectively. If the height of the external border of the paracone and metacone
is taken into account and compared with the worn inner parts of these two
cusps and the equally well-worn protocone and hypocone, it appears that these
cusps formed no more of a true bilophodont tooth than do the cusps in other
species of Heliscomys; in each of the species the cusps generally are separate
entities. II . gregoryi is reported to have an "incipient tendency to form lophs,"
Galbreath: A New Heteromyid Rodent 293
and H. hatchcri does the same when worn, but by union with the anterior
cingulum. If cusps in H. tenuiccps do form lophs, the process is definitely not
by union of the cusps with the anterior cingulum. The transverse median
valley is deep and divides the tooth on the buccal side. The anteroposterior
valleys are shallow and hanging, and can be said to exist only as indentations
between the two sets of cusps. The paracone and metacone are much higher
than the other two cusps, but much of this disparity in height may be the
result of greater wear on the protocone and hypocone; H. gregoryi agrees with
H. tenuiceps in these respects. Possibly the protocone and hypocone were
much larger than the paracone and metacone. The internal cingulum of Ml
exhibits only one large cusp opposite the medial end of the transverse valley,
and shows no evidence of having been divided into two cusps. It is barely
possible that there may have been two cusps and that wear makes it appear
that there was only one. I doubt that there were two cusps because the
cingulum is still so high (as high as the outer edges of the paracone and meta-
cone) as to suggest that it is only slightly worn. Posteriorly this single cusp
in the cingulum is united with the hypocone. Anteriorly the cusp is con-
fluent with an anterior cingulum that is small, but, nevertheless, plainly visible
as it crosses the occlusal face of the tooth to the paracone. There is some
reason to believe that there was a posterior cingulum, but wear, which has
obliterated even the posterior wall of the hypocone, prevents my being certain
about this. This cingulum is absent in H. gregoryi and present in H. hatcheri.
M2 compares favorably with Ml except for the following differences: The
protocone and hypocone are equal to the paracone and metacone in area, but
not in height; although the internal cingulum is undivided, there is no evi-
dence of a cusp as in Ml. Here, too, the cingulum is as high as the paracone
and metacone. Possibly the cingulum was confluent with the hypocone. The
internal cingulum continues around the margin of the tooth to the paracone
as an anterior cingulum which is sharper and plainer than the anterior cingulum
on Ml. There is no evidence of a posterior cingulum.
M3 shows a great amount of wear, and the occlusal pattern is not too clear.
The median transverse valley is reduced almost to a pit, and the paracone and
metacone are divided by a small notch. The protocone and paracone, the
latter being much higher, are larger than the metacone which is reduced in
size, and not all this difference in size can be the result of wear. The hypo-
cone is absent. The internal cingulum is as high as the paracone and shows
no evidence of division into two cusps, but in M3 this character is apparently
variable for H. gregoryi does not have the internal cingulum divided and
H. hatcheri has it markedly so. A slight anterior arm of the internal cingulum
may have reached forward to the anterior face of the protocone. Wear pre-
vents knowing whether a crest surrounds the tooth completely, or only on
three sides.
In size the teeth of H. tenuiceps average twenty per cent larger than any of
the upper teeth of H. gregoryi, H. hatcheri, or the Pipestone Springs specimen,
and exceed any of the known lower teeth including those of H. vetus and H.
senex by twenty-five per cent or more. Inasmuch as the upper teeth rarely
exceed the lower in length in all the related genera of heteromyids, it is as-
sumed that a similar relationship existed between the upper and lower molars
294 University of Kansas Publs., Mus. Nat. Hist.
of //. tenuiceps and, therefore, that this species can be distinguished by its
large size. The relative size of the premolars and molars is the same in H.
tenuiceps as in other species of Heliscomys. However, within the framework
of this similar relationship there are two differences. P4 of H. tenuiceps is
relatively larger than the P4 of H. gregoryi, and relatively smaller than the
P4 of H. hatcheri. The width of the molars is relatively greater in H. tenuiceps
and H. gregoryi than in H . hatcheri.
MEASt'REMENTS
(In millimeters)
U. K. M. N. H.
(Vert. Paleo.)
No. 7702
Height of skull at M2 7.48
Length from anterior end of nasals to rear of M3 15.41
Length of nasal bones 10 . 50
Width of rostrum at highest point of root canal 3.97
Interorbital width 4.39
Estimated length of skull 25 . 00
I, anteroposterior length 1 .56
I, transverse width 0 . 63
P4-M3 crown length 3.75
P4-M3 alveolar length 3.80
P4, anteroposterior length * 1 . 05
P4, transverse width 1 . 08
Ml, anteroposterior length 0.93
Ml , transverse width 1 . 17
M2, anteroposterior length 0.93
M2. transverse width 1.14
M3, anteroposterior length 0.78
M3, transverse width 0.93
Discussion. — Heliscomys tenuiceps shows beyond any doubt that
the heteromyid pattern of skull was developed by mid-Oligoeene
times, and in this species was already undergoing lateral compres-
sion. The major change later made in heteromyid skulls is broaden-
ing of the dorsal surface of the skull in the interorbital area.
The complete confirmation of Wood's (1939) statement that the
"sciuromorph" zygomasseteric structure had been developed by this
time in the heteromyid rodents as it had been in the early Eomyids
is demonstrated in this specimen. Further, it is to be noted that the
infraorbital canal is not sciuridlike, but has been forced forward on
the rostrum, as in the Geomyoidea.
In some ways this skull shows similarities to Florentiamys loomisi
Wood, of the early Miocene, which aid in determining the relation-
ship of that unusual genus to Heliscomys and to the heteromyids in
general. When Wood described Florentiamys the peculiar combi-
nation of characters found in this animal prompted him to speculate
that: (1) It was a typical heteromyid which had secondarily de-
* This and the following measurements at occlusal surface.
Galbreath: A New Heteromyid Rodent 295
veloped cingula; (2) its cheek teeth were nearer the primitive pat-
tern than were those of any other known fossil heteromyid, and
that Heliscomys represented a simplification in the reduction of the
cingula; or (3) it was not a heteromyid, but a parallel development
from the 'Taramys" stock. Wood favored the second possibility.
Now that a part of the skull of one species of Heliscomys is known,
the undivided internal cingulum that is confluent with the hypocone,
the lateral compression of the deep rostrum, and the general simi-
larity to the heteromyids appear as points in common between the
two skulls, and demonstrate the closeness of Florentiamys to the
heteromyids. However, the specimen does not contribute anything
new to use in choosing between Wood's first two postulates. In the
writer's opinion the undivided internal cingulum is a primitive con-
dition that has survived in Florentiamys and Heliscomys tenuiceps.
This common character together with the laterally compressed
rostrum leads me to think that structurally, H. tenuiceps is a link
between Florentiamys and the ancestral form of Heliscomys. Ad-
mittedly P4 of Florentiamys seems far from the Heliscomys pattern,
but I think that this highly specialized structure could have been
derived from Heliscomys or a common ancestor.
LITERATURE CITED
McGrew, Paul O.
1941. Heteromyids from the Miocene and Lower Oligocene. Geol. Ser. of
Field Mus. Nat. Hist., vol. 8, pp. 55-57, 1 fig.
Wood, Albert E.
1933. A New Heteromyid Rodent from the Oligocene of Montana. Jour.
Mamm, vol. 14, pp. 134-141, 5 figs.
1935. Evolution and Relationship of the Heteromyid Rodents with New
Forms from the Tertiary of Western North America. Annals of the
Carnegie Mus., vol. 24, pp. 73-262, 157 figs.
1937. Part II. Rodentia, in The Mammalian Fauna of the White River
Oligocene; by William Berryman Scott and Glenn Lowell Jepsen.
Trans. Amer. Phil. Soc, ns., vol. 28, pp. 155-269, figs. 8-70, pis. 23-33.
1939. Additional Specimens of the Heteromyid Rodent Heliscomys from
the Oligocene of Nebraska. Amer. Jour. Sci., vol. 237, pp. 550-561,
11 figs.
Transmitted March 1, WJ,S.
n
22 3312
Speciation in the Brazilian Spiny Rats
(Genus Proechimys, Family Echimyidae)
BY
JOAO MOOJEN
PS. CO&P. ZOOL
LIBRARY
JAN 30 Id
University of JLansas Publications
Museum of Natural History
Volume 1, No. 19, pp. 301-406, 140 figures in text
December 10, 1948
University of Kansas
LAWRENCE
1948
UNIVERSITY OF KANSAS PUBLICATIONS
The University of Kansas Publications, Museum of Natural His-
tory, are offered in exchange for the publications of learned societies
and institutions, universities and libraries. For exchanges and in-
formation, address the Exchange Desk, University of Kansas Li-
brary, Lawrence, Kansas, U. S. A.
Museum of Natural History. — E. Raymond Hall, Chairman, Editorial Com-
mittee.
This series contains contributions from the Museum of Natural History.
Cited as Univ. Kans. Publ., Mus. Nat. Hist.
Vol. 1. 1. The pocker gophers (genus Thomomys) of Utah. By Stephen D.
Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces pluvialis Cope, and noteworthy
records of other amphibians and reptiles from Kansas and Okla-
homa. By Hobart M. Smith. Pp. 85-89. August 15, 1946.
3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp.
93-96, 1 figure in text. August 15, 1946.
4. Hybridization between two species of garter snakes. By Hobart M.
Smith. Pp. 97-100. August 15, 1946.
5. Selected records of reptiles and amphibians from Kansas. By John
Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.
6. Kyphosis and other variations in soft-shelled turtles. By Hobart
M. Smith. Pp. 117-124. July 7, 1947.
7. Natural history of the prairie vole (Mammalian genus Microtus).
By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6,
1947.
8. The postnatal development of two broods of great horned owls
(Bubo virginianus) . By Donald F. Hoffmeister and Henry W.
Setzer. Pp. 157-173, 5 figures in text. October 6, 1947.
9. Additions to the list of the birds of Louisiana. By George H.
Lowery, Jr. Pp. 177-192. November 7, 1947.
10. A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-
216. November 29, 1947.
11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R.
and E. Raymond Hall. Pp. 217-236, 2 figures in text. November
29, 1947.
12. A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest
and E. Raymond Hall. Pp. 237-244, 6 figures in text. December
10, 1947.
13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By
Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure
in text. December 10, 1947.
14. A new pocket gopher (Thomomys) and a new spiny pocket mouse
(Liomys) from Michoacan, Mexico. By E. Raymond Hall and
Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
<S-NA-L
Speciation in the Brazilian Spiny Rats
(Genus Proechimys, Family Echimyidae)
BY
JOAO MOOJEN
University of Kansas Publications
Museum of Natural History
Volume 1, No. 19, pp. 301-406, 140 figures in text
December 10, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Edward H. Taylor
Volume 1, No. 19, pp. 301-406, 1 plate, 140 figures in text
Published December 10, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3343
Speciation in the Brazilian Spiny Rats
M0S(C^Sui^r3echimys, Family Echimvidae)
By
JOXO MOOJEN
!';
CONTENTS
P\(iE
Introduction 305
Methods and Terminology 305
Acknowledgments 308
Paleontology 308
Speciation 311
Subgeneric variation 312
Specific variation in the subgenus Proechimys 314
Subspecific variation in the subgenus Proechimys 317
Specific variation in the subgenus Trinomys 320
Subspecific variation in the subgenus Trinomys 322
Taxonomic Characters 323
Size and proportions of external parts 323
Pelage . 324
Skull 326
Incisive foramen 320
Teeth 327
Habits 330
Changes with Age 331
Genus Proechimys 333
Artificial Key to Subgenera and Species 334
Subgenus Proechimys 338
Proechimys goeldii 338
Proechimys goeldii steerei 338
Proechimys goeldii goeldii 340
Proechimys semispinosus 342
Proechimys semispinosus liminalis 343
Proechimys semispinosus amphichoricus 34- 1
Proechimys semispinosus kermiti 345
(303)
304 University of Kansas Publs., Mus. Nat. Hist.
PAOE
Proechimys longicaudatus 346
Proechimys longicaudatus brevicauda 349
Proechimys longicaudatus boimensis 350
Proechimys longicaudatus longicaudatus 351
Proechimys longicaudatus leucomystax 352
Proechimys longicaudatus roberti 353
Proechimys guyannensis 355
Proechimys guyannensis villicauda. 355
Proechimys guyannensis ribeiroi 361
Proechimys guyannensis hyleae 361
Proechimys guyannensis nesiotes 363
Proechimys guyannensis leioprimna 364
Proechimys guyannensis oris 365
Proechimys guyannensis arescens 366
Proechimys guyannensis riparum 367
Proechimys guyannensis arabupu 369
Subgenus Trinomys 369
Proechimys dimidiatus 371
Proechimys iheringi 373
Proechimys iheringi iheringi 378
Proechimys iheringi bonafidei 378
Proechimys iheringi gratiosus 379
Proechimys iheringi panema 380
Proechimys iheringi denigratus 381
Proechimys iheringi paratus 382
Proechimys setosus 384
Proechimys setosus setosus 385
Proechimys setosus elegans 387
Proechimys albispinus 388
Proechimys albispinus albispinus 390
Proechimys albispinus sertonius 391
Incerta Sedis 392
Proechimys myosuros 392
Conclusions 393
Table of Measurements 395
Literature Cited 400
Fig. 1. Proechimys dimidiatus (Giinther). Live female on lefl and male on
right. X Vz. From Tingua, Nova Iguassu, Rio de Janeiro, Brazil. Photo-
graphed in spring (August or September) of 1H42 by author.
INTRODUCTION
The spiny-rats included in the genus Proechimys are common in
almost every forest of South America above the Tropic of Capri-
corn, and in Central America northward to approximately 12° N,
in Nicaragua. In size and proportions they are similar to the brown
rat Rattus norvegicus but actually they belong to a very different
suborder of rodents — the Hystricomorpha. The hystricomorphs are
represented in South America by a large variety of animals, of which
capybaras, agoutis and cavies are common representatives.
The pelage of the spiny-rats has a large number of flattened,
spinelike hairs, especially on the back. The color ranges through
different tints and shades of reddish-brown more or less evenly dis-
tributed on the upper parts; the underparts are usually pure white,
sharply contrasting with the brown color above. The tail is bi-
colored, brown above and white below.
The spiny-rats live in forests of different types, generally in the
proximity of water. Shelter is usually procured under boulders,
stumps or masses of roots. The reproductive rate is low; on the
average, there are only two young per litter and only two litters
per year.
Sixty-odd names have been given to species and subspecies of
Proechimys in the last hundred and fifty years and no serious re-
vision of the taxonomy of the genus was undertaken in the last
century. The purpose of the present work is to provide means of
understanding species and subspecies within the genus and to de-
scribe the different kinds known to occur within the confines of
Brazil.
METHODS AND TERMINOLOGY
Pelage. — It was found advisable to use a standardized nomenclature for
hairs. The names here proposed are a choice of those used in the literature,
with the suffix "form" as an element of uniformity. I feel that it would be
advantageous if everyone adopted a similar universal system in mammalogy.
The names listed below are used as nouns and are considered as English
versions which could easily be adapted to different languages. These names
may be complemented with adjectives as needed. Examples are lanceolate
aristiforms, spinous aristiforms, and woolly setiforms.
Aristiforms: The most conspicuously developed hairs in a three-layered
pelage or the corresponding hairs in a simpler pelage. Names previously used
for these hairs are: guard hair, leithaar and jarre.
(305)
306 University of Kansas Publs., Mus. Nat. Hist.
Setiforms: Common to all species and most numerous throughout the pel-
age; second in conspicuousness, being the dominant hairs in the middle layer.
Synonyms are : over hairs, grannenhaare and soies.
Villiforms: The smallest hairs in the three-layered pelage. Synonyms are:
underfill-, wollhaar and duvet.
Vibrissi forms: The vibrissae proper, or any typically sensory hair.
Teeth. — The tri tubercular nomenclature was abandoned because of over-
whelming difficulties; more research on the Hystricomorpha is certainly needed
before the tritubercular nomenclature can be applied with confidence. The
following names are used for features of the molariform teeth :
Main fold: The inner or lingual fold in the upper molariform teeth and
outer or labial fold in the lower molariform teeth.
Counlerfold: Any outer or labial fold in the upper or any inner or lingual
fold in the lower molariform teeth.
For incisors Thomas (1921:141) is followed: opisthodont, orthodont and
proodont depending on the angle between the exposed part of incisors and
the ventral surface of the rostrum.
The capital letters P and M designate premolars and molars, respectively,
of the upper jaws; lower case letters p and m designate corresponding teeth
in the lower jaws.
Measurements. — Measurements of skins were used only when provided by
the collector. The length of the hindfoot is intended to be always cum unguis,
but in a few instances it is impossible to be sure whether the collector in-
cluded the nail. Length of tail was used only when the tail seemed not to
be mutilated. Ear measurements taken by collectors are scarce. In spite of
the apparent usefulness of length of ear, it was found to be inadvisable to
take the measurement on the dry skins.
The following measurements of the skull are used in the tables:
Greatest length: From the anteriormost part of the nasals to the posterior-
most part of the supraoccipital.
Condylo-incisive length: From the anterior face of one incisor, at the al-
veolus, to the posteriormost part of the exoccipital condyle of the same side.
Zygomatic breadth: Maximum distance across zygomata in a plane per-
pendicular to longitudinal axis of the skull.
Length of nasals: Maximum length of one or both, whichever is the greater.
lnterorbital constriction: Least width between the orbits on top of the
.skull.
Palatilar length: From the posterior face of an incisor, at the alveolus, to
the nearest part of the posterior edge of the palatine bone.
Crown length of cheekteeth: From the anterior border of P4 to the pos-
terior border of M3.
In the accounts of species, measurements of aristiforms and setiforms are
used. The hairs measured were taken from the middorsal region and outer
thighs, and the measurements are means.
All specimens of which measurements are here recorded, as for example in
Moojen: Brazilian Spiny Rats 307
the tables, are fully adult; each specimen shows some wear on each of the
four upper molariform teeth unless otherwise indicated.
Capitalized color terms are after Ridgway "Color Standards and Color
Nomenclature," Washington, D. C, U. S. A., 1912. One setiform was taken
from the animal and placed over the rectangles in Ridgway's charts and the
examination made under a microscope with low (X7) magnification and
natural light. This method was found to give the most satisfactory results.
The following abbreviations are used for names of institutions:
AMNH — American Museum of Natural History.
CNHM — Chicago Natural History Museum.
DZ — Departamento de Zoologia da Secretaria de Agricultura, Sao Paulo,
Brazil.
MCZ — Museum of Comparative Zoology at Harvard College.
MN — Museu Nacional, Brazil.
MZ — Museum of Zoology, University of Michigan.
SEPFA — Servico de Estudos e Pesquisas sobre a Febre Amarela, Brazil.
USNM— United States National Museum.
UZM — Universitets Zoologiske Museum, Copenhagen.
308 University of Kansas Plbls., Mus. Nat. Hist.
ACKNOWLEDGMENTS
Approximately two thousand skins and skulls were assembled at the Mu-
seum of Natural History, University of Kansas, through the cooperation of
the authorities in the various institutions of North America, Brazil and Den-
mark, as listed immediately above. This comprehensive material was used
to obtain a more complete understanding of the group, and for the loan of
these specimens I am extremely grateful to the authorities of each of the in-
stitutions.
First of all I acknowledge the encouragement given me in the Proechirnys
project by Heloisa Alberto Torres, Director of the Museu Nacional, Rio de
Janeiro. I extend my thanks also to Stephen D. Durrant, of the University
of Utah, for helpful corrections in the preparation of the manuscript; to Mrs.
Virginia Cassell Unruh, for the preparation of the drawings of the skulls; to
Miss Alice M. Bruce for assistance in drawing the maps; and to my daughter,
Julieta, for help in assembling data and for typing.
Dr. Remington Kellogg, Curator of Mammals in the United States Na-
tional Museum, and the late Dr. Wilfred H. Osgood, formerly Curator Emer-
itus of the Department of Zoology in the Chicago Natural History Museum,
generously permitted me to use their private lists of South American mam-
mals. These lists contain much unpublished data, as for example, proof, in
Kellogg's list, that Proechimys guyannensis (E. Geoffroy Saint-Hilaire, 1803)
antedates P. cayennensis (Desmarest, 1817). I register here my gratitude to
both these zoologists and acknowledge other critical assistance from Dr. Kel-
logg.
The John Simon Guggenheim Memorial Foundation awarded me a fellow-
ship for which I am deeply grateful. This expression of the Foundation's in-
terest in education and good neighborliness made possible the completion of
the present paper.
Finally I desire to express my deepest gratitude to Professor E. Raymond
Hall, Director of the Museum of Natural History and Chairman of the De-
partment of Zoology at the University of Kansas whose untiring aid and guid-
ance has enabled me to terminate this study.
PALEONTOLOGY
The only known, significant, fossil Proechimys comes from de-
posits in the limestone caves of Lagoa Santa, Minas Gerais, Brazil.
These deposits, of Late Pleistocene or Recent age, were extensively
studied by P. W. Lund and the results published in a series of
French and Danish papers. F. Ameghino (1934:110) studied an-
other fauna from a deposit of similar age in the cave of Iporanga,
Sao Paulo, Brazil. Proechimys is recorded in his account under the
inclusive specific name fuliginosus.
The molariform teeth of the fossil described by Lund (1841:
pi. 21, fig. 14) show's its close relationship to the living form P. s.
Moojen: Brazilian Spiny Rats 309
elegans (Lund) which still inhabits the same region. It belongs in
the more specialized subgenus Trinomys which seems to have been
derived from Proechimys. Trinomys has the main fold in the molars
always greatly developed and the fold tends to set apart one lamina
in the occlusal surface. The Lagoa Santa fossil, like some specimens
of the living subspecies, has a small main fold in P4. However, the
main fold is large in all upper molars and in the lower molariform
teeth which are notably specialized in the extreme reduction of the
number of counterfolds to only one.
One hypothesis concerning the evolution of the genus is that a
more primitive group of Proechimys lived in all of the Central
Plateau of Brazil in the Pleistocene Time. The climatic conditions
at that time might have been such as to support large forests but,
since the Pleistocene, these climatic conditions may have changed
from humid to the present drier conditions, which support the
dominant, savanna, floral climax. Actually the extinct fauna from
the caves includes animals which have disappeared from the area
and now live only in more humid areas, as for example Myocastor,
which has shifted to the lowlands to the west and south.
Possibly climatic changes were responsible for the faunal shift
from the region that is now a plateau in Central Brazil. This
climatic change may have resulted from the gradual uplift of the
eastern part of the continent, This uplift prevents part of the
trade winds which come from the east from carrying the same
amount of moisture inland as they did previously. In fact, the
Andean revolution, even if it occurred as late at Late Tertiary,
would have had no perceptible influence on the amount of water
precipitated on the more eastern parts of the continent. Oliveira
and Leonardos (1943:617) point out that after a Cretaceous sub-
mersion of the central part of Brazil, there was a general uplift.
The authors (op. cit. :689) mention the presence of continental
Cretaceous deposits in the Central Plateau of Brazil, in support of
these changes, and state that "pelo menos em certas zonas do litoral
a elevagao do continente prolongou-se ate o Pleistocene "
Berry (1942:373) concluded, among other things, that there was
an southward extension "in South America of equatorial floras in
the lower Miocene." and (op. c?£.:372) that . . . "east of the
Andean Axis in the south temperate zone there was a normal
mesophytic flora . . . instead ... of present day large
steppes."
My idea is that a tropical forest still covered the Central Plateau
of Brazil in (early?) Pleistocene times and that populations of
310 University of Kansas Publs., Mus. Nat. Hist.
Proechimys of a primitive type, similar to P. g. steerei, for example,
lived in that extensive forest-climax. The gradual uplift of the
plateau, however, gradually brought about drier conditions in this
region. As a result a large cliscral change was initiated, which
shifted the forest-climax to the more humid eastern escarpments and
lowlands that were gradually being developed, while the savanna
climax was being established on the plateau. Eventually the effect
of the decreasing moisture was locally accentuated by the erosion
of the sandstones (Oliveira and Leonardos, 1943:690) in north-
eastern Brazil, thus depriving it of a natural reservoir of rain water.
An arid belt was developed which now constitutes an efficient geo-
graphic barrier to the distribution of many kinds of animals.
One marginal species may have shifted eastward with the forest-
climax to effect the Recent distribution. The eastern species be-
came completely isolated from the main group, accumulated muta-
tions, and evolved into the subgeneric type Trinomys. The generic
trend that gave rise to Trinomys probably remained more stable as
far as supraspecific changes are concerned. The lack of barriers in
the distributional area of the original group favored the dispersal
and submergence of mutations and, therefore, there was but little
further supraspecific evolution. The speciation in both subgenera
finally resulted from gradual differentiation of varying populations
since they show combinations of the generic biotypes and possess
few truly qualitative characters.
The cliseral changes in the Central Plateau, which developed the
dry belt, a barrier, might explain the evolution of a few more
supraspecific groups of mammals, as indicated by the presence of
similar forms in the Amazonian region and in Southeastern Brazil.
Among these Echimys and Phyllomys, in the same family with
Proechimys, show differences that are parallel to those observed in
Proechimys. One of these parallel changes is the increased lamina-
tion of the cheekteeth. Although Echimys, from the Amazonian
region, has upper molariform teeth with the four laminae fused,
Phyllomys has the four laminae completely separated.
None of the genera known from the Upper Oligocene and Miocene
of Argentine deposits seems to be directly ancestral to Proechimys.
Moojkn: Brazilian Spiny Rats 311
SPECIATION
The detection of differences of systematic worth between popula-
tions of animals, represented by skins and skulls, is a step prelimi-
nary to deducing the factors responsible for the differences. Ordi-
narily the factors which cause heritable differences have to do with
geographic isolation and adaptation to ecological conditions. When
differences in the structure of the animal are known, a person is
led to speculate on the factors which could cause them. For one
thing, does the observed degree of difference tend to isolate animals
possessing the "new" character from the other animals? It would
seem to me that the isolation once started by one of these differ-
ences tends to be accentuated with time and the difference itself
thus then becomes a factor responsible for further differentiation.
Whether or not transition from one character to another occurs
gradually, in its geographic expression, and thus whether or not
intergradation occurs between two subspecies, can be ascertained
by the analysis of a series of population-samples appropriately dis-
tributed geographically. If two characters of systematic worth are
known to blend in one part of the geographic range of a subgenus,
and if the same two characters are seen in two other populations,
far removed geographically from each other and without any
samples of annectent populations to provide actual evidence of
intergradation, then such intergradation is to be inferred.
The available collections of Proechimys mostly were made hap-
hazardly with the result that there are extensive areas from which
no specimens as yet are available. Thus, actual proof of intergrada-
tion is often lacking in areas where it almost certainly occurs. In
some extensive areas, however, many samples, from relatively reg-
ular intervals, have been available and they provide genuine proof
of intergradation. These instances have served as a guide for esti-
mating whether other samples should be considered to be full species
or instead merely subspecies of the same species.
Lack of intergradation in any of the characters may be accepted
as the criterion of full species. Where two populations occupying
the same range (sympatric populations) show different qualitative
characters, they almost certainly do not crossbreed. Furthermore
the characters that distinguish such kinds of nonintergrading ani-
mals are likely to be considered as of full specific value when de-
tected in far distant parts of the range of the subgenus.
312 University of Kansas Publs., Mus. Nat. Hist.
In a genus that is widespread and continuously distributed, it it
useful to know which characters always distinguish full species and
which ones, sometimes or always, distinguish only subspecies, since
in a population from a small island, there is, ordinarily, less indivi-
dual variation than in a corresponding population from the main-
land or a larger island; under certain circumstances a person might
be tempted to give specific rank to the population when its charac-
ters actually are analogous to those separating subspecies else-
where.
Sometimes it is convenient to recognize species-groups, a system-
atic category without nomenclatural status, intermediate between
the species and the subgenus. When there are two groups of species
not sharply separated, including one species whose characters over-
lap those of each of the two groups, it would seem most appropriate
to recognize only species-groups instead of subgenera. When, on
the other hand, the two groups of species have mutually exclusive
characters and a species with intermediate characters is unknown,
the two groups of species can conveniently be accorded separate
subgeneric rank.
SUBGENERIC VARIATION
A few characters are common to one group of species and other
features are common to a second group. The most striking of these
features is the character of the main fold in the molariform teeth.
In one group the fold transversely crosses the crown of the tooth
and in the other it extends scarcely halfway across. No specimen
is intermediate in this respect. These two groups, furthermore, are
separated geographically by an important barrier, the arid belt that
starts in the northeastern littoral of Brazil (Ceara), and that ex-
tends south and southwesterly, more or less accompanying the Sao
Francisco River in the Plateau, to about 20° S. Proechimys is
thought not to inhabit this arid belt. At the latitude of 20° S the
conditions become more suitable for Proechimys, especially along
the rivers which flow eastward, but there the Plateau is replaced
by mountains: the Serra Geral at the west, and Serra da Manti-
queira at the south; these ranges are bare of forests at higher ele-
vations. Two groups of species of Proechimys are, therefore, kept
geographically isolated: one group lives in southeastern Brazil, and
the other lives in a large area to the west which starts at 21° S in
Paraguay and Brazil and widens northward and includes, farther
west, central and northern Brazil and all the South American coun-
Moojen: Brazilian Spiny Rats 313
tries above 21° S, as well as Central America northward to southern
Nicaragua.
The two groups which are here treated as subgenera may be desig-
nated as follows :
Trinomys — main fold deep: aristiforms well-developed on the
rump and outer thighs; tail no less than 75 per cent of length of head
and body ; skull without ridges across the parietals ; no conspicuous
groove for transmission of nerve inside infraorbital foramen; molari-
form teeth decreasing in size from premolar to third molar; 1 to 3
counterfolds in the molariform teeth.
Proechimys — main fold shallow: aristiforms not developed on
rump and outer thighs; tail less than 75 per cent of length of head
and body; groove for transmission of nerve present in infraorbital
foramen of several subspecies; molariform teeth increasing in size
from premolar to second molar; 2 to 5 counterfolds in molariform
teeth.
Most of these characters vary but do not overlap. Subgeneric
rank is here accorded to the two groups of Proechimys characterized
immediately above.
The primary cause of the subgeneric differentiation is thought to
have been geologic changes in the continental area. As already
pointed out (see Paleontology), decreasing humidity in the Central
Plateau of Brazil may have caused a migration southwestward of one
or more of the species along with the forests. Once isolated geo-
graphically, the species probably differentiated at an accelerated
rate.
The fact that a much larger number of subspecies occupies the
larger geographic range of the subgenus Proechimys would not be
sufficient to prove that this subgenus, Proechimys, is nearer to the
primitive group than Trinomys, the subgenus occupying the smaller
range with fewer subspecies. The paleontological evolution of the
rodents, however, consistently points to teeth with a larger number
of counterfolds (as seen in Proechimys) as the primitive condition.
The extension of the main fold, tending to set apart one lamina in
each upper molariform tooth, seems to be a specialization; reduc-
tion in the size of the head and body, increase in length of tail and
decreasing size of molars posteriorly also may be specializations. The
main point, however, is to establish if Trinomys is a relic group
rather than a "differentiated" one. If an intermediate form were
known which connected Trinomys with one species of Proechimys
more than with another or even if Trinomys itself more closely re-
314 University of Kansas Publs., Mus. Nat. Hist.
sembled one of the groups of species of the subgenus Proechimys than
it did another, we would assume that divergence and selection ac-
counted for the subgeneric variation. The lack of any such connect-
ing link favors the first idea, namely that Trinomys differentiated
rapidly with the aid of geographic variation.
If Trinomys is, as I am inclined to consider it, the result of "differ-
entiation," its subgeneric features are to be admitted as "new" and
therefore the most primitive species in the genus should be found in
the subgenus Proechimys.
It is a matter of common sense to admit the two groups considered
above as subgenera rather than genera. Since the two structural
plans were established they would, and do, act as different sources of
variation. On the other hand, the morphological differences do not
give the two groups an amount of morphological differences that
would justify full generic rank for each.
SPECIFIC VARIATION IN THE SUBGENUS
PROECHIMYS
Most of the described forms in the subgenus were initially named
as distinct full species. More recently, however, in accordance with
the ideas now prevalent in systematic work, many of the named
kinds were reduced to the rank of subspecies. Tate first made a geo-
graphic arrangement (1935:399-400) and later (1939:177-178) pro-
visionally synonymised several named kinds of Proechimys with
Proechimys "cayennensis cayennensis." A similar tendency was
clearly displayed by Ellerman (1940:115-122) who allocated 29
names, out of 33 (in the subgenus, as here understood), to the species
Proechimys guyannensis and gave full specific rank to four other
named kinds. Osgood (1944) also had the same viewpoint; that is
to say, he appeared to have the idea that there were only two full
species in the subgenus in Brazil — admitting this orally — and con-
sequently he synonymised some full species where two or more oc-
curred in the same place, thinking that he was dealing with indi-
vidual, rather than specific, differences. Evidently the number of
species in the subgenus cannot be great because the known kinds
show few patterns worthy of specific designation and therefore the
majority of the existing names should be suspected of having no
more than subspecific value. Nevertheless none of the above writers
presented real evidence in support of his arrangement.
Criteria for the recognition of full species are most easily recog-
nized where two or more different species live together. In the
Moojen: Brazilian Spiny Rats .''.I.")
literature, P. goeldii and P. "oris" were mentioned by Thomas
(1912:89) as having been collected in the same place; P. mincae
and P. canicollis, by H. H. Smith (in Allen, 1904:440); P. "leu-
comystax," from Utiariti, by Miranda Ribeiro (1914:42) and P.
"longicaudatus," from the same place, by Allen (1916:569) were
other examples. In these, and other alleged instances of two or
more kinds occurring together, detailed study of the specimens con-
cerned was necessary to learn the true facts. Also with the oppor-
tunity to compare collections from several different places, new
facts emerged. P. longicaudatus, as it was conceived of by Allen,
was a composite species, but in one locality, Utiariti, Ribeiro and
Allen actually were dealing with two distinct species.
The species, or subspecies belonging to different species, living
together are: goeldii and hylcae, at Fazenda Paraiso; goeldii and
riparum in Manaus; boimensis and hyleae in Tauari; leucomystax
and villicauda in Utiariti; mincae and canicollis in Bonda; gularis
and hendeei on the banks of Rio Napo (''same trap lines," accord-
ing to P. Hershkovitz, In Litt.). Study of samples of the above
named pairs of kinds of Proechimys showed the following specific
differences: goeldii is large with narrow aristiforms, has a large and
strongly built skull, with four counterfolds in one or more upper
molars: hyleae is smaller, has wide aristiforms, smaller skull with
less pronounced ridges, and never has more than three counterfolds
in the upper molariform teeth; riparum closely resembles hyleae;
boimensis has thin aristiforms, small skull and no more than three
counterfolds in the upper molariform teeth in contrast to hyleae,
already discussed; leucomystax closely resembles boimensis; villi-
cauda closely resembles both hyleae and riparum; mincae is similar
to hyleae-riparum-villicauda; canicollis has the number of counter-
folds in all molars reduced to two; gularis is large, has a strongly
built and ridged skull, some upper molariform teeth with four coun-
terfolds and wide aristiforms; hendeei closely resembles leu-
comystax and boimensis.
The evidence obtained from study of specimens where two or
more species occurred together was applied to the remaining sam-
ples and the geographic distribution was worked out. As a result
the arrangement below was made, including all valid kinds already
named and those here newly named from Brazil. The names of
kinds I do not consider as belonging to the subgenus (and genus)
are excluded. These are Echimys macrourus Jentink, not seen, and
Proechimys cayennensis hoplomyoides Tate (= genus Hoplomys).
The application of names is tentative, however, because the types
316 University of Kansas Publs., Mrs. Nat. Hist.
deposited in Europe have not been seen. An asterisk denotes the
forms not seen by me.
Proechimys guyannensis : arabupu, arescens, bolivianus, cherriei, chrys-
aeolus, gvaiiae, o'cotmelli, guyannensis*, hyleae, leioprimna, mincae,
nesiotes, ochraceus, oris, poliopus, ribeiroi, riparum, trinitatis, urichi,
vacillator* , villicauda, warreni.
Proechimys longicmidatus : boimensis, brevicauda, elassopus, hendeei, leu-
comystax, longicaudatus, nigrojulvus, pachita, rattinus*, roberti, securus,
simonsi.
Proechimys semispinosus : amphichoricus, burrus, calidior, centralis, chiriqui-
nus, colombianus, decumanus, goldmani* , gorgonae, gularis, hilda*, ignotus.
kermiti, liminalis, panamensis, rosa*, rubellus, semispinosus.
Proechimys goeldii: goeldii, steerei.
Proechimys canicollis.
Proechimys guyannensis appears to be more plastic than any other
species. In size of animal, width of aristiforms, color and number of
counterfolds in the cheekteeth, it shows marked response to varia-
tions in geographic conditions. Proechimys longicaudatus is appar-
ently less plastic; only the number of counterfolds shows marked
variation. Proechimys semispinosus varies much within its range.
Proechimys goeldii seems to be relatively uniform. Proechimys
canicollis shows relatively little variation throughout its range but
probably is divisible into two or more subspecies.
The primitive Proechimys probably was large with a short tail,
narrow aristiforms, strongly built skull, and five counterfolds in each
molariform tooth. Primitiveness here is inferred from characters
which now are of general occurrence in the whole group as opposed to
those restricted in geographic occurrence.
It is a curious fact that in this genus, populations from small
islands are more primitive than populations on the mainland. Ap-
parently a small population restricted to a small island tends to re-
vert to the primitive type. The homozygous condition will tend
toward a generalized genotype and the disappearance of secondary
biotypes. P. i. iheringi on the Island of Sao Sebastiao averages
larger, has thinner aristiforms, and a stronger skull than the same
subspecies on the mainland, and the cheekteeth usually have two
and three counterfolds. The same subspecies on the mainland has
no more than two counterfolds. Proechimys semispinosus gorgonae
and Proechimys semispinosus ignotus, living on Gorgona and San
Jose islands, respectively, are both characterized by large size, short
tails, strong and conspicuously ridged skulls, and cheekteeth fre-
quently with four and five counterfolds. On the mainland, closely
related subspecies, like P. s. panamensis, chiriquinus and gularis, far
Moojen: Brazilian Spiny Rats 317
less frequently have four counterfolds in more than one or two teeth.
More striking still is the population-sample of gularis from the island
of Llunchi, in the Rio Napo, eastern Ecuador. In it there is a higher
ratio of cheekteeth with four counterfolds than there is in the samples
from the banks of the river.
The two insular forms, P. s. gorgonae and P. s. ignotus, referred to
as primitive in the discussion above, have wide aristiforms, which is
contrary to what would be expected in a primitive Proechimys. Sup-
posing, however, as actually seems to be the fact, that narrowness of
the aristisforms depends on an increased number of genes, we deduce
that the population from the mainland, that gave rise to the popula-
tions of the islands, did not have all of the genes necessary to make
the artistiforms narrow. In fact the subspecies known on the main-
land, near the aforementioned islands, have wide aristiforms.
Another point which favors the idea that narrow aristiforms result
from an increased number of genes is that, generally, the aristiforms
are narrow in any species whose geographic range is extensive and
relatively uniform.
Proechimys goeldii is the species which has the largest number of
characters that are judged to be primitive, and it may be the oldest
stock. P. semispinosus, P. longicaudatus and P. guyannensis may
have been derived from an early splitting of the genus or they may
have branched off the main stem at different times. P. canicollis,
however, seems clearly to be an offshoot of P. guyannensis; canicollis
shows greater resemblance to guyannensis than to any other species.
P. g. vacillator is another close relative of P. guyannensis with the
number of counterfolds almost as much reduced as in P. canicollis.
Conceivably, vacillator is a full species, but the reduction in number
of counterfolds in the teeth more probably expresses only one extreme
of a gradient, as will be discussed below.
SUBSPECIFIC VARIATION IN THE SUBGENUS
PROECHIMYS
In spite of the lack of specimens from areas in which Proechimys
certainly occurs, it is evident that the genus has great plasticity and
that the number of subspecies will be greatly increased as additional
material is studied. Only perfunctory examination of samples from
outside the area of Brazil shows me that there are several unnamed
subspecies there. My impression is that Allen's trinitatis, of Trini-
dad, the genotype of Proechimys, will eventually be split.
There are two main lines of subspeciation in Proechimys guyan-
nensis. The one south of the Amazon River includes P. g. boliv-
2—3343
318 University of Kansas Publs., Mus. Nat. Hist.
ianus, in Bolivia, P. g. villicauda, and P. g. ribeiroi occurring on the
divide of the headwaters of the Amazon and Parana rivers, in
Brazil, and P. g. hyleae in the lower TapajOz and P. g. nesiotes in
the lower Tocantins. All six subspecies have a large number of
counterfolds in the molariform teeth. In these six subspecies, p4
has four counterfolds and the lower molars have three each. To-
ward the northeastern coast the number of counterfolds decreases
to three in p4 and to two in the lower molars, as in P. g. arescens,
P. g. leioprimna and P. g. oris.
In northern South America, north of the Amazon River, the sub-
species with the greatest number of counterfolds is P. guyannensis
warreni (known from only the Demerara River area) ; p4 has four
counterfolds and the lower molars have three each. The number
decreases in all the adjacent populations: P. g. guyannensis, in the
Guianas, P. g. trinitatis, and P. g. urichi (going westward from the
Guianas to Venezuela) have the counterfolds reduced to three in
p4, but the lower molars still have the same number of counter-
folds, namely, three, although there is a tendency for them to co-
alesce; farther west, on the coast, the number decreases to three
counterfolds in p4 and to only two in the lower molars as in P. g.
guairae and P. g. mincae. Subspecies south of the coast show the
same reduction of counterfolds, P. g. cherriei and P. g. oconnelli
being examples ; P. g. ochraceus and P. g. poliopus have the reduc-
tion carried to the upper molars, M3 having usually only two
counterfolds ; P. g. chrysaeolus in the valley between the Madalena
and the Cauca rivers seems to be somewhat isolated and shows re-
version to three counterfolds in the lower molariform teeth ; directly
southward of the range of P. g. warreni the number of counterfolds
decreases to three in all lower cheekteeth (population at Ayan-
Tepuy, southern Venezuela), and then to three in p4 and to two
in the lower molars, as in P. g. arabupu on the Brazilian side of
Mount Roraima, and the reduction is extended to the upper molars
in P. g. vacillator.
On the north bank of the Amazon, the only population of P. g.
hyleae known to me (from Obidos) has four counterfolds in p4 and
three in the lower molars; P. g. riparum, from Manaus, also on the
north bank of the Amazon, has three counterfolds in p4 and two
counterfolds in the lower molars. P. g. hyleae occurs also on the
south bank of the Amazon. P. g. riparum, therefore, may be the
northern part of the southern cline, instead of the southern end of
the northern cline.
Moojen: Brazilian Spiny Rats 319
The whole picture, as outlined above, may be explained by as-
suming that the species P. guyannensis differentiated somewhere on
the Central Plateau of South America, with three counterfolds in
each upper molariform tooth, four counterfolds in the lower pre-
molar and three counterfolds in the lower molars. The species
might have extended its range to the Guianas and then all the bio-
types with reduced number of counterfolds might have slowly de-
veloped by natural selection. The gradient is, broadly, from sub-
species with greater number of counterfolds in more humid areas,
to a gradually lessening number of counterfolds in less humid areas.
Proechimys longicaudatus is limited in the south to the head-
waters of the Parana River drainage, where the subspecies P. I.
roberti and P. I. longicaudatus are found. The species ranges north-
ward through the Tapajoz drainage, with P. I. leucomystax in the
headwaters and P. I. boimensis in the lower course. To the north-
west and west the species is represented in Bolivia by P. I. securus;
P. I. elassopus, P. I. simonsi, P. I. pachita, and P. I. hendeei occur in
Peru and P. I. brevicauda in Peru and Brazil; and P. I. nigrofulvus
occurs in Ecuador. Again in P. longicaudatus it seems that the num-
ber of counterfolds follows a gradient from more humid areas with
four counterfolds in p4, as seen in nigrofulvus, pachita, simonsi,
elassopus and brevicauda, decreasing to three or four in securus, to
three in longicaudatus, but with m3 having only two counterfolds in
leucomystax and roberti. P. I. boimensis, widely separated in the
lower Tapajoz (no samples being known from the intervening-
range) may be the end of a cline started by leucomystax with only 2
counterfolds in m3 and ending to the nortrrward with four counter-
folds in m3. Over the same area the counterfolds in p4 increase
from 3 to 4.
Of Proechimys goeldii I have had inadequate material but there
seems to be a similar gradient in it which may be traced from P. g.
steerei to P. g. goeldii. P. g. steerei has four counterfolds in more
upper molars than occurs in the other subspecies.
Proechimys semispinosus has its wide range in the mountainous,
western area of South America, the headwaters of the Amazon
drainage and northward in Central America and the nearby Pacific
Islands. In these populations a gradient may exist in the number of
counterfolds wrhich is varied in every population. The highest num-
ber seems to occur in the populations from northern Peru and
Ecuador, decreasing from there in all directions, except in the
Panamanian and Columbian islands. In gross examination, it seems
that the size of the animals increases to the northwards.
320 University of Kansas Publs., Mus. Nat. Hist.
SPECIFIC VARIATION IN THE SUBGENUS
TRINOMYS
Some specific characters are duplicated in each of the two sub-
genera; that is to say, there are some parallel developments and
they give the common generic stock its biotypical variability.
Among these parallel developments are the width of the aristiforms,
the amount of pigment in the agouti-colored seti forms, and the shape
of the nasal bones. Other characters, however, appear in one sub-
generic group and not in the other. The specific variation will be
discussed separately for each subgenus.
The aristiforms are narrow and soft in P. dimidiatus and in the
other species are wide and stiff, and on the outer thighs and rump
some are light-colored. P. albispinus has the maximum number of
light-colored aristiforms; they are present over the sides and back.
This species has, however, a type of aristiforms unique in the genus —
the clavate type. The tail is longer in P. iheringi and P. setosus than
in P. dimidiatus and P. albispinus; the longer type is associated with
a penicillate tip suggesting an adaptation to arboreal habit. The
skull and nasals are longer in P. dimidiatus and P. iheringi than in
P. setosus and P. albispinus. In the latter two species the longi-
tudinal dorsal outline of the skull is conspicuously convex as opposed
to slightly convex in the other two species. The palate is longest in
P. dimidiatus and P. iheringi extending posteriorly to the level of
the second molars ; it is slightly shorter in P. setosus and shortest in
P. albispinus where it does not extend behind the level of the first
molars. The incisors are opisthodont in P. dimidiatus and P.
iheringi and orthodont in P. setosus and P. albispinus and even
proodont in one part of the last species.
The molariform teeth have a large number of counterfolds in
both P. dimidiatus and P. iheringi, although the number varies but
little in the first species and much in the second. The variation
in P. iheringi decreases in populations of increasingly more northern
geographic distribution; in both P. setosus and P. albispinus the
number of counterfolds is greatly reduced ; there is only one in most
specimens of P. albispinus. The incisive foramen is small and nearly
round in P. dimidiatus, larger and elongate in P. iheringi, very
narrow and fissurelike in both P. setosus and P. albispinus.
The characters of Trinomys, as briefly outlined above, seem to be
the result of one original species having split first into four species
which provide a gradient for certain characters. Subsequently one
of these four species, P. iheringi, split into six subspecies and another
Moojen: Brazilian Spiny Rats 321
gradient, parallel to the first, and involving the same characters, is
to be seen.
The interrelationship among the species is evident, not only be-
cause they have the same subgeneric characters, but because the
full species themselves provide successive steps in a stairway of in-
creasing specialization from P. dimidiatus to P. albispinus.
Morphologically P. dimidiatus and P. iheringi are sometimes dif-
ficult to distinguish, especially on the basis of cranial features.
Nevertheless close attention to the small, nearly round, incisive
foramen of P. dimidiatus versus the larger, more elongate foramen
in P. iheringi will permit separation of the two. However, the two
species live in the same place and one is led to infer that there may
be greater differences in their physiology than in their morphology.
In fact Dr. H. W. Laemmert, from the Service de Estudos e
Pesquisas Sobre a Febre Amarela in Brazil, informs me that while
P. dimidiatus was highly susceptible to the virus of yellow fever
(18 out of 24 with virus in circulation), P. iheringi showed a lower
rate of susceptibility (3 out of 25 with virus in circulation). P.
longicaudatus roberti, belonging in the other subgenus, showed no
susceptibility at all.
At Teresopolis, Estado do Rio de Janeiro, the two species were
found in two different forests, only a few kilometers apart, but
dimidiatus lived at a higher elevation, where the humidity was re-
markably higher. Naturally the plant associations were different
in the two forests. This seeming ecological adaptation of the two
kinds of Proechimys may explain why P. iheringi ranges farther
north; the forests to the northward are less humid.
One of the four species, P. setosus, subspecies elegans, was used
by Winge (1941:80, 82) as representative of the genus Proechimys
when he was estimating the relationships of that genus. Because
Cercomys, with four crests in each of its cheekteeth, was, on other
grounds, regarded by him (op. cit.: 80) as". . . the most prim-
itive genus within the group.", and because he noted in P. s. elegans
4 crests in P4 and in some first molars, he concluded that Proechimys
was "very closely related to Cercomys." His conclusion seems to
be correct, but actually other species of Proechimys (subgenus Trin-
omys) , for example, P. dimidiatus, have four or more crests in each
cheektooth, and, therefore, may be considered as more closely re-
lated to Cercomys than is P. setosus. If a large number of crests
indicates primitiveness, P. dimidiatus, always with four, is more
primitive than any other species in the subgenus Trinomys. Also,
the large skull, long hind foot, short tail and thin aristiforms of P.
dimidiatus, in my opinion, are primitive characters.
322 University of Kansas Publs., Mus. Nat. Hist.
SUBSPECIFIC VARIATION IN THE SUBGENUS
TRINOMYS
One of the species of Trinomys, Proeehimys iheringi, is here sub-
divided into six subspecies which show a clinal variation. P. i. iher-
ingi, in the southernmost part of the range of the species (Ilha de
Sao Sebastiao), has three counterfolds in the upper cheekteeth of
almost every young specimen but one of these counterfolds, since it
is small, very shallow, and disappears after little wear, is probably
in the process of disappearance; all lower cheekteeth have two
counterfolds or, rarely, m3 has only one. P. i. bonafidei is the next
subspecies northward, where it was collected at 850m altitude
(Fazenda Boa Fe). This subspecies still has two counterfolds in
all the upper cheekteeth; only 3 out of 16 specimens fail to have
these counterfolds coalesced in one or more of the teeth. In the
lower cheekteeth the coalescence is evident in 18 per cent of the
specimens. P. i. gratiosus, from Floresta da Caixa Dagua (alt.
750m), geographically is well removed from bonafidei (more than
two degrees north), and no samples were obtained from the interven-
ing area. It shows such great reduction in the counterfolds that the
existence of intermediate populations is clearly suggested. Every
upper cheektooth of this subspecies has the two counterfolds coa-
lesced and in 40 per cent of the specimens M3 has only one counter-
fold; in the lower cheekteeth 60 per cent of the specimens have only
one counterfold in m3. P. i. panema, occurring approximately 100
kilometers to the northward of P. i. gratiosus (lowland form), has
one counterfold in M3 in only 20 per cent of the specimens but the
lower third molar has only one counterfold in 80 per cent of the spec-
imens. In P. i. denigratus, from about 3 degrees north of the range
of P. i. panema, the reduction is proportionately greater: P4 now is
the only upper cheektooth with two counterfolds in every specimen;
all molars tend to have only one; p4 has also two counterfolds but
all lower molars have only one.
The relative size of the tail also varies in a cline from south to
north. Its length is approximately 87 per cent of the length of the
head and body in P. iheringi; 88 per cent in bonafidei; 99 per cent
in gratiosus; 100 per cent in panema; and 103 in denigratus.
One of the subspecies, P. i. paratus, however, seems to be com-
pletely out of the dental cline. It was collected in the near prox-
imity of the type locality of P. i. gratiosus, at an elevation of 120m
lower. This subspecies has two counterfolds in all molariform teeth
Moojen: Brazilian Spiny Rats 323
and only one of the two specimens known shows these counterfolds
coalesced in P4 and Ml. The sample, 2 specimens, is too small to
be trustworthy; hence it is impossible satisfactorily to account for
the break in the clinal variation. Conceivably two full species are
involved, but I prefer at present to defer decision on this problem
until such time as more evidence is accumulated.
P. setosus is poorly represented, both of the available skins being
faded. Furthermore, no type locality is known for the subspecies
P. s. setosus.
P. albispinus has only two known subspecies: P. a. albispinus,
living in a region of higher humidity, is slightly the darker and has
subapical zones of the setiforms on the sides Ochraceous-Tawny ;
P. a. sertonius, living in a much drier region, has the same subapical
zone Ochraceous-Buff. The number of specimens of P. a. sertonius
is so few that no gradient can be detected, even if one exists.
TAXONOMIC CHARACTERS
Size and Proportions of External Parts
Absolute size of head and body, tail, hindfoot and ear are useful
in distinguishing subgenera and subspecies and to some extent in
differentiating species.
The length of head and body is large to medium in Proechimys
and medium to small in Trinomys. The tail is long to medium in
Trinomys and short in Proechimys. The longest tail, 242 mm, is
found in P. i. denigratus, and the shortest tail, 123 mm, in P. g.
steerei. The relative length of tail also provides gradients or clines.
In every species, males surpass females in average size. Never-
theless, the largest animals are usually females. How this para-
doxal fact is to be accounted for, I am not sure, but it may be that
the animals grow as long as they live and that females have more
chances to survive longer since the care of the young keeps them
closer to shelter.
Color. — Upper parts vary from Buckthorn Brown to Ochraceous-
Buff. Dark color ordinarily is correlated with an environment of
higher degree of humidity and light color with lower humidity.
However, species may be found in similar conditions of humidity
but differing in color. Proechimys albispinus albispinus, for exam-
ple, a light-colored form, is found in areas where the rainfall
averages 1,000 to 1,500 mm of annual precipitation, in the isohygra
of 80 per cent relative humidity. These conditions actually are
similar to those where P. dimidiatus, of darker color, is found. The
324 University of Kansas Publs., Mus. Nat. Hist.
subspecies albispinus, however, ranges mostly over a dry area and
the fact that it occurs also in a moist area without appreciable
change in color is difficult to explain.
Insular populations are usually darker or richer in color than
corresponding continental populations. On a small island, uni-
formity of environment and inbreeding may be responsible for an
accumulation of characters for richness of color.
Pelage
The pelage provides most useful taxonomic characters. Except-
ing the vibrissiform hairs, all of the elements of the pelage have a
common feature, the flattened shape. The hair constellation (cf.
Toldt, 1935) on the upper and lateral surfaces is composed of hairs
of two main types: aristiforms (guard hairs) and setiforms (over
hairs).
The aristiforms are wide, strong, and have the dorsal (= ante-
rior) margins raised, forming a wide shallow longitudinal groove
on the dorsal face of the hair. The tip is a filament that usually
is lacking in aristiforms which are especially strong. Wear prob-
ably removes these tips. The aristiforms have the bases whitish or
grayish and the amount of pigment gradually increases distally to
a dark brown or blackish shade. On the dorsal and lateral surfaces
of the head the aristiforms are small and narrow but gradually
increase in length and width caudad on the animal. The maximum
development is reached in the middorsal region, from where they
decrease in size and number toward the lateral surfaces or caudad.
This decrease in the development of the aristiforms, however, is not
uniformly gradual. Generally, the aristiforms become increasingly
conspicuous in a middorsal band, but they extend to the sides and
onto the outer sides of the thighs ; the band narrows rapidly on the
rump. In the subgenus Trinomys, where the aristiforms attain
their maximum development, they are still strong and conspicuous
on the rump and sometimes around the base of the tail. In
Proechimys the aristiforms do not extend caudad from the hips.
Also, in Trinomys, besides the ordinary lanceolate type, there are
some aristiforms on the dorsal surface with a clavate shape; the
base is wide and the distal part narrow. This parallels the condi-
tions in the pelage of the most spiny species in the genus Echimys,
Echimys chrysurus (Lichtenstein).
The recently named subspecies Proechimys cayennensis hop-
lomyoides Tate, 1939, shows an extraordinary development of the
Moojen: Brazilian Spiny Rats 325
aristiforms on the back and sides such as occurs in the genus
Hoplomys. Actually the small bulla, wide basisphenoid and tooth
structure add to the possibility of hoplomyoides being a true Hop-
lomys, and worn teeth might have been responsible for the difficulty
which Tate had in allocating the form to the proper genus. How-
ever, the narrow braincase is more nearly like that of Proechimys
than that of Hoplomys. The intermediate nature of hoplomyoides
argues for including the genus Hoplomys as a subgenus of Pro-
echimys.
Species with narrow aristiforms have a rather soft and flexible
pelage, while those with wide aristiforms have harsh, spiny pelage.
The aristiforms vary in width from 0.45 to 1.3 mm, depending upon
the species or subspecies.
Animals with narrow aristiforms tend to have a more or less
uniform coloration throughout the dorsal parts. The blackish distal
parts of the aristiforms regularly interline the ground color made by
the subapical zone of the setiforms. If, on the contrary, wide
aristiforms occur, the dorsal surface is conspicuously marked by
the wide blackish lines among spots of color formed by the subapical
zones of the setiforms. No clinal variation was detected in width
of aristiforms but geographic variation in width was noted; for
example, the subspecies of P. iheringi differ in this respect.
The setiforms are narrow and flattened but are without pro-
nouncedly raised margins. The setiforms are usually bicolored on
the dorsal and lateral surfaces of the animals, with a subapical zone
of some reddish-brown color, like Ochraceous-Orange or Ochraceous-
Buff. They are whitish or gray on the basal parts and gradually
blacken toward the tip, but have a reddish subapical zone. Common
exceptions to this pattern are setiforms without subapical zones;
these appear on the dorsal surface among setiforms which are
normal in possessing distinctive subapical zones. Also there are
setiforms without blackened tips on the lateral surfaces. Due to
their relative abundance and subapical color, these setiforms are
responsible for the dominant color on the upper parts. Like the
aristiforms, they are longer and wider in the middorsal region of the
animal and are gradually less developed on the remainder of the
upper parts. Actually there is more than one type of setiform in
the hair constellation; they vary in length, width and color. At-
tention was not given, however, to every type of setiform.
The ventral surface of the body and the inner sides of the legs
are uniformly covered by short setiforms, thinner and more sparsely
326 University of Kansas Publs., Mus. Nat. Hist.
distributed on the inner side of the legs. These setiforms are usually
uniformly white in color or, sometimes, the distal parts are buff or
more richly colored.
Vibrissiforms are scattered on the dorsal and lateral surfaces of
the body, and in penicillate arrangements on the head. They are
lunger than the pelage proper, have a nearly circular cross-section
and are blackish in color.
Skull
The absolute size of the skull is proportionate to bulk of the body.
The supraorbital and parietal ridges are especially developed in the
P. semispinosus group, where they extend across the parietals to the
interparietals. In all members of the subgenus Proechimys, these
ridges extend onto the parietal region. In Trinomys, however, they
do not extend so far posteriorly as the parietal, but only onto the
squamosal.
The rostrum varies from slender to stout. Elongate rostra are
common in Proechimys; Trinomys has a short blunt rostrum.
The infraorbital foramen commonly has a ventral groove for
nerve transmission in many forms of Proechimys but Trinomys
almost always lacks this groove. Presence or absence of the groove
is a subspecific character in the subgenus Proechimys.
The jugals are dorso-ventrally wide in Trinomys except in the
species P. setosus. In Proechimys a dorso-ventrally narrow jugal
is the rule, but P. canicollis has an especially wide jugal. A
postorbital process appears on the jugo-squamosal suture and is here
called postorbital process of the zygoma. In Proechimys it is more
or less weakly developed and shows no variation of systematic
worth. In Trinomys, on the other hand, this process varies in a
clinal way (P. iheringi) and stages of the gradient characterize
populations of subspecific rank.
Linear and spatulate shape of the humular process of the pterygoid
constituted specific characters for Thomas, but there is so much in-
dividual variation in the shape of this process in almost every popu-
lation that it has not been used in the present account.
The mesopterygoid finterpterygoid) fossa in almost every speci-
men extends anteriorly to the level of Ml or M2 in Trinomys, and
to M3 in Proechimys. Exceptions may occur, as in P. hendeei,
where the fossa extends to the level of M2.
Incisive Foramen
The shape and dimensions of the incisive foramen long have been
recognized as providing specific characters. Large size of the fora-
men is probably correlated with the requirement for a large amount
Moojen: Brazilian Spiny Rats 327
of moisture reaching Jacobson's organ in the nasopalatine space;
the moistening of the sensory epithelium is certainly involved.
There seems to be a certain correlation between small size of the
incisive foramen and high degree of humidity in the environment.
Shapes and dimensions of the foramen appear as simple or multiple
biotypes and provide characters which can be employed to dif-
ferentiate subspecies, species and even subgenera. Usually a char-
acter, say a general shape, occurs in nearly all populations of a
given subspecies but the particular shape seems to be more closely
correlated with ecological conditions, especially humidity. Animals
which live far away from large rivers usually have larger foramina
than animals which live close to rivers.
Both the premaxilla and the maxilla develop processes which
form a sheath for the vomer. This vomerine sheath forms a bridge
which longitudinally crosses the incisive foramen; the structure of
this bridge varies widely. Sometimes the maxillary part is not de-
veloped and the sheath is incomplete posteriorly; sometimes this
maxillary part is very slender and merely touches the premaxillary
part. The premaxillary part, however, is always well developed.
Teeth
Considered by itself the variation in the tooth pattern can lead
to erroneous conclusions as to differentiation of species, because the
number of folds on the occlusal face of a tooth and the depth of
certain folds may be subject to great individual variation as shown
by examination of more than one large series of specimens of the
same kind, age and sex from a single locality. Also there are geo-
graphic gradients or clines, in number of folds. Nevertheless the
variation in number of folds, when measured at sufficient intervals
along a cline, may provide quantitative characters useful in differ-
entiating subspecies.
The main fold involves both the occlusal face of the tooth and
the side wall. The counterfolds, which are smaller counterparts of
the main fold, in most instances also implicate the wall of the tooth
opposite to that marked by the main fold, but are to be seen mostly
on only the occlusal face of the tooth. Unerupted teeth with the
crowns unworn and other teeth which had barely broken through
the gums were ground down to permit the making of drawings of
the surfaces at different levels. This study revealed that the main
fold is deepest in the wall of the tooth. The development of the
main fold varies in two different ways: in all samples from south-
eastern and eastern Brazil it is strongly developed, deeply grooves
328 University of Kansas Plbls., Mus. Nat. Hist.
/../
^_^wSf^55a**>7»,
Fios. 2-17. Second left upper molar of the two subgenera Proechimys and Trinomys. All
X 8. Anterior border of tooth is at the top of each figure (Nos. 9 and 17 excepted). Note
especially that main fold is short in Proechimys and long in Trinomys.
Figs. 2-9. Proechimys {Proechimys) semispinosus liminalis, female, MN no. 6243, Rio
Quichito. Fig. 2, unworn crown. Figs. 3-8, cross sections at 0.5 mm. intervals, showing
changes in the main fold and counterfolds at increasing depths as the tooth was ground down.
Fig. 8 is 3 mm. below surface shown in fig. 2. Fig. 9, posterior view with proximal end of
the tooth open showing basal ends of folds. Later in life the proximal end closes and three
roots are formed.
Figs. 10-17. Proechimys (Trinomys) iheringi denigratus, female, SEPFA no. 17060,
Mata do Ribeirao da Fortuna. Figs. 10-16 corresponding to figs. 2-8. Fig. 17, postero-
ventral view with proximal end of the tooth open and part of walls cut away, showing basal
ends of folds. Later in life, as in Proechimys, the proximal end closes and three roots are
formed.
Moojen: Brazilian Spiny Rats 329
the tooth through its crown and, in younger individuals, completely
divides the occlusal surface of the tooth. As use wears down the
crown, the main fold soon becomes separated from the opposite
wall and then gradually shortens toward its basal portion. In the
other type, common to animals of all the remaining part of the
range of the genus, the main fold is rather short, never reaching the
opposite wall. In this case, however, one of the counterfolds usually
appears almost opposite the main fold in such a way that in non-
erupted or just-erupted teeth the main fold and one counterfold
may be connected by a shallow groove that may give the impression
of extension of the main fold and, therefore, lead to false interpre-
tations. Closer examination shows that the counterfold which ap-
parently meets the main fold is really situated anteriorly or pos-
teriorly to it, in upper or lower teeth respectively. One subspecies
in the subgenus Trinomys differs from the general characteristics of
the subgenus in sometimes showing a small main fold in P4 whereas
it is large in all other cheekteeth. The structural differences here
mentioned in the main fold were never before recognized. There-
fore, the meaning of "quadrilaminate" pattern, "three outer folds"
or "three inner folds" (of authors) is not consistent insofar as the
two groups are concerned.
Writers have more or less tacitly admitted three as the usual
number of counterfolds present in the upper molariform teeth. El-
lerman, for example (1940:117), states: "Upper cheekteeth nor-
mally with three outer and one inner folds each, these soon becom-
ing isolated as islands. A few species, which will be discussed
below, vary slightly in pattern." Thomas (1921:140) erected the
subgenus Trinomys on the basis of the upper molariform teeth hav-
ing only three laminae. Actually a meticulous study of widely
varying samples shows that the number of counterfolds may vary
from one to five, the usual number being three or four. One of the
most important facts to record on this subject is that young spec-
imens with slightly worn molariform teeth are more apt than either
adult or younger specimens to show the maximum number of coun-
terfolds. Usually nonworn teeth show rounded crests and valleys
of different depth. For example, it is common to see one continuous
groove giving the impression of a main fold crossing the occlusal
surface transversely. The slightest wear of the occlusal surface,
however, shows that really there are two valleys instead of one.
The two are the main fold and one counterfold. In this case, it
would be easy to confuse the two types of teeth, one with the main
330 University of Kansas Publs., Mtjs. Nat. Hist.
fold short and the other with the main fold extending almost all the
way across the occlusal surface.
Some of the counterfolds are especially shallow and tend to dis-
appear in an early stage of wear, and adult individuals may have
these folds completely worn away. Advanced wear usually develops
a cuplike occlusal surface with only the remains of the main fold and
also remains of one or more counterfolds represented by small enamel
islands (Figs. 2 to 17) . In the form Proechimys iheringi iheringi, for
example, every tooth shows three counterfolds in the upper molari-
form teeth of individuals in which the wear is not advanced. This
number, however, is less in all or part of the molariform teeth of
older individuals.
Adjacent counterfolds may appear to be coalesced in many in-
stances. Coalescence is more likely to be seen in species where a
wider variation in the number of the counterfolds is involved and it
appears as a gradient in the reduction of the number of counterfolds.
Of great importance, as a general feature of molariform teeth, is
the relative size as related to the geographical distribution, showing,
again, a natural division in the genus. In all forms of southeastern
Brazil the premolars are larger than the first molars, the first molars
are larger than the second molars, and the second molars are larger
than the third molars. The forms from central and northern Bahia,
Brazil, have the molariform teeth more or less the same size. The
forms from the remaining part of the area occupied by the genus have
premolars smaller than the first molars, the first molars smaller than
the second molars, but the second molars larger than third ones.
HABITS
P. dimidiatus was studied in the field and laboratory. P. dimidia-
tus in captivity showed regular diurnal activity, coming out of the
nest for food at intervals. Individuals were fed a cereal mixture and
nuts of small size. The animals usually buried the nuts in the sand
of the outer cage. While holding the nut with the mouth and front
feet, the animal patted the sand rapidly, thus burying the nut, and
it then pushed more sand over the place with the front feet.
Sometimes the emergence from the nest is followed by a long yawn-
ing and stretching ceremony. The animal spreads the fore and hind
legs widely apart, while the back is curved down and the head and
tail turn upward. Then one of the hind legs is stretched backward
and, at the same time, the mouth is opened widely and the tail is
moved in an undulatory fashion. The operation may be repeated
using the other legs, or not.
Moojen: Brazilian Spiny Rats 331
P. dimidiatus was regularly found in climax forest. The best
shelter and nesting ground was usually under boulders, commonly
not farther than 10 meters from water. The entrance to the nest was
kept clean. No more than two adult animals (male and female)
were captured in the same shelter, and only a few times were young
captured in the same place with adults. Nesting places were located
also at the bases of trees and near fallen logs where litter accumu-
lates.
Records of animals kept in captivity show that the species dimidi-
atus survives more than two years. Specimen MN no. 5448 J1 was
adult when captured by the Servicos de Estudos e Pesquisas sobre
a Febre Amarela on December 5, 1938, and died on January 17, 1942.
Therefore, it lived for more than 1,139 days.
Proechimys dimidiatus, in Rio de Janeiro, as well as P. i. bonafidei
and P. i. iheringi which live in the same region, were found breeding
from September to November and from March to May. Proechimys
longicaudatus roberti, in Anapolis, Goiaz, was found breeding from
July to November and from January to March. P. g. hyleae and
P. g. leioprimna in the lower Tapajoz and lower Tocantins rivers,
Para, were found breeding in January.
It seems that in the Central Plateau and southeastern Brazil,
Proechimys has two litters per year, one in the early spring and a
second in the late summer. The number of young per litter varies
from 1 to 5, although 2 is the usual number.
CHANGES WITH AGE
Juveniles. — The animals are born with a thick pelage of thin aristi-
forms and thin setiforms. The color is uniformly blackish brown.
The nose, hands, feet, ears and tail are pinkish ; P4 and Ml are al-
ready erupted and the second molars are included in the bony alveoli.
The incisors are orthodont; the rostrum is short and the braincase is
wide. The posterior part of the skull is greatly curved dorsally. No
change is noticed in the pelage before the second molars erupt and
become functional.
Adolescents. — As soon as the second molars become functional,
the pelage starts molting on the back. The thin aristiforms are still
in place but the aristiforms of the adult pelage may be noticed
growing under them in an oval patch which extends from behind
the shoulders caudad to the hips. At that age the first, agouti-
colored aristiforms appear on the mystacial region, immediately
behind the vibrissiforms. The rostrum gradually lengthens and
the braincase appears to become less inflated.
X12
University of Kansas Publs., Mus. Nat. Hist.
By the time the third molars erupt, the aristiforms start showing
among the setiforms which are now changing to agouti color in the
same area on the back, while the thin aristiforms of the juvenal
stage disappear. The agouti setiforms are appearing also poste-
riorly from the mystacial region to the sides of the head and neck
and, at the same time, on the frontal region. The patch of glossy
aristiforms and setiforms on the back is sharply differentiated from
the dull juvenal pelage of the sides and rump. In a later stage the
area of agouti setiforms on the sides of the neck extends to the
outer sides of the arms and finally reaches the area on the back
where the agouti setiforms were already developed.
Adults. — When the third molars become functional, the agouti
setiforms are in place except for those on the upper sides of the
neck. The aristiforms have now extended over their normal area
of distribution. As soon as the third molars show wear, the pre-
molars and first molars have the counterfolds isolated in the oc-
clusal surfaces as enamel islands. Wearing gradually isolates all
counterfolds.
Senile individuals. — Progression of wear soon eliminates the signs
of the shallowest counterfolds from the occlusal surface and finally
the tooth is reduced to a short crown with a cuplike occlusal sur-
face completely filled with dentine. Only the main fold usually
remains; it is more or less reduced in size.
From the records of the Servigos de Estudos e Pesquisas sobre a
Febre Amarela, the following data for males of Proechimys longi-
caudatus roberti were obtained:
Age
Number
of
cheek-
teeth
Weight
in grams
Length
of head
and
body
Length
of tail
Length
of
hind
foot
One day old
2
2
2
3
3
4
4
20.5
26.0
85.0
115.0
180.0
200.0
360.0
70
110
150
175
195
223
230
53
60
105
120
135
158
170
24
17 days old
28
Juvenile
39
Adolescent
45
Adolescent (older)
45
Adult
48
Senile individual
48
The weights and measurements (except for one- and 17-day-old
animals) represent averages of specimens of the different ages.
Moojen: Brazilian Spiny Rats
333
Genus Proechimys J. A. Allen
Genotype. — Echimys trinitatis Allen and Chapman, by original
designation.
Proechimys Allen and Chapman, 26 December 1899, Bull. Amer. Mus. Nat.
Hist., 12(20) :264, orig. description; Tate, 1935, Bull. Amer. Mus. Nat. Hist.,
68(5) :398; Ellerman, 1940, The families and genera of living rodents, Brit.
Mus. (Nat. Hist.), 1:115.
General characters. — Muriform echimyids of medium size; pelage with
flattened and lanceolate and sometimes clavate aristiforms, varying greatly in
width and distributed over most of the dorsal surface from shoulders to hips
or base of tail; setiforms also flattened, evenly distributed throughout; entire
ventral surface and inner sides of legs white or, rarely, invaded by some buffy
color; tail shorter than, equal to, or slightly longer than, head and body, bi-
colored and with a few bristles between scales, sometimes heavily penicillated ;
feet long and narrow; pollex rudimentary; hallux shorter than fifth toe; ears
wide and long; mammae 2 — 1 = 6.
Skull. — Generally elongate and strongly built, with supraorbital ridges well
developed, frequently extending across parietals toward occipital region;
zygomatic arches variable in depth, always with postorbital process; infra-
orbital foramen with or without lower groove for transmission of nerve; in-
cisive foramen usually large; vomerine sheath complete or incomplete; mes-
opterygoid fossa extending forward at least to plane of third molars; bullae
large; angular process of mandible turned upward.
Figs. 18-21. Occlusal views of the upper left and lower right molariform teeth of the
two subgenera of the genus Proechimys. Anterior end of the tooth row at the top of draw-
ing. All X 6.
Figs. 18-19. Proechimys (Proechimys) goeldii steerei, sex ?, USNM no. 105537, "Hyutana-
ham." Upper teeth at left (fig. 18).
Figs. 20-21. Proechimys (Trinomys) dimidiatus, male, MN no. 6256, Pedra Branca.
Teeth. — Incisors opisthodont, orthodont or proodont, not grooved; upper
molariform teeth with a main internal fold and one to five external counter-
folds which usually appear as enamel islands in worn teeth, these counterfolds
barely implicating the lateral wall; lower molariform teeth with folds as in
the upper molariform teeth except that they are reversed and the number of
internal counterfolds is usually fewer in the molars.
3—3343
334 University of Kansas Publs., Mus. Nat. Hist.
Artificial Key to the Subgenera and Species
1. (a) Tail less than 90 per cent of head and body; aristi-
forms not evident on outer thighs and rump; skull
with ridges across parietals; size of upper cheekteeth
increasing from P4 to M2 ; main fold small.
subgenus Proechimys, 2
(b) Tail 90 per cent or more of head and body; aristi-
forms evident on outer thighs and rump; skull with
no ridges across parietals; size of upper cheekteeth
decreasing from P4 to M3; main fold large.
subgenus Trinomys, 5
2. (a) One or more upper molars with four counterfolds . . . 3
(b) Upper molars with no more than three counterfolds, 4
3. (a) Aristiforms wide (more than 0.7 mm).
P. semispinosus, p. 342
(b) Aristiforms narrow (less than 0.7 mm) ... .P. goeldii, p. 338
4. (a) Aristiforms wide (0.9 mm or more), or narrow (0.6
to 0.7 mm) but then with only two counterfolds in
each lower molar P. guyannensis, p. 355
(6) Aristiforms narrow (0.5 to 0.65 mm) but with one or
more lower molars having three counterfolds.
P. longicaudatus, p. 346
5. (a) Aristiforms narrow (0.5 mm) and limber; no differ-
entiated light-colored aristiforms on outer thighs and
rump; incisive foramen short and widest posteri-
orly P. dimidiatus, p. 371
(6) Aristiforms 0.6 mm or more and stiff; differentiated
light-colored aristiforms on outer thighs and rump;
incisive foramen elongated and constricted posteri-
orly 6
6. (a) Skull large, more than 50 mm in total length, incisors
opisthodont P. iheringi, p. 373
(b) Skull small, less than 49 mm in total length, incisors
orthodont or proodont 7
7. (a) No clavate aristiforms, tail with white tip, P. setosus, p. 384
(b) Clavate aristiforms among the ordinary ones, tail
without white tip P. albisyinus, p. 388
Moojen: Brazilian Spiny Rats
335
0
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70 40
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20
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Fig. 22. Map showing the distribution of the two subgennra of the genus Procchimys.
336
University of Kansas Publs., Mus. Nat. Hist.
Fig. 23. Map showing the geographic ranges of four species of the genus Proechimys.
Moojen: Brazilian Spiny Rats
337
Fio. 24. Map showing the geographic ranges of four species of the genus Proechimys.
338 University of Kansas Publs., Mus. Nat. Hist.
Subgenus Proechimys J. A. Allen
General characters. — Pelage with lanceolate aristiforms limited to an area
on the dorsal surface between the shoulders and the hips; length of tail less
than 90 per cent of length of head and body; skull with conspicuous ridges;
extension of supraorbital ridges always evident on parietals; infraorbital fora-
men usually with separate groove for transmission of nerve; palate usually
extended posteriorly as far as third molars; incisors opisthodont; molariform
teeth with a small main fold, never extended transversely to opposite wall in
occlusal surface of tooth; usually one counterfold anterior to main fold in
upper molariform teeth and posterior to main fold in lower molariform teeth;
premolars usually smaller than first molars, first molars smaller than second
molars but second molars larger than third molars.
Proechimys goeldii Thomas
General characters. — Size large; tail short; aristiforms narrow and soft,
usually concealed in pelage by setiforms; general color of upper parts some
tint of orange, gradually becoming lighter on sides with no conspicuous, dark
longitudinal band on back; feet dark; ventral surface of body and inner side
of legs white but sometimes with some buff locally; skull broad and strongly
built but not conspicuously ridged; zygomatic expanse great and rostrum not
elongate; incisive foramen narrow; bullae large and inflated; upper molari-
form teeth with three to four counterfolds, M3 ordinarily with four; lower
premolars with four, and molars with three, counterfolds.
Proechimys goeldii steerei Goldman
Proechimys steerei Goldman, Proc. Biol. Soc. Washington, 24:238, 28
November 1911 (original description); Goldman, 1912, Proc. Biol. Soc.
Washington, 25:186; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :400;
Ellerman, 1940, The families and genera of living rodents, Brit. Mus.
(Nat, Hist.), 1:119; Osgood, 1944, Zool. Ser. Field Mus. Nat. Hist.,
29(13) :204.
Type locality. — Hyutanaham, Upper Purus, Ldbrea, Amazonas, Brazil.
Type: United States National Museum, no. 105535, adult male; collected in
1901 by Prof. J. B. Steere.
Range. — Known only from the type locality and Porto Velho.
Diagnosis. — Upper parts Mars Orange on back, grading to Ochraceous-
Tawny on sides; zygomatic breadth narrow; nasals short; incisive foramen
narrow and short; vomerine sheath complete and thick; upper molars usually
with four counterfolds.
Pelage. — Aristiforms on middorsal region: Grayish basally, gradually
blackening toward tip; total length, 16 to 19 mm; maximum width, 0.5 mm.
Setiforms on middorsal region: Grayish on basal third, gradually blackening
toward tip but interrupted by a Mars Orange, subapical zone 1.5 mm long;
total length, 16 to 19 mm ; maximum width, 0.06 mm. Setiforms on outer
thighs: Whitish basally, gradually blackening toward tip but interrupted by
Orange Rufous or Ochraceous-Tawny, subapical zone; total length, 14 to 16
mm; maximum width, 0.05 mm.
Skull. — Large and strong; rostrum rather pointed posteriorly; supraorbital
ridges not much expanded and extending across anterior half of parietals;
Moojen: Brazilian Spiny Rats
339
Figs. 25, 27. Proechimys goeldii steerei, sex ?, "Hyutanaham," USNM no. 105537. X 1.
Figs. 26, 28. Proechimys goeldii goeldii, female, AMNH no. 3748S. X 1.
Figs. 29, 30. Proechimys goeldii steerei, sex ?, USNM no. 105537, "Hyutanaham." X 1.
Figs. 31, 32. Proechimys goeldii goeldii, female, AMNH no. 37488, Fazenda Paraiso. X 1.
340 University of Kansas Publs., Mus. Nat. Hist.
infraorbital foramen without groove for transmission of nerve, or groove obso-
lete ; zygomatic arches slender ; postorbital process of zygoma involving mostly
squamosal; incisive foramen short and narrow (4.5x2.5 mm) with margins
almost parallel or tapering gradually caudad and extending toward palate as
ridges; posterior margin of incisive foramen approximately 2.5 mm anterior
to premolars; vomerine sheath complete, with both elements well-developed;
mesopterygoid fossa never extending anterior to middle of M3; bullae large,
well inflated and with shallow grooves.
Teeth. — Upper molariform teeth: P4 with three counterf olds ; upper molars
with four counterf olds each or, less commonly, three. Lower premolars with
four counterf olds; lower molars with three each.
Comparisons. — From P. g. goeldii, steerei differs in: Back and sides with
more reddish; narrower interorbitally and across zygomata; palatilar length
less and nasals shorter; maxillary part of vomerine sheath thicker; usually
four instead of three counterfolds in M3.
Remarks. — This subspecies is clearly related to P. goeldii. One
skull from Porto Velho, Rio Madeira, Guapore, Brazil (CNHM no.
21558) may belong to an unnamed subspecies but is provisionally
included here.
In the field notes of Professor Joseph Beal Steere, an entry for
no. 72 reads: "Big white bellied wood rats x two young found in
nest of grass on the ground with the two young — much darker
young female." No. 77 in his field notes corresponds to the type
specimen.
Specimens examined. — Total number, 4, from Brazil, as follows: Amazonas. Labrea,
Hyutanaham, 3 (USNM); Territ. Guapore, Porto Velho, 1 (CNHM).
Proechimys goeldii goeldii Thomas
Proechimys goeldii Thomas, June 1905, Ann. Mag. Nat. Hist., 15 (ser.
7) :587, (orig. descr.) ; Thomas, 1912, Ann. Mag. Nat. Hist., 9 (ser. 8):89;
Thomas, 1920, Ann. Mag. Nat. Hist., 6 (ser. 9) :277, Tate, 1935, Bull.
Amer. Mus. Nat. Hist., 68(5) :400; Osgood, 1944, Zool. Ser. Field Mus.
Nat. Hist., 29(13) :199.
Proechimys cayennensis goeldii, Ellerman, 1940, The families and gen-
era of living rodents, Brit. Mus. (Nat. Hist.), 1:121.
Type locality. — Santarem, Santarem, Para, Brazil. Type: British Museum
(Nat. Hist.), no. 5.1.25.6, adult female; presented by Dr. E. A. Goeldi.
Range. — Margins of the Amazon, between Jamunda and Tapajoz rivers.
Diagnosis. — Upper parts Ochraceous-Tawny ; wide across zygomata; nasals
of moderate length; incisive foramen long and narrow; vomerine sheath com-
plete but maxillary part slender; first and second upper molars with four
counterfolds.
Pelage. — Arislijorms on middorsal region: Whitish basally and gradually
blackening toward tip which is extended as long, thin filament; total length,
22 to 24 mm; maximum width, 0.5 mm. Setijorms on middorsal region:
Gray basally, gradually blackening toward tip but interrupted by Ochraceous-
Tawny, subapical zone 3.3 mm long; total length, 19 to 21 mm; maximum
Moo j en: Brazilian Spiny Rats 341
width, 0.06 mm. Setijorms on outer thighs: Whitish basally, gradually black-
ening toward tip but interrupted by Ochraceous-Tawny, subapical zone 3 mm
long; total length 14 to 16 mm; maximum width, 0.04 mm.
Skull. — Large and strong; nasals pointed posteriorly; supraorbital ridges
moderately developed and extended caudad across anterior third of parietals;
zygomatic arches strong; postorbital process of zygoma involving mostly
squamosal; incisive foramen elongate and narrow (5 to 6.5x2.3 mm) with
margins more or less parallel and raised to form ridges which extend poste-
riorly to within 3 mm of plane of premolars; vomerine sheath complete, with
maxillary part thin and extended caudad as medial crest; mesopterygoid fossa
extending forward as far as posterior faces of second molars or slightly short
thereof; bullae large and inflated.
Teeth. — Molariform teeth large, P4-M3 averaging more than 9 mm in
length. Upper molariform teeth: P4 and M3 with three counterfolds ; Ml
and M2 with four counterfolds each. In lower teeth, p4 with four counter-
folds and each molar with three counterfolds.
Comparisons. — Differences from P. g. steerei are given in the account of that
subspecies.
Remarks. — Specimens from the type locality were not available.
Specimens from Fazenda Paraiso, Faro, were relied upon as repre-
sentative of the subspecies. These agree with the type according to
Thomas (1912:89). However, the skin of the type was changed in
color by preservative (Thomas, 1905:587) and the best skin he
saw was from Faro (1912:89).
Thomas (1920:277) applied the name goeldii also to specimens
from Manacaparu, a place a short distance above Manaus on the
SolimSes (Amazon) River and from Acajutuba, near Manaus, on
the Negro River. In referring to these specimens (2 from Mana-
caparu and 2 from Acajutuba) Thomas (loc. cit.) said "Five molar
laminae are frequently, if not invariably, present among these
specimens." He did not, however, mention whether or not the
number of laminae was constant in both M2 and M3. One speci-
men from Acajutuba, in the collection of Museu Nacional (MN
no. 1973 $ ) , actually has five laminae in M3, but the specimens in
the American Museum from Faro agree absolutely with Thomas'
original description of goeldii.
Osgood (1944:199) doubted that goeldii was a valid species.
Evidence that Osgood's doubt was unjustified is furnished by the
fact that Thomas (1912:89) pointed out that his specimen from
Faro agrees with the type. Likewise, my two specimens from Faro
agree with the type insofar as it has been described. Thomas
(1912:89) mentioned two additional skulls from the type locality
which, he stated, agree with the type which was received from the
Museu Goeldi, Para.
342 University of Kansas Publs., Mus. Nat. Hist.
Specimens examined. — Total number, 4, from Brazil as follows: Para, Faro, Faro, Fazenda
Paraiso, 2 (AMNH) ; Amazonas, Manaus, Manaus, 1 skull (AMNH); Amazonas, Manaus,
Acajutuba, 1 (MN).
Additional record*. — Total number, 7 (British Museum), from Brazil, as follows: Para,
Santarem, Santarem (Thomas, 1912:89; 1920:277), 3; Amazonas, Manaus, Acajutuba
(Thomas, 1920:277), 2; Manacaparu, Manacaparu (Thomas, 1920:277), 2.
Proechimys semispinosus (Tomes)
General characters. — Size large; tail short and hairy; aristiforms wide and
stiff, especially well-developed on back; general color on upper parts some
shade of ochraceous, usually much darker on back and forming a conspicuous
38
Figs. 33, 30. Proechimys semispinosus liminalis, female, MN no. 6253, Rio Quichito.
Type. X 1.
Figs. 34, 37. Proechimys semispi?iosus amphichoricus, male, AMNH no. 77020, Mount
Duida. Type. XL _
Figs. 35, 38. Proechimys semispinosus kermiti, female, AMNH no. 37124, Lower Rio
Solimoes. Type. X L2 (from photograph).
Moojen: Brazilian Spiny Rats
343
dorsal band; feet dark; ventral surfaces and inner sides of legs white; skull
elongate and strong with ridges well developed; incisive foramen long and
narrow; bullae large; usually four counterfolds in M3 and M2; usually three
but sometimes four counterfolds in Ml and even P4; lower premolar with
four and lower molars with three counterfolds.
33 \|
Figs. 39, 40. Procchimys semispinosus liminalis, female, MN no. 6253, Rio Quichito.
Type. X 1.
Figs. 41, 42. Proechimys semispinosus amphichoricus, male, AMNH no. 77020, Mount
Duida. Type. X 1.
Figs. 43, 44. Proechimys semispmosus kermiti, female, AMNH no. 37124, Lower Rio
Solimoes. Type. X 1-2 (from photograph).
Proechimys semispinosus liminalis subspecies nova
Type locality. — Rio Quichito, affluent from the south of the Javari River,
near Benjamin Constant, Benjamin Constant, Amazonas, Brazil. Type:
Museu Nacional, no. 6253. adult female, collected in August, 1942, by E.
Parko.
Range. — Known only from the type locality.
Diagnosis. — Color uniformly dark, setiforms marked with Ochraceous-
Tawny; skull wide across zygomata; nasals short; prepalatilar part of skull
long; incisive foramen long and narrow; vomerine sheath incomplete or com-
plete; M2 and M3 almost always with four counterfolds; Ml more rarely
with four counterfolds.
Pelage. — Aristijorms on middorsal region: Gray basally, gradually blacken-
ing toward tip which is generally extended as a filament; total length, 21 to
23 mm; maximum width, 0.9 to 1 mm. Setiforms on middorsal region: Gray
basally, gradually blackening toward tip but interrupted by Ochraceous-
344 University of Kansas Publs., Mus. Nat. Hist.
Tawny, subapical zone 3 mm long; total length, 22 to 24 mm; maximum
width, 0.06 mm. Setiforms on outer thighs: Whitish basally, gradually
blackening toward tip but interrupted by Ochraceous-Buff, subapical zone
2.5 mm long; total length, 13 to 15 mm; maximum width, 0.08 mm; some
with gray base, blackening gradually toward tip, without any subapical zone;
some with Light Ochraceous-Tawny, subapical zone.
Skull. — Large and strongly built throughout; supraorbital ridges expanded
and thick, extending, in old specimens, across parietals to anterior angles of
interparietals; interparietal ridges always conspicuous; rostrum elongated;
nasals blunt posteriorly; zygomatic arches strong; infraorbital foramen with
weakly-developed groove for transmission of nerve; postorbital process of
zygoma involving mostly squamosal; incisive foramen averaging 6x2.7 mm,
widest in middle part and posteriorly constricted, with raised margins which
do not extend across maxillae as ridges; posterior margin of incisive foramen
approximately 1.5 mm anterior to plane of premolars; vomerine sheath in-
complete or, sometimes, complete but always with maxillary part slender;
mesopterygoid fossa not extending forward past centers of third molars;
bullae moderately developed.
Teeth. — Upper molariform teeth: P4 always with three counterf olds ; Ml
with three counterfolds in 9 of 10 specimens and four counterfolds in re-
mainder; M2 with four counterfolds in 7 specimens, three counterfolds in
remainder; M3 with four counterfolds in 6 specimens, three counterfolds in
remainder. Lower premolar always with four, and molars with three, counter-
folds.
Comparisons. — From P. s. semispinosus, liminalis differs in: darker color;
wider aristif orms ; greater percentage of upper molars with four counterfolds.
From P. s. amphichoricus, liminalis differs in: lighter upper parts of almost
uniform color instead of with conspicuous, blackish, middorsal, longitudinal
band; more strongly built skull; longer incisive foramen; vomerine 6heath
usually incomplete instead of always complete.
Specimens examined. — Total number, 10 (MN) from the type locality.
Proechimys semispinosus amphichoricus subspecies nova
Type locality. — Mount Duida, Esmeralda, Amazonas, Venezuela; altitude
325 m. Type: American Museum of Natural History, no. 77020, adult male;
collected 16 October 1920 by Olalla Bros.
Range. — Headwaters of Negro and Orinoco rivers, along boundary of Brazil
and Venezuela.
Diagnosis. — Color dark, blackish on middorsal area; subapical zone of seti-
forms on back Buckthorn Brown, but many with distal parts black; skull
broad across zygomata; nasals long; prepalatilar area of skull long; incisive
foramen long and narrow; vomerine sheath complete; upper molars usually
with four counterfolds but P4 usually with only three.
Pelage. — Aristijorms on middorsal region: Grayish basally, gradually black-
ening toward tip; total length, 18 to 20 mm; maximum width, 0.8 to 1.0 mm.
Setiforms on middorsal region: Gray basally, gradually blackening toward tip
but interrupted by a light (16 i), Buckthorn Brown, subapical zone 2 mm
long; total length, 18 to 22 mm; maximum width, 0.03 mm. Most of them,
Moojen: Brazilian Spiny Rats 345
however, whitish basally, gradually blackening toward tip without any dis-
tinctively-colored, subapical zone; total length, 24 to 26 mm; maximum width,
0.5 mm. Setijorms on outer thighs: Whitish basally, gradually blackening
toward tip but interrupted by an Ochraceous-Buff, subapical zone 3.5 mm
long; black tip short; total length, 17 to 19 mm; maximum width, 0.05 mm.
Skull. — Large and slender; rostrum elongate; nasals bluntly pointed pos-
teriorly; supraorbital ridges thick (but not expanded) and extending across
parietals but almost obsolete in middle part of parietals; infraorbital fora-
men with weakly-developed groove for transmission of nerve; postorbital
process of zygoma involving mostly squamosal; incisive foramen 5.5x2.8 mm
wide in anterior third, with margins constricted posteriorly and extending as
ridges approximately 2 mm beyond posterior margin of incisive foramen;
posterior margin of incisive foramen approximately 2.5 mm anterior to pre-
molars; vomerine sheath complete with maxillary part weak and premaxillary
part extending posteriorly beyond middle of incisive foramen; mesopterygoid
fossa extending forward as far as middle of M3; bullae well inflated and
elongated.
Teeth. — P4 with four counterfolds in one of five specimens and with three
in remainder; Ml with four counterfolds in three of five specimens and with
three in remainder; M2 with three counterfolds in one specimen and with
four in all four remaining specimens; M3 always with four counterfolds.
Lower premolars with four counterfolds and lower molars with only three.
Comparisons. — The subspecies is easily distinguishable from P. s. annularis
by : larger number of black setiforms on back, forming an almost black longi-
tudinal band; more elongate skull; larger and longer bulla; longer incisive
foramen which is more constricted posteriorly.
Specimens examined. — Total number, 6 (AMNH), as follows: Venezuela, territ. Ama-
zonas, Esmeralda, Mt. Duida, altitude 325 m., 4; Venezuela, territ. Amazonas, Rio Cassi-
quiare, Quemapure, 1 ; Brazil, Amazonas, Sao Gabriel, Rio Uaupes or Caiari, Tatu, 1.
Proechimys semispinosus kermiti Allen
Proechimys kermiti Allen, 30 December 1915, Bull. Amer. Mus. Nat.
Hist., 34(22) :629 (orig. descr.) ; Allen, 1916, Bull. Amer. Mus. Nat. Hist.,
35(30) :569; Tate, 1935, Bull. Amer. Mus. Nat. Hist, 68(5) :400; Eller-
man, 1940, The families and genera of living rodents, Brit. Mus. (Nat.
Hist.), 1:119.
Type locality. — Lower Rio Solimoes (up the Solimoes 50 to 60 miles on the
north bank of the river), Manacaparu, Amazonas, Brazil. Type: American
Museum of Natural History, no. 37124, adult female ; collected 20 April, 1914,
by Leo E. Miller (Roosevelt Brazilian Expedition).
Range. — Known only from type locality.
Diagnosis. — Upper parts Tawny, with darker longitudinal band on back,
gradually becoming Ochraceous-Buff on sides; zygomata widely spread; nasals
long; incisive foramen long; vomerine sheath incomplete; only M3 with four
counterfolds.
Pelage. — Aristiforms on middorsal region: Grayish basally, gradually black-
ening toward tip; total length, 18 to 20 mm; maximum width, 0.8 mm. Seti-
forms on middorsal region: Grayish basally, gradually blackening toward tip
but interrupted by Tawny, subapical zone 2 mm long; total length, 18 to 20
346 University of Kansas Publs., Mus. Nat. Hist.
mm; maximum width, 0.06 mm; some blackened toward tip without siHj-
apical zone. Setiforms on outer thighs: Whitish basally, gradually blackening
toward tip but interrupted by Ochraceous-Buff, subapical zone 2.5 mm long;
total length, 18 to 20 mm; maximum width, 0.05 mm.
Skull— Large, elongate, and strongly built; rostrum not conspicuously
elongated; nasals bluntly pointed posteriorly; supraorbital ridges wide and
extending posteriorly across parietals almost to level of interparietal; infra-
orbital foramen with moderate development of groove for transmission of
nerve; zygomatic arches slender; postorbital process of zygoma involving
mostly squamosal; incisive foramen 6.5 mm long and 2.7 mm wide, wider in
anterior third and gradually constricted posteriorly, with margins extended
toward palate as ridges; vomerine sheath incomplete, maxillary part thread-
like; mesopterygoid fossa extending forward as far as anterior third of m3;
bullae large and well inflated.
Teeth. — P4 with three counterf olds ; M3 with four counterf olds ; Ml and
M2 with three counterf olds. Lower premolars with four counterf olds ; lower
molars with three counterfolds.
Comparisons.— Prom P. s. amphichoricus, kermiti differs in: upper parts
Tawny instead of Buckthorn Brown; incisive foramen longer and wider;
vomerine sheath incomplete; only M3 instead of usually all molars, with
four counterfolds. From P. s. liminalis, kermiti differs in : upper parts Tawny
instead of Ochraceous-Tawny; aristiforms narrower; M3 only, instead of
usually M2 and M3, with four counterfolds.
Specimens examined. — Only the type.
Proechimys longicaudatus (Rengger)
General characters. — Size medium to large; tail short; aristiforms Ions and
narrow; general color on upper parts Ochraceous-Buff to Ochraceous-Orange,
finely and uniformly lined with blackish and not forming evident dark band
on back; feet dorsally white or gray; underparts of body and inner sides of
legs white; skull elongate and slender with moderate ridges; incisive foramen
of medium size; vomerine sheath complete or incomplete; bullae large and
elongate; upper molariform teeth with three counterfolds; lower molariform
teeth with three counterfolds but commonly one or two molars have only two
although premolar may have four.
Remarks. — The identity of "Echimys longicaudatus Rengger"
can be ascertained only after samples have been collected in the
area indicated by Rengger: "unter dem ein und zwansigsten Breit-
engrade" in Paraguay. Of the samples available to me, those from
Urucum, in western Brazil, are geographically nearest the type lo-
cality. North of Urucum, both in Brazil and Bolivia, two species
of Proechimys live together and one of them is the same species as
that at Urucum. Of the two species found to the northward in
Brazil and Bolivia, the one that ranges farther south probably will
occur at the locality indicated by Rengger. Provisionally, there-
fore, the name longicaudatus is allocated to the Urucum sample (see
Osgood, 1944:198). In fact, the lack of a type specimen and the
Moojen: Brazilian Spiny Rats
347
general nature of Rengger's description make "Echimys longicau-
datus" a nomen vanum. If two species are found living together
in the region of northern Paraguay indicated by Rengger it prob-
ably will be impossible to be sure to which one his vague descrip-
tion applies.
The form from Urucum, to which the name Proechimys longi-
caudatus is here applied, is undoubtedly closely related to Pro-
echimys leucomystax Ribeiro, from Utiariti, on the Rio Papagaio
and also to P. roberti and P. boimensis, all from Brazil. P. longi-
caudatus is used as the name of the species because it is the oldest
of the four names.
Figs. 45, 48. Proechimys longicaudattts boimensis, male, MCZ no. 30881, Boim. X 1.
Figs. 46, 49. Proechimys longicaudatvs longicaudattts, male, AMNH no. 37085, Urucum.
X 1.
Figs. 47, 50. Proechimys longicaudatus leucomystax, male, AMNH no. 37509, Tapirapoa.
X 1.
348 University of Kansas Publs., Mus. Nat. Hist.
Figs. 51, 52. Proechimys longicaudatus roberti, male, MN no. 6233, Pouso Alto, Goiaz.
X 1.
Figs. 53, 54. Proechimys longicaudatus boimensis, male, MCZ no. 30881, Boim. X 1-
Figs. 56, 56. Proechimys longicaudatus longicaudatus, male, AMNH no. 37085, Urucum.
X 1.
Figs. 57, 58. Proechimys longicaudatus leucomystax, male, AMNH no. 37509, Tapirapoa.
X 1'
Figs. 59, 60. Proechimys longicaudatus roberti, male, MN no. 6233, Pouso Alto, Goiaz.
X 1.
Moojen: Brazilian Spiny Rats 349
Proechimys longicaudatus brevicauda (Giinther)
Echimys brevicauda Giinther, 1 April 1877, Proc. Zool. Soc. London for
1876, (49) :748, fig. 9.
Proechimys brevicauda Ihering, 1904, Rev. Mus. Paulista, S. Paulo,
6:422; Osgood, 1914, Zool. Ser. Field Mus. Nat. Hist., 10(12): 168;
Thomas, 1924, Ann. Mag. Nat. Hist., 13 (ser. 9):534; Thomas, 1927, Ann.
Mag. Nat. Hist., 19 (ser. 9):553; Thomas, 1927, Ann. Mag. Nat. Hist.,
20 (ser. 9):604; Thomas, 1928, Ann. Mag. Nat. Hist., 2 (ser. 10):262;
Thomas, 1928, Ann. Mag. Nat. Hist., 2 (ser. 10):292; Tate, 1935, Bull.
Amer. Mus. Nat. Hist., 68 (5):399; Osgood, 1944, Zool. Ser. Field Mus.
Nat. Hist., 29 (13) :201.
Proechimys cayennensis brevicauda Ellerman, 1940, The families and
genera of living rodents, Brit. Mus. (Nat. Hist.), 1:120.
Type locality. — Chamicuros, Rio Huallaga, Peru. Type: British Museum
(Nat. Hist.), no. 69.3.31.7,9; Lectoparatype : British Museum (Nat. Hist.),
no. 66.1.29.8, from Upper Amazons (E. Barttet), selected by Thomas (1900:
301).
Range. — Region of the headwaters and upper courses of the Jurua. and
Ucaiali rivers, eastern Peru and northwestern Brazil.
Diagnosis. — Upper parts Tawny and blackish without marked longitudinal
band on back; underparts buffy or white; aristiforms narrow; skull slender;
incisive foramen wide; vomerine sheath complete; molariform teeth with
three counterfolds, except p4 with four and m3 with only two.
Pelage. — Aristiforms on middorsal region: Blackish basally, gradually
blackening toward tip which is extended as a long filament; total length, 18
to 20 mm; maximum width, 0.65 mm. Setijorms on middorsal region: Gray
basally, gradually blackening toward tip but interrupted by Tawny, subapical
zone 1.2 mm long; total length, 19 to 21 mm; maximum width, 0.1 mm. Seti-
jorms on outer thighs: Whitish basally, gradually blackening toward tip but
interrupted by Ochraceous-Tawny, subapical zone 3 mm long.
Skull. — Slender but not elongated ; nasals tapering posteriorly ; interparietals
wide; supraorbital ridges not much extended and faintly shown across parie-
tals; jugals dorsoventrally "wide" (3.5 mm); postorbital process of zygoma
weakly developed; incisive foramen 5.5x3 mm, oval, with posterior borders
raised to form ridges which extend toward premolars; vomerine sheath com-
plete, with maxillary part laterally compressed and extended toward palate
as ridge ; mesopterygoid fossa extending forward as far as third molars ; bullae
large and well inflated.
Teeth. — Molariform teeth with three counterfolds, except p4 which has four
and m3 which has only two counterfolds.
Comparisons. — From P. I. longicaudatus, brevicauda differs in: upper parts
Tawny instead of Ochraceous-Buff ; lower premolar with four instead of three
counterfolds; m3 only, instead of both ml and m3, with two counterfolds.
From P. I. boimensis, brevicauda differs in: upper parts Tawny instead of
Ochraceous-Orange; aristiforms wider; m3 with two instead of three counter-
folds.
Specimens examined. — Total number, 3 (DZ), from Brazil, Amazonas, Joao Pessoa, Rio
Jurua.
4—3343
350 University of Kansas Publs., Mus. Nat. Hist.
Proechimys longicaudatus boimensis J. A. Allen
Proechimys boimensis Allen, 24 July, 1916, Bull. Amer. Mus. Nat. Hist.
35(27) :523; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :400; Ellerman,
1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.),
1:119.
Proechimys cayennensis Osgood, 1944, Zool. Ser. Field Mus. Nat. Hist.,
29(13) :199.
Type locality. — Boim, Rio Tapajoz, Santarem, Para, Brazil. Type: Ameri-
can Museum of Natural History, no. 37486. adult male; "October 10, 1911 (ex
Museu Goeldi)."
Range. — Along lower course of Tapajoz River.
Diagnosis. — Upper parts Ochraceous-Orange ; incisive foramen posteriorly
constricted; mesopterygoid fossa sharply pointed anteriorly; p4 with four
counterfolds, remaining molariform teeth with three counterfolds.
Pelage. — Aristijorms on middorsal region: Gray basally, gradually blacken-
ing toward tip which is extended as a long filament; total length, 16 to 18
mm; maximum width, 0.5 mm. Setiforms on middorsal region: Gray basally,
gradually blackening toward tip, but interrupted by short (1.5 mm), Ochrac-
eous-Orange, subapical zone; total length 18 to 20 mm; maximum width 0.09
mm. Setijorms on outer thighs: White basally, gradually blackening toward
tip but interrupted by Ochraceous-Orange, or Ochraceous-Buff, subapical zone
3.5 mm long; total length 15 to 17 mm; maximum width, 0.06 mm.
Skull. — Elongate and slender; rostrum slender and long; nasals evenly
pointed posteriorly; supraorbital ridges weak and barely reaching anterior-
most part of parietals; zygomatic arches slender; infraorbital foramen with
uroove for nerve-transmission obsolete; postorbital process of zj'goma weak
and involving mostly squamosal; incisive foramen 5 mm long and 2.5 to 3
mm wide, oval, with margins not much extended caudad as ridges; posterior
margin of incisive foramen approximately 2 mm anterior to premolars; vom-
erine sheath complete with maxillary part slender; mesopterygoid fossa sharply
pointed anteriorly and extending forward as far as anterior part of third molar;
bullae of medium size, smooth and more or less triangular in contour.
Teeth. — Crown length of upper molariform teeth 7 to 7.5 mm; all upper
molariform teeth with three counterfolds; lower premolar with four counter-
folds; lower molars with three counterfolds each.
Comparisons. — From three of the other four subspecies found in Brazil,
boimensis differs in having four, instead of three, counterfolds in the lower
premolars. Differences from P. I. brcvic.auda are indicated in the account of
that subspecies.
Remarks. — The material available from Boim, Tapajoz, is rather
poor and of no great significance. The Museu Nacional has one
specimen from Boim which agrees with the type in the American
Museum of Natural History. The Museum of Comparative Zoology
at Harvard College, however, has one specimen in its collection
(MCZ no. 30888 $ ), adult, also from Boim, in which the color
pattern is different although agreement with the type specimen is
shown in cranial characters and width and length of hairs. This
Moojen: Brazilian Spiny Rats 351
specimen is much darker than the other specimens (of orange-tint)
assigned to this subspecies. However, in other samples of this
species, similarly dark animals have been noted. It seems that the
orange tint is gained only in old age. Another specimen (MCZ no.
30878 $ ) agrees in all respects with the type of the subspecies but
the locality recorded on the label is Cameta, on the Tocantins River.
Possibly the subspecies has a range so wide as to include Cameta
but I suspect that the data on the label is incorrect as to locality.
The collector marked J on the label although the specimen is a 5 .
The mistake might have occurred through labeling of specimens at
a time later than that at which they were collected. The collector
was in both localities on more than one occasion.
Specimens examined. — Total number, 5, from Brazil, Para, as follows: Santarem, Rio
Tapajoz. Boim. 3 (AMNH type. MCZ 1, MN 1); Porto de Moz, Tauari, 1 (MCZ); Cameta
(?) 1 (MCZ).
Proechimys longicaudatus longicaudatus (Rengger)
Echimys longicaudatus Rengger, 1830, Naturgeschichte der Saeugethiere
von Paraguay, p. 236.
Loncheres myosuros Lichtenstein, 1832, Darstellung neuer odor wenig
bekannter Saugethiere, pi. 36 and text.
Echimys myosuros Is. Geoffroy Saint-Hilaire, 1840, Mag. Zool., Ann. 2
(ser. 2) :15, 17; Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20) :261.
Echimys cay ennen sis Pictet, 1841, Mem. Soc. Phvs. Hist. Nat., Geneve,
9:145; Waterhouse, 1848, Nat. Hist. Mammalia, 2:334.
Proechimys longicaudatus Thomas, 1901, Ann. Mag. Nat. Hist., 8 (ser.
7):532; Thomas, 1904, Proc. Zool. Soc. London, p. 240; Allen, 1916, Bull.
Amer. Mus. Nat. Hist., 35(30) :569; Tate, 1935, Bull. Amer. Mus. Nat.
Hist., 68(5) :400.
Proechimys cayennensis longicaudatus Ellerman, 1940, The families
and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:121; Osgood,
1944, Zool. Ser. Field Mus. Nat. Hist., 29(13) :198.
Type locality. — Northern Paraguay ("unter dem ein und swansigsten Brei-
tengrade"). Type: Apparently no type specimen was preserved.
Range. — Western Mato Grosso, Brazil, and northern Paraguay.
Diagnosis. — Upper parts almost uniformly Ochraceous-Buff; incisive fora-
men widest posteriori}^ ; vomerine sheath complete; p4 and m2 with three
counterfolds; ml and rn3 usually with two counterfolds.
Pelage. — Aristijorms on middorsal region: Dark gray, gradually blackening
toward tip that has long filament; total length 20 to 22 mm; maximum width
0.6 mm. Setijorms on middorsal region: Whitish basally, gradually blackening
toward tip but interrupted by Ochraceous-BufT, subapical zone; blackish tip
extended (3.5 mm) and thin; total length 19 to 20 mm; maximum width 0.00
mm. Setijorms on outer thighs: Whitish basally, progressively grayish or
blackish toward tip but interrupted by Light Ochraceous-Buff or Ochraceous-
Buff, subapical zone; total length 18 to 20 mm; maximum width 0.06 mm.
Skull. — Slender; supraorbital ridge notably raised; bullae large, elongate,
smooth and inflated; jugals narrow; postorbital process of zygoma of medium
size and constructed entirely of jugal; incisive foramen wide and large (5.5 x
352 University of Kansas Publs., Mus. Nat. Hist.
3 mm), being wider posteriorly than anteriorly and with posterior margins
raised; vomerine sheath complete, maxillary part slender and laterally flat-
tened; mesopterygoid fossa extending forward as far as centers, or anterior
margins, of third molars; posterior palatine foramina on plane with posterior
faces, or centers, of second molars.
Teeth. — Upper molariform teeth always with three counterfolds. Lower
molariform teeth: p4 and m2 always with three counterfolds, sometimes the
two anterior ones coalesced in m2; ml with three counterfolds in one specimen
(33 per cent) ; m3 with only two counterfolds in all specimens.
Comparisons. — Differences from P. I. leucomystax and P. I. roberti are given
in the accounts of those subspecies.
Specimens examined. — Total number, 3 (2 AMNH, 1 CNHM), from Brazil, Mato Grosso,
Corumbd, Urucum.
Proechimys longicaudatus leucomystax Ribeiro
Proechimys leucomystax Ribeiro, May, 1914, Commissao de linhas
telegraphicas estrategicas de Matto Grosso ao Amazonas, Annexo no. 5.
Hit. Nat., Zool., Mammiferos, p. 42, pi. 24 (orig. descr.) ; Tate, 1935,
Bull. Amer. Mus. Nat. Hist., 68(5) :400; Ellerman, 1940, The families and
genera of living rodents, Brit. Mus. (Nat. Hist.), 1:119.
Type locality. — Utiariti, Rio Papagaio, Diamantion, Mato Grosso, Brazil.
Type: Museu Nacional, no. 2212, adult, skull only, collected on 5 May 1909,
by Prof. A. Miaranda Ribeiro, is here designated lectotype. See remarks.
Range. — Serra dos Parecis, Mato Grosso, Brazil.
Diagnosis. — Ochraceous-Buff , richly lined with blackish, on upper parts ; some
setiforms completely blackened distally; incisive foramen regularly ovoid;
vomerine sheath incomplete; upper molariform teeth and lower premolar with
three counterfolds; m2 with three counterfolds but ml and m3 usually with two
counterfolds.
Pelage. — Aristijorms on middorsal region: Grayish basally, gradually black-
ening toward tip which is extended as a long filament; total length 21 to 22
mm; maximum width 0.65 mm. Setiforms on middorsal region: Whitish
basally, gradually blackening toward tip but interrupted by Ochraceous-Buff,
subapical zone; some setiforms grayish basally and gradually blackening to-
ward distal portion without any colored, subapical zone; total length 20 to
22 mm, maximum width 0.06 mm. Setiforms on outer thigh: White basally,
gradually becoming gray toward tip but interrupted by Light Ochraceous-Buff,
subapical zone; some setiforms gray basally and blackening toward tip, but
interrupted by Ochraceous-Buff, subapical zone; total length 15 to 18 mm;
maximum width 0.08 mm.
Skull. — Slender; supraorbital ridges conspicuous; bullae large, smooth and
inflated, with slight, transverse groove; jugals narrow; postorbital processes of
zygomata small and involving only squamosal; incisive foramen of medium
size but narrow (5x2.5 mm), regularly oval and with margins uplifted pos-
teriorly; vomerine sheath incomplete but maxillary part projecting forward and
sometimes almost reaching premaxillary part; mesopterygoid fossa reaching
forward as far as centers of third molars; posterior palatine foramina at plane
of posterior faces of second molars or slightly anterior thereto.
Moojen: Brazilian Spiny Rats 353
Teeth. — Upper molariform teeth with three counterfolds. Lower molariform
teeth: p4 and m2 always with 3 counterfolds; ml and m3 with 2 counterfolds.
Comparisons. — From P. I. longicaudatus, leucomystax differs in : upper parts
richly lined, instead of scarcely lined, with blackish ; incisive foramen narrower,
and regularly oval instead of widest anteriorly.
Remarks. — Proechimys leucomystax was described mainly on the basis of the
Utiariti specimen, here designated lectotype of the species. The specimen from
the Juina River is younger, as stated by Ribeiro in his description. Ribeiro
mentions the skin of the specimen from Utiariti as "em muito mao estado" and
I presume it was discarded as it has not been found in the collection of the
Museu Nacional, Brazil.
Specimens examined. — Total number, 6, from Brazil, Mato Grosso : Caceres, Salto
Sepotube, 2 (MN) ; Caceres, Tapirapoa, Rio Sepotuba, 2 (AMNH) ; Diamantino, Utiariti,
Rio Papagaio, 1 skull (MN) ; Diamantino, Rio Juina, 1 (MN).
Proechimys longicaudatus roberti Thomas
Proechimys roberti Thomas, December, 1901, Ann. Mag. Nat. Hist.,
8 (ser. 7):531 (orig. descr.) ; Thomas, 1904, Ann. Mag. Nat. Hist., 14 (ser.
7):195; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):140; Tate, 1935,
Bull. Amer. Mus. Nat. Hist., 68(5) :400.
Proechimys cayennensis roberti Ellerman, 1940, The families and gen-
era of living rodents, Brit. Mus. (Nat. Hist.), 1:121.
Type locality. — Rio Jordao, Araguari, Minas Gerais, Brazil; 960 meters alt.
Type: British Museum (Nat. Hist.), no. 1.11.3.62, old male, collected
8 August, 1902, by A. Robert; original number, 705.
Range. — Western Minas Gerais and southern Goiaz.
Diagnosis. — Color almost uniformly Ochraceous-Orange on upper parts;
setiforms long and narrow; incisive foramen long and wide; vomerine sheath
usually complete; upper molariform teeth and lower premolar with three
counterfolds; m3 with two, and ml and m2 with two or three, counterfolds.
Pelage. — Aristijorms on middorsal region: Whitish basally, gradually black-
ening toward tip; total length, 21 to 23 mm; maximum width, 0.6 mm. Seti-
forms on middorsal region: Whitish on basal half, gradually blackening to-
ward tip but interrupted by an Ochraceous-Orange, subapical zone 6 to 7 mm
long; total length 25 to 30 mm; maximum width, 0.05 mm. Setiforms on
outer thighs: Whitish on basal half, gradually becoming gray and then black-
ish toward tip but interrupted by wide, Ochraceous-Buff, subapical zone, the
tip being brownish or blackish; total length, 22 to 23 mm; maximum width,
0.04 mm.
Skull. — Slender; supraorbital ridges bent outward and making sharp angle
at point of frontosquamosal suture, then continuing backward parallel to each
other and extending across parietals; squamosal taking small part in supra-
orbital ridges; bullae large (11x8 mm), inflated, with shallow depressions;
incisive foramen not especially long but wide (5x3 to 3.5 mm), widest in
anterior third and constricted posteriorly; vomerine sheath usually complete,
with maxillary part reduced to slender threadlike process or, less commonly,
missing; mesopterygoid fossa extending forward as far as centers of third
molars; zygomatic arches strong with jugals of medium dorsoventral width
(approximately 3.5 mm) ; postorbital process of zygoma weakly developed
354
University of Kansas Publs., Mis. Nat. Hist.
and involving only squamosal; posterior palatine foramina on plane of pos-
terior margins of first molars or slightly anterior thereto.
Teeth. — Upper molariform teeth usually with three counterfolds (92.5 per
cent of 42 specimens) ; M2 with four counterfolds in 5 per cent of specimens;
Fig. 61. Map showing the geographic ranges of the subspecies of three species of the
subgenus Proechimys in Brazil.
M3 with two counterfolds in 2.5 per cent of specimens. Lower molariform
teeth: p4 usually with three counterfolds (97.5 per cent of 39 specimens);
rarely with four (2.5 per cent) ; ml with three counterfolds in 58 per cent and
two counterfolds in 42 per cent of specimens; m2 with three counterfolds in
61 per cent and two counterfolds in 39 per cent of specimens; m3 always with
onlv two counterfolds.
Moojen: Brazilian Spiny Rats 355
Comparisons. — From P. 1. boimensis, roberti differs in: wider incisive fora-
men; lower premolar with three, and one or two lower molars with only two,
counterfolds. instead of lower premolar with four and all lower molars with
three counterfolds. From P. longicaudatus, roberti differs in: upper parts
Ochraeeous-Orange instead of Ochraeeous-Buff; incisive foramen wider in
posterior third than in anterior third.
Remarks. — This subspecies seems to be adapted to forests of post-
climactic conditions which is probably typical of most valleys and
margins of the rivers in southern Goiaz and western Minas Gerais.
It was found in Goiaz usually in riparian forests with climactic as-
sociations or in some advanced stage of the sere. The animals also
make incursions into nearby open areas or crops of corn.
Specimens examined. — Total number, 52, from Brazil, as follows: Minas Gerais, Aragnari,
Rio Jordao (affluent of Parnaiba), 960 meters alt., 2 (1 CNHM, 1 DZ) ; Goiaz, Anapolis,
1010 meters alt., 38 (MN); Goiaz, Pouso Alto, 768 meters alt., 11 (MN); Goiaz, Tio Sao
Miguel, 2 (MN).
Proechimys guyannensis (E. Geoffroy)
.1/ us guyannensis E. Geoffroy Saint-Hilaire, 1803, Catalogue des mam-
miferes du Museum d'Histoire Naturelle, Paris, p. 194.
Echimys cayennensis Desmarest, 1817, Nouv. Diet. Hist. Nat., Paris,
nouv. ed., 10:59.
Proechimys cayennensis Allen, 1899, Bull. Amer. Mus. Nat. Hist.,
12(20) :261, 264; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :399; El-
lerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat.
Hist.), 1:120.
General characters. — Size medium to large ; tail usually short ; aristiforms
narrow to wide; general color of setiforms on back ranging from Tawny to
Ochraceous-Buff and becoming gradually lighter on sides; no conspicuous dark
longitudinal band on back; upper parts of hands and feet white to light
brown; underparts white, including inner sides of legs; skull elongate and not
conspicuously ridged; vomerine sheath complete or incomplete; upper pre-
molar with three counterfolds and molars with two or three ; lower premolar
with three or four counterfolds, and lower molars with two or three.
Proechimys guyannensis villicauda subspecies nova
Type locality. — Tapirapoa, Rio Sepotuba, Cdceres, Mato Grosso, Brazil.
Type: Museu Nacional, no. 1932, adult male (color faded); collected on 2
February, 1909, by Prof. A. Miranda Ribeiro; original number, 788 A.
Range. — Serra dos Parecis, headwaters of Paraguai and Tapajoz rivers.
Diagnosis. — Aristiforms wide and stiff; general color on upper parts Ochra-
ceous-Orange ; incisive foramen long; vomerine sheath incomplete or complete;
lower premolar with four counterfolds, remaining molariform teeth with three
counterfolds.
Pelage. — Aristiforms on middorsal region: Whitish basally, gradually be-
coming gray toward tip, with distal fourth blackish and ending as a long fila-
ment; total length, 22 to 23 mm; maximum width, 1 mm. Setiforms on mid-
dorsal region: Whitish on basal half, gradually blackening toward tip but inter-
rupted by Ochraeeous-Orange, subapical zone 5 mm long; total length, 26 to 30
356 University of Kansas Publs., Mus. Nat. Hist.
mm; width, 0.04 to 0.12 mm. Setiforms on outer thighs: Whitish on basal half,
gradually blackening toward tip but interrupted by Ochraceous-Buff to Ochra-
ceous — Orange subapical zone; total length 20 to 23 mm; width, 0.03 to 0.18 mm.
Skull. — Strong ; supraorbital ridges raised and extending across anterior fourth
of parietals; nasals elongate; bullae rounded, inflated, with shallow grooves;
postorbital process of zygoma weakly developed and constructed entirely of
jugal; incisive foramen elongate and narrow (5.5x2.5 mm), posteriorly con-
stricted with posterior margins elevated above surface of bones; vomerine
sheath incomplete or complete but, when complete, with maxillary part filiform
and delicate; mesopterygoid fossa extending forward as far as middle, or even
anterior, parts of third molars; posterior palatine foramina on plane of centers,
or even anterior faces, of second molars.
Teeth. — Upper molariform teeth with three counterfolds. Lower molariform
teeth: premolar with four counterfolds, molars with three counterfolds, some-
times with two folds coalesced in center of tooth.
Comparison. — From P. g. ribeiroi, and P. g. bolivianus, villicauda differs in
wider aristiforms. From P. g. ribeiroi, villicauda further differs in: larger and
wider incisive foramen; vomerine sheath incomplete or complete instead of
always complete and thick. From P. g. bolivianus, villicauda differs in : lower
premolars always, instead of rarely, with four counterfolds, and m3 always with
three counterfolds instead of usually with only two counterfolds.
Specimens examined. — Total number, 4, from Brazil, Mato Grosso, as follows : Cdceres.
Tapirapoa, 3 (MN) ; Diamantino, Rio Papagaio, Utiaritl, 1 (AMNH).
Fig. 62. Proechimys guyannensis villicauda, male, MN no. 1932, Tapirapoa. Type X 1.
Fig. 63. Proechimys guyannensis ribeiroi, male, MN no. 1935, Rio Doze de Outubro.
Type. X 1.
Fig. 64. Proechimys guyannensis hyleae, male, MCZ no. 30887, Tauari. Type. X 1.
Fig. 65. Proechimys guyannensis nesiotes, male, CNHM no. 19496, Ilha de Manapiri.
Type. X 1.
Fig. 66. Proechimys guyannensis leioprimna, female, CNHM no. 19503, Cameta. Type.
X 1.
Fig. 67. Proechimys guyannensis oris, male, CNHM no. 19495, Providencia. X 1.
Fig. 68. Proechimys guyannensis arescens, male, CNHM no. 26440, Fazenda Inhuma.
Paratype. X 1.
Fig. 69. Proechimys guyannensis riparum, female, AMNH no. 143018, Manaus. Type.
X 1.
Fig. 70. Proechimys guyannensis arabupu, male, AMNH no. 75816, Arabupu. Type. X 1.
Moojen: Brazilian Spiny Rats
357
/T\
358 University of Kansas Publs., Mus. Nat. Hist.
Moojen: Brazilian Spiny Rats
359
Fig. 71. Proechimys guyannensis villicauda, male, MN no. 1932, Tapirapoa. Type. X 1.
Fig. 72. Proechimys guyannensis ribeiroi, male, MN no. 1935, Rio Doze de Outubro.
Type. X 1.
Fig. 73. Proechimys guyannensis hyleae, male, MCZ no. 30887, Tauari. Type. X 1.
Fig. 74. Proechimys guyannensis nesiotes, male, CNHM no. 19496, Ilha de Manapiri.
Type. X 1.
Fig. 75. Proechimys guyannensis leioprimna, female. CNHM no. 19503, Cameta. Type.
X 1.
Fig. 76. Proechimys guyannensis oris, male, CNHM no. 19495, Providencia. X 1.
Fig. 77. Proechimys guyannensis arescens, male, CNHM no. 26440, Fazenda Inhuma.
Paratype. X 1.
Fig. 78. Proechimys guyannensis riparum, female, AMNH no. 143018, Manans. Type.
X 1.
Fig. 79. Proechimys guyannensis arabupu, male, AMNH no. 75816, Arabupu. Type. X 1.
Figs. 80, 81. Proechimys guyannensis villicauda, male, MN no. 1932, Tapirapoa. Tvpe.
X 1.
Figs. 82, 83. Proechimys guyannensis ribeiroi, male, MN no. 1935, Rio Doze de Outubro.
Type. X 1.
Figs. 84, 85. Proechimys guyannensis hyleae, male, MCZ no. 30887, Tauari. Type. X 1.
Figs. 86, 87. Proechimys guyannensis nesiotes, male, CNHM no. 19496, Ilha de Manapiri.
Type. X 1.
Figs. 88, 89. Proechimys guyannensis leioprimna, female, CNHM no. 19503. Type. X 1.
360 University of Kansas Publs., Mus. Nat. Hist.
Figs. 90, 91. Proechimys guyannensis oris, male, CNHM no. 19495, Providencia. X 1.
Figs. 92, 93. Proechimys guyannensis arescens, male, CNHM no. 26440, Fazenda Inhuma.
Paratype. X 1.
Figs. 94, 95. Proechimys guyannensis riparvm, female, AMNH no. 143018, Manaus.
Type. X 1.
Figs. 96, 97. Proechimys guyannensis arabupu, male, AMNH no. 75816, Arabtipu. Type.
X 1.
Moojen: Brazilian Spiny Rats 361
Proechimys guyannensis ribeiroi subspecies nova
Type locality. — Rio 12 de Outubro, affluent of the Camarare, Mato Grosso,
Mato Grosso, Brazil; about 190 kilometers west of Utiariti; altitude 414 meters.
Type: Museu Nacional, no. 1935, adult male (colors faded); collected on 20
June, 1909, by Prof. A. Miranda Ribeiro; original number G.
Range. — Known only from the type locality.
Diagnosis. — Aristijorms wide and stiff; incisive foramen small and narrow;
vomerine sheath complete and thick; p4 with four counterfolds; remaining
molariform teeth with three counterfolds.
Pelage. — Aristijorms on middorsal region: Whitish basally, gradually black-
ening toward tip; total length, 19 to 22 mm; maximum width, 0.8 mm. Seti-
forms on middorsal region: Whitish on basal half, gradually blackening toward
tip but interrupted by subapical zone probably of some tint of ochraceous ; total
length, 22 to 24 mm; maximum width, 0.06 mm. Setiforms on outer thighs:
Whitish basally, gradually blackening toward tip but interrupted by probably
light ochraceous, subapical zone; total length 14 to 16 mm; maximum width,
0.03 mm.
Skull. — Slender; supraorbital ridges low; bullae ovate with shallow grooves;
postorbital process of zygoma almost obsolete and involving mostly jugal; in-
cisive foramen short and narrow (4x2 mm), constricted posteriorly and with
posterior margins raised; vomerine sheath complete and thick; mesopterygoid
fossa extending forward as far as posterior faces of second molars; posterior
palatine foramina on plane with centers of second molars.
Teeth. — Upper molariform teeth with three counterfolds. Lower molariform
teeth: p4 with four counterfolds; molars with three counterfolds which some-
times are fused.
Compaiisons. — From P. g. bolivianus, ribeiroi differs in: aristiforms wider;
incisive foramen shorter and narrower; vomerine sheath complete and thick,
instead of complete or incomplete and not thick; p4 always with four, instead
of usually only three, counterfolds and lower molars always with three, instead
of sometimes with only two, counterfolds in m3. Differences from P. g. villi-
cauda are given in the account of that subspecies.
Remarks. — The name ribeiroi is given in honor of the late Pro-
fessor Alipio Miranda Ribeiro, in recognition of his important work
in Brazilian vertebrate zoology.
Specimens examined. — Total number, 2 (MN), from Brazil, Mato Grosso, Mato Grosso,
Rio 12 de 0\itubro ; altitude, 414 meters.
Proechimys guyannensis hyleae subspecies nova
Type locality.— Tauari, Rio Tapajoz, Porto de Moz, Para, Brazil; approxi-
mately 87 kilometers south of Santarem. Type: Museum of Comparative
Zoology at Harvard College, no. 30887, adult male; collected on 19 January,
1934, by A. M. Olalla; original number 7288.
Range. — Region of lower Tapajoz River and banks of Amazon up to the
Jamunda River.
Diagnosis. — Aristiforms conspicuously wide and stiff; general color on up-
per parts Tawny; incisive foramen long and oval; vomerine sheath complete
362 University of Kansas Publs., Mus. Nat. Hist.
but with maxillary part slender or. sometimes, incomplete ; p4 with four
counterfolds, rarely three ; remaining molariform teeth with three counter-
folds.
Pelage. — Aristijorms on middorsal region: Whitish basally, gradually be-
coming blackish toward tip; total length, 19 to 21 mm; maximum width, 1.1
mm. Seliforms on middorsal region: a. Gray basally, gradually blackening
toward tip but interrupted by wide (5 to 6 mm) Tawny, subapical zone; some
are whitish basally and gradually become sooty brown toward tip except for
same type of subapical zone (tip only slightly darker than subapical zone) ;
b. With the same type described above and some completely blackish, with
the base gray; total length 22 to 25 mm; maximum width, 0.1 mm. Setijorms
on outer thighs: Whitish basally, gradually becoming gray and then blackish
toward tip but interrupted by long, Ochraceous-Tawny, subapical zone; tip,
itself, blackish brown; sometimes this type appears with some lighter ones
and sometimes with completely blackish setiforms; total length 16 to 18 mm;
maximum width 0.3 mm.
Skull. — Medium in size and slender; cranium narrow and not increasing
much in breadth posteriorly ; rostrum stout, laterally thick, with masseteric
crest well-developed; nasals pointed posteriorly; supraorbital ridges broad
but barely extended across parietals; zygomatic arches strong; postorbital
process of zygoma involving mostly squamosal ; incisive foramen long and
narrow (5.5 to 6x2.5 mm), oval and extending posteriorly to point only 2 mm
anterior to premolars; vomerine sheath complete with maxillary part usually
slender; mesopterygoid fossa extending forward as far as centers of third
molars; bulla of medium size, well inflated and with shallow grooves on sur-
face.
Ti < Ih. — Upper molariform teeth with three counterfolds. Lower premolar
with four counterfolds or, sometimes (20 per cent of 15 specimens), with only
three ; lower molars with three counterfolds.
Comparisons. — From P. g. oris and P. g. ncsiotes, hyleae differs in: wider
aristiforms; general color on upper parts Tawny, instead of Ochraceous-Orange ;
vomerine sheath not always complete, instead of always complete. From P. g.
oris, hyleae differs in: p4 usually with four, instead of only three counter-
folds and all molars with three, instead of only lower molars with three coun-
terfolds. From P. g. nesiotes, hyleae differs in: p4 usually, instead of always.
with four counterfolds; color on back Tawny instead of Ochraceous-Orange;
aristiforms wider.
Remarks. — This subspecies shows greater variability than any
other in this species. There are two types of coloration. The most
common type of coloration is dark, with Tawny, subapical zones in
the setiforms of the middorsal region and many completely black
setiforms; in the other type the subapical zone is still Tawny but
there are no black setiforms. One specimen from Obidos, on the
north bank of the Amazon, completely agrees in the characteristics
of color and skull with the reddish type and suggests either that
there is an extension of the range of the subspecies along the lower
Moojen: Brazilian Spiny Rats 363
course of the Tapajoz or that there are two subspecies, in which
event the animals from Tauari are intergrades between hyleae and
an unnamed, tawny-colored subspecies occurring to the southward.
Between 13 and 23 January, 1934, A. M. Olalla collected 10 adult
females, 6 of which contained embryos. Three of the females had
2 embryos each, two had 3 embryos each and one had only 1 em-
bryo. At this same time and place only ten per cent of specimens
obtained were not fully adult,
Specimens examined. — Totai number, 21, from Brazil, Para, as follows: Porto de Moz,
Tauari, right bank of Tapajoz, approximately 85 kilometers south of Santarem, 20 (19 MCZ,
1 CNHM); Obido*, Obidos, 1 (MCZ).
Proechimys guyannensis nesiotes subspecies nova
Type locality. — Ilha de Manapiri, Rio Tocantins, Para, Brazil. Type: Chi-
cago Natural History Museum, no. 19406, adult male; collected on 9 December,
1910, by Dr. Emilia Snethlage; original number, 12.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms wide and stiff; general color on upper parts Ochra-
ceous-Orange ; incisive foramen long, with parallel borders; vomerine sheath
complete and thick; p4 with four counterfolds, remaining molarifoim teeth with
three counterfolds.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blackening
toward tip; total length, 18 to 19 mm; maximum width, 0.9 mm. Sctiforms on
middorsal region-: Gray basally, gradually blackening toward tip but inter-
rupted by Ochraceous-Orange, subapical zone 4 mm long; total length 16 to 19
mm; maximum width, 0.06 mm. Setiforms on outer thighs: Whitish basallj',
gradually becoming gray and blackish toward tip but interrupted by Ochra-
ceous-Buff, subapical zone 3 mm long ; total length, 13 to 15 mm ; maximum
width, 0.03 mm.
Skull. — Of medium size; rostrum short; nasals pointed posteriorly; postor-
bital ridges extending caudad acro>s anterior fifth of parietals; zygomatic
archo^ strong; jugal with process in posterior part of masseteric fossa; postor-
bital process of zygoma involving mostly squamosal; incisive foramen elongate,
narrow (5x2.3 mm), and parallel sided; posterior margin of incisive foramen
approximately 3 mm anterior to premolars; margins of foramen raised to form
ridges; vomerine sheath complete, of almost uniform width and set deeply in
foramen; mesopterygoid fossa extending forward as far as centers of third
molars; bullae of medium size and inflated.
Teeth. — Upper molariform teeth with three counterfolds; p4 with four
counterfolds; ml-3 with three counterfolds.
Comparison. — From P. g. oris, nesiotes differs in: Aristiforms conspicuously
wider; incisive foramen shorter and narrower, with borders parallel instead of
posteriorly constricted; posterior margin of incisive foramen farther from pre-
molars; p4 with four, instead of three, counterfolds; lower molars with three
instead of two counterfolds. From P. g. leiopiimna, nesiotes differs in : Incisive
foramen with parallel borders instead of oval; p4 with four instead of three
counterfolds; m3 always with three instead of two counterfolds.
364 University of Kansas Publs., Mus. Nat. Hist.
Remarks. — Dr. E. Snethlage mentions the type as having been collected at
night in the forest.
Specimens examined. — Total number, 3 (MCZ, CNHM, MN), from Brazil, Para, Tocantins
River, Una de Manapiri.
Proechimys guyannensis leioprimna subspecies nova
Type locality. — Cameta, left bank of Tocantins River, near its mouth,
Cameta, Para, Brazil. Type: Chicago Natural History Museum, no. 19503,
adult female; collected on 21 January, 1911, by Dr. Emilia Snethlage; original
number, 35.
Range. — Known only from type locality but probably extending westward
toward Xingu River.
Diagnosis. — Aristiforms wide and stiff; general color on upper parts Ochra-
ceous-Orange, incisive foramen moderately long; oval; vomerine sheath com-
plete; all molariform teeth with three counterfolds, except lower, third molar
which has only two.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip which is extended as a short filament; total length, 19 to 21 mm;
maximum width, 0.8 to 1 mm. Setijorms on middorsal region: Whitish basally,
gradually blackening toward tip but interrupted by Ochraceous-Orange, sub-
apical zone 2 to 3 mm long; total length, 17 to 20 mm; maximum width 0.1
mm. Setijorms on outer thighs: Whitish basally, becoming gradually gray and
then blackish toward tip but interrupted by Ochraceous-Buff, subapical zone;
blackish tip short; total length 13 to 15 mm; maximum width 0.06 mm.
Skull. — Of medium size; rostrum relatively short; nasals with posterior
borders rounded; postorbital ridges extending across anterior fourth of parietals;
zygomatic arches moderately strong; postorbital process of zygoma involving
both jugal and squamosal; incisive foramen of medium length (4 to 5 mm)
and narrow (about 2.5 mm), oval and extending caudad to a plane approxi-
mately 2 mm anterior to premolars; vomerine sheath complete, with premaxillae
forming approximately anterior % of sheath; maxillary part of sheath short
but well-developed ; mesopteiygoid fossa extending forward as far as centers of
third molars; bullae of medium size but well-inflated.
Teeth. — Upper molariform teeth with three counterfolds. Lower molari-
form teeth with three counterfolds, except third molar which has only two
Comparison. — From P. g. oris, leioprimna differs in: conspicuously wider
aristiforms; shorter and narrower incisive foramen; lower molariform teeth
with three counterfolds (except m3 with only two), instead of lower molars
with only two counterfolds. Differences from P. g. nesiotes are given in the
account of that subspecies.
Remarks. — The paratype was collected in an "igarape," depres-
sion usually invaded by the river waters ; the paratype, collected on
18 January, 1911, had two large embryos.
Specimens examined. — Total number, i (2 CNHM, 2 AMNH), from Brazil, Pari, Cameta.
Moojen: Brazilian Spiny Rats 365
Proechimys guyannensis oris Thomas
Proeohimys oris Thomas, September, 1904, Ann. Mag. Nat. Hist., 14
(ser. 7):195; Thomas, 1905, Ann. Mag. Nat. Hist., 15 (ser. 7):587;
Thomas, 1912, Ann. Mag. Nat. Hist., 9 (ser. 8):89; Tate, 1935, Bull.
Amer. Mus. Nat. Hist., 68:400; Osgood, 1944, Zool. Ser. Field Mus. Nat.
Hist, 29:199.
Proechimys cayennensis oris Ellerman, 1940, The families and genera
of living rodents, Brit. Mus. (Nat. Hist.), 1:121.
Type locality. — Igarape-assu, E. F. B., near Belem, Igarape-assu, Para,
Brazil. Type: British Museum (Nat. Hist.), no. 4.7.4.78, old male; collected
on 6 March, 1904, by Alphonse Robert; original number, 1818.
Range. — Probably most of the region on south bank of Amazon River, be-
tween Tocantins (west) and Gurupi River (south).
Diagnosis. — Aristiforms narrow but somewhat stiff; color on upper parts
Ochraceous-Orange ; incisive foramen long and wide, conspicously constricted
posteriorly; posterior margin of incisive foramen close to plane of premolars;
vomerine sheath complete but maxillary part threadlike; upper molariform
teeth and lower premolar with three counterfolds; lower molars with only
two counterfolds.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip, which is extended as a filament; total length, 16 to 17 mm;
maximum width, 0.6 to 0.7 mm. Setiforms on middorsal region: Gray basally,
gradually becoming blackish toward tip but interrupted by Ochraceous-Orange,
subapical zone 2 to 4 mm long; total length, 18 to 20 mm; maximum width
0.06 mm. Setiforms on outer thighs: Whitish basally, gradually becoming
blackish toward tip but interrupted by Ochraceous-Buff, subapical zone; total
length, 15 to 16 mm; maximum width, 0.04 mm.
Skull. — Of medium size; supraorbital ridges well developed and extending
across anterior fifth of parietals; zygomatic arches strong; jugal with mas-
seteric fossa deep and with well-developed posterior process; postorbital zygo-
matic process involving mostly squamosal; incisive foramen long (6 to 7 mm),
widest anteriorly (2.5 to 3.2 mm) ; but narrowing posteriorly to less than 1 mm
and extending caudad almost to plane of premolars; vomerine sheath com-
plete but maxillary part delicate and threadlike; mesopterygoid fossa extend-
ing forward as far as third molars or posterior parts of second molars; bullae
large and inflated.
Teeth. — Each upper molariform tooth with three counterfolds. In lower
jaw, premolar with three, and molars with only two, counterfolds.
Comparisons. — Differences from the subspecies with adjoining ranges are
given in the accounts of those subspecies.
Remarks. — Thomas (1912:89) extended the known range of the
subspecies to Faro, on the Jamunda River, on the left bank of the
Amazon, and to Boim, on the Tapajoz River, as well as to Bene-
vides, E. F. Braganca, near Belem. It seems to me that the speci-
mens from Faro should be referred provisionally to Proechimys
guyannensis hyleae; the specimens from Boim are "more brightly
5—3343
366 University of Kansas Publs., Mus. Nat. Hist.
rufous" (Thomas, loc. cit.) and could be referred to Allen's P.
boimensis, described in 1914, but P. guyannensis hyleae probably
lives in the same place and only an examination of the specimens,
which I have not seen, would permit of certainly allocating the
specimens to their correct species. The specimens from Benevides
are more certainly P. g. oris.
E. Snethlage collected one specimen in a garden (Providencia,
E. F. B.). However, according to the personnel of the Brazilian
Health Service, the animals are strictly forest dwellers although
they do make excursions into more open places.
Specimens examined. — Total number, 3, from Brazil, Para, as follows: Providencia, E. F.
B., approximately 15 kilometers east from Belem, 1 (CNHM) ; Tanaquara, near Belem, 1
(MN) ; Rio Guama, near Belem, 1 (AMNH).
Additional record. — Brazil, Para, Benevides, E. F. B., approximately 100 kilometers north-
east of Belem (Thomas, 1912:89).
Proechimys guyannensis arescens Osgood
Proechimys cayennensis arescens Osgood, 12 July 1944, Zool. Ser. Field
Mus. Nat. Hist., 29(13) :198.
Type locality. — Fazenda Inhuma, below Santa Filomena, upper Rio Par-
naiba, Vitoria do Alto Parnaiba, Maranhao. Brazil. Type: Chicago Natural
History Museum, no. 26441, adult male; collected on 5 August, 1925, by
Heinrich E. Snethlage.
Range. — Region including the valleys of the Turi-assu aud Parnaiba rivers,
Maranhao, Brazil.
Diagnosis. — Aristiforms moderately wide and not conspicuously stiff; gen-
eral color of upper parts near (15'a) Ochraceous-Orange ; incisive foramen
long and wide; vomerine sheath complete or incomplete; upper molariform
teeth and lower premolar with three counterfolds ; lower molars with only two.
Pelage. — Aristiforms on middorsal region: Whitish basally, gradually
blackening toward tip; total length, 19 to 21 mm; maximum width, 0.7 mm.
Setijorms on middorsal region: Whitish basally or on basal half, gradually
becoming gray and then blackish toward tip, but interrupted by long (5 to
6 mm) subapical zone near (15'a) Ochraceous-Orange; total length, 15 to
16 mm; maximum width 0.05 mm. Seliforms on outer thighs: Whitish on
basal half, gradually becoming gray and then blackish toward tip but inter-
rupted by Ochraceous-Buff, subapical zone; tip sometimes not conspicuously
darker than subapical zone; total length 18 to 25 mm; maximum width,
0.03 mm.
Skull. — Medium in size, not elongated; nasals pointed posteriorly; supra-
orbital ridges strong and thick, extending caudad across anterior third of
parietals; zygomatic arches strong; postorbital process of zygoma involving
only squamosal; incisive foramen 5 by 2.7 mm, oval and extending caudad
to plane approximately 2 mm anterior to premolars; posterior margins of in-
cisive foramen not forming a ridge; vomerine sheath complete and with max-
illary part slender and threadlike, or incomplete, in which event, maxillary
part not extended enough to join premaxillary process; mesopterygoid fossa
Moojen: Brazilian Spiny Rats 367
extending forward as far as centers of third molars; bulla large and more or
less triangular in its peripheral outline.
Teeth. — Upper molariforrn teeth with three counterfolds each. Lower pre-
molar with three counterfolds; lower molars with two counterfolds.
Comparisons. — From P. g. oris, arescens differs in : Color of upper parts
lighter and more uniform; incisive foramen oval instead of conspicuously con-
stricted posteriorly; posterior margin of incisive foramen farther from pre-
molars.
Remarks. — One specimen from Turi-assu (MN) has been iden-
tified by 0. Thomas as "P. oris" (his own handwriting is on the
label) and the subspecies is really closely related to oris.
Specimens examined. — Total number, 3, from Brazil, Maranhao, as follows: Yitoria do
Alto Pamaiba, Fazenda Inhuma (below Santa Filomena), 2 (CNHM) ; Alto da Alegria, Turi-
assu, 1 (MN).
Proechimys guyannensis riparum subspecies nova
Type locality. — Manaus, Manaus, Amazonas, Brazil. Type: American Mu-
seum of Natural History, no. 143018, adult female; collected 6 March, 1943.
Range. — Known only from type locality but probably extending northward
and eastward.
Diagnosis. — Aristiforms wide and stiff; upper parts Ochraceous-Tawny ; in-
cisive foramen short, wide, and oval; vomerine sheath incomplete; upper
molariforrn teeth and lower premolar with three counterfolds; lower molars
with only two counterfolds.
Pelage. — Aristiforms on middorsal region: Gray basally gradually blacken-
ing toward tip; total length, 18 to 20 mm; maximum width, 0.9 mm. Seti-
forms on middorsal region: Whitish basally, gradually blackening toward tip
but interrupted by 2 mm long, Ochraceous-Tawny, subapical zone; total
length 20 to 22 mm; maximum width, 0.04 mm. Some are whitish basally
and gradually become black toward tip with no subapical zone. Setiforms on
outer thighs: Whitish basally, gradually becoming gray and then blackish
toward tip but interrupted by Ochraceous-Buff, subapical zone 3 to 4 mm
long; tip not conspicuously dark; total length, 17 to 19 mm; maximum width,
0.03 mm.
Skull. — Of medium size and slender; rostrum slender; nasals rounded pos-
teriorly; supraorbital ridges well developed and barely extended onto ante-
riormost part of parietals; zygomatic arches slender; infraorbital foramen
with well-developed groove for nerve transmission; postorbital process of
zygoma almost obsolete and involving mostfy squamosal; incisive foramen
short and wide (4.5x3.3 mm), oval and with posterior margins raised to form
ridges which extend toward premolars; posterior margin of incisive foramen
approximately 2.5 mm anterior to premolars; vomerine sheath incomplete,
with only short, premaxillary part; mesopterygoid fossa extending forward as
far an anterior parts of third molars; bullae large and inflated, with more or
less triangular outline.
Teeth. — Crown length of well worn P4-M3, 6.8 mm; upper molariforrn
368
University of Kansas Ptjbls., Mus. Nat. Hist.
teeth with three counterfolds each. Lower premolar with three counterfolds ;
lower molars with two counterfolds.
Comparisons. — From P. g. oris and P. g. hyleae, riparum differs in : Shorter
and wider incisive foramen; vomerine sheath incomplete, instead of some-
times incomplete. From P. g. oris, riparum differs in: Upper parts Ochrace-
ous-Tawny instead of Ochraceous-Orange ; aristiforms conspicuously wider.
70 40
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100 70 40
Fig. 98. Map showing the geographic ranges of the subspecies of Proechimys guyannensis in
Brazil.
From P. g. hyleae, riparum differs in: Aristiforms narrower; upper parts
Ochraceous-Tawny instead of Tawny; lower premolars with three, instead of
four, counterfolds; lower molars with two, instead of three, counterfolds.
Specimen) examined. — Type only.
Moojen: Brazilian Spiny Rats 369
Proechimys guyannensis arabupu subspecies nova
Type locality.— Arabupu, Mount Roraima, Boa Vista, Territ. Rio Branco;
about 1540 meters altitude. Type: American Museum of Natural History,
no. 75816, adult male ; collected by Dr. G. H. H. Tate on 30 December, 1927 ;
original number, 4716.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms conspicuously wide and stiff; color on upper parts
dark, near (15';) Ochraceous-Tawny ; incisive foramen widest in anterior
third; vomerine sheath complete, sometimes incomplete; upper molariform
teeth and lower premolar with three counterf olds ; lower molars with two
counterfolds.
Pelage. — Aristiforms on middorsal region: Whitish basally, gradually
blackening toward tip; total length, 19 to 22 mm; maximum width, 1.1 mm.
Setiforms on middorsal region: Gray basally, gradually blackening toward tip
but interrupted by dark (15';) Ochraceous-Tawny, subapical zone 3 mm
long; some completely blackish on distal parts; total length, 20 to 23 mm;
maximum width, 0.07. Setiforms on outer thighs: Whitish basally, gradually
blackening toward tip but interrupted by Ochraceous-Buff or Light Ochra-
ceous-Buff, subapical zone; total length, 20 to 23 mm; maximum width,
0.07 mm.
Skull. — Size medium; nasals pointed posteriorly; supraorbital ridges prom-
inent and slightly extended caudad onto anterior half of parietals; groove for
transmission of nerve in infraorbital foramen weakly developed; zygomatic
arches strong; postorbital process of zygoma involving mostly squamosal;
incisive foramen approximately 5.5 x 2.7 mm, widest in anterior third and
constricted posteriorly, with posterior margin about 1 mm anterior to plane
of premolars; vomerine sheath complete, with premaxillary part expanded
and maxillary part notably slender and sometimes lacking; mesopterygoid
fossa in some specimens extending forward as far as middle parts of second
molars; bullae large and inflated.
Teeth. — Upper molariform teeth with three counterfolds each. Lower pre-
molar with three counterfolds; molars with only two.
Comparisons. — From P. g. warreni, arabupu differs in: Narrower aristi-
forms; narrower incisive foramen; lower premolar with three instead of four
counterfolds; lower molars with two, instead of three, counterfolds. From
P. g. oris, arabupu differs in: Aristiforms wider; posterior margin of incisive
foramen farther from plane of premolars; upper parts dark (15';) Ochraceous-
Tawny, instead of Ochraceous-Orange.
Remarks. — The sample is fairly uniform.
Specimens examined. — Total number, 6 (AMNH), from Brazil, Territorio do Rio Branco,
Boa Vista, Mount Roraima, Arabupu ; approximately 1540 m. altitude.
Subgenus TRINOMYS Thomas
Genotype. — Echimys albispinus Is. Geoffroy Saint-Hilaire, 1838; by
original designation.
Trinomys Thomas, July 1921, Ann. Mag. Nat. Hist., 8 (ser. 9) :140
(orig. descr.); Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :401; Eller-
man, 1940, The families and genera of living rodents, Brit. Mus. (Nat.
Hist.), 1:115.
370
University of Kansas Publs., Mus. Nat. Hist.
General characters. — Pelage of upper parts with lanceolate and, sometimes,
clavate aristiforms extending over most of rump and onto thighs; tail 86 to
103 per cent of length of head and body; tail sometimes white-tipped and
Fig. 90. Map showing the geographic ranges of the subspecies of three species of the subgenus
Trinomys.
sometimes penicillate; skull small, with ridges moderately developed; supra-
orbital ridges involving no part of parietals; infraorbital foramen with no
separate groove for transmission of nerve; mesopterygoid fossa extending for-
ward to level of second or first molars; incisors opisthodont, orthodont or
proodont; molariform teeth, in occlusal view, with main fold large and usu-
Moojen: Brazilian Spiny Rats 371
ally reaching opposite wall; no counterfold anterior to main fold in upper
molariform teeth and usually no counterfold posterior to main fold in lower
molariform teeth; premolars larger than first molars, first molars larger than
second molars and second molars larger than third molars; four molariform
teeth of nearly equal size in some animals.
Thomas (1921:140) erected the subgenus Trinomys, including in it the
species albispinus and setosus and stated that "the primary distinction between
these [ Trinomys and Proechimys] lies in the number of laminae present in
the cheekteeth — four in Proechimys, three in Trinomys." The distinction is
valueless as a subgeneric character, not only because the character is not con-
si ant in the species in the subgenus but also because there is subspecific vari-
ation in number of laminae in the cheekteeth. Proechimys albispinus, how-
ever, shares with three other species common characters, as listed above, and
the name Trinomys will, therefore, apply to this group of species, since Pro-
n-himys albispinus is the genotype.
Proechimys dimidiatus (Giinther)
Echimys dimidiatus Giinther, 1 April 1877, Proc. Zool. Soc. London,
1876(4) :747.
Proechimys dimidiatus Allen, 1899, Bull. Amer. Mus. Nat. Hist,, 12(20) :
264; Ribeiro, 1905, Arch. Mus. Nac. Rio de Janeiro, 13:187; Thomas,
1921, Ann. Mag. Nat. Hist., 8 (ser. 9) :141; Tate, 1935, Bull. Amer. Mus.
Nat. Hist., 68(5) :400; Ellerman, 1940, The families and genera of living
rodents, Brit. Mus. (Nat. Hist.), 1:122.
Type locality. — Unknown; probably southwestern Rio de Janeiro, Brazil
(see Remarks).. Type: British Museum (Nat. Hist.), no. 51.7.21.24; pre-
sented by Lord Derby.
Range. — Rio de Janeiro, from the southern limit of the state northward to
and including the Distrito Federal.
General characters. — Size large ; tail averaging 80 per cent of head and
body; aristiforms narrow and soft (0.4 to 0.5 mm wide), imparting a non-
spiny character to the pelage ; general color of upper parts Ochraceous-Buff ,
finely lined with blackish brown, gradually becoming lighter on sides; ventral
surface of body and inner sides of legs white; feet dorsally white but with a
sepia-colored stripe along outer margin; tail brownish-black above and white
below, but white sometimes extended to upper side in distal part; skull broad
with no conspicuous ridges; jugals deep with transverse ridge usually con-
spicuous; postorbital process of zygoma involving only squamosal; incisive
foramen short and wide posteriorly; vomerine sheath complete in 95 per
cent of specimens and with maxillary part thick; posterior palatine foramina
at plane of first molars or slightly anterior to them; bullae moderately de-
veloped; in juvenal specimens, each upper molariform tooth with three coun-
terfolds. but posteriormost counterfold small; in adult specimens, posterior-
most counterfold disappearing in 50 per cent of fourth premolars and first
molars, in 20 per cent of second molars, and in 15 per cent of third molars;
lower molariform teeth with two counterfolds in almost every juvenal speci-
men, this number, in adult animals, decreasing in m3 to one in 20 per cent
of specimens but rarely being reduced in other teeth.
372 University of Kansas Ptjbls., Mus. Nat. Hist.
General characters. — Aristiforms soft and narrow, ranging from 15 to 19 mm
in total length and 0.4 to 0.5 mm in maximum width; pelage generally non-
spiny and not harsh; length of tail ranging from 20 per cent shorter than
head and body to as long as, or slightly longer than, head and body; ears
rather small (23 to 25 mm).
Figs. 100-103. Proechimys dimidiatus, male, MN no. 5452, Tijuca. X 1.
Color. — General color of back and sides results from uniform mixture of
black distal parts of aristiforms with Ochraceous-Buff of subapical zone of
setiforms. Dorsally, from nose caudad to rump, mixture appears brownish-
black, lined with Ochraceous-Buff; toward sides, amount of Ochraceous-Buff
gradually increases and resultant color is much lighter brown than on back.
On outer parts of arms and legs, color turns gradually to sepia toward distal
parts and finally to uniform sepia on wrists and ankles, this color extending
to outer dorsal parts of hands and feet; on ankles, sepia forms complete ring,
as usual in the genus. Tail blackish-brown on upper parts, this stripe grad-
ually tapering toward tip where dark brown hairs form small pencil; white of
under side of tail sometimes seen also entirely around distal part, short of
tip which remains dark brown. Ventral surfaces wholly white, from upper
lips caudad including inner surfaces of legs.
Hairs. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip that has long, fine filament; total length 16 to 19 mm; maxi-
mum width 0.5 mm. On outer thigh whitish basally, gradually blackening
toward tip; some with Ochraceous-Buff, subapical zone; total length 13 to
Moojen: Brazilian Spiny Rats 373
15 ram; maximum width 0.25 mm. Setiforms on middorsal region: Whitish
on basal half, gradually blackening toward tip, but interrupted by Ochraceous-
Buff, subapical zone; some with Light Ochraceous-Buff, subapical zone and
short, blackish zone on tip; total length 12 to 14 mm; maximum width
0.02 mm. Setiforms on outer thighs: Whitish on basal half, then gradually
becoming gray on middle part and finally Light Ochraceous-Buff on distal
third, or with tip blackish and Ochraceous-Buff, subapical zone.
Skull. — Elongate and broad with no conspicuous crests; rostrum rather
stout; jugals deep with transverse crest usually well-developed; zygomatic
postorbital process conspicuous and formed entirely of squamosal; incisive
foramen short and wide posteriorly; vomerine sheath complete in great ma-
jority of specimens, its maxillary part wide and strong; posterior palatine
foramina on plane with front of Ml or slightly farther forward; bullae rather
small and elongate.
Teeth. — P4 with three secondary folds in all juvenal specimens, but pos-
teriormost fold small and disappearing in 50 per cent of adult specimens; Ml
with 3 outer folds in juveniles and also disappearing in 50 per cent of adults;
M2 with three outer folds in juveniles, but only 20 per cent remaining in
adults; M3 with 3 outer folds in 50 per cent of juveniles, decreasing to 15
per cent in adults. Lower molariform teeth: p4 with 2 secondary folds; ml
with 2 secondary folds in 90 per cent of adults and in all juveniles; m2 with
2 secondary folds in 98 per cent of adults and in all juveniles; m3 with 2
secondary folds in 81 per cent of adults, remaining percentage with only one
counterfold, and with 2 secondary folds in all juveniles.
Remarks. — Samples studied of P. dimidiatus are notably uniform
throughout the geographic range of the species. The few biotypes
detected seemed unworthy of subspecific rank.
In discussing the type locality of the species, Thomas (1921:141)
states: "We know that its donor did obtain a number of specimens
from Rio Janeiro, and the skull agrees so closely with those of two
examples from Itatiaia, near to the Rio-Minas frontier, collected
and presented by Prof. J. P. Hill, that I have no hesitation in re-
ferring the latter to Gunther's species."
Specimens examined.— Total number, 211 (MN), from Brazil as follows: Rio de Janeiro;
Parati, Pedra Branca (400 m.), 113; Mangaratiba, Fazenda do Rubiao (750 m.), 3; Fazenda
do Tenente (700 m.), 4; Fazenda da Lapa (450 m.), 13; Teresopolis, Fazenda Guinle (960 m.)(
61; Nova Iguassu, Barro Branco (20 m.), 16; Distrito Federal, Tijuca, 1.
Additional records. — Rio de Janeiro, Itatiaia (Thomas, 1921:141); Rio de Janeiro, Zona
da mata, Mont-Serrat, Serra do Itatiaia (Ribeiro, 1905:187).
Proechimys iheringi Thomas
General characters. — Size large; tail long; aristiforms generally wide and
stiff; general color on upper parts and sides a combination of blackish from
tips of aristiforms with cinnamon ground color from subapical zones of seti-
forms; darker band on middorsal line; differentiated light-colored aristiforms
conspicuous on outer sides of thighs and rump; usually rufous tint on neck
and postauricular region; underparts white; tail with white tip, usually ac-
374 University of Kansas Publs., Mus. Nat. Hist.
centuated by white brush; feet white on dorsal surface; hind feet slightly
darker on outer sides; skull elongate and smooth; jugals wide dorso-ventrally ;
incisive foramen elongate; upper molariform teeth usually with one to five
counterfolds, number varying with subspecies; lower premolar always with
two counterfolds and lower molars always with one or two counterfolds.
Remarks. — As a whole, the samples of the populations of the
species do not afford a satisfactory record of the distribution; my
concept of the group may be changed when further collections are
made in localities geographically intermediate between those from
which specimens now are known. If some of the forms prove to be
physiologically isolated, they may deserve treatment as full species
according to the conventional standards of systematic zoology.
P. pone met, for example, does not seem to be geographically iso-
lated from P. gratiosus. P. denigratus, at the northernmost known
occurrence of the species, actually represents a striking jump in the
cline, although collections from intermediate regions may provide
intermediate structural stages. Further collecting may also prove
that the southern form, P. iheringi iheringi, is completely isolated
from the rest of the group. However, these samples are certainly
more related to each other than any one of them is to that of the
other species found in the same range, namely P. dimidiatus, and
all the forms in question, therefore, seem best arranged as sub-
species of one full species. A clinal variation certainly exists among
these forms and the most striking differences correspond to larger
geographical distances.
Moojen: Brazilian Spiny Rats
375
/T\
Fig. 104.
Fio. 105.
Fig. 106.
Type. X 1.
Fig. 107.
Fig, 108.
tuna. Type.
Fig. 109.
Braz. Type.
Proechimys iheringi iheringi, female, MN no. 6453, Ilha de Sao Sebastiao. X 1.
Proechimys iheringi bonafidei, male, MN no. 6183, Fazenda Boa Fe. Type. X 1.
Proechimys iheringi gratiosus, male, MN no. 4024, Floresta da Caixa Dagua.
Proechimys iheringi panema, female, MN no. 8288, Campinho. Type. X 1.
Proechimys iheringi denigratus, male, MN no. 8500, Mata do Ribeirao da For-
X 1-
Proechimys iheringi paratus, female, MN no. 4012, Floresta da Capela de Sao
X 1.
376
University of Kansas Publs., Mus. Nat. Hist,
Fig. 110.
Fig. 111.
Fig. 112.
Type. X 1.
Fio. 113.
Fio. 114.
tuna. Type.
Fio. 116.
Braz. Type.
Proechimys iheringi iheringi, female, MN no. 6453, Ilha de Sao Sebastiao. X 1.
Proechimys iheringi bonafidei, male, MN no. 6183, Fazenda Boa Fe. Type. X 1.
Proechimys iheringi gratiosus, male, MN no. 4024, Floresta da Caixa Dagua.
Proechimys iheringi panema, female, MN no. 8288, Campinho. Type. X 1.
Proechimys iheringi denigratus, male, MN no. 8500, Mata do Ribeirao da For-
X 1.
Proechimys iheringi paratus, female, MN no. 4012, Floresta da Capela de Sao
X 1.
Moojen: Brazilian Spiny Rats
377
x i.
Figs. 116, 117. Proechimys iheringi iheringi, female, MN ho. 6453, Ilha de Sao Sebastiao.
Figs. 118, 119. Proechimys ihering
Type. X 1.
Figs. 120, 121. Proechimys ihering
Dagua. Type. X 1.
Figs. 122, 123. Proechimys ihering
X 1.
Figs. 124, 125. Proechimys iheringi
Fortuna. Type. X 1-
Figs. 126, 127. Proechimys iheringi
Sao Braz. Type. X 1.
i bonafidei, male, MN no. 6183, Fazenda Boa Fe.
i gratiosus, male, MN no. 4024, Floresta da Caixa
i panema, female, MN no. 8288, Campinho. Type.
i denigratus, male, MN no. 8500, Mata do Ribeirao da
i paratus, female, MN no. 4012, Floresta da Capela de
378 University of Kansas Publs., Mrs. Nat. Hist.
Proechimys iheringi iheringi Thomas
Proechimys iheringi Thomas, August. 1911, Ann. Mag. Nat. Hist., 8
(ser. 8) :252 (orig. descr.) ; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser.
9):141; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :400; Ellerman,
1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.),
1:122.
Type locality. — Island of Sao Sebastiao (off Sao Paulo), Formosa, Sao
Paulo, Brazil. Type: British Museum (Nat. Hist.), no. 2.8.25.5, adult male,
presented by the Sao Paulo Museum.
Range. — Littoral and islands of Sao Paulo and Rio de Janeiro.
Diagnosis. — Aristiforms narrow; tail shorter than head and body; setiforms
Cinnamon-Buff; incisive foramen short; vomerine sheath complete; upper
molariform teeth with two or three counterf olds ; lower molariform teeth with
two counterfolds, rarely one in m3.
Pelage. — Aristijorms on middorsal region: Gray basally, gradually blacken-
ing toward tip; total length 18 to 23 mm; maximum width, 0.6 mm. Aristi-
jorms on outer thighs: Gray basally, blackening distally toward tip; some
differentiated with Cinnamon-Buff tip. Setiforms on middorsal region: Gray
basally, gradually blackening toward tip but interrupted by a Cinnamon-Buff,
subapical zone 3 mm long; total length, 16 to 20 mm; maximum width,
0.06 mm. Setiforms on outer thighs: Gray basally, gradually blackening to-
ward tip but interrupted by Cinnamon-Buff, subapical zone or with Cinnamon-
Buff continuous to tip.
Skidl. — Slender; bullae small and well inflated; jugal dorsoventrally wide
with transverse ridge inconspicuous; incisive foramen short. 3.5x2.5 mm;
vomerine sheath complete; mesopterygoid fossa extending forward as far as
middle parts of second molars; postorbital process of zygoma small, formed
by both jugal and squamosal; posterior palatine foramina at plane of pre-
molars; interorbital breadth narrow.
Teeth. — Upper molariform teeth with two or three counterfolds (when un-
worn usually three and rarely four) ; sometimes only one counterfold in M3
and sometimes counterfolds fused in molars. Lower molariform teeth with
two counterfolds, rarely one in m3.
Comparisons. — From P. i. bonafidei and P. i. gratiosus, iheringi differs in:
Incisive foramen shorter; vomerine sheath complete, instead of usually in-
complete; setiforms Cinnamon-Buff, instead of Ochraceous-Buff ; upper molar-
iform teeth with two or three separate counterfolds, instead of having coun-
terfolds fused or reduced to one or two; aristiforms narrower in iheringi than
in bonafidei.
Specimens examined.- Total number, 25, from Brazil, as follows: Sao Paulo, Formosa,
Ilha de Sao Sebastiao, 9 (DZ 6, MN 2, MCZ 1); Sao Paulo, Mogi das Cruzes, Alto da Sena,
alt. 900 in., 2 (DZ); Sao Paulo, Ubatuba, alt. 10 m., 4 (2 DZ, 2 MN) ; Rio de Janeiro,
Angra dos Rets, 2 (MN); Rio de Janeiro, Angra dos Reis, Ilha Grande, 7 (5 DZ, 1 MCZ,
1 MN).
Proechimys iheringi bonafidei subspecies nova
Type locality. — Fazenda Boa Fe, Teresopolis, Rio de Janeiro, Brazil; alt.
850 meters. Type: Museu Nacional, no. 6183, adult male; collected on 18
August, 1942, by G. Pereira; SEPFA no. M 14663.
Moojen: Brazilian Spiny Rats 379
Range. — Known only from the type locality.
Diagnosis. — Aristiforms wide and stiff; tail shorter than head and body;
setiforms Ochraceous-Buff ; incisive foramen long; vomerine sheath incom-
plete, or rarely complete; molariform teeth with two counterfolds usually fused.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip; total length, 22 to 26 mm; maximum width, 0.8 mm. Aristi-
forms on outer thighs: Gray basally, gradually blackening toward tip but in-
terrupted by Ochraceous-Buff subapical zone; some Ochraceous-Buff to tip;
total length, 18 to 20 mm; maximum width, 0.7 mm. Setiforms on middorsal
region: Gray basally, gradually blackening toward tip but interrupted by
Ochraceous-Buff, subapical zone; total length, 17 to 20 mm; maximum width,
0.06 mm. Setiforms on outer thighs: Gray basally, gradually blackening to-
ward tip but interrupted by Ochraceous-Buff, subapical zone; only a short
blackened tip.
Skull. — Large, with elongate rostrum; bullae large and well inflated; jugals
with transverse ridge inconspicuous; post orbital process of zygoma small,
formed mostly by squamosal ; incisive foramen elongated (5.5 x 2.5 mm) ;
vomerine sheath incomplete or, if complete, with maxillary part thin and deli-
cate; posterior palatine foramen at plane of first molars; mesopterygoid fossa
extending forward as far as middle parts of second molars.
Teeth. — Upper molariform teeth with two counterfolds; these completely
separated in 3 of 16 specimens; two counterfolds coalesced in all three molars
in 6 specimens; counterfolds coalesced in only two molars in 3 specimens;
counterfolds coalesced in only one molar in 4 specimens. Lower molariform
teeth with two counterfolds which are completely separated in 13 of 16 speci-
mens; counterfolds coalesced in only one molar in 2 specimens; counterfolds
coalesced in all three molars in one specimen.
Comparisons. — From P. i. gratiosus, bonafidei differs in: Aristiforms wider;
tail shorter; molariform teeth with two counterfolds instead of one or two.
Differences from P. i. iheringi are given in the account of that subspecies.
Remarks. — Of females with embryos two were captured in April
and one in September. The embryos number 2, 1, 2. Young were
captured mostly in April, but two were taken in July. Male gonads
seemed to be most active in March, April and September. The
animals lived in a second growth forest, approaching the climax.
The rainfall was more than 1600 mm annually, and the mean an-
nual temperature was 18.5° centigrade.
Specimens examined. — Total number, 18 (MN), from Brazil, Rio de Janeiro, Teresopolis,
Fazenda Boa Fe.
Proechimys iheringi gratiosus subspecies nova
Type locality. — Floresta da Caixa Dagua, Santa Teresa, Espirito Santo,
Brazil; altitude 750 meters. Type: Museu Nacional, no. 4024, adult male;
collected on 25 May, 1940. by C. Lako; SEPFA no. M 6911.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms narrow; tail of same length as head and body;
380 University of Kansas Publs., Mus. Nat. Hist.
setiforms Ochraceous-Buff; incisive foramen long; vomerine sheath usually
incomplete; upper molariform teeth with one or two counterf olds ; lower
molariform teeth with two count erf olds, except that m3 usually has only one.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually black-
ening toward tip; total length, 21 to 27 mm; maximum width, 0.6 mm. Aris-
tiforms on outer thighs: Gray basally, gradually blackening toward middle,
and Ochraceous-Buff on distal half; total length, 18 to 21 mm; maximum
width, 0.5 mm. Setiforms on middorsal region: Gray basally, gradually black-
ening toward tip but interrupted by short, Ochraceous-Buff, subapical zone;
total length, 18 to 20 mm; maximum width, 0.06 mm. Setiforms on outer
thighs: Gray basally, gradually blackening toward middle, and distal part
Ochraceous-Buff or with only tip blackened; total length, 14 to 16 mm; maxi-
mum width, 0.05 mm.
Skull. — Slender; bullae small but well-inflated; upper edge of jugals deeply
concave; transverse ridge of jugals conspicuous; postorbital process of zygoma
small, involving only squamosal; incisive foramen elongate (5x2.5 mm);
vomerine sheath almost always incomplete, and maxillary part lacking or,
when present, slender; mesopterygoid fossa extending forward as far as middle
of second molars; posterior palatine foramina at plane of front border of first
molars or slightly anterior thereto.
Teeth. — Upper molariform teeth with two counterfolds in 10 of 16 speci-
mens and only one in remainder; these folds commonly coalesced; M3 with
only one counterfold in 6 specimens, and 2 counterfolds in remainder. Lower
molariform teeth with two counterfolds in 6 specimens and in 10 of them m3
has only one counterfold.
Comparisons. — From P. i. panema, gratiosus differs in: Lower molariform
teeth with only one counterfold in smaller percentage of specimens; incisive
foramen shorter; aristiforms narrower; setiforms Ochraceous-Buff instead of
Cinnamon. Differences from iheringi and paratus are given in the accounts
of those subspecies.
Remarks. — All the animals were captured in climax forest.
Specimens examined. — Total number, 16 (MN), from Brazil, Espirito Santo, Santa Teresa,
Floresta da Caixa Dagua, altitude 750 meters.
Proechimys iheringi panema subspecies nova
Type locality. — Campinho, Colatina, Espirito Santo, Brazil; altitude 500
meters. Type: Museu Nacional, no. 8288, adult female; collected on 15 July,
1942, by C. Lako.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms moderately wide; tail of approximately same
length as head and body; setiforms Cinnamon; incisive foramen moderately
long and narrow; vomerine sheath incomplete; upper molariform teeth with
two counterfolds, but m3 most frequently with one.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually black-
ening toward tip; total length, 21 to 23 mm; maximum width, 0.8 mm. Aris-
tiforms on outer thighs: Gray, some gradually blackening toward tip and
others with distal part Cinnamon; total length, 17 to 19 mm; maximum
width, 0.7 mm. Setiforms on middorsal region: Gray, gradually blackening
Moojen: Brazilian Spiny Rats 381
»
toward tip, but interrupted by Cinnamon, subapical zone; total length, 18 to
20 mm; maximum width, 0.06 mm. Setijorms on outer thighs: Gray, grad-
ually blackening toward middle, and Cinnamon on all of distal parts or with
tip blackish; total length, 13 to 15 mm; maximum width, 0.09 mm.
Skull. — Strong, with jugals dorso-ventrally wide; interorbital region and
cranium wide; bullae well inflated; transverse ridge of jugals not well-devel-
oped; postorbital process of zygoma small and formed only of squamosal;
incisive foramen 4.7x2.2 mm; vomerine sheath always incomplete, with max-
illary part reduced to small process; mesopteiygoid fossa extending forward
as far as middle of second molars or only slightly short thereof; posterior
palatine foramina at plane of front of first molars.
Teeth. — All upper molariform teeth with two counterfolds in 4 speci-
mens; one having only one counterfold in M3; 3 with counterfolds coalesced
in one or two molars. Lower molariform teeth with two counterfolds in one
specimen, these counterfolds not coalesced; m3 with one counterfold in 4
specimens and with two in one specimen.
Comparisons. — Differences from P. denigratus and P. i. paratus are given in
the accounts of those animals.
Specimens examined. — Total number, 5 (MN), from Brazil, Espirito Santo, Colatina,
Campinho ; altitude 500 meters.
Proechimys iheringi denigratus subspecies nova
Type locality. — Mata do Ribeirao da Fortuna, 40 kilometers west of Ilheus,
Itabuna, Bahia, Brazil. Type: Museu Nacional, no. S500. adult male; col-
lected 16 March, 1945.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms wide and stiff; tail longer than head and body;
setiforms near (15"o) Cinnamon; incisive foramen long and narrow; vomerine
sheath complete; premolars with two counterfolds, upper molars with one or
two, and lower molars with only one.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip; total length, 20 to 22 mm; maximum width, 1.1 mm. Aristi-
forms on outer thighs: Gray basally, gradually blackening toward tip or with
distal part near (15"a) Cinnamon; total length, 14 to 16 mm; maximum
width, 0.5 mm. Setiforms on middorsal region: Gray basally, gradually
blackening toward tip but interrupted by near (15"a) Cinnamon, subapical
zone 4 mm wide; total length, 18 to 20 mm; maximum width, 0.05 mm.
Setiforms on outer thighs: Gray basally, gradually blackening toward tip but
interrupted by wide, near (15"a) Cinnamon, subapical zone.
Skull. — Slender; nasals short; bullae large and well-inflated; jugals with
conspicuous transverse ridge; postorbital process of zygoma conspicuous, spini-
form and formed almost exclusively by jugal; incisive foramen elongated and
narrow (5x1.8 mm); vomerine sheath complete and formed almost exclu-
sively by premaxillae; maxillary part of this sheath short and in most speci-
mens the two parts of sheath completed by vomer itself; mesopterygoid fossa
extending forward as far as middle of second molars and in some skulls as
far as anterior border of second molars; posterior palatine foramina at ante-
rior plane of first molars.
6—3343
382 University of Kansas Publs., Mtjs. Xat. Hist.
Teeth. — Upper molariform teeth: P4 always with two counter!" olds ; Ml
with two counterfolds in 65 per cent of specimens but anterior couuterfold
poorly developed; rest of specimens with only one counterfold in Ml; M2
with two counterfolds in 50 per cent of specimens and only one in remainder;
M3 with two counterfolds in only 17 per cent of specimens, and remainder
with only one. Lower molariform teeth: p4 always with two counterfolds;
molars always with only one counterfold.
Comparisons. — From P. i. panema. denigratus differs in: Each lower molar
with only one, instead of with more than one, counterfold; incisive foramen
longer and narrower; vomerine sheath complete instead of incomplete; aris-
tiforms conspicuously wider; tail longer.
Remarks.— One female (SEPFA no. M 17060) captured on 9
January, 1944, gave birth to two females on 26 January, 1944.
Each of these young measured 177 mm in total length and weighed
27.8 g. On 4 March, 1944, their measurements were: head and
body, 120,120; tail, 120,130; hind foot, 32,33; ear, 21,22; skull: —
total length, 36.0,35.0; condyloincisive length, 29.0,29.1; zygomatic
breadth, 19.1,18.5; length of nasals, 12.5,11.6; interorbital constric-
tion, 9.3,8.8; cranial breadth. 16.4,16.9; palatilar length, 11.5,10.5;
crown length of P4 and Ml, 4.3,4.3 mm.
The forest where the animals were captured has a high percentage
of deciduous trees in spite of the heavy rainfall in this region. All
of the animals were trapped near water. Young were captured
from January to May. Most animals have a conspicuous Cinnamon
patch on the nuchal region.
Specimens examined. — Total number, 34 (SEPFA 33, MN j), from Brazil, Bahia, Itabuna,
Mata do RJbeirao da Fortuna.
Proechimys iheringi paratus subspecies nova
Type locality. — Floresta da Capela de Sao Braz, Santa Teresa, Espirito
Santo, Brazil; altitude 630 meters. Type: Museu Nacional, no. 4012, adult
female; collected on 24 September, 1940, by Dr. H. W. Laemmert; SEPFA
no. M 8447.
Range.— Known only from the type locality.
Diagnosis. — Aristiforms wide and stiff; tail 96 per cent of head and body;
color on setiform Cinnamon-Buff; incisive foramen short and moderately
wide; vomerine sheath complete; all molariform teeth with two counterfolds.
Pelage. — Aristiforms on middorsal region : Gray basally, gradually blacken-
ing toward tip; total length, 24 to 26 mm; maximum width, 1.3 mm. Aristi-
forms on outer thighs: Gray basally, gradually blackening toward middle, and
distal parts near (15"c) Pinkish Cinnamon; total length, 18 to 20 mm; maxi-
mum width 0.8 mm. Setifoims on middorsal region: Gray basally, gradually
blackening toward tip but interrupted by Cinnamon-Buff, subapical zone;
total length 14 to 16 mm; maximum width, 0.06 mm.
Skull. — Slender; bullae large and well-inflated; jugals with conspicuous.
Moo j en: Brazilian Spiny Rats
383
transverse ridge; postorbital process of zygoma moderately developed and
involving only squamosal; incisive foramen short and narrow (4.1x2.1 mm);
vomerine sheath complete, with maxillary part short and thick; mesopterygoid
Fig. 128. Map showing the geographic ranges of the subspecies of Proechimys iheringi.
fossa extending forward as far as posterior parts of second molars; posterior
palatine foramina at plane of premolars.
Teeth. — Upper and lower molariform teeth with two counterfolds. Coun-
terfolds coalesced in P4 and Ml of one specimen.
384 University of Kansas Publs., Mus. Nat. Hist.
Comparisons. — From P. i. gratiosus and P. i. panema, paralus differs in:
all molariform teeth with two, instead of some with fewer, counterfolds;
vomerine sheath complete and thick instead of usually incomplete; incisive
foramen shorter and narrower; aristiforms conspicuously wider; setiforms
Cinnamon-Buff instead of Ochraceous-Buff and Cinnamon, respectively. Tail
96 per cent of head and body in paratus instead of 100 per cent as in panema.
Remarks. — The animals were captured in climax forest.
Specimens examined. — Total number, 3 (MN), from Brazil, Espirito Santo, Santa Teresa,
Floresta da Capela de Sao Braz ; altitude 630 meters.
Proechimys setosus (Desmarest)
General characters. — Size medium; tail approximately same length as head
and body; aristiforms moderately wide; feet rather large; ears of medium
size; color on upper parts and sides sepia gradually changing to Ochraceous-
Tawny; few differentiated, light-colored aristiforms present on outer thighs
and rump; under surface of body and inner sides of legs white; tail with
white tip and conspicuous, white pencil; feet white dorsally; skull short and
smooth, somewhat flattened in interorbital region; jugals narrow dorso-ven-
trally; incisive foramen moderately long and notably narrow; vomerine
sheath complete and slender; postorbital process of zygoma spinelike and
involving mostly jugal; premolars usually with two counterfolds; molars with
only one counterfold, rarely two in Ml or in M3.
Remarks. — The specimens available are undoubtedly faded and,
therefore, the colors mentioned above for the upper parts and sides
may not correspond to the colors of unfaded pelages. Desmarest
(1817:59) describes the color of setosus as similar to that of the
"Echimys de Cayenne" {Proechimys guyannensis) but being more
"rousse." Is. Geoffroy Saint-Hilaire (1840:52) describes the same
animal as being "d'un brun roussatre" on the upper parts.
The Proechimys from Lagoa Santa, Minas Gerais, "Echimys"
elegans Lund, is certainly related to P. iheringi as well as to P. al-
bispinus. From P. iheringi, elegans differs in having a smaller skull
with shorter rostrum, narrower incisive foramen, and orthodont in-
cisors. On the other hand the restricted distribution of the aristi-
forms in the pelage and the white, penicillate tail are points of
resemblance to iheringi. From P. albispinus, elegans differs in hav-
ing a less spinous pelage and longer tail with white pencil instead
of a brown pencil. The skulls, however, are similar, except for the
fact that elegans does not have proodont incisors as albispi?ius some-
times does. Thomas (1921:141) states, after describing the skull of
the type of setosus, that "Specimens corresponding to this animal
have been obtained at Lagoa Santa, Minas, by Lund and others,
and at Bahia." Thomas, however, would not have referred to speci-
mens from "Bahia" as being comparable to elegans had they not
Moojen: Brazilian Spiny Rats 385
been different from albispinus which he discussed in the same paper.
Also, he would not have confused ''specimens comparable to ele-
gans" with a subspecies of P. iheringi {P. i. denigratm, from south-
ern Bahia) which has opisthodont instead of orthodont incisors.
Since French collectors sent material to Europe at the beginning of
the 19th century from (southern?) Bahia, possibly setosus came
from there.
In the collection of the American Museum of Natural History
there is one specimen (AMNH no. 16140) of Proechimys, included
in the so-called Maximilian Collection. The characters of this
specimen agree closely with those of the specimens from Lagoa
Santa. The locality of capture of specimen no. 16140 is unknown,
but it is reasonable to assume that Prince Maximilian zu Wied ob-
tained it somewhere along his route of travel through southeastern
Bahia. Wied (1826:445) mentions "L[oncheres]. myosuros Licht."
as "am Parahyba, am Peruhype und Belmonte," which greatly in-
creases the possibility of its having come from southern Bahia. The
close similarity to elegans of Wied's specimen indicates that the
locality of capture possibly was in the region of the less humid, low
escarpments of southern Bahia.
My conclusion is that Wied's specimen corresponds closely to
setosus and, tentatively, I identify it as such. "Echimys elegans"
due to the relationships mentioned above is here considered to be
a subspecies of setosus.
Among the species described in earlier times, and whose identity
was never ascertained, "Echinomys" fuliginosus Wagner seems to
be synonymous with setosus. Wagner describes the animal as hav-
ing a tail "apicis versus pilis albidis vestita" and the figure of the
cheekteeth (1844, pi. 239 D) shows a typical trilaminate condition
which occurs commonly in elegans. Moreover, the tail of fuligi-
nosus is only 9 per cent shorter than the head and body and the
aristiforms of this subspecies are moderately wide.
Proechimys setosus setosus (Desmarest)
Echimys setosus Desmarest (Geoffroy's MS), 1817, Nouv. Diet. Hist.
Nat. nouv. ed., 10:59 (orig. descr.) ; Is. Geoffroy Saint-Hilaire, 1838,
Comptes Rendus Acad. Sci., Paris, 6(26) :886; Is. Geoffroy Saint-Hilaire,
1840, Mag. Zool., Paris, (ser. 2, annee 2):12, 33, 52; Allen, 1899, Bull.
Amer. Mus. Nat. Hist., 12(20) :257, 261.
Echimys cayennensis Pictet, 1841, Mem. Soc. phys. Hist. Nat., Geneve,
9:145; Waterhouse, 1848, Nat. Hist. Mammalia, 2:334.
Echinomys fuliginosus Wagner, 1843, Schreber's Saugethiere, suppl.
3:343; Wagner, 1844, Schreber's Saugethiere, suppl. 4, pi. 39 D.
Proechimys setosus Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20) :
264; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9) :141; Tate, 1935, Bull.
386
University of Kansas Publs., Mus. Nat. Hist.
Amer. Mus. Nat. Hist., 68(5) :401; Ellerman, 1940, The families and
genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.
Proechimys fuliginosus Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :
400; Ellerman. 1940, The families and genera of living rodents, Brit. Mus.
(Nat, Hist.), 1:119.
Type locality. — Unknown; see remarks under P. selosus. Type: Museum
d'Histoire Naturelle, Paris, no. A. 7787 (Thomas, 1921:141), "very imperfect."
Diagnosis. — Aristiforms wide; P4 and Ml with two counterfolds; p4 with
two counterfolds, one anterior to main fold.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually black-
ening toward tip; total length, 18 to 20 mm; maximum width, 0.8 mm.
Aristojorms on outer thighs: Color much faded; total length, 15 to 17 mm;
maximum width, 0.3 mm. Setiforms on middorsal region: Color faded; total
length. 16 to 18 mm; maximum width, 0.04 mm. Setiforms on outer thighs:
Color faded; total length, 10 to 13 mm; maximum width. 0.03 mm.
Figs. 129-132. Proechimys setosus elegant, sex
Lagoa Santa. X 1.
Skull. — Short; rostrum short and stout; length of nasals approximately
15 mm (broken); bullae roundish, smooth and well-inflated; jugals dorso-
ventrally narrow (3.1 mm) with strong transverse ridge; postorbital process
of zygoma spiniform, slender and involving mostly jugal; incisive foramen
narrow (3.8 x 1.7 mm) and narrowest in posterior part; vomerine sheath com-
plete; posterior palatine foramina obsolete; inesopterygoid fossa extending
forward as far as middle of second molars.
Teeth. — Incisors orthodont. P4 with two counterfolds; Ml with two coun-
terfolds but anterior one notably small; M2 and M3 with only one counter-
fold each. In lower jaw: p4 with two counterfolds, one anterior to main
fold; molars with only one counterfold.
Moojen: Brazilian Spiny Rats 387
Comparisons. — From P. s. elegans, setosus differs in: Ml with two counter-
fold* as opposed to only one; M3 with one counterfold instead of sometimes
with two coimterf olds ; p4 with one counterfold anterior to main fold and
another posterior, instead of both counterfolds posterior.
Remarks. — The measurements above were taken from the Maxi-
imilian specimen mentioned above. Measurements of the type were
given by Desmarest as: head and body, 5% inches, tail about ft1/?
inches. Is. Geoffroy Saint-Hilaire (1838:886) corrects these meas-
urements to: head and body 195 mm; tail (part missing), 170 mm.
Proechimys setosus elegans (Lund)
ElcJmnys]. elegans Lund, 1S41, Kong. Danske Videnskab. Selsk. natur-
vidensk. math. Afhandl., Kjobenhavn, 8:99 (orig. descr.).
Loncheres elegans Lund, 1841, Kong. Danske Videnskab. Selsk. natur-
vidensk. math. Afhandl., Kjobenhavn, 8:245, 266, 294; Wagner, 1843,
Wiegman's Archiv f. Naturg., Berlin, 2 (Jahrg. 9) :47.
Echimys cayennensis Waterhouse, 1848, Nat. Hist., Mammalia, 2:337.
Echinomys cajennensis Winge, 1888, Jordfundne og nulevende Gnavere
(Rodentia), E Museo Lundii, Kjobenhavn, 1(3) :71, pi. 6, figs. 5-6, pi. 7,
fig. 1.
Proechimys setosus Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9) :141.
Proechimys elegans Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :400;
Ellerman, 1940, The families and genera of living rodents, Brit Mus
(Nat. Hist.), 1:119.
Type locality. — Lagoa Santa, Nova Lima, Minas Gerais. Brazil. Type:
Syntypes in Universitets Zoologiske Museum. Kjobenhavn; collected by
P. W. Lund.
Range. — Known only from the type locality.
Diagnosis. — Aristiforms wide; P4 usually with two counterfolds; M3 some-
times with two counterfolds; p4 with two counterfolds anterior to main fold.
Pelage. — Aristiforms on middorsal region: Gray basally, gradually blacken-
ing toward tip; total length, 18 to 20 mm; maximum width, 0.7 mm. Aristi-
forms on outer thighs: Gray basally, gradually blackening toward tip which is
Cinnamon; total length, 15 to 17 mm; maximum width, 0.3 mm. Setiforms
on middorsal region: Whitish basally, gradually blackening toward tip, but
interrupted by Cinnamon, subapical zone; total length, 17 to 19 mm; maxi-
mum width. 0.04 mm. Setiforms on outer thighs: Whitish basally, gradually
blackening toward tip but interrupted by near (15"a) Cinnamon, subapical
zone; total length, 10 to 12 mm; maximum width, 0.03 mm.
Skull. — Short; rostrum short but not stout; length of nasals 17 mm; bullae
large, smooth, and well-inflated; jugals with conspicuous, transverse ridge;
postorbital process of zygoma long, spiniform and constructed entirely of
jugal; incisive foramen narrow (4x1.7 mm); vomerine sheath complete and
slender; posterior palatine foramina obsolete; mesopterygoid fossa extending
anteriorly as far as middle parts of second molars.
Teeth. — Incisors orthodont. P4 usually with two counterfolds, rarely with
three; upper molars with only one counterfold, but M3 sometimes with two,
posterior one being vestigial. Lower molariform teeth: p4 with two counter-
folds, both being anterior to main fold; molars with only one counterfold.
388 University of Kansas Publs., Mus. Nat. Hist.
Comparisons. — Differences from P. s. setosus are given in the account of
that subspecies.
Remarks. — According to Lund, these animals are found in the
vicinity of small pools, swim well in spite of not having webbed
toes, at night go after food and climb the corn stalks, and have
their nests in the grass at the margins of the pools.
Specimens examined. — Total number, 2 (UZM), from Brazil, Minas Gerais, Nova Lima,
Lagoa Santa.
Proechimys albispinus (Is. Geoffroy)
General characters. — Size small; tail of same length as head and body or
slightly less; feet small; ears of medium size; color of upper parts Ochraceous-
Tawny gradually changing to Ochraceous-Buff on sides; differentiated, light-
colored aristiforms on back, sides, rump and at base of tail; clavate aristi-
forms on back with Ochraceous-Tawny or Ochraceous-Buff, subapical zone;
underparts of body and inner sides of legs white; tail blackish above, white
below, with no white tip; hands and feet white on dorsal parts and some
specimens darker on outer margins of feet; skull short and smooth, somewhat
flattened in frontal region; jugal dorso-ventrally wide and with moderately
conspicuous transverse ridge; postorbital process of zygoma well developed
and involving both jugal and squamosal; bullae large and smooth; incisive
foramen short and narrow; vomerine sheath incomplete or complete; molari-
form teeth with only one counterfold; incisors orthodont or proodont.
Remarks. — A good series from Macaco Seco, Andarai, Bahia,
agrees closely with the form first described (albispinus) from the
Island Madre de Deus, in Todos os Santos Bay, Bahia. Compared
with topotypes of P. albispinus sertonius, the animal from Macaco
Seco in general color is more Ochraceous-Tawny and has a narrower
skull with orthodont incisors. Specimens from Bonfim, northeastern
Bahia, on the other hand, agree with Thomas' albispinus sertonius,
from Lamarao, being browner and having broader skulls than P. a.
albispinus and having proodont, instead of orthodont, incisors. The
range of each of the two subspecies is, therefore, fairly extensive.
The insular form extends to the less rainy, continental area and the
form from Lamarao ranges northward (NNW) in the same type of
highly deciduous forest, the "caatinga."
The species albispinus is certainly the most specialized form of
the entire genus for drier habitats. In addition to the general
adaptations described above, it is noteworthy for having both
lanceolate and clavate aristiforms. The latter type has a wide basal
part and an abruptly narrowed, distal part. The same development
is seen in the genus Echimys, where highly spinous forms, like
Echimys paleacea (Lichtenstein), show the same two types of aristi-
forms.
Moojen: Brazilian Spiny Rats
389
134
139
Figs. 133, 135. Proechimys albispinus albispinus, male, CNHM no. 20409, Macaco Seco. X 1.
Figs. 134, 136. Proechimys albispinus sertcmius, male, MN no. 6454, Bonfim. X 1.
Figs. 137, 138. Proechimys albispinus albispinus, male, CNHM no. 20409, Macaco Seco. X 1.
Figs. 139, 140. Proechimys albispinus sertonius, male, MN no. 6454, Bonfim. X 1.
390 University of Kansas Publs., Mus. Nat. Hist.
Proechimys albispinus alhispinus (I*. Geoffroy)
Echimys albispinus Is. Geoffrov Saint-Hilaire, 25 June 1838, Comptes
Rendus Acad. Sci., Paris, 6(26) :886; Is. Geoffroy, August, 1838, Ann. Sci.
Nat. Paris, 10 (ser. 2): 125; Is. Geoffroy, 1840, Mag. Zool., Paris (ser. 2,
annee 2. livr. 13) :33, 53. pi. 26. pi. 29 (figs. 1. 2, 3) ; Allen, 1899, Bull.
Amer. Mus. Nat. Hist., 12(20) :261.
Echinomys fuliginosus Wagner, 1843. Schreber's Siiugethiere, suppl.,
3:343.
Echimys albispinosus Waterhouse, Nat. Hist., Mammalia, 2:341.
Proechimys albispinus Allen, 1899. Bull. Amer. Mus. Nat. Hist., 12(20) :
264; Thomas, 1911, Ann. Mag. Nat. Hist., 8 (ser. 8) :252.
Proechimys albispinus albispinus Thomas, 1921, Ann. Mag. Nat. Hist.,
8 (ser. 9): 141; Tate, 1935. Bull. Amer. Mus. Nat. Hist., 68(5) :401; Eller-
man, 1940. The families and genera of living rodents, Brit. Mus. (Nat.
Hist,), 1:122.
Type locality. — Ilha Madre de Deus, Ilaparica (near Salvador). Bahia,
Brazil. Type: Museum d'Histoire Naturelle, Paris, no. A 7669, "skull . . .
practically perfect" (Thomas, 1921:142).
Range. — Island Madre de Deus, Macaco Seco, Andarai and probably islands
of the bay of Todos os Santos and valley of the Paraguassu River.
Diagnosis. — Aristiforms wide ; color on setiforms Ochraceous-Tawny on up-
per parts and sides; incisors orthodont; molariform teeth with one counter-
fold, p4 rarely with two.
Pelage. — Aristiforms on middorsal region: Lanceolate aristiforms, with
basal part whitish, gradually blackening toward tip ; total length, 25 to 28 mm ;
maximum width, 1.2 mm; clavate aristiforms with base whitish, gradually
blackening toward tip but interrupted by Ochraceous-Tawny, subapical zone.
Aristiforms on outer thighs: Whitish on basal half, gradually blackening to-
ward tip; total length, 24 to 26 mm; maximum width, 0.9 mm. Some are
whitish basally. gradually blackening toward distal part but distal fourth or
fifth near (15';') Ochraceous-Tawny. Setiforms on middorsal region: Whitish
basally. gradually blackening toward tip but interrupted by Ochraceous-
Tawny, subapical zone; total length, 20 to 23 mm; maximum width, 0.1 mm.
Some setiforms almost completely whitish. Setiforms on outer thighs: Whit-
ish basally, gradually blackening toward tip but interrupted by near (15';)
( )chraceous-Tawny, subapical zone; total length, 18 to 20 mm; maximum
width. 0.06 mm.
Skull. — Narrow; bullae small and smooth; jugals dorso-ventrally wide with
conspicuous transverse ridge; postorbital process of zygoma well-developed
and formed by jugal and squamosal; posterior palatine foramina obsolete;
incisive foramen narrow and short; vomerine sheath complete or incomplete
bul premaxillary part at a level lower than that of maxillary part (when skull
is viewed from ventral face) ; mesopterygoid fossa extending forward as far as
anterior borders of second molars.
Teeth. — Incisors orthodont; molariform teeth with only one counterfold,
excepl that p4 rarely has two counterfolds.
Comparisons. — From P. a. serlonius, albispinus differs in: sides of body
darker; incisors orthodont as opposed to proodont; p4 rarely with two coun-
ts folds instead of alwavs with one counterfold.
Moojen: Brazilian Spiny Rats 391
Remarks. — The localities where P. a. albispinus has been collected have a
forest climax with a moderate percentage of deciduous trees.
Specimens examined. — Total number, 19 (18 CMNH, 1 MCZ), from Brazil, Bahia,
Andarai, Macaco Seco.
Proechimys albispinus sertonius Thomas
Proeokimys albispinus sertonius Thomas, July, 1921, Ann. Mag. Nat.
Hist., 8 (ser. 9) :142 (orig. descr.) ; Tate, 1935, Bull. Amer. Mus. Nat. Hist.,
68(5)'-401; Ellerman, 1940. The families and genera of living rodents,
Brit, Mus. (Nat. Hist.), 1:122.
Type locality. — Lamarao ("about 70 miles north of Bahia City"), Ituiutaba,
Bahia, Brazil; altitude 300 meters. Type: British Museum (Nat, Hist.),
no. 3.9.5.S6. adult male; collected on 16 June, 1903, by Alphonse Robert;
original number, 1508.
Range. — Known from the type locality and Bonfim; probably occupies val-
leys of Jacuipe and the Itapicuru rivers and littoral between them.
Diagnosis. — Aristiforms wide; color of setiforms Ochraceous-Tawny on
back, grading to Ochraceous-Buff on sides; incisors proodont; no molariform
tooth with more than one counterfold.
Pelage. — Aristiforms on middorsal region: Lanceolate aristiforms whitish
basally, gradually blackening toward tip; total length, 23 to 27 mm; maxi-
mum width, 1.3 mm. Clavate aristiforms, and some lanceolate ones, whitish
basally, gradually blackening toward tip but interrupted by Ochraceous-
Tawny, subapical zone. Some clavate aristiforms without subapical zone but
blackened in distal part; total length, 23 to 24 mm; maximum width, 0.7 mm.
Aristiforms on outer thighs: Whitish basally, gradually blackening toward tip
but interrupted by Light Ochraceous-Buff, subapical zone; tip slightly darker;
some whitish basally, grayish in middle and light yellowish toward tip; total
length, 20 to 22 mm; maximum midth, 0.9 mm. Setiforms on middorsal re-
gion: Whitish basal part succeeded by grayish, then by long, light, yellowish
band, which becomes Light Ochraceous-Buff, and blackish tip; total length,
26 to 29 mm; maximum width, 0.15 mm. Setiforms on outer thighs: Whitish
basal part succeeded by grayish, then Light Ochraceous-Buff, subapical zone
and blackish tip; total length, 18 to 20 mm; maximum width. 0.13 mm.
Skull. — Broad; bullae small and smooth; jugals dorso-ventrally "wide,"
with conspicuous transverse ridge; postorbital process of zygoma well-devel-
oped and formed by both jugal and squamosal; incisive foramen narrow and
short; vomerine sheath incomplete or complete but premaxillary part on a
lower level than maxillary part (when skull is viewed from ventral face) ;
mesopterygoid fossa extending forward as far as anterior faces of second
molars.
Teeth. — Incisors proodont; molariform teeth with only one counterfold.
Comparisons. — Differences from P. a. albispinus are given in the account of
that subspecies.
Remarks. — Localities where samples were collected are typical
"caatinga" forest, a climax of mainly deciduous trees; cacti are also
common in the region.
Specimens examined. — Total number, 10, from Brazil, Bahia, as follows: Ituiutaba,
Lamarao, 4 (1 DZ, 1 CMNH, 1 MCZ, 1 USNM) ; Bonfim, Bonfim, 6 (5 DZ, 1 MN).
392 University of Kansas Publs., Mus. Nat. Hist.
INCERTA SEDIS
Proechimys myosuros (Lichtenstein)
Loncheres myosuros Lichtenstein, 1818, Das zoologische Museum der
Universitat zu Berlin, (2): 18 (nomen nudum); Lichtenstein, 1820. Ab-
handl. K. Akad. Wissensch., Berlin (1818-1819) :192, pi. 1, fig. 2 (orig.
descr.); Wied, 1826, Beitrage zur Naturgeschichte von Brasilien, 2:445.
Mus leptosoma Brants, 1827, Het geslacht der Muizen door Linnaeus
opgesteld . . ., Berlyn, p. 150; Lichtenstein, 1830, Darstellung neuer
oder wenig bekannter Saugethiere, Berlin, Heft 7, pi. 36. fig. and text
pages.
Mus cinnamoneus Lichtenstein, 1830, Darstellung neuer oder wenig
bekannter Saugethiere, Berlin, Heft 7, pi. 36.
Echimys myosuros Is. Geoffroy Saint-Hilaire, 1838, Comptes Rendus
Acad. Sci., 6(26) :886, and 1840, Mag. Zool., Paris (ser. 2, annee 2):15,
33, 53; Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20) :261.
Echinomys leptosoma Wagner, 1843, Schreber's Saugethiere, suppl.,
3:341.
Echinomys myosuros Burmeister, 1854, Syst. ubersicht Thiere Brasiliens,
p. 199.
Proechimys setosus Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9) :141.
Proechimys myosuros Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5) :
400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus.
(Nat. Hist.), 1:119.
Proechimys leptosoma Tate, 1935, Bull. Amer. Mus. Nat. Hist.. 68(5) :
400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus.
(Nat. Hist.), 1:119.
General characters. — Aristiforms wide (1'") and numerous on dorsal parts
of body; tail longer (9") than head and body (8"); hind feet short (1"6'").
Color (According to Lichtenstein and Brants' descriptions). — Black between
ears; dark brown on middorsal line with reddish tinge on front and upper
side of neck ; posteriorly from shoulders there is a greasy shine added to color ;
this dark, dorsal band widens posteriorly, there encroaching on sides of body;
sides lighter brown, sparsely marked with dark brown lines, from sides of
head caudad to, and including, outer surfaces of hind legs; outer sides of
forelegs colored like outer sides of hind legs; ankles ringed with brown. Tail
blackish above, whitish below. Upper surfaces of hands and feet white.
Skull. — No description of skull or teeth found.
Remarks. — Thomas (1921:140-143) summarized the available in-
formation on the forms from southeastern Brazil and synonymised
myosuros with setosus. The description of leptosoma by Brants ap-
plies to this same species except in a few features. However, neither
Brants nor Lichtenstein described the tail of myosuros as having a
white tip or even as having a heavily pencilled tip, although Wag-
ner (1843:342) in redescribing Lichtenstein's species indicated that
the tail had a white pencil. He gave also measurements of the head
and body, and tail, which do not agree with the original measure-
ments given by either Lichtenstein or Brants. Lichtenstein, in the
Moojen: Brazilian Spiny Rats 393
original description, gave the type locality of myosuros as Bahia,
and stated that it was collected by Freireiss. Brants also gave
Bahia as the type locality for leptosoma. The names leptosoma
Brants and cinnamoneus Lichtenstein are evidently no more than
duplicate names for myosuros, as pointed out by Thomas (1921:
141) and subsequent writers. Specimens referred to any of these
forms, therefore, could take the earlier name myosuros until identi-
fied otherwise. Lichtenstein later (1830, text for plate 36, fig. 2)
added Sao Paulo State to the known range of the species, mention-
ing specimens collected there by Sello. Probably, therefore, Wag-
ner redescribed leptosoma using a composite sample; the white tip
on the tail could occur in any race of P. iheringi from Sao Paulo.
Considering the short hind feet and the wide aristiforms, Pro-
echimys myosuros probably will eventually prove to be related to
albispinus; perhaps it will prove to be a synonym of albispinus.
CONCLUSIONS
1. The genus Proechimys is divisible into two subgenera. In all
Brazil there are four full species of each subgenus, or 8 species
in all. All but one of these are divisible into subspecies of
which there are 29, making a total of 30 kinds in Brazil; 14 of
these are here newly named.
2. It is new information, I think, that: (1) One main fold extend-
ing entirely across the worn crown of the molariform tooth is
peculiar to Trinomys; in the subgenus Proechimys, apparent
complete division of the crown surface is accomplished by a
short main fold meeting a counterfold originating on the op-
posite side of the tooth; (2) progressive decrease in size of
molariform teeth from P4 to M3 is peculiar to the subgenus
Trinomys; in the subgenus Proechimys, M2 is largest and the
teeth are progressively smaller anteriorly.
3. In the one species, Proechimys albispinus, which has the widest
distribution of aristiforms on the body of any species in the
genus, some of the aristiforms are clavate. Clavate aristiforms
occur in the most spiny species of the related genus Echimys.
4. In subspecies of any one full species the incisive foramen is
larger in animals which inhabit arid areas than in those which
inhabit humid areas. Possibly increased area of moist mucosa
associated with Jacobson's organ is required in arid areas for
maintenance of the necessary keenness of smell.
394 University of Kansas Publs., Mus. Nat. Hist.
5. The number of counterfolds in the molariform teeth vary in
clinal fashion. Their variation is in response to humidity. In-
creasing humidity is correlated with increasing number of folds,
and decreasing humidity is correlated with decreasing number
of folds.
6. Clinal variation correlated with increasing humidity is to
be seen also in longer tail and darker color of pelage.
7. The primitive Proechimys probably was large, with a short
tail, narrow aristiforms, strongly built skull, and five counter-
folds in each molariform tooth.
8. Geographic isolation appears to have been a factor in the
establishment of the two subgenera; the arid belt along the
Sao Francisco River and northward to Ceara appears to be
uninhabited by Proechimys and constitutes a barrier separating
the two subgenera, Proechimys and Trinomys.
9. This arid belt probably developed relatively early, since in
deposits of late Pleistocene age, remains of the subgenus
Trinomys have been found in the area where the subgenus
still occurs.
10. The most primitive types occur at the periphery of the range
of the genus.
11. Populations from small islands tend to be more primitive than
populations on the mainland. Insular populations develop a
homozygous condition with resultant disappearance of second-
ary biotypes.
12. Insular animals ordinarily are larger than their mainland coun-
terpart.
Moojen: Brazilian Spiny Rats 395
TABLE OF MEASUREMNTS
Table 1. — Measurements (in millimeters) of adults of Proechimys
£ .2 -is
-3 ~ ,C S ^~ -G O
--I £ ~ Zt S_ .g _Sjg - g-.g J g.|
--° r- r- t- ~ 2?_3 — £ oS"c - "S+3 « 5
— . -C .c .^ .^ c o;^ >hn sea « £ w -^ c &
II f II ll f{ |J II I II | II
_; j _! >j O O n j £ PhO
P. 0. steerei, $ $ Hyutanaham
USNM 105535 218 123 48 17 53.5 44.0 25.2 19.3 11.7 18.2 8.2
USNM 105530 217 135 50 ... 55.2 45.3 25.7 20.7 11.4 19.2 8.0
?
USNM 105537 56.3 44.9 24 0 20 4 11.4 19 3 8 7
P. g. goeldii, $ Fazenda Paraiso
AMNH37489 218 ... 52 20 55.1 44.9 27.0 22.1 12.0 18.6 9.4
9
AMNH 37488 228 157 49 22 57.3 47 6 27 9 22 1 12 4 20 3 9.6
P. s. Uminalis, $ $ Rio Quichito
Mean 229 145 43 21 58.4 47.6 27.3 21.8 12.9 20.1 8.3
Maximum 250 ... 45 24 61.7 50.0 28.9 23.0 13.8 21.2 8.8
Minimum 210 ... 40 18 53.3 44.3 25.5 19.4 12.0 18.5 7.9
No. of specimens 5 1 5 5 5 5 5 5 5 5 5
5 $
MN6253 215 150 43 20 57.5 46.5 28.3 21.5 12.0 19.4 8.7
MX 62.50 • 213 .. 42 20 60.8 49 7 22 0 13 7 210 9.1
P. s. amphichoricus, $ $ Esmeralda
AMNH77000 252 163 53 ... 60.8 50.3 27.5 24.4 13.7 20.7 9.4
AMNH 76994 260 160 54 25.5 22.8 12.6 9.5
AMNH77020 250 181 57 ... 62.0 52.0 27.3 25.9 13.8 21.5 9.6
5
AMNH 76999 235 149 50 24 0 19 0 9.3
P. s. kermiti, 9 Lower Solimoes
AMNH 37124 210 ... 55 . . . 65.2 53.7 29.2 27.6 13.5 21.4 9.0
P. I. brevicauda, $ Joao Pessoa
DZ900 245 147 48 ... 58.3 46.6 26.5 22.5 11.6 18. 2 8.2
P. I. boimensis, $ Boim
MCZ30881 220 160 50 ... 54.6 44.2 24.8 21.0 11.7 17.3 7.5
$
MN 1976 1S2 140 45 ... 52.6 42.2 24.3 20.8 11.4 16.8 7.6
9 Cameta
MCZ 30878 240 ... 48 ... 55.1 45 0 25.2 22.4 11.5 18 4 7.6
P. I. longicaxidatus, $ Urucum
CNHM26732 229 121 48 21.5 11.5 18.5 8.3
99
AMNH 37085 210 150 44 ... 51.4 42.5 24.3 19.5 10.8 17.1 8.9
AMNH 37086 210 ... 50 . . . 48.5 40.9 23.7 17.1 10.2 17.1 8.3
396 University of Kansas Publs., Mus. Nat. Hist.
Table 1. — Measurements (in millimeters) of adults of Proechimys — Continued
* -o 5 -a "3 r. ■$ °
•c>>«-S^g a „ a -£■£ § ^,-g
.. .t! _ _ — o *!_ -a _?3_e _ .So — a Si
I
.-o ^.v. ■* g ,s_* ■? .ax: «- .-e.o r* as
o
oo o o os ^3 ^j- ■£*» o •g-e
pJ3 XJ jSjj^^ ai^4 ^, Ea Sal XI fc; w
T
e§ a a£ si j;o = .2 mx> a So -2 g-3
* 55 a t -O >> oj a i8!^
P. Z. leucomystax, $ Tapirapoa
AMNH37509 230 150 43 ... 50.7 42.2 24.5 18.4 10.3 17.5 8.1
9
AMNH37510 210 ... 42 ... 48.1 41.3 23.0 16.9 9.9 17.2 8.1
9 Utiariti
MN2212 48.0 39.9 18.3 11.2 15.9 8.0
$ Salto Sepotuba
MN 1936 202 147 44 ... 52.6 43.0 23.6 19.2 10.7 17.8 7.8
P. I. roberti, $ $ Anapolis
Mean 208 159 45 21 52.7 43.5 24.9 20.7 13.1 17.2 7.9
Maximum 235 190 55 25 56.1 47.8 27.1 23.7 12.0 19.1 8.2
Minimum 170 135 36 18 48.1 40.0 22.8 18.2 10.6 15.6 7.6
No. of specimens 16 14 16 16 11 11 11 11 11 11 11
99
Mean 219 149 44 20 51.1 42.3 24.1 20.0 10.7 17.2 8.0
Maximum 290 155 48 24 55.5 45.4 25.8 21.5 11.1 17.7 8.7
Minimum 195 125 40 18 48.9 40.3 23.1 19.1 10.5 16.6 7.7
No. of specimens 10 8 10 10 7 7 7 7 7 7 7
P. g. villicauda, $ $ Tapirapoa
MN 1932 225 145 47 ... 55.6 45.5 26.8 24.0 12.0 18.1 8.9
MN 1934 215 162 50 ... 56.2 46.0 26.1 21.3 12.0 18.6 8.4
$ Utiariti
AMNH 57544 250 200 55 24.3 13.1 19.9 9.1
P. g. ribeiroi, $ Rio 12 de Outubro
MN 1935 190 134 47 ... 50.1 41.0 24.3 20.0 11.5 15.9 8.1
P. g. hyleae, $ $ Tauari
Mean 248 146 52 ... 58.1 47.6 27.1 22.9 12.1 19.4 8.5
Maximum 260 174 53 ... 59.0 47.8 .... 23.4 13.2 20.0 8.8
Minimum 217 143 51 ... 57.2 46.5 22.4 11.1 18.8 8.3
No. of specimens 4 34... 2 2 1 2 2 2 2
99
Mean 229 149 50 ... 54.3 44.9 25.8 21.0 11.8 18.1 8.5
Maximum 270 168 54 ... 56.1 46.3 27.4 23.0 13.4 19.3 9.0
Minimum 190 132 49 ... 51.5 42.9 24.5 19.1 11.1 17.1 7.9
No. of specimens 10 9 10 . . . 9 11 11 10 10 10 10
P. g. nesiotes, $ Ilha de Manapiri
CNHM 19496 201 133 47 20 52.7 42.7 25.1 19.5 11.1 17.8 8.0
$
MN 1975 200 152 47 21 52.1 42.6 25.8 19.5 12.3 18.3 8.0
Moojen: Brazilian Spiny Rats 397
Table 1.— Measurements (in millimeters) of adults of Proechimys — Continued
■ sp m - ■*•
-a s 3 ■- "3 ., ~ °
.s >, -^ — tuiJg o a 73 .3 a- Sf-£
J3-0 J ^ ^g 8-g It g^ ^ ■§« £ at
*% «, •&! s| «" Is le * fc§ - l-S
3 3 ffl <u Jc J? >> oj c Jr r$
3 JJJO O N J ~ * o
P. #. leioprimna, 9 $ Cameta
AMNH 37484 192 151 41 22 54.8 44.9 25.4 19.2 12.6 18.4 8.4
CNHM 19503 189 164 47 22 54.4 43.7 26.2 20.5 12.6 18.2 8.2
CNHM 19536 189 ... 43 22
P. g. oris, $ Providencia
CNHM 19495 230 170 45 24 56.1 47.1 25.8 22.0 11.4 17.4 8.3
$ Tanaquara
MN 1974 230 175 49 23 53.2 43.0 23.8 20.1 10.5 17.9 7.7
$ Rio Guama
AMNH 37487 205 142 42 21
P. g. arescens, $ $ Fazenda Inhuma
CNHM26440 206 149 51 24 54.7 44.1 26.3 21.4 12.1 18.7 8.3
CNHM28441 191 164 51 ... 55.6 45.0 25.7 22.4 11.7 18.7 8.7
P. g. riparum, 9 Manaus
AMNH 143018 225 ... 44 20 52.6 43.2 24.0 20.5 11.0 17.2 6.7
P. g. arabupu, $ $ Arabupu
AMNH 75816 243 220 56 ... 59.2 48.7 27.0 23.8 12.9 17.8 8.7
AMNH 75819 230 181 52 ... 55.0 46.0 26.0 22.5 12.3 16.9 8.3
AMNH 75815 228 198 52
99
AMNH75810 226 170 48 ... 53.9 45.6 25.6 21.0 12.0 16.1 8.3
AMNH75823 209 188 48 ... 53.4 43.9 24.5 21.5 11.7 16.4 8.3
AMNH 75817 204 167 47 ... 51.1 43.1 21.6 11.3 16.6 8.2
P. dimidiatus, $ $ Pedra Branca
Mean 199 170 46 ... 52.4 43.5 26.2 19.5 12.1 16.4 8.3
Maximum 220 195 50 ... 56.4 47.1 27.5 21.5 13.6 18.0 8.7
Minimum 180 150 44 ... 48.1 40.4 24.6 17.5 11.0 14.4 7.4
No. of specimens 19 18 19 . . . 45 46 45 45 46 46 46
5 9
Mean 197 162 44 ... 51.8 42.9 25.8 19.4 11.8 16.3 8.3
Maximum 230 180 46 ... 55.1 45.9 27.4 22.0 13.0 18.4 8.9
Minimum 165 145 42 ... 48.6 39.5 23.8 17.6 10.7 14.8 7.7
No. of specimens 14 12 14 . . . 42 44 44 42 44 41 44
P. i. iheringi, $ $ Ilha de Sao Sebastiao
Mean 207 197 48 ... 54.5 44.6 26.2 19.9 12.0 18.3 8.3
Maximum 220 205 50 ... 55.0 45.2 27.1 20.4 12.8 18.7 8.5
Minimum 196 190 46 ... 53.5 43.7 25.9 19.3 10.9 17.5 8.0
No. of specimens 5 25... 5 5 4 4 5 5 5
9 9
MN 6453 228 185 46 ... 54.3 44.5 25.9 20.5 11.0 18.9 8.2
DZ 2095 205 180 46 ... 53.2 42.9 26.4 18.9 11.1 17.0 8.2
DZ2525 205 ... 46 ... 56.9 45.8 27.8 20.9 12.5 18.9 8.2
7—3343
398
University of Kansas Publs., Mus. Nat. Hist.
Table 1. — Measurements (in millimeters) of adults of Proechirnys — Continued
-I
c
J)
■a
-3
■£■»
1.8
■~
<*- o
o c
•S S
►J J
a
a
s
a
J
T3 C
1 Q>
O"
0
C3T)
5 a)
o
ho
c
bo
g
o 2 "
So S3
w Ah
■5*
C -_
P73
P. j. bonafidei, $ $ Fazenda Boa Fe
Mean 211 186 50 25 53.3 43.4 25.6 20.2 12.1
Maximum 220 194 54 26 55.8 45.2 26.3 21.7 13.2
Minimum 200 176 47 24 50.7 41.0 24.2 19.1 11.0
No. of specimens 7 5 8 8 4 5 5 7 7
Mean 209 185 52 25 52.6 44.4 26.6 20.0 12.4
Maximum 226 203 55 27 56.9 46.4 28.6 21.4 13.2
Minimum 185 153 50 22 44.4 42.3 24.9 18.0 11.4
No. of specimens 7 7 7 7 6 6 7 7 7
P. i. gratiosus, $ $ Floresta da Caixa Dagua
Mean 193 191 48 ... 51.2 41.4 25.5 18.5 11.7
Maximum 200 200 49 ... 51.7 42.2 27.0 18.9 12.0
Minimum 185 175 47 ... 50.5 40.4 24.6 18.0 11.1
No. of specimens 5 55... 5 5 5 5 5
29
Mean 204 175 49 24 50.5 42.1 26.0 18.4 11.4
Maximum 220 190 50 26 52.6 43.1 26.6 19.5 12.1
Minimum 195 160 47 22 48.4 41.0 25.3 17.5 10.7
No. of specimens 52544 4 4 4 4
P. i. paratus, $ $ Floresta da Capela de Sao Braz
MN4023 203 195 54 28 51.2 41.7 25.4 18.3 10.4
MN 5458 190 170 51 27
9
MN4012 220 210 54 29 52.2 42.3 25.4 19.1 12.3
P. i. panema, $ $ Campinho
MN8286 215 ... 45 23 54.0 45.1 27.7 19.5 13.4
MN8284 195 ISO 43 23 51.5 41.8 24.5 18.1 11.4
22
MN8288 190 190 46 21 51.6 42.8 25.3 18.1 11.7
MN8287 200 190 46 23 52.6 43.6 27.2 18.4 12.5
MN8285 190 ... 45 24 50.0 41.1 26.6 19.4 12.3
P. i. denigratus, $ $ Mata do Ribeirao da Fortuna
Mean 197 218 52 24 51.5 42.2 25.7 18.3 11.3
Maximum 217 242 54 28 55.4 45.3 27.0 20.3 12.4
Minimum 190 204 50 21 48.7 39.5 23.7 16.8 10.4
No. of specimens 10 99 10 8 8 8 8 8
22
Mean 201 207 52 24 49.1 41.2 25.0 18.3 11.2
Maximum 225 225 54 28 54.1 44.6 25.7 21.5 11.8
Minimum 180 175 49 20 48.2 39.5 23.5 17.4 10.5
No. of specimens 12 12 12 12 8 8 8 8 8
16.3
8.5
17.4
9.1
14.8
8.1
7
7
16.4
8.6
17.0
9.2
15.5
8.2
7
7
16.3
8.0
16.7
8.2
15.6
7.9
5
5
16.3
7.9
17.5
8.2
15.3
7.6
4
4
16.2 8.4
17.5 8.7
16.4
8.1
16.0
7.6
15.7
7.9
16.7
8.3
16.1
8.1
16.0
8.2
17.6
8.5
15.0
8.0
8
8
16.0
7.8
17.1
8.3
15.5
7.5
8
7
Moojen: Brazilian Spiny Rats 399
Table 1. — Measurements (in millimeters) of adults of Proechimys — Concluded
55 — 2 ef ^^
a a *» a) M qS
_ «U — ' Qg
o -S'C <- "-Js
03 — J4
o
•n
T3
J3
>
8
J3 >,
'3
g
^3
8s
s
'3
.9
ih"3
*^t
«-■ o
Sjz
o
o
|8
o a
•° s
■fig
<v
-4- CO
OJ3
c 05
O u,
0 §
0)
a
as
o;o
2-°
i-9
J
.J
1-1
O
o
N
4
3 2 a a
iJ w Ph O
P. s. seiosus, ? No locality
AMNH 16140 39.8 25.2 .... 11.0 15.9 7.6
P. s. elegans, $ Lagoa Santa
UZM H82 190 190 47 25 24.3 17.5 11.2 16.6 7.7
?
UZM L104 48.6 40.8 24.8 16.6 10.3 15.9 7.7
P. a. albispinus, $ S Macaco Seco
Mean 175 162 40 ... 45.9 39.7 23.6 15.6 10.6 15.7 7.6
Maximum 193 173 47 ... 48.2 42.0 24.9 17.1 11.6 17.5 8.2
Minimum 165 147 38 ... 44.2 38.1 22.7 14.8 9.7 14.3 7.2
No. of specimens 9 8 9 ... 11 12 12 14 13 14 13
CNHM 20402 187 171 40 ... 46.0 40.8 23.8 14.9 11.3 16.0 7.4
CNHM 20408 47.6 40.7 24.6 17.1 11.2 15.9 8.2
P. a. sertonius, $ Lamarao
*NHM 18196 190 160 36 25 11.7
$ Boiifim
*IN 6454 185 175 37 ... 45.7 39.8 24.2 15.5 10.7 16.1 7.5
$9
DZ2636 190 150 35 ... 43.8 37.6 22.8 14.7 9.6 14.5 ...
DZ2635 175 155 38 ... 46.7 40.6 23.3 16.9 10.2 16.1 7.4
400 University of Kansas Publs., Mus. Nat. Hist.
LITERATURE CITED
Allen, J. A.
1904. Mammals from the District of Santa Marta, Colombia, collected by
Mr. Herbert H. Smith, with field notes by Mr. Smith. Bull. Amer.
Mus. Nat. Hist., 20 :407-468, November 28, 1904.
1916. Mammals collected on the Roosevelt Brazilian Expedition, with
field notes by Leo E. Miller. Bull. Amer. Mus. Nat. Hist., 35:559-
610, August 9, 1916.
Ameghino, F.
1934. Notas sobre una pequena coleccion de huesos de mamiferos, proce-
dentes de las grutas calcareas de Iporanga en el Estado Sao Paulo
(Brasil), Obras completas y Correspondencia Cientifica de Floren-
tino Ameghino, La Plata, 17:103-153.
Berry, E. W.
1942. Mesozoic and Cenozoic plants of South America, Central America
and the Antilles. Proc. Eighth Amer. Sci. Congress, 4:365-373.
Desmarest, A. G.
1817. Nouveau Dictionaire d'Histoire Naturelle, ed. 2, 10:59.
Ellerman, J. R.
1940. The families and genera of living Rodents. Vol. 1, Rodents other
than Muridae, pp. xxvi -f- 689, 189 text figs. British Museum (Nat-
ural History). June 8, 1940.
Geoffroy Saint-Hilaire, Isidore
1S40. Notice sur les rongeurs epineux, designes par les auteurs sous les
noms d'Echimys, Loncheres Hctcromys et Nelomys. Mag. Zool.
(Guerin-Meneville), s. 2, Ann. 2, pp. 1-57, pis. 20-29.
Lichtenstein, M. H. C.
1830 (1827-34). Darstellung neuer oder wenig bekannter Saugethiere, 65
Arten, etc., pp. 118, 50 pis. colored.
Lund, P. W.
1841. Blik paa Brasiliens Dyreverden for sidste Jordomvaeltning, af Dr.
Lund. Kongelige Danske Videnskabernes Selskabs naturvidenska-
belige og mathematiske. Fortsaettelse af Pattedyrene. Kjobenhavn,
2:217-272, pis. 14-24.
OLrvEiRA, A. I., and Leonardos, O. H.
1943. Geologia do Brasil. Min. da Agricultura. Rio de Janeiro, 26 + 813,
202 figs., photographs, 1 map.
Osgood, Wilfred H.
1944. Nine new South American rodents. Zool. Ser. Field Mus. Nat. Hist.,
29:191-204, July 12, 1944.
Ribeiro, Alipio de Miranda
1914. Historia Natural. Zoologia. Mammiferos. Commissao de Linhas
Telegraficas, Estrategicas de Matto Grosso ao Amazonas. Annexo
no. 5, Rio de Janeiro, pp. 1-49 + 1-3, pis. 25, May, 1914.
Moojen: Brazilian Spiny Rats 401
Ridgway, Robert
1912. Color standards and color nomenclature, iv + 44 pp., 53 pis. Pub-
lished by the author, Washington, D. C.
Tate, G. H. H.
1935. The taxonomy of the genera of Neotropical hystricoid rodents. Bull.
Amer. Mus. Nat. Hist, 68:295-447, June 12, 1935.
1939. The mammals of the Guiana Region. Bull. Amer. Mus. Nat. Hist.,
76:151-229, October 20, 1939.
Thomas, Oldfield
1900. Descriptions of new rodents from western South America. Ann. and
Mag. Nat. Hist., 6(ser. 7) :294-302, September, 1900.
1905. New Neotropical Molossus, Conepatus, Nectomys, Proechimys, and
Agouti, with a note on the genus Mesomys. Ann. and Mag. Nat.
Hist, 15(ser. 7) : 584-591, June, 1905.
1912. On small mammals from the Lower Amazon. Ann. and Mag. Nat.
Hist, 9(ser. 8) :84-90, January, 1912.
1920. On mammals from the Lower Amazons in the Goeldi Museum,
Para. Ann. and Mag. Nat. Hist, 6(ser. 9) :266-283, September, 1920.
1921. On the spiny rats of the Proechimys group from Southeastern Brazil.
Ann. and Mag. Nat. Hist, 8(ser. 9): 140-143, July, 1921.
Wagner, Andreas
1843. Beschreibung einiger neuer Nager, welche auf der Reise des Herrn
Hofrats v. Schubert gesammelt wurden, mit Bezugnahme auf einige
andere verwandte Formen. Abhandl. Akad. Wiss. math.-phys. CI,
3(4): 173-216, pi. 2.
Wagner, J. A.
1844 (1774-1846). In Schreber's Die Saugethiere . . . , 7 pts. including
text and pis, colored (347).
Wied, Maximilian zu
1826. Beitrage zur Naturgeschichte von Brazilien, vol. 2, Mammalia, 600
pp., 6 pis, Weimar.
Winge, Herltjf
1888. Jordf undue og nulevende Gnavere (Rodentia) fra Lagoa Santa,
Minas Geraes, Brasilien. E Museo Lundii Afhandlinger, 1(3): 1-178,
pis. 1-8.
1941. The interrelationships of the mammalian genera. Translated from
Danish by E. Deichmarm and G. M. Allen. K0benhavn, 3 vols.
(vol. 1, xii + 418, 3 pis, 1941; vol. 2, 376 pp., 1941; vol. 3, 308 pp.,
1942.
Transmitted, December 1, 1947.
□
22-3343
15. A new hylid frog from eastern Mexico. By Edward H. Taylor.
Pp. 257-264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Oklahoma.
By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kan-
sas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure
in text. August 16, 1948.
18. A new species of heteromyid rodent from the Middle Oligocene of
northeastern Colorado with remarks on the skull. By Edwin C.
Galbreath. Pp. 285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (Genus Proechimys, Family
Echimyidae). By Joao Moojen. Pp. 301-406, 140 figures in text.
December 10, 1948.
Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
S-/VA-L
v
Three New Beavers from Utah
By
STEPHEN D. DURRANT and HAROLD S. CRANE
mUS. 200L
LIBRARY
flAR -8 1950
UNIVERSITY
University of Kansas Publications
Museum of Natural History
Volume 1, No. 20, pp. 407-417, 7 figs, in text
December 24, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Edward H. Taylor
Volume 1, No. 20, pp. 407-417, 7 figs, in text
December 24, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
I94B
22-3716
Three
MUS. tm?. ZOOL
riAR -S 19! )
i .
NeW Beavers from Jtah
By
STEPHEN D. DURRANT AND HAROLD S. CRANE
The subspecific identity of beavers from Utah seems never to
have been carefully investigated. With the exception of the name
Castor canadensis repentinus applied to animals from Zion and
Parunuweap canyons by Presnall (1938:14), all other writers from
1897 until the present time, have used for animals from Utah, the
name combination Castor canadensis frondator Mearns, the type of
which is from Sonora, Mexico. Study of specimens of beavers from
Utah, accumulated in the collections of the Museum of Zoology,
University of Utah, proves these animals to be far more variable
than formerly supposed, and discloses the existence of three hitherto
unnamed kinds, which are named and described below.
We recognize the need for caution in proposing new names for
American beavers, because the transplanting of these animals from
one watershed to another may have permitted the animals of a
given area to change genetically, say, through hybridization, and
may also have altered the geographic distribution of the several
kinds. The officials of the Utah State Fish and Game Commission
have assured us' that such transplants have not occurred in the
areas where these three new kinds are found, and further that
nowhere in the state have transplants been made from one major
drainage system to another; such transplants as have been made were
only within the same major drainage system.
The capitalized color terms used in this paper are after Ridgway,
Color Standards and Color Nomenclature, Washington, D. C, 1912.
All measurements are in millimeters. We are indebted to the offi-
cials of the United States National Museum for the loan of com-
parative materials.
Castor canadensis pallidus new subspecies
Type. — Female, adult, skin and skull, number 719, Museum of Zoology, Uni-
versity of Utah; Lynn Canyon, 7,500 ft., Boxelder County, Utah; September
7, 1932 ; collected by W. W. Newby.
Range. — Known only from the Raft River Mountains.
Diagnosis. — Size small; tail and hind foot short (see measurements). Color
(type): Pale, upper parts uniformly Ochraceous-Buff; underfur Snuff Brown;
underparts uniformly Light Buff, grading to Light Ochraceous-Buff at base of
tail; underfur Light Drab; front and hind feet Light Ochraceous-Buff. Skull:
Rostrum short; nasals broad (breadth averaging 54 per cent of length), con-
(409)
410 University of Kansas Pubs., Mus. Nat. Hist.
stricted posteriorly and barely projecting posteriorly beyond premaxillae;
zygomatic arches robust, but not widely spreading (zygomatic breadth 77 per
cent of basilar length) ; mastoid breadth 73 per cent of zygomatic breadth ;
anterolateral margin of orbit narrow (6.2) ; occipital condyles visible from
dorsal view; condjdobasal length greater than occipitonasal length; upper
incisors narrow (Orange Chrome in color) ; eoronoid processes high and wide ;
cheek teeth narrow.
Measurements. — Measurements of the type are as follows: Total length,
1040; length of tail, 380; length of hind foot, 157; length of ear, 35; occipitonasal
length, 129.1; basilar length, 116.6; mastoid breadth, 65.6; interorbital breadth,
23.6; length of nasals, 43.3; zygomatic breadth, S9.7; breadth of nasals, 23.4;
alveolar length of upper molariform teeth, 30.4.
Comparisons. — From topotypes and near topotypes of Castor canadensis
taylori, C. c. pallidus differs as follows: Size smaller; tail and hind foot
shorter. Color: Markedfy lighter throughout. Skull: Nasals shorter and
wider (breadth of nasals averages 54 per cent of length of nasals, as opposed
to 46 per cent); nasals barely projecting posteriorly beyond premaxillae; ros-
trum shorter; zygomatic breadth relative to basilar length less; mastoid breadth
actually as well as relatively greater; interorbital breadth greater; occipitonasal
length shorter rather than longer than cond3'lobasal length; tympanic bullae
smaller; eoronoid process higher and wider; cheek teeth narrower.
From specimens of Castor canadensis baileyi, from 20 miles north northeast
of Elko, Elko County, Nevada, C. c. pallidus differs as follows: Body smaller;
tail longer; hind foot shorter; ears shorter: Color: Markedly lighter through-
out. Skull: Larger; nasals shorter and wider (breadth of nasals averages
54 per cent of length of nasals as opposed to 41 per cent) ; nasals barely pro-
jecting posteriorly beyond premaxillae; rostrum broader; zygomatic breadth
relative to basilar length less; mastoid breadth actually as well as relatively
greater; occipitonasal length less rather than greater than condj'lobasal length;
tympanic bullae smaller; eoronoid process higher and wider; cheek teeth
narrower.
From one topotype and two specimens of Castor canadensis repentinus,
from the Colorado River at Yuma, Yuma County, Arizona, C. c. pallidus differs
as follows: Tail and hind foot shorter. Color: Lighter throughout. Skull:
Narrower; nasals shorter and wider (breadth of nasals averages 54 per cent
of length of nasals as opposed to 47 per cent) ; nasals barely projecting pos-
teriorly beyond premaxillae; rostrum shorter; z3Tgomatic breadth relative to
basilar length less; mastoid breadth actually as well relatively greater;
tympanic bullae narrower and smaller; eoronoid process higher and wider;
check teeth narrower.
From one specimen of Castor canadensis concisor, from Trappers Lake,
Garfield County, Colorado, and from the original description of that sub-
species (Warren and Hall, 1939: 35S), C. c. pallidus differs as follows: Size
smaller. Color: Markedly lighter throughout. Skull: Smaller, narrower;
nasals shorter and wider (breadth of nasals averages 54 per cent of length of
nasals as opposed to 48 per cent) ; rostrum shorter; zygomatic breadth relative
to basilar length less; mastoid breadth relative to zygomatic breadth greater;
tympanic bullae narrower and smaller; jugals narrower; distal end of meatal
Durrant: Three New Beavers from Utah 411
tube smaller; coronoid process shorter and wider; angular process shorter and
rounded rather than nearly pointed; cheek teeth narrower.
From the type and near topotypes of Castor canadensis rostralis, C. c.
pallidas differs as follows: Size smaller; tail and hind foot shorter. Color:
Markedly lighter throughout. Skull: Smaller and narrower; rostrum shallower
and narrower; posterior end of nasals more constricted and barely projecting
posteriorly beyond premaxillae; zygomatic breadth relative to basilar length
less; mastoid breadth actually as well as relatively greater; dorsal surface of
lacrimal bone larger; tympanic bullae narrower; coronoid process higher and
wider; angular process not projecting so far caudad; check teeth narrower.
From the type and near topotypes of Castor canadensis duchesnei, C. c.
pallidas differs as follows: Size smaller; tail and hind foot shorter. Color:
Lighter throughout. Skull: Shorter, narrower and less massive; nasals shorter
and wider (breadth of nasals averages 54 per cent of length of nasals as op-
posed to 46 per cent) ; nasals barely projecting posteriorly beyond premaxillae;
rostrum shorter and narrower; zygomatic breadth relative to basilar length
less; mastoid breadth actually as well as relatively greater; tympanic bullae
narrower and smaller; coronoid process higher and wider; angular process not
projecting so far caudad; cheek teeth narrower.
Remarks. — The Raft River Mountains of extreme northwestern
Utah, where C. c. pallidus occurs, are the only mountains of the
state within the drainage of the Snake River. The Snake River
proper lies 50 miles to the northward in Idaho and contains another
kind of beaver, C. c. taylori (Davis, 1939 : 273) . Although occurring
within the same- drainage as C. c. taylori, C. c. pallidus is as distinct
from it as from any other named kind. The relationships of C. c.
pallidus, as indicated by the short rostrum and short, wide nasals,
are rather more with C. c. rostralis of the Wasatch Mountains, than
with C. c. taylori.
The pale color of the animals belonging to C. c. pallidus was noted
at the time of capture, and is the same in the young specimen
(625 mm. total length) as in the type, an adult.
Spechnens examined. — Total, 2, distributed as follows: Boxelder County: Raft River.
5 mi. S Yost, Raft River Mountains, 6,000 ft., 1; Lynn Canyon, Raft River Mountains,
7,f>00 ft., 1.
Castor canadensis rostralis new subspecies
Type. — Male, young adult, skin and skull, number 5199, Museum of Zoology,
University of Utah; Red Butte Canyon, Fort Douglas, 5,000 ft., Salt Lake
County, Utah; October 13, 1947; collected by Harold S. Crane and Clifton M.
Greenhalgh, original number 446 of Crane.
Range. — Known from the western streams of the Wasatch Mountains;
probably occurs in all streams draining westward into the basin of Pleistocene
Lake Bonneville.
Diagnosis. — Size large; tail and hind foot long (see measurements). Color
(type): Upper parts Snuff Brown, purest on head; underfur Brownish Black
412 University of Kansas Pubs., Mus. Nat. Hist.
(2) ; base of tail Cinnamon Buff; hind feet Carob Brown; ears Blackish Brown
(2) ; underparts Auburn, grading posteriorly to Cinnamon Buff; underfur Light
Drab. Skull: Large, massive; nasals short and broad (breadth averaging 54 per
cent of length) and moderately convex transversely ; rostrum deep and broad ;
ventral surface of rostrum moderately concave dorsally ; dorsal surface of
lacrimal bone small; frontal region generally flat; zygomatic arches robust
and widely spreading (zygomatic breadth averaging 82 per cent of basilar
length).
Measurements. — Measurements of the type and average and extreme cranial
measurements of 6 unsexed adults from Charleston, are, respectively, as fol-
lows: Total length, 1,330; length of tail, 470; length of hind foot, 170; length
of ear, 34; occipitonasal length, 128.2, 134.3 (142.1-129.5); basilar length, 112.4,
117.2 (128.2-113.2); mastoid breadth, 62.5, 64.3 (68.9-60.2); interorbital breadth,
27.9, 26.0 (26.9-26.0); zygomatic breadth, 91.3, 93.8 (105.8-90.8); length of
nasals, 44.9, 43.9 (51.3-41.5); breadth of nasals, 24.5, 23.7 (25.7-22.0); alveolar
length of upper molariform teeth, 28.3, 30.9 (32.5-28.7).
Comparisons. — From topotypes and near topotypes of Castor canadensis
taylori, C. c. rostralis differs as follows: Color: Darker on upper parts owing
to darker underfur, guard hairs actually lighter. Skull: Longer; nasals shorter
and wider (breadth of nasals averages 54 per cent of length of nasals as opposed
to 47 per cent) ; extension of nasals posterior to premaxillae less ; rostrum
shorter, broader and deeper; dorsal surface of lacrimal bone smaller; zygomatic
breadth relative to basilar length greater; mastoid breadth relative to zygo-
matic breadth less; coronoid process shorter; coronoid and condyloid processes
farther apart and space between them shallower.
From one topotype and two specimens from the Colorado River at Yuma,
Yuma County, Arizona, of Castor canadensis rcpentinus, C. c. rostralis differs
as follows: Size larger; tail longer. Color: Darker throughout. Skull:
Longer; nasals shorter and wider (breadth of nasals relative to length of
nasals averages 54 per cent as opposed to 47 per cent) ; extension of nasals
posterior to premaxillae less; rostrum shorter, deeper and wider; zygomatic
breadth relative to basilar length greater; mastoid breadth actually as well as
relatively greater; dorsal surface of lacrimal bone smaller; coronoid and
condylar processes farther apart and space between them shallower.
From specimens of Castor canadensis baileyi, from 20 miles north north-
east of Elko, Elko County, Nevada, C. c. rostralis differs as follows: Size
larger; tail and hind foot longer. Color: Darker throughout. Skull: Larger
in all measurements taken; nasals markedly wider (breadth of nasals relative
to length of nasals averages 54 per cent as opposed to 41 per cent) ; extension
of nasals posterior to premaxillae less; dorsal surface of lacrimal bone smaller;
mastoid breadth relative to zygomatic breadth less.
From one specimen of Castor canadensis concisor, from Trappers Lake,
Garfield County, Colorado, and from the original description of that subspecies
(Warren and Hall, 1939 : 358), C. c. rostralis differs as follows: Color: Guard
hairs lighter; underfur darker (blackish as opposed to brownish). Skull:
Longer and narrower; nasals broader and shorter (breadth of nasals averages
54 per cent of length of nasals as opposed to 48 per cent) ; dorsal surface of
lacrimal bone smaller; distal end of mcatal tube smaller; distal end of angular
Durrant: Three New Beavers from Utah 413
process rounded rather than pointed; coronoid process shorter; coronoid and
condylar processes farther apart and space between them shallower.
Among known kinds of Castor canadensis, C. c. rostralis is most like Castor
canadensis duchesnei, from which the former subspecies differs as follows: Tail
and hind foot longer. Color: Darker throughout. Skull: Nasals shorter and
wider (breadth of nasals averages 54 per cent of length of nasals as opposed to
46 per cent) ; nasals less arched transversely ; rostrum shorter, deeper and
broader; ventral surface of rostrum less concave dorsally; dorsal surface of
lacrimal bone smaller.
For comparison with Castor canadensis pallidas, see account of that sub-
species.
Remarks. — Animals from Kamas, in the drainage of the Weber
River, are intergrades between C. c. rostralis and C. c. duchesnei, but
their short, wide nasals and wide rostra make them referable to C. c.
rostralis.
The available specimens of C. c. rostralis are all from streams
which ultimately empty into Great Salt Lake, which is in the north-
ern part of the basin of Pleistocene Lake Bonneville. Some streams
drain into the Lake Bonneville Basin without emptying into Great
Salt Lake proper. Beavers from these streams, we suspect, when
they become known, will be found to be related to C. c. rostralis.
Specimens examined. — Total, 16 (2 skins and skulls, 14 skulls only), distributed as fol-
lows: Summit County: Kamas, 5,500 ft., 6. Salt Lake County: Red Butte Canyon, Fort
Douglas, 5,000 ft., 1; Millcreek Canyon, 6 mi. above mouth, 7,000 ft., 1. Wasatch County:
Charleston, Heber Valley, 5,500 ft., 8.
Castor canadensis duchesnei new subspecies
Type.— Male, young adult, skin and skull, number 4625, Museum of Zo-
ology, University of Utah; Duchesne River, 10 miles northwest of Duchesne,
5,600 ft., Duchesne County, Utah; September 23, 1946; collected by Dave
Thomas, original number 160 of K. R. Kelson.
Range. — Drainage of the Duchesne and White rivers in Utah and Colorado.
Diagnosis. — Size large; tail long (see measurements). Color (type): Upper
parts Sayal Brown, purest on head, grading to Cinnamon Buff at base of tail;
underfur Fuscous; hind feet Burnt Umber; ears Fuscous Black; underparts
Tawny Olive; underfur Smoke Gray. Skull: Large, massive; nasals long,
slender (breadth averaging 46 per cent of length) and markedly convex trans-
versely; rostrum long and attenuate; zygomatic arches heavy and widely
spreading (zygomatic breadth averaging 81.5 per cent of basilar length) ; ventral
surface of rostrum markedly concave dorsally, especially immediately behind
upper incisors; nasals extend posterior to premaxillae.
Measurements. — Measurements of the type and average and extreme cranial
measurements of 9 unsexed adults, from Currant Creek, are, respectively, as
follows: Total length, 1,176; length of tail, 458; length of hind foot, 165; length
of ear, 33; occipitonasal length, 123.6, 132.1 (138.5-122.3); basilar length, 98.6.
114.4 (125.8-99.2); mastoid breadth. 60.4, 65.1 (67.2-64.1); interorbital breadth,
414
University of Kansas Pubs., Mus. Nat. Hist.
Figs. 1-4 Dorsal views of skulls of Castor canadensis. X Vz
Fig. 1. Castor canadensis rostralis, male, young adult, no. 5199 (holotype), Mus. Zool.,
Iniv. Utah.
Fig. 2. Castor canadensis pallidus, female, adult, no. 719 (holotype), Mus. Zool., Univ.
Utah.
Fio. 3. Castor canadensis duchesnei, male, young adult, no. 4625 (holotype), Mus. Zool.,
Univ. Utah.
Fig. 4. Castor canadensis concisor, male, adult, no. 2090, Mus. Nat. Hist., Univ. Kansas.
from Trappers Lake, Garfield County, Colorado, obtained by L. L. Dyche, October 22, 1891.
Durrant: Three New Beavers from Utah
n:»
23.0, 25.1 (26.1-23.7) ; zygomatic breadth, 88.3, 94.2 (99.7-89.5) ; length of nasals,
46.1, 48.4 (51.5-46.2) ; breadth of nasals, 20.5, 22.5 (24.5-18.8) ; alveolar length of
upper molariform teeth, 28.9, 29.9 (32.2-26.5) .
Comparisons. — From topotypes and near topotypes of Castor canadensis
Laylori, C. c. duchesnci differs as follows: Color: Guard hairs lighter, underfill-
darker. Skull: Nasals narrower; rostrum narrower; mastoid breadth relative to
zygomatic breadth less; zygomatic breadth relative to basilar length greater;
tympanic bullae narrower and smaller; cheek teeth narrower.
From specimens of Castor ca7ia<lensis baileyi, from 20 miles north northeast
of Elko, Elko County, Nevada, C. c. duchesnei differs as follows: Size larger;
tail and hind foot longer. Color: Guard hairs lighter, underfur darker. Skull:
Figs. 5-7 Lateral views of left side of skulls of Castor canadensis. X ¥-i
Fig. 5. Castor canadensis rostralis. male, young adult, no. 5199 (holotype), Mus. Zoo!..
Univ. Utah.
Fig. 6. Castor canadensis pallidus, female, adult, no. 719 (holotype), Mus. Zool., Univ.
Utah.
Fig. 7. Castor canadensis duchesnei, male, young adult, no. 4625 (holotype), Mus. Zool.,
Univ. Utah.
416 University of Kansas Pubs., Mus. Nat. Hist.
Larger in all measurements taken ; nasals broader and longer (breadth of nasals
averages 46 per cent of length of nasals as opposed to 41 per cent) ; rostrum
broader and longer; mastoid breadth relative to zygomatic breadth less; tym-
panic bullae larger.
From one specimen of Castor canadensis concisor, from Trappers Lake, Gar-
field County, Colorado, and from the original description of that subspecies
(Warren and Hall, 1939: 358), C. c. duchesnei differs as follows: Color:
Lighter throughout. Skull: Nasals more convex transversely; rostrum nar-
rower; ventral border of rostrum more concave dorsally, especially immediately
behind upper incisors; distal end of meatal tube smaller; angular process shorter
and rounded rather than pointed; cheek teeth smaller.
Among known subspecies of Castor canadensis, C. c. duchesnei is most like
Castor canadensis repentinus, but differs from the latter as follows: Size larger;
hind foot shorter. Color: Darker throughout. Skull: Basilar length less;
mastoid breadth greater; nasals shorter and narrower; extension of nasals pos-
terior to premaxillae less; nasals more convex transversely; cheek teeth smaller.
For comparisons with Castor canadensis pallidus and Castor canadensis ros-
tralis, see accounts of those subspecies.
Remarks. — The extent of the range of C. c. duchesnei within the
drainage of the White River is not definitely known. Three animals
from 9% miles southwest of Pagoda Peak, Rio Blanco County, Colo-
rado, from the drainage of the White River, are intergrades between
C. c. concisor and C. c. duchesnei. They are like the latter subspecies
in shape and length of the nasals, less expanded distal end of the
meatal tube and the rounded angular process, and it appears best,
pending the acquisition of more material, to refer them to C. c.
duchesnei. Another specimen, number 2090, Museum of Natural
History, University of Kansas, from Trappers Lake, Garfield
County, Colorado, at the headwaters of the White River, and only
16 miles distant from the three aforementioned animals, is, however,
nearly typical of C. c. concisor. Relying upon the original description
(Warren and Hall, 1939: 358), this animal is like C. c. concisor in
size and shape of the jugals, in size of the distal end of the meatal tube
and in the pointed end of the angular process. Warren and Hall
{loc. cit.) noted that animals assignable to C. c. concisor occurred
throughout the mountainous parts of Colorado, and recorded them
from the headwaters of nearly all the major rivers of that state.
Apparently C. c. concisor also occurs in the headwaters of the White
River, while the main part of the river is inhabited by animals re-
ferable to C. c. duchesnei.
Specimens examined. — Total, 15 (4 skins and skulls, 11 skulls only), distributed as follows:
Utah: Wasatch County: Currant Creek, Strawberry Valley, 6,000 ft., 11. Duchesne County:
Duchesne River, 10 mi. NW Duchesne, 5,600 ft., 1. Colorado: Rio Blanco County: 9%
mi. SW Pagoda Peak, 7,700 ft., 3 (Museum of Natural History, University of Kansas).
Duerant: Three New Beavers from Utah 417
LITERATURE CITED
Davis, William B.
1939. The Recent mammals of Idaho. The Caxton Printers, Caldwell
Idaho, 400 pp., 2 full page half-tones, 33 figs, in text, April 5, 1939.
Presnall, C. C.
1938. Mammals of Zion-Bryce and Cedar Breaks. Zion-Bryce Mus. Bull.,
2:1-20, January, 1938.
Warren, Edward R. and Hall, E. Raymond.
1939. A new subspecies of beaver from Colorado. Journ. Mamm., 20:358-362,
1 map, August 14, 1939.
Transmitted, May 15, 1948.
□
2-2-371(5
Two New Meadow Mice from Michoacan
Mexico
BY
E. RAYMOND HALL
ftSUS. CO^P. 20GL
LIBRARY
MAR -8 13! ;
University of Kansas Publications
Museom of Natural History
Volume 1, No. 21, pp. 423-427, 6 figs, in text
December 24, 1948
University of Kansas
LAWRENCE
1948
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Edward H. Taylor
Volume 1, No. 21, pp. 423-427, 6 figs, in text
December 24, 1948
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1948
22-3717
MAR -8 19' I
Two New Meadow Mice from Michoacan, Mexico
By
E. RAYMOND HALL
In preparing a list of kinds of mammals of which specimens have
been saved from the Mexican state of Michoacan, two heretofore
unrecognized subspecies of the Mexican meadow mouse, Microtus
mexicanus, have been found. Names for these and descriptions are
given below.
Microtus mexicanus fundatus new subspecies
Type. — Male, adult, skin and skull; No. 100637, Univ. California Mus. Vert.
Zool.; 3V2 mi. S Patzcuaro, 7,900 ft., Michoacan, Mexico; March 9, 1943; ob-
tained by E. R. Hall, original no. 5882.
Range. — Known only from the vicinity of the type locality.
Diagnosis. — Size large (see measurements) ; color brown overlain with cin-
namon; nasals expanded distally with premaxillary borders concave laterally;
posterior border of orbit inclined posterolaterally ; preorbital region and
interparietal region depressed; incisive foramina narrow; zygomatic arches
parallel; tympanic bullae much inflated.
Comparisons. — Among named subspecies of Microtus mexicanus, M. m.
fundatus most closely resembles M. m. mexicanus but differs as follows: Larger
in all parts measured; pelage with slightly less buffy color and with the buffy
color that is present of a slightly lighter tint; posterior two-thirds of pre-
maxillary border of each nasal concave rather than straight ; posterior border of
orbit forming more acute angle with sagittal plane of posterior part of skull;
superior outline of nasals straight rather than depressed in posterior part;
tympanic bullae more inflated both in vertical and horizontal planes.
From M. c. salvus, the subspecies next to the westward, jundatus differs as
follows: Averaging larger in all parts measured; less reddish on upper parts;
underparts with more reddish color but the reddish of lighter tint; perineal
region buffy instead of plumbeous; nasals with premaxillary borders laterally
concave rather than straight; superior outline of skull with nasal segment
sloping anteroventrally and interparietal segment sloping posteroventrally
rather than straight; posterior margin of orbit inclined posterolaterally thus
forming an acute angle, instead of a right angle, with sagittal plane of brain-
case; zygomatic arches parallel rather than bowed outward; incisive foramina
narrower; tympanic bullae more inflated in vertical plane.
Relying on Bailey's (N. Amer. Fauna, 17:55, 1900) description of Microtus
Julviventer, jundatus differs in much larger tympanic bullae.
Remarks. — The series of 59 specimens includes individuals of
several ages of both sexes. This has been a great advantage in mak-
ing comparisons with individuals of geographically adjoining sub-
species since individuals of the same age and sex could be compared.
(425)
426
University of Kansas Pubs., Mus. Nat. Hist.
When the skulls are laid top-side down on a flat surface the occiput
is much higher than in salvus.
Our specimens, taken in the dry season, were trapped mostly in
runways beneath a dense growth of grass underneath a rail fence.
Specimens examined.- — Total, 59, distributed, with respect to the town of Patzcuaro, as
follows: 31/2 mi. S, 7,900 ft., 9; 4 mi. S, 7,S00 ft,, 16; 5 mi. S, 7,800 ft., 26; 9 mi. SE,
8,000 ft., 8.
4 6
Figs. 1-6. Skulls of the type specimens of two subspecies of
Microtus viericanus. XI.
Figs. 1-3. Microtus mcxicanus Jundatus.
Figs. 4-6. Microtus mcxicanus salvus.
Microtus mexicanus salvus new subspecies
Type. — Female, adult, skin and skull; No. 52099, Chicago Natural History
Museum; Mount Tancitaro, 11,400 ft., Michoacan, Mexico; July 19, 1941;
obtained by F. C. Wonder, original No. 1163.
Range. — Known only from Mount Tancitaro at elevations of 7,800 to 11,400
feet.
Diagnosis. — Size medium (see measurements) ; color brown overlain with
cinnamon; premaxillary borders of nasals straight; superior outline of skull
nearly straight; posterior margin of orbit at right angle with long axis of skull;
zygomatic arches bowed outward; incisive foramina wide; tympanic bullae
small.
Comparisons. — From topotypes of its nearest relative, Microtus mcxicanus
phaeus Merriam, salvus differs as follows: Tail shorter; pelage with slightly
more buffy (reddish) pigment which, nevertheless, is of a lighter tint; pre-
maxillary borders of nasals straight rather than concave; superior outline of
skull more nearly straight (less convex) ; inferior border of rostrum more
nearly straight (less concave distally) ; palatal fossae uniformly shallower.
Comparison with M. m. jundatus is made in the account of that subspecies.
Hall: Two New Meadow M . 427
Remarks. — The degree of difference between M. m. salvus and
M. m. jundatus exceeds that between M. m. salvus and M. m. phaeus
or that between M. m. jundatus and M. m. mexicanus.
Specimens examined. — Total, 14, from Mount Tancitaro, Miehoacan, distributed, accord-
ing to elevation above sea level, as follows: 11,400 feet, 8; 11,000 ft., 2; 7,800 ft., 1; no
elevation recorded, 3.
Measurements of the two subspecies. — Average and extreme measurements
of ten adult males of M. m. fundatus and eight adult males of M. m. salvus,
are, respectively, as follows: Total length, 147(135«158), 141(134-146); length
of tail, 33.4(30-39), 29.6(25-32); length of hind foot, 21.1(20-22), 20.0(18-21);
eondylobasal length, 27.7(27.0-29.0), 25.9(25.5-26.2); occipitonasal length, 27.5
(26.8-2S.5), 25.7(25.3-26.0) ; length of nasals, 8.0(7.7-8.4), 7.5(8.9-8.1) ; zygomatic
breadth, 16.4(15.9-17.2), 15.0(14.6-15.3); interorbital breadth, 3.7(3.5-3.9), 3.5-
(3.3-3.6); mastoid breadth, 12.7(12.2-13.2), 11.7 (11.7-11.7); alveolar length of
upper molar series, 7.0(6.8-7.3), 6.7(6.5-6.9); width of rostrum, 5.7(5.1-5.9),
5.3(5.3-5.3); palatilar length, 13.3(12.7-14.3), 12.6(11.9-12.9). Of M. m. salvus
only two specimens yield data for the first, second, sixth and eighth cranial
measurements named above.
For the loan of comparative material I am grateful to Dr.
Hartley H. T. Jackson and Mr. Stanley P. Young of the Biological
Surveys Collection in the United States National Museum, Messrs.
Karl P. Schmidt and Collin C. Sanborn of the Chicago Natural
History Museum, and for assistance with the field work to the John
Simon Guggenheim Memorial Foundation and to Miss Annie M.
Alexander.
Transmitted Jane 1, 1948.
□
22-3717
An Annotated Check List of the Mammals
of Michoacan, Mexico
BY
E. RAYMOND HALL and BERNARDO VILLA R.
iiiu-i.
1AR -8 @50
University of Kansas Publications
Museum of Natural History
Volume 1, No. 22, pp. 431-472, 2 plates, 1 figure in text
December 27, 1949
University of Kansas
LAWRENCE
1949
UNIVERSITY OF KANSAS PUBLICATIONS
The University of Kansas Publications, Museum of Natural His-
tory, are offered in exchange for the publications of learned societies
and institutions, universities and libraries. For exchanges and in-
formation, address the Exchange Desk, University of Kansas Li-
brary, Lawrence, Kansas, U. S. A.
Museum of Natural History. — E. Raymond Hall, Chairman, Editorial Com-
mittee.
This series contains contributions from the Museum of Natural Histoiy.
Cited as Univ. Kans. Publ., Mus. Nat. Hist.
Vol. 1. 1. The pocket gophers (genus Thomomys) of Utah. By Stephen D.
Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces pluvialis Cope, and noteworthy
records of other amphibians and reptiles from Kansas and Okla-
homa. By Hobart M. Smith. Pp. 85-89. August 15, 1946.
3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp.
93-96, 1 figure in text. August 15, 1946.
4. Hybridization between two species of garter snakes. By Hobart M.
Smith. Pp. 97-100. August 15, 1946.
5. Selected records of reptiles and amphibians from Kansas. By John
Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.
6. Kyphosis and other variations in soft-shelled turtles. By Hobart
M. Smith. Pp. 117-124. July 7, 1947.
7. Natural history of the prairie vole (Mammalian genus Microtus).
By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6,
1947.
8. The postnatal development of two broods of great homed owls
(Bubo virginianus) . By Donald F. Hoffmeister and Henry W.
Setzer. Pp. 157-173, 5 figures in text. October 6, 1947.
9. Additions to the list of the birds of Louisiana. By George H.
Lowery, Jr. Pp. 177-192. November 7, 1947.
10. A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-
216. November 29, 1947.
11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R.
and E. Raymond Hall. Pp. 217-236, 2 figures in text. November
29, 1947.
12. A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest
and E. Raymond Hall. Pp. 237-244, 6 figures in text. December
10, 1947.
13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By
Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure
in text. December 10, 1947.
14. A new pocket gopher (Thomomys) and a new spiny pocket mouse
(Liomys) from Michoacan, Mexico. By E. Raymond Hall and
Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
An Annotated Check List of the Mammals
of Michoacan, Mexico
BY
E. RAYMOND HALL and BERNARDO VILLA R.
University of Kansas Publications
Museum of Natural History
Volume 1, No. 22, pp. 431-472, 2 plates, 1 figure in text
December 27, 1949
MUS. COM?. Z08L
LIBRARY
riAR -8 1350
llil Ml >• ■
r LUU - .-
University of Kansas
LAWRENCE
1949
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Edward H. Taylor, Robert W. Wilson
Volume 1, No. 22, pp. 431-472, 2 plates, 1 figure in text
December 27, 1949
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD V01LAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1849
22-6113
tfUS. ! (OOL
LS;
flAR -8 19!
"
An Annotated Check |^ist of the Mammals of Michoacan,
Mexico
-■
By
E. RAYMOND HALL and BERNARDO VILLA R.
INTRODUCTION
When General Lazaro Cardenas was President of the Republic
of Mexico, encouragement was given by his administration to lin-
guistic groups of native American peoples to record in printed form,
eventually in their native languages, accounts of their cultural ac-
complishments and accounts of the natural resources of the regions
concerned. For the Tarascan ''Empire" centering in the state of
Michoacan, a committee of Mexicans and citizens of the United
States of America was formed to forward these aims. Under the
leadership of ethnologists on the committee, especially Professor
Daniel Rubin F. de la Borbolla and Professor Ralph L. Beals, invi-
tations to cooperate in the studies were extended to biologists. One
of us (Hall) was invited to investigate the fauna of native wild
mammals. In 1943, assisted by a fellowship which Hall at that
time held from the John Simon Guggenheim Memorial Foundation,
and with support from Miss Annie M. Alexander, through the Uni-
versity of California Museum of Vertebrate Zoology, most of March
— March 3 to March 29, 1943 — was spent in the state of Michoacan.
Bernardo Villa R. of the Instituto de Biologia de la Universidad
de Mexico was a member of the party from March 23 to 27. Pre-
viously, March 4 to 22, Roberto Alcantar from the Universidad de
Michoacan, in Morelia, participated in the field work. Mr. J. R.
Alcorn was active in the collecting from the beginning until he
entrained for the United States on March 24. The remainder of
the field party was made up of E. Raymond Hall, his wife Mary F.
Hall, and their three sons, William Joel, Hubert H., and Benjamin
D. Hall.
From March 4 to 15 we collected at, and in the vicinity of, Patz-
cuaro. We were housed in two cottages kindly made available by
Sr. Efrain Buenrostro, in Campo Turista Janitzio, 200 meters north-
west of the railroad station in Colonia Revolucion. The shore of
Lake Patzcuaro, the cultivated fields surrounded by stone fences,
and the oak and pine forests roundabout provided varied habitats.
From March 16 to 23 we collected in the territory 1 to 6 miles
(435)
436 University of Kansas Publs., Mus. Nat. Hist.
south of Tacambaro, making our headquarters in the Europa Hotel,
in the town. The steep main street of Tacambaro with native pines
at the upper end descends to plantings of bananas and sugar cane
at the lower end. Our collecting all was done below (south of) the
town in the semitropical country and none at all was done above
(north of) the town.
From March 24 to 27 (three night's trapping) we collected in the
vicinity of Zamora, making our headquarters in rooms diagonally
across the street intersection from the Hotel Fenix.
The resulting specimens, approximately 650 in number, were de-
posited in the Museum of Vertebrate Zoology at the University of
California at Berkeley.
A noteworthy coincidence is that on the very day, February 26,
on which we crossed the international border into Mexico at Laredo,
the beginning of the new volcano, Paricutin, was announced in the
daily press. Our collecting of mammals in Michoacan was nearly
all done in sight of the towering white plume of this rapidly height-
ening volcanic cone and frequently our traps were thickly dusted
with its wind-borne ash. Our eagerness at that time to have sta-
tions established for observing the effects on vertebrates of the
deposition of ash, was gratified in that Dr. Robert T. Hatt inde-
pendently had the same idea and such observations at appropriate
places and times were begun by him and staff members of the Mu-
seum of Zoology of the University of Michigan. One of us, Villa,
was privileged to share in these observations in the spring of 1947.
This continuing interest in the mammals of Michoacan has made
it seem, to us, the more desirable to place on record our findings
as to kinds and occurrence of species. In doing this we have ex-
amined the collections made previously on Cerro Tancitaro and
vicinity by the field party led by Mr. Harry Hoogstraal from the
University of Illinois and the Chicago Natural History Museum.
The specimens of mammals collected by this field party are in the
Chicago Natural History Museum and we are obliged to Mr. Karl
P. Schmidt, Mr. Colin C. Sanborn and the late Dr. Wilfred H.
Osgood for the privilege of studying this material.
Drs. William H. Burt and Emmet T. Hooper, of the Museum of
Zoology of the University of Michigan, lent to us for examination
five specimens of bats, of as many species, which they had taken
in Michoacan. Drs. Remington Kellogg and Henry W. Setzer have
provided us with data on specimens of deer and peccary from
Michoacan which are in the United States National Museum.
Hall and Villa R.: Mammals of Michoacan 437
Specimens in the Institute of Biology of the University of Mexico
have been used. Financial provision by the Kansas University
Endowment Association has enabled us to obtain specimens needed
for comparison from other parts of Mexico.
In addition to the materials mentioned above we have used pub-
lished references to mammals of Michoacan and have prepared the
following lists of kinds of mammals positively known to us to
occur in the Mexican state of Michoacan. It is noteworthy that
specimens recorded in the literature from Acambaro, Michoacan,
no longer are to be ascribed to Michoacan, since a relocation of the
boundary between the states of Michoacan and Guanajuato, places
Acambaro in the latter state.
Our aims were: (1) To record kinds of mammals positively
known from the state, under the correct scientific name, and vernac-
ular names in English, Spanish, and Tarascan. The first Tarascan
name is given in the spelling used by Tarascans followed by the
phonetic equivalent in English in parentheses. (2) To indicate the
geographic range of each kind in the state, and, (3) To record
miscellaneous information which it is thought probably will be
useful in one way or another to other students whose work certainly
will lengthen the list of kinds of mammals known from Michoacan
and otherwise add to our knowledge of them.
Several kinds of bats, of which we lack records, certainly occur
in Michoacan. Four or five kinds of cats (genus Felis), species of
the genera Potos, Lutra, Tayra, Grison, and several other kinds of
mammals of which we now lack positive record, also probably occur
there; the list of kinds, we expect, will number more than one
hundred species and subspecies when more intensive collecting has
been done in the state. In all, we have positive record of 85 kinds
of native, wild mammals of which specimens have been examined
or recorded from Michoacan. Distances and elevations here are
recorded either in the metric system or in the English system, ac-
cording to the system used on the labels of the specimens concerned.
Unless otherwise indicated, catalogue numbers of more than 100,000
are of specimens in the University of California Museum of Verte-
brate Zoology and numbers of less than 100,000 are of the Chicago
Natural History Museum.
438
University of Kansas Publs., Mus. Nat. Hist.
i
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Fig. 1. Map of the state of Michoacan showing place names mentioned in the
text.
ACCOUNTS OF SPECIES AND SUBSPECIES
Didelphis mesamericana mesamericana Oken
Opossum; Spanish, Tlacuache; Tarascan, Ujkuri (Ukuri)
Didlelphys] . mes-americana Oken, Lehrbuch d. Naturgesch, pt. 3,
2:1152, 1S16, type from northern Mexico.
Didelphis mes-amcricana, Allen, Bull. Amer. Mus. Nat. Hist., 16:256,
August 18, 1902.
Range. — Statewide.
Specimens examined, 7: nos. 100063-100067, 100074, 51396, distributed by lo-
calities as follows: Patzcuaro (3 mi. N, 6,700 ft., 1; 2 mi. W, 6,700 ft, 2; 5 mi.
S, 7,800 ft, 1), 4; Tancitaro, 1; 1% mi. S Tacambaro, 5,700 ft, 1; 1 mi. E and
6 mi. S Tacambaro, 1.
Remarks. — The coarse overhair is white all the way to the base
in three specimens but is black in its distal two thirds (white in basal
third) in four specimens. The overhair, six centimeters anterior to
the base of the tail, is 83(80-85) mm. long in the three gray speci-
mens (those with white overhair) and 68(64-72) mm. long in the
black specimens. The ears and all four feet are black. The tail
is black in its proximal half and white in its distal half except in
Hall and Villa R.: Mammals of Michoacan 439
one specimen in which the distal half is almost as dark as the
proximal half. Of the two largest specimens, one is a female from
1% mi. S Tacambaro and the other, a male is from 6 mi. S of the
same place. Measurements are: Total length, ^ 810, § 786;
length of tail, ^360, $ 348; length of hind foot, — , 58; condylo-
basal length, 110.0; 99.6; zygomatic breadth, 68.5; 59.6; length of
nasals, 59.7, 45.0. The tail amounts to 48, 48 and 47 per cent of the
total length in specimens from Patzcuaro; 50 per cent in one from
Tancitaro; 45 and 44 per cent in two from Tacambaro. The sub-
species mesamericana probably intergrades with Didelphis virginiana
virginiana by way of D. m. texensis and D. v. pigra, as Davis
(1944:375) and other writers suggest, in which case the proper name
of the subspecies mesamericana would be Didelphis virginiana mes-
americana. Until intergradation is actually demonstrated, it seems
best to use the name D. m. mesamericana.
Most of our specimens were caught in steel traps, at meat baits,
set for small carnivores.
Marmosa canescens canescens (Allen)
Murine Opossum; Spanish, Raton Tlacuache
Didelphis (Micoureus) canescens Allen, Bull. Amer. Mus. Nat. Hist.,
5:235, September 22, 1893, type from Santo Domingo de Guzman, Isthmus
of Tehuantepec, Mexico.
Marmosa canescens Allen, Bull. Amer. Mus. Nat. Hist., 9:58, March
15, 1897.
Range. — Below Quercus belt, probably throughout western half of state.
Specimen examined, 1 : no. 100062, 1 mi. E and 6 mi. S Tacambaro, 4,000 ft.
Remarks. — The one unsexed subadult measures 5.5 mm. from Ml
to M3 inclusive, which measurement is near the minimum that
Tate (1933: table 1, Sec. 5) records for this subspecies but larger
than the maximum that he (loc. cit.) records for the subspecies
sinaloae which occurs to the northward of Michoacan. Tate (op.
cit. :141) lists two other specimens from Los Reyes. Our specimen
was caught in a mouse trap set in dry grass between a sugar cane field
and a patch of banana trees.
Sorex saussurei saussurei Merriam
Saussure Shrew; Spanish, Musarana
Sorex saussurei Merriam, Proc. Biol. Soc. Washington, 7:173, Septem-
ber 29, 1892, type from north slope Sierra Nevada de Colima, approxi-
mately 8,000 feet, Jalisco.
Range. — In and above Quercus belt, probably throughout northeastern half
of state.
Specimens examined, 14: nos. 8688, 52131-52141, 100076, 100077, distributed
by localities as follows: Patzcuaro, 1; 4 mi. S Patzcuaro, 7,800 ft., 2; Mount
Tancitaro (7,800 ft., 8; 9,500 ft., 1; 9,600 ft., 1; 10,000 ft., 1), 11.
440 University of Kansas Publs., Mus. Nat. Hist.
Remarks. — Two males from Mount Tancitaro, with much worn
teeth, catalogue nos. 52132 and 52138, measure, respectively, as fol-
lows: Total length, 122, 114; length of tail, 46, 43; length of hind
foot, 15, 14; condylobasal length, 18.4, 18.3; palatal length, 8.0, 7.3;
cranial breadth, 9.4, 9.2; least interorbital breadth, 3.7, 3.8; maxil-
lary breadth, 5.5, 5.5; maxillary tooth-row, 6.8, 6.7. The long palate
in no. 52132 and the broad brain case in both specimens appear to be
only individual variations or possibly variations correlated with the
advanced age of the two animals since in other features they do not
differ from specimens which are smaller in these two dimensions.
Jackson (1928:156) records specimens of this shrew from Mount
Patamban and Nahuatzin.
Cryptotis pergracilis pergracilis (Elliot)
Short-tailed Shrew; Spanish, Musarafia Colicorta
Blarina pergracilis Elliot, Field Columb. Mus., publ. 71, zool. ser.,
3:149, February, 1903, type from Ocotlan, Jalisco, Mexico.
Cryptotis pergracilis pergracilis, Miller, Proc. Biol. Soc. Washington.
24:223, October 31, 1911.
Range. — Probably statewide.
Specimen examined, 1: no. 1721 B. Villa R. from Colonia Ibarra, Patzcuaro.
Remarks. — Our one specimen, originally a mount, proves to have
a crushed brain case. The specimen was saved on March 10, 1944,
by P. Luna, who in March, 1943, told one of us (Hall) that many
of these shrews fell into the cement fish-rearing tanks at the biologi-
cal station situated at Colonia Ibarra, a suburb of Patzcuaro, on the
shore of Lake Patzcuaro. We are indebted to Dr. H. H. T. Jackson
for examining our specimen and assigning a name to it.
Balantiopteryx plicata Peters
Sac-winged Bat; Spanish, Murcielago S'acoptero;
Tarascan word for bat is Huasis (Wasis)
Balantiopteryx plicata Peters, Monatsber. k. preuss. Akad. Wiss.
Berlin, p. 476, 1867, type from Puntaarenas, Costa Rica.
Range. — Statewide.
Specimens examined, 2: nos. 52224, 52225, from Apatzingan, 1,040 ft.
Remarks. — This bat is a cave dweller, not infrequently found
roosting with other species.
Glossophaga soricina leachii (Gray)
Long-tongued Bat; Spanish, Murcielago Siricotero
M onophyllus leachii Gray, Voyage of the Sulphur, Zool., 1:18, 1844,
type from Realejo, Nicaragua.
Glossophaga soricina leachii, Miller, Proc. U. S. Nat. Mus., 46:419,
December 31, 1913.
Hall and Villa R.: Mammals of Michoacan 441
Range. — Statewide.
Specimens examined, 4: nos. 11377, Univ. Kan., and alcoholic specimens
nos. 950-952 B. Villa R. field numbers, I.B. (specimens in Instituto do Biologia,
Univ. de Mexico), distributed by localities as follows: Hacienda El S'abino,
Michoacan, approximately 25 mi. S Uruapan, 1 ; El Guayabo, 34 kms. S Urua-
pan, 3.
Remarks. — Specimens from El Guayabo were taken in a natural
cave which they shared with Desmodus rotundus murinus and Arti-
beus planirostris planirostris.
The length of the thumb averages 7.4 mm. (7.0 to 7.7). As com-
pared with G. s. alticola from northeast Tlaxcala according to the
description given by Davis (1944:377), our specimens agree with
alticola in length of thumb. In all other characters they correspond
to leachii.
Choeronycteris mexicana Tschudi
Long-tongued Bat; Spanish, Murcielago Carilargo
Choeronycteris mexicana Tschudi, Fauna Peruana, p. 72, 1844, type
from Mexico.
Range . — Statewide .
Specimens examined, 4: nos. 100078-100081, from 2 mi. W Patzcuaro, 7,700 ft.
Remarks. — A colony of 20 or more bats of this species was found
in a natural cave. Four were caught by hand as they flew about
after we disturbed them. We returned on the following day, but
found that all the bats had left.
Leptonycteris nivalis nivalis (Saussure)
Leaf-nosed Bat; Spanish, Murcielago Lengiiilarga
M[ = Ischnoglossa]. nivalis Saussure, Revue et Magasin de Zoologie,
12(ser. 2) :492, November, 1860, type from near snow line on Mount Ori-
zaba.
Leptonycteris nivalis, Miller, Proc. Biol. Soc. Washington, 13:126, April
6, 1900.
Range. — Probably middle and higher altitudes through state.
Specimen examined, 1: no. 91911, Univ. Michigan Mus. Zool., from 1050 m.,
12 miles (on Huetamo Road) south of Tzitzio.
Remarks. — The subspecific name L. n. nivalis is tentatively ap-
plied to this specimen in the absence of an opportunity to compare
it directly with the holotype or topotypes of Leptonycteris nivalis
yerbabuenae Martinez and Villa (1940:291). Unfortunately, the
materials on which this name, L. n. yerbabuenae, was based all
were destroyed in 1945 or 1946 while Villa was absent from the
Institute of Biology of the University of Mexico.
442 University of Kansas Publs., Mus. Nat. Hist.
Artibeus planirostris planirostris (Spix)
Big Leaf-nosed Bat; Spanish, Murcielago Zapotero
Phyllostoma planirostre Spix, Simiarum et vespertilionum Brasilien-
sium, p. 66, 1823, type from suburbs of Bahia, Brazil.
Artibeus planirostris, Dobson, Catal. Chiroptera, British Mus., p. 515
(part), 1878.
Range. — Probably southwestern part of state.
Specimen examined, 1 : no. 945b B. Villa R., field no., I. B., El Guayabo,
34 kms. S Uruapan.
Measurements. — Head and body, 89.0 mm.; total length of skull to front of
upper canines, 28.0; mastoid breadth, 15.5; zygomatic breadth, 17.8; maxillary
width across first upper molars, 13.2; breadth across cingula of upper canines, 7.9;
greatest length of one ramus of lower jaw including anteriormost incisor tooth,
19.4; length of upper tooth-row, anterior border of canine to posterior border
of M2, 10.4; length of lancet (nose-leaf), 9.0; width of lancet, 6.5; width of
horseshoe, 9.0; forearm, 57.3; 3rd metacarpal, 52.6; 1st (basal) phalanx, 16.0;
2nd (middle) phalanx, 26.7; 3rd (distal) phalanx, 18.5; 4th metacarpal, 50.7;
1st (basal) phalanx, 14; 2nd phalanx, 18.8; 5th metacarpal, 54; 1st (basal)
phalanx, 11.2; 2nd phalanx, 13.2; lower leg, 22.9; foot with claws, 15.2; calcar,
6.5.
Remarks. — Our single specimen, a female, was caught on July
28, 1945, by my (B. Villa's) father, Andres Villa, in a natural cave,
roosting with the individuals of Glossophaga s. leachii. The north-
ernmost locality in Mexico from which A. p. planirostris previously
has been recorded is El Papayo, in the state of Guerrero (Andersen,
1908:238), approximately 225 kilometers to the southward. A. p.
planirostris and Artibeus jamaicensis closely resemble each other
but A. planirostris may be recognized by the presence of a minute
M3 which is absent in A. jamaicensis. Specimen no. 945b has M3
present on both sides of the upper jaw. From Artibeus hirsutus,
known from La Salada, Michoacan, approximately 40 miles north
and slightly to the east of El Guayabo, our specimen differs in the
apparently hairless tibia and interfemoral membrane. The measure-
ments, of no. 945b, recorded above, if compared with those given
by Andersen (1908:246) are seen mostly to fall within the range
recorded for A. hirsutus. Where measurements are outside this
range, they fall within the range of those of the larger A. p. plani-
rostris. We recognize that the Mexican species of Artibeus are not
well understood, at least by us.
Artibeus hirsutus K. Andersen
Leaf-nosed Bat; Spanish, Murcielago Zapotero
Artibeus hirsutus K. Andersen, Ann. and Mag. Nat. Hist., 18(ser. 7) :
420, December, 1906, type from La Salada, Michoacan.
Range. — Known from western part of state.
Hall and Villa R.: Mammals of Michoacan 443
Remarks. — From Michoacan, Andersen (1908:247) examined
three specimens, all from the type locality.
Desmodus rotundus murinus Wagner
Vampire Bat; Spanish, Vampiro
D[esmodus] . murinus Wagner, Schreber's Saugthiere, Suppl., 1 :377, 1840,
type from Mexico.
Desmodus rotundus murinus, Osgood, Field Mus. Nat. Hist., publ. 155,
zobl. ser., 10:63, January 10, 1912.
Range. — Statewide, except rare or absent at higher altitudes.
Specimens examined, 6: nos. 944-949 B. Villa R. field numbers, I.B. El
Guayabo, 34 kms. S Uruapan.
Remarks. — This species is colonial; usually it is found in large
numbers in favorite roosting sites, mainly in natural caves. Four
of our specimens, caught in July, are females and two are young
males. One, female, no. 944, has one embryo of 40 mm. in length.
Myotis yumanensis lutosus Miller and Allen
Yuma Myotis; Spanish, Murcielago de Yuma
Myotis yumanensis lutosus Miller and Allen, U. S. Nat. Mus., Bull.,
144:72, May 25, 1928, type from Patzcuaro, Michoacan.
Myotis yumanensis, Miller, N. Amer. Fauna, 13:67, October 16, 1897.
Range. — Known only from Patzcuaro and El Molino.
Remarks. — Originally recorded by Miller (1897:67) from Patz-
cuaro, the animals from central Mexico were named as a new sub-
species by Miller and Allen (1928:72) who record one specimen
from El Molino.
Myotis velifer velifer (Allen)
Cave Bat; Spanish, Murcielago Vespertino
Vespertilio velifer J. A. Allen, Bull. Amer. Mus. Nat. Hist., 3:177,
December 10, 1890, type from Santa Cruz del Valle, Guadalajara, Jalisco.
Myotis velifer velifer, Allen and Miller, U. S. Nat. Mus. Bull., 144:89,
May 25, 1928.
Range. — Statewide.
Specimens examined, 17: nos. 100083-100099, from 3 mi. NW Patzcuaro,
6,700 ft.
Remarks. — Our specimens were taken on March 12, 1943, from
a crevice in the wall of an abandoned chapel where 35 or more
individuals of both sexes were living. Miller (1897:59) records
the species from Patzcuaro and Miller and Allen (1928:91) record
it from there and also from Lake Chapala, La Palma, Acambaro
(now in Guanajuato) and Negrete.
444 University of Kansas Publs., Mus. Nat. Hist.
Myotis thysanodes thysanodes Miller
Fringed-tailed Myotis; Spanish, Murcielago Colirugosa
Myotis thysanodes Miller, N. Amer. Fauna, 13:80, October 16, 1897,
type from Old Fort Tejon, Kem County, California.
Range. — Known only from Patzcuaro.
Remarks. — In the original description five specimens are recorded
from Patzcuaro and Miller and Allen (1928:127) mention the same
locality of occurence.
Myotis californicus mexicanus (Saussure)
California Myotis; Spanish, Murcielago de California
Vlespertilio]. mexicanus Saussure, Rev. et Mag. de Zool., 12 (ser. 2):
282, 1860, type from somewhere in the warmer part of the state of Mexico.
Myotis calij ornicus mexicanus, Miller, N. Amer. Fanua, 13:73, October
16, 1897.
Range. — Known in Michoacan only from Patzcuaro.
Remarks. — Specimens from Patzcuaro are recorded by Miller
and Allen (1928:160).
Eptesicus fuscus miradorensis (H. Allen)
Big Brown Bat; Spanish, Murcielago Fusco
Slcotophilus]. miradorensis H. Allen, Proc. Acad. Nat. Sci. Philadelphia,
p. 287, 1866, type from Mirador, "Veracruz.
Eptesicus fuscus miradorensis, Miller, N. Amer. Land Mamm., 1911,
p. 62, December 31, 1912.
Range. — Probably statewide.
Specimen examined, 1: no. 91909, Univ. Michigan, Mus. Zool., from Rancho
Escondido, one mile north of Apo, 6,000 feet elevation, June 29, 1947, female
adult, taken by W. H. Burt.
Lasiurus borealis mexicanus (Saussure)
Red Bat; Spanish, Murcielago Rojizo
A[talapha~\. mexicana Saussure, Rev. et Mag. de Zool., 13(2) :97,
March, 1861, type probably from Veracruz, Puebla or Oaxaca.
Lasiurus borealis mexicanus, Miller, N. Amer. Fauna, 13:111, October
16, 1897.
Range. — Probably larger part of state.
Specimen examined, 1 : no. 89446, Univ. Michigan, Mus. Zool., from Nuevo
San Juan (Los Conejos), 5 mi. SW Uruapan, May 23, 1945, by W. H. Burt.
Remarks. — This specimen, a male with much worn teeth, answers
well to the description of L. b. mexicanus except that the minute
premolar between the canine and fourth premolar is missing on
each side of the upper jaw. This, however, seems the less remark-
able after examination of 18 skulls of L. b. borealis from the United
States in two of which these minute premolars are likewise absent;
one of the two specimens from the United States has unworn teeth
and the other much worn teeth.
Hall and Villa R.: Mammals of Michoacan 445
Lasiurus cinereus cinereus (Beauvois)
Hoary Bat; Spanish, Murcielago Pardo
Vespertilio cinereus (misspelled linereus) Beauvois, Catal. Raisonne
Mus. Peale, Philadelphia, p. 18, 1796, type locality, Philadelphia, Pennsyl-
vania.
Lasiurus cinereus, H. Allen, Monogr. N. Amer. Bats, Smithsonian
Misc. Colls., 7(1) :12, June, 1864.
Range. — Higher elevations throughout state.
Specimen examined, 1: no. 89456, Univ. Michigan Mus. Zool., from Barranca
Seca, May 6, 1945, adult male with much worn teeth, obtained by W. H. Burt.
Corynorhinus rafinesquii mexicanus G. M. Allen
Long-eared Bat; Spanish, Murcielago Narigudo
Corynorhinus megalotis mexicanus Allen, G. M., Bull. Mus. Comp.
Zool., 60:347, April, 1916, type from "near Pacheco," Chihuahua.
Corynorhinus rafinesquii mexicanus, Miller, U. S. Nat. Mus., Bull.,
128:83, April 29, 1924.
Corynorhinus macrotis pallescens, Miller, N. Amer. Fauna, 13:52,
October 16, 1897.
Range. — Known from only Patzcuaro.
Remarks. — Miller (1897:53) recorded one specimen from Patz-
cuaro and Allen (1916:349) merely alludes to Miller's record.
Tadarida mexicana (Saussure)
Mexican Free-tailed Bat; Spanish, Murcielago Coludo
Molossus mexicanus Saussure, Rev. et Mag. de Zool., 12:283, July,
1860, type from Cofre de Perote, 13,000 feet, Veracruz.
Tadarida mexicana, Miller, Bull. U. S. Nat. Mus., 128:86, April 29, 1924.
Range. — Statewide.
Specimens examined, 12: nos. 100100-100111, distributed by localities as fol-
lows: 1 mi. N Zamora, 5,450 ft., 1; 3 mi. N Patzcuaro, 6.800 ft,, 3; 3 mi. NW
Patzcuaro, 6,700 ft., 3; Isla Janitzio, Lago de Patzcuaro, 6,600 ft., 5.
Remarks. — This species is widespread in Mexico, ranging from
sea level to high elevations as at the type locality. In Michoacan
most of our specimens were shot as they flew about at early dusk.
The five from Isla Janitzio were shot as they clung to the roof of a
cave along with scores of other individuals of the same species.
Eumops underwoodi underwoodi Goodwin
Mastiff Bat; Spanish, Murcielago Mastin
Eumops underwoodi Goodwin, Amer. Mus. Novitates, 1075:2, June
27, 1940, type from El Pedrero, 6 km. N Chinaela, approximately 3,000
ft. elevation, Dept. La Paz, Honduras.
Range. — Known only from Tancitaro Mtn.
Specimen examined, 1: no. 89461, Univ. Michigan Mus. Zool., from Rancho
Escondido, 2 mi. N Apo, Tancitaro Mtn., $ ad. with much worn teeth, taken
June 11, 1945, by W. H. Burt.
Remarks. — Selected measurements of this specimen are: Total
length, 158; ear from notch, 32; mastoid breadth, 16.1; width across
446 University of Kansas Publs., Mus. Nat. Hist.
crowns of M3, 12.6; maxillary tooth-row (from anterior face of
canine above cingulum to posterior face of M3), 11.8. The total
length is less than in E. underwoodi or than in Eumops sonoriensis
Benson (1947:133); the other measurements given above exceed
those of E. sonoriensis and equal or approach those of E. under-
woodi. The ears seem not to be connected across the forehead ; the
color is near (I) Bister above and slightly lighter on the underparts.
The specimen is clearly intermediate in size, as it also is geo-
graphically, between Eumops underwoodi underwoodi Goodwin and
Eumops underwoodi sonoriensis Benson and gives basis for arrang-
ing these two named kinds as subspecies of a single species as Ben-
son (1947:134) suggested might prove to be necessary. We are not
certain whether this specimen should be referred to the subspecies
underwoodi or sonoriensis and probably this uncertainty will re-
main until the range of individual variation in underwoodi is known.
Procyon lotor hernandezii Wagler
Raccoon; Spanish, Mapache; Tarascan, Apatze (Apatz)
Pr[ocyori] hernandezii Wagler, Isis, 24:514, 1831, type from Valley of
Mexico, according to Nelson and Goldman (Proc. Biol. Soc. Washington,
44:17, February 21, 1931).
Procyon lotor hernandezi, Allen, Bull. Amer. Mus. Nat. Hist., 3:176,
December 10, 1890.
Range. — Statewide.
Specimens examined, 2: no. 100113 from 10 mi. ESE Zamora, 5,500 ft., 1;
no. 52220 from 15 kms. W Apatzingan, 1,040 ft., 1.
Remarks. — In allusion to its habit of washing its food, in cap-
tivity at least, before eating it, the Spanish speaking people often
refer to this species as ositos labadores. The specimen from 10 mi.
ESE Zamora is a skull without lower jaws or indication of sex.
Because the racoons damage corn in the roasting ear stage the
animals are disliked by the farmers, a score of whom sometimes
band together in an organized hunt to kill the animals. Dogs are
especially trained to hunt them. In Michoacan no use is made of
the pelts.
Nasua narica molaris Merriam
Coati; Spanish, Pizote; Tarascan, Amatze (Amatz)
Nasua narica molaris Merriam, Proc. Biol. Soc. Washington, 15:68,
March 22, 1902, type from Manzanillo, Colima; Goldman, Proc. Biol. Soc.
Washington, 55:79, June 25, 1942.
Range. — Probably all but higher parts of state.
Remarks. — We have no positive record of this animal which
Goldman (1942:79) writes "is widely distributed from Jalisco south
through Colima, Michoacan, ... to southwestern Oaxaca."
Hall and Villa R.: Mammals of Michoacan 447
In the parts of Michoacan visited by us the Spanish name tejon
instead of pizote was used for this animal although in parts of
Mexico where the badger (Taxidea) occurs, tejon is, we understand,
the name used for the badger.
Bassariscus astutus consitus Nelson and Goldman
Ring-tailed Cat; Spanish, Cacomixtle
Bassariscus astutus consitus Nelson and Goldman, Jour. Washington
Acad. Sci., 22:487, October 19, 1932, type from La Salada, 40 mi. S Urua-
pan, Michoacan.
Range. — Probably greater part, or all, of state.
Specimen examined, 1: no. 100112 from 3 mi. NW Patzcuaro, 6,700 ft.
Remarks. — La Salada and three miles northwest of Patzcuaro
are the two localities represented by actual specimens. A live ani-
mal, at night, was seen one mile east and four miles south of
Tacambaro at 4,500 feet elevation. The young female from three
miles northwest of Patzcuaro was trapped at a break in a stone
fence.
Ring-tailed cats live in the stone walls, crevices and rocky ledges,
around corn fields and pasture lands.
Mustela frenata leucoparia (Merriam)
Weasel; Spanish, Comadreja; Tarascan, Apasr or Apatzee (Apatz)
Putorius jrenatus leucoparia Merriam, N. Amer. Fauna, 11:29, June
30, 1896, type from Patzcuaro, Michoacan.
Mustela frenata leucoparia, Miller, Bull. U. S. Nat. Mus., 79:100, De-
cember 31, 1912.
Range. — Probably statewide.
Specimens examined, 6: in Biological Surveys Collection of U. S. Nat. Mus.,
nos. 120304, 125972, 34914/47179, 36855/49239, and 34915/47180; 2014 B. Villa R.,
I.B., distributed as follows: Zamora, 1; Los Reyes, 1; Patzcuaro, 4.
Remarks. — This subspecies of weasel is notable for having, among
American weasels of any kind, the maximum amount of white on
the head. When collecting at Patzcuaro we saw no live specimens
but were shown several from there that had been recently mounted
by P. Luna. He regarded the animal as not especially rare.
Spilogale angustifrons angustifrons Howell
Spotted Skunk; Spanish, Zorrillo Manchado
Spilogale angustifrons Howell, Proc. Biol. Soc. Washington, 15:242,
December 16, 1902, type from Tlalpam, D. F.
Range. — Probably all of state except low costal area.
Specimen examined, 1 : no. 100126, 3 mi. NW Patzcuaro, 6,700 ft.
Remarks. — The short tail of our specimen, an adult male, is note-
worthy as perhaps also is the breadth between the orbits. External
448 University of Kansas Publs., Mus. Nat. Hist.
measurements are 338, 101, 39. It weighed 308 grams. Selected
cranial measurements are: Basilar length, 44.1; zygomatic breadth,
32.4; postpalatal length, 26.6; least interorbital breadth, 13.8; height
of cranium, 16.0. The specimen was trapped in a hole in a stone
fence. Howell (1906:23), under the name Spilogale gracilis, re-
corded another male from Patzcuaro.
Mephitis macroura macroura Liechtenstein
Hooded Skunk; Spanish, Zorrillo or Mofeta Rayada; Tarascan,
Cuitziqui (Kweetzeke)
Mephitis macroura Lichtenstein, Darstellung Neuer oder wenig be-
kannter Saugethiere pi. 46, 1832, type from mountains northwest of
Mexico City.
Range. — Statewide.
Specimens examined, 12: nos. 100114-100125, distributed with reference to
Patzcuaro, as follows: 3 mi. NW, 6,700 ft., 1; 2 mi. W, 7,600 and 7,000 ft., 2;
3% mi. S, 7,900 ft, 1; 4 mi. S, 7,800 ft., 2; 5 mi. S, 7,800 ft, 5; 9 mi. SE, 8,000
ft, 1.
Remarks. — Skunk tracks were abundant in all localities around
Patzcuaro. Most of our specimens were caught in steel traps, some
along the edges of cornfields, others along the highway and along
the pole fences. Tarascan friends at Colonia Revolucion were eager
to have the bodies of the skunks which we caught. They regarded
the skunks as a delicacy and told us that this food was reputed to
be good for a person's blood and complexion.
Conepatus mesoleucus nelsoni Goldman
Hog-nosed Skunk; Spanish, Zorillo Real
Conepatus mesoleucus nelsoni Goldman, Jour. Mamm, 3:41, February
8, 1922, type from Armeria (near Manzanillo), Colima, 200 ft. altitude.
Range. — Probably greater part, or all, of state.
Specimen examined, 1 : no. 52217, Tancitaro, 6,000 ft.
Remarks. — The name C. m. nelsoni is tentatively used for the
single skin, which is without skull or indication of sex.
Urocyon cinereoargenteus colimensis Goldman
Gray Fox; Spanish, Zorra Gris; Tarascan, Cumihuatz (Cumewatz)
Urocyon cinereoargenteus colimensis Goldman, Jour. Washington Acad.
Sci, 28:495, November 15, 1938, type from 3 mi. W city of Colima, 1,700
ft. elevation.
Range. — Statewide.
Specimens examined, 2: no. 100127, from 1 mi. E and 6 mi. S Tacambaro,
4,000 ft, and no. 51393 from Apatzingan.
Remarks. — The female from southeast of Tacambaro, caught on
March 20, 1943, had two embryos, 28 mm. in length. This female
PLATE 4
Fig. 1. Panoramic view of Lake Patzcuaro
Fig. 2. Stuffed skins, in dorsal view, of 6 males of Mephitis macroura
macroura, all trapped within a radius of 5 miles of Patzcuaro, to show the
amount of individual variation in color-pattern. X Mo- Photo by W C
Matthews.
PLATE 5
•MHRm^Jp?
Fk;. 1. Ungrazed pasture with oaks on slope of El Estribo, 7,700 feet eleva-
tion, two miles west of the town of Patzcuaro, Michoacan. Several species of
rodents. Liomys, Sigrnodon, and Peromyscus were taken abundantly in the
grass in the immediate foreground. Photo March 16. 1943, by Mary F. Hall.
Fig. 2. Xerophitic vegetation, eleven miles west of Zamora, Michoacan.
5,750 feet elevation, where rodents were trapped. Photo March 26, 1943, bv
Mary F. Hall.
Hall and Villa R.: Mammals of Michoacan 449
was trapped near a small stream. Goldman (1938:497) reported
7 specimens of U. c. colimensis from the following localities in the
state: La Huacana, 1; La Salada, 2; Los Reyes, 1; Mount Tan-
citaro, 1; Patzcuaro, 2.
Canis Iatrans cagottis (Hamilton Smith)
Coyote; Spanish, Coyote; Tarascan, Jihuatz (Hewatz)
Lyciscus cagottis Hamilton Smith, Jardine's Naturalist's Library,
Maram., 9:164, 1839, type from Rio Frio between city of Mexico and
Puebla.
Canis Iatrans cagottis, Nelson, Proc. Biol. Soc. Washington, 45:224,
November 26, 1932.
Range. — Probably most of state.
Remarks. — On March 27 or 28, 1943, in Morelia, at a gasoline
filling station, one of us (Hall) saw a freshly killed coyote tied on
the bumper of the automobile of a Medical Doctor. In response to
inquiry about the animal the Doctor said that he killed it some 15
miles northeast of town.
Lynx rufus escuinapae Allen
Bobcat; Spanish, Gato del Monte; Tarascan, Misicpapu (misicpapu)
Lynx ruff us escuinapae Allen, Bull. Amer. Mus. Nat. Hist., 19:614, No-
vember 14, 1903, type from Escuinapa, Sinaloa.
Range. — Probably all of state above the Tropical Life-zone.
Specimen examined, 1: no. 47818 (U. S. Nat. Mus., Biol. Surv. Coll.), Patz-
cuaro.
Remarks. — The skull of the male from Patzcuaro agrees well with
those of topotypes of L. r. escuinapae even to the elongate tympanic
bullae. Because of their elongation the bullae resemble, in some
degree, those of the ocelot. Natives told us that the gato del monte
was resident around Patzcuaro. Four miles south of Patzcuaro in
a cornfield at the edge of an area grown up to oak trees and brush,
tracks were seen that our Indian companion identified as those of
the gato del monte.
Citellus variegatus variegatus (Erxleben)
Rock Squirrel; Spanish, Ardilla de Pedregal; Tarascan,
Kuaraki (Kuaraki)
[Sciurus] variegatus Erxleben, S'yst. Regni, Anim., 1:421, 1777; type
locality fixed as Vallev of Mexico near the city of Mexico, by Nelson,
Science, N. S., 8:898, December 23, 1898.
[Citellus] variegatus, Elliot, Field Columb. Mus. Pub., zool. ser. 4:148,
1904.
Range. — Probably in all semi-arid, rocky habitats of the state.
Specimens examined, 11: nos. 100128-100135; 51385-51387, distributed by lo-
450 University of Kansas Publs., Mus. Nat. Hist.
calities as follows: 1 mi. N Zamora, 5,450 ft., 1; 3 mi. NW Patzcuaro, 6,700
ft., 5; 4 mi. S Patzcuaro, 7,800 ft., 1; IV2 mi. S Tacambaro, 5,700 ft., 1; Mount
Tancitaro, 1; Pedregal, Tancitaro, 1; Tancitaro, 1.
Remarks. — Rock squirrels were seen along rock fences, around
Patzcuaro, where they are fairly common. On July 17 and 18,
1947, at San Juan Parangaricutiro, one of us (Villa) saw these
squirrels running over the newly formed lava bed which was still
emitting vapors and which in places (between boulders) was
emitting heat detectable by the collectors. This is only additional
evidence of the animal's strong predilection for rocks, boulders and
cliffs, which has earned for it, in parts of the western United States,
the vernacular name "rock squirrel" and in Mexico "Ardilla de
Pedregal."
Howell (1938:138) reported specimens from the following lo-
calities: Acambaro, 1; Los Reyes, 1; Mount Tancitaro, 2; Patz-
cuaro, 12; Querendaro, 1; Zamora, 2.
Citellus adocetus adocetus Merriam
Lesser Tropical Ground Squirrel ; Spanish, Cuiniqui ; Tarascan,
Kuaraki (Kuaraki)
Citellus adocetus Merriam, Proc. Biol. Soc. Washington, 16:79, May
29, 1903, type from La S'alada, 40 miles south of Uruapan, Michoacan.
Range. — Southern part of state in arid tropical land.
Specimens examined, 4: nos. 52000, 52001, 51388, 51389, distributed by lo-
calities as follows: "Near Tancitaro," 2; Acahuato, 1; Apatzingan, 1,040 ft., 1.
Remarks. — Ground squirrels of this species are fairly abundant
in the arid tropical parts of the state. Their burrows are usually
found on stony areas along small ravines or under mesquite (Pro-
sopis juliflora) thickets. The name cuiniqui in use by the Spanish
speaking population is merely a corruption of the Tarascan name.
Cuiniqui, therefore, is a particular kind of ardilla terricola (ground
squirrel).
Sciurus poliopus nemoralis Nelson
Michoacan Squirrel; Spanish, Ardilla arboricola; Tarascan,
Uakui (Wakqe)
Sciurus albipes nemoralis Nelson, Proc. Biol. Soc, Washington, 12:151,
June 3, 1898, type from Patzcuaro, Michoacan.
Sciurus poliopus nemoralis Nelson, Proc. Washington Acad. Sci., 1:50,
May 9, 1899.
Range. — Pine and oak forests of most of state.
Specimens examined, 3: nos. 2102 and 2103 Louisiana State University from
20 mi. E Morelia, 7,300 ft., and no. 1369 B. Villa R. from V/2 km. N San Juan
2,250 meters.
Remarks. — Tree squirrels of this kind have been reported by
Nelson (1899:51) from Patzcuaro and Nahuatzin.
Hall and Villa R.: Mammals of Michoacan 451
The young specimen, no. 2102, J1 , has the tail slender, resembling
somewhat that of the S. p. senex from the southward. The under-
pays of the female are Warm Buff, more clearly so on the underside
of the legs.
Sciurus poliopus senex Nelson
Michoacan Squirrel; Spanish, Ardilla Arboricola; Tarascan,
Uakui (Wakqe)
Sciurus poliopus senex Nelson, Proc. Biol. Soc. Washington, 17:148,
October 6, 1904, type from La Salada, 40 mi. S Uruapan.
Range. — Lowlands in southern part of state.
Specimens examined, 3: nos. 52004, 52014, 52015, distributed by localities as
follows: Tancitaro, 6,000 ft., 1; Apatzingan, 1,040 ft., 2.
Remarks. — Specimen number 52014, £ , represents the melanistic
phase of this subspecies.
The upper parts of the hind legs in this specimen are slightly
grizzled. The upper side of the tail is vermiculated with whitish
and the underside of the tail is black.
Thomomys umbrinus pullus Hall and Villa
Southern Pocket Gopher; Spanish, Tuza Serrana; Tarascan,
Cunu (Como)
Thomomys umbrinus pullus Hall and Villa, Univ. Kansas Publ., Mus.
Nat. Hist., 1:251, July 26, 1948, type from 5 miles south Patzcuaro, 7,800
ft., Michoacan.
Range. — Known only from pine-covered rolling land three to five miles south
of Patzcuaro.
Specimens examined, 17: nos. 100136-100152, distributed by localities as fol-
lows: 3 mi. S Patzcuaro, 7,800 ft., 1; 4 mi. S Patzcuaro, 7,800 ft., 10; 5 mi. S
Patzcuaro, 7,800 ft., 6.
Remarks. — Most of these pocket gophers were caught in areas
supporting a good growth of pine trees in the same places where
the much larger pocket gopher, Cratogeomys gymnurus, lived. Con-
cerning the individual designated as the type specimen, H. H. Hall
(field notes) writes that when he was making an excavation to re-
veal the gopher's burrow (5 inches below the surface), he dug deeper
than was necessary and broke into the burrow of a Cratogeomys
directly below. Another of us (E. R. Hall) had the same experience
where the burrow of a Thomomys was approximately six inches
below ground and that of a Cratogeomys approximately 16 inches
below the surface of the ground. At the time this arrangement
led us to wonder if Thomomys was in some sense a "parasite" on
the larger Cratogeomys by levying on food stores, if Cratogeomys
has any, but we found no evidence that such was the case and from
452 University of Kansas Publs., Mus. Nat. Hist.
our subsequent trapping concluded that the two-story arrangement
was accidental and not the rule. The habit of burrowing at different
levels probably was one factor which permitted the two kinds of
pocket gophers to live in the same area. The average weight of
these gophers was 86 grams in males and 74 grams in females, or
only an eighth as much as in Cratogeomys.
Cratogeomys gymnurus imparilis (Goldman)
Plains Pocket Gopher; Spanish, Tuza Llanera; Tarascan,
Cumu (Como)
Platygeomys gymnurus imparilis, Goldman, Jour. Mamm., 20:89, Feb-
ruary 14, 1939, type from Patzcuaro, 7,000 ft., Michoaean.
Platygeomys lylorhinus Merriam, N. Amer. Fauna, 8:167, pi. 13, fig.
1, January 31, 1895.
Range. — Patzcuaro and Tacambaro, as now known.
Specimens examined, 14: nos. 100153-100166, distributed by localities as fol-
lows: 2 mi. W Patzcuaro. 7.700 ft., 2; 3 mi. S Patzcuaro, 7,800 ft., 1; 4 mi. S
Patzcuaro, 7,800 ft, 1; 5 mi. S. Patzcuaro, 7.800 ft, 6; 9 mi. SE Patzcuaro, 8,000
ft, 1: 1% mi. S Tacambaro, 5,700 ft, 1; 1M> mi. S Tacambaro, 5,700 ft, 2.
Remarks. — Burrows were common in cultivated fields and along
the roads and trails on the southern and southeastern side of Lake
Patzcuaro. In the vicinity of Tacambaro we noted burrows only
in the area between one and a half and two miles south of town
where two specimens were taken. As mentioned in the immediately
preceding account, the small Thomomys umbrinus pullus and the
large Cratogeomys were found in the same area. The color of our
specimens varies from Cinnamon-Brown through Prouts Brown
and in some specimens is Fuscous Black.
Hooper (1946:397) has shown that the genus Platygeomys is not
generically distinct from the earlier named Cratogeomys. From
independent study of specimens not examined by Hooper we have
satisfied ourselves that he is correct in synonymizing Platygeomys
under Cratogeomys. Average and extreme weights of 4 of each sex
from 2 to 5 miles south of Patzcuaro are: $ , 683 (562-819) ; $ , 558
(438-707) grams.
Cratogeomys angustirostris (Merriam)
Plains Pocket Gopher; Spanish, Tuza Llanera; Tarascan,
Ciimu (Como)
Platygeomys tylorhinus angustirostris Merriam, Proc. Biol. Soc. Wash-
ington, 16:81, May 29, 1903, type from Patamban, 10,000 ft, Michoaean.
Platygeomys angustirostris, Goldman, Jour. Mamm, 20:90, February
14, 1939.
Range. — Known only from the type locality.
Hall and Villa R.: Mammals of Michoacan 453
Cratogeomys varius (Goldman)
Plains Pocket Gopher; Spanish, Tuza Llanera; Tarascan,
Cumu (Como)
Platygeomys varius Goldman, Jour. Mamm., 20:90, February 14, 1939,
type from Uruapan, about 6,000 ft., Michoacan.
Range. — Known only from the type locality.
Remarks. — Specimens from localities intermediate between Urua-
pan, the type locality of C. varius, and the known localities of oc-
curence of Cratogeomys gynmurus imparilis are much needed to
ascertain if C. varius is specifically different from C. g. imparilis,
or merely subspecifically different.
Zygogeomys trichopus tarascensis Goldman
Michoacan Pocket Gopher; Spanish, Tuza de Michoacan; Tarascan,
Cumu (Como)
Zygogeomys trichopus tarascensis Goldman, Proc. Biol. Soc. Washing-
ton. 51 :211, December 23, 1938, type from six miles southeast of Patzcuaro,
8,000 ft., Michoacan.
Zygogeomys trichopus Merriam, N. Amer. Fauna, 8:196, January 31,
1895.
Range. — "Known only from the upper slopes of the mountains in the vi-
cinity of the type locality" (Goldman, 1938:211).
Remarks. — As we drove an automobile from Patzcuaro to Tacam-
baro we noted mounds made by pocket gophers along the road in the
highest part of the pass and supposed that these mounds were made
by Zygogeomys although we took no specimens of any kind of
pocket gopher in the pass.
Zygogeomys trichopus trichopus Merriam
Michoacan Pocket Gopher; Spanish, Tuza de Michoacan; Tarascan,
Cumu (Como)
Zygogeomys trichopus Merriam, N. Amer. Fauna, 8:196, pi. 6, 14-18,
January 31, 1895, type from Nahuatzin, Michoacan.
Range. — Altitudinally from 6,000 feet to 11,800 feet on Mountains Tancitaro,
Patamban, and at Nahuatzin.
Specimens examined, 9: nos. 51970-51978, all from Mount Tancitaro, dis-
tributed by altitude as follows: 6,000 ft., 5; 7,800 ft., 3; 10,500 ft., 1.
Remarks. — The upper parts of specimens available to us are rich
Seal-Brown and glossy. The chin, and in most specimens, the upper
side of the hind feet are white; the irregular white patch of the
throat is present only in two young females, numbers 51974 and
51978.
454 University of Kansas Publs., Mus. Nat. Hist.
Liomys pictus plantinarensis Merriam
Western Spiny Pocket Mouse; Spanish, Raton Espinoso Occidental;
Tarascan word for mouse is Jeyaqui (Hayake)
Liomys -plantinarensis Merriam, Proc. Biol. Soc. Washington, 15:46,
March 5, 1902, type from Plantinar, Jalisco.
Liomys pictus plantinarensis, Goldman, N. Amer. Fauna, 34:37, Sep-
tember 7, 1911.
Range. — Northwestern part of state in semitropical areas.
Remarks. — Goldman (1911:38) records specimens from Los
Reyes, noting that in some cranial features they suggest intergrada-
tion between L. p. plantinarensis and L. p. parviceps.
Liomys pictus parviceps Goldman
Western Spiny Pocket Mouse; Spanish, Raton Espinoso Occidental
Liomys parviceps Goldman, Proc. Biol. Soc. Washington, 17:82, March
21, 1904, type from La S'alada, "40 miles south of Uruapan, Michoacan."
Liomys pictus parviceps Goldman, N. Amer. Fauna, 34:38-39, Sep-
tember 7, 1911.
Range. — Southern part of state in semitropical and tropical areas.
Specimens examined, 28: nos. 100185-100199, 52072-52084, distributed by lo-
calities as follows: Apatzingan, 1,040 ft., 13; 1 mi. E and 2% mi. S Tacambaro,
4,700 ft., 4; 4 mi. S and 1 mi. E Tacambaro, 4,500 ft., 5; 1 mi. E and 6 mi. S
Tacambaro, 4,000 ft., 6.
Remarks. — Most measurements show a sexual dimorphism in
this subspecies. Adult males are 15 per cent larger in external
measurements except that the foot is approximately the same.
Cranial measurements average approximately 5 per cent larger in
males except that the breadth of the rostrum and length of the
maxillary tooth-row are slightly less. South and east of Tacambaro
our specimens all were taken in dry semitropical country, where
bananas and sugar cane were the principal crops grown. This sub-
species has been recorded also from La Huacana, Michoacan, as
well as from La Salada, the type locality, by Goldman (1911:39).
Liomys irroratus jaliscensis (Allen)
Northern Spiny Pocket Mouse; Spanish, Raton Norteno
Heteromys jaliscensis Allen, Bull. Amer. Mus. Nat. Hist., 22:251, July
25, 1906, type from Las Canoas, approximately 20 mi. W Zapotlan, 7,000
ft., Jalisco.
Liomys irroratus jalicensis, Goldman, N. Amer. Fauna, 34:60, Sep-
tember 7, 1911.
Range. — Northwestern part of state.
Specimens examined, 3 : nos. 120273-120275 (U. S. Nat. Mus.) from Zamora
Remarks. — As explained in detail by Hall and Villa (1948:254)
these specimens from Zamora are intergrades between L. i. jaliscensis
Hall and Villa R.: Mammals of Michoacan 455
and L. i. acutus and with almost equal propriety could be referred
to either subspecies.
Liomys irroratus acutus Hall and Villa
Northern Spiny Pocket Mouse; Spanish, Raton Norteno
Liomys irroratus acutus Hall and Villa, Univ. Kansas Publ., Mus. Nat
Hist., 1:253, figs. 4-6, July 26, 1948, type from 2 mi. W Patzcuaro, 7,700
ft., Michoacan.
Liomys irroratus alleni, Goldman, N. Amer. Fauna, 34:57, September
7, 1911, part.
Range. — Patzcuaro and vicinity.
Specimens examined, 16: nos. 100170-100184 and 50356 (U. S. N. M.), dis-
tributed, with reference to Patzcuaro, as follows: 3 mi. NW, 6,700 ft., 1; 2 mi.
W, 7,700 ft., 5; 2 mi. W, 6,700 ft., 2; Patzcuaro itself, 1; 5 mi. S, 7,800 ft., 7.
Remarks. — None of the eight females contained embryos. Two
adult males weigh, in grams, 71.5 and 65.1; the average and extreme
weights for five adult females are 50.8 (44.8-61.8).
Liomys irroratus alleni (Coues)
Northern Spiny Pocket Mouse; Spanish, Raton Norteno
Heteromys alleni Coues, Bull. Mus. Comp. Zool., 8:187, March, 1881,
type from Rio Verde, San Luis Potosi, Mexico.
Liomys irroratus alleni Goldman, N. Amer. Fauna, 34:56, September
7, 1911.
Range. — Northeastern part of state.
Specimens examined, 5: nos. 50325-50329 (U. S. Nat. Mus.) from Queren-
daro.
Remarks. — The specimens from Querendaro are not typical of the
subspecies L. i. alleni in that the shape of the interparietal bone and
width of the basisphenoid bone are almost exactly intermediate
between the conditions obtaining in typical L. i. alleni and topotypes
of L. i. acutus.
Reithrodontomys megalotis saturatus Allen and Chapman
Western Harvest Mouse; Spanish, Raton Orejudo
Reithrodontomys saturatus Allen and Chapman, Bull. Amer. Mus. Nat.
Hist,, 9:201, June 16, 1897, type from Las Vigas, Veracruz.
Reithrodontomys megalotis saturatus, Howell, N. Amer. Fauna, 36:36,
June 5, 1914.
Range. — Northeastern part of state.
Specimens examined, 12: nos. 100202-100212, 100273, from 1 mi. N Zamora,
5,450 ft.
Remarks. — Howell (1914:37) referred nine specimens from Na-
huatzin to this race and two from the same place to the subspecies
R. m. zacatecae. Our specimens from Zamora agree with topotypes
456 University of Kansas Publs., Mus. Nat. Hist.
of R. m. saturatus and with specimens of that subspecies from the
Valley of Mexico in dark color and large size.
If our specimens of Reithrodontomys megalotis are correctly iden-
tified, subspecifically, R. m. zacatecae ranges southward around the
western end of the geographic range of R. m. saturatus.
Where R. megalotis and R. fulvescens occur together, we find the
skull of the latter to be distinguishable by: a median spine on the
posterior border of the hard palate (truncate in R. megalotis) ; an
S-shaped instead of a C-shaped pattern on the worn occlusal face
of the last lower molar; and two re-entrant angles, on the lateral
side on the worn occlusal surface of the third upper molar, reaching
halfway across the tooth whereas in R. megalotis the anterior re-
entrant angle is wanting or extends less than a third of the way
across the crown surface of the tooth.
Reithrodontomys megalotis zacatecae Merriam
Western Harvest Mouse; Spanish, Raton Orejudo
Reithrodontomys megalotis zacatecae Merriam, Proc. Washington
Acad. Sci., 3:557, November 29, 1901, type from Valparaiso Mountains,
Zacatecas.
Range. — From northwestern part of state south through its central part.
Specimens examined, 53: nos. 100217-100269; distributed bv localities, with
reference to Patzcuaro, as follows: 3 mi. N, 6,700 ft., 3; 3 mi. N, 6.800 ft., 1;
3V2 mi. S, 7,900 ft., 3; 4 mi. S, 7,800 ft, 31 ; 5 mi. S, 7,800 ft, 9; 9 mi. SE, 8,000
ft., 6.
Remarks. — Howell (1914:40) has referred four specimens from
Patamban to this subspecies, and our large series from the vicinity
of Patzcuaro is also referred to R. m. zacatecae because of small
size and reddish (less blackish) color. We lack typical specimens
of R. m. zacatecae from the type locality for comparison and our
knowledge of zacatecae is derived from Howell's (1914:39) descrip-
tion of it.
Average measurements of 5 adult males of R. m. zacatecae from
the Patzcuaro area showing much wear on the teeth compared with
those of five specimens from the Zamora area, of corresponding sex
and age of R. m. saturatus reveal the smaller size of R. m. zacatecae:
Total length, 157, 166; length of tail, 84, 84; length of hind foot,
19.6, 20.1; length of ear from notch in flesh, 14.8, 14.0; basilar
length, 16.2, 16.6; length of nasals, 8.3, 8.5; zygomatic breadth,
11.1, 11.5; mastoid breadth, 9.9, 10.2; breadth of rostrum, 3.8, 4.0;
interorbital construction, 3.2, 3.1; alveolar length of maxillary tooth-
row, 3.5, 3.5; length of rostrum, 7.7, 8.0.
Hall and Villa R.: Mammals of Michoacan 457
Reithrodontomys fulvescens tenuis Allen
Fulvous Harvest Mouse; Spanish, Raton Moreno
Reithrodontomys tenuis Allen, Bull. Amer. Mus. Nat. Hist., 12:15,
March 4, 1899, type from Rosario, Sinaloa.
Reithrodontomys fulvescens tenuis, Howell, N. Amer. Fauna, 36:45,
June 5, 1914.
Range. — Western part of state.
Specimens examined, 27: nos. 100213-100216. 100274-100277, 100293-100311,
distributed bv localities as follows: 11 mi. W Zamora, 5,750 ft.. 2; 6^2 mi. W
Zamora, 5,950 ft., 2; 6 mi. W Zamora, 5,950 ft,, 4; 1% mi. SSE Tacambaro,
5,700 ft,, 2; 1% mi. S Tacambaro, 5,700 ft., 1; 1 mi. E and 2% mi. S Tacam-
baro, 7; 4 mi. S and I mi. E Tacambaro, 4,700 ft., 5; 1 mi. E and 5 mi. S
Tacambaro, 4,000 ft., 1; 1 mi. E and 6 mi. S Tacambaro, 4,000 ft., 3.
Remarks. — Of the 19 specimens from the vicinity of Tacambaro,
only two have the underparts reddish as does R. f. inexspectatus.
The upper parts are less reddish than in R. f. inexspectatus but
more reddish and less blackish than in R. /. toltecus from the valley
of Mexico or than in R. f. toltecus from Zamora. The external
measurements and cranial measurements are less than in R. f.
toltecus or R. f. inexspectatus and are as small as, or even smaller
than, those of R. f. tenuis to the northward or than those of R. f.
nelsoni to the westward. Relying only on printed descriptions of
R. f. tenuis and R. f. nelsoni, we are inclined to refer our specimens
to R. /. tenuis although the reddish color, we suppose, is evidence of
intergradation with R. f. nelsoni and R. f. inexspectatus.
The four skins from Zamora are gray, as opposed to reddish, both
above and below and in this respect they agree with the description
of R. f. tenuis. They are lighter-colored (grayer) than either R. f.
toltecus or R. f. inexspectatus. The four specimens from Zamora
are larger than animals from the vicinity of Tacambaro and average
slightly smaller than topotypes of R. f. inexspectatus.
By identifying our specimens as R. f. inexspectatus and R. f.
tenuis, we are left without any specimens that we, ourselves, have
examined, which are referable to the subspecies R. f. toltecus. The
specimens from Los Reyes which Howell (1914:47) referred to R. /.
toltecus have not been seen by us, and we guess, on the criteria
used by us, that the animals would be referable to R. f. tenuis.
Because Dr. Emmet T. Hooper has a revisionary study of the
Mexican Reithrodontomys underway, we have not attempted to
bring together all of the pertinent material from different collections
as would be required for an ideally thorough analysis of the geo-
graphic variation in Reithrodontomys megalotis and Reithro-
dontomys fulvescens.
458
University of Kansas Publs., Mus. Nat. Hist.
As illustrative of statements made concerning the average size of
Reithrodontomys fulvescens, the following measurements, all of
specimens with much wear on each of the molar teeth, are recorded.
Locality
OJ £
§
k%
•2°
£ k
A
1
H
s
0
a
c
a
c
■a
o
M
a
-I
ho
B
A
03
M
o
J3
03
0)
t-
X!
o
ca
0
be
>>
S)
.a
-*j
-a
a)
S
Xl
T3
'3
■8
cc
<5
1
- i
— 01
H *
£H
5!
Vail. Mex. . .
3
dd
200
114
22.0
17.6
9.3
12.0
10.7
3.7
Patzcuaro . .
5
dd
170
101
20.6
16.7
8.7
11.6
10.6
3.7
100215
d*
153
82
19.0
15.9
7.8
10.9
10.3
3.5
100275
c?
184
101
21.0
16.5
8.0
11.1
10.4
3.6
Tacambaro .
5
do*
159
91
19.4
15.5
8.0
10.5
9.6
3.3
Vail. Mex...
3
9 9
184
103
21.0
16.6
8.7
11.4
10.6
3.6
Patzcuaro. .
5
9 9
182
104
21.6
16.8
8.6
11.6
10.5
3.6
Zamora ....
5
9 9
159
91
19.0
16.0
8.4
11.4
10.0
3.5
Tacambaro .
5
9 9
149
87
18.6
14.9
7.7
10.4
9.5
3.4
Reithrodontomys fulvescens inexspectatus Elliot
Fulvous Harvest Mouse; Spanish, Raton Moreno
Rhithrodontomys inexspectatus Elliot, Field Columb. Mus. zool. ser.,
3:145, February, 1903, type from Patzcuaro, Michoacan.
Reithrodontomys fulvescens toltecus, Howell, N. Amer. Fauna, 36:51,
June 5, 1914, part.
Range. — Central Michoacan; limits of range unknown.
Specimens examined, 15: nos. 100278-100292, distributed by localities as fol-
lows: 3 mi. N Patzcuaro, 6,800 ft,. 6; 3 mi. NW Patzcuaro, 6,700 ft., 3; 2 mi.
W Patzcuaro, 7,600 ft., 2; 2 mi. W Patzcuaro, 7,700 ft., 4.
Remarks. — Howell (1914:51) made R. inexspectatus Elliot a syn-
onym of R. f. toltecus and perhaps we should follow him in this.
The facts are that in our large series from the vicinity of Patzcuaro,
the upper parts are more reddish than in R. /. toltecus from the
valley of Mexico, and more reddish than in R. f. tenuis if we correctly
interpret Howell's (op. c?'£.,:45) description of R. f. tenuis. In the
color of the underparts the series is, to us, indistinguishable from
topotypical toltecus and therefore has more reddish underparts than
Hall and Villa R.: Mammals of Michoacan 459
R. /. tenuis, as we know R. /. tenuis, from Howell's (loc. cit.) descrip-
tion. In size, the series from Patzcuaro is intermediate between
R. f. tenuis and R. f. toltecus but nearer the latter.
Reithrodontomys chrysopsis chrysopsis Merriam
Volcano Harvest Mouse; Spanish, Raton Dorado
Reithrodontomys chrysopsis Merriam, Proc. Biol. Soc. Washington,
13:152, June 13, 1900, type from Mount Popocatepetl, Mexico.
Reithrodontomys chrysopsis chrysopsis, Howell, N. Amer. Fauna, 36:66,
June 5, 1914.
Range.— Patamban, 1,200 feet elevation as now known.
Remarks. — Howell (1914:68) lists nine specimens from Patam-
ban. We have not examined these specimens. He listed at the same
time seven of the specimens from Tancitaro, but we have found
specimens from Tancitaro to be of another subspecies, R. c. seclusus.
Accordingly, we are in doubt as to whether the mice from Patamban
are subspecies chrysopsis, seclusus, or an unnamed subspecies and
our use here of the name Reithrodontomys chrysopsis chrysopsis
for them is, of course, provisional.
Reithrodontomys chrysopsis seclusus Hall and Villa,
Volcano Harvest Mouse; Spanish, Raton Dorado
Reithrodontomys chrysopsis seclusus Hall and Villa, Proc. Biol. Soc.
Washington, 62: 163, August 23, 1949, type from Mount Tancitaro, 7,800 ft,
Michoacan.
Reithrodontomys chrysopsis chrysopsis, Howell, N. Amer. Fauna, 36:66,
June 5, 1914, part.
Range. — Known only from Mount Tancitaro, from 6,000 feet elevation up
to at least 11,000 feet.
Specimens examined, 22: nos. 51407-51411, 52110-52126, all from Mount
Tancitaro, distributed by altitude as follows: 6,000 ft., 5; 7,800 ft, 10; 10,500
ft, 1; 11,000 ft, 1; no altitude recorded, 5.
Remarks. — The 22 specimens in the Chicago Natural History
Museum are remarkably uniform in color in spite of differences in
age ; 17 are so young as to have the first upper molar only slightly
worn and 5 are adults. In preparing the original description of
R. c. seclusus, known only from specimens in the summer pelage,
comparison of color was made with only the winter pelage of R. c.
chrysopsis and it was pointed out that the differences noted in color
between the two subspecies might be seasonal rather than sub-
specific. A summer specimen of R. c. chrysopsis (K. U. 17980,
taken on June 15, 12 km. ESE Amecameca, 11,500 ft,), is available
as the present account is being written. In direct comparison with
the original material of R. c. seclusus, all in summer pelage, and
460 University of Kansas Publs., Mus. Nat. Hist.
in comparison with a specimen of R. c. chrysopsis in winter pelage
(January 18), from 30 km. E Amecameca, the summer pelage of no.
17980 is more blackish than the winter pelage and therein more
closely resembles that of R. c. seclusus. The same is true of the
more sparsely haired tail. The ears, however, are blackish as in
winter-taken R. c. chrysopsis. This feature and also the large size
and different cranial proportions of specimen no. 17980 are in ac-
cordance with the differences between R. c, chrysopsis and R. c.
seclusus as recorded in the original description of the latter.
Baiomys taylori analogus (Osgood)
Northern Pygmy Mouse ; Spanish, Raton Pigmeo Norteno
Pcromyscus taylori analogus Osgood, N. Amer. Fauna, 28:256, April
17, 1909, type from Zamora, Michoacan.
Baiomys taylori analogies, Miller, N. Amer. Land. Mamm., 1911, p.
137, December 31, 1912.
Range. — Northwestern part of state.
Specimens examined, 35: nos. 100331-100365, distributed by localities as fol-
lows: 8 mi. N Zamora, 5,500 ft.. 2; 11 mi. W Zamora, 5,750 ft., 1; 6x/2 mi. W
Zamora, 5,950 ft., 2; 4 mi. W Zamora, 5,450 ft,, 1; 3 mi. NW Patzcuaro, 6,700
ft., 21; 3 mi. N Patzcuaro, 6,800 ft,, 5; 2 mi. W Patzcuaro, 6,700 ft., 3.
Remarks.— Osgood (1909:257) has recorded this subspecies also
from Acambaro and Los Reyes. We found that these animals got
caught in our traps almost as often in the daytime as at night and
concluded that they were less nocturnal, or at any rate more diurnal,
than any of the other cricetine mice at the places in Michoacan
where we trapped.
Baiomys musculus musculus (Merriam)
Tropical Pygmy Mouse; Spanish, Raton Pigmeo Tropical
Sitomys musculus Merriam, Proc. Biol. Soc. Washington, 7:170, Sep-
tember 29, 1892, type from Colima, Colima.
Peromyscus musculus, Allen and Chapman, Bull. Amer. Mus. Nat.
Hist,, 9:203, June 16, 1897.
Baiomys musculus, Meams, II. S. Nat. Mus. Bull., 56:381, April 13,
1907.
Range.— Dry, tropical, southwestern parts of state.
Specimens examined, 41: nos. 100366-100406; distributed by localities as fol-
lows- 1% mi. S Tacambaro, 5,700 ft., 1; 1 mi. E and 2V2 mi. S Tacambaro,
4 700 ft 11 • 4 mi. S and 1 mi. E Tacambaro, 10; 1 mi. E and 5 mi. S Tacam-
baro, 4,000 ft., 1; 6 mi. S Tacambaro, 4,000 ft,, 4; 1 mi. E and 6 mi. S Tacam-
baro, 4,000 ft., 14.
Remarks.— Osgood (1909:258) has recorded this subspecies also
from La Huacana and La Salada. Three adult males with much
worn teeth weigh, in grams, 8.3, 9.3, and 10.8. Weights of three
adult, nonpregnant, females are 8.1, 9.4, and 9.7. None of our 13
females was pregnant.
Hall and Villa R.: Mammals of Michoacan 461
Peromyscus maniculatus labecula Elliot
Deermouse; Spanish, Raton Cuatralvo
Peromyscus labecula Elliot, Field Columb. Mus., zool. ser., 3:143,
February (March?), 1903, type from Ocotlan, Jalisco.
P[eromyscus]. slonoriensis'] . labecula, Osgood, Proc. Biol. Soc. Wash-
ington, 17:57, March 21, 1904.
Range. — Northwestern part of state.
Specimens examined, 59: nos. 51402-51406. 52175, 52176, 100407-100447,
100484; and 1366-1368, 1370-1372, 1374-1377. of Bernardo Villa R., distributed
bv localities as follows: Tancitaro, 7; 11 mi. W Zamora, 5,750 ft., 3; 2 mi. W
Patzcuaro, 7.700 ft., 4; ZV2 mi. S Patzcuaro, 7,800 ft., 2; 4 mi. S Patzcuaro,
7.800 ft., 3; 5 mi. S Patzcuaro, 7,800 ft,, 24; 9 mi. SE Patzcuaro, 8,000 ft., 6;
iy2 km. N San Juan, 2,250 M., 3; 1 km. NNE San Juan, 2,250 M., 7.
Remarks. — The zygomatic arches are less widely flaring in speci-
mens from Tancitaro than in those from Patzcuaro.
Peromyscus perfulvus Osgood
Marsh Mouse; Spanish, Raton Brenero
Peromyscus perfulvus Osgood, Jour. Mamm., 26:299, November 14,
1945, type from 10 kilometers west of Apatzingan, 1,040 ft., Michoacan.
Range. — Known only from the semitropical Life-zone in western Michoacan.
Specimens examined, 5: nos. 100593, 100595, 100597, 100598, and 100600, all
from 1 mi. E and 6 mi. S Tacambaro, 4,000 ft.
Remarks. — J. R. Alcorn took these specimens between fields of
sugar cane in- tall (5 ft, high) grass growing in a belt 10 feet or so
wide along side a stream, which a person could step across. The
one adult, no. 100597, was recognized at the time of capture as
different from any other species known to us, by reason of the long,
unicolored, sparsely-haired tail and nearly clear Cinnamon Rufous
color above, white underparts, white feet, and dark brown ears. The
four other specimens in darker immature pelage are plumbeous and
cinnamon whereas immature individuals of comparable age of
Peromyscus banderanus caught in the same place are plumbeous
above and lack the cinnamon color. The immature animals of the
two species differ in color more than do the adults. Osgood (1945
:300) has recorded eleven specimens from the type locality. Our
one adult weighs 52.4 grams.
Peromyscus boylii evides Osgood
Brush Mouse; Spanish, Raton de Chaparral
Peromyscus spicilegus evides Osgood, Proc. Biol. Soc. Washington,
17:64, March 21, 1904, type from Juquila, Oaxaca.
Peromyscus boylei evides Osgood, N. Amer. Fauna, 28:152, April 17,
1909.
Range. — Northern part of state.
Specimens examined, 22: nos. 100450-100471, distributed by localities as fol-
462 University of Kansas Publs., Mus. Nat. Hist.
lows: 1V> mi. SSE Tacambaro, 5,700 ft., 16; 1% mi S Tacambaro, 5,700 ft.,
1 ; 1 mi. E and 2V2 mi. S Tacambaro, 4,700 ft,, 5.
Remarks. — Osgood (1909:153) recorded 31 specimens from Los
Reyes. Weights recorded by the collectors of our specimens for
13 males from the vicinity of Tacambaro, are 24.8 (17.2-37.1) and
for 9 females 25.0 (20.0-31.5) grams. These weights include those
of subadults as well as those of adults.
Peromyscus boylii levipes Merriam
Brush Mouse; Spanish, Raton de Chaparral
Peromyscus levipes Merriam, Proc. Biol. Soc. Washington, 12:123,
April 30, 1898, type from Mount Malinche, Tlaxcala.
Peromyscus boylei levipes, Osgood, N. Amer. Fauna, 28:153, April 17,
1909.
Range. — Probably central-eastern part of state.
Remarks. — Osgood (1909:155) records one specimen of this sub-
species from Patzcuaro.
Peromyscus hylocetes Merriam
Woods Mouse; Spanish, Raton Ocotero
Peromyscus hylocetes Merriam, Proc. Biol. Soc. Washington, 12:124,
April 30, 1898, type from Patzcuaro, 7,000 feet, Michoacan; Osgood, N.
Amer. Fauna, 28:159, pi. 3, fig. 8, April 17, 1909.
Range. — Mountainous parts of state.
Specimens examined, 24: nos. 100472, 100542, both from 9 mi. SE Patzcuaro
at 8,000 feet altitude; 1358-1365 of Bernardo Villa R., from Cerro Curitzaran,
3.5 km. NNW San Juan, 2,200 M.; 52178, 52185, 52187-52192, 52197, 52198,
52203, 52213-52215, from Tancitaro at elevations of 6,000 to 10,000 ft.
Remarks. — All of the specimens were taken in pine forest. The
old male from nine miles southeast of Patzcuaro weighs 43.7 grams
and the younger male from there 35 grams. The adults from
Tancitaro have longer diastemae and some have the braincase more
prolonged posteriorly, than in specimens from the vicinities of San
Juan and Patzcuaro but individual variation is considerable and we
are unable to differentiate some of the adults from Tancitaro from
those from elsewhere.
Peromyscus truei gratus Merriam
Piiion Mouse; Spanish, Raton Pinonero
Peromyscus gratus Merriam, Proc. Biol. Soc. Washington, 12:123, April
30, 1898, type from "Tlalpam," D. F.
Peromyscus sagax Elliot, Field Columb. Mus., Chicago, zool. ser., 3:142,
February, 1903, type from La Palma, Michoacan.
Peromyscus pavidus Elliot, Field Columb. Mus., Chicago, zool. ser.,
3:142, February, 1903, type from Patzcuaro, Michoacan.
Peromyscus zelotes Osgood, Proc. Biol. Soc. Washington, 17:67, March
21, 1904, type from Querendaro, Michoacan.
Hall and Villa R.: Mammals of Michoacan 463
Peromyscus truei gratus, Osgood, N. Amer. Fauna, 28:173, April 17,
1909.
Range. — Northern part of state.
Specimens examined, 42: nos. 100448, 100473-100483, 100485-100509, 8700,
8702, 8703, 8896, 8897, distributed by localities as follows: 8 mi. N Zamora,
5,500 ft., 2; 11 mi. W Zamora, 5,750 ft., 11; SVi mi. W Zamora, 5,950 ft., 4;
6 mi. W Zamora, 5,950 ft., 4; Rio Duaro, 9 mi. E Zamora, 5,500 ft., 1; 3 mi. N
Patzcuaro, 6,800 ft., 1; 3 mi. NW Patzcuaro, 6,700 ft., 10; V& mi. NW
Patzcuaro, 6,700 ft., 1; 2 mi. W Patzcuaro, 6,700 ft., 3; Patzcuaro (Chicago Nat.
Hist. Mus.), 5.
Remarks. — The ear measured from the notch is shortest, 19 (18-
21), at Zamora, intermediate, 21 (19-23), at Patzcuaro, and longest,
21.8 (20-23), at the type locality of gratus in the Valley of Mexico.
Peromyscus melanophrys zamorae Osgood
Blackish Mouse; Spanish, Raton Obscuro
Peromyscus melanophrys zamorae Osgood, Proc. Biol. Soc. Washing-
ton, 17:65, March 21, 1904, type from Zamora, Michoacan; N. Amer.
Fauna, 28:187, April 17, 1909.
Range. — Northern part of state.
Remarks. — Insofar as we know, this mouse has been taken in
Michoacan only at the type locality.
Peromyscus banderanus banderanus Allen
Tarascan Mouse; Spanish, Raton Tarasco
Peromyscus banderanus Allen, Bull. Amer. Mus. Nat. Hist., 9:51,
March 15, 1897, type from Valle de Banderas, Nayarit; Osgood, Jour.
Mamm., 26:300, November 14, 1945.
Peromyscus banderanus vicinior Osgood, N. Amer. Fauna, 28:209, 210,
April 17, 1909, part.
Range. — Northern and eastern parts of state.
Remarks. — Specimens from Los Reyes referred to the subspecies
P. b. vicinior by Osgood (1909:209-210) were later characterized
by Osgood (1945:300) as agreeing with specimens from Zitacuaro,
and Osgood (loc, tit.) thought that those from both Los Reyes and
Zitacuaro were not P. b. vicinior but possibly P. b. banderanus. He
had this material set aside for further study when he showed it to
one of us (Hall) in 1945. It was his intention to revise the entire
species (P. banderanus) but so far we know never did this before
his death.
Peromyscus banderanus vicinior Osgood
Tarascan Mouse; Spanish, Raton Tarasco
Peromyscus banderanus vicinior Osgood, Proc. Biol. Soc. Washington,
17:68, March 21, 1904, type from La Salada, Michoacan.
Range. — Hot valleys of western part of state.
Specimens examined, 53: nos. 100543-100592, 100594, 100596, 100599, dis-
tributed by localities, from Tacambaro, as follows: IVi mi. S and 1 mi. E,
464 University of Kansas Publs., Mus. Nat. Hist.
4,700 ft., 21 ; 4 mi. S and 1 mi E, 4,500 ft., 10; 6 mi. S, 4,000 ft., 6; 6 mi. S and
1 mi. E, 4,000 ft., 16.
Remarks. — There is much variation in size in our animals. The
three largest males weigh, in grams, 67.5, 50.3, 48.9 and correspond-
ing figures for the two heaviest, nonpregnant, females are 53.5 and
48.3 grams. Of the 14 adult females, only one was recorded as
having embryos; it had two embryos each 24 millimeters in crown
to rump length. Where we trapped among big boulders and among
the roots of trees of the genus Ficus, Peromyscus banderanus vicin-
ior was the only species of the genus taken. Peromyscus boylii
evides occurred in the less tropical vegetation, altitudinally and
zonally above P. b. vicinior.
Peromyscus melanotis Allen and Chapman
Black-eared Deermouse; Spanish, Raton Montanero
Peromyscus melanotis Allen and Chapman, Bull. Amer. Mus. Nat.
Hist., 9:203, June 16, 1897, type from Las Vigas, 8,000 ft., Veracruz.
Range. — Higher mountains throughout state.
Specimens examined, 33: nos. 51397-51401, 52142-52166, 52172-52174, from
Tancitaro.
Remarks. — The elevation recorded on the label of one specimen
is 9,000 feet and on the labels of other specimens is no lower than
10,500 feet and on some is as high as 12,000 feet. The elevation of
capture is not recorded for two specimens. Osgood (1909:112)
previously recorded the species from 12,000 feet elevation on Mount
Tancitaro.
Oryzomys couesi regillus Goldman
Tropical Rice Rat; Spanish, Rata Arrocera Tropical; Tarascan word
for rat is Jeyaquihuiri (Hayakewire), or Sarisi
Oryzomys couesi regillus Goldman, Proc. Biol. Soc. Washington, 28:129,
June 29, 1915, type from Los Reyes, Michoacan; Goldman, N. Amer.
Fauna, 43:37, September 23, 1918.
Range. — Plateau region of Northeast Michoacan.
Specimens examined, 22: nos. 100601-100622, distributed by localities as fol-
lows: 1 mi. N Zamora, 5,450 ft., 2; 4 mi. W Zamora, 5,450 ft,, 1; 4 mi. S
Patzcuaro, 7,800 ft., 1; 1 mi. E and 6 mi. S Tacambaro, 4,000 ft., 18.
Remarks — The tooth-row is longest in the specimens from Za-
mora, shortest in those from Tacambaro and intermediate in length
in the one specimen from Patzcuaro. The shorter tooth-row at the
lower elevation (Tacambaro), we interpret as intergradation with
Oryzomys couesi mexicanus. In color the specimens from Tacam-
baro are, to us, indistinguishable from those from Zamora and
Patzcuaro but the color is notably darker than that of specimens
from the vicinity of Apatzingan which are here referred to the sub-
Hall and Villa R.: Mammals of Michoacan 465
species Oryzomys couesi mexicanus. The largest male, fully adult
from Zamora weighs 82.9 grams and the largest one from the
vicinity of Tacambaro weighs 73.6 grams.
Oryzomys couesi mexicanus Allen
Tropical Rice Rat; Spanish, Rata Arrocera Tropical
Oryzomys mexicanus Allen, Bull. Amer. Mus. Nat. Hist., 9:52, March
15, 1897, type from Hacienda San Marcos, 3,500 ft., Tonila, Jalisco.
Oryzomys couesi mexicanus, Goldman, N. Amer. Fauna, 43:33, Sep-
tember 23, 1918.
Range. — Semitropical and tropical western part of the state.
Specimens examined, 14: nos. 52018-52023, 52063-52070, from 1,040 ft., 10
kms,. W Apatzingan.
Remarks. — These specimens are notably paler and have shorter
tooth-rows than those referred to O. c. regillus.
Oryzomys fulvescens lenis Goldman
Fulvous Rice Rat; Spanish, Rata Arrocera Pigmea
Oryzomys fulvescens lenis Goldman, Proc. Biol. Soc. Washington,
28:130, June 29, 1915, type from Los Reyes, Michoacan; N. Amer. Fauna,
43:91, September 23, 1918.
Range. — Semitropical parts of state.
Remarks. — The type and one topotype so far as we know are the
only specimens of this species to have been obtained from the state.
The size is hardly larger than that of a large Reithrodontomys.
Sigmodon melanotis Bailey
Fulvous Cotton Rat; Spanish, Rata Algodonera Leonada
Sigmodon melanotis Bailey, Proc. Biol. Soc. Washington, 15:114, June
2, 1902, type from Patzcuaro, 7,000 ft., Michoacan.
Range. — Central Michoacan, as now known.
Specimens examined, 8: nos. 100623-100626, 52089-52092, distributed by lo-
calities as follows: 2 mi. W Patzcuaro (7,400 and 7,700 ft.), 2; 3*6 mi. S
Patzcuaro, 7,800 ft., 2; Tancitaro, 6,000 ft., 4.
Remarks. — This species was taken along with the species S.
hispidus two miles west of Patzcuaro, and can be easily distinguished
from the latter by the dark reddish as opposed to grayish color of
the upperparts and by the shorter hind foot (less instead of more
than 32.5 mm.).
Sigmodon hispidus mascotensis Allen
Hispid Cotton Rat; Spanish, Rata Algodonera Setosa
Sigmodon mascotensis Allen, Bull. Amer. Mus. Nat. Hist., 9:54, March
15, 1897, type from San Sebastian, near Mascota, Jalisco.
Sigmodon hispidus mascotensis, Bailey, Proc. Biol. Soc. Washington,
15:108, June 2, 1902.
Range. — Larger part of state.
466 University of Kansas Publs., Mus. Nat. Hist.
Specimens examined, 4: nos. 100629, 3 mi. N Patzcuaro, 6,700 ft.; 100630, 2
mi. W Patzcuaro, 7,700 ft,; 100632, 1% mi. S Tacambaro, 5,700 ft.; 100631, 6 mi.
S and 1 mi. E Tacambaro, 4,000 ft.
Remarks. — Bailey (1902:109) records a "very large" specimen
from Querendaro. Our specimens have shorter molariform tooth-
rows than do those from nearer the type locality, for example, those
from Tuxpan, Las Canoas, and Artenkiki, all three places in Jalisco.
Sigmodon hispidus atratus Hall
Hispid Cotton Rat; Spanish, Rata Algodonera Setosa
Sigmodon hispidus atratus Hall, Proc. Biol. Soc. Washington, 62:149,
August 23, 1949, type from QV2 mi. W Zamora, 5,950 ft., Michoacan.
Range. — Known only from Zamora and the type locality.
Specimens examined, 2: nos. 100628 (the holotvpe), GV2 mi. W Zamora, 5,950
ft.; 120268 (U. S. Nat. Mus., Biol. Surveys Coll.), Zamora.
Remarks. — When the present account first was prepared our speci-
men from six and a half miles west of Zamora was tentatively re-
ferred to S. h. mascotensis. Subsequently a second specimen, from
Zamora, was found. It agreed with the specimen from six and a
half miles west of Zamora. Inasmuch as the second specimen agrees
with the first and since each of the two differs from any previously
described kind, a name and description were published in time to be
inserted in the present account. From S. h. mascotensis, S. h. atratus
differs in shorter hind foot, darker upper parts, more densely haired
tail, shorter skull, more convex dorsal longitudinal outline of skull,
posteriorly constricted anterior palatine foramina instead of parallel-
sided foramina, and shorter and less decurved anterior process of
maxillary arm of zygoma.
Neotomodon alstoni alstoni Merriam
Volcano Mouse ; Spanish, Raton de Los Volcanes
Neotomodcm alstoni Merriam, Proc. Biol. Soc. Washington, 12:128,
April 30, 1898, type from Nahuatzin, 8,500 ft., Michoacan.
Range. — Higher mountains of state.
Specimens examined, 22: nos. 52179-52184, 52186, 52193-52196, 52199, 52200,
52204-52212, all from Mount Tancitaro, distributed bv localities as follows:
7.800 ft,, 5; 7,850 ft., 3; 10.000 ft,, 4; 10,200 ft., 5; 10.500 ft,, 1; 10,800 ft., 1;
11,000 ft, 2; 11,400 ft, 1.
Remarks. — The taking of specimens on Mount Tancitaro extends
the known geographic range of Neotomodon approximately 75 kilo-
meters to the southwestward ; the westernmost locality previously
known was Nahuatzin, the type locality.
Hall and Villa R.: Mammals of Michoacan 467
Nelsonia goldmani Merriam
Dwarf Wood Rat; Spanish, Rata Montcra Minuscula
Nelsonia goldmani Merriam, Proc. Biol. Soc. Washington, 16:80,
May 29, 1903, type from Mount Tancitaro, Michoacan.
Range. — Known only from the type locality.
Remarks. — In the original description three specimens are re-
corded from the type locality.
Neotoma latifrons Merriam
White-throated Wood Rat; Spanish, Rata Montera Frentuda
Neotoma latifrons Merriam, Proc. Biol. Soc. Washington, 9:121, July
2, 1894, type from Querendaro, Michoacan.
Range. — Known only from the type locality.
Neotoma ferruginea tenuicauda Merriam
Ferruginous Wood Rat; Spanish, Rata Ferruginosa
Neotoma tenuicauda Merriam, Proc. Biol. Soc. Washington, 7:169,
September 29, 1892, type from north slope of Sierra Nevada de Colima,
12,000 ft., Colima.
Neotoma ferruginea tenuicauda, Howell, N. Amer. Fauna, 31:73, Octo-
ber 19, 1910.
Range. — Probably all but southern tropical part of state.
Specimens examined, 4: nos. 100633 from 9 mi. SE Patzcuaro, S.000 ft., and
52177, 51390, 51391, from Tancitaro, the elevation being given as 7,850 ft. on no.
52177.
Remarks. — Our one specimen from 9 miles southeast of Patzcuaro
was caught in a small steel trap set at a meat bait.
Microtus mexicanus salvus Hall
Mexican Meadow Mouse; Spanish, Metorito
Microtus mexicanus salvus Hall, Univ. Kansas Publ., Mus. Nat. Hist.,
1:426, December 24, 1948, type from Mount Tancitaro, 11,400 ft., Micho-
acan.
Range. — Known only from Mount Tancitaro at elevations of 7,800 to 11,400
feet.
Specimens examined, 14: nos. 51412, 51413, 52093, 52095-52099, 52101, 52103-
52107, all from Mount Tancitaro, distributed by elevations as follows: 11,400
ft,, 8; 11,000 ft., 2; 7,800 ft., 1; no elevation recorded, 3.
Microtus mexicanus fundatus Hall
Mexican Meadow Mouse; Spanish, Metorito
Microtus mexicanus fundatus Hall, Univ. Kansas Publ., Mus. Nat.
Hist., 1 :425, December 24, 1948, type from ZVi mi. S. Patzcuaro, 7,900 ft.,
Michoacan.
Range. — Central part of state.
Specimens examined, 59: nos. 100636-100694, distributed, with reference to
the town of Patzcuaro, as follows: 3% mi. S, 7,900 ft., 9; 4 mi. S, 7,800 ft, 16;
5 mi. S, 7,800 ft, 26; 9 mi. SE, 8,000 ft, 8.
Remarks. — Of the 23 females, only one was pregnant. It had two
468 University of Kansas Publs., Mus. Nat. Hist.
embryos. Average and extreme weights of ten adults of each sex,
are: males, 37.8 (31.5-48.2); females, 38.0 (31.0-48.6) grams. Our
specimens were trapped in well-defined runways beneath a rail
fence where there was a growth of grass sufficient to make a cover
for the runways. Bailey (1900:54-55) has recorded under the name
Microtus mexicanus phaeus specimens from Nahuatzin which may
be referable to the subspecies M. m. fundatus.
Mus musculus subsp.?
House Mouse; Spanish, Raton Casero
Mus musculus Linnaeus, Systema Naturae, ed. 10, 1:62, 1758, type from
Upsala, Sweden.
Range. — Probably throughout state.
Specimens examined, 4 : nos. 100696-100699, of which one is from Tacambaro,
5,700 ft., and 3 are from 4 mi. S and 1 mi. E Tacambaro, 4,500 ft,
Remarks. — In each of our specimens the belly is dark, approxi-
mately as dark as the back. The specimens caught by us were
living in the wild ; that is to say, they were not caught in and around
buildings. Elliot (1903:141) records the species from Patzcuaro.
Rattus rattus alexandrinus (Geoffroy)
Black Rat; Spanish, Rata Negra
Mus alexandrinus Geoffroy, Catal. Mammif. du Mus. Nat. d'Hist.,
Paris, p. 192, 1803, type from Alexandria, Egypt.
R[attus~\. rattus alexandrinus, Hinton, Jour. Bombay Nat. Hist. Soc,
26:63, December 20, 1918.
Range. — Probably throughout tropical and subtropical parts of state; re-
corded also from Patzcuaro.
Specimens examined, 3: nos. 52027, 52033 from Tancitaro and 8900 from
Patzcuaro.
Remarks. — These specimens answer well to the description of
R. r. alexandrinus except that no. 8909, taken in May, 1901, by F. E.
Lutz, has yellowish underparts suggestive of Rattus rattus frugi-
vorous. In the town of Tacambaro we saw a freshly killed rat of this
species which was all black.
Sylvilagus floridanus subcinctus (Miller)
Florida Cottontail; Spanish, Conejo de Florida
Lepus floridanus subcinctus Miller, Proc. Acad, Nat. Sci. Philadelphia,
p. 386, October 5, 1899, type from Hacienda El Molino, Negrete, Micho-
acan.
Sylvilagus floridanus subcinctus, Lyon, Smiths. Misc. Coll., 45:336,
June 15, 1904; Nelson, N. Amer. Fauna, 29:180, August 31, 1909.
Range. — Northeastern part of state.
Remarks. — Nelson (1909:181) records specimens from Acambaro,
Querendaro and the type locality.
Hall and Villa R.: Mammals of Michoacan 469
Sylvilagus floridanus restrictus Nelson
Florida Cottontail; Spanish, Conejo de Florida
Sylvilagus floridanus restrictus Nelson, Proc. Biol. Soc. Washington,
20:82, July 22, 1907, type from Zapotlan, Jalisco; Nelson, N. Amer. Fauna,
29:181, August 31, 1909.
Range. — Forested areas of non-tropical part of state except northeastern
part.
Remarks. — Nelson (1909:183) records specimens from Los Reyes,
Patzcuaro and Mount Tancitaro. This species and the Mexican
cottontail are favorite small game for the rural peoples.
Sylvilagus cunicularis cunicularis (Waterhouse)
Mexican Cottontail; Spanish, Conejo Mexicano
Lepus cunicularis Waterhouse, Nat. Hist. Mammalia, 2:132, 1848, type
from Zacualpan (probably in state of Mexico) .
Sylvilagus cunicularis, Nelson, N. Amer. Fauna, 29:239, August 31,
1909.
Range. — Probably all of state except tropical coastal areas where another
subspecies of the same species probably will be found to occur.
Specimens examined, 2: nos. 51965, 51966, from Tancitaro, one specimen
labeled as taken at 6,000 feet altitude.
Remarks. — Nelson (1909:241) has recorded this rabbit also from
Patzcuaro.
Lepus callotis Wagler
•White-sided Jack Rabbit; Spanish, Liebre
Lepus callotis Wagler, Naturliches System der Amphibien, p. 23, 1830,
type from southern end of Mexican Tableland; Nelson, N. Amer. Fauna,
29:122, August 31, 1909.
Range. — Approximately northeastern half of state.
Remarks. — Nelson (1909:124) records specimens from Los Reyes
and Querendaro; we did not see any animals of this species in our
own field work.
Tayassu angulatus humeralis Merriam
Collared Peccary; Spanish, Jabali del Collar; Tarascan, cuchjeramba
(cucheramba) or Juateanapu (whatalanapu)
Tayassu angulatus humeralis Merriam, Proc. Biol. Soc. Washington,
14:122, July 19, 1901, type from Armeria, Colima.
Range. — Approximately southwestern half of state.
Remarks. — Under date of October 11, 1948, Henry W. Setzer
{in litt.) states that four specimens of this species, in the Biological
Surveys Collection in the United States National Museum, were
taken at La Salada, by Nelson and Goldman, and bear catalogue
numbers 126156, 126157, 126158 and 126159. No. 126158 is a fe-
male taken on March 19, 1903. The other three specimens are
470 University of Kansas Publs., Mus. Nat. Hist.
males taken on March 17, 1903. We did not see any animals of
this species in our own field work, and the only materials from
Michoacan actually examined by one of us (Hall) are the skulls
of nos. 126156 and 126158, referred to above, from the Biological
Surveys Collection.
Odocoileus virginianus sinaloae Allen
White-tailed Deer; Spanish, Venado Cola Blanca; Tarascan,
Asiini (Ashumi)
Odocoileus sinaloae Allen, Bull. Amer. Mus. Nat. Hist., 19:613, No-
vember 14, 1903, type from Esquinapa, Sinaloa.
Range. — Probably statewide.
Remarks. — Through the courtesy of Dr. A. Remington Kellogg
we learn that in a manuscript on the deer of the Odocoileus virgini-
anus group, he and the late Major E. A. Goldman had recorded
specimens, in the Biological Surveys Collection of the United States
National Museum, as follows: Nahuatzin, 8,500 ft., nos. 35924/
48232, and 35925/48233; Los Reyes, 5,000 ft., no. 165673; Patzcuaro,
7,000 ft., no. 35535/47819; and Uruapan, 4,500 ft,, no. 13060. We
have not anywhere seen the name combination Odocoileus virgini-
anus sinaloae but from the original description we judge that
Odocoileus sinaloae is to be arranged as a geographic race of the
wide-ranging species Odocoileus virginianus as that species is now
understood.
Dasypus novemcinctus mexicanus Peters
Nine-banded Armadillo; Spanish, Armadillo; Tarascan, Isingu (Esingo)
Dasypus novemcinctus mexicanus Peters, Monatsber. k. preuss. Akad.
Wissensch. Berlin, p. 180, 1864 (name restricted by Bailey, N. Amer.
Fauna, 25:52, October 24, 1905, to the subspecies occurring at Colima).
Range. — Probably statewide.
Specimens examined, 2; nos. 51392 from Tancitaro and 51964 from Apatz-
ingan, 1,040 ft.
Remarks. — The female from Tancitaro is immature as indicated
by the wide-open sutures between the bones of the skull which in
over-all length is only 72.8 mm. The male from Tancitaro is older
and the over-all length of the skull is 98.33 mm. Geographic con-
siderations alone are responsible for our use of the subspecific name
mexicanus; we do not know the morphological features which dis-
tinguish viexicanus from other named subspecies.
Hall and Villa R.: Mammals of Michoacan 471
LITERATURE CITED
Andersen, K.
1908. A monograph of the Chiropteran genera, Uroderma, Enchisthenes,
and Artibeus. Proc. Zool. Soc. London, for 1908:204-319, text figs.
40-58.
Allen, G. M.
1916. Bats of the genus Corynorhinus. Bull. Mus. Comp. Zool., Harvard
College, 60:333-356, 1 pi., April, 1916.
Bailey, V.
1900. Revision of American voles of the genus Microtus. N. Amer. Fauna,
17:1-88, 5 pis., 17 figs, in text, June 6, 1900.
1902. Synopsis of the North American Species of Sigmodon. Proc. Biol.
Soc. Washington, 15:101-116, June 2, 1902.
Benson, S. B.
1947. Description of a mastiff bat (genus Eumops) from Sonora, Mexico.
Proc. Biol. Soc. Washington, 60:133-134, December 31, 1947.
Davis, W. B.
1944. Notes on Mexican mammals. Jour. Mamm, 25:370-403, 1 fig. in
text, December 12, 1944.
Elliot, D. G.
1903. A list of a collection of Mexican mammals with descriptions of
some apparently new forms. Field Columb. Mus. Pub. No. 71, zool.
ser., 3 (no. 8): 141-149, February, 1903.
Goldman, E. A.
1911. Revision of the spiny pocket mice (genera Heteromys and Liomys)
N. Amer. Fauna, 34:1-70, 3 pis. 6 figs, in text, September 7, 1911. .
1938. List of the gray foxes of Mexico. Jour. Washington Acad. Sci.,
28:494-498, November 15, 1938.
1942. Notes on the coatis of the Mexican mainland. Proc. Biol. Soc.
Washington, 55:79-82, June 25, 1942.
Hall, E. R.
1948. Two new meadow mice from Michoacan, Mexico. Univ. Kansas
Publ. Mus. Nat. Hist., 1 :423-427, 6 figs, in text, December 24, 1948.
1949. A new subspecies of cotton rat, Sigmodon hispidus, from Michoacan.
Mexico. Proc. Biol. Soc. Washington, 62:149-150, 3 figs, in text, Au-
gust, 23, 1949.
Hall, E. R., and Villa-R., B.
1948. A new pocket gopher (Thomomys) and a new spiny pocket mouse
(Liomys) from Michoacan, Mexico. Univ. Kansas Publ., Mus. Nat.
Hist., 1:249-255, 6 figs, in text, July 26, 1948.
1949. A new harvest mouse from Michoacan, Mexico. Proc. Biol. Soc.
Washington, 62:163-164, August 23, 1949.
472 University of Kansas Publs., Mus. Nat. Hist.
Hooper, E. T.
1946. Two genera of pocket gophers should be congeneric. Jour. Mamm.,
27:397-399, November 25, 1946.
Howell, A. H.
1906. Revision of the skunks of the genus Spilogale. N. Amer. Fauna,
26:1-55, 10 pis., November 24, 1906.
1914. Revision of the American harvest mice (Genus Reithrodontomys) .
N. Amer. Fauna, 36:1-97, 7 pis., 6 figs, in text, June 5, 1914.
1938. Revision of the North American ground squirrels, with a classifica-
tion of North American Sciuridae. N. Amer. Fauna, 56:1-256, 32
pis. (some colored), 20 figs, in text, May 18, 1938.
Jackson, H. H. T.
1928. A taxonomic revision of the American long-tailed shrews. . . . N.
Amer. Fauna, 51:vi + 238, 13 pis., 24 figs., July 24, 1928.
Martinez, L, and Villa-R., B.
1940. Segunda contribucion al conocimiento de los murcielagos Mexica-
nos— II Estado de Guerrero. Anales d. Inst. Biol., (Univ.) Mexico.
11:291-361, illustrated, 1940.
Miller, G. S., Jr.
1897. Revision of the North American bats of the family Vespertilionidae.
N. Amer. Fauna, 13:1-140, 3 pis., 40 figs, in text, October 16, 1897.
Miller, G. S., Jr., and Allen, G. M.
1928. The American bats of the genera Myotis and Pizonyx. U. S. Nat.
Mus. Bull., 144:viii + 218, 1 pi., 1 fig., 13 maps, May 25, 1928.
Nelson, E. W.
1899. Revision of the squirrels of Mexico and Central America. Proc.
Washington Acad. Sci., 1:15-110, 2 pis., May 9, 1899.
1909. The rabbits of North America. N. Amer. Fauna, 29:1-314, 13 pis.,
19 figs, in text, August 31, 1909.
Osgood, W. H.
1909. Revision of the mice of the American genus Peromyscus. N. Amer.
Fauna, 28:1-285, 8 pis., 12 figs., April 17, 1909.
1945. Two new rodents from Mexico. Jour. Mamm., 26:299-301, Novem-
ber 14, 1945.
Tate, G. H. H.
1933. A systematic revision of the marsupial genus Marmosa, with a dis-
cussion of the adaptive radiation of the murine opossums {Mar-
mosa). Bull. Amer. Mus. Nat, Hist, 66:1-250, pis. 1-26, 29 figs, in
text, August 10, 1933.
Transmitted August 30, 1948.
22-6113
15. A new hylid frog from eastern Mexico. . By Edward H. Taylor.
Pp. 257-264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Oklahoma.
By Edwin C. Galbrcath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kan-
sas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure
in text. August 16, 1948.
18. A new species of heteromyid rodent from the Middle Oligocene of
northeastern Colorado with remarks on the skull. By Edwin C.
Galbreath. Pp. 285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (Genus Proechimys, Family
Echimyidae). By Joao Moojen. Pp. 301-406, 140 figures in text.
December 10, 1948.
20. Three new beavers from Utah. By Stephen D. Durrant. Pp. 407-
417, 7 figures in text. December 24, 1948.
21. Two new meadow mice from Michoacan, Mexico. By E. Raymond
Hall. Pp. 423-427, 6 figures in text. December 24, 1948.
22. An annotated check list of the mammals of Michoacan, Mexico.
By E. Raymond Hall and Bernardo Villa R. Pp. 431-472, 2 plates,
1 figure in text. December 27, 1949.
Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
s
NA-L.
Subspeciation in the Kangaroo Rat,
Dipodomys ordii
BY
HENRY W. SETZER
„Z00L
LIBRAE
HAR -8 I9:38
University of J£ansas Publications
Museum of Natural History
Volume 1, No. 23, pp. 473-573, 27 figures in text, 7 tables
December 27, 1949
University of Kansas
LAWRENCE
1949
UNIVERSITY OF KANSAS PUBLICATIONS
The University of Kansas Publications, Museum of Natural His-
tory, are offered in exchange for the publications of learned societies
and institutions, universities and libraries. For exchanges and in-
formation, address the Exchange Desk, University of Kansas Li-
brary, Lawrence, Kansas, U. S. A.
Museum op Natural History. — E. Raymond Hall, Chairman, Editorial Com-
mittee.
This series contains contributions from the Museum of Natural History.
Cited as Univ. Kans. Publ., Mus. Nat. Hist.
Vol. 1. 1. The pocket gophers (genus Thomomys) of Utah. By Stephen D.
Durrant. Pp. 1-82, 1 figure in text. August 15, 1946.
2. The systematic status of Eumeces pluvialis Cope, and noteworthy
records of other amphibians and reptiles from Kansas and Okla-
homa. By Hobart M. Smith. Pp. 85-89. August 15, 1946.
3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp.
93-96, 1 figure in text. August 15, 1946.
4. Hybridization between two species of garter snakes. By Hobart M.
Smith. Pp. 97-100. August 15, 1946.
5. Selected records of reptiles and amphibians from Kansas. By John
Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.
6. Kyphosis and other variations in soft-shelled turtles. By Hobart
M. Smith. Pp. 117-124. July 7, 1947.
7. Natural history of the prairie vole (Mammalian genus Microtus).
By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6,
1947.
8. The postnatal development of two broods of great horned owls
(Bubo virginianus) . By Donald F. Hoffmeister and Henry W.
Setzer. Pp. 157-173, 5 figures in text. October 6, 1947.
9. Additions to the list of the birds of Louisiana. By George H.
Lowery, Jr. Pp. 177-192. November 7, 1947.
10. A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-
216. November 29, 1947.
11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R.
and E. Raymond Hall. Pp. 217-236, 2 figures in text. November
29, 1947.
12. A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest
and E. Raymond Hall. Pp. 237-244, 6 figures in text. December
10, 1947.
13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By
Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure
in text. December 10, 1947.
14. A new pocket gopher (Thomomys) and a new spiny pocket mouse
(Liomys) from Michoacan, Mexico. By E. Raymond Hall and
Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948.
(Continued on inside of back cover.)
Subspeciation in the Kangaroo Rat,
Dipodomys ordii
BY
HENRY W. SETZER
University of Kansas Publications
Museum of Natural History
Volume 1, No. 23, pp. 473-573, 27 figures in text, 7 tables
December 27, 1949
M$. CBKP. ZQOL
LIBRARY
MAR -8 1950
UNIVERSITY ,
University of Kansas
LAWRENCE
1949
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Edward H. Taylor, Robert W. Wilson
Volume 1, No. 23, pp. 473-573, 27 figures in text, 7 tables
December 27, 1949
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA, KANSAS
1949
22-6114
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HAR -8 19! s
Subspeciation in the Kapgaroo Rat,
Dipodomys ordii
By
HENRY W. SETZER
CONTENTS
PAGE
Introduction 477
Methods and Acknowledgments 478
Paleontology 480
Phylogeny of the Species of the Genus 484
Dispersal of the Several Species 498
Subspeciation 499
Accounts of Subspecies 511
Dipodomys ordii 511
Dipodomys ordii richardsoni 511
Dipodomys ordii oklahomae 514
Dipodomys ordii compactus 515
Dipodomys ordii sennetti 517
Dipodomys ordii evexus 518
Dipodomys ordii medius 519
Dipodomys ordii obscurus 521
Dipodomys ordii terrosus 523
Dipodomys ordii jremonti 524
Dipodomys ordii uintensis 525
Dipodomys ordii sanrafaeli 526
Dipodomys ordii panguitchensis 527
Dipodomys ordii monoensis 528
Dipodomys ordii ordii 530
Dipodomys ordii luteolus 533
Dipodomys ordii extractus 534
Dipodomys ordii chapmani 536
Dipodomys ordii montanus 538
Dipodomys ordii cinderensis 540
Dipodomys ordii jetosus 541
Dipodomys ordii utahensis 543
Dipodomys ordii columbianus 544
Dipodomys ordii idoneus 546
Dipodomys ordii priscus 547
Dipodomys ordii celeripes 549
Dipodomys ordii cineraceus 550
Dipodomys ordii marshalli 551
Dipodomys ordii inaquosus 552
(475)
476 Contents — Concluded
PAGE
Dipodomys ordii attenuatus 553
Dipodomys ordii juscus 555
Dipodomys ordii longipes 556
Dipodomys ordii pallidtis 558
Dipodomys ordii nexilis 559
Dipodomys ordii cupidineus 561
Dipodomys ordii palmeri 562
Conclusions 563
Tables of Measurements , 565
Literature Cited 571
Setzer: Subspeciation in Kangaroo Rat 477
INTRODUCTION
The geographic range of the kangaroo rats, genus Dipodomys,
extends from southern Canada south to the southern limits of the
Mexican Tableland and from the Pacific Coast east to the eastern
limits of the Great Plains in Kansas, Oklahoma and Nebraska.
These animals are usually restricted to sandy soils in semiarid re-
gions. The species Dipodomys ordii, with which this account is
primarily concerned, is, to the best of my knowledge, almost exclu-
sively confined to sandy areas.
Since 1841, when Gray gave the generic name Dipodomys to the
kangaroo rats, basing the name on the four-toed species Dipodomys
phillipsi, several other generic names have been applied. Fitzinger,
in 1867, used the name Perodipus for those animals with five toes
on the hind foot, designating Dipodomys agilis as the type of his
genus. In 1890, Merriam proposed the generic name Dipodops
with Dipodomys agilis as the type, apparently being unaware of
Fitzinger's name, Perodipus. Trouessart, in 1897, through what was
an apparent lapsus calami, applied the generic name Cricetodipus
Peale to all of the species of the then known genera Perodipus and
Dipodomys, but Trouessart later, 1904 or 1905, in his Supplementum,
corrected this lapsus and used the names Dipodomys and Perodipus.
Grinnell (1919:203) showed that some of the four-toed Dipodomys
had five toes on one hind foot and four on the other and that Perod-
ipus must fall as a synonym of the earlier generic name Dipodomys
which was to be applied to all of the kangaroo rats.
Dipodomys ordii was named by Woodhouse in 1853, from speci-
mens from El Paso, Texas, but between that time and 1919 the
name ordii was used in combination with all of the generic names
mentioned above (see synonymies under the accounts of the sub-
species).
The nearest approach to a revision of the genus was Grinnell's
(1922) "A Geographical Study of the Kangaroo Rats of California."
Since that time, Hall and Dale (1939) revised the D. microps group
and Durrant and Setzer (1945) reported upon the kangaroo rats
of Utah. The present paper is a review of the species Dipodomys
ordii. Some of the objectives in this review have been to learn:
(1) What kinds of kangaroo rats are subspecies of the species
Dipodomys ordii; (2) the limits of geographic range of this full
species; (3) the extremes of color, and of size and shape of the skull
478 University of Kansas Ptjbls., Mus. Nat. Hist.
in this one species; (4) the significance of different sizes, shapes
and colors; (5) the reasons for the existence, or formation, of se-
lected subspecies; and (6) the relationships of Dipodomys ordii to
other species in the genus.
METHODS AND ACKNOWLEDGMENTS
Available specimens were arranged according to geographic origin.
These were segregated as to sex and then under each sex by age.
Individual variation was next measured in each of several samples
in which individuals were of like geographic origin, sex, age and
season. Finally, comparable materials were arranged geographi-
cally for detection of variations of systematic worth. Following
preliminary studies of material thus arranged, additional specimens
were collected from critical areas.
When fully adult animals (see next paragraph) were segregated
as to sex, and then measured, the degree of secondary sexual varia-
tion was found to be less than the degree of individual variation;
therefore in the tables of indices, no distinction as to sex has been
made.
The only external measurements of the animals used were those
recorded by the collectors on the labels attached to the skins. These
measurements were total length, length of tail and length of hind
foot. Measurements of the ear have not been used since they were
not in all instances recorded by collectors and since measurements
of dry ears proved to be unsatisfactory. Only measurements of
fully adult specimens have been used. The term fully adult is
applied only to those specimens in which the auditory bulla is
shiny and translucent, the permanent P4 is fully erupted and
worn, and the tail is fully striped and penicillate. No one of these
characters alone was accepted as proof of adulthood but only the
three in combination.
The following measurements of the skull have been used in the
tables:
Greatest length. — From the most anterior tip of the nasals to the most
posterior projection of the auditory bullae.
Greatest breadth across bullae. — From the most lateral projection of the
auditory bulla on one side to the corresponding position on the other bulla.
Breadth across maxillary arches. — Greatest breadth across arches in a
plane perpendicular to the long axis of the skull.
Width of rostrum. — Width of the premaxillae and the nasals taken imme-
diately anterior to the upper incisors (not greatest width of nasals which is
attained farther anteriorly).
Setzer: Subspeciation in Kangaroo Rat 479
Length of nasals. — Maximum length of a nasal bone.
Least inter orbital breadth. — Least width between the orbits immediately
posterior to the lacrimal processes.
Basilar length. — From the anterior margin of the foramen magnum to the
posterior border of the alveolus of one of the upper incisors.
Capitalized color terms are from Ridgway, "Color Standards and Color No-
menclature," Washington, D. C, 1912. Color determinations were made by
comparing a masked area of pure color on the side of the animal with a
masked rectangle of named color on Ridgway's plates in natural light always
from the same angle.
Abbreviations used for specimens examined from the various collections are
as follows:
AMNH — American Museum of Natural History.
BYU — Brigham Young University.
CNHM— Chicago Natural History Museum.
CM — Carnegie Museum.
CMNH— Colorado Museum of Natural History.
DJC — Dixie Junior College.
DRD— Donald R. Dickey Collection.
KU — Museum of Natural History, University of Kansas.
LACM — Los Angeles County Museum.
MHS— Collection of Myron H. Swenk.
MVZ — Museum of Vertebrate Zoology, University of California.
OU — Museum of Zoology, University of Oklahoma.
RH — Collection of Ross Hardy.
UM — Museum of Zoology, University of Michigan.
UN — Museum of Natural History, University of Nebraska.
USAC— Utah State Agricultural College.
USES — United States Biological Surveys Collection.
USNM— United States National Museum.
UU — Museum of Zoology, University of Utah.
TCWC— Texas Cooperative Wildlife Collection.
This study is based on 3,732 specimens which were assembled at
the Museum of Natural History, University of Kansas, or studied
at other institutions. For the loan of this material and for the op-
portunity afforded for its study, I am extremely grateful to the
authorities of each of these institutions and to the owners of the
private collections.
Acknowledgement is made to the Office of Research and Inven-
tions of the United States Navy for assistance with the field work
which permitted the acquisition of essential specimens from several
of the critical geographic areas while the author was research assist-
ant on a larger over-all project (N6 ori-164-T02) of which the
determination of the geographic range of this rodent species, a po-
tential host of tularemia, was one facet. Tularemia was not de-
tected in this genus.
480 University of Kansas Publs., Mus. Nat. Hist.
I extend my thanks also to Professor Stephen D. Durrant, of the
University of Utah, for helpful corrections in the preparation of the
manuscript; to Mrs. Virginia Cassell Unruh for the preparation of
the drawings; to Professor E. Raymond Hall, of the University of
Kansas, for guidance in the study and critical assistance with the
manuscript; to Professors H. H. Lane and Worthie H. Horr for
valued suggestions; to Mr. J. R. Alcorn for providing specimens for
dissection when he was working under the University of Kansas
endowment fund; and to the other friends and associates who have
given of their time and criticism.
PALEONTOLOGY
The family Heteromyidae was defined by Wood (1935:81) essen-
tially as follows: Cheek teeth brachydont to hypsodont and even
rootless; usually six cusps per molar, three on each loph; enamel
rarely divided into two plates, never reduced to one; skull light,
thin and papery; mastoids inflated, mastoidal breadth often great-
est, never appreciably less than zygomatic breadth; interorbital
space wider than rostrum; palate nearly horizontal and little if any
below level of zygomata; nasals extended beyond incisors; zygo-
mata slender, with greatly reduced malar, almost, or quite, abutting
against tympanic; frontals and parietals broad, with latter reach-
ing, or nearly reaching, orbits; frontal trapezoidal; parietal quad-
rate, to pentagonal or triangular; interparietal primitively large,
secondarily reduced; squamosal mostly or entirely confined to
orbit; tympanic vesicular and inflated, in some forms highly in-
flated; mastoids inflated and bullous, reaching top of skull, and
forming part of occipital surface; occipitals contracted and lim-
ited in area on occiput, but extending onto dorsum of skull; coro-
noid processes small, inclined caudad and lying below level of con-
dyle; jaw small and weak with large, everted angle; tail as long as,
or longer than, head and body; claws of manus elongate, fossorial,
but forelimb slender; pelage usually coarse and frequently spinose;
ears and eyes large; body murine in form; locomotion in many
forms saltatorial.
This characterization of the family includes all of the members
of the subfamilies Perognathinae, Heteromyinae and Dipodomyinae
as well as the genus Microdipodops which I am disinclined to place
with any of the three subfamilies. Apparently it is more closely
related to the subfamily Perognathinae.
The subfamily Dipodomyinae, which contains the genera Dipod-
omys, Prodipodomys and Cupidinimus, might be characterized
Setzer: Subspeciation in Kangaroo Rat 481
after Coues' (1875) original description of the subfamily as follows:
Cheek teeth progressively hypsodont, in Dipodomys becoming ever-
growing; enamel progressively interrupted, eventually reduced to
anterior and posterior plates; upper and lower third molars reduced
in size; tooth pattern rapidly destroyed, leaving only an enamel
oval; upper incisors smooth (some fossils) or grooved (living forms) ;
progressive expansion of the auditory bullae and increase in sal-
tatorial ability; pterygoid fossa double; calcaneal-navicular or even
calcaneal-cuneif orm articulation ; tail tufted.
Owing to the fact that so little paleontological material is known
and because even that is fragile and not easily accessible for study,
knowledge of the fossil representatives has been drawn primarily
from the literature, especially from Wood's (1935) account.
Heteromyids are known from the Chadron formation, of early
Oligocene age, in which a single tooth was found. In the Orellan
stage of the mid-Oligocene where the genus Heliscomys occurs, it is
notably generalized, in comparison with other members of the fam-
ily, but it may not be ancestral at all. The lower premolar is tri-
cuspidate and the first and second molars are quadritubercular with
a broad cingulum. The teeth are bunodont and brachydont, with
the cusps not uniting to form lophs. Wood (1935:78) shows
Mookomys formicorum (from the Arikeean) as the next hetero-
myid in the evolutionary sequence and postulates that this species
arose from Heliscomys gregoryi. Mookomys is judged by Wood
to be the common ancestral form of the perognathines and the
dipodomyines.
Cupidinimus, the genus next in line, is characterized by smooth
upper incisors; lower molars with incipient H-pattern; cheek teeth
progressively hypsodont and lophate (but always rooted) ; and cal-
caneal-navicular articulation.
The time range of this genus is from the late Miocene (Niobrara
River, Local Fauna) of Nebraska to the medial Pliocene, Thousand
Creek (Hemphillian) of Nevada.
Hibbard (1937:462) described Dipodomys kansensis from the
Ogallala formation (Hemphillian age) of Kansas. He redescribed his
species, and made it the type of the new genus Prodipodomys (Hib-
bard, 1939:458), differentiating it from Dipodomys on the basis
of the three-rooted p4, double-rooted ml and m2 and the single
rooted m3. It is shown to be closely allied to Dipodomys by the
form and position of a large foramen posterior and labial to m3, and
by the development of the masseteric ridge.
482 University of Kansas Publs., Mus. Nat. Hist.
The next youngest heteromyid fossils which have been described
are of the genus Prodipodomys? from Arizona. Gidley (1922:123)
described Dipodomys minor from the Benson (Blancan) which
Gazin (1942:486) refers to the genus Prodipodomysl. Wood (1935:
156) described Dipodomys gidleyi from the Curtis (Pleistocene).
Both of these species are primitive as regards dentition; that is to
say, the enamel ring of the tooth is complete and lacks any sign of
a break. The limb bones of D. gidleyi show lesser saltatorial abil-
ity, and therefore appear to be more primitive, than those of any
living Dipodomys.
Several heteromyids which have not been assigned to any genus
are known. Wilson (1939:36-37) recognized some from the Ava-
watz (Clarendonian) and the Ricardo (Clarendonian) . Another,
possibly of the genus Diprionomysl , from the Barstow (Barstovian)
was described by Wood (1935:197) as follows: "The general shape
of the tooth as figured strongly suggests either one of the most ad-
vanced species of Dipodomys or else a Geomyid. ... It is
much more advanced than are any known contemporary heteromy-
ids, and compares fairly well with such late Tertiary and Pleistocene
geomyids as have been described. It certainly is not referable to any
known heteromyid genus other than Dipodomys, and should proba-
bly be called a Geomyid." Wilson (loc. cit.) refers to these specimens
as Dipodomyine (?) n. gen. and sp. If these specimens referred to
by Wood and Wilson are true heteromyids then a change in the
phylogenetic scheme proposed by Wood (1935) would be necessary.
Wilson (loc. cit.) says, referring to the Avawatz specimen, "The
cheek teeth are very hypsodont but are apparently not persistent in
growth, . . . Wide enamel breaks are present in M/l dividing
the enamel into anterior and posterior bands. The enamel of P/4 is
complete in the present stage of wear, but an examination of the
tooth indicates that breaks would develop with additional attrition
at the buccal and lingual margins of the metalophid, and at the buc-
cal border of the protolophid. The incisor is of the slender heter-
omyid type."
Wood (1935:118) in referring to the ancestry of Cupidinimus
with regard to the grooving of the incisors says: "The philosophy
of evolution which would prohibit its derivation from Mookomys,
because of the grooved incisors in the latter genus, would require a
separate line leading back at least to the Lower Miocene."
In view of the above statements, it is conceivable that additional
material will be found carrying the dipodomyine line back into the
Setzer: Subspeciation in Kangaroo Rat
483
early Miocene. Perhaps the line involving Mookomys and Cupidini-
mus which was regarded by Wood as the line of descent, is merely
an aberrant side branch that parallels in its structures the main
line of evolution of the dipodomyines (Figure 1).
Recent
Pleistocene
Dipodomys agilis
Dipodomys gidleyi
c
o
o
a.
Blanco n
Prodipodomys? minor
Hemphilhon
— ,> Cupidinimus magnus
Prodipodomys konsensis y^
Clarendonian
Avawatz specimen
V _
o>
c
<D
O
O
s
Barstovion
Cupidinimus nebroskensis /
*< \ /
Hemingfordion
^ \ /
Arikoreean
Mookomys formicorum
c
O
O
5
/
Orellan
Heliscomys gregoryi
Chadronian
■
Fig. 1. Phylogeny of the Dipodomyines (modified after Wood, 1935).
As Wilson (1939:37) says: "Indeed it is hard to recognize such
a form as Cupidinimus nekraskensis as directly ancestral to Di-
podomys in view of the occurrence of the much more advanced
Avawatz specimen in deposits that are at most only slightly later
than those in which the former is found. The kangaroo rats were
apparently much farther along in their development by lower
Pliocene time than heretofore supposed."
Wood (1935:78) suggested that Dipodomys gidleyi gave rise to
Dipodomys spectabilis and Dipodomys ordii, and Dipodomys minor
gave rise to Dipodomys compactus. However, my own study indi-
cates that Dipodomys compactus is conspecific with Dipodomys
ordii and should stand as Dipodomys ordii compactus. Consequently
a different phyletic arrangement than that proposed by Wood (loc.
484 University of Kansas Publs., Mus. Nat. Hist.
tit.) is required. Since D. compactus is more closely allied to
Prodipodomys? minor than D. ordii is to D. gidleyi, it is possible
that P.? minor gave rise to D. ordii and that D. spectabilis is the
end product of the phyletic trend of D. gidleyi (Figure 1).
The trend of phyletic development in the dipodomyines has been
toward the saltatorial habit. To acquire this habit from a scamper-
ing ancestor, certain morphological modifications were necessary.
Among these modifications were a lengthening of the tail, a length-
ening of the hind legs, the development of a calcaneal-navicular-
ectocuneiform contact instead of a calcaneal-navicular contact for
additional strength in leaping, a shortening of the forelimb, an
increase in size and inflation of the mastoid and tympanic portions
of the skull with a consequent reduction in size of the interparietal
region and the fusion of certain of the cervical vertebrae. Late
Miocene (Cupidinimus) and Pliocene (Avawatz specimen and
Prodipodomys) forms had acquired certain of these morphological
modifications that are present in the modern genus Dipodomys.
PHYLOGENY OF THE SPECIES OF THE GENUS
Representatives of nine species of Dipodomys were dissected in
an attempt to determine the degree of specialization and the relative
systematic position of each species.
The myology was found to agree in detail as to origin, insertion
and innervation with that of Dipodomys spectabilis as reported by
Howell (1932). The only variation noted in the muscular system
was the size of the individual muscles in those animals of widely
divergent body size.
Dipodomys ordii is the most generalized and Dipodomys deserti
is the most specialized of the kangaroo rats (see Table 1), as judged
by the osteology. Information gained by the study of the viscera
of the various species supports this judgment. The visceral mass is
relatively loose in D. ordii, but is markedly compact in D. deserti.
This compactness appears to be brought about by the foreshorten-
ing of the mesenteries which support the entire gut and by the closer
apposition of the large intestine to the caecum; both the intestine
and caecum occupy a ventral position in the abdominal cavity. In
Dipodomys ordii the entire visceral mass is loosely interconnected
and the caecum is relatively small as compared to the tightly com-
pact viscera and the large caecum in Dipodomys deserti. Another
striking feature is the size, proportion and position of the liver. In
all animals dissected, even in D. deserti, the right lobe of the liver
Setzer: Subspeciation in Kangaroo Rat 485
descends and forms a capsule around the anterior end of the right
kidney. In the Ord kangaroo rat, the bulk of the liver lies on the
right side of the body cavity. That is to say, there is a greater bulk
of the liver on the right side and it is situated more dorsad than in
any of the other species examined. In the most specialized condi-
tion, as in Dipodomys deserti, the bulk of the liver is almost equal
on the right and left sides, and instead of having the greater bulk
situated dorsally as in D. ordii it is cup-shaped, with the dorsal and
ventral parts of approximately equal size and situated on almost
the same transverse plane. The entire mass of the liver is concave
posteriorly.
TABLE 1
Skeletal Indices of Dipodomys
a
3
.2 oa
a s § | Is S
a £ 6 h £ o
ordii 144.5 57.2 127.75 60.55 88.4 63.4
microps 138.5 56.17 132.3 57.27 90.95 60.8
panamintinus...' 146.1 55.3 132.0 57.5 90.5 60.8
agilis 147.0 55.05 133.65 57.25 94.55 62.65
heermanni 142.9 54.2 135.9 55.35 92.2 60.93
ingens 142.9 54.1 130.6 56.2 89.65 66.2
spectabilis 140.9 53.05 133.9 54.2 95.6 64.6
phillipsii 163.4 55.05 137.85 58.97 101.5 64.5
merriami 160.75 53.85 137.5 57.35 99.75 63.9
nitratoides 155.0 54.1 137.4 57.0 98.25 65.5
deserti 149.5 53.4 139.4 54.9 96.6 67.6
The right kidney is variable in position in reference to the left.
In all species the right kidney lies anterior to the left but in some,
D. deserti and D. ingens, it is markedly anterior.
In Dipodomys agilis, D. merriami and D. deserti there are small
to large patches of lymphoid tissue on the caecum. These patches
were not noted in any of the other species examined and I do not
know their function. In the three above mentioned species, however,
486 University of Kansas Publs., Mus. Nat. Hist.
the large intestine is shorter in proportion to the small intestine
than in any other species except D. heermanni (see Table 2) and
with the exception of D. heermanni, D. venustus and D. ordiv the
actual measurements are less.
Inasmuch as little is known of the food habits of the various
species of kangaroo rats, any ascription of adaptive significance
to the varying proportions of the digestive system would be only
speculative.
Midgley (1938) describes the visceral anatomy of D. ordii and
D. microps. Except for the differences here noted the description
of the viscera as given by Midgley (loc. cit.) applies to the rest of
the species studied.
TABLE 2
Visceral Measurements (in Millimeters) of Dipodomts
■— f .5
e C M t- .^ V
X C a "O o
5
-
Large intestine. . . .
432
290
464
397
237
413
374
419
430
Small intestine ....
165
126
220
195
131
228
207
255
274
Percent of small to
large intestine. .
38.2
43.4
47.5
49.2
55.2
55.3
55.4
60.9
63.7
From the differences noted in the skeleton, in the entire visceral
mass, and in the shape and position of the liver it appears that as
a saltator becomes more specialized skeletally, there is a concur-
rent compacting and aligning of the viscera into a more or less
bilaterally balanced mass. It seems that this alignment is for a
stabilization in leaping. It seems reasonable that the individual
that has a loose and unconsolidated visceral mass, or in which the
viscera or a least the heaviest part of the viscera is relatively uni-
lateral, would be thrown slightly off balance at the end of the jump.
This would place the animal at a slight disadvantage before being
able to make the next jump. Howell (1944:40) comments on the
fact that kangaroo rats often land off balance, "owing apparently
to clumsy use of the tail." Possibly the unilaterality of the vis-
ceral mass plus a shorter tail and a more clumsy use of that organ
accounts for the off balance landings which Howell has observed.
The skeleton, particularly of the appendages, shows the most
Setzer: Subspeciation in Kangaroo Rat 487
modification, ranging from a relatively generalized to a specialized
condition. Skeletal indices, as established by Howell (1944:199)
have been used in estimating the amount of such specialization.
These indices are obtained by dividing the length of one segment
of a limb by the length of another segment and are expressed in
percentages. The Femorotarsalmetatarsal and Cranial indices are
not from Howell (loc. cit.).
The Humeroradial index (radius/humerus X 100) in the gen-
eralized animal is theoretically 100 because the humerus and radius
are of the same length. In kangaroo rats, which are saltators, the
index rises to more than 100 owing to the lengthening of the radial
component.
The Intermembral index (humerus and radius/femur and tibia
X 100) in a generalized animal is theoretically 100, but, as Howell
(1944:205) points out, the index in generalized mammals is prob-
ably nearer 75. If the hind leg elongates at the expense of the fore-
limb the animal will be a better saltator and the skeletal elements
will yield a lower intermembral index.
The Femorotibial or Crural index (tibia/femur X 100) expresses
the development of the tibia as an adaptation to the saltatorial
habit and in generalized animals would be expected to be 100. As
an adaptation to saltation the tibia would elongate at the expense
of the femur and the index would be more than 100. The degree
of divergence from 100 would be an expression of the degree of
saltatorial ability.
The Tibioradial index (radius/tibia X 100) in the generalized
animal also would be expected to approximate 100 but it is doubtful
if any living mammals, except brachiating kinds, yield an index of
more than 75. In saltators, the index is low because of the elonga-
tion of the hind appendages, whereas the forelimbs do not change
their length or are shortened.
The Femorotarsalmetatarsal index (tarsometatarus/femur X 100)
in the generalized condition would be less than 50 and an index
approaching 100 would indicate a specialization for saltation owing
to the elongation of the tarsometatarsal elements.
The Cranial index (breadth across bullae/length of skull X 100)
reflects the development of the auditory or mastoid region of the
skull as an adaptation for more acute hearing and possibly for more
delicate balance. In heteromyids, the generalized condition would
be represented by an index of 50 or less, and as the width across
the bullae increases, the index rises toward 100.
488 University of Kansas Publs., Mus. Nat. Hist.
Setzer: Subspeciation in Kangaroo Rat 489
Figs. 2-10. Showing the compacting of the visceral mass; liver at the top,
small intestine and caecum at the bottom. All figures approximately X 1-
Fig. 2. Dipodomys ordii inaquosus, $ , adult, no. 23365, KU; 7 mi.
W Fallon, Churchill County, Nevada; trapped 27 October, 1945.
Fig. 3. Dipodomys panamintinus mohavensis, $ , adult, no. 22094,
KU; IY2 mi. N Mojave, Kern County, California; 3 February, 1948
Fig. 4. Dipodomys heermanni morroensis, $ , adult, no. 22082,
KU; S side Mono Bay, 4 mi. S Morro, San Luis Obispo County, Cali-
fornia; 25 January, 1948.
Fig. 5. Dipodomys ingens, 2, adult, no. 22069, KU; 25 mi. SW
Mendota, San Benito County, California; 2 February, 1948.
Fig. 6. Dipodomys agilis perplexus, 2, adult, no. 22091, KU; l%o
mi. N Monolith, Kern County, California; 3 February, 1948.
Fig. 7. Dipodomys venustus sanctiluciae, $ , adult, no. 22071, KU;
V/2 mi. S Jolon, Monterey County, California; 26 January, 1948.
Fig. 8. Dipodomys spectabilis spectabilis, 2, adult, no. 22110, KU;
5 mi. NE Willcox, Cochise County, Arizona; 19 January, 1948.
Fig. 9. Dipodomys merriami merriami, 2, adult, no. 23366, KU;
E side Carson Lake, Churchill County, Nevada; 2 October, 1945.
Fig. 10. Dipodomys deserti deserti, $ , adult, no. 23364, KU; 15 mi.
WSW Fallon, Churchill County, Nevada; 3 November, 1945.
TABLE 3
Relative Specializations of the Species for each Index
o S 1! "S " — p
s S § J i g i S
a £ o h (S o <
ordii 5 11115 2.33
microps 1 2 4 5 4 2 3.0
panamintinus 6 3 3 3 3 1 3.1
agilis 7 4 5 6 6 4 5.3
heermanni 3 6 7 9 5 3 5.5
ingens 4 7 2 8 2 10 5.5
spectabilis 2 11 6 11 7 8 7.5
phillipsii 11 5 10 2 11 7 7.6
merriami 10 9 9 4 10 6 8.0
nitratoides 9 8 8 7 9 9 8.5
deserti 8 10 11 10 8 11 9.6
The figure 1 represents the least specialized condition for the index, while
the figure 11 represents the most specialized condition. The remainder of the
numbers indicate the relative degree of specialization of each species for each
index.
2—6114
490 University of Kansas Publs., Mus. Nat. Hist.
The species that have been examined are listed in Table 3 in
increasing order of specialization from top to bottom.
Usually animals of extreme morphological specialization are
much restricted environmentally. Attempts to correlate the relative
evolutionary position of the various species, as indicated by the
degree of specialization interpreted from the indices with that of
habitus has proven unsuccessful. For example, Dipodomys mer-
riami, which is third from the top in the list as arranged above, is
neither restricted to loose sandy soil as is D. deserti nor to brush as
are D. agilis and D. venustus. D. merriami does, nevertheless, in-
habit a variety of habitats from loose sandy soils to rather hard
rocky ground. Throughout the genus there is, however, a general
trend toward increased specialization as the animals adopt the more
open desert environment, as is indicated by the elongation of the
tail and hind appendage and increase in size of the auditory region
of the skull. A marked difference is noted in the size of the pinna
of the ear in the various species. Generally, those species having
small pinnae inhabit open desert country while those with large
pinnae inhabit brushy country. This is in direct contradistinction
to the hares and rabbits in which the small-eared kinds are brush
dwellers whereas the large-eared kinds are inhabitants of open
country. Three possible explanations for hares and rabbits having
this specialization of the pinnae are: (1) To enable the open-
dwelling animals with the larger pinnae to hear more readily the
approach of an enemy when it is yet far away, while the brush-
living forms, which rely for escape on a short dash into cover, do
not need so large a "funnel"; (2) large pinnae have been developed
by those animals which live in the open desert as an aid in dissi-
pating the body heat; (3) large pinnae in brush-dwelling animals
would be a decided disadvantage in rapid movement through the
brush. Grinnell (1922:20) points out that animals with large pin-
nae usually have small auditory bullae and conversely, animals
with small pinnae have large bullae. This compensatory factor,
implying an auditory function, appears to be inoperative in D.
panamintinus mohavensis which has small ears and small bullae
and in D. elephantinus which has large ears and large auditory
bullae. Grinnell (loc. tit.) suggests that several additional factors
enter into the problem, such as the amount of digging each species
must do to gain safety, the texture of the soil for burrowing, the
extent of forage area and the type of cover in connection with the
mode of attack of predators. Of these factors, perhaps the most
important are the two first mentioned.
Setzer: Subspeciation in Kangaroo Rat
491
Figs. 11-15. Ventral views of skulls showing the degree of development of
the auditory bullae and the configuration of the pterygoid fossae. All figures
approximately X 1.
Fig. 11. Dipodomys ordii compactus, $, adult, no. 646, TCWC;
19 mi. S Port Aransas, Mustang Island, Nueces County, Texas; 24
April, 1939.
Fig. 12. Dipodomys ordii oklahomae, 9, adult, no. 265456, USBS;
21/4 mi. S Norman, Cleveland County, Oklahoma; 21 March, 1934.
Fig. 13. Dipodomys ordii richardsoni, $, adult, no. 15995, KU; 1
mi. S Lamar, Prowers County, Colorado; 8 September, 1945.
Fig. 14. Dipodomys ordii nexilis, 9, adult, no. 149941, USBS; 5 mi.
W. Naturita, Montrose County, Colorado; 20 July, 1907.
Fig. 15. Dipodomys deserti deserti, 5, adult, no. 18670, KU; 14 mi.
WSW Fallon, Churchill County, Nevada; 3 November, 1945.
492 University of Kansas Publs., Mus. Nat. Hist.
Figs. 16-20. Dorsal views of skulls showing the degrees of inflation of the
auditory bullae and the correlation of large bullae with small interparietal.
All figures approximately X 1-
Fig. 16. Dipodomys ordii compactus, for data see Fig. 11.
Fig. 17. Dipodomys ordii oklahomae, for data see Fig. 12.
Fig. 18. Dipodomys ordii richardsoni, for data see Fig. 13.
Fig. 19. Dipodomys ordii nexilis, for data see Fig. 14.
Fig. 20. Dipodomys deserti deserti, for data see Fig. 15.
Setzer: Subspeciation in Kangaroo Rat 493
Wood (1935:155), on the basis of structure of the teeth, listed
the species which he examined in the following increasing order of
specialization: Dipodomys compactus (now Dipodomys ordii com-
pactus), D. nitratoides, D. merriami, D. ordii, D. agilis, D. herr-
manni, D. spectabilis, and D. deserti. This arrangement is at va-
riance with that of Grinnell (1922) who listed the species in order
of increasing specialization as: Dipodomys herrmanni, D. panamin-
tinus, D. ingens, D. spectabilis, D. merriami, D. nitratoides, D.
ordii, D. agilis, D. venustus, D. microps and D. deserti. As noted,
the only agreement between the two arrangements is the placing
of D. deserti as the most specialized. Relying on skeletal indices
alone, I would accord the same position to D. deserti but would not
arrange the other species as have either Wood or Grinnell.
In this study, the amount of specialization of each species, as
indicated by the skeleton, was determined by assigning consecutive
numbers from 1 to 11 to each species in its place in each index, and
then totaling and averaging these arbitrary numbers (Table 3). It
will be noted that there is a tendency for each species to occupy
the same relative position in each of the indices.
It is felt, however, that a more nearly correct arrangement, ac-
cording to degree of specialization, is obtained by using the six
skeletal indices plus the information obtained from the study of
the viscera. On this basis the species may be arranged from least
to most specialized as follows: Dipodomys ordii, D. microps, D.
panamintinus, D. agilis, D. herrmanni, D. ingens, D. spectabilis, D.
phillipsii, D. merriami, D. nitratoides and D. deserti.
Grinnell (1922:95-96) arranged the Recent species of Dipodomys
in nine groups. Davis (1942:332) also proposed an arrangement
of nine groups in which he combined the Compactus and Ordii
groups of Grinnell, established a new Elator group by removing
Dipodomys elator from Grinnell's Phillipsii group, and in linear
arrangement, Davis shifted the Spectabilis and what remained of
the Phillipsii groups to new positions. Burt (1936:152) arranged
Grinnell's groups into three (unnamed) groups solely on the basis
of the structure of the baculum. In the arrangement proposed by
Grinnell, two of his nine groups contained only one species each,
one other, the Microps group, has since been shown to contain only
one species and another, the Compactus group, contained only kinds
which are, by me, regarded only as subspecies of Dipodomys ordii.
To my mind neither Davis nor Burt added to or fundamentally
changed the basic concepts as set forth in 1922 by Grinnell. Owing
494 University of Kansas Publs., Mus. Nat. Hist.
to the paucity of material at that time, especially from areas of
intergradation, Grinnell's groupings and arrangement were as nearly
natural as could be expected. With the accumulation of additional
material and with the knowledge that certain kinds treated by
Grinnell as full species are in actuality subspecies, it is felt that the
several species of kangaroo rats can best be arranged in six groups
which, from the least to the most specialized, are as follows:
Ordii Group. — Composed of the subspecies of Dipodomys
ordii and Dipodomys microps. Grinnell placed these two
species in separate groups ; Burt on characters of the baculum
alone placed D. microps with Dipodomys deserti and Dipod-
omys spectabilis. Within the single species D. ordii, I find
that the difference in shape and size of the baculum between
the subspecies of D. ordii is as great as the difference which
Burt (1936:154-155) found between the full species D. agilis
and D. microps. The characters of the baculum are an aid, but
not in and of themselves an adequate basis, for determining
the natural relationships of the groups of species. Certainly
the remainder of the morphological differences between D.
deserti and D. microps are so great that I doubt that the
similarity in the baculum is signficant, at least in this one
instance. The chisel-shaped lower incisors of D. microps
appear to be a specialization. They may enable D. microps
to utilize more woody types of vegetation than can D. ordii.
Both species occupy the same territory over much of their
geographic range, probably because they eat different kinds
of food.
Panaminitinus Group. — Composed of all the now known sub-
species of Dipodomys panamintinus and the species Dipod-
omys stephensi, if the latter is a full species. This group was
included by Grinnell in the Heermanni group, with which it
agrees in broadness of the maxillary arches and the configur-
ation of the penis bone, but on the basis of the degree of spe-
cialization, as indicated by the indices (see Table 1), I feel
that the Panamintinus group is more properly placed after
the Ordii group and should be separated from the Heermanni
group. Actually, animals in the Panamintinus group are
intermediate between those of the Ordii and Heermanni
groups.
Heermanni Group. — Composed of the subspecies of Dipodomys
heermanni and Dipodomys agilis, the species Dipodomys in-
gens, Dipodomys venustus and Dipodomys elephantinus. D.
Setzer: Subspeciation in Kangaroo Rat 495
ingens even though larger in linear measurements than any
of the other kinds included in this group, has almost the
same degree of specialization as does D. heermanni. D.
agilis, even though somewhat less specialized than the other
kinds placed in this group, by the general nature of the
indices, by the form of the visceral mass and to some degree
by the shape of the baculum, shows itself properly to belong
with this group. The species D. venustus, judged by char-
acters of the visceral anatomy, also belongs here rather than
with some other group or as a separate group. From the
appearance of the visceral mass, D. venustus is somewhat
more specialized than either D. heermanni or D. agilis, but
D. venustus does show its affinities with this group. The
species D. elephantinus has not been examined as thoroughly
as have the other species but the external morphology and the
configuration of the cranium place it with this group.
Spectabilis Group. — Composed of the subspecies of Dipodomys
spectabilis. In two of the six indices, D. spectabilis shows a
high degree of specialization toward saltation, but in the
other four indices it shows a relatively low degree of speciali-
zation or is average for the genus. Burt (1936:155) placed
D. spectabilis with D. deserti on the basis of the baculum
alone. ■ I have not examined D. nelsoni but place it with this
group as did also Grinnell and Davis.
Merriami Group. — Composed of the subspecies of Dipodomys
merriami, Dipodomys nitratoides and Dipodomys phillipsii,
and the species Dipodomys platycephalus, Dipodomys mar-
garitae, Dipodomys insularis, Dipodomys mitchelli, Dipodo-
mys ornatus and Dipodomys elator. I have not examined
five of these species. However, the indices and characters of
the viscera indicate that the three species first mentioned are
closely allied. Owing to the lack of known intergradation
between the three, I judge that they should be retained as
full species, but the difference in degree of morphological
specialization is no more than would be expected between
subspecies. I have examined no specimens of Dipodomys
elator but from what I know of its morphology, I think that
Grinnell better indicated its relations in allying it with D.
phillipsii than did Davis in erecting a new group for it on
the basis of linear measurements.
Deserti Group. — Composed of Dipodomys deserti which has
only two subspecies. In all morphological respects, D. deserti
is the most specialized species in the genus as shown by the
496
University of Kansas Publs., Mus. Nat. Hist.
reduced number (4) of toes on the hind foot, the bilateral
arrangement of the viscera, the extreme development of the
auditory region of the skull and by developing, early in life,
the hiatus in the enamel wall of each molariform tooth.
The parallel arrangement below emphasizes the differences and
similarities between Grinnell's (1922) arrangement and the one pro-
posed in the present paper.
Grinnell's arrangement
Heermanni Group
Dipodomys heermanni
Dipodomys morroensis
Dipodomys mohavensis
Dipodomys leucogenys
Dipodomys panamintinus
Dipodomys stephensi
Dipodomys ingens
Spectabilis Group
Dipodomys spectabilis
Dipodomys nelsoni
Phillipsii Group
Dipodomys phillipsii
Dipodomys perotensis
Dipodomys ornatus
Dipodomys elator
Merriami Group
Dipodomys merriami
Dipodomys nitratoides
Dipodomys platycephalus
Dipodomys margaritae
Dipodomys insidaris
Dipodomys mitchelli
Ordii Group
Dipodomys ordii
Compactus Group
Dipodomys compactus
Dipodomys sennetti
Agilis Group
Dipodomys agilis
Dipodomys venustus
Dipodomys elephantinus
Microps Group
Dipodomys microps
Dipodomys levipes
Deserti Group
Dipodomys deserti
Were in Heermani Group Above
Present arrangement
Heermanni Group
Dipodomys heermanni
Dipodomys agilis
Dipodomys ingens
Dipodomys venustus
Dipodomys elephantinus
Spectabilis Group
Dipodomys spectabilis
Dipodomys nelsoni
Now in Merriami Group Below
Merriami Group
Dipodomys merriami
Dipodomys nitratoides
Dipodmnys platycephalus
Dipodomys margaritae
Dipodomys insularis
Dipodomys mitchelli
Dipodomys phillipsii
Dipodomys ornatus
Dipodomys elator
Ordii Group
Dipodomys ordii
Dipodomys microps
Now in Ordii Group Above
Now in Heermanni Group Above
Now in Ordii Group Above
Deserti Group
Dipodomys deserti
Panamintinus Group
Dipodomys panamintinus
Dipodomys stephensi
Setzer: Subspeciation in Kangaroo Rat
497
Names of the subspecies are omitted from the groups named above
and only the names of full species, as understood by Grinnell and
as understood now, have been used. It will be noted that the phylo-
genetic order follows that of Grinnell rather than the one proposed
herein.
The fossil record of the kangaroo rats is so scanty that one can
but speculate on the evolutionary sequence. Wood (1935) presented
a diagnosis of the early phyletic history up to and through Cupidini-
mus; this is probably as correct as can be made. I cannot, however,
share his view that the recent species of Dipodomys have originated
from a descendant of Cupidinimus nebraskensis ; instead, I think
that the Recent species originated from some unknown ancestor in
the southwest.
deserti
nitratoides
v mem'ami
{ phillipsii
spectabilis
ingens
heermanni
venustus
elephantinus
panpmintinus
stephensi
microps
ordii
Fig. 21. Diagrammatic representation of the relationships and history of
the Recent species Dipodomys.
In view of the foregoing evidence it seems best to estimate the
relationships and history of the various species and groups of spe-
cies only as far back as the early Pleistocene (see Figure 21) . Inas-
much as faunas of fossil mammals from the mid-Pleistocene contain
few, if any, Recent species (see Hibbard, 1937:193), the living spe-
cies of Dipodomys have probably had a geologic history no longer
than the period of time which has elapsed since the middle Pleisto-
cene, or at the earliest the early Pleistocene. Of the Recent species,
only Dipodomys agilis is known as a fossil ; it was found in the late
498 University of Kansas Publs., Mrs. Nat. Hist.
Pleistocene tar pits of California. Under the heading "Dipodomys
near ingens," however, Schultz (1938:206) recorded remains of
kangaroo rats from the tar seeps of McKittrick in the San Joaquin
Valley of California.
DISPERSAL OF THE SEVERAL SPECIES
If we assume the region of origin and center of dispersal of a
group of animals to be the one in which the greatest numbers of the
most specialized species of a given genus are found, then the northern
Tableland of Mexico and the adjoining region of the United States
in southeastern California and southwestern Nevada is the region of
origin and the center of dispersal for the genus Dipodomys. Di-
podomys deserti, Dipodomys merriami, Dipodomys panamintinus,
Dipodomys microps, Dipodomys phillipsii and Dipodomys ordii are
found in the region mentioned. That the aforementioned region may
be the center of differentiation for this genus is further indicated by:
First, the finding, in this region, of saline deposits of Cenozoic (Mio-
cene) age, indicating aridity, which is thought to have been one of
the essential stimuli for the development of the saltatorial habit in
the genus Dipodomys; second, the recovery of the advanced hete-
romyids from the Avawatz and Ricardo of the Clarendonian (Plio-
cene) of this same region; and third, the present abundance and di-
versification of kangaroo rats in this same geographic region which
has been more or less arid since Miocene time.
A secondary center of evolution has been the low, hot interior
valleys and adjacent foothills of central California where Dipodo-
mys ingens, Dipodomys heermanni, Dipodomys venustu-s, Dipodo-
mys agilis, Dipodomys elephantinus and Dipodomys nitratoides are
now found. Although there are as many species as in the principal
center of origin, the amount of specialization and adaptive radia-
tion in California is not so great. Probably during the Quaternary,
when the process of mountain building was actively under way the
animals that had reached central California from the parental cen-
ter became isolated by the emergence of the Tehachapi Mountains.
This mountain range separated the California animals from popu-
lations farther south and east. As a result, D. nitratoides was dif-
ferentiated from D. merriami, and D. heermanni underwent an evo-
lution of its own which resulted in animals having either four or five
toes on the hind foot. At the same time Dipodomys ingens de-
veloped there and has since been undergoing an evolution parallel to
that of the large-sized species, Dipodomys spectabilis. The two
species have paralleled each other not only in large size but to some
Setzer: Subspeciation in Kangaroo Rat 499
extent in habits such as building large mounds that are kept free
of vegetation and in occupying areas of rather hard clayey soil.
Structurally, however, D. ingens has not yet become quite so spe-
cialized as D. spectabilis, probably because D. ingens has had less
time in which to become so. A second species, if it is a full species,
Dipodomys elephantinus, has also been isolated in central Cali-
fornia but has not attained so high a degree of specialization as
D. ingens. It is interesting to note that in each of the two stocks,
two large-sized species have been evolved. In the parental stock
the two species are Dipodomys deserti and Dipodomys spectabilis;
the former is the most specialized species in the genus. In the stock
isolated in California, however, even though two large species have
been formed they are still below the average in degree of specializa-
tion for the genus. As noted elsewhere in this paper, the species
from these low, hot valleys, excepting D. nitratoides, are all closely
related one to another. Dipodomys venustus and Dipodomys ele-
phantinus are either closely related species or possibly only sub-
species of one species, Dipodomys agilis.
It is worthy of note that as the distance away from the center of
differentiation increases, the number of species decreases. For
example, in the northern Great Basin there are only two species
(Dipodomys .ordii and Dipodomys microps) and farther eastward,
on the eastern side of the Rocky Mountains, there is only the one
species, Dipodomys ordii. In north-central Texas, Dipodomys ela-
tor, perhaps a relict species, is found occupying an area farther east
than that occupied by Dipodomys ordii at that latitude.
Dipodomys ordii, Dipodomys phillipsii and Dipodomys merriami
occupy the southern portion of the range of the genus. Instead of
being generalized at this southern part of the periphery of the
range as are the kinds found on the other parts of the periphery of
the range of the genus, these three southern kinds are notably
specialized there in the south. The subspecies D. o. palmeri which
occurs in the area, is the most specialized of the species Dipodomys
ordii; and Dipodomys phillipsii and Dipodomys merriami stand
high in the scale of specialization with respect to the other species
of the genus. The reason for this is not clear.
SUBSPECIATION
Dipodomys ordii is, without question, a valid species if one ac-
cepts Mayr's (1942:120) definition that "Species are groups of
actually or potentially interbreeding natural populations, which are
reproductively isolated from other such groups." D. ordii is not
500 University of Kansas Publs., Mus. Nat. Hist.
known to hybridize with other species where their geographic
ranges are adjacent or overlap. The first part of the definition "ac-
tually or potentially interbreeding populations" is substantiated by
the 35 recognizable subspecies which can be defined as "a complex
of interbreeding and completely fertile individuals which are mor-
phologically identical or vary only within the limits of individual,
ecological and seasonal variability. The typical characters of this
group of individuals are genetically fixed and no other geographic
race of the same species occurs within the same range" (after
Rensch, 1934; from Mayr, 1942:106). Thus we find that certain
populations of individuals differ from others and that in geographic
areas between two of these populations, individuals (intergrades)
are found which resemble those of both populations. In another in-
stance, a population may be geographically isolated yet in its
characters it may be recognizable as a subspecies without actual
intergradation because of slight degrees of difference, or a group
may be different from another without being geographically sep-
arated and may or may not show intergradation.
Subspeciation in Dipodomys ordii almost certainly has been ef-
fected, by means of mutations, under the influence of natural selec-
tion. Natural selection enhanced by geographic and e c o 1 o g i c
isolation, probably has retained mutations of evolutionary signifi-
cance, thus permitting the development of the many recognizable
subspecies.
In the subspecies of Dipodomys ordii the color ranges from pale
to dark. The difference in color is as pronounced as that between
the full species D. deserti and D. heermanni. The lightest-colored
subspecies are Dipodomys ordii celeripes, D. o. extractus and D. o.
compactus; the darkest are D. o. obscurus and D. o. palmeri.
There is a marked tendency for intergrades between a light-
colored subspecies, such as D. o. celeripes, and a dark-colored kind,
such as D. o. utahensis, to show varying degrees of blending in
color. The insular population, D. o. compactus, has, however, two
distinct color phases, a light phase and a slightly darker phase, and
shows no tendency toward blending. In other kinds of mammals,
blending of color is known to be the result of the action of multiple
alleles, but in the insular kangaroo rat (D. o. compactus) the color
appears to be the result of either a reduced multiple allelomorphic
complex or even a unit factor. The two color phases of this insular
subspecies, which might be an expression of a unit factor, more
probably are specializations in which the multiple alleles for color
Setzer: Subspeciation in Kangaroo Rat 501
have been reduced. The probability that there is either a unit factor
or a reduced number of alleles at work is suggested by the taking
of more dark-colored than light-colored animals and by the absence
of blending of color. This insular population has undoubtedly been
derived from the mainland kangaroo rat, D. o. sennetti, which has
the usual range of variation but, to my knowledge, there are no in-
dividuals of D. o. sennetti so light as the darkest animals of D. o.
co?npactus from the islands.
Populations from a given locality are remarkably stable in color
except the animals from Samalayuca, Chihuahua, which vary in
color from individuals almost as light as D. o. compactus to ani-
mals that approach D. o. ordii in darkness of pelage.
The subspecies of D. ordii show no noticeable variation in the
extent of the hip stripe, supraorbital and postauricular spots, basal
white ring of the tail, lateral stripes of the tail or the extent of white
on the venter and feet. There is, however, variation in the degree
and extent of the arietiform facial markings. In Dipodomys ordii
utahensis, D. o. cupidineus, D. o. obscurus and D. o. fuscus these
markings are pronounced. In D. o. celeripes, D. o. pallidus, D. o.
compactus and D. o. attenuatus these markings are either obliter-
ated or nearly so.
In Dipodomys ordii, color does not seem to be correlated with
amount of moisture or geography, but rather with color of soil. For
example, all animals from the Bonneville Basin of western Utah,
are light colored as are the soils; animals from the San Rafael Des-
ert of eastern Utah are reddish, as is the soil. More striking ex-
tremes of this are shown by D. o. compactus of Padre and Mustang
islands, Texas, which is pale-colored as is the sand on which it
lives, and D. o. medius from east-central New Mexico and western
Texas, which is reddish as is the soil there, which is derived from
Permian rocks. In localities where alkaline soils are present, kan-
garoo rats may be found with a roseaceous cast to the pelage as a
result of the action of the alkaline salts on the pigment of the hair.
The roseaceous color is lost when the animal sheds the old pelage.
In the dorsal and ventral stripes of the tail, I find as much varia-
tion in the species D. ordii as Grinnell (1922: Fig. E, p. 14) recorded
in the whole genus. In D. o. obscurus, D. o. fuscus and D. o. utahensis
the stripes are complete to the distal end of the tail and dark, whereas
in D. o. pallidus and D. o. celeripes the ventral stripe is either absent
or nearly so and the dorsal stripe is pale.
Color as a taxonomic character is valuable in a broad sense, and
502 University of Kansas Publs., Mus. Nat. Hist.
is useful in placing an individual or a group of individuals in the
subspecies to which they pertain. In most subspecies studied, color
was quite uniform throughout the range of the animals, but in D. o.
ordii and D. o. columbianus color is so variable that cranial features
were relied on almost exclusively for the final diagnosis.
Among the subspecies of Dipodomys ordii there is relatively little
variation in the length of the head and body. The smallest measure-
ment is 95.5 mm. in D. o. idoneus and the largest is 118.3 mm. in
D. o. richardsoni. The shortest tail is found to be 112.0 mm. in
D. o. celeripes and the longest is 154.7 mm. in D. o. terrosus. The
length of the hind foot varies from 35.0 mm. in D. o. idoneus to 44.5
mm. in D. o. nexilis.
Allen's Rule is not operative in the species D. ordii. According
to this rule, shorter tails and smaller feet in conjunction with a
large body would be expected as the more northerly limits of the
species are approached, and conversely, smaller body and larger
appendages would be expected as the southerly limits of the species
are approached. This is not the case, however, since the subspecies
D. o. terrosus ranges farthest north and has the longest tail, whereas
D. o. celeripes, found in the central part of the range of the species,
has the shortest tail. Again, in regard to the hind foot, the shortest
is found in D. o. idoneus which is at the extreme south of the range
of the species, whereas the longest hind foot is found in D. o. nexilis
which occupies a nearly central position in the range.
Long tail and long hind foot would seem to be specializations for
saltation and the two would be expected to be correlated. Actually
there is no significant correlation in D. ordii. D. o. celeripes, in which
the hind foot is near the mean for the species (39.8 as opposed to
the mean of 40.7), has the shortest tail. D. o. compactus has a short
tail (117.0 mm.) but a medium-sized hind foot. D. o. nexilis and
D. o. terrosus have both a long hind foot and long tail.
Cranial measurements vary less, probably because one person can
measure a series with a uniformly subjective error. External meas-
urements, however, are liable to a greater degree of subjective error.
The total length of the skull varies from 35.4 mm. in D. o. attenuatus
to 41.3 mm. in D. o. terrosus. In no one series of adults from one
locality, however, is the variation so marked as it is for the species
as a whole. The usual range of variation in length of skull in any
given series is not, as a rule, more than 2.5 mm.
Cranial indices (breadth across bullae/length of skull X 100) as
established for random samples of the different species of the genus
Setzer: Subspeciation in Kangaroo Rat 503
(exclusive of D. ordii) ranged from 60.8 to 67.6. In the subspecies
of D. ordii the same index varies from 59.7 to 65.2 with an average
of 63.4. In other words, the degree of specialization indicated by
this one index, in a few subspecies of D. ordii, is almost as great
as that in Dipodomys deserti, which on the basis of total morphology
appears to be the most specialized species in the genus. Also, on the
basis of this same index, some subspecies of D. ordii are more gen-
eralized than is any other species in the genus.
There is a general tendency for the nasals to decrease in length
and the rostrum to decrease in width as the southern limits of the
range of D. ordii are approached. In ascertaining the decrease in
length of the nasals an index was obtained as follows: nasals/inter-
orbital width X 100 (see Table 4). The width of the rostrum, how-
ever, does not decrease in the same degree, nor at the same rate, as
does the length of the nasals. This decrease in length of the nasals
and in width of the rostrum may be correlated with the mean annual
relative humidity of the environment. It is known (Howell and
Gersh, 1936:8) that desert rodents, more exactly kangaroo rats,
have a water retention mechanism in the kidneys and walls of the
urinary bladder which enables them more efficiently to conserve
metabolic water. The significance of the decrease of the area of the
nasal mucosa, which seems to be related to relative aridity, is not
yet properly understood.
In no cranial feature other than shortened nasals and narrowed
rostrum, does Dipodomys ordii show a gradation such that it might
be termed a cline. Other parts of the skull that were measured do
not vary greatly.
Perhaps the greatest amount of variation in the skull is in features
which are not readily measurable by the usual physical means. The
shape and size of the pterygoid fossae vary from almost round to
rather ovoid in a given series of animals from one locality; the size
and configuration of the zygomatic arches vary from slender to
robust and from straight to curved laterally ; the size of the lacrimal
processes varies much in any given series, as do also the degree of
expansion of the distal end of the nasals, the convexity of the brain-
case and the curvature of the upper incisors. In all instances where
these features varied much, one size or shape was more pronounced
in the series than any other size or shape. Thus, when comparisons
were made, the size and certain shapes were the criteria used in
assigning the animals under consideration to a given subspecies.
Subspeciation in Dipodomys ordii seems to have been influenced
504 University of Kansas Publs., Mus. Nat. Hist.
by water barriers. It is known (Grinnell, 1922:28) that kangaroo
rats lack the ability to swim. Large stable rivers such as the Colo-
rado, Snake and Columbia serve as effective barriers to further
dispersal of kangaroo rats. Streams that freeze over in the winter
months, however, are not efficient barriers. This is indicated by the
"blending" of morphological characters of D. o. nexilis and D. o.
sanrafaeli along the Green River which freezes over.
Any mountain which has vegetational belts above the Transition
Life-zone would serve as a barrier to the dispersal of these ani-
mals. The Uinta Mountains, lying in an east-west direction, are
interposed between the ranges of D. o. priscus and D. o. uintensis.
The high Wasatch Mountains and their associated outliers, lying
in a north-south direction in Utah, serve as an efficient barrier to
the east-west movement of kangaroo rats and as a result, the sub-
species east of the mountain mass are remarkably different from
those to the west.
TABLE 4
Proportionate Decrease of Nasals
B
2
u
«**
o
■v
£
**-
o
g
3
!
C J3
is
g *
a
1
a
55
o
o
t-H
X
"3
13
o
u
a
— -
4.1
14.75
13.5
91.6
4.35
13.9
12.95
93.1
4.3
14.35
13.75
94.8
4.1
13.5
12.65
93.8
3.5
13.3
13.0
97.7
3.7
13.2
13.75
103.5
3.3
12.8
13.0
101
LI
terrosus . . .
luteolus. . .
evexus
montanus.
ordii
idoneus . . .
palmeri . . .
The first three columns represent the actual measurements of the various
elements; the fourth column is the index established.
Six different complexes (groups) of subspecies of D. ordii have
probably arisen as a result of geographical separation.
The Great Plains complex consisting of D. o. richardsoni, D. o.
oklahomae, D. o. evexus, D. o. terrosus, D, o. luteolus, D. o. priscus
Setzer: Subspeciation in Kangaroo Rat 505
and D. o. medius are, with the exception of D. o. priscus, inhabitants
of the high plains grassland habitat. D. o. priscus inhabits the red
Desert of Wyoming.
The Gulf Coast complex, comprising D. o. sennetti and D. o. com-
pactus are separable from all others by small auditory bullae and
short tail. D. o. compactus probably has differentiated from D. o.
sennetti since the cutting off, by wave action, from the mainland,
of the islands on which D. o. compactus lives.
The Mexican complex consisting of D. o. obscurus, D. o. fuscus,
D. o. idoneus and D. o. palmeri have probably differentiated by
natural selection acting on fortuitous variations, but I lack first
hand knowledge of the region concerned.
The southwestern complex consists of D. o. chapmani, D. o. ex-
tractus, D. o. attenuatus and D. o. ordii. D. o. attenuatus and D. o.
chapmani are subspecifically distinct owing to geographic isolation,
although both kinds show intergradation where their ranges ap-
proach that of D. o. ordii.
The western desert complex, composed of D. o. monoensis, utahen-
sis, cineraceus, columbianus, cinderensis, jetosus, celeripes, mar-
shalli, inaquosus, pallidas, panguitchensis and fremonti was isolated
from the other complexes of D. ordii by the Quaternary uplift of
the Wasatch Mountain mass, consisting of the Wasatch, Fish Lake
and San Pitch mountains and the Wasatch, Aquarius, Paunsaugunt
and Kaiparowits plateaus, and the concurrent reestablishment of
drainage systems. The drainages are those of the Colorado and
Columbia rivers and that of the Snake River from Blackfoot, Idaho,
to the junction with the Columbia. D. o. fremonti has been isolated
on the upper reaches of the Fremont River which arises from the
eastern side of the Wasatch Divide. D. o. panguitchensis has been
isolated in Panguitch Valley as a result of the canyons formed by
the Sevier River in Utah. D. o. cineraceus, although its subspecific
and insular status are in doubt, appears to have been isolated on
Dolphin Island, Great Salt Lake, Utah.
The intermontane complex consisting of D. o. montanus, longipes,
cupidineus, nexilis, sanrafaeli and uintensis, like the western desert
complex, has become separated from the remainder of the subspecies
of the species D. ordii by Quaternary geologic events. D. o. cupi-
dineus has been cut off by the gorges of the Colorado River to the
south and the Virgin River to the north. D. o. sanrafaeli is sepa-
rated from D. o. uintensis by the Tavaputs Plateau and by the Roan
and Book cliffs, and is separated from the range of D. o. nexilis by
3—6114
;,()<;
University of Kansas Publs., Mus. Nat. Hist.
cupidineus
nexilis
I
longipes
montanus'-
pallidus
inoquosus
marshalli
cinerqceus
cele'ripes
columbianus
utahensis
fetbsus
cinderensis
monoensis
fremonti
i
sanrafaeli
uintensis
ponguitchensis'
attenuates
paimeri
fuscus
idoneus
pnscus
chapman/
extractus
I
ordii
luteolus
obscurus
terrosus
I
medius
evexus
oklahomae
richardsoni
Fig. 22. An arrangement, according to morphological indices, of the sub-
species of Dipodomys ordii.
the Colorado River although there is intergradation between Z). o.
nexilis and D. o. sanrafaeli. D. o. longipes has been separated from
the rest of this intermontane complex by the San Juan and Colorado
rivers, but to the east it intergrades freely with adjacent subspecies.
D. o. montanus has been relatively isolated in the San Luis Valley of
Colorado and New Mexico, but in the southern part of its range it
does show intergradation with other subspecies.
The complexes mentioned above are represented graphically in
Figure 22, in a way that expresses some of my ideas as to their
genetic relationships.
The indices used to determine the amount of specialization that
each complex of subspecies has undergone are as follows:
The Body index (head and body/length of tail X 100) is the ex-
pression of the elongation of the tail as an organ of balance while
the length of the head and body remain relatively constant. As
the tail elongates the index decreases and as the tail becomes shorter
the index increases.
Setzer: Subspeciation in Kangaroo Rat 507
The Pedal index (hind foot/head and body X 100) is the expres-
sion of the development of the hind foot as an element essential for
the saltatorial habit. As the hind foot elongates the index will
increase; elongation of the hind foot is interpreted as a specializa-
tion.
The Cranial index (breadth across bullae/length of the skull X
100) reflects the degree of development of the tympanic or mastoid
region, or both, and is thought to be an adaptation for more acute
hearing and possibly for more delicate balance. Inflation of the
tympanic bullae is thought to be a specialization. As the auditory
bullae become more inflated, the index increases toward 100.
The Bullar index (width of maxillary arches/breadth across bul-
lae X 100) also expresses the degree of inflation of the auditory
bullae. In a generalized mammal, at least in the heteromyids, the
index would be 100, but as the auditory bullae become larger the
index will decrease from 100.
In attempting to arrange the subspecies of D. ordii according to
degree of specialization, the geographic positions of the subspecies
have been considered along with the information provided by the
above-mentioned indices. These indices were used in the same way
as were the indices for the species of the genus. In Tables 5 and 6
and in the accounts and maps the subspecies are arranged from the
least specialized to the most specialized.
508
University of Kansas Publs., Mus. Nat. Hist.
TABLE 5
Indices for the Subspecies of DIPODOMYS ORDII
71
V
6
3
richardsoni 88 . 85
oklahomae 86 . 75
compactus 127. 7
sennetti 94 . 25
evexus 80 . 1
medius 80 . 4
obscurus
terrosus 75 . 25
f remonti 80 . 55
uintensis 77 . 2
monoensis 85 . 4
ordii 79.05
luteolus 75.0
extractus 83 . 65
chapmani 75 . 05
montanus 80 . 4
cinderensis 85 . 1
fetosus 81.8
utahensis 80 . 2
columbianus 78 . 5
idoneus 72 . 3
priscus 74 . 9
celeripes 91 . 85
cineraceus 75 . 5
marshalli 81 . 5
inaquosus 78 . 05
attenuates 73 . 5
fuscus 79.8
longipes 75 . 7
pallidus 76.9
nexilis 77. 1
cupidineus 73. 15
palmeri 72 . 25
34.35
60.95
88.25
35.5
61.7
90.25
37.25
59.75
88.35
34.0
62.85
85.95
35.7
60.5
92.9
33.7
63.55
85.9
62.95
86.4
35.05
61.6
86.85
34.7
62.9
85.5
35.3
62.3
86.0
36.4
63.4
85.6
37.6
62.75
86.9
37.05
62.35
86.3
34.35
64.3
84.25
36.35
62.9
85.65
36.15
64.25
82.5
37.2
65.15
84.75
38.85
63.95
83.95
36.95
64.45
84.35
37.55
64.25
84.9
36.6
64.2
85.0
39.45
62.3
84.95
38.75
65.0
84.25
39.1
63.9
84.8
37.3
65.2
83.0
37.9
64.25
83.05
37.35
64.0
83.4
39.0
64.3
83.2
37.1
64.3
82.75
40.75
64.35
84.65
40.7
64.95
78.45
39.1
64.1
80.85
37.15
65.1
80.45
Setzer: Subspeciation in Kangaroo Rat 509
TABLE 6
Numerals (derived from Table 5) are Indicative of the Relative Degree of
Specialization of the Subspecies of DIPODOM YS ORDI1
>>
"3
■a
a
u
03
8
-a
o
•a
e
a
3
£
«
^
0
pa
<
richardsoni 4
oklahomae 5
compactus 1
sennetti 2
evexus 15
medius 12
obscurus
terrosus 25
fremonti 11
uintensis 20
monoensis 6
ordii 17
luteolus 27
extractus 8
chapmani 26
montanus 13
cinderensis 7
fetosus 9
utahensis 14
columbianus 18
idoneus 31
priscus 28
celeripes 3
cineraceus 24
marshalli 10
inaquosus 19
attenuatus 29
fuscus 16
longipes 23
pallidus 22
nexilis 21
cupidineus 30
palmeri 32
3
3
4
3.5
8
5
2
5.0
19
1
3
6.0
2
10
10
6.0
8
2
1
6.5
1
15
11
9.75
12
7
10.0
6
4
6
10.25
5
12
14
10.5
7
7
9
10.75
12
14
13
11.25
23
9
5
13.5
15
8
8
14.5
4
25
22
14.75
11
11
12
15.0
10
23
30
19.0
18
32
19
19.0
26
17
24
19.0
14
28
21
19.25
22
21
17
19.5
13
20
15
19.75
30
6
16
20.0
25
30
23
20.25
28
16
18
21.5
20
33
28
22.75
24
22
27
23.0
21
18
25
23.25
27
24
26
23.25
16
26
29
23.5
32
27
20
25.25
31
29
33
26.0
29
19
31
27.25
17
31
32
28.0
510 University of Kansas Publs., Mus. Nat. Hist.
Fig. 23. Distribution of subspecies of Dipodomys ordii.
1. D.o. richardsoni 13. D.o. monoensis 25. D.o. celeripes
2. D.o. oklahomae
3. D. o. compactus
4. D. o. sennetti
5. D. o. evexus
6. D.o. medius
7. D. o. obscurus
8. D.o. terrosus
9. D. o. panguitchensis
10. D.o. uintensis
11. D.o. sanrajaeli
12. D. o. jremonti
14. D. o. ordii
15. D.o. luteolus
16. D. o. exlractus
17. D.o. chapmani
18. Z).o. montanus
19. D.o. cinderensis
20. D.o. fetosus
21. D.o. utahensis
26. D. o. cineraceus
27. D.o. marshalli
28. D. o. inaquosus
29. D. o. attenuatus
30. D.o. juscus
31. D.o. longipes
32. D. o. pallidus
33. D. o. nexilis
22. D.o. columbianus 34. D.o. cupidineus
23. D. o. idoneus 35. D. o. palmeri
24. D.o. prisons
Setzer: Subspeciation in Kangaroo Rat 511
Dipodomys ordii
Ord Kangaroo Rat
Dipodomys ordii is a medium sized, relatively short-tailed, five-
toed species of a color about average for the genus. As in other
members of the genus, the hind legs and feet are disproportionately
long as an adaptation to the saltatorial mode of progression. The
upperparts are buffy, reddish or blackish, depending on the sub-
species, but the entire ventral surface, dorsal surfaces of the hind
feet, supraorbital and postauricular spots, forelimbs, hip stripes,
lateral stripes of the tail and the tail at the base are pure white.
The skull has a relatively short rostrum, moderate to large auditory
bullae, relatively wide interparietal, relatively wide maxillary arches
and grooved upper incisors.
The only other five-toed kangaroo rats with which Dipodomys ordii,
at places, shares its geographic range, are Dipodomys panamintinus
and Dipodomys microps. Dipodomys ordii can be distinguished
from Dipodomys panamintinus by smaller size (for instance the
hind foot is shorter instead of longer than, 44 mm.) and narrower
expanse of maxillary arches in relation to breadth across the audi-
tory bullae, and from Dipodomys microps by the awl-shaped, in-
stead of chis-el-shaped, lower incisors.
The species D. ordii is divisible into 35 subspecies, accounts of
which follow:
Dipodomys ordii richardsoni (Allen)
Dipodops richardsoni Allen, Bull. Amer. Mus. Nat. Hist., 3:277, June
30, 1891.
Dipodomys phillippi, Knox, Trans. Kansas Acad. Sci., 4:22, 1875, (part
— the part from Osborne, Kansas).
Dipodormjs phillipsi ordi, Coues and Allen, Monogr. North American
Rodentia, p. 542, 1877 (part — the part from Ft. Cobb, Oklahoma).
Perodipus richardsoni, Allen, Bull. Amer. Mus. Nat. Hist., 7:260, Au-
gust 21, 1895 (part — the part from Pendennis, Kansas).
Cricetodipus richardsoni, Trouessart, Catalogus Mammalium, 1 :581,
1897.
Perodipus montanus richardsoni, Bailey, N. Amer. Fauna, 25:144, Oc-
tober 1905 (part — the part from Canadian, Texas).
Perodipus ordii richardsoni, Goldman, Proc. Biol. Soc. Washington,
30:113, May 23, 1917.
Dipodomys ordii richardsoni, Grinnell, Journ. Mamm., 2:96, May 2,
1921.
Type.— Male, no. 3025/2345, Amer. Mus. Nat. Hist.; on one of the sources
of the Beaver River, Beaver County, Oklahoma; obtained on October 26,
1887, by Jenness Richardson and John Rowley, Jr. (After Allen, original
description, type not seen.)
512 University of Kansas Publs., Mus. Nat. Hist.
Range. — Southwestern Nebraska, eastern Colorado, northeastern New Mex-
ico, Panhandle of Texas, and western parts of Oklahoma and Kansas; mar-
ginal localities are: in Nebraska: Bladen, Haigler; in Colorado: Olney; in
New Mexico: Clayton; in Texas: 6 mi. S and 1 mi. W Quitaque, Vernon;
in Oklahoma: 3 mi. S Cleo Springs; in Kansas: Medora.
Diagnosis. — Size large (see measurements). Color dark; entire dorsal sur-
face Cinnamon-Buff, purest on sides and flanks, upper parts suffused with
black; arietiform markings, pinnae of ears, plantar surfaces of hind feet,
dorsal and ventral stripes of tail, blackish; in some specimens the ventral
stripe of tail does not extend to tip of pencil. Skull large; rostrum short and
wide; nasals long; zygomata relatively heavy; auditory bullae well inflated
and wide; thus with short rostrum giving appearance of nearly equilateral
triangle; upper incisors long and robust.
Comparisons. — From Dipodomys ordii luteolus, D. o. richardsoni differs
as follows: Size smaller in external measurements except length of body
which is longer; color darker, except on plantar surfaces of hind feet and
dorsal and ventral stripes of the tail which are lighter; ventral stripe of tail,
in most specimens, continuous to end of pencil, whereas in D. o. luteolus
ventral stripe is present on only proximal two-thirds; skull larger in all
measurements taken; zygomatic arch heavier; auditory bullae relatively as
well as actually more inflated; external auditory meatus egg-shaped as con-
trasted to nearly round in D. o. luteolus; pterygoid fossae rounded as com-
pared to ovate in D. o. luteolus.
From Dipodomys ordii oklahomae, D. o. richardsoni differs as follows:
Size larger in all measurements taken; color darker in all pigmented areas;
skull larger in all respects ; auditory bullae larger and more inflated ventrally ;
jugal straight or nearly so instead of bowed laterad; pterygoid fossae smaller;
nasals straight instead of inflated as a "bulb" distally.
For comparisons with D. o. montanus and D. o. evexus, see accounts of
those subspecies.
Remarks. — This race of Dipodomys ordii is readily distinguished
from Dipodomys ordii evexus, from the valley of the upper Arkansas
River, by larger size, larger skull and lighter color. Intergradation
with Dipodomys ordii luteolus occurs rather freely in northeastern
Colorado, as indicated by specimens from 3 miles northeast of Fitz-
simmons, 6 miles east and 1 mile north of Denver and Barr Lake.
These specimens resemble D. o. richardsoni in light color, greater
inflation of the auditory bullae and the shape of the pterygoid fossae
but resemble D. o. luteolus, to which they are here referred, in the
length of the nasals, the least interorbital width and in the external
measurements. In the southern part of the range of D. o. richardsoni
intergradation occurs with Dipodomys ordii medius, as at 6 miles
southwest of Muleshoe, Texas. Specimens from there have the long,
wide rostrum and narrow skull of D. o. richardsoni but in the sum
total of their characters more closely resemble D. o. medius. At
Setzer: Subspeciation in Kangaroo Rat 513
Texline, Texas, the animals show intergradation in the length and
shape of the nasals and degree of convexity of the cranium but are
referable to D. o. richardsoni.
In fine, intergradation occurs at all points where the range of
D. o. richardsoni touches that of any other geographic race. No one
series of it is as uniform as are most series of specimens of other
known races. Dipodomys ordii richardsoni shows a mixture of
characters. Nevertheless, each of the populations studied has char-
acters in most of the animals that make this form recognizable as a
taxonomic unit — a unit that seems, as yet, not to have become
stabilized even in the central parts of its range.
Coues and Allen (1877:542) list specimens from Fort Cobb, Ar-
kansas. It is known that the Post Office Department, for adminis-
trative purposes, attached certain towns and military installations
in Indian Territory (now Oklahoma) to the State of Arkansas.
Thus it is apparent that Fort Cobb, Arkansas, as recorded by Coues
and Allen (loc. tit.) is Fort Cobb, Oklahoma. Specimens from Fort
Cobb would be expected to be D. o. richardsoni.
Specimens examined. — Total, 351, distributed as follows:
Nebraska: Adams County: Bladen, 10 (AMN1I). Dundy County: Haigler, 1 (USBS).
Colorado: Crowley County: Olney, 1 (USBS). Kiowa County: Chivington, 3 (USBS).
Otero County: 18 mi. S La Junta, 4 (AMNH); Higbee, 1 (USBS). Bent County: 4 mi. SE
Las Animas, 4100 ft., 3 (MVZ). Prowers County: Lamar, 9 (LACM); 1 mi. S Lamar,
4000 ft., 11 (KU). ' Baca County: Gaumes Ranch, 4600 ft., NW Corner, 1 (USBS).
Kansas: Cheyenne County: 23 mi. NW St. Francis, 5 (KU). Rawlins County: 2 mi.
NE Ludell, 2 (KU); iy2 mi. W Ludell, 1 (KU). Wallace County: Lacey Ranch, 4% mi. E
and 9 mi. S Wallace, 1 (KU); unspecified, 2 (KU). Logan County: 5 mi. W Elkader, 2
(KU); unspecified, 1 (UM). Gove County: unspecified, 1 (KU). Trego County: Banner,
8 (USNM); Parrington Ranch, 12 mi. S Collyer, 2 (KU); unspecified, 8 (USNM). Ellis
County: Ellis, 1 (USBS). Lane County: Pendennis, 10 (USBS). Hamilton County: Cool-
idge, 2 (CNHM); 1 mi. E Coolidge, 5 (KU). Pawnee County: 1 mi. S Larned, 4 (KU) ;
2 mi. S and M mi. W Larned, 2 (KU); 3 mi. S and IY2 mi. W Larned, 10 (KU). Edwards
County: Kinsley, 3 (USBS); 3y2 mi. E Kinsley, 5 (KU); S side Arkansas River, 2 mi. S
Kinsley, 1 (KU); 1 mi. W and 3>/2 mi. S Kinsley, 9 (KU). Stafford County: Little Salt
Marsh, 15 mi. N and 3 mi. E Stafford, 2 (KU). Reno County: Medora, 1 (UM); 2 mi. W
and V2 mi. S Medora, 4 (KU). Kiowa County: 5 mi. N Belvidere, 1 (KU). Pratt County:
Cairo, 2 (USBS). Sedgwick County: Wichita, 6 (AMNH). Morton County: 10 mi. N and
3 mi. E Elkhart, 34 (KU). Seward County: 1 mi. E Arkalon, 7 (KU); Liberal, 1 (KU) ;
unspecified, 1 (KU). Meade County: Meade, 1 (USNM); 13 mi. SW Meade, 13 (6 AMNH;
7 KU); 17 mi. SW Meade, 2 (KU). Clark County: 12 mi. S Ashland, 1 (UM); unspecified,
3 (KU). Barber County: Medicine Lodge, 4 (USBS); 1 mi. W Aetna, 3 (KU); % mi. W
Aetna, 2 (KU); Aetna, 3 (KU); 1 mi. SW Aetna, 1 (KU); 1% mi. SW Aetna, 1 (KU);
1 mi. S Aetna, 5 (KU) ; unspecified, 2 (KU). Harper County: 4y2 mi. NE Danville, 12 (KU) ;
2 mi. NE Runnymede, 3 (KU).
New Mexico: Union County: Clayton, Apache Canyon, 1 (USIBS). Quay County:
Glenrio, 10 (LACM).
Oklahoma: Cimmaron County: Kenton, 1 (CM). Beaver County: iy2 mi. N Beaver,
7 (KU); Beaver River, 8 (7 AMNH; 1 CNHM). Harper County: 3 mi. S of Englewood,
Kansas, 2 (MVZ); 4V2 mi. N Laverne, 1 (UM). Woods County: 2 mi. W Edith, 1 (USBS);
Alva, 12 (UM); Waynoka, 18 (UM); 3 mi. SW Waynoka, 1 (USBS). Alfalfa County: 4
mi. SE Cherokee, 1 (USBS). Ellis County: Shattuck, 1 (USBS). Woodward County:
Woodward, 9 (USBS). Major County: 3 mi. S Cleo Springs, 1 (USBS).
514 University of Kansas Publs., Mus. Nat. Hist.
Texas: Dallam County: Texline, 8 (USBS). Lipscomb County: Lipscomb, 3 (USBS)
Hemphill County: 17 mi. NE Canadian, 1 (MVZ); 1 mi. W Canadian, along Red Deer River,
12 (MVZ); y2 mi. W Canadian, along Red Deer River, 7 (MVZ); Canadian, 5 (USBS).
Oldham County: Tascosa, 6 (USBS). Wheeler County: 1 mi. W Mobeetie, 2 (MVZ); Mo-
beetie, 8 (USBS); Wallace Ranch, SW Wheeler County, 1 (TCWC). Hall County: Newlin,
1 (USBS). Wilbarger County: Vernon, 5 (USBS). Floyd County: 6 mi. S and 1 mi. W
Quitaque, 1 (I'M),
Dipodomys ordii oklahomae Trowbridge and Whitaker
Dipodomys oklahomae Trowbridge and Whitaker, Journ. Mamm., 21:
343, August 14, 1940.
Dipodomys ordii oklahomae, Davis, Journ. Mamm., 23:332, August 14,
1942.
Type.— Female, young adult, no. 265454, U. S. Nat. Mus., Biol. Surv. Coll.
(formerly Univ. of Oklahoma, Mus. Zool., no. 14517) ; north bank of South
Canadian River, 2*4 mi. S Norman, Cleveland County, Oklahoma; obtained
on March 16, 1934, by H. L. Whitaker, original no., X-catalog no. 29312 of
U. S. Nat. Mus.
Range. — Known only from the South Canadian River Valley west of Minco,
Canadian County; and east to Lexington, Cleveland County, Oklahoma.
Diagnosis. — Size medium (see measurements). Color light, entire dorsal
surface near (c) Vinaceous-Buff, paler on sides with great suffusion of white;
arietiform markings, pinnae of ears, plantar surfaces of hind feet, proximal
ventral portion of tail and dorsal stripe on tail, brownish. Skull of medium
size; rostrum wide; nasals short, projecting but slightly anteriorly to incisors;
zygomatic processes of maxillae heavy; bullae not greatly inflated.
Comparisons. — Dipodomys ordii oklahomae differs from D. o. richardsoni
as follows: Size smaller; color lighter in all pigmented areas; ventral stripe of
tail extending only one-fourth the length rather than three fourths or to end
of tail; skull smaller in all measurements taken; rostrum heavier; auditory
bullae less inflated; pterygoid fossae larger; braincase slightly more inflated;
nasals more expanded distally; interparietal region wider.
From Dipodomys ordii ordii, D. o. oklahomae differs in: Size larger in all
measurements taken; color lighter in all pigmented areas; ventral stripe of
tail extending one fourth length of tail rather than to end; skull larger in all
respects; rostrum heavier; zygomatic arches heavier; bullae more inflated
ventrally; cutting edge of upper incisors wider; pterygoid fossae larger; brain-
case more vaulted; nasals more expanded distally; orbital region larger; in-
terparietal region wider.
Remarks. — Trowbridge and Whitaker named this kangaroo rat
as a full species. The diagnostic characters were the length and
breadth of the rostrum and the relatively great inflation of the audi-
tory bullae. Also, Dipodomys oklahomae was not known to inter-
grade with any other named kinds. Davis (1942:332) treated D.
oklahomae as a subspecies of the earlier named species Dipodomys
ordii. Certain characters in specimens from the type series of both
D. o. richardsoni and D. oklahomae, such as the shape and configu-
ration of the nasals, the over-all proportion of the skull, tooth pat-
tern and body proportions through individual variation overlap and
Setzer: Subspeciation in Kangaroo Rat 515
indicate that these two groups of animals belong to the same species,
even though animals from intermediate geographic areas are not
available to show actual intergradation. My findings corroborate
Davis' conclusion that D. oklahomae should stand as Dipodomys
ordii oklahomae. In spite of the great similarities shown by the
two groups of animals there are still sufficient diagnostic characters
between the two groups to enable them to be segregated easily as
valid subspecies.
Dipodomys ordii oklahomae is, for some unknown reason, re-
stricted to a limited geographic range. Specimens examined from
the upper reaches of the South Canadian River, farther westward,
are all referable to D. o. richardsoni rather than, as would be ex-
pected, to D. o. oklahomae since the habitat for these animals is
continuous from the type locality of D. o. oklahomae to the upper
reaches of the South Canadian River. In length and shape of the
nasals, degree of inflation of the auditory bullae and width of the
interorbital region these specimens from the upper reaches of the
South Canadian River are intergrades between D. o. richardsoni
and D. o. medius. The range of D. o. medius lies to the south of
that of D. o. richardsoni and to the southwest of that of D. o. okla-
homae.
The present range of D. o. oklahomae, as now understood, is the
most eastern part of the range of the species Dipodomys ordii and
of the genus Dipodomys. The existence of D. o. oklahomae in this
area is a precarious one since its habitat is limited in extent and is
periodically flooded.
Although no specimens are known from the area where intergra-
dation between D. o. oklahomae and D. o. richardsoni would be ex-
pected to occur, it would seem that when animals from this region
become available, intergradation will be shown to occur.
Specimens examined. — Total, 8, all from Oklahoma, distributed as follows: Grady County:
4 mi. N Minco, 1 (USBS). Cleveland County: 2% mi. S Norman, 7 (6 OU ; 1 USBS).
Dipodomys ordii compactus True
Dipodomys compactus True, Proc. U. S. Nat. Mus., 11:160, January 5,
1889.
Cricetodipus compactus, Trouessart, Catalogus Mammalium, 1:581,
1897.
Perodipus compactus, Elliot, Field Columbian Museum, Zool. Ser.,
2:240, 1901.
Dipodomys ordii compactus, Davis, Journ. Mamm., 23:332, August 14,
1942.
Type. — None designated but Poole and Schantz (1942:406) assumed it to
be a female, no. 19665/35227, only the skin found, from Padre Island, Cameron
County, Texas. April 3, 1888. Purchased from C. K. Worthen.
516 University of Kansas Publs., Mtjs. Nat. Hist.
Range. — Padre and Mustang islands, Cameron County, Texas.
Diagnosis. — Size medium (see measurements); tail short. Color light; en-
tire dorsal surface Light Oehraceous-Buff, purest on sides and flanks, upper
parts but lightly suffused with black. A lighter color phase has entire dorsal
surface Cartridge Buff, purest on sides and flanks, upper parts but lightly
washed with black. In both phases, cheeks white; pinnae of ears, plantar sur-
faces of hind feet, dorsal stripe of tail, ventral stripe of tail (in most specimens)
present on proximal third of tail only, brownish. Skull small; rostrum nar-
row and long; nasals long; auditory bullae inflated, but greatest breadth
across bullae only slightly more than breadth across zygomatic processes of
maxillae; interparietal region wide.
Comparisons. — From Dipodomys ordii sennetti, D. o. compactus differs in:
Size slightly less; color lighter in all pigmented areas; skull smaller; auditory
bullae slightly less inflated; interorbital width less; interparietal region wider;
nasals longer.
From Dipodomys ordii attenuatus, D. o. compactus differs in: Body larger;
tail shorter; normal color phase darker, and lighter color phase lighter; skull
larger; rostrum wider and longer; nasals longer; interorbital region wider;
auditory bullae relatively as well as actually less inflated ; interparietal region
wider; pterygoid fossae large and round as opposed to small and ovoid.
Compared with Dipodomys ordii medius and Dipodomys ordii ordii, D. o.
compactus is smaller, lighter in color, and has less inflated auditory bullae and
a smaller skull.
Remarks. — This subspecies of Dipodomys ordii was originally
described as Dipodomys compactus by True in 1889 and stood as
a full species until Davis (1942:332) relegated it to subspecific
status under Dipodomys ordii. Davis (op. cit.) observed close re-
semblances in external proportions, size of mastoid bullae, width
of supraoccipital, and size and shape of the interparietal, between
Dipodomys ordii and Dipodomys sennetti and therefore concluded
that they were only subspecies of one species. He observed that
the difference between Dipodomys compactus and Dipodomys sen-
netti was of approximately the same degree as that between Di-
podomys sennetti and Dipodomys ordii. From this he concluded
that all three were subspecies of the one species Dipodomys ordii.
In any sizeable sample of Dipodomys sennetti there are crania
closely resembling those of Dipodomys ordii ordii and others closely
resembling those of Dipodomys compactus. The external propor-
tions of both D. sennetti and D. compactus are duplicated in D.
ordii from El Paso and conversely, specimens with the proportions
of typical D. o. ordii occur in populations of D. sennetti and D. com-
pactus. Thus, it appears that Davis' usage of the name Dipodomys
ordii compactus should stand although there may be a hiatus in
geographic occurrence between D. ordii and D. sennetti, as of course
there is between D. sennetti and D. compactus.
Setzer: Subspeciation in Kangaroo Rat 517
In D. o. compactus there is a complete enamel ring around the
occlusal surface of each molarif orm tooth ; in D. o. ordii this ring is
incomplete lingually on each of the molariform teeth and labially
on the first three, and in some individuals of D. o. sennetti the
enamel ring is complete and in others it is incomplete labially and
lingually as in D. o. ordii.
Specimens examined. — Total, 44, all from Texas, distributed as follows: Nueces County:
19 mi. S Port Aransas, Mustang Island, 27 (17 TCWC; 10 MVZ) ; Mustang Island, 17
(LACM).
Dipodomys ordii sennetti (Allen)
Dipodops sennetti Allen, Bull. Amer Mus. Nat. Hist., 3:226, April 29,
1891.
Cricetodipus sennetti, Trouessart, Catalogus Mammalium, 1 :581, 1897.
Perodipus sennetti, Elliott, Field Columbian Museum, Zool. Ser.,
2:239, 1901.
Dipodomys ordii sennetti, Davis, Journ. Mamm., 23:332, August 14,
1942.
Type. — Male, no. 3478/2733. Amer. Mus. Nat. Hist.; near Brownsville,
Cameron County, Texas; obtained on March 9, 1888, by J. M. Priour. (After
Allen, original description, type not seen.) Type locality recorded by Bailey
(1905:145) as "Santa Rosa, 85 mi. SW Corpus Christi."
Range. — Southern Texas, south of Corpus Christi; marginal localities, all
in Texas are: Somerset, 8 mi. NE Los Angeles, 8 mi. E Encinal, Santa Rosa,
28 mi. E Raymondville, 2 mi. S Riviera.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face (c) between Pinkish Buff and Cinnamon-Buff, purest on sides and
flanks, upper parts mixed with black ; arietiform markings, pinnae of ears,
dorsal and ventral stripes of tail, plantar surfaces of hindfeet, brownish-black.
Skull small; auditory bullae but slightly inflated in relation to size of skull;
nasals slightly flaring distally; premaxillae extending but slightly posterior to
nasals; interorbital width relatively great; external auditory meatus small;
rostrum relatively long and wide; zygomatic arches relatively heavy.
Comparisons. — From Dipodomys ordii ordii, D. o. sennetti differs in: Size
smaller, tail shorter; color darker; skull smaller; nasals longer; rostrum
wider; auditory bullae less inflated; external auditory meatus smaller;
pterygoid fossae more rounded; zygomatic arches heavier.
From Dipodomys ordii medius, D. o. sennetti differs as follows: Size
smaller; color darker, but with less red in pelage; skull markedly smaller in
all respects.
From Dipodomys ordii compactus, D. o. sennetti differs in: Size somewhat
less; color darker; skull with total length greater; orbit smaller; least in-
terorbital width greater; braincase more inflated; width across auditory bullae
more; interparietal region wider; external auditory meatus larger; medial
part of audital portion (see Howell, 1932) of auditory bullae larger.
Remarks. — Dipodomys sennetti, along with Dipodomys com-
pactus, was regarded by Davis as conspecific with Dipodomys ordii.
Reasons for placing these two kinds of kangaroo rats as subspecies
of D. ordii are given in the account of Dipodomys ordii compactus.
518 University of Kansas Publs., Mrs. Nat. Hist.
This subspecies is known only from north of the Rio Grande
which may serve as a barrier to the spread of the animal into
northern Tamaulipas.
Specimens examined. — Total, 20, all from Texas, distributed as follows: Atascosa County:
Somerset, 2 (TCWC). LaSalle County: 8 mi. NE Los Angeles, 1 (TCWC) ; 8 mi. E Encinal,
1 (TCWC). Kleberg County: 2 mi. S Riviera, 9 (TCWC). Jim Hogg County: Hebronville,
3 (LACM). Brooks County: Falfurrias, 2 (LACM). Willacy County: 28 mi. E Raymond-
ville, 2 (TCWC).
Dipodomys ordii evexus Goldman
Dipodomys ordii evexus Goldman, Journ. Washington Acad. Sci.,
23:468, October 15, 1933.
Perodipus montanus richardsoni Warren, Mammals of Colorado, p. 76,
1910 (part — the part from Salida, Colorado).
Type.— Male, adult, no. 150990, U. S. Nat. Mus. Biol. Surv. Coll.; Salida,
Chaffee County, Colorado (altitude 7000 ft.) ; obtained on November 10,
1907, by Merritt Cary, original no. 1245.
Range. — Upper Arkansas River Valley of south-central Colorado, from
Salida to Pueblo.
Diagnosis. — Size large (see measurements). Color dark, entire dorsal surface
between (16") Pinkish Cinnamon and Cinnamon-Buff, purest on sides and
flanks, upper parts strongly suffused with black; arietiform markings, pinnae
of ears, plantar surfaces of hind feet and dorsal and ventral stripes of tail,
blackish. Skull of medium size; rostrum short and wide; nasals short; audi-
tory bullae but slightly inflated; braincase but slightly vaulted.
Comparisons. — From Dipodomys ordii richardsoni, D. o. evexus differs as
follows: Size smaller in all measurements taken; color darker; ears darker,
dorsal and ventral stripes on tail darker, arietiform markings darker and more
distinct, plantar surfaces of hind feet darker; skull smaller in all measure-
ments; length, as expressed in percentage of width of skull, greater in D. o.
evexus (66 per cent in D. o. evexus, 62 per cent in D. o. richardsoni which
gives the appearance of a long, narrow skull as contrasted with a rather short,
wide skull); auditory bullae less expanded laterally, posteriorly and ventrally;
interparietal region relatively wider in proportion to greatest width across
auditory bullae; cutting edge of upper incisors narrower; pterygoid fossae
smaller and more nearly circular.
Compared with Dipodomys ordii luteolus, D. o. evexus differs as follows :
Size somewhat smaller in external measurements; color darker in all pig-
mented areas; skull smaller in two of the seven measurements taken but in
the other five measurements somewhat larger; auditory bullae less inflated;
cutting edge of upper incisors narrower; zygomatic arch heavier; pterygoid
fossae smaller and more nearly circular; external auditory meatus ovoid as
contrasted to nearly circular; paroccipital processes smaller.
From Dipodomys ordii nexilis, D. o. evexus differs in: Color darker;
rostrum wider and shorter; interorbital region wider; breadth across maxillary
arches greater; auditory bullae less inflated; interparietal region larger;
zygomatic arches heavy and bowed laterad; molariform teeth smaller; cut-
ting edge of upper incisors narrower.
Setzer: Subspeciation in Kangaroo Rat 519
For comparison with Dipodomys ordii montanus see account of that sub-
species.
Remarks. — This race of kangaroo rat, described from the upper
Arkansas River Valley, closely resembles Dipodomys ordii luteolus
but differs in darker color, slightly smaller body and larger skull.
No evidence of intergradation with any other race was noted.
To the south the range of D. o. evexus is separated from that of D. o.
montanus by a high, transverse ridge of the Rocky Mountains which
is inhospitable to these animals. Much territory inhospitable to
Dipodomys intervenes also between the ranges of D. o. evexus and
D. o. luteolus, but there are areas connecting the northern part of
the range of D. o. evexus and the southwestern part of the known
range of D. o. luteolus, in which Dipodomys may occur. If kangaroo
rats occur in these areas it is to be expected that they will show
intergradation between the two subspecies concerned.
Specimens examined. — Total, 24, all from Colorado, distributed as follows: Chaffee
County: Salida, 10 (3 USBS ; 7 AMNH). Fremont County: Canyon City, 13 (USBS).
Pueblo County: Pueblo, 1 (USBS).
Dipodomys ordii medius new subspecies
Perodipus montanus richardsoni, Bailey, N. Amer. Fauna, 25:144, Oc-
tober, 1905 (part — the part from Santa Rosa, New Mexico).
Type.— Male,. no. 118526, U. S. Nat. Mus. Biol. Surv. Coll.; Santa Rosa,
Guadalupe County, New Mexico; obtained on October 5, 1902, by Jas. H.
Grant, original no. 565.
Range. — From north-central New Mexico, southeastward to west-central
Texas; marginal localities are, in New Mexico: 15 mi. N Ojo Caliente, Gallina
Mts., Deer Creek, San Pedro; in Texas: 20 mi. N Monahans, Colorado, 7 mi.
E Post, 6 mi. SW Muleshoe.
Diagnosis. — Size medium (see measurements). Color dark; entire dorsal
surface (14") between Orange-Cinnamon and Cinnamon, purest on sides and
flanks, dorsal surface lightly washed with black; arietiform markings, pinnae
of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail, brownish-
black. Skull of medium size; nasals long; medial mastoid portion of auditory
bullae well inflated caudad; braincase vaulted; external auditory meatus
small; rostrum short and truncate; medial auditory portion of auditory bullae
relatively little inflated; pterygoid fossae ovate; zygomatic arches slender and
relatively straight; junction of jugal and zygomatic process of maxilla heavy.
Comparisons. — From Dipodomys ordii richardsoni, D. o. medius differs as
follows: Tail longer; hind foot shorter; color darker; arietiform markings
more distinct; white lateral stripes of tail narrower; ventral stripe of tail in
most specimens complete to end of pencil; postauricular spots less pronounced;
hip stripe narrower and in some specimens almost obliterated; skull smaller
in all measurements taken; angle of dorsal extension of premaxilla with zygo-
matic process of maxilla more nearly 90°; braincase more vaulted; medial
520 University of Kansas Publs., Mus. Nat. Hist.
mastoid portion of auditory bullae more inflated, and coming to more of a
point; medial auditory portion of auditory bullae more inflated ventrally;
rostrum shorter and narrower; external auditory meatus smaller.
From Dipodomys ordii montanus, D. o. medius* differs in: Color lighter in
all pigmented areas; skull larger in all respects; rostrum shorter and heavier;
bullae more inflated; zygomata, while nearly straight, are bowed slightly
laterally; pterygoid fossae more ovate; foramen magnum larger; pterygoid
foramina smaller.
Compared with Dipodomys ordii ordii and Dipodomys ordii sennetti, D. o.
medius is larger and darker. The skull is also larger in all measurements
taken.
Compared with Dipodomys ordii longipes, D. o. medius is darker and smaller.
Remarks. — This hitherto undescribed race of Dipodomys ordii
can readily be distinguished from any of its near neighbors by the
characters set forth under diagnosis and comparisons.
Intergradation is noted with D. o. ordii, D. o. longipes, D. o.
montanus and D. o. richardsoni. Among named races D. o. medius
shows closest affinities with D. o. richardsoni but the two are easily
separable. The northwestern extremity of the range of D. o. medius
is an area of integradation in which no specimens are clearly of one
subspecies or the other. In specimens from 5 miles east of Abiquiu,
New Mexico, three-way intergradation occurs. These animals are
like D. o. medius in size, D. o. longipes in color and their cranial
proportions are as in D. o. montanus. At Deer Creek, New Mexico,
and at Monahans, Texas, the animals show intergradation in size
of body and agree with D. o. ordii in cranial proportions. In speci-
mens from 6 miles southwest of Muleshoe, Texas, intergradation
with D. o. richardsoni in the shape of the skull and width of the
rostrum is noted. In the sum total of characters studied, however,
these specimens are referable to D. o. medius.
Specimens examined. — Total, 129, distributed as follows:
IS'ew Mexico: Rio Arriba County: 2 mi. SE El Rito, 2 (KU) ; Rio Alamosa, 15 mi.
N Ojo Caliente, 1 (USBS) ; 5 mi. E Abiquiu, 4 (USBS) ; Rinconada, 5 (USBS); Espanola,
C (USBS). Sandoval County: 12 mi. NW Alameda, 5500 ft., 3 (MVZ). Sante Fe County:
Seton's Ranch, near Santa Fe, 1 (USBS); 8 mi. SW Santa Fe, 8 (KU); San Pedro, 3
(USBS). San Miguel County: Pecos, 2 (USBS); 3 mi. S Pecos, 2 (USBS); Rowe, 6
(LACM). Bernallilo County: Bear Canyon, Sandia Mountains, 7 (USBS); Pajarito, 3
(MVZ). Guadalupe County: Santa Rosa, 10 (USBS). Lincoln County: 44 mi. NW Ro -
well, 5 (MVZ). De Baca County: 8 mi. N Fort Sumner, 9 (USBS). Roosevelt County:
Kenna, 4 (LACM). Curry County: 4 mi. W and 2% mi. N Clovis, 1 (MVZ). Chaves
County: 40 mi. N Roswell, 2 (USBS); 35 mi. N Roswell, 2 (USBS); 15 mi. NE Roswell,
8 (LACM); Stinking Springs Lake, 3 (USBS).
Texas: Bailey County: (i mi. SW Muleshoe, 5 (I'M); !' mi. SW Muleshoe, 2 (I'M).
Garza County: 7 mi. E Post, 5 (UM). Martin County: Stanton, 4 (USBS). Howard
County: G mi. NE Coahoma, 7 (UM); 1 mi. S Coahoma, 1 (UM) ; 5 mi. W Big Springs,
2400 ft., 1 (MVZ). Mitchell County: Colorado, 5 (USBS). Winkler County: 20 mi. N
Monahans, 1 (USBS). Ward County: Monahans, 1 (USBS).
Setzer: Subspeciation in Kangaroo Rat 521
Dipodomys ordii obscurus (Allen)
Perodipus obscurus Allen, Bull. Amer. Mus. Nat. Hist., 19:603, No-
vember 12, 1903.
Dipodomys ordii obscurus, Grinnell, Journ. Mamm, 2:96, May 2, 1921.
Type.— Male, adult, no. 20957, Amer. Mus. Nat. Hist.; Rio Sestin, north-
western Durango, Mexico; obtained on April 13, 1903, by J. H. Batty. (Type
not seen.)
Range. — Northwestern and northern Durango, Mexico; marginal localities
are: Rosario, Rio Sestin, Mt. San Gabriel, Rio del Bocas, Villa Ocampo.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face {16") between Pinkish Cinnamon and Cinnamon-Buff, purest on sides,
flanks and cheeks, upper parts strongly suffused with black; arietiform mark-
ings, plantar surfaces of hind feet, pinnae of ears, dorsal and ventral stripes
of tail, brownish. Skull of medium size, nasals long and flared distally; ros-
trum long and narrow; interorbital region relatively narrow; auditory bullae
less inflated than in Dipodomys ordii palmeri; interparietal region narrow ; zygo-
matic arches heavy and bowed laterad; pterygoid fossae ovoid; braincase but
slightly vaulted.
Comparisons. — From Dipodomys ordii palmeri, D. o. obscurus differs in:
Size larger; color lighter; nasals shorter and more flared distally; interorbital
width less; lacrimal processes larger; auditory bullae less inflated; pterygoid
fossae ovoid as opposed to subcircular; zygomatic arches heavier; rostrum
shorter and wider.
From Dipodomys ordii ordii, D. o. obscurus differs as follows: Size smaller;
color darker; skull smaller; nasals longer; rostrum narrower and shorter; in-
terorbital width greater; interparietal region narrower; narrower across audi-
tory bullae; zygomatic arches heavier and more bowed laterally; pterygoid
fossae more ovoid; breadth across maxillary arches greater; external auditory
meatus smaller.
With Dipodomys ordii attenuatus and Dipodomys ordii sennetti, D. o.
obscurus needs no comparison since it is larger and darker than either of those
subspecies and can readily be told from the latter by the greater expansion
of the auditory bullae.
For comparison with Dipodomys ordii juscus see account of that subspecies.
Remarks. — D. o. obscurus seemingly is not a far-ranging subspe-
cies. The only examples referable to it come from a relatively re-
stricted area of Durango, Mexico. One specimen from Rio del
Bocas, Durango, is not typical and shows the characters described
for the animals from Chihuahua City and from Casas Grandes. I
have considered the possibility that this specimen is an intergrade
between D. o. obscurus and an unnamed subspecies ranging to the
northeastward. The other specimens in the series from Rio del
Bocas are typical of D. o. obscurus.
Specimens examined. — Total, 69, all from Durango, distributed as follows: Rosario, 20
(AMNH); Villa Ocampo, 5 (AMNH) ; Rio Sestin, 30 (28 AMNH; 2 CNHM) ; Mt. San
Gabriel, 2 (AMNH); Rio del Bocas, 11 (AMNH); Rancho Santuario, 1 (AMNH).
4—6114
522
University of Kansas Publs., Mus. Nat. Hist.
105
® TYPE LOCALITY
• SPECIMEN EXAMINED
A ADDITIONAL RECORD
Fig. 24. Known occurrences and probable geographic range of the subspe-
cies of Dipodomys ordii in the southeastern fourth of the range of the species.
1. D.o. richardsoni 7. D.o. obscurus 30. D.o. juscus
2. D. o. oklahomae 14. D. o. ordii
3. D.o. compactus 16. D.o. extractus
4. D.o. sennelti 23. D.o.idoneus
6. Z). o. medium 29. D. o. attenuatus
31. D.o. longipes
35. D.o. palmeri
Setzer: Subspeciation in Kangaroo Rat 523
Dipodomys ordii terrosus Hoffmeister
Dipodomys ordii terrosus Hoffmeister, Proc. Biol. Soc. Washington,
55:165, December 31, 1942.
Dipodomys phillipsi ordi, Coues and Allen, Monogr. North American
Rodentia, p. 541, August, 1877 (part — the part from Yellowstone River,
Montana).
Perodipus montanus richardsoni, Cary, N. Amer. Fauna, 49:124, De-
cember, 1926 (part — the part from Glendive, Montana).
Type. — Male, no. 93477, Mus. Vert. Zool., Univ. California; Yellowstone
River, 5 mi. W Forsyth, 2,750 ft., Rosebud County, Montana; obtained on
June 2, 1940, by J. R. Alcorn, original no. 1528.
Range. — Extreme southwestern Saskatchewan and southeastern Alberta,
eastern half of Montana, northern Wyoming and probably extreme western
North Dakota; marginal localities are: 50 mi. W Swift Current, Saskatche-
wan; "near Medicine Hat," Alberta; in Wyoming, Sheep Creek, and 23 mi.
SW Newcastle; in Montana, Medicine Rocks (14 mi. NE Ekalaka), and
Glendive.
Diagnosis. — Size large (see measurements). Color dark, entire dorsal sur-
face near (c) Ochraceous-Buff, purest on sides and flanks; upper parts mixed
with black; arietiform markings, pinnae of ears, dorsal and ventral stripes of
tail and plantar surfaces of hind feet brownish-black. Skull large; rostrum
short, wide and deep; braincase slightly vaulted; auditory bullae markedly
inflated ventrally; zygomatic arches heavy and bowed laterad; upper incisors
long and robust.
Comparisons. — From Dipodomys ordii priscus, D. o. terrosus differs as fol-
lows: Size larger, in all measurements taken, except for length of hind feet,
which is less; color darker in all pigmented areas; skull larger in all parts
measured except width of interparietal, which is less; auditory bullae more
inflated ventrally; zygomatic processes of maxillae wider; rostrum deeper and
shorter.
From Dipodomys ordii richardsoni, D. o. terrosus differs as follows: Size
larger; color darker in all pigmented areas; ventral stripe of tail extending
farther distally; skull larger except in width across auditory bullae, which is
the same.
For comparison with Dipodomys ordii luteolus see account of that sub-
species.
Remarks. — As noted in the comparisons, D. o. terrosus is larger
and darker than D. o. priscus, D. o. luteolus or D. o. richardsoni, its
closest geographic neighbors, and does not resemble any of them,
but rather resembles D. o. longipes and D. o. evexus in size and ap-
pearance, both of which are distantly removed geographically.
Like other subspecies of the species D. ordii, D. o. terrosus prefers
sandy soils to those of any other type. Two miles east and 1 mile
south of Forsyth, Montana, animals were trapped on lenses of
sandy soil. These lenses alternated with areas of black loam of sim-
ilar size. It was noteworthy that burrows were found only in the
areas of sandy soil, although paths used by the rats when foraging
524: University of Kansas Publs., Mus. Nat. Hist.
did extend onto and several crossed the lenses of black loam. We
were not permitted to excavate any of these burrows, but conver-
sation with farmers of the immediate vicinity indicated that the
burrows were not deep. An eight-inch disc would frequently plow
out nests and food caches. It was said that each of several caches
contained as much as a peck of wheat.
Intergradation was noted in animals from 23 miles southwest of
Newcastle and Arvada, Wyoming. In animals from both localities
the pterygoid fossae are more as in D. o. luteolus but referable
to D. o. terrosus. The specimens from Arvada, although immature,
possessed cranial characters which were intermediate between those
of D. o. terrosus and D. o. luteolus but the specimens are referable
to the former.
Specimens examined. — Total, 74, distributed as follows:
Montana: Petroleum County: 24 mi. N Roundup, 8 mi. SW Flatwillow, 2 (UM).
Garfield County: Jordan, 10 (1 UM; 2 MVZ ; 7 AMNH). Dawson County: Glendive, 9
(USNM). Musselshell County: Harvey Ranch, Melstone, 3 (MVZ). Rosebud County: Yel-
lowstone River, 5 mi. W Forsyth, 2750 ft., 7 (MVZ); 2 mi. E and 1 mi. S Forsyth, 2600
ft., 8 (KU). Custer County: Miles City, 1 (USBS). Yellowstone County: Billings, 2 (1
USBS; 1 MVZ). Big Horn County: Fort Custer, 1 (USBS); Crow Agency, 1 (USBS).
Powder River County: Powderville, 4 (USBS). Carter County: Medicine Rocks, 16 mi. N
Ekalaka, 2 (USBS); Medicine Rocks, 14 mi. N Ekalaka, 2 (USBS); Clark's Fork, 1 (USBS).
Wyoming: Big Horn County: Dry Creek, 10 mi. W Germania, 1 (USBS); 3 mi. E
Germania, 1 (USBS); Greybull, 2 (USBS); Bighorn Basin, 1 (USBS). Sheridan County:
Arvada, 8 (USBS). Campbell County: Little Powder River, 1 (USBS). Weston County:
Newcastle, 2 (USBS); 23 mi. SW Newcastle, 4 (USBS). Fremont County: Wilson's Ranch,
Sheep Creek, S base Owl Creek Mountains, 1 (USBS).
Additional records. — Canada (Anderson, 1946: 131): Alberta: near Medicine Hat, 1;
Saskatchewan: near Shackleton, 45-50 mi. NW Swift Current, 1; near Tompkins, 50 mi. W
Swift Current, 1.
Dipodomys ordii fremonti Durrant and Setzer
Dipodomys ordii fremonti Durrant and Setzer, Bull. Univ. Utah, 35
(no. 26) :21 June 30, 1945.
Type. — Female, no. 15661, Carnegie Museum, Pittsburgh, Pennsylvania; Tor-
rey, 7000 ft., Wayne County, Utah; obtained on July 19, 1938, by W. F. and
F. H. Wood, original no. 1562.
Range. — Known only from the type locality.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face Cinnamon-Buff, purest on sides, flanks and cheeks; upper parts strongly
suffused with black; arietiform markings, pinnae of ears, plantar surfaces of
hind feet, dorsal and ventral stripes of tail, brownish. Skull small; upper
incisors long; rostrum deep; jugal bowed laterally; diastema long; upper
molariform tooth-row long.
Comparisons. — From Dipodomys ordii panguitchensis, D. o. fremonti differs
in: Color lighter in all pigmented areas, particularly ears which are light
brown in D. o. fremonti and black in D. o. panguitchensis ; skull larger in all
measurements taken; upper incisors longer; rostrum deeper; auditory bullae
deeper; jugal bowed laterally rather than straight; diastema longer.
Setzer: Subspeciation in Kangaroo Rat 525
From Dipodomys ordii cupidineus, longipes, ncxilis, uintensis and sanrajaeli,
D. o. fremonti can readily be distinguished by its smaller size and generally
darker color.
Remarks. — This subspecies of Dipodomys ordii inhabits the upper
reaches of the Fremont River in west-central Wayne County, Utah.
D. o. fremonti appears to be isolated and is known only from the
type locality. D. o. fremonti is so remarkably different from any
other subspecies of Dipodomys ordii that a long period of isolation
from the ancestral stock (which probably gave rise also to Dipodo-
mys ordii utahensis and Dipodomys ordii panguitchensis) is in-
dicated. Although intergradation is not known to occur with other
kinds, differentiation has not progressed far enough for these ani-
mals to be recognized as a distinct species.
The subspecies closest, geographically, to D. o. fremonti is D. o.
cupidineus from which D. o. fremonti differs more than from any of
the other named forms.
Specimens examined. — Total, 9, from Utah, as follows: Wayne County: Torrey, 7000 ft.,
9 (CM).
Dipodomys ordii uintensis Durrant and Setzer
Dipodomys ordii uintensis Durrant and Setzer, Bull. Univ. Utah, 35
(no. 26) :27, June 30, 1945.
Perodipus longipes, Allen, Bull. Amer. Mus. Nat. Hist., 8:246, Novem-
ber 1896 (part — the part from Uncompahgre Indian Reservation, Utah).
Dipodomys ordii luteolus, Moore, Journ. Mamm, 11:88, February 1930
(part — the part from Vernal, Utah).
Type.— Male, adult, no. 11634, Carnegie Museum, Pittsburgh, Pennsylvania;
Red Creek, 6,700 ft., 2 mi. N Fruitland, Duchesne County, Utah ; obtained on
August 15, 1936, by J. K. and M. T. Doutt, original no. 3433.
Range. — Uinta basin of the White, Green and Duchesne river drainage in
northeastern Utah; marginal occurrences are: 2 mi. N Fruitland, 10 mi. S
Ouray, Vernal.
Diagnosis. — Size large (see measurements); hind foot short. Color dark;
entire dorsal surface, near (c) Cinnamon-Buff, purest on sides and flanks, with
moderate suffusion of black on upper parts ; cheeks white ; arietiform markings,
pinnae of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail,
brownish. Skull large; frontomaxillary suture convex mediad; lacrimal proc-
ess large; styloid process projects, on ventral surface of tympanic bulla, beyond
middle of external auditory meatus; nasals flared distally.
Comparisons. — From Dipodomys ordii priscus, D. o. uintensis differs in:
Hind foot shorter; color darker; styloid process projects on ventral part of
tympanic bulla well anterior to middle of external auditory meatus as opposed
to projecting to middle; depth of foramen magnum, expressed in percentage
of width across posterior margin of occipital condyles, greater (86 per cent in
D. o. uintensis and 81 per cent in D. o. priscus) ; frontomaxillary suture convex
mediad as opposed to nearly straight; lacrimal processes larger; nasals more
flared distally.
526 University of Kansas Publs., Mus. Nat. Hist.
From Dipodomys ordii nexilis, D. o. uintensis differs as follows: Size
smaller; color lighter; interorbital breadth greater; frontom axillary suture
convex mediad as opposed to concave; lacrimal processes larger; nasals more
flared distally; narrower across auditory bullae; basal length greater; zygo-
matic arches bowed laterad as opposed to relatively straight.
From Dipodomys ordii longipes, D. o. uintensis differs as follows: Size
smaller; color darker; auditory bullae wider, longer and deeper; frontomaxillary
suture convex mediad as opposed to nearly straight; greatest breadth across
auditory bullae less.
For comparison with Dipodomys ordii sanrajaeli see account of that sub-
species.
Remarks. — This large, rather dark race inhabits the desert valleys
of the White, Green and Duchesne rivers in northeastern Utah. The
race nearest geographically, as well as morphologically, is Dipod-
omys ordii prisons. Intergradation occurs with the latter subspecies
at Vernal, Uintah County, in cranial measurements and in color.
On the basis of color alone D. o. uintensis can be distinguished from
D. o. sanrajaeli, the geographic race to the south. Specimens from
Jensen are intermediate in color and cranial measurements between
Dipodomys ordii nexilis and D. o. uintensis but are referable to the
latter.
Specimens examined. — Total, 40, all from Utah, distributed as follows: Duchesne County:
Red Creek, 6700 ft., 2 mi. N Fruitland, 4 (CM); 10 mi. S Myton, 1 (UU); 20 mi. S My-
ton, 1 (RH). Uintah County: Vernal, 1 (BYU); 20 mi. E Ouray, 5 (CM); Junction Green
and White rivers, 4800 ft., 2 mi. S Ouray, 5 (CM) ; Pariette Bench, 5000 ft., 8 mi. S Ouray,
8 (CM); Desert Springs, 10 mi. S Ouray, 4 (CM); Pariette Bench, 12 mi. S Ouray, 2 (CM);
Jensen, 5 (BYU); E side Green River, 3 mi. S Jensen, 4 (CM).
Dipodomys ordii sanrafaeli Durrant and Setzer
Dipodomys ordii sanrafaeli Durrant and Setzer, Bull. Univ. Utah, 35
(no. 26) :26 June 30, 1945.
Dipodomys ordii longipes, Stanford, Journ. Mamm., 12:360, November,
1931 (part — the part from King's Ranch, Utah).
Type. — Female, adult, no. 4612, Museum of Zoology, University of Utah;
\y/<2, mi. N Price, 5567 ft., Carbon County, Utah; obtained on June 5, 1940,
by Ross Hardy and H. Higgins, original no. 1901.
Range. — East-central Utah, east into west-central Colorado. Marginal oc-
currences are: in Utah, 12 mi. E Price, 1% mi. N Price, Notom, King's
Ranch, 12 mi. SW Green River, 16 mi. NW Moab; in Colorado, State Line
and Grand Junction.
Diagnosis. — Size large (see measurements). Color dark, entire dorsal sur-
face Cinnamon-Buff, purest on sides and flanks with but slight suffusion of
black on upper parts; cheeks white; arietiform markings, pinnae of ears,
plantar surfaces of hind feet, dorsal and ventral stripes of tail, brownish-black.
Skull large; pterygoid fossae ovoid; lacrimal processes small; width across
maxillary arches relatively great; auditory bullae well inflated; diastema short.
Comparisons. — From Dipodomys ordii longipes, D. o. sanrafaeli differs as
follows: Size smaller; color lighter, more cinnamon, pinnae of ears lighter;
Setzer: Subspeciation in Kangaroo Rat 527
skull smaller; auditory bullae smaller; pterygoid fossae ovoid rather than
round; wider across occipital condyles; narrower across zygomatic processes
of maxillae.
From Dipodomys ordii cupidineus, D. o. sanrafaeli can be recognized by its
larger size, lighter color and larger skull.
For comparisons with Dipodomys ordii nexilis, Dipodomys ordii priscus and
Dipodomys ordii uintensis see accounts of those subspecies.
Remarks. — Inter-gradation between Dipodomys ordii cupidineus
and D. o. sanrafaeli is noted in the intermediate size of body in a
single specimen from Notom. Intergradation in color and cranial
characters occurs between Dipodomys ordii nenlis and D. o. sanra-
faeli in specimens from 16 miles northwest of Moab. All these
specimens, however, are referable to D. o. sanrafaeli.
Animals from that part of the range of D. o. sanrafaeli west of
the Green River are typical while those to the east of the river are
all intergrades. Animals from 16 miles northwest of Moab, Utah,
and from three localities in Colorado, even though intergrades with
D. o. nexilis, are all referable to D. o. sanrafaeli. It appears that
the Green River does not act as a complete barrier in this area since
in the winter it occasionally freezes over, thus allowing the animals
to cross. It is thought that kangaroo rats do not hibernate but
remain more or less active throughout the winter. Man-made con-
veniences, such as bridges, might also serve as means of dispersal,
permitting these animals to cross otherwise prohibitive barriers.
Where there are no bridges across the Green River, farther to the
south, the rats apparently do not cross the river ; steep, rocky canyon-
walls and the lack of ice on the water in winter lessen the chances
of small mammals crossing from one side to the other.
Specimens examined. — Total, 30, distributed as follows:
Utah: Carbon County: 12 mi. NE Price, 2 (CM); 3 mi. NE Price. 1 (RH) ; 1% mi.
N Price, 2 (1 RH; 1 UU) ; Wellington, 1 (RH). Emery County: "San Rafael, 21 mi. out,"
1 (USAC); 12 mi. SW Green River, 2 (CM). Grand County: 1 mi. E Green River, 1
(MVZ) ; 16 mi. NW Moab, 2 (CM). Wayne County: Notom, 1 (BYU). Garfield County:
King's Ranch, 4800 ft., 3 (2 UU ; 1 USAC).
Colorado: Mesa County: State Line, 11 (MVZ); Fruita, 1 (USBS) ; Grand Junction,
2 (USBS).
Dipodomys ordii panguitchensis Hardy
Dipodomys ordii panguitchensis Hardy, Proc. Biol. Soc. Washington,
55 :90, June 25, 1942.
Type.— Male, adult, no. 4375, Museum of Zoology, University of Utah; one
mile south of Panguitch, 6666 ft., Garfield County, Utah; obtained on August
31, 1940, by Ross Hardy, original no. 2151.
Range. — Known only from the type locality.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face near Olive-Brown, purest on sides and flanks, upper parts strongly suf-
528 University of Kansas Publs., Mus. Nat. Hist.
fused with black; cheeks white; arietiform markings, pinnae of ears, plantar
surfaces of hind feet, dorsal and ventral stripes of tail which are wider than
white lateral stripes, blackish. Skull small; rostrum relatively short and wide;
interorbital region wide; interparietal region wide; foramen magnum elongate
dorsoventrally ; pterygoid fossae ovoid.
Comparisons. — From Dipodomys ordii utahensis, which it closely resembles,
D. o. panguitchensis differs in: Size larger; color darker; interparietal region
wider; foramen magnum elongate dorsoventrally as opposed to nearly round;
pterygoid fossae ovoid as opposed to nearly round.
This subspecies can be distinguished from Dipodomys ordii fetosus, Dipodo-
mys ordii celeripes and Dipodomys ordii cupidineus by its darker color and
generally larger size.
For comparisons with Dipodomys ordii cinder ensis and Dipodomys ordii
jremonti see accounts of those subspecies.
Remarks. — This geographic race inhabits the upper reaches of
the Sevier1 River Valley in the vicinity of Panguitch, Utah. Natural
barriers to kangaroo rats, such as the Cedar Mountains to the west,
high plateau country to the south, the Paunsaugunt Plateau to the
east and the narrow canyons of the Sevier River to the north pre-
vent these animals from extending their range or from coming into
physical contact with any adjacent geographic races. This isola-
tion has resulted in a fairly stable population. Some animals,
however, show characters, such as the width of the rostrum, and
the shape and length of the nasals which are intermediate between
those of topotypes of D. o. utahensis and the type series of D. o.
panguitchensis.
Specimens examined. — Total, 3, all from Utah, distributed as follows: Garfield County:
1 mi. S Panguitch, 6666 ft., 3 (2 RH; 1 UU).
Dipodomys ordii monoensis (Grinnell)
Perodipus monoensis Grinnell, Univ. California Publ. Zool., 21 :46,
March 29, 1919.
Dipodomys ordii monoensis, Grinnell, Journ. Mamm., 2:96, May 2,
1921.
Type. — Female, adult, no. 27002, Museum of Vertebrate Zoology, Univer-
sity of California; Pellisier Ranch, 5 mi. N Benton Station, 5600 ft., Mono
County, California; obtained on September 21, 1917, by J. Dixon, original
no. 6384.
Range. — Northeastern Inyo and Mono counties, California, north to south-
ern Pershing County and east to eastern Nye County, Nevada; marginal oc-
currences are: in California, 5 mi. N Benton Station and Deep Spring Valley;
in Nevada, Arlemont, 2 mi. NW Palmetto, 1 mi. N Beatty, 5 mi. W White
Rock Spring, Big Creek at Quinn Canyon Mts., 2% mi. S Lock's Ranch, 2
mi. S Millett P. O., 13V2 mi. NW Goldfield, Fingerrock Wash, Eastgate,
Vi mi. NE Toulon, 21 mi. W and 2 mi. N Lovelock, V2 mi. S Pyramid Lake,
West Walker River in Smith's Valley, and 10 mi. S Yerington.
Setzer: Subspeciation in Kangaroo Rat 529
Diagnosis. — Size medium (see measurements). Color pale, entire dorsal
surface (c) between Pinkish Buff and Cinnamon-Buff, purest on sides, flanks
and cheeks, with but slight suffusion of black in upper parts; pinnae of ears,
plantar surfaces of hind feet, dorsal and ventral stripes of tail, brownish.
Skull medium in size; rostrum relatively long and narrow; nasals relatively
short; interorbital region narrow; interparietal region relatively wide; lacrimal
processes small; auditory bullae relatively small; pterygoid fossae circular;
zygomatic arches robust and relatively straight; foramen magnum nearly
circular.
Comparisons. — From Dipodomys ordii columbianus, D. o. monoensis differs
as follows: Size larger; color lighter; skull larger; rostrum longer and nar-
rower; interorbital region narrower; breadth across auditory bullae less; lacri-
mal processes larger; braincase less vaulted; auditory bullae more inflated
ventrally; pterygoid fossae smaller; zygomatic arches more robust; cutting
edge of upper incisors wider.
From Dipodomys ordii jetosus, D. o. monoensis differs in: Hind foot shorter;
color lighter; skull smaller; rostrum shorter and narrower; interorbital width
less; interparietal region larger; lacrimal processes smaller; auditory bullae
less inflated.
For comparison with Dipodomys ordii inaquosus see account of that sub-
species.
Remarks. — This subspecies retains all of its diagnostic characters
throughout nearly all parts of its geographic range. Intergradation
occurs in animals from the southern end of Pyramid Lake, Big
Smoky Valley and near Toquima Peak, Nevada; these animals, al-
though typical of D. o. monoensis in coloration, resemble D. o.
columbianus cranially. Three-way intergradation between D. o.
columbianus, D. o. jetosus and D. o. monoensis is noted in animals
from east-central Nye County, Nevada. These animals resemble
D. o. monoensis in size, D. o. jetosus in color and resemble D. o.
columbianus in certain cranial features. These animals are referred
to D. o. monoensis. Animals from Toulon, Nevada, in the inflation
of the auditory bullae, the vault of the braincase, the color and the
total length show intergradation with D. o. inaquosus but are refer-
able to D. o. ??ionoensis.
Specimens examined. — Total, 264, distributed as follows:
California: Mono County: Pellisier Ranch, 5 mi. N Benton Station, 17 (7 DRD; 10
MVZ); Benton, 5639 ft., 2 (1 LACM ; 1 MVZ); Taylor Ranch, 2 mi. S Benton Station,
5300 ft., 2 (MVZ). Inyo County: Deep Springs Valley, 1 (LACM).
Nevada: Washoe County: y2 mi. S Pyramid Lake, 3950 ft., 1 (MVZ); 1% mi. N
Wadsworth, 4100 ft., 2 (MVZ). Pershing County: 21 mi. W and 2 mi. N Lovelock, 4000
ft., 2 (MVZ); 3% mi. NNE Toulon, 3900 ft., 1 (MVZ); 3 mi. NNE Toulon, 3900 ft., 6
(MVZ); Vi mi. NE Toulon, 1 (MVZ); Toulon, 3930 ft., 5 (MVZ). Churchill County:
Truckee Canal, 2 mi. SW Hazen, 4000 ft., 1 (MVZ); 1 mi. NW Soda Lake, 4000 ft., 2
(MVZ); 1 mi. S Soda Lake, 4000 ft., 1 (MVZ); 5 mi. W Fallon, 1 (MVZ); 4 mi. W Fallon,
4000 ft., 3 (MVZ); 1 mi. W Mountain Well, 5350 ft., 3 (MVZ); Eastgate, 4400 ft., 13
(MVZ). Lyon County: 6 mi. N Fernley, 1 (MVZ); 1 mi. SE Wadsworth, 4200 ft., 7
(MVZ) ; % mi. N Fernley Underpass, Fernley, 1 (MVZ) ; y2 mi. N Fernley Underpass, Fern-
530 University of Kansas Publs., Mus. Nat. Hist.
ley, 1 (MVZ); Wilson Canyon, 8 mi. NE Wellington, 4700 ft., 1 (MVZ); West Walker River,
Smith's Valley, 4700 ft., 4 (MVZ); 10 mi. S Yerington, Mason Valley, 4500 ft., 6 (MVZ).
Mineral County: 8 mi. SE Schurz, 4100 ft., 18 (MVZ); Fingerrock Wash, Stewart Valley,
5400 ft., 4 (MVZ); Cat Creek, 4 mi. W Hawthorne, 4500 ft., 1 (MVZ); Huntoon Valley,
5700 ft., 1 (MVZ). Nye County: 2 mi. S Millett P. O., 5500 ft., 1 (MVZ); 4 mi. SE Mil-
lett P. O., 5500 ft., 11 (MVZ); 5 mi. SE Millett P. O., 5500 ft., 5 (MVZ); 4 mi. S Millett
P. O., 5500 ft., 2 (MVZ); Millman Ranch, Moore Creek, 6400 ft., 19 mi. SE Millett P. O.,
9 (MVZ); Meadow Creek Ranger Station, Toquima Mts., 2 (MVZ); Monitor Valley, 9 mi.
E Toquima Mts., 7000 ft., 19 (MVZ); Fish Spring Valley, Y2 mi. N Fish Lake, 6500 ft.,
2 (MVZ); Railroad Valley, 2% mi. S Lock's Ranch, 5000 ft., 5 (MVZ); Hot Creek Valley
3% mi. E Hot Creek, 5650 ft., 1 (MVZ); Hot Creek Valley, % mi. S Hot Creek, 5900 ft.,
1 (MVZ); 5y2 mi. NE San Antonio, 5700 ft., 1 (MVZ); San Antonio, 5400 ft., 2 (MVZ);
9 mi. W and 3 mi. S Tybo, 6200 ft., 2 (MVZ); Ralston Valley, 15% mi. NE Tonopah, 5800
2 (MVZ); Railroad Valley, 2% mi. S Lock's Ranch, 5000 ft., 5 (MVZ); Hot Creek Valley
Creek, 5800 ft., 1 (MVZ); Ralston Valley, 34 mi. E and 1 mi. N Tonopah, 5650 ft., 2
(MVZ); Old Mill, N end Reveille Valley, 6200 ft., 6 (MVZ); 1% mi. S Silverbow, Kawich
Mountains, 1 (MVZ); 5 7/10 mi. SE Kawich, 2 (MVZ); 5 mi. W White Rock Spring, 6950
ft., Belted Range, 2 (MVZ); 1 mi. N Beatty, 1 (DRD). Esmeralda County: UY2 mi. NW
Goldfield, 4850 ft., 3 (MVZ); 7 mi. N Arlemont, 5500 ft., 6 (MVZ); Arlemont, 11 (MVZ);
Mouth Palmetto Wash, 7 (DRD); 2 mi. NW Palmetto, 7 (DRD); 1 mi. NW Palmetto, 1
(DRD); Palmetto, 7 (DRD); 1 mi. SE Palmetto, 7 (DRD); Pigeon Spring, 6400 ft., 1
(MVZ); Indian Spring, Mt. Magruder, 20 (DRD).
Dipodomys ordii ordii Woodhouse
D(ipodomys) ordii Woodhouse, Proc. Acad. Nat. Sci. Philadelphia,
6:224, 1853.
Dipodomys phillipsi ordi, Coues and Allen, Monogr. North American
Rodentia, p. 541, 1877 (part — the part from El Paso, Texas).
Dipodops ordii, Merriam, N. Amer. Fauna, 4:42, October, 1890 (part —
the part from El Paso, Texas).
Cricetodipus ordii, Trouessart, Catalogus Mammalium, 1 :581, 1897.
Perodipus ordi, Elliot, Field Columbian Museum, Zool. Ser., 2:238,
1901.
Perodipus montanus richardsoni, Bailey, N. Amer. Fauna, 25:144, Oc-
tober, 1905 (part — the part from Carlsbad, New Mexico).
Perodipus ordii, Goldman, Proc. Biol. Soc. Washington, 30:113, May
23, 1917.
Type. — None designated. Species characterized from specimens obtained
by Dr. Woodhouse at El Paso, Texas.
Range. — Southeastern Arizona, southern New Mexico, western Texas and
north-central Mexico; marginal occurrences are: in Arizona, 20 mi. NE Calva,
Oracle and Calabasas; in Sonora, Nogales; in Chihuahua, Casas Grandes, Cor-
ralitos and Santa Rosalia; in Texas, 16 mi. E Van Horn and 30 mi. N Van
Horn; in New Mexico, 40 mi. N Roswell, 40 mi. SE Corona and Mangos
Valley.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face {16") between Pinkish Cinnamon and Cinnamon-Buff, purest on sides
and flanks, upper parts suffused with black; arietiform markings, pinnae of
ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail, brownish-
black. Skull of medium size; rostrum narrow and relatively long; braincase
flattened; auditory bullae but slightly inflated; zygomatic arches slender and
straight; upper incisors short and narrow.
Comparisons. — From Dipodomys ordii richardsoni, D. o. ordii differs as
follows: Size smaller; color somewhat lighter; dorsal and ventral stripes of
tail brownish instead of blackish; skull smaller in all measurements taken.
Setzer: Subspeciation in Kangaroo Rat 531
From Dipodomys ordii montanus, D. o. ordii differs in: Size smaller; color
lighter; skull narrower across bullae and rostrum; wider interorbitally ; all
other measurements taken the same, thus imparting to the skull, in dorsal
view, a longer, narrower appearance.
From Dipodomys ordii chapmani, D. o. ordii differs in: Size smaller; color
lighter; skull smaller; zygomatic arches narrower at suture of jugal and zygo-
matic process of maxillary; rostrum narrower; upper incisors smaller; nasals
shorter; least interorbital width less.
For comparisons with Dipodomys ordii sennetti and Dipodomys ordii com-
pactus see accounts of those subspecies.
Remarks. — Intergradation occurs with Dipodomys ordii chap-
mani, Dipodomys ordii longipes, Dipodomys ordii medius and Di-
podomys ordii attenuatus. Only along the Rio Grande near El Paso,
Texas, are individuals of a population uniform. On the periphery of
the range, specimens from a given locality may resemble D. o. ordii
or D. o. longipes or may be intermediate between these two sub-
species.
Animals from the Organ Mountains near Globe Springs, 20 miles
north of Cliff, New Mexico, and those labeled with reference to Van
Horn, Texas, on the average are darker, have a wider expanse across
the auditory bullae, a wider interorbital region and, in most speci-
mens, more distally flared nasals than the norm of other popula-
tions. This deviation from the normal is interpreted not as inter-
gradation with any other subspecies but rather as individual
variation in a given population.
Specimens from 40 miles southeast of Corona and 10 miles north-
east of Socorro, New Mexico, show intergradation between D. o.
ordii and D. o. montanus in size of body, configuration of nasals and
cranial size. For a discussion of intergradation with D. o. longipes,
D. o. medius and D. o. chapmani see the remarks in the accounts of
those subspecies.
Specimens from Cananea and Santa Cruz, Sonora, Casas Grandes
Viejo, Chihuahua City and Colonia Diaz, Chihuahua, are not typi-
cal of D. o. ordii but rather show the influence of some race probably
to the southeast that is as yet unknown. In a series of nine speci-
mens from near Casas Grandes Viejo, Chihuahua, four individuals
are typically D. o. ordii, four resemble an apparently undescribed
form and one specimen is intermediate between D. o. ordii and the
unnamed subspecies. This unnamed race differs from D. o. ordii in
having darker pelage, darker dorsal stripe on tail, larger body,
wider interorbital region, longer skull, greater breadth across the
bullae, less vaulted braincase, more robust zygomatic arches and
the foramen magnum more deeply notched on both the dorsal and
532 University of Kansas Publs., Mus. Nat. Hist.
ventral rims. It seems that these animals mentioned above should
be referred to D. o. ordii at least until such time as material becomes
available from eastern Chihuahua, northern Coahuila and north-
eastern Durango.
Specimens examined. — Total, 451, distributed as follows:
Arizona: Gila County: Rice, 2 (USBS). Maricopa County: Marinette, 8 (CAS). Pinal
County: Oracle, 4 (USBS). Graham County: 20 mi. NE Calva, 5 (USBS); Fort Grant, 11
(7 USBS; 4 MVZ). Pima County: Ft. Lowell, 3 (DRD) ; 11% mi. S Tucson, 1 (MVZ) ;
Continental, 20 (8 USBS; 12 DRD); La Osa, 11 (USNM) ; Babiquivari Mountains 10 mi.
N International Boundary, 1 (DRD). Santa Cruz County: 2 mi. S Tumacacori Mission, 1
(DRD); 7 mi. N Patagonia, 4500 ft., 12 (CAS); 3 mi. N Patagonia, 1 (CAS); Calabasas,
7 (6 USBS; 1 USNM). Cochise County: 8 mi. W Bowie, 1 (LACM) ; Wilcox, 4163 ft.,
25 (12 USBS; 5 UM; 4 MVZ; 4 USNM); 12 mi. SE Dos Cabezos, 2 (UM); 1 mi. WSW
Chiricahua National Monument, 5000 ft., 1 (MVZ); Portal, 4500 ft., 2 (USBS); Fairbank,
8 (3 AMNH; 5 CNHM); Mouth Pinery Canyon, 4 (USBS).
New Mexico: Torrance County: Gran Quivira, Mesa Jumanes, 6 (USBS). Catron
County: Mangos Valley, 3 (USBS); Alma, 3 (USBS); Pleasanton, 7 (USBS). Socorro
County: Gallina Mountains, 1 (USBS); 10 mi. NE Socorro, 2 (USBS); 3 mi. N Socorro, 3
(MVZ); Socerro, 1 (USBS); Range, 2 mi. SW Socerro, 4700 ft., 2 (MVZ); Lava Mesa; S
Clyde, 4300 ft., 1 (MVZ); Dry Creek, 3 (USBS); San Augustine Plain, 12 mi. N Monica
Springs, 2 (USBS). Lincoln County: Guyo Canyon, 40 mi. SE Corona, 1 (USBS); 4 tni.
W Carrizozo, 2 (UM). Grant County: Gila, 5 (USBS); Cactus Flat, 20 mi. N Cliff, 3
(USBS); Cliff, Gila River, 4470 ft., 1 (USBS); Silver City, 2 (USBS); Redrock, 2 (USBS);
9 mi. N Faywood, 2 (USBS); Hachita, 1 (USBS); Dog Spring, 11 (USNM); Deer Creek,
Culberson Ranch, 2 (USBS). Sierra County: Fairview, 6500 ft., 1 (USBS); Cuchillo, 4700
ft., 3 (USBS); Lake Valley, 5000 ft., 3 (USBS). Otero County: Tularosa, 5 (USBS); 10
mi. SW Tularosa, 2 (CNHM); Quartz Sands, SW Tularosa, 2 (MVZ); White Sands, 12 mi.
W Alamogordo, 1 (MVZ); 2 mi. S Alamogordo, 2 (UM); 3 mi. S Alamogordo, 1 (UM);
5 mi. S Alamogordo, 1 (UM) ; 9 mi. SW Alamogordo, 1 (UM); 12 mi. SW Alamogordo, 2
(UM); 15 mi. SW Alamogordo, 1 (LACM); White Sands, 18 mi. SW Alamogordo, 7 (MVZ);
19 mi. SW Alamogordo, 3 (UM) ; White Sands National Monument, 24 (13 CNHM; 11
LACM); y2 mi. SW Escondido, 4000 ft., 2 (MVZ); Jarilla, 1 (USBS). Hidalgo County:
10 mi. NW Lordsburg, 6 (LACM); 4 mi. NW San Luis Pass, 5200 ft., Animas Valley, 5
(MVZ). Luna County: Deming, 11 (USBS). Dona Ana County: Garfield, 4 (USBS); 6
to 8 mi. NE Las Cruces, 1 (CAS); 15 mi. W Las Cruces, 1 (LACM); 11 mi. W Las Cruces,
2 (CAS); Las Cruces, 3 (USBS); Organ Mountains, near Globe Springs, 2 (USBS); Coe's
Ranch, El Paso Road, 35 mi. N El Paso, Texas, 2 (USBS); V* mi. N Strauss, 2 (CAS);
1 mi. E Strauss, 4100 ft., 7 (MVZ) ; 35 mi. W El Paso, Texas, 2 (USNM) ; 20 mi. W
El Paso, Texas, 1 (USNM); 10 mi. W El Paso, Texas, 1 (USNM); Mexican Boundary, near
monument 5, Lat. 31° 47'; Long. 30° 15', 13 (USNM). Eddy County: 3 mi. NW Carlsbad,
7 (MVZ); 2 mi. E Carlsbad, 1 (KU); Eddy, 10 (USBS).
Texas: El Paso County: 3 mi. NE City Limits, El Paso, 3764 ft., 15 (12 MVZ; 3
TCWC); El Paso, 3 (USBS); near El Paso, 7 (USNM). Hudspeth County: 1 mi. NW old
Fort Hancock, 3900 ft., 3 (MVZ); Fort Hancock, 4 (USNM); 4 mi. NW Sierra Blanca, 1
(LACM). Culberson County: 35 mi. N Van Horn, 5 (TCWC); 30 mi. N Van Horn, 1
(TCWC); 16 mi. E Van Horn, 3 (TCWC); 16 mi. SE Van Horn, 5 (TCWC); Kent, 1
(USBS). Reeves County: 5 mi. E Toyahvale, 1 (USBS). Jeff Davis County: 14 y2 mi. S
Fort Davis, 2 (UM). Presidio County: 10 mi. NE Marfa, 1 (UM).
Sonora: Nogales, 2 (USNM); Spnta Cruz, 4 (USNM); 5 mi. N Cananea, 4750 ft., 4
(MVZ); Alamo Wash, 35 mi. NW Magdalena, 5 (DRD); Sonora, 2 (AMNH).
Chihuahua: 4.3 mi. W Casas Grandes Viejo, 5000 ft., 8 (MVZ); 1.5 mi. W Casas
Grandes Viejo, 1 (MVZ); Casas Grandes, 2 (USBS); Gallego, 1 (USBS); Colonia Diaz, 6
(USBS); Las Trincheras, 9 mi. S by road Boquillos de Condios, 4 (MVZ); Santa Rosalia,
6 (USBS) ; Chihuahua, 7 (USBS) ; 5 mi. SE Chihuahua, 5250 ft., 4 (MVZ) ; Corallitos, 4 (1
USBS; 3 MVZ).
Setzer: Subspeciation in Kangaroo Rat 533
Dipodomys ordii luteolus (Goldman)
Perodipus ordii luteolus Goldman, Proc. Biol. Soc. Washington, 30:112,
May 23, 1917.
Dipodomys phillipsi ordi, Coues and Allen, Monogr. of N. Amer. Ro-
dentia, p. 541, August, 1877 (part — the part from Niobrara River, Ne-
braska) .
Perodipus montanus richardsoni, Cary, N. Amer. Fauna, 33:141, Au-
gust 17, 1911 (part — the part from Sterling, Colorado).
Dipodomys ordii luteolus, Grinnell, Journ. Mamm., 2:96, May 2, 1921.
Type.— Male, adult, no. 160408, U. S. Nat. Mus. Biol. Surv. Coll.; Casper,
Natrona County, Wyoming; obtained on September 2, 1909, by Merritt Cary,
original no. 1690.
Range. — Southeastern Wyoming, northeastern Colorado, northwestern half
of Nebraska and southwestern South Dakota; marginal occurrences are: in
Wyoming, Casper, Sun and Ft. Steele; in Colorado, Loveland, Hugo and
Akron; in Nebraska, Birdwood Creek, Neligh and Valentine; in South Dakota,
Batesland and Buffalo Gap.
Diagnosis. — Size medium (see measurements). Color light, entire dorsal sur-
face between Light Ochraceous-Buff and Ochraceous-Buff, purest on sides and
flanks; upper parts but lightly washed with black; arietiform markings, plantar
surfaces of hind feet, pinnae of ears and dorsal and ventral stripes of tail,
blackish. Skull medium in size; jugal weak; braincase slightly inflated; nasals
slightly flared distally.
Comparisons. — From Dipodomys ordii terrosus, D. o. luteolus differs as fol-
lows: Size smaller, except that tail and ear are longer; color lighter in all
pigmented areas; skull smaller in every measurement taken; auditory bullae
less inflated; zygomatic processes of maxillae smaller; cutting edge of upper
incisors narrower; zygomatic arch weaker.
From Dipodomys ordii priscus, D. o. luteolus differs in: Size larger except
hind foot which is shorter; dorsal and ventral stripes of tail, plantar surfaces
of hind feet, arietiform markings and pinnae of ears, in most specimens, darker;
auditory bullae less inflated; nasals shorter; rostrum wider; total length of
skull shorter; zygomatic arch weaker; foramen magnum more ovate.
From Dipodomys ordii evexus, D. o. luteolus differs as follows: Hind foot
longer; color lighter in all pigmented areas; auditory bullae more inflated;
pterygoid fossae more expanded laterally; width across maxillary arches less;
interorbital region narrower; zygomatic arch weaker; external auditory meatus
almost round as opposed to ovoid.
Comparison with Dipodomys ordii richardsoni is made in account of that
subspecies.
Remarks. — Dipodomys ordii luteolus resembles D. o. priscus in
size and color but can readily be told from it and D. o. richardsoni
when specimens from the central portions of the ranges of the sub-
species are compared. At and near the periphery of the range, es-
pecially in that part which adjoins the range of D. o. richardsoni,
intergradation occurs. Specimens from Kennedy, Perch and Neligh,
Nebraska, approach D. o. richardsoni in the shape of the pterygoid
534 University of Kansas Publs., Mus. Nat. Hist.
fossae and nasal bones, but in all other characters they resemble
D. o. luteolus to which subspecies they are here referred. Speci-
mens from Loveland and 20 miles east of Avalo, Colorado, show
intergradation with D. o. richardsoni in the width of the rostrum
and size of the zygomatic arch but are referable to D. o. luteolus.
No specimens here referred to D. o. luteolus were found to inter-
grade with D. o. priscus.
Specimens examined.— Total, 250, distributed as follows:
Sonth Dakota: Perkins County: 9 mi. N Bison, 8 (MVZ). Meade County: Smiths-
ville, 2 (USBS). Jackson County: 20 mi. SSE Phillip, in Haakon County, 1 (MVZ). Custer
County: Elk Mountain, 1 (MHS) ; Buffalo Gap, 2 (USNM). Bennett County: Big Spring
Canyon, Batesland, 7 (CNHM) ; Rosebud Indian Agency, 1 (USBS).
Wyoming: Fremont County: 2% mi. W Shoshoni, 80 (KU): Granite Mountain, 1
(UM). Natrona County: 1 mi. NE Casper, 19 (KU); Casper, 7 (USBS); Sun, 3 (USBS).
Converse County: Douglas, 2 (USBS). Niobrara County: Van Tassel Creek, 1 (CM). Carbon
County: Fort Steele, 1 (USBS). Goshen County: Rawhide Butte, 1 (USBS).
Nebraska: Sioux County: Warbonnet Canyon, 6 (MHS); Glen, 1 (MHS); Agate, 3
(1 UM; 2 CNHM); Jim Creek, 1 (MHS). Box Butte County: 16 mi. NE Alliance, 4
(UM). Sheridan County: 12 mi. S Gordon, 3 (MVZ); 5 mi. N Antioch, 4 (UM) ; 4 mi. N
Antioch, 3 (UM). Cherry County: Valentine, 9 (6 USBS; 2 USNM; 1 NU) ; 15 mi. E
Gordon, 1 (CNHM); Niobrara River, 3 (USBS); 18 mi. NW Kennedy, 3 (2 USBS; 1
MHS); Valentine Lake Refuge, 1 (NU); Kennedy, 10 (3 MVZ; 5 USBS; 2 USNM); 30 mi.
5 Wood Lake, 4 (CNHM); Cherry, 7 (NU); near Clark's Canyon, 7 (USBS). Rock County:
Perch, 12 (10 AMNH ; 2 CNHM). Antelope County: Neligh, 8 (1 USBS; 7 MHS).
Scotts Bluff County: Mitchell, 1 (MHS). Hooker County: Kelso, 5 (UM). Thomas
County: Halsey, 1 (MHS); Dismal River, 2 (USBS). Custer County: 1 mi. NW Gavin,
6 (2 MVZ; 4 UM) ; Gavin, 3 (UM); 8 mi. NW Anselmo, 1 (UM) ; 7 mi. NW Anselmo,
1 (UM). Lincoln County: Birdwood Creek, 2 (USBS); Brady, 4 (NU); Hackberry Lake,
13 (UM).
Colorado: Larimer County: Loveland, 8 (USBS). Weld County: 20 mi. E Avalo,
3 (USBS); 8 mi. E Pawnee Buttes, 1 (USBS); Greeley, 1 (USBS). Morgan County: 30 mi.
S Fort Morgan, 1 (UM). Logan County: Sterling, 8 (USBS). Adams County: Barr Lake,
4 (CMNH); 6 mi. E and 1 mi. N Denver, 1 (CMNH); 3 mi. NE Fitzsimmons, 1 (CMNH).
Washington County: Akron, 2 (UM). Lincoln County: Limon, 1 (USBS); Hugo, 1
(USBS). El Paso County: Colorado Springs, 2 (MVZ).
Dipodomys ordii extractus new subspecies
Type.— Male, adult, no. 76562, Museum of Vertebrate Zoology, University
of California; 1 mi. E Samalayuca, 4500 ft., Chihuahua, Mexico; obtained on
May 15, 1937, by William B. Richardson, original no. 2148.
Range. — Known only from the type locality.
Diagnosis. — Size medium (see measurements). Color pale, entire dorsal
surface Pinkish Buff, purest on sides and flanks, dorsal surface but slightly suf-
fused with black; pinnae of ears, plantar surfaces of hind feet, dorsal and
ventral stripes of tail light-brownish. Skull medium in size; nasals long;
rostrum relatively as well as actually wide; least interorbital breadth wide;
auditory bullae relatively little inflated; braincase but slightly vaulted; ex-
ternal auditory meatus ovoid; zygomatic arches relatively robust and but
slightly bowed laterad; pterygoid fossae subcircular; lacrimal processes small;
foramen magnum deeply notched on dorsal border.
Comparisons. — From Dipodomys ordii ordii, D. o. extractus differs as
follows: Size larger; color lighter; arietiform marks absent in D. o. extractus
but pronounced in D. o. ordii; skull larger; rostrum wider; nasals longer;
Setzer: Subspeciation in Kangaroo Rat
535
• SPECIMEN EXAMINED
A ADDITIONAL RECORD
® TYPE LOCALITY
Fig. 25. Known occurrences and probable geographic range of the subspe-
cies of Dipodomys ordii in the northeastern fourth of the range of the species.
1. D.o. richardsoni 8. D.o. terrosus 18. D.o. montanus
2. D.o. oklahomae 10. D.o. uintensis 24. D.o. prisons
5. D.o. evexus 11. D.o. sanrajaeli 31. D.o. longipes
6. D.o. medius 15. D.o. luteolus 33. D.o. nexilis
536 University of Kansas Publs., Mus. Nat. Hist.
maxillary arches wider; auditory bullae more inflated laterally; braincase less
vaulted; pterygoid fossae subcircular as opposed to circular; foramen magnum
more deeply evaginated dorsally and ventrally.
From Dipodomys ordii obscurus, D. o. extractus differs as follows: Size
larger; color lighter; skull larger in all respects, notably in least interorbital
width, greatest length of skull and breadth across auditory bullae; zygomatic
arches less robust and straight as opposed to robust and bowed laterally;
pterygoid fossae subcircular as opposed to circular; auditory bullae more
inflated ventrally.
From Dipodomys ordii idoneus, D. o. extractus differs in: Size larger;
color lighter; skull larger in all measurements taken except least interorbital
width and breadth across maxillary arches which are smaller; nasals longer
and narrower; braincase less vaulted; zygomatic arches less robust and more
nearly straight; auditory bullae less inflated ventrally.
Remarks. — This subspecies of Dipodomys ordii is the palest of
any of the known races from Mexico. It inhabits an area of light-
colored sandy soil at the type locality.
The only subspecies with which D. o. extractus really needs com-
parison is Dipodomys ordii ordii from which it differs as described
above. The series which is herein described contains two individuals
that resemble D. o. ordii cranially and three that resemble it in color
although they are somewhat paler. One specimen, MVZ no. 76560,
is the palest of any of the series, has a short tail and in these respects
resembles the pale form of Dipodomys ordii compactus. It is ap-
parent from the skull of this latter individual that it is not D. o.
compactus but D. o. extractus.
Dipodomys ordii ordii inhabits, in most of its range, dark-colored
soils, whereas D. o. extractus inhabits light-colored soils. Possibly
the differentiation that has occurred is a result of an ecological
separation much as occurs with Perognathus on the white sands and
black lavas of the Tularosa Basin of New Mexico (see Benson,
1933).
Specimens examined. — Total, 14, all from Chihuahua, as follows: 1 mi. E Samalayuca,
4500 ft., 14 (MVZ).
Dipodomys ordii chapmani Mearns
Dipodomys chapmani Mearns, Bull. Amer. Mus. Nat. Hist., 2:291,
February 21, 1890.
Cricetodipus chapmani, Trouessart, Catalogus Mammalium, 1:581,
1897.
Perodipus chapmani, Elliot, Field Columbian Museum, Zool. Ser.,
2:237, 1901.
I'< rodipus ordii chapmani, Goldman, Proc. Biol. Soc. Washington,
30:113, May 23, 1917.
Dipodomys ordii chapmani, Grinnell, Journ. Mamm., 2:96, May 2,
1921.
Setzer: Subspeciation in Kangaroo Rat 537
Cotypes. — Male, no. 2400, and female, no. 2398, Amer. Mus. Nat. Hist.;
Fort Verde, Yavapai County, Arizona; obtained on January 26, 1887, and
October 1, 1885, respectively, by Edgar A. Mearns. (Type not seen.)
Range. — Central Arizona from the Grand Canyon of the Colorado River,
southeastward to, and probably beyond, Fort Verde; marginal occurrences
are: lower end Prespect Valley, Kirkland, Prescott, Camp Verde, Bill
Williams Mountain.
Diagnosis. — Size medium (see measurements). Color dark; entire dorsal
surface between (10") Pinkish Cinnamon and Cinnamon-Buff, purest on sides
and flanks, upper parts strongly suffused with black; arietiform markings,
pinnae of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail,
blackish. Skull medium in size; rostrum long and narrow; nasals long; audi-
tory bullae slightly distended; braincase vaulted.
Comparisons. — For comparisons with Dipodomys ordii cupidineus, Dipod^
omys ordii longipes and Dipodomys ordii ordii see accounts of those sub-
species.
Remarks. — This subspecies of Dipodomys ordii is relatively iso-
lated from other subspecies of the species. The Colorado River with
its deep canyons is interposed between the ranges of D. o. chapmani
and D. o. cupidineus. A high range of mountains separates D. o.
chapmani from the range of D. o. longipes. The only race with
which D. o. chapmani probably comes into contact is D. o. ordii to
the southeast. No specimens are available from the southeastern
part of the range of D. o. chapmani. Animals from the northwestern
part of the range of D. o. ordii, without exception, have dark-colored
pelage which is characteristic of D. o. chapmani. In the shape of
the nasals, vault of the braincase, interorbital width and auditory
bullae, animals from Rice, Arizona, resemble D. o. chapmani, but
in other characters resemble D. o. ordii to which subspecies they
are referred. Animals from 20 miles northeast of Calva, Arizona,
are dark-colored and are intermediate between the dark D. o.
chapmani and the pale D. o. ordii in size of the cranium and body
but are referable to D. o. ordii. In the northwestern part of the
range of D. o. chapmani the animals are not typical in that they
have a shorter, wider rostrum and a heavier zygomatic arch than
topotypes.
Specimens examined. — Total, 90, all from Arizona, distributed as follows: Mohave
County: lower end Prospect Valley, 4500 ft., Grand Canyon, 7 (USBS) ; Kinsman, 9
(LACM). Coconino County: Aubrey Valley, 10 mi. S Pine Spring, 4 (USBS); Bill Wil-
liams Mountain, 1 (USNM). Yavapai County: 3 mi. N Fort Whipple, 11 (USBS); Fort
Whipple, 4 (2 USBS; 2 USNM); Willow Creek, 4 mi. N Prescott, 4 (MVZ) ; 1 mi. W
Camp Verde, 1 (MVZ); Camp Verde. 33 (15 USBS; 3 MVZ; 3 AMNH; 9 LACM; 3
DRD); y2 mi. S Camp Verde, 4 (MVZ); Kirkland, 9 (USBS); Turkey Creek, 3 (USBS).
5— €114
538 University of Kansas Publs., Mus. Nat. Hist.
Dipodomys ordii montanus Baird
Dipodomys montanus Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:334,
April, 1855.
Perodipus montanus, Merriam, Proc. Biol. Soc. Washington, 17:140,
July 14, 1904.
Perodipus ordii montanus, Goldman, Proc. Biol. Soc. Washington,
30:113, May 23, 1917.
Dipodomys ordii montanus, Grinnell, Journ. Mamm., 2:96, May 2,
1921.
Type.— Male, no. 490/1631, U. S. Nat. Mus.; near Fort Massachusetts,
Costilla County, Colorado; obtained in 1853 by F. Kreutzfeldt (Pacific Rail-
road Survey).
Range. — The San Luis Valley of south-central Colorado and north-central
New Mexico; marginal occurrence are: in Colorado, 22 mi. E Mosca, Sa-
guache, Alamosa and Antonito; in New Mexico, 4 mi. SW Cimmaron.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face between (c) Vinaceous-Buff and Avellaneous, purest on sides and flanks,
upper parts strongly suffused with black; arietiform markings, pinnae of ears,
plantar surfaces of hind feet and dorsal and ventral stripes of tail, blackish.
Skull small; rostrum slender; interorbital width narrow; auditory bullae
strongly inflated; braincase but slightly vaulted; interparietal region wide;
zygomatic arch bowed laterally.
Comparisons. — From Dipodomys ordii richardsoni, D. o. montanus differs
as follows: Size smaller; color darker in all pigmented areas; ventral stripe
of tail continuous to end of pencil in all specimens, whereas in D. o. richardsoni
the stripe is incomplete in most specimens. In all specimens the dorsal and
ventral stripes of the tail are darker in D. o. montanus; skull smaller in all
measurements taken; jugal straight or nearly so, as contrasted to bowed
laterally; pterygoid fossae shallower and more ovoid, that is to say, wings of
pterygoid flare laterally; ventral projection of auditory bullae more pro-
nounced; rostrum, immediately in front of zygomatic process of maxilla,
nearly parallel to median plane along long axis of skull, while in D. o. rich-
ardsoni this union forms a decided obtuse angle; paroccipital processes smaller;
foramen magnum smaller and more ovoid as opposed to larger and more
circular.
From Dipodomys ordii evexus, D. o. montanus differs as follows: Size
smaller in all measurements taken except width across auditory bullae which
is greater; color darker, ventral stripe of tail continuous to end of pencil;
least width of supraoccipital less (2.5 mm. in D. o. montanus and 3.5 mm. in
D. o. evexus); extension of premaxillae posterior to nasals greater; braincase
more inflated dorsally; bullae relatively as well as actually larger and more
inflated ventrally; pterygoid fossae narrower anteroposteriorly and wider lat-
erally than in D. o. evexus; jugal straight or nearly so as contrasted to bowed
laterally; lacrimal processes smaller.
For comparison with Dipodomys ordii medius see account of that sub-
Bpeciea.
Remarks. — This subspecies of Dipodomys ordii appears to be re-
stricted to the San Luis Valley of Colorado and New Mexico. In-
Setzer: Subspeciation in Kangaroo Rat
539
( Ire "lilll
30
25 0 25 75 125 175 MILES
' i 1
20bL
120
115
110
® TYPE LOCALITY
0 SPECIMEN EXAMINED
▲ ADDITIONAL RECORD
Fig. 26. Known occurrences and probable geographic range of the subspe-
cies of Dipodomys ordii in the southwestern fourth of the range of the species.
14. D.o. ordii 17. D.o. chapmani 31. D.o. longipes
540 University of Kansas Publs., Mus. Nat. Hist.
tergradation, between Dipodomys ordii ordii and D. o. montanus,
is noted in animals from Gran Quivira, Mesa Jumanes, New Mexico.
These animals have the frontomaxillary suture and extension of
the premaxilla as in D. o. ordii and the nasals and inflation of the
auditory bullae as in D. o. montanus. In the majority of char-
acters they are, however, referable to the latter. Intergradation is
noted also in animals from 4 miles southwest of Cimmaron, New
Mexico, in that they resemble D. o. richardsoni in the shape of the
nasals and the width and length of the rostrum but in the remainder
of the characters studied resemble D. o. montanus to which they
are referred.
Baird, in the original description, gives "near Fort Massachusetts"
as the type locality for this subspecies. Miller (1923) lists the type
locality as 'Tort Massachusetts (now Fort Garland)." Fort Mas-
sachusetts and Fort Garland are two different ex-army posts. Old
Fort Massachusetts was situated on what is now the Trinchera
Ranch which is north of the town of Fort Garland. Old Fort Gar-
land is approximately one-half mile south of the town of Fort Gar-
land. The two abandoned army posts were no more than twelve
miles apart. In 1946, at neither abandoned post could I find any
habitat suitable for kangaroo rats.
Specimens examined. — Total, 148, distributed as follows:
Colorado: Saguache County: Saguache, 1 (USBS). Alamosa County: 1.6 mi. NE
Medano Springs Ranch Headquarters, 8 (MVZ) ; Great Sand Dunes National Monument,
10 (MVZ); 3 mi. S Great Sand Dunes National Monument, 4 (MVZ); Medano Ranch, 15
mi. NE Mosca, 11 (2 USBS; 9 MVZ); 22 mi. E Mosca, 5 (AMNH); Alamosa, 1 (UM).
Conejos County: Antonito, 1 (USBS); 7 mi. E Antonito, 3 (USBS); 12 mi. E Antonito,
2 (USBS). Costilla County: 4V2 mi. N Blanca, 8100 ft., 3 (MVZ); 1 mi. NE Blanca, 7950
ft., 1 (MVZ); 3 mi. NW Fort Garland, 6 (CMNH) ; 1 mi. NW Fort Garland, 3 (MVZ);
15 mi. W Fort Garland, 7800 ft., 4 (KU) ; Fort Garland, 64 (31 USBS; 28 LACM ; 5
MVZ); 5 mi. SSE Fort Garland, 4 (AMNH); 5 mi. S Fort Garland, 2 (AMNH).
New Mexico: Colfax County: 4 mi. SW Cimmaron, 5 (AMNH); 3 mi. SE Cimmaron,
8 (AMNH); iy2 mi. W Philmont Ranch Headquarters, 2 (AMNH).
Dipodomys ordii cinderensis Hardy
Dipodomys ordii cinderensis Hardy, Proc. Biol. Soc. Washington,
57:53, October 31, 1944.
Type.— Male, no. 4611, Mus. Zool., Univ. Utah; at approximately 4,000 ft.
on sandy soil, immediately north of the northern of two large cinder cones in
Diamond Valley, 10 mi. N St. George, Washington County, Utah; obtained on
February 13, 1944, by Ross Hardy, original no. 2690.
Range. — Diamond Valley, Washington County, Utah, north through Moun-
tain Meadows, east as far as Cedar City, Iron County, Utah, north through
the Escalante Desert to Lund, Iron County, and west almost to the Utah-
Nevada boundary; marginal occurrences, all in Utah are: 11 mi. SE Lund; N
end Mountain Meadows; Diamond Valley; 10 mi. W Cedar City; 4tl> mi. NW
Summit and 6 mi. W Parowan.
Setzer: Subspeciation in Kangaroo Rat 541
Diagnosis.— Size small (see measurements). Color dark, upper parts near
Buffy Brown, strongly suffused with black, purest on sides and flanks; pinnae
of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail, arieti-
f orm markings, black. Skull small ; rostrum narrow ; auditory bullae relatively
as well as actually well inflated; external auditory meatus small; interparietal
region narrow.
Comparisons. — From Dipodomys ordii fetosm, D. o. cinderensis differs in:
Size smaller; color darker; skull smaller in all measurements taken.
From Dipodomys ordii panguitchensis, D. o. cinderensis differs as follows:
Size smaller; color lighter; skull smaller except for auditory bullae which are
larger; external auditory meatus smaller; maxillary tooth-row longer.
Compared with Dipodomys ordii longipes and Dipodomys ordii cupidineus,
D. o. cinderensis is smaller, darker and can easily be distinguished.
For comparisons with Dipodomys ordii celeripes and Dipodomys ordii
utahensis see accounts of those subspecies.
Remarks. — This race confined to Utah, and inhabiting northern
Washington County and most of Iron County, is restricted almost
exclusively to areas of loose shifting sand. Intergradation with
Dipodomys ordii fetosus occurs near Lund. The animals from this
locality intergrade in size of body and in color, but in the majority
of cranial characters resemble D. o. cinderensis. In all characters
studied D. o. cinderensis appears to be more closely related to Di-
podomys ordii utahensis than to any other subspecies with which it
comes into actual contact.
Animals from near Parowan have both cranial and external char-
acters by which they could be referred to either D. o. panguitchensis
or D. o. cinderensis. Owing to the existence of the physical barrier
(Cedar Mountains) separating the two populations, they are re-
ferred to D. o. cinderensis. Dipodomys ordii cinderensis is not a
well differentiated race but appears to possess characters sufficiently
diagnostic to distinguish it from contiguous forms.
Specimens examined. — Total, 74, all from Utah, distributed as follows: Iron County: 11
mi. SE Lund, 50 (46 RH; 4 MVZ) ; 4y2 mi. NW Summit and 6 mi. W Parowan, 9 (RH);
10 mi. W Cedar City, 1 (USAC) ; 5 mi. W Cedar City, 1 (USAC) ; Cedar City, 2 (BYU).
Washington County: Diamond Valley, 9 (RH); N end Mountain Meadows, 2 (RH).
Dipodomys ordii fetosus Durrant and Hall
Dipodomys ordii jetosus Durrant and Hall, Mammalia, 3:14, March,
1939.
Type. — Female, adult, no. 48451, Museum of Vertebrate Zoology, University
of California; 2 mi. N Panaca, 4,800 ft., Lincoln County, Nevada; obtained
on June 24, 1931, by Ward C. Russell, original no. 1658.
Range. — Southeastern Nevada and western Beaver and Millard counties,
Utah; marginal occurrences in Nevada, 2 mi. SE Pioche, 15 mi. WSW Sunny-
side, 16y2 mi. WSW Sunnyside, 14 mi. NNE Sharp, 8V2 mi. NE Sharp, 15 mi.
542 University of Kansas Publs., Mus. Nat. Hist.
S Groom Baldy, 10 mi. E Crystal Spring, Panaca; in Utah, Pine Valley, 50 mi.
W Milford and 5 mi. S Garrison.
Diagnosis. — Size medium (see measurements). Color dark, entire dorsal
surface (16") between Pinkish Cinnamon and Cinnamon-Buff, purest on sides
and flanks with strong admixture of black in upper parts; cheeks white; pinnae
of ears, arietiform markings, plantar surfaces of hind feet, dorsal and ventral
stripes of tail, blackish. Skull medium in size; rostrum wide and short; inter-
orbital region narrow; braincase but slightly vaulted; lacrimal processes large;
zygomatic arches weak and bowed laterally; pterygoid fossae large and sub-
circular.
Comparisons. — From Dipodomys ordii utahensis, D. o. fetosus differs as
follows: Hind foot longer; color lighter; skull larger; rostrum, relatively as
well as actually, shorter and wider; nasals longer and wider; interorbital region
narrower; braincase less vaulted; interparietal region narrower; auditory
bullae more inflated laterally, posteriorly and ventrally; foramen magnum
smaller.
From Dipodomys ordii ccleripes, D. o. fetosus differs as follows: Larger;
color darker; arietiform markings present; skull larger; rostrum longer and
wider; nasals longer and less inflated distally; interorbital region wider;
auditory bullae somewhat more inflated; zygomatic arches more robust.
For comparisons with Dipodomys ordii monoensis, D. o. columbianus, D. o.
pallidus, D. o. panguitchensis and D. o. cinderensis see accounts of those sub-
species.
Remarks. — Dipodomys ordii fetosus has a small geographic range.
Hall (1946:416) points out that the interorbital breadth in topo-
types is less than in those from any other part of the range. This
feature, therefore, is not useful in characterizing this subspecies.
Intergradation with Dipodomys ordii columbianus is noted in the
greater total length and the somewhat darker color in animals from
Coal Valley and Garden Valley, Nevada. In the part of the range
which lies in Utah, none of the animals is typical and all are inter-
grades in color and cranial characters with Dipodomys ordii
celeripes.
Specimens examined. — Total, 136, distributed as follows:
Nevada: Nye County: White River Valley, 15 mi. WSW Sunnyside, 5500 ft., 26
(MVZ); White River Valley, 16V2 mi. WSW Sunnyside, 5500 ft., 6 (MVZ); 14 mi. NNE
Sharp, 1 (MVZ); Garden Valley, 15 mi. NE Sharp, 1 (MVZ); Garden Valley, lty2 mi. NE
Sharp, 3 (MVZ); 8'/2 mi. NE Sharp, 17 (MVZ). Lincoln County: E side Coal Valley, 14
mi. N Seeman Tass, 4850 ft., 4 (MVZ); 2 mi. SE Pioche, 1 (MVZ); E side Coal Valley,
10 mi. N Seeman Pass, 1 (MVZ); Desert Valley, 20 mi. SW Pioche, 1 (MVZ); 2 mi. N
Panaca, 4800 ft., 17 (16 MVZ; 1 CAS); Desert Valley, 21 mi. W Panaca, 10 (MVZ); Crys-
tal Spring, Pahranagat Valley, 4000 ft., 2 (MVZ); 10 mi. E Crystal Springs, 5000 ft., 1
(MVZ); 15 mi. S Groom Baldy, 1 (MVZ).
Utah: Millard County: 5 mi. S Garrison, 5400 ft., 12 (MVZ); Pine Valley, Sec. 33,
T. 25 S, R. 17 W, 5000 ft., 16 (UU); Warm Cove, Sec. 34, T. 25 S, R. 18 W, 5500 ft., 2
(UU); Desert Range Experiment Station, 50 mi. W Milford, 5252 ft., 10 (9 UU; 1 BYU).
Setzer: Subspeciation in Kangaroo Rat 543
Dipodomys ordii utahensis (Merriam)
Perodipus montanus utahensis Merriam, Proc. Biol. Soc. Washington,
17:143, July 14, 1904.
Perodipus ordii utahensis, Goldman, Proc. Biol. Soc. Washington,
30:113, May 23, 1917.
Dipodomys ordii columbianus, Grinnell, Journ. Mamra., 2:96, May 2,
1921.
Dipodomys ordii columbianus, Hall, Univ. California Publ. Zool., 37:5,
April 10, 1931 (part— the part from Ogden, Utah.)
Tfype.— Male, adult, no. 55115, U. S. Nat. Mus. Biol. Surv. Coll.; Ogden,
Weber County, Utah; obtained on July 15, 1893, by Vernon Bailey, original
no. 4085.
Range. — Between Great Salt Lake and the Wasatch Mountains from ex-
treme northern Utah, south to northern Sevier County, Utah; marginal occur-
rences, all in Utah, are: Promontory Point, Ogden, St. John, Cedar Valley,
Nephi, Aurora, Spring City, Provo and 4 mi. N Draper.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face (a) between Cinnamon-Buff and Clay Color, purest on sides, flanks and
cheeks, with heavy admixture of black in upper parts; arietiform markings,
pinnae of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail
blackish. Skull small; rostrum short and narrow; interorbital region moderately
wide; interparietal large, spatulate anteriorly; lacrimal processes relatively
large; braincase vaulted; zygomatic arches robust and straight; auditory
bullae relatively, as well as actually, greatly inflated.
Comparisons. — From Dipodomys ordii marshalli, D. o. utahensis differs as
follows: Size smaller; color darker; skull smaller; rostrum longer and nar-
rower; interorbital region wider; lacrimal processes larger; interparietal region
larger; auditory bullae less inflated; palate longer and narrower.
From Dipodomys ordii pallidus, D. o. utahensis differs in: Size smaller;
color darker; skull smaller; rostrum shorter and narrower; interorbital region
narrower; auditory bullae less inflated laterally, posteriorly and ventrally;
interparietal region larger; lacrimal processes smaller; foramen magnum with
ventral notch as opposed to rounded ventrally; pterygoid fossae circular as
opposed to subcircular; palate longer and narrower.
From Dipodomys ordii cinderensis, D. o. utahensis differs as follows: Size
larger, except hind foot which is shorter; color somewhat lighter, more reddish;
skull larger; rostrum longer; nasals longer; braincase more vaulted; pterygoid
fossae circular as opposed to subcircular; auditory bullae less inflated laterally
but more inflated ventrally.
For comparisons with Dipodomys ordii columbianus, D. o. jetosus, D. o. mar-
shalli, D. o. panguitchensis and D. o. cineraceus see accounts of those sub-
species.
Remarks. — The systematic status of D. o. utahensis has long been
in doubt. Several workers have, at various times, considered it to be
indistinguishable from Dipodomys ordii columbianus. Durrant and
Setzer (1945:29) recognized it as a valid subspecies and gave diag-
nostic characters by which it could be distinguished from D. o.
columbianus.
544 University of Kansas Publs., Mus. Nat. Hist.
Intergradation has been noted in specimens from the eastern and
southeastern shores of Great Salt Lake. In color and the width of
the auditory bullae these specimens are intermediate between D. o.
utahensis and Dipodomys ordii marshallv but in the sum total of
characters are referable to the latter. Specimens from 20 miles south-
west of Nephi and from U. B. (Yuba) Dam are darker and inter-
mediate in rostral width and breadth of the auditory bullae but are
referable to Dipodomys ordii celeripes.
Specimens examined. — Total, 91, all from Utah, distributed as follows: Box Elder County:
Promontory Point, 1 (USNM). Weber County: Ogden, 4293 ft., 20 (7 BYU ; 7 MVZ;
4 USNM); Little Mountain, 1 (USNM). Davis County: Antelope Island, Great Salt Lake,
4250 ft., 5 (USNM). Tooele County: Bauer, 4500 ft., 6 (UU); St. John, 4300 ft., 4 (UU);
Little Valley, Sheeprock Mountains, 5500 ft., 1 (UU) ; Clover Creek, Onaqui Mountains,
5500 ft., 1 (UU). Salt Lake County: Plain, 4 mi. N Draper, 4500 ft., 1 (UU). Utah
County: Fairfield, Cedar Valley, 4800 ft., 15 (9 BYU; 6 UU); W Lake Mountains, 9
(BYU); Provo, 4510 ft., 10 (MVZ); Sand Dunes W Curtis Station, 4 (BYU). Juab
County: Nephi, 2 (1 USNM ; 1 MVZ) ; 4 mi. W Nephi, 1 (RH). San Pete County: Spring
City, 4 (LACM). Sevier County: 1 mi. W Aurora, 5190 ft., 6 (5 UU; 1 USNM).
Dipodomys ordii columbianus (Merriam)
Perodipus ordi columbianus Merriam, Proc. Biol. Soc. Washington,
9:115, June 21, 1894.
Dipodomys phillipii, Baird, U. S. P. R. R. Expl. and Surveys, Gen.
Rept., p. 412, 1857 (part— the part from Ft. Walla Walla, O. T.).
Cricetodipus ordii columbianus, Trouessart, Catalogus Mammalium,
1:581, 1897.
Perodipus ordii columbianus, Goldman, Proc. Biol. Soc. Washington,
30:113, May 23, 1917.
Dipodomys ordii columbianus, Grinnell, Journ. Mamm., 2:96, May 2,
1921.
Type.— Female, adult, no. 24181/31594, U. S. Nat. Mus., Dept. of Agricul-
ture, Coll.; Umatilla, Plains of Columbia, Umatilla County, Oregon; obtained
on October 18, 1890, by Clark P. Streator, original no. 386.
Range. — Southeastern Washington, eastern Oregon, southwestern Idaho,
northwestern, northern and most of northeastern Nevada; marginal occur-
rences are: in Washington, 2 mi. SW Paterson, 4 mi. E Burbank and Wal-
lula; in Oregon, 7 mi. E Madras, 2 mi. E Prineville and Guano Valley; in
California, Eagleville, Red Rock, Observation Peak, 5 mi. E Litchfield, Honey
Lake, Vinton; in Nevada, 3% mi. E Flanigan, 6 mi. S Pahrurn Peak, 2% mi.
E and 11 mi. N Gerlach, 2 mi. SW Quinn River Crossing, 1 mi. SE Tuscarora,
5 mi. N Beowawe, V2 mi. S Beowawe, 21/£ mi. NE Smiths Creek Ranch, Bells
Ranch, 5 mi. W Austin, 8 mi. W Eureka, 4 mi. S Shoshone, 5 mi. SE Greens
Ranch, and 22 mi. N Deeth; in Idaho, 8 mi. W Rodgerson, 6 mi. SW Amer-
ican Falls, 4 mi. NE American Falls, 5 mi. NW Michaud, Arco, Hammett and
Payette; in Oregon, Umatilla.
Diagnosis. — Size medium (see measurements). Color dark, entire dorsal
surface (16") between Pinkish Cinnamon and Cinnamon-Buff, purest on sides
and flanks with strong admixture of black on upper parts; cheeks white;
arietiform markings, pinnae of ears, plantar surfaces of hind feet, dorsal and
ventral stripes of tail, blackish. Skull medium in size; rostrum relatively
Setzer: Subspeciation in Kangaroo Rat 545
long and narrow; nasals long and expanded distally; interorbital region nar-
row; lacrimal processes small; braincase but slightly vaulted; auditory bul-
lae relatively greatly inflated; pterygoid fossae circular; zygomatic arches
robust.
Comparisons. — From Dipodomys ordii utahensis, D. o. columbianus differs
as follows : Color lighter, dorsal and ventral stripes of tail brownish as opposed
to black; ventral stripe of tail, in most specimens, incomplete to end of pencil
as opposed to complete; lacrimal processes smaller; extension of premaxillae
dorsad wider ; interparietal generally narrower, smaller, quadrangular and trun-
cate anteriorly, as opposed to spatulate and rounded anteriorly; braincase
less vaulted; zygomatic arches bowed laterally as opposed to straight; styloid
processes larger; cutting edge of upper incisors narrower.
From Dipodomys ordii fetosus, D. o. columbianus differs in: Hind foot
shorter; color generally darker; skull larger; rostrum shorter and wider; nasals
somewhat shorter; braincase more vaulted; auditory bullae less inflated; inter-
parietal region wider; ventral border of foramen magnum more rounded (less
indented) ; zygomatic arches straight as opposed to bowed laterally.
For comparisons with Dipodomys ordii inaquosus, Dipodomys ordii monoen-
sis and Dipodomys ordii celeripes see accounts of those subspecies.
Remarks. — In most subspecies of Dipodomys ordii having large
geographic ranges, there is a certain amount of variation, but D. o.
columbianus remains relatively stable throughout its large range.
Where extreme variation is noted in D. o. columbianus it is ob-
viously the result of intergradation. This intergraclation is noted
between Dipodomys ordii monoensis and D. o. columbianus in Lyon
County, Nevada, where the animals are referable to the former race
cranially but to the latter on the basis of color. Animals from Kel-
ton, Utah, are in color as Dipodomys ordii marshalli and cranially
combine characters of D. o. columbianus, D. o. utahensis and D. o.
marshalli. The sum total of their characters places them with D. o.
marshalli. Durrant and Setzer (1945:29), and the present writer
in this account, do not record any animals from Utah as belonging
to D. o. columbianus. Nevertheless, the influence of D. o. colum-
bianus is seen in the animals from Kelton, Utah.
Specimens examined. — Total, 588, distributed as follows:
Washington: Benton County: Blalock Island, 2 mi. SW Paterson, 2 (MVZ). Walla
Walla County: 4 mi. E Burbank, 4 (MVZ); 2 mi. SSE Burbank, 27 (MVZ); Wallula, 7 (4
MVZ; 3 DRD).
Oregon: Gilliam County: Arlington, 1 (MVZ). Morrow County: 2V2 mi- SW Irigon,
6 (MVZ); Cecil, 1 (MVZ). Umatilla County: Umatilla, 10 (MVZ). Jefferson County: 7
mi. E Madras, 2 (DRD). Crook County: 2 mi. NE Prineville, 1 (MVZ); 7 mi. W Prine-
ville, 8 (MVZ); 4 mi. SW Prineville, 23 (MVZ); Crooked River, at mouth of Bear Creek,
8 (MVZ). Lake County: Fort Rock, 1 (DRD) ; NE edge Alkali Lake, 8 (MVZ) ; 9 mi. S
Adel, mouth 20 Mile Creek, 1 (MVZ); Guano Ranch, Guano Valley, 2 (DRD). Harney
County: Malheur Lake, 9 (LACM) ; Narrows, 1 (DRD) ; 1 mi. S Narrows, 3 (MVZ) ; 5 mi.
SW Narrows, 19 (MVZ); Smith Creek, 10 mi. SE Diamond, 1 (MVZ); Lake Alvord, 1
(MVZ); 1% mi. E Denio, 4 (MVZ). Malheur County: 3 mi. N Vale, 2 (MVZ); 8 mi. N
Jordan Valley, 1 (MVZ); y2 mi. S Rome, 5 (MVZ); 1 mi. S Rome, 11 (MVZ).
546 University of Kansas Publs., Mus. Nat. Hist.
Idaho: Payette County: Payette, 1 (DRD). Butte County: Arco, 1 (MVZ). Elmore
County: Hammett, 1 (MVZ). Bingham County: 5 mi. E Shelley, 1 (MVZ); Blackfoot,
5 (MVZ); 3 mi. S Blackfoot, 6 (MVZ); Aberdeen, 1 (MVZ). Owyhee County: S bank
Snake River, Homedale, 13 (MVZ); Indian Cove, 4 (MVZ); 5 mi. SE Murphy, 3 (MVZ);
Castle Creek, 8 mi. S Oreana, 1 (MVZ); S Fork Owyhee River, 12 mi. N Nevada line,
1 (MVZ). Minidoka County: Acequia, 8 (MVZ); 4 mi. N Rupert, 2 (MVZ). Twin Fails
County: Salmon Creek, 8 mi. W Rodgerson, 7 (MVZ). Power County: 5 mi. NW Michaud,
4 (MVZ) ; 4 mi. NE American Falls, 4 (MVZ) ; 6 mi. SW American Falls, 10 (MVZ).
California: Modoc County: Eagleville, 12 (CAS); 2 mi. E Eagleville, 7 (CAS); 2U
mi. E Eagleville, 24 (CAS); 3 mi. E Eagleville, 6 (CAS); 5 mi. E Eagleville, 3 (CAS).
Lassen County: 2 mi. W Red Rock P. O., 1 (MVZ); 1 mi. W Red Rock P. O., 4 (MVZ);
Dransfield, 6 mi. E Ravendale, 5300 ft., 4 (CAS); 7 mi. E Ravendale, 1 (MVZ); 7 mi. N
Observation Peak, 21 (MVZ) ; 6 mi. N Observation Peak, 7 (MVZ) ; Observation Peak, 31
(MVZ); Pete's Valley, 4 (MVZ); 5 mi. E Litchfield, 12 (MVZ); 6 mi. W Wendell, 3975 ft.,
3 (CAS); 8 mi. SW Wendell, 5 (CAS); Honey Lake, 2 (LACM); 4^ mi. WNW Stacy,
5 (MVZ); 3 mi. NW Warm Springs, 1 (MVZ). Plumas County: Beckwith, 1 (DRD); 2 mi.
WNW Vinton, 2 (MVZ) ; Vinton, 18 (14 LACM ; 4 MVZ) ; 1 mi. E Vinton, 7 (MVZ) ; near
Vinton, 21 (DRD).
Nevada: Washoe County: 3 mi. N Vya, 3 (MVZ); 4% mi. NE Painted Point, 4
(MVZ); 8Vz mi. E Vya, 1 (MVZ); 3 mi. E Painted Point, 1 (MVZ); Long Valley Ranch,
3 mi. S Vya, 1 (MVZ); 10y2 mi. S Vya, 1 (MVZ); 11 mi. S Vya, 2 (MVZ); 13 mi. S Vya,
4 (MVZ); Hausen, 4800 ft., 3 (MVZ); 10 mi. SE Hausen, 4675 ft., 7 (MVZ); 2% mi. E
and 11 mi. N Gerlach, 4050 ft., 4 (MVZ); Smoke Creek, 9 mi. E California line, 3900 ft.,
5 (MVZ); 40° 28' N Lat., 6 mi. E California line, 4000 ft., 3 (MVZ); Horse Canyon, 3 mi.
NW Pahrum Peak, 5000 ft., 1 (MVZ); Fox Canyon, 6 mi. S Pahrum Peak, 4800 ft., 4
(MVZ); N side Sand Pass, 3950 ft., 2 (MVZ); 4 mi. NW Flanigan, 4200 ft., 1 (MVZ); 3y2
mi. NW Flanigan, 4200 ft., 1 (MVZ); 3V2 mi. E Flanigan, 2 (MVZ); 2% mi. SW Pyramid,
4300 ft., 2 (MVZ). Humboldt County: 1 mi. S Denio, Oregon, 4200 ft., 6 (MVZ); Quinn
River Crossing, 4100 ft., 1 (MVZ). Elko County: Mary's River, 22 mi. N Deeth, 3 (MVZ);
1 mi. SE Tuscarora, 5900 ft., 2 (MVZ); 5 mi. W Halleck, 5200 ft., 7 (MVZ); 3 mi. W Hal-
leck, 5300 ft., 2 (MVZ). Lander County: 1 mi. E Battle Mountain, 1 (MVZ); Reese River
Valley, 6 mi. N Austin, 2 (MVZ); Malloy Ranch, 5 mi. W Austin, 3 (MVZ); 2y2 mi. NE
Smiths Creek Ranch, 5800 ft., 1 (MVZ); Campbell Creek, 6900 ft., 3 (MVZ); Campbell
Creek Ranch, 5500 ft., 8 (MVZ). Eureka County: 5 mi. N Beowawe, 7 (MVZ); y2 mi. S
Beowawe, 1 (MVZ); Pine Creek, 2 mi. E Palisades, 7 (MVZ); Evans, 4 (MVZ); Winzell,
3 (MVZ); 4 mi. SE Romano, Diamond Valley, 1 (MVZ); 8 mi. W Eureka, 12 (MVZ).
White Pine County: 5 mi. SE Greens Ranch, Steptoe Valley, 1 (MVZ); Cherry Creek,
6600 ft., 2 (MVZ); 1 mi. E Illipah, 6100 ft., 3 (MVZ); 5V2 mi. SE Ely, 6500 ft., 5 (MVZ);
Spring Valley, 5900 ft., 4 mi. S Shoshone, 6 (MVZ). Nye County: Bells Ranch, Reese
River, 6890 ft., 1 (MVZ).
Dipodomys ordii idoneus new subspecies
Type. — Male, adult, no. 90029, Museum of Zoology, University of Michigan;
San Juan, 12 mi. W Lerdo, 3,800 ft., Durango, Mexico; obtained on March 1,
1946, by F. and F. Hammerstrom.
Range. — Known only from the type locality.
Diagnosis. — Size small (see measurements). Color pale, entire dorsal surface
(c) between Light Ochraceous-Buff and Ochraceous-Buff, purest on sides, flanks
and cheeks; upper parts but slightly washed with black; arietiform markings,
lateral parts of pinnae of ears, plantar surfaces of hind feet, dorsal and ventral
stripes of tail, brownish; medial part of pinnae of ears flesh-colored. Skull
medium in size; rostrum wide and relatively long; braincase but slightly in-
flated; least interorbital width great; interparietal region relatively narrow;
upper incisors strongly recurved; zygomatic arches relatively heavy and
slightly bowed laterally; auditory bullae greatly inflated ventrally, laterally
and posteriorly.
Comparisons. — From Dipodomys ordii ordii, D. o. idoneus differs in: Size
Setzer: Subspeciation in Kangaroo Rat 547
smaller; color lighter; skull larger; nasals longer and somewhat more inflated
distally as opposed to straight; interorbital width greater; breadth across
maxillary arches greater; interparietal region narrower; auditory bullae more
inflated posteriorly, laterally and ventrally and projecting anteriorly into orbit;
extension of nasals anterior to upper incisors less; zygomatic arches heavier
and more bowed laterally; paroccipital processes smaller; foramen magnum
elongate as opposed to nearly circular.
From Dipodomys ordii obscurus, D. o. idoneus differs as follows: Size
smaller; color lighter; skull larger, rostrum wider; nasals shorter and more
expanded distally; interorbital region wider; width across maxillary arches
greater; interparietal region narrower; auditory bullae more inflated laterally,
posteriorly and ventrally; zygomatic arches shorter, heavier and less bowed
laterally.
From Dipodomys ordii palmeri, D.o. idoneus differs in: Size smaller; color
lighter; skull larger; rostrum shorter and wider; nasals shorter and more
expanded distally; interorbital region wider; auditory bullae more inflated
laterally, posteriorly and ventrally; zygomatic arches heavier.
From Dipodomys ordii fuscus, D. o. idoneus differs as follows: Size smaller;
color lighter; skull smaller; rostrum narrower and shorter; interorbital width
greater; braincase more vaulted; nasals longer; interparietal region generally
wider; auditory bullae less inflated; breadth across maxillary arches greater;
zygomatic arches heavier and more bowed laterally; pterygoid fossae ovoid
as opposed to subcircular; occipital condyles more anterior to dorsal evagi-
nation when viewed ventrally.
Dipodomys ordii idoneus differs from Dipodomys ordii attenuatus in larger
size, generally darker color and larger skull.
Remarks. — This subspecies of Dipodomys ordii is paler than
either of the subspecies with adjoining geographic ranges. Meas-
urements of the skull, in proportion to external measurements, are
larger than in any other subspecies of Dipodomys ordii. Cranially,
this new subspecies is more closely allied to Dipodomys ordii fuscus
but in color appears to be much closer to Dipodomys ordii attenu-
atus from which it differs markedly in cranial features.
Specimens examined. — Total, 3, all from Durango, distributed as follows: San Juan, 12
mi. W Lerdo, 3 (UM).
Dipodomys ordii priscus Hoffmeister
Dipodomys ordii priscus Hoffmeister, Proc. Biol. Soc. Washington,
55:167, December 31, 1942.
Perodipus longipes, Allen, Bull. Amcr. Mus. Nat. Hist., 8:246, Novem-
ber, 1896 (part — the part from Kinney Ranch, Wyoming).
Perodipus montanus richardsoni, Cary, N. Amer. Fauna, 33:141, Au-
gust 17, 1911 (part — the part from Lay, Colorado).
Periodipus [Dipodomys] ordii luteolus, Svihla, Journ. Mamm.; 12:262,
August, 1931 (part — the part from Linwood, Utah).
Type.— Male, no. 89119, Mus. Vert. Zool., Univ. California; Kinney Ranch,
21 mi. S Bitter Creek, 7100 ft., Sweetwater County, Wyoming; obtained on
September 16, 1939, by Donald T. Tappe, original no. 766.
Range. — Southwestern Wyoming, extreme northeastern Utah and north-
f>4s University of Kansas Publs., Mus. Nat. Hist.
western Colorado; marginal occurrences are: in Wyoming, 7 mi. N Ft.
Washakie and 10 mi. SW Granger; in Colorado, Lay and Snake River south
of Sunny Peak.
Diagnosis. — Size medium (see measurements). Color light, entire dorsal
surface between (c) Light Ochraceous-Buff and Ochraceous-Buff, purest on
sides, flanks and cheeks; upper parts slightly suffused with black; arietiform
markings, pinnae of ears, plantar surfaces of hind feet, dorsal and ventral
stripes of tail, grayish to blackish in different individuals. Skull medium in
size; rostrum long and narrow; nasals long; auditory bullae relatively nar-
row and skull long, giving appearance of much narrower skull than is actually
the case; zygomata weak; upper incisors moderately wide at cutting edge.
Comparisons. — From Dipodomys ordii uintensis, D. o. prisons differs as
follows: Hind foot longer; color lighter in all pigmented areas, and skull
smaller in all measurements taken.
From Dipodomys ordii sanrafaeli, D. o. prisons differs in : Color decidedly
less red in upper parts ; ventral stripe of tail not continuous to end of pencil ;
nasals longer; auditory bullae less inflated.
For comparisons with Dipodomys ordii luteolus and Dipodomys ordii ter-
rosus see accounts of those subspecies.
Remarks. — The characters of this subspecies are stable through-
out most of its geographic range. It is not known to intergrade
with adjacent forms. One specimen, a male, available from the
Snake River, south of Sunny Peak, Colorado, however, is not typ-
ical of D. o. prisons, in that it has a wider and deeper rostrum than
any other specimen from the entire range. Between D. o. priscus and
D. o. luteolus, the skull, but not the color, provides diagnostic dif-
ferences.
The Red Desert of Wyoming is mostly sandy but there are large
exposures of rock and gravel. At the type locality of D. o. priscus
rockier soil predominates but is interspersed with stabilized sand
dunes where the kangaroo rats are abundant. Traps set on the
rocky soils, in a continuous line between the dunes, yielded no
kangaroo rats.
Kangaroo rats in the area of the Kinney Ranch, Wyoming, ap-
parently form a good source of food for owls, since many bones
(jaws and limb elements) were found in owl pellets at the bases of
cliffs approximately 5 miles northeast of the ranch buildings.
Specimens examined. — Total, 72, distributed as follows:
Wyoming: Fremont Count}/: 7 mi. N Fort Washakie, 1 (USBS); Fort Washakie, 1
(USBS); Wind River, 1 (USBS). Sweetwater County: Eden, 1 (USBS); 5 mi. E Rock
Springs, 1 (UM); 10 mi. SW Granger, 13 (UM) ; Bitter Creek, 9 (6 AMNH; 3 CNHM) ;
Kinney Ranch, 21 mi. S Bitter Creek, 18 (MVZ); Shell Creek, 25 mi. S Bitter Creek, 3
(CM); 30 mi. S Bitter Creek, 3 (KU) ; 33 mi. S Bitter Creek, 8 (KU). Carbon County:
20 mi. W Baggs, 2 (USBS).
Utah: Daggett County: E bank Green River, 4 mi. E Linwood, 4 (CM).
Colorado: Moffat County: Snako River, -outh of Sunny Peak, 1 (USBS); 20 mi. NW
Sunbeam, 4 (CM); Bear River, Lay, 1 (USBS). Rio Blanco County: W side White River,
1 mi. N Rangely, 1 (CM).
Setzer: Stjbspeciation in Kangaroo Rat 549
Dipodomys ordii celeripes Durrant and Hall
Dipodomys ordii celeripes Durrant and Hall, Mammalia, 3:10, March,
1939.
Dipodomys ordii columbianus, Hall, Univ. California Publ. Zool., 37:5,
April 10, 1931.
Type. — Male, adult, no. 1956, Museum of Zoology, University of Utah;
Trout Creek, 4600 ft., Juab County, Utah; obtained on May 5, 1937, by
Stephen D. Durrant, original no. 1168.
Range. — Eastern Nevada, western and west-central Utah, east to eastern
Sevier County, Utah; marginal occurrences are: in Nevada, Tecoma, Cobre
and 8 mi. SE Mt. Moriah; in Utah, E side Clear Lake, U. B. (Yuba) Dam,
10 mi. SW Nephi, and Trout Creek.
Diagnosis. — Size small (see measurements). Color pale, entire dorsal sur-
face Pinkish Buff, purest on sides, flanks and cheeks, with but slight suffusion
of black in upper parts; pinnae of ears, plantar surfaces of hind feet, dorsal
and ventral stripes of tail, brownish. Skuil small; rostrum relatively short
and wide; nasals relatively short; lacrimal processes small; braincase but
slightly vaulted; auditory bullae but little inflated; interorbital region nar-
row; styloid processes project on ventral surface of auditory bullae beyond
middle of external auditory meatus; zygomatic arches robust and bowed
laterally; pterygoid fossae ovoid.
Comparisons. — From Dipodomys ordii marshalli, D. o. celeripes differs as
follows: Size smaller, hind foot longer, tail shorter; color lighter; skull
smaller; rostrum wider and shorter; interorbital region narrower; auditory
bullae less inflated; lacrimal processes smaller; palate shorter; pterygoid
fossae larger.
From Dipodomys ordii cinderensis, D. o. celeripes differs as follows: Size
smaller, tail shorter; color lighter; rostrum wider and shorter; nasals shorter;
interorbital width less; auditory bullae less inflated; foramen magnum wider
and shorter; pter3rgoid fossae more circular.
From Dipodomys ordii columbianus, D. o. celeripes differs in: Size smaller;
color lighter; rostrum wider and shorter; interorbital width less; interparietal
region wider; auditory bullae, generally, less inflated; upper incisors longer and
more robust; pterygoid fossae larger.
For comparisons with Dipodomys ordii jetosus, Dipodomys ordii pallidus
and Dipodomys ordii panguitchensis see accounts of those subspecies.
Remarks. — It appears that wherever the range of D. o. celeripes
meets that of an adjoining race, intergradation occurs freely.
Practically all of the Nevadan specimens referable to this subspecies
are intergrades with Dipodo?nys ordii columbianus in size and color.
Specimens from west of Delta, Utah, are intermediate in size be-
tween Dipodomys ordii pallidus and D. o. celeripes but their cranial
characters are more as in D. o. celeripes. Intergrades with Dipod-
omys ordii utahensis are discussed under the account of that sub-
species.
The characters differentiating D. o. celeripes from any contiguous
subsppcies are not present in every specimen even in the type series.
550 University of Kansas Publs., Mus. Nat. Hist.
Specimens examined. — Total, 96, distributed as follows:
Nevada: Elko County: 13 mi. N Montello, 5000 ft., 2 (MVZ) ; iy2 mi. N Tecoma,
4900 ft., 6 (MVZ); Tecoma, 4900 ft., 8 (MVZ); Cobre, 6100 ft., 3 (MVZ); Salt Springs,
4200 ft., 1 (MVZ). White Pine County: 2 mi. W Smith Creek Cave, 6300 ft., Mt. Moriah,
2 (MVZ); Hendry Creek, 8 mi. SE Mt. Moriah, 6200 ft., 1 (MVZ).
Utah: Toole County: Clifton Flat, 7 mi. SW Gold Hill, 6149 ft., 4 (UU); Parrish
Ranch, 5 mi. N Ibapah, 5175 ft., 1 (UU); Ibapah, 5000 ft., 23 (UU). Juab County: Trout
Creek, 4600 ft., 21 (19 UU ; 1 USAC ; 1 MVZ); 20 mi. SW Nephi, 2 (UU) ; U. B. (Yuba)
Dam, Sevier River, 5000 ft., 10 (UU). Millard County: 4 mi. S Gandy, 5000 ft., 1 (MVZ) ;
Smith Creek, 6 mi. S Gandy, 5400 ft., 2 (MVZ); Hendry Creek, 5000 ft., 17 mi. S Gandy,
4 (MVZ); Oak City, 5000 ft., 1 (UU) ; White Valley, 60 mi. W Delta, 1 (UU); 35 mi. W
Delta, 1 (UU); East side Clear Lake, 4600 ft., 4 (3 UU; 1 USAC); 2 mi. E Clear Lake,
4600 ft., 2 (UU).
Dipodomys ordii cineraceus Goldman
Dipodomys ordii cineraceus Goldman, Journ. Mamm., 20:352, August
14, 1939.
Type.— Male, no. 263890, U. S. Nat. Mus. Biol. Surv. Coll.; Dolphin Island,
Great Salt Lake, 4250 ft., Box Elder County, Utah; obtained on June 4, 1938,
by "William H. Marshall, original no. 57.
Range. — Dolphin Island, Great Salt Lake, Utah.
Diagnosis. — Size medium (see measurements). Color pale, upper parts
near Pale Pinkish Buff, but lightly washed with black, purest on sides and
flanks; arietiform markings, pinnae of ears, plantar surfaces of hind feet, dorsal
and ventral stripes of tail, brownish. Skull medium in size; rostrum short;
nasals relatively short; interparietal region relatively wide; auditory bullae
but slightly inflated.
Comparisons. — From Dipodomys ordii utahensis, D. o. cineraceus differs in:
Size smaller; color lighter; nasals longer; auditory bullae larger; total length
of skull greater.
From Dipodomys ordii marshalli, D. o. cineraceus differs in: Size smaller;
color darker; skull larger in all measurements taken.
Remarks. — The race D. o. cineraceus along with Dipodomys
microps russeolus is found on Dolphin Island, Great Salt Lake, Utah.
Dolphin Island is connected with the mainland by a bar that could
allow animals from the island to disperse onto the mainland. There
are no animals from the mainland, however, that are referable to
D. o. cineraceus or to D. m. russeolus. The animals available from
the mainland, at Kelton, although geographically near Dolphin
Island, are referable to D. o. marshalli and to D. m. bonnevillei.
The paucity of material from the mainland, adjacent to Dolphin
Island, may be significant. Future trapping in this area is needed
before we can be certain that D. o. cineraceus is isolated.
Specimens examined. — Total, 2, both from Utah, as follows: Box Elder County: Dolphin
Island, Great Salt Lake, 4250 ft., 2 (USNM).
Setzer: Subspeciation in Kangaroo Rat 551
Dipodomys ordii marshalli Goldman
Dipodomys ordii marshalli Goldman, Proc. Biol. Soc. Washington,
50:223, December 28, 1937.
Type.— Female, adult, no. 262655, U. S. Nat. Mus. Biol. Surv. Coll.; Bird
Island, Great Salt Lake, 4,300 ft., Tooele County, Utah; obtained on June 22,
1937, by W. H. Marshall, X-catalog no. 27969.
Range.— Bird, Carrington, Badger and Stansbury islands, Great Salt Lake;
around western edge of Great Salt Lake north to Kelton, Box Elder County,
Utah; around southern and southeastern edge of the lake to mouth of the
Jordan River; marginal occurrences are: all in Utah, Kelton, 2 mi. W Grants-
ville, 14 mi. W Salt Lake City and Bird Island, Great Salt Lake.
Diagnosis. — Size medium (see measurements). Color pale, entire dorsal
surface near Pinkish Buff, purest on sides, flanks and cheeks with but slight
suffusion of black on upper parts; arietiform markings, pinnae of ears, plantar
surfaces of hind feet, and dorsal and ventral stripes of tail, brownish. Skull
medium in size; rostrum narrow and long; pterygoid fossae ovoid; cutting
edge of upper incisors narrow; external auditory meatus round and small;
jugal straight or nearly so.
Comparisons. — From Dipodomys ordii pallidus, D. o. marshalli differs in:
Hind foot smaller ; color darker, arietiform markings more distinct ; dorsal and
ventral stripes of tail more pronounced; skull smaller; palate shorter and
wider; jugals lighter; external auditory meatus smaller; cutting edge of upper
incisors narrower; nasals shorter and less flared distally.
From Dipodomys ordii utahcnsis, D. o. marshalli differs in: Size smaller;
color lighter; skull smaller, except least interorbital width and depth of
auditory bullae which are greater; cutting edge of upper incisors narrower;
nasals less flared distally.
For comparison with Dipodomys ordii celeripes see account of that sub-
species.
Remarks. — This subspecies was described from Bird Island,
Great Salt Lake, and was thought by Marshall (1940:153), who
collected the specimens, to be restricted to that island. However,
specimens taken over a period of years on the nearby mainland, by
students from the University of Utah, are referable to this sub-
species. Although these animals from the mainland are referable
to D. o. marshalli, they have some characteristics of D. o. utahensis
from the eastern and southern mainland. Four specimens from Kel-
ton, also on the mainland, are not typical of D. o. marshalli; never-
theless, more than half of their characters of taxonomic worth are as
in D. o. marshalli to which the animals are referred.
Great Salt Lake has not been so effective in isolating the animals
living on the islands as heretofore has been thought. After these
supposedly isolated kinds of animals from Great Salt Lake were
named, some other workers have shown several of the kinds to have
552 University of Kansas Publs., Mus. Nat. Hist.
extensive ranges on the mainland. Some of the kinds ranging also
on the mainland are: Dipodomys ordii marshalli, Dipodomys mi-
crops subtenuis and Peromyscus crinitus pergracilis. Of the animals
named from the islands, I suppose that those which require the
semiarid habitat found in this area will all be found to have large
ranges on the adjacent mainland and that each of the kinds of ani-
mals which do not require the above habitat, and which lived in
this region during the Pleistocene, will be found to be restricted to
the island from which it was named.
Specimens examined. — Total, 47, all from Utah, distributed as follows: Box Elder County:
Kelton, 4300 ft., 7 (5 MVZ; 2 UU). Tooele County: 2 mi. W Grantsville, 1 (CAS); Bird
Island, Great Salt Lake, 4300 ft., 1 (USNM) ; Carrington Island, Great Salt Lake, 4300 ft.,
1 (USNM); Stansbury Island, Great Salt Lake, 4300 ft., 10 (6 UU; 4 USNM). Salt Lake
Comity: 18 mi. W Salt Lake City, 4260 ft., 16 (UU); 17 mi. W Salt Lake City, 4320 ft.,
7 (UU); 16 mi. W Salt Lake City, 4300 ft., 3 (UU); 14 mi. W Salt Lake City, 4300 ft.,
1 (UU).
Dipodomys ordii inaquosus Hall
Dipodomys ordii inaquosus Hall, Proc. Biol. Soc. Washington, 54:58,
May 20, 1941.
Type. — Male, adult, no. 73580, Museum of Vertebrate Zoology, University
of California; 11 mi. E and 1 mi. N Jungo, 4,200 ft., Humboldt County,
Nevada; obtained on July 26, 1936, by Ward C. Russell, original no. 5026.
Range. — Southeastern Humboldt and northern Lander counties, Nevada;
marginal occurrences, all in Nevada, are: 7 mi. N Winnemucca, 8 mi. E and
1 mi. N Jungo, 15 mi. SW Winnemucca, 23 mi. NW Battle Mountain, Izen-
hood, and 18 mi. NE Iron Point.
Diagnosis. — Size medium (see measurements). Color pale, entire dorsal sur-
face Pinkish Buff, purest on sides and flanks with but slight suffusion of
black on the upper parts; cheeks white; arietiform marks indistinct, pinnae
of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail, brown-
ish. Skull medium in size; rostrum short and wide; nasals relatively long
and expanded distally; braincase but slightly vaulted; interorbital width
narrow; extension of premaxillae posterior to nasals relatively great; inter-
parietal relatively large; pterygoid fossae subcircular; foramen magnum
cvoid; auditory bullae relatively greatly expanded ventrally.
Comparisons. — From Dipodomys ordii columbianus, D. o. inaquosus differs
as follows: Size larger; color lighter; skull larger; nasals longer; rostrum
longer and wider; interobital region narrower; braincase more vaulted; inter-
parietal larger; auditory bullae more inflated ventrally; pterygoid fossae sub-
circular as opposed to nearly circular; zygomatic arches more robust.
From Dipodomys ordii monoensis, D. o. inaquosus differs in: Size larger;
color lighter; skull larger; nasals longer; rostrum wider; interorbital width
greater; braincase more vaulted; auditory bullae more inflated laterally,
ventrally and posteriorly; interparietal region narrower; pterygoid fossae
subcircular as opposed to circular; lacrimal processes larger; foramen magnum
ovoid as opposed to circular.
Setzer: Subspeciation in Kangaroo Rat 553
Remarks. — In pallor D. o. inaquosus closely approaches Dipodo-
mys ordii celeripes from eastern Nevada and western Utah. The
two populations are separated, however, by a large area inhabited
by the darker Dipodomys ordii columbianus. Specimens from 18
mi. NE Iron Point, Nevada, which Hall (1946:413) refers to D. o.
columbianus, are intergrades in color, size and cranial characters
and here are referred to D. o. inaquosus instead of to D. o. colum-
bianus. Animals from Toulon are intergrades between D. o. ina-
quosus and Dipodomys ordii monoensis but are referable to the
latter.
Specimens examined. — Total, 47, all from Nevada, distributed as follows : Humboldt
County: 18 mi. NE Iron Point, 4600 ft., 1 (MVZ) ; 7 mi. N Winnemucca, 4400 ft., 4
(MVZ); 1 mi. N Winnemucca, 4600 ft., 8 (MVZ); 5 mi. NE Golconda, 7 (MVZ); 3 mi. SW
Winnemucca, 4500 ft,, 1 (MVZ); 4 mi. SW Winnemucca, 4500 ft., 1 (MVZ); 5 mi. SW
Winnemucca, 4600 ft., 2 (MVZ); 8 mi. E and 1 mi. N Jungo, 4200 ft., 6 (MVZ); 11 tni.
E and 1 mi. N Jungo, 4200 ft., 7 (MVZ); 10 mi. SW Winnemucca, 4500 ft., 1 (MVZ);
23 mi. NW Battle Mountain, 4 (MVZ). Pershing County: 15 mi. SW Winnemucca, 1
(MVZ). Lander County: Izenhood, 2 (MVZ); 3 mi. S Izenhood, 2 (MVZ).
Dipodomys ordii attenuates Biyant
Dipodomys ordii attenuatus Bryant, Occas. Papers, Museum of Zool.,
Louisiana State Univ., no. 5:65, November 10, 1939.
Type. — Male, no. 80429, Museum of Vertebrate Zoology, University of
California; Mouth of Santa Helena Canyon, 2146 ft., Big Bend of the Rio
Grande, Brewster County, Texas; obtained on November 19, 1938, by Adrey
E. Borell, original no. 5581.
Range. — Big Bend region of Brewster County, Texas.
Diagnosis. — Size small (see measurements). Color light, entire dorsal sur-
face between Pinkish Buff and Pinkish Cinnamon, purest on sides and flanks,
upper parts but lightly mixed with black; cheeks and narrow outer margin of
ear, pure white; arietiform markings absent; pinnae of ears, dorsal and ventral
stripes of tail, brownish; plantar surfaces of hind feet, blackish. Skull small;
rostrum long and narrow; nasals short; interparietal longer than wide; audi-
tory bullae longer than deep.
Comparisons. — Compared with Dipodomys ordii ordii, D. o. attenuatus
differs in: Size smaller; color lighter; skull smaller; nasals thinner; rostrum
narrower; auditory bullae less inflated.
From Dipodomys ordii richardsoni, D. o. attenuatus differs in smaller size
and lighter color. Skull smaller in all respects.
For comparison with Dipodomys ordii sennetti see account of that sub-
species.
Remarks. — This small race of Dipodomys ordii was known for-
merly only from the type locality. This study has shown it to
range northward and slightly westward. At four places, 6, 20, and
47 miles south of Marathon and 10 miles west of San Vicente, Texas,
6—6114
554 University of Kansas Publs., Mus. Nat. Hist.
® TYPE LOCAI ITY
• SPECIMEN EXAMINED
A ADDITIONAL RECORO
Fig. 27. Known occurrences and probable geographic range of the subspe-
cies of Dipodomys ordii in the northwestern fourth of the range of the species.
9. D.o. panguitchensis
10. D. o. uinlensis
11. D.o. sanrafaeli
12. D. o. jremonti
13. D.o. monoensis
17. D.o. chapmani
19. D.o. cinderensis
20. D.o. jetosus
21. D. o. utahensis
22. D.o. columbianus
24. D. o. priscus
25. D. o. celeripes
26. D.o. cineraceus
27. D.o. marshalli
28.
D.o.
inaquosus
31.
D.o.
longipes
32.
D.o.
pallidus
33.
D .o.
nexilis
34.
D.o.
cupidineus
Setzer: Subspeciation in Kangaroo Rat 555
this race can be said to be typical, in that specimens from these
places agree with those from the type locality. At two other locali-
ties, 5 miles east of Toyahvale and Kent, Texas, animals show an
intergrading tendency toward Dipodomys ordii ordii. The animals
from these latter places are darker colored and are intermediate in
size and cranial characters. Since these animals show more resem-
blance to D. o. ordii in the majority of characters they are referred
to that race, rather than to D. o. attenuatus.
Specimens examined. — Total, 14, all from Texas, distributed as follows: Brewster County:
6 mi. S Marathon, 1 (USBS) ; 20 mi. S Marathon, 1 (USBS) ; Cooper's Well, 47 mi. S
Marathon, 4 (MVZ) ; 10 mi. W San Vicente, 1 (UM) ; Johnson's Ranch, Big Bend Rio
Grande, 4 (2 TCWC; 2 UM) ; mouth Santa Helena Canyon, 3 (1 TCWC; 2 MVZ).
Dipodomys ordii fuscus new subspecies
Type.— Male, adult, no. 93SS6, U. S. Nat. Mus., Biol. Surv. Coll.; Juamave,
Tamaulipas, Mexico; obtained on June 3, 1898, by E. W. Nelson and E. A.
Goldman, original no. 12437.
Range. — Nuevo Leon, extreme northern Zacatecas, Tamaulipas and north-
era San Luis Potosi, Mexico; marginal occurrences are: Coahuila, La Ven-
tura; Tamaulipas, Tula, Juamave and Nuevo Laredo.
Diagnosis. — Size medium (see measurements). Color dark, entire dorsal
surface (a) between Cinnamon-Buff and Clay Color, purest on sides, flanks
and cheeks, upper parts strongly suffused with black; posterior surfaces and
fold of pinnae of ears, white; arietiform markings, plantar surfaces of hind
feet, inside of pinnae of ears, dorsal and ventral stripes of tail, blackish. Skull
of medium size; nasals relatively short; interorbital width narrow; auditory
bullae relatively as well as actually greatly inflated posteriorly, laterally and
ventrally; braincase vaulted; zygomatic arches light and straight; pterygoid
fossae ovoid; rostrum short and wide.
Comparisons. — From Dipodomys ordii palmeri, D. o. fuscus differs in:
Size larger; color lighter; skull larger; rostrum shorter and wider; interorbital
width less; nasals shorter; auditory bullae more expanded laterally, posteriorly
and ventrally; interparietal region narrower; pterygoid fossae more circular;
upper incisors wider at cutting edge; zygomatic arches straight rather than
bowed laterally.
From Dipodomys ordii obscurus, D. o. fuscus differs as follows: Size
larger; color lighter; skull larger; rostrum wider and shorter; interorbital
region wider; width across maxillary arches greater; auditory bullae more
expanded laterally, posteriorly and ventrally; zygomatic arches light and
straight instead of heavy and bowed laterally; pterygoid fossae more nearly
circular; braincase more vaulted; foramen magnum smaller.
From Dipodomys ordii ordii, D. o. fuscus differs in: Size larger; color
darker; skull larger; rostrum shorter and wider; nasals short and more flared
distally; interorbital width greater; auditory bullae more inflated posteriorly,
laterally and ventrally; interparietal region narrower; width across maxillary
arches greater; pterygoid fossae larger and more nearly circular; braincase
more vaulted.
From Dipodomys ordii sennetti, D. o. fuscus can be distinguished by larger
556 University of Kansas Publs., Mus. Nat. Hist.
size, longer tail, darker color, and cranially by larger auditory bullae, longer
nasals, wider interorbital region and greater width across maxillary arches.
Remarks. — This dark subspecies of Dipodomys ordii is most close-
ly allied to Dipodomys ordii palmeri from which it can be dis-
tinguished by the characters set forth under comparisons.
It seems that the Rio Grande serves as a barrier to the dispersal
southward of kangaroo rats from the north side and vice versa. D.
o. fuscus ranges from the type locality to Nuevo Laredo and has not
been found on the north side of the river, being replaced there, up-
river, by Dipodomys ordii medius and down-river by Dipodomys
ordii sennetti.
Specimens examined. — Total, 53, distributed as follows :
Coahuila: San Juan Neponuceno, 5 mi. N La Ventura, 4 (MVZ); La Ventura, 7
(USBS).
Tamaulipas: Nuevo Laredo, 2 (USBS); Miquihuana, 14 (4 USBS; 9 MCZ; 1
AMNH); Juamave, 8 (USBS); Tula, 5 (UM); 3 mi. N Lulu, 5 (MVZ): Lulu, 8 (MVZ).
Dipodomys ordii longipes (Merriam)
Dipodops longipes Merriam, N. Amer. Fauna, 3:72, September 11,1890.
Perodipus ordii, Allen, Bull. Amer. Mus. Nat. Hist., 5:71, April 28,
1893 (part — the part from Bluff, Utah).
Cricetodipus longipes, Trouessart, Catalogus Mammalium, 1:581, 1897.
Perodipus longipes, Elliot, Field Columbian Museum, Zool. Ser., 2:239,
1901.
Perodipus ordii longipes, Goldman, Proc. Biol. Soc. Washington, 30:113,
May 23, 1917.
Dipodomys ordii longipes, Grinnell, Journ. Mamm., 2:96, May 2, 1921.
Dipodomys ordii cleomophila Goldman, Journ. Washington Acad. Sci.,
23:469, October 15, 1933, type from 5 miles northeast of Winona, Coco-
nino County, Arizona.
Type.— Male, no. 17703/24639, U. S. Nat. Mus. Dept. Agric. Coll.; foot of
Echo Cliffs, Painted Desert, Arizona; obtained on September 22, 1889, by C.
Hart Merriam, original no. 512.
Range, — Southeastern Utah, northeastern Arizona to immediately south of
the Little Colorado River, northwestern New Mexico and extreme southwest-
ern Colorado; marginal occurrences are: in Arizona, 20 mi. NE Lees Ferry,
Foot Echo Cliffs in Painted Desert, 5 mi. NE Winona, Winslow and Hol-
brook; in New Mexico, 10 mi. SW Quemado, Riley, Laguna, Chama Canyon
and Shiprock.
Diagnosis. — Size large (see measurements). Color dark, entire dorsal sur-
face between {16") Pinkish Cinnamon and Cinnamon-Buff, purest on sides and
flanks with but slight suffusion of black on upper parts; cheeks white; arieti-
form markings, pinnae of ears, plantar surfaces of hind feet, dorsal and ventral
stripes of tail, brownish. Skull large; rostrum long and narrow; nasals long
and thin; auditory bullae greatly inflated; styloid processes project on ven-
tral surface of auditory bullae beyond middle of external auditory meatus.
Comparisons. — From Dipodomys ordii chapmani, D. o. longipes differs as
follows: Size larger; color lighter; skull larger in all measurements taken;
auditory bullae much more inflated.
Setzer: Subspeciation in Kangaroo Rat 557
From Dipodomys ordii cupidineus, D. o. longipcs differs in: Size larger;
color lighter; rostrum longer and narrower; nasals longer; interorbital width
greater; breadth across zygomatic processes of maxillae greater; auditory
bullae more inflated; zygomatic arches heavier and more bowed laterally.
For comparisons with Dipodomys ordii medius and Dipodomys ordii nexilis
see accounts of those subspecies.
Remarks. — The large reddish Dipodomys ordii longipes is notably
distinct from those subspecies which have adjoining ranges. The sub-
species closest, geographically and morphologically, is Dipodomys
ordii nexilis with which D. o. longipes intergrades, in size of body
and color of pelage, north of the San Juan River, Utah. This inter-
gradation is probably the result of a migration of D. o. longipes
around the head waters of, or above the place in, the river where
there is constantly flowing water. It is probably impossible for
these animals to cross a constantly flowing stream as wide as the
San Juan is in its lower reaches. The San Juan River, however, in
1896, and again in 1934, was so low that it ceased to flow below
Aneth (Gregory, 1938:6, 19). Consequently, at these times, Dipod-
omys could have crossed the river from one side to the other, at
least above the Goosenecks. Intergradation is noted also with
Dipodomys ordii medius at Chama Canyon, New Mexico, where the
color is like that of D. o. medius, although the cranial characters are
as in D. o. longipes; specimens from Chama Canyon are referred to
the latter. Ten miles southwest of Quemado and at Riley, New
Mexico, intergradation with Dipodomys ordii ordii occurs in the
shape of the nasals and the intermediate size of the skull. In color
and external measurements these animals more closely resemble D.
o. longipes.
Goldman (1933:469) described Dipodomys ordii cleomophila
from 5 miles northeast of Winona, Coconino County, Arizona, and
gave as distinctive characters "upper parts distinctly darker,
. . . black facial markings more distinct; skull slightly differ-
ent." These comparative statements were made in reference to
Dipodomys ordii longipes. While it is true that the animals which
Goldman examined are darker and show some slight cranial dif-
ferences, these animals, in my opinion, are not sufficiently distinct
to warrant their separation from D. o. longipes. In the degree of
expansion of the auditory bullae, the width of the rostrum, the
length of the nasals and interorbital width, animals from 5 miles
northeast of Winona (2 specimens) are within the range of in-
dividual variation shown by D. o. longipes. The color is darker,
being more as in Dipodomys ordii chapmani. Specimens examined
558 University of Kansas Publs., Mus. Nat. Hist.
from 10 miles north of Angell, Springerville, Tanner Tank and
Cedar Ranch Wash, Arizona (listed by Goldman as referable to
D. o. cleomophila) , and those from Wupatki Ruins, Arizona, agree
with D. o. longipes except that the width across the maxillary arches
is greater. This greater width across the maxillary arches and the
darker color of animals from Winona and Wupatki Ruins, in my
opinion, afford insufficient basis for the recognition of the subspecies
Dipodomys ordii cleomophila Goldman which, therefore, is placed
as a synonym of Dipodomys ordii longipes (Merriam).
Specimens examined. — Total, 244, distributed as follows:
Utah: San Juan County: 1 mi. N Bluff, 3500 ft., 1 (UU); Va mi. N Bluff, 3300 ft., 7
(UU); Bluff, 2 (UU); Johns Canyon, San Juan River, 5150 ft., 2 (UU).
Arizona: Coconino County: 20 mi. NE Lees Ferry, 1 (USNM); 2 mi. S Endische, 2
(MVZ); 2 mi. SE Endische Spring, Navajo Mountain, 1 (MVZ); V2 mi. W Navajo Spring,
Echo Cliffs, 4 (MVZ); 5 mi. S Navajo Mountain, 2 (MVZ); 7 mi. SE Navajo Mountain,
1 (MVZ); Moa Vae, 10 (MVZ); 2 mi. S Grand Canyon Bridge, 1 (USBS); Tuba, 4 (1
USBS; 3 AMNH); 12 mi. above mouth of Moencopi Wash, 2 (USBS); 14 mi. NW Cedar
Ridge, 1 (USBS); 5 mi. N Cameron, 1 (MVZ); Painted Desert, 6 mi. NE Cameron, 1
(MVZ); 5 mi. E Little Colorado River on Tuba road, 2 (MVZ); Cameron, 4200 ft., 17 (8
LACM; 9 MVZ); Wupatki Ruins, 4 (MVZ); Deadmans Flat, 6400 ft., NE San Francisco
Mt., 3 (MVZ); 3 mi. NW Winona, 4 (USBS); 5 mi. NE Winona, 2 (USBS); 10 mi. N
Angell, Walnut Tank, 8 (USBS); Cedar Ranch Wash, 3 mi. above mouth, 5 (USBS); Tan-
ner Tank, 2 (USBS); Tanners Crossing, Painted Desert, 1 (USBS). Navajo County: Right
fork, Segi-ot-Sosie Canyon, 11 mi. NW Kayenta, 2 (MVZ) ; Bat Woman Canyon, 13 mi. W
Kayenta, 1 (MVZ); Kayenta, 4 (USBS); Segi Canyon, 12 mi. WSW Kayenta, 1 (MVZ)
Segi Canyon, 13 mi. WSW Kayenta, 1 (MVZ); Oraibi, 6000 ft., 9 (4 USBS; 5 MVZ)
Walpi, 2 (USBS); Keam Canyon, 12 (6 USBS; 3 USNM; 3 MVZ); Winslow, 6 (USBS)
Zuni Well, 7% mi. N Adamana, 9 (MVZ); Holbrook, 10 (USBS); Long Canyon, 3 (AMNH)
Bubbling Springs, 2 (AMNH). Apache County: Dinnehotso, 5000 ft., 3 (MVZ); Lukachu-
kai, 1 (USBS); Chin Lee, 6 (USBS); 9 mi. SW Chin Lee, 1 (USBS); 7 mi. from mouth,
Canyon de Chelly, 1 (USBS); Fort Defiance, 2 (USNM); Ganado, 1 (USBS); Springerville,
2 (USBS); 3 mi. SE Springerville, 5 (USBS); unspecified, 4 (2 USBS; 2 USNM); Trash
Tank, Grand Canyon, 2 (USBS).
New Mexico: San Juan County: Ship Rock, 4 (USBS); Fruitland, 9 (USBS);
Blanco, 1 (USBS); Chaco Canyon National Monument, 4 (MVZ). Rio Arriba County:
Chama Canyon, 1 (USBS). McKinley County: 2 mi. W Mariano Lake, 3 (MVZ); Gallup,
6 (USBS); Wingate, 6 (USBS); Thoreau, 2 (USBS); Zuni River, 1 (USBS). Sandoval
County: Juan Tafoya, 1 (USBS). Valencia County: 8 mi. SE Grants, 7 (UM); 9 mi. SSE
Grants, 5 (UM) ; 2y2 mi. E El Morro P. O., Zuni Mts., 1 (LACM); Laguna, 1 (USBS);
Laguna Indian Reservation, 7 (LACM). Catron County: Cougar Ranch, NE Adams Dig-
gings, 1 (CAS); 10 mi. SW Quemado, 1 (USBS). Socorro County: 1 mi. S Bernardo, 1
(MVZ); Riley, 1 (USBS).
Dipodomys ordii pallidas Durrant and Setzer
Dipodomys ordii pallidus Durrant and Setzer, Bull. Univ. Utah, 35
(no. 26) :24, June 30, 1945.
Type.— Male, adult, no. 3526, Museum of Zoology, University of Utah; Old
Lincoln Highway, 18 mi. SW Orr's Ranch in Skull Valley, 4,400 ft., Tooele
County, Utah; obtained on June 6, 1940, by S. D. Durrant, original no. 1905.
Range. — Low valleys of west-central Utah in Tooele, Juab and Millard
counties; marginal occurrences are: all in Utah, 18 mi. SW Orr's Ranch, 7 mi.
S Fish Springs, Hinckley and Lynndyl.
Diagnosis. — Size medium (see measurements). Color light, entire dorsal
surface Light Pinkish Cinnamon, purest on sides and flanks, with but slight
Setzer: Subspeciation in Kangaroo Rat 559
suffusion of black on upper parts; cheeks white; arietiform markings, pinnae
of ears, plantar surfaces of hind feet, dorsal and ventral stripes of tail, brownish.
Skull large; auditory bullae long, wide and well inflated; external auditory
meatus elongate with a notch on the dorsal border; nasals long and flaring dis-
tally.
Comparisons. — From Dipodomys ordii celeripes, D. o. pallidus differs in:
Size larger; color generally darker, although some specimens are lighter;
skull larger; nasals longer and more flared distally; interobital width greater;
auditory bullae larger; external auditory meatus larger.
From Dipodomys ordii jetosus, D. o. pallidus differs in: Size larger; color
lighter; ventral stripe of tail indistinct as opposed to pronounced; skull
larger; nasals longer and more flared distally; auditory bullae larger; external
auditory meatus larger; palate shorter and broader.
For comparisons with Dipodomys ordii marshalli and Dipodomys ordii
utahensis see accounts of those subspecies.
Remarks. — Among named subspecies of Dipodomys ordii, D. o.
pallidus most closely resembles, morphologically, D. o. marshalli,
its nearest geographic neighbor to the north and east. Intergra-
dation with Dipodomys ordii utahensis is noted in color and inter-
mediacy of body size of specimens from Clover Creek. The ma-
jority of cranial characters, however, show these animals to be
referable to D. o. utahensis rather than to D. o. pallidus. Specimens
taken at Lynndyl and Hinckley show intergradation in size of body,
length and configuration of the nasals and the degree of inflation
of the auditory bullae between Dipodomys ordii celeripes and D.
o. pallidus. The majority of characters studied show these latter
animals to be referable to D. o. pallidus. Specimens from 35 and
60 miles west of Delta, in size of body and shape and inflation of
the auditory bullae, are intermediate between D. o. pallidus and
D. o. celeripes. These specimens are here referred to D. o. celeripes.
Specimens examined.- — Total, 34, all from Utah, distributed as follows: Tooele County:
Old Lincoln Highway, 18 mi. SW Orr's Ranch in Skull Valley, 4400 ft., 9 (UU). Juab
County: Fish Springs, 4400 ft., 4 (UU); 7 mi. S Fish Springs, 4400 ft., 4 (UU). Millard
County: 1 mi. N Lynndyl, 4768 ft., 5 (UU); Lynndyl, 4768 ft., 1 (UU); Hinckley, 4600 ft.,
11 (UU).
Dipodomys ordii nexilis Goldman
Dipodomys ordii nexilis Goldman, Journ. Washington, Acad. Sci.,
23:470, October 5, 1933.
Perodipus longipes, Warren, Mammals of Colorado, p. 77, 1910 (part —
the part from Coventry, Colorado).
Dipodomys ordii longipes, Moore, Journ. Mamm., 10:260, August, 1929
(part — the part from Monticello, Utah).
Type.— Male, adult, no. 149938, U. S. Nat. Mus. Biol. Surv. Coll.; 5 mi.
W Naturita, Montrose County, Colorado; obtained on July 20, 1907, by
Merritt Cary, original no. 1068.
Range. — West-central Colorado, southwest into San Juan County, Utah,
north of the San Juan River; northwest into Grand County, Utah, to the
560 University of Kansas Publs., Mus. Nat. Hist.
Colorado River; and westward probably as far as the Colorado River; marginal
occurrences are: in Utah, Cisco, 18 mi. NE Moab, and Blanding; in Colo-
rado, 5 mi. W Naturita, and Coventry.
Diagnosis. — Size medium (see measurements). Color dark, entire dorsal
surface between (a) Cinnamon-Buff and Clay Color, purest on sides and
flanks, upper parts strongly suffused with black; arietiform markings, pinnae
of ears, plantar surfaces of hind feet, dorsal and ventral tail-stripes blackish.
Lateral white stripes of tail less than one fourth of diameter of tail. Skull
large; rostrum long and narrow; nasals long; auditory bullae strongly in-
flated; braincase not inflated; pterygoid fossae ovoid.
Comparisons. — From Dipodomys ordii sanrajaeli, D. o. nexilis differs in:
Color darker; skull smaller in all measurements taken; auditory bullae no-
ticeably smaller; width across zygomatic processes of maxillae less.
From Dipodomys ordii longipes, D. o. nexilis differs in: Size smaller; color
darker in all respects; rostrum wider; breadth across auditory bullae less;
interorbital width greater; breadth across zygomatic processes of maxillae
less; cutting edge of upper incisors wider; zygomatic arch lighter and more
nearly straight.
Comparison with Dipodomys ordii evexus is made in account of that sub-
species.
Remarks. — This animal is apparently not abundant at any place
in its range. Two different attempts, by me, to obtain topotypes
were unsuccessful. A single specimen was obtained and that was
only a skull saved from a mutilated animal that was taken away
from a rattlesnake. The habitat at the type locality is such that
a person would not expect it to be inhabited by kangaroo rats. The
soil is a heavy clay with a generous admixture of stones but in
isolated spots there are light sandy soils which should be suitable
for kangaroo rats. Even so, 500 traps set in the area of the type
locality over a period of two nights yielded no Dipodomys.
This subspecies of Dipodomys ordii, inhabiting west-central Colo-
rado and southeastern Utah, is darker than any other subspecies
with which its range comes in contact. The races to both the north
and south are larger, with the exception of the hind foot which is
longer in D. o. nexilis than in either D. o. sanrajaeli or D. o.
longipes. The affinities of D. o. nexilis are with D. o. longipes
rather than with any of the other known subspecies of Dipodomys
ordii.
Intergradation between D. o. nexilis and D. o. longipes is shown
by animals from Bluff, San Juan County, Utah, which, however,
are referable to the latter. In animals from sixteen miles northwest
of Moab, Grand County, Utah, there is intergradation in size of
skull and in color between D. o. nexilis and D. o. sanrajaeli. The
specimens are referred to D. o. sanrajaeli.
Setzer: Subspeciation in Kangaroo Rat 561
D. o. nexilis is apparently prevented from extending its range to
the northward by the presence there of D. o. sanrafaeli. To the
south D. o. nexilis encounters D. o. longipes and the San Juan River.
To the east it is limited by the Rocky Mountains and to the west
by the deep canyons of the Colorado River.
Warren (1942:183) did not apply the name D. o. nexilis to Ord
kangaroo rats in Colorado. He listed specimens from Montrose,
Montezuma and Mesa counties as being referable to D. o. longipes.
The specimens from Montezuma County, Colorado, probably are
D. o. longipes and those from Montrose and Mesa counties (for
which Warren gives no precise localities) are D. o. nexilis.
Specimens examined. — Total, 35, distributed as follows :
Utah: Grand County: Cisco, 4 (CM); 18 mi. NE Moab, 6000 ft., 1 (UU). San Juan
County: 15 mi. N Monticello, 4 (MVZ); Blanding, 1 (UU).
Colorado: Delta County: Hotchkiss, 1 (USBS). Montrose County: Paradox, 1 (DRD) ;
Bedrock, 4 (AMNH) ; 5 mi. W Naturita, 2 (USBS); Coventry, 3 (1 USBS; 2 AMNH).
Dipodomys ordii cupidineus Goldman
Dipodomys ordii cupidineus Goldman, Joum. Washington Acad. Sci.,
14:372, September 19, 1924.
Type.— Male, no. 243093, U. S. Nat. Mus. Biol. Surv. Coll.; Kanab Wash,
at southern boundary of Kaibab Indian Reservation, Arizona; obtained on
October 12, 1922, by E. A. Goldman, original no. 23384.
Range. — Northeastern Arizona and south-central Utah; marginal occur-
rences are: in Utah, Escalante, near Paria and S of Virgin; in Arizona, 6 mi.
N Wolf Hole, 20 mi. S Wolf Hole, 5 mi. S Trumbull Spring and 10 mi. S
Jacobs Pools in Houserock Valley.
Diagnosis. — Size large (see measurements). Color relatively dark, entire
dorsal surface between (16") Pinkish Cinnamon and Cinnamon-Buff, purest
on sides and flanks, upper parts washed with black ; arietif orm markings, pinnae
of ears, dorsal and ventral stripes of tail, plantar surfaces of hind feet, blackish.
Skull large; auditory bullae well inflated; interorbital width relatively narrow;
rostrum long and narrow; nasals long and slightly flared distally; pterygoid
fossae ovoid; external auditory meatus small and ovoid; jugal light and
relatively straight.
Comparisons. — From Dipodomys ordii chapmani, D. o. cupidineus differs in:
Size larger; color lighter in all pigmented areas; skull larger; rostrum shorter
and wider; nasals shorter; auditory bullae more inflated, particularly median
tympanic portion; least interorbital width less.
From Dipodomys ordii nexilis, D. o. cupidineus differs as follows: Size
smaller; color lighter in all pigmented areas; skull smaller; rostrum relatively
longer and narrower; least interorbital width less; auditoiy bullae less inflated;
breadth across maxillary arches less; foramen magnum circular as opposed to
ovoid; cutting edge of upper incisors narrower; zygomatic arches lighter.
For comparisons with Dipodomys ordii cinderensis, Dipodomys ordii pan-
guitchensis and Dipodomys ordii longipes see accounts of those subspecies.
562 University of Kansas Ptjbls., Mus. Nat. Hist.
Remarks. — This subspecies is prevented from intergrading, to
the east, with D. o. longipes by the deep canyons of the Colorado
River, with D. o. cinderensis by the Virgin River and with D. o.
panguitchensis by a series of high mountain ranges. Intergradation
does, however, occur with D. o. sanrajaeli in animals from Escalante,
Garfield County, Utah, but they are referable to D. o. cupidineus.
The Colorado River, farther to the north, again serves as a barrier
in preventing intergradation between D. o. nexilis and D. o.
cupidineus.
Specimens examined. — Total, 122, distributed as follows:
Utah: Garfield County: Mouth of Calf Creek, Escalante River, 3 (BYU); Ten Mile
Spring, 3 (BYU); Escalante, 3 (BYU). Washington County: near Short Creek road, south
of town of Virgin, 18 (RH). Kane County: near Paria, 1 (UU); Willow Tank Springs, 7
(BYU); 8 mi. NW Kanab, 4800 ft., 1 (UU); near Sand Dunes, 7 (RH) ; Kanab, 15 (12
MVZ; 2 BYU; 1 UU); 1 mi. S Kanab, 4400 ft., 2 (UU).
Arizona: Mohave County: near S Boundary Kaibab Indian Reservation, 4 (MVZ) ; 6
mi. N Wolf Hole, 8 (MVZ); 4 mi. N Wolf Hole; 3 mi. NW Diamond Butte, 7 (USBS) ;
20 mi. S Wolf Hole, 1 (USBS) ; 1 mi. W Diamond Butte, 1 (USBS) ; 6 mi. N Mt. Trumbull,
1 (USBS); Nixon Springs, 6250 ft., W Base Mt. Trumbull, 6 (3 USBS; 3 MVZ); 5 mi. S
Trumbull Spring, 1 (USBS). Coconino County: Fredonia, 3 (USBS); 2 mi. W Lees Ferry,
3 (MVZ); Jacobs Pools, 4000 ft., Houserock Valley, 15 (MVZ); 10 mi. S Jacobs Pools,
Houserock Valley, 2 (MVZ); Houserock Valley, 10 (USBS).
Dipodomys ordii palmeri (Allen)
Dipodops ordii palmeri Allen, Bull. Mus. Comp. Zool., 8:187, 1881.
Cricetodipus ordii palmeri, Trouessart, Catalogus Mammalium, 1 :581,
1897.
Perodipus ordii palmeri, Goldman, Proc. Biol. Soc. Washington, 30:113,
May 23, 1917.
Dipodomys ordii palmeri, Grinnell, Journ. Mamm., 2:96, May 2, 1921.
Cotypes. — Two males, adults, nos. 5886 and 5887, Mus. Comp. Zool.; San
Luis Potosi, Mexico; obtained on May 1, 1878, and September 1, 1878, re-
spectively, by Dr. Edward Palmer. (Types not seen.)
Range. — Eastern Zacatecas, Aguascaliente, northern Jalisco, San Luis Potosi,
Hidalgo, and probably Queretaro; marginal occurrences are: Zacatecas, Cani-
tas and Berriozobal; Aguascaliente, 1 mi. N Chicalote; Jalisco, 9 mi. N Encar-
nacion; Guanajuato, Celaya; Hidalgo, Irolo.
Diagnosis. — Size small (see measurements). Color dark, entire dorsal sur-
face (h) between Cinnamon and Sayal Brown, purest on sides and flanks,
upper parts strongly suffused with black; posterior surfaces and fold of pinnae
of ears, white; arietiform markings, plantar surfaces of hind feet, inside of
pinnae of ears, dorsal and ventral stripes of tail, blackish. Skull small;
nasals long; rostrum long and narrow; interorbital region relatively wide;
degree of inflation of auditory bullae about average for species; zygomatic
arches light and bowed laterally; pterygoid fossae subcircular; braincase but
slightly vaulted.
Comparisons. — From Dipodomys ordii ordii, D. o. palmeri, differs as follows:
Size somewhat larger; color darker; skull larger; nasals longer and flaring dis-
fcally; interorbital width greater; narrower across auditory bullae; interparietal
region narrower; breadth across maxillary arches greater; zygomatic arches
Setzer: Subspeciation in Kangaroo Rat 563
bowed laterally as opposed to relatively straight; pterygoid fossae more nearly
circular; rostrum longer and narrower.
From Dipodomys ordii sennetti, D. o. palmeri differs in larger size, darker
color, longer tail and longer skull.
For comparisons with Dipodomys ordii obscurus and Dipodomys ordii
juscus see accounts of those subspecies.
Remarks. — Specimens from Berriozobal, Zacatecas, in the width
of the rostrum and the configuration of the nasals, are intermediate
between Dipodomys ordii juscus and D. o. palmeri. In color these
animals are more nearly like typical representatives of D. o. palmeri.
The animals from Jalisco and Guanajuato are not typical but the
characters differentiating them from D. o. palmeri are merely modi-
fications of such slight degree that they all have been placed with
that subspecies.
Specimens examined. — Total, 56, distributed as follows:
Zacatecas: Canitas, 3 (USBS); Berriozobal, 10 (USBS).
San Luis Potosi: Potrero Santa Ana, 7.6 mi. S Matehuala, 2 (MVZ) ; Jesus Maria, 12
(USBS) ; 2 mi. NW San Luis Potosi, 3 (MVZ).
Aguascalientes : 1 mi. N Chicalote, 1900 m., 4 (MVZ).
Jalisco: 9 mi. N Encarnacion, 1900 m., 1 (MVZ).
Guanajuato: Celaya, 8 (USBS).
Hidalgo: Ixmiquilpan, 1 (USBS); Irolo, 12 (USBS).
CONCLUSIONS
1. There are thirty-five recognizable subspecies of the species
Dipodomys ordii of which four are herein, for the first time,
recognized by name. Three subspecies, D. o. oklahomae, D.
o. sennetti and D. o. compactus, previously were regarded by
most authors as full species.
2. The species Dipodomys ordii is divisible into six complexes,
or groups, of subspecies on both geographic and morphological
bases.
3. Dipodomys ordii is the most generalized Recent species of the
genus.
4. The extremes of geographic variation in Dipodomys ordii are
greater than in any other species of the genus.
5. Color, at least in Dipodomys ordii, does not seem to be corre-
lated with amount of moisture but rather with color of soil.
6. Clinal variation, from north to south, is shown in the decreas-
ing length of the nasals. This decrease in length of nasals and
resultant decrease in size of the nasal chamber may be corre-
lated with the decrease of humidity of the environment.
7. Subspeciation has been enhanced by the late Quaternary moun-
tain building which was prevalent over the western United
States.
564 University of Kansas Publs., Mus. Nat. Hist.
8. In general, the most primitive kinds of Dipodomys ordii occur
at the periphery of the range of the species.
9. Natural selection plus geographical and ecological isolation
has undoubtedly been operative in speciation and in sub-
speciation.
10. Dipodomys deserti is found to be the most specialized species
in the genus.
11. Six, rather than nine, groups of species are recognized, on the
basis of morphology, as comprising the genus Dipodomys.
12. The center of dispersal for the genus Dipodomys appears to
have been in the southwestern United States and the adjoining
part of Mexico. A secondary center of differentiation is ap-
parent in the low, hot valleys of central California.
13. Parallel development of species is noted between Dipodomys
of the parental center and Dipodomys of the isolated valleys of
central California; Dipodomys in the California center is the
less specialized.
14. The trend of the dipodomyines, as indicated by the fossil rec-
ord, has been toward a saltatorial specialization with conse-
quent morphological changes.
15. The morphological change in the direction of saltatorial spe-
cialization is clearly evident in the compacting and aligning
of the viscera as well as in the lengthening of the distal seg-
ments of the hind legs and the tail, tufting of the tail, enlarge-
ment of the auditory bullae, shortening of the neck and fusion
of the cervical vertebrae for stability and other modifications
of the skeleton.
Setzer: Subspeciation in Kangaroo Rat
565
TABLE 7
Measurements (in Millimeters) of Dipodomys ordii
-^>
O
o
**1
o
JS
CO
0)
j3
S
"3
j3
HJ
tu
Z3
03
+3
T3
c
13
CS *
co 2
O *
c* ^
03 ca
n
CD g
3
fan
■*-»
CO
o
S3
CO
03
C
• IH
*H
o
M
CD
a
' — i
OS
i
c
CD
H-»
o
-a
c
o
till
c
EQ ,
il
-*■* CD
m to
O
O
.4
H->
M
a
HH>
•IH H^
■p-d
03 £
o
<u
t. ■"
is
<p
CD
H
Hi
^
0
o
oq
tJ
►J
n
Z).
D. o. richardsoni Vfa mi.
N Beaver
Oklahoma
(KU)
cf Mean (4)
Maximum
251.2
256
243
131.5
136
133
40.
41
40
5
41.0
41.8
40.4
24.7
25.1
24.4
22.0
22.3
21.5
4.3
4.5
4.2
14.9
15.4
14.6
13.0
13.2
12.7
26.3
27.3
26.0
9 KU 17962
KU 17963
253
246
255
135
133
136
42
40
41
40.6
39.4
41.2
25.4
24.0
25.5
22.6
22.1
4.6
4.3
4.5
15.1
14.1
15.1
14.2
12.5
13.8
25.7
25.6
KU 17964
26.2
D. o. oklahomae 2% mi. S Norman, Oklahoma (USBS)
c? Mean (4)
239.7
254
227
245
234
130.0
140
125
133
120
39.5
42
37
40
39
39.0
40.3
38.0
39.1
38.7
23.9
24.6
23.0
24.5
23.8
21.5
22.4
20.9
22.0
21.7
4.3
4.4
4.2
4.4
4.4
13.7
14.3
13.4
14.4
14.3
13.3
13.6
12.8
13.5
13.5
24.9
Maximum
25.8
Minimum
23.9
9 USBS 265456
25.3
OU 20140
24.8
D. o. com-pactus 19 mi. S Port Aransas, Mustang Island, Texas (TCWC)
c? Mean (10)
234.6
251
228
231.0
235
224
119.5
134
112
116.0
120
110
39.3
41
38
39.0
41
38
37.0
37.9
35.6
37.1
38.2
36.0
21.8
22.5
21.3
22.4
22.9
21.7
19.4
20.8
18.2
19.7
20.5
19.1
4.1
4.2
3.9
4.0
4.1
3.8
13.8
14.6
12.9
13.9
14.7
13.2
11.9
12.3
11.6
24.2
Maximum
25.3
9 Mean (7)
22.8
24.2
Maximum
24.8
23.8
o. sennetti 2 mi. S Riviera, Texas (TCWC)
d"
Mean (5)
217.6
222
208
218.3
226
209
112.2
115
104
112.0
115
108
35.8
38
34
36.0
37
35
37.2
38.2
36.3
37.2
37.7
36.7
23.4
24.1
23.0
23.3
23.9
22.8
20.1
20.7
19.4
20.1
20.8
19.0
4.0
4.3
3.8
4 0
4.1
3.9
13.6
14.4
13.0
13.9
14.4
13.3
13.1
13.2
12.6
13.1
13.4
12.6
24.2
Maximum
24.6
Minimum
23.8
9
Mean (4)
24.1
Maximum
Minimum
24.4
23.5
D. o. evexus Salida,
cf AMNH 28802
Colora
do (AMNH)
267 0 i-in n
42.0
43
39
42.2
43
42
39.3
23.0
23.5
22.1
21.5
21.6
22.1
20.8
4.2
4.4
4.2
4.3
4.4
4.2
25.0
AMNH 28804
271
252
261.0
264
258
156
137
147.0
149
144
14.6
ii.6
25.8
AMNH 28805
24.5
9 Mean (4)
38.0
38.1
37.9
23.5
24.0
22.7
14.1
14.2
14.0
13.4
13.8
13.0
24.6
25.0
24.3
D. o. medins Santa Rosa, New Mexico (USBS)
c^1 Mean (6)
Maximum
258.5
266
251
261.0
143.3
151
132
145.0
38.0
40
37
40.0
39.5
40.6
38.9
40.4
38.1
25.2
25.9
24.8
25.4
24.3
21.4
22.5
20.6
22.2
20.8
4.1
4.2
4.0
4.5
4.1
14.2
15.2
13.5
15.7
13.5
12.9
13.1
12.8
13.5
13.5
24.8
25.6
9 USBS 118527
24.1
25.7
USBS 127310
566 University of Kansas Publs., Mus. Nat.
TABLE 7 — Continued
Hist.
+3
o
0
0
J3
a;
s
"3
j3
C
9)
a
-^
o
X3
o
-a
M
C
V
+3
- —
43 .
.fi.fi
to to
o w
"Si
3
Eh
-*>
oa
o
M
O
J3
43
OS
in
03
C
*^<
o
J3
3
hi
o
U
0)
43
J3
43
M
a
0)
■4
03
03
"c3
+3
O
to
c
M
M
C
a)
oq-h
0)
H
►J
l-l
0
o
H
h-j
hJ
«
D. o. obscurus Rio Sestin, I
cf Mean (6)
)urango
(AMN1
3)
36.3
38.0
35.2
36.5
36.0
35.9
22.9
23.4
22.6
22.8
22.3
22.7
19.7
21.0
19.1
19.9
19.6
19.4
3.7
3.9
3.6
3.8
3.5
3.4
13.1
13.5
11.9
13.1
13.0
12.6
12.6
13.0
12.4
12.9
12.3
22.8
23.9
21.7
9 AMNH 20945
22.3
AMNH 20951..
22.5
AMNH 20958..
22.7
D. o. terrosus Jordan, Montana
c? AMNH 41442
280.0
267.0
279.0
265.0
273.0
155.0
155.0
162.0
149 0
154.0
44.0
40.0
41.0
40.5
41.0
42.7
40.5
40.8
41.4
41.3
26.5
24.8
25.7
25.4
25.1
23.6
21.1
21.6
22.2
22.4
4.0
4.0
4.4
4.3
4.0
15.2
14.5
14.6
14.7
14.9
14.5
13.1
13.4
13.0
13.8
27.0
AMNH 41443
25.1
AMNH 41444
25.9
9 AMNH 41441
26.8
MVZ 25658
26.0
D. o. fremonti Torrey, Utah
d1 CM 15663
250.0
248.0
258.0
252.0
141.0
136.0
141.0
140.0
40.0
38.0
39.0
39.0
37.0
37.3
37.5
36.8
23.3
23.8
23.5
23.5
19.3
19.8
19.9
20.5
3.6
3.8
3.7
3.7
13.5
13.7
13.8
13.5
12.0
11.6
11.9
ll.fi
23.2
CM 15670
23.5
9 CM 15666
23.5
CM 15667
23.2
D. o. uintensis 2 mi. N Fruitland, Utah
c? CM 11634
253.0
260.0
140.0
150.0
40.0
41.0
37.9
38.2
23.3
24.2
20.3
20.6
3.9
4.0
13.5
13.2
12.5
12.5
23.9
CM 11640. .
24.5
D. o. sanrafaeli 12 mi. S'W Green River, Utah
9
CM 15649.
CM 15647.
265.0
253.0
144.0
138.0
41.0
42.0
14.1
13.5
D. o. panguitchensis 1 mi. S Panguitch, Utah
d" RH2151
RH2152
9 RH2153.
257.0
252.0
240.0
145.0
135.0
132.0
41.0
40.0
38.0
13.3
13.5
12.7
12.0
11.9
11.3
D. o. monoensis 5 mi. N Benton Station, California (MVZ)
rf1 MVZ 26993..
232.0
240.0
230.0
228.0
240.0
220.0
125.0
125.0
122.0
125.4
137.0
120.0
40.0
39.0
39.0
38.4
39.0
38.0
36.6
36.8
37.2
36.6
37.2
36.0
23.7
23.0
23.2
23.1
24.2
22.6
20.0
19.8
20.1
19.7
20.0
19.3
3.7
3.6
3.6
3.5
3.7
3.4
13.0
13.4
13.4
13.1
13.2
13.0
11.5
12.0
11.8
12.2
11.5
23.4
MVZ 26995
23.3
MVZ 26997
23.9
9 Mean (5)
Minimum
23.4
23.6
23.0
Setzer: Subspeciation in Kangaroo Rat
567
TABLE 7— Continued
-M
o
OQ
o
*4->
o
,z",
,J3
|
C3
JO
a!
■v
IS
fcfl
e3j5
3
u
oa
o
3
3
O
J3
Ml
M
■M
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J2J2
* §
13
C
0>
o
Xi
o
+3
^3
1> o
■gg
o
5
o
A
C-fl
a
o
C
01
£«
2jS
»>£
c3 &
a;
09
08
H
1-1
i-l
a
o
pq
£
►J
1-5
m
D. o. ordii Near El Paso, Texas (USNM)
Mean (5)..
Maximum.
Minimum .
Mean (6) . .
Maximum.
Minimum .
235.2
245 0
222.0
128.0
136.0
120.0
37.4
39.0
35.0
37.1
38.1
36.3
37.3
38.0
36.3
24.1
24.8
23.4
24.0
24.1
24.0
19.6
20.5
18.8
19.6
20.0
19.1
3.7
3.8
3.5
3.7
3.8
3.6
13.0
13.2
12.8
13.3
13.5
13.0
12.7
13.3
12.1
12.8
13.4
12.4
D. o. luteolus 1 mi. NE Casper, Wyoming (KIT)
Mean (12) .
Maximum.
Minimum .
Mean (7) . .
Maximum.
Minimum.
265.6
281.0
254.0
152.2
163 0
145.0
42.2
43.0
42.0
38.9
39.5
37.5
24.1
25.0
23.8
20.8
22.1
19.9
4.3
4.4
4.2
13.9
14.5
13.0
13.0
13.7
12.5
260 7
269.0
250.0
148.0
153.0
139.0
41.0
43.0
40.0
38.6
40.5
37.6
24.2
25.7
23.0
20.9
21.9
20.0
4.3
4.4
4.2
13.9
14.9
13.3
12.9
13.8
12.5
D. o. extractus 1 mi. E Samalayuca, Chihuahua (MVZ)
cf
Mean (11).
Maximum .
Minimum .
MVZ 76568....
MVZ 76569....
MVZ 76570....
235.1
251.0
224.0
128.2
142.0
121.0
37.3
39.0
35.0
37.5
38.6
36.2
24.1
24.6
23.2
20.1
20.8
19.3
3.5
3.8
3.3
13.2
13.8
12.6
13.0
13.5
12.4
251.0
244.0
230.0
132.0
140.0
120.0
39.0
37.0
37.0
39 6
37.3
35.7
25.4
23.6
23.4
22.2
20.0
19.5
3.7
3.6
3.2
13.8
13.5
12.4
14.1
12.8
12.1
D. o. chapmani Camp Verde, Arizona (USBS)
cf Mean (6)
Maximum
Minimum
Mean (4) . .
Maximum.
Minimum .
258.0
268.0
244.0
148.0
157.0
138.0
39.0
41.0
38.0
38.9
40.1
38.1
24.2
24.7
23.6
21.1
22.1
20.5
3.8
3.9
3.8
14.2
14.8
13.3
13.5
13.9
13.1
255.4
262.0
245.0
151.0
152.0
144.0
38.2
39.0
37.0
37.1
37.7
36.0
23.9
24.3
23.7
20.1
20.1
20.1
3.8
3.9
3.6
13.4
13.7
13.1
13.5
14.9
12.6
D. o. montanus Fort Garland, Colorado (USBS)
cf
Mean (11).
Maximum.
Minimum .
Mean (11).
Maximum.
Minimum .
252.0
140.5
41.0
37.7
24.2
20.1
4.1
13.6
12.8
263.0
150.0
42 0
39.0
25.0
21.0
4.5
14.0
13.2
233.0
131.0
39.0
36.6
23.6
19.0
3.9
12.7
12.4
256.0
141.0
40.8
37.2
23.9
19.6
4.1
13.4
12.5
259.0
145.0
42.0
38.4
24.4
21.4
4.3
14.1
13.3
237.0
132.0
40.0
36.0
23.0
18.2
3.9
12.3
11.9
D. o. cinderensis 11 mi. SE Lund, Utah
MVZ 102059.
MVZ 102056.
MVZ 102057.
MVZ 102058.
244.0
237.0
229.0
241.0
135.0
130.0
122.0
127.0
41.0
41.0
41.0
40.0
35.7
36.4
36.3
36.8
23.2
23.7
23.5
24.0
19.2
20.6
20.2
20.0
3.6
3.5
3.6
3.6
12.2
13.0
13.0
13.2
11.2
12.1
11.5
11.7
568
University of Kansas Publs., Mus. Nat. Hist.
TABLE 7— Continued
•4->
o
o
o
Q
DO
c
CO
"c3
a
CD
'3
60
-0
o
M
T3 o
f 3
^2
<0 £
o ^
11
J-
CO
o
0
03
OQ
o
IS
p
a
s
03
0
c
g m
8 s
s*
53
03
oi
H
t-1
hJ
O
o
pq
h-3
ij
n
Z). o. fetosus 2 mi. N Panaca, Nevada (MVZ)
d"
Mpan (4)
242.5
249.0
233.0
229 0
235.0
224.0
133.7
140.0
126.0
125 5
129.0
122.0
42.3
43.0
42.0
40.6
41.5
40.0
37.4
37.7
36.8
35.8
37.0
34.3
23 7
24 0
23.2
23.3
24.1
23.0
20.0
20.3
19.5
19.5
20 0
19.0
3.7
3.8
3.7
3 6
3.6
3.6
13 3
13 9
12.7
12.8
13.3
11.9
11.7
12.0
11.5
11 3
11.6
11.0
23.7
23.8
9
Mean (4)
23.6
22.5
23.2
21.9
D. o. utahensis Ogden, Utah
MVZ 44005
243.0
238 0
240 0
241.0
133.0
133 0
137 0
134.0
40.5
38.0
39 0
40.0
35.9
36.1
35.6
36.4
23.1
22 9
23 i 5
19.2
19.8
19.8
3.5
3.7
3 6
3.6
12.8
13.5
12.8
13.3
12.4
12.2
11.7
12 3
22.4
MVZ 44006
22.9
MVZ 44007
22.4
MVZ 44008
D. o. idoneus 12 mi. W Lerdo, Durango
D. o. -priscus 33 mi. S Bitter Creek, Wyoming (KU)
D. o. dneraceiis Dolphin Island, Great Salt Lake, Utah
D. o. columbianus Umatilli
d1 MVZ 45317
l, Oregc
243.0
236 0
240.0
239 0
251.0
n
140 0
137.0
132 0
130 0
140.0
40.0
40 0
40 0
39.0
41.0
37.2
36.8
36.4
35.8
36.6
23.6
23.5
23.4
22.7
23.8
20 0
20.1
19.9
19.8
19.6
3.8
3.6
3 7
3.7
3.6
13 5
13.2
13.0
13.2
13.3
12.2
12.1
11.9
12.0
23.4
MVZ 45318
23.4
MVZ 45322
23.3
9 MVZ 45314
22.7
MVZ 45315
23.1
cf UM 90027
35 0
35.0
37.3
37.2
24.3
23.5
20.7
20.0
3.8
3.6
13.0
13.5
14.1
13.4
23.1
UM 90029
230.0
129.0
23.3
c? Mean (7)
Maximum
259.0
265.0
251.0
257.0
264.0
249.0
148.0
152 0
144.0
147.0
152 0
138 0
44.0
45.0
43.0
43.0
45.0
40.0
39.1
40.4
38.0
39.4
40 4
38.1
24.3
25.1
23.7
24.6
25.2
23.5
20.7
21.2
20.0
20 8
21.9
20.1
4.1
4.3
4.0
4.2
4.3
4.1
14.3
15.2
13.8
14.3
14.9
14.0
13.1
13.6
12.7
13.1
13 3
12.7
24.9
25.5
Minimum
9 Mean (4)
Minimum
23.7
24.7
25.2
24.0
D. o. celeripes Trout Creek
d1 Mean (4)
, Utah
208.7
220 0
203.0
219.0
223.0
(UU)
109 0
115 0
100 0
110 0
120.0
40.2
41 0
39.0
39 0
40.0
35.3
36.6
34.0
35.3
37.0
22.9
23.6
22.1
23.4
23.6
19.3
20 2
18.4
19.1
20.4
3.6
3.8
3.5
3.6
3.6
12.6
13.5
11.7
11.9
13.1
11.2
11.5
11.1
11.8
12.1
9 UU 1957
22.4
UU 1961
23.9
9 USNM 263893
228.0
230.0
129 0
132.0
39.0
38.0
37.2
37.1
23.4
23.7
19.7
20.1
3.5
3.6
13.6
13.4
11.6
11.9
23.0
USNM 263894
23.6
Setzer: Subspeciation in Kangaroo Rat
569
TABLE 7— Continued
-M
o
o
o
pd
OQ
S
,4
a
03
~a
J3
49
T3
J3
M
OS <S
O «
B M
a> g
3
M
03
o
3
CO
c
13
M
O
CO
J3
■p
M
a
s
hi
M
ra
03
B
C
-*>
o
O
A
•**
M
c
o
J3
M
C
i —
cp m
CD o
** u
B o
£=3
o
o
M
w
£3
CO
CJ3
03 •?
B &
H
►J
h5
a
0
ffl
hJ
hH
M
Z). o. marshalli Stansbury Island, Great Salt Lake, Utah
<? UU 2968
238.0
241.0
128.0
136.0
40.0
40.0
36.4
36.3
35.9
23.5
23.7
23.5
19.2
19.9
3.5
3.7
3.6
13.0
12.9
12.9
12.0
11.9
23.2
UU 2969
23.0
9 UU 2972
1
23.1
D. o. inaquosus 11 mi. E and 1 mi. N Jungo, Nevada (MVZ)
0* Mean (4)
243.0
247.0
238.0
242.0
231.0
254.0
137.0
140.0
131.0
139.0
130.0
138.0
40.0
40.5
39.0
40.5
40.5
41.0
36.6
36.8
36.4
36.4
36.3
38.2
23.5
24.0
23.3
23.9
23.2
23.9
19.8
20.0
19.5
19.2
19.0
20.0
3 7
3.8
3.6
3.7
3.5
3.6
13.5
13.8
13.1
13.2
12.7
13.9
11.8
12.1
11.5
11.4
11.0
12.2
23.1
Maximum
23.5
Minimum
23.0
9 MVZ 73577
22.9
MVZ 73579
23.2
MVZ 73582
D. o. attenuatus Johnson's
d" TCWC 3633
Ranch,
235.0
237.0
245.0
238.0
Texas
126.0
136.0
143.0
140.0
41.0
37.0
37.3
38.0
37.3
35.5
35.7
34.6
24.1
22.1
22.3
22.5
20.1
18.8
19.1
18.5
3.7
3.5
3.6
3.5
13.7
13.2
12.4
11.9
12.2
12.9
13.0
23.4
TCWC 3634
22.3
UM 79121
23.3
9 UM 79122
21.9
D. o. fuscus Juamave, Tamaulipas (USBS)
d" Mean (4)
9 Mean (4)
248.0
254.0
241.0
244.7
254.0
240.0
149.0
152.0
146.0
146.2
155 0
138.0
38.5
40.0
38.0
38.5
40.0
38.0
37.6
38.3
37.0
37.4
37.7
37.2
24.1
24.7
23.5
24.1
24.5
23.7
20.2
20.4
19.7
19.9
20.5
19.2
3.6
3.7
3.6
3.4
3.6
3.3
12.6
13.1
12.3
12.9
13.0
12.7
12.7
12.9
12.5
12.7
12.9
12.4
23.6
23.7
23.6
23.2
Maximum
23.4
Minimum
23.2
D. o. longipes Kayenta, Arizona
o" USBS 247915.
USBS 247916.
9 USBS 247552 .
255.0
242.0
255.0
145.0
138.0
135.0
41.0
40.0
40.0
39.3
38.4
39.9
25.5
25.4
25.7
20.7
20.6
21.2
3.9
3.8
4.0
13.8
13.3
13.8
13.0
12.7
D. o. pallidum 18 mi. SW Orris Ranch, Utah (UU)
d" Mean (4)
236.6
240.0
230.0
229.0
131.3
134.0
128.0
132.0
42.3
43.0
41.0
40.0
37.6
37.9
37.0
36.6
24.2
24.6
23.7
23.5
20.4
20.9
19.8
20.0
3.8
4.0
3.7
3.7
13.4
13.6
12.9
12.5
11.8
12.1
11.5
12.0
23.6
Maximum
24.2
23.4
9 UU 3528
D. o. nexilis 5 mi. W Naturita, Colorado
d" USBS 149940
261.0
265.0
156.0
142.0
44.0
45.0
38.7
40.5
25.1
26.3
18.9
21.5
4.1
4.2
13.8
14.4
11.4
13.1
24.6
9 USBS 149941
25.3
7—6114
570 University of Kansas Publs., Mus. Nat. Hist.
TABLE 7— Concluded
-^
o
o
1*4
o
A
CO
a
"3
a
'o)
43
1c
T3 <u
ts.2
cu~
t- 3
3
Lh
CO
o
as
CO
S
c
o
14
till
M
■M
^-.
—
XI jo
«~.
C
C
CD
o
us
0
J3
18
O
o
J3
-h>
C J
b
^^
43
-*->
-^ 3
-*J
■h>T3
0)
cj
M
m
-d
M
a
O
C
c
g§
c3 ?
CO
03
H
hJ
hJ
o
o
m
hJ
►J
M
£>.
o. cupidineus Houserock Valley, Arizona (USBS)
Moan (6) . .
Maximum.
Minimum .
Mean (4) . .
Maximum.
Minimum .
239 0
244.0
225.0
244.7
247.0
240.0
138 0
39 8
36.9
24.1
19.3
3.8
12.8
12.0
146.0
41.0
37.5
24.9
20.1
4.0
13.0
12.6
130.0
38.5
36.3
23.5
18.7
3.6
12.6
11.7
141.0
40 0
37.4
24.2
19.7
3.8
13.2
12.3
147.0
40 5
37.7
24.4
20.0
3.9
13.5
12.6
137.0
39.5
37.1
23.9
19.5
3.7
12.9
12.0
23.6
24.3
23.2
23.9
24.6
23.7
D. o. palmeri Jesus Maria, San Luis Potosi (USBS)
cf
Mean (6) . .
Maximum.
Minimum .
Mean (6) . .
Maximum.
Minimum.
242.0
255.0
229.0
239.5
252.0
225.0
141.6
37.6
36.7
23.6
20.0
3.3
12.9
12.9
157.0
39 0
37.3
24.4
20.8
3 6
13.6
13.4
129.0
35.0
35.9
23.2
19.4
3.0
12.5
12.6
139.6
37.1
36.4
23.8
19.9
3.4
12.8
13.1
148 0
38 0
37.3
23.9
20.0
3.6
13 3
13.2
128.0
36.0
35.9
23.7
19.5
3.3
12.2
13.0
22.6
23.0
22.0
22.7
23.5
22.0
Setzer: Subspeciation in Kangaroo Rat 571
LITERATURE CITED
Anderson, R. M.
1947. Catalogue of Canadian Recent mammals. National Museum of
Canada, Bull. 102, Biol. Ser., 31, pp. v + 238, January 24, 1947.
Bailey, V.
1905. Biological survey of Texas. N. Amer. Fauna, 25:1-222, 24 figs.,
Government Printing Office, October 24, 1905.
Benson, S. B.
1933. Concealing coloration among some desert rodents of the southwest-
ern United States. Univ. California Publ. Zool., 40:1-70, 2 pis., 8
figs, in text, June 13, 1933.
Burt, W. H.
1936. A study of the baculum in the genera Perognathus and Dipodomys.
Journ. Mamm, 17:145-156, May 14, 1936.
Coues, E.
1875. A critical review of the North American Saccomyidae. Proc. Acad.
Nat. Sci. Philadelphia, 1875:272-327, 3 figs, in text, August 31, 1875.
Coues, E., and Allen, J. A.
1877. Monographs of North American Rodentia, pp. x + 1081, 77 figs.,
1877.
Davis, W. B. .
1942. The systematic status of four kangaroo rats. Journ. Mamm., 23:
328-333, August 13, 1942.
Durrant, S. D., and Setzer, H. W.
1945. The distribution and taxonomy of kangaroo rats (genus Dipodo-
mys) of Utah. Bull. Univ. Utah, 35(26) :l-39, 2 figs., 4 maps, June
30, 1945.
Gazin, C. L.
1942. The late Cenozoic vertebrate faunas from the San Pedro Valley,
Arizona. Proc. U. S. Nat. Mus., 92:475-518, 2 pis., 9 figs., 1942.
GlDLEY, J. W.
1922. Preliminary report on fossil vertebrates of the San Pedro Valley,
Arizona, with descriptions of new species of Rodentia and Lago-
morpha. U. S. Geol. Surv. Prof. Papers, 131-E, pp. 119-131, 2 pis.,
1922.
Goldman, E. A.
1933. New mammals from Arizona, New Mexico, and Colorado. Journ.
Washington Acad. Sci., 23:463-473, October 15, 1933.
Gregory, H. E.
1938. The San Juan Country: a geographic and geologic reconnaissance
of southeastern Utah. U. S. Geol. Surv. Prof. Papers, 188, pp.
v + 123, 26 pis., 4 figs., 1938.
572 University of Kansas Publs., Mus. Nat. Hist.
Grinnell, J.
1919. Four new kangaroo rats from west-central California. Proc. Biol.
Soc. Washington, 32:203-206, December 31, 1919.
1922. A geographical study of the kangaroo rats of California. Univ.
California Publ. Zool., 24(1) :1-124, 7 pis., 24 figs., June, 1922.
Hall, E. R., and Dale, F. H.
1939. Geographic races of the kangaroo rat Dipodomys microps. Occas.
Papers, Mus. Zool., Louisiana State Univ., 4:47-63, 2 figs., Novem-
ber 10, 1939.
Hall, E. R.
1946. Mammals of Nevada, pp. xi + 710, frontispiece, 11 pis., 473 figs.,
Univ. California Press, Berkeley and Los Angeles, California, July
1, 1946.
Hjbbard, C. W.
1937. Additional fauna of Edson Quarry of the Middle Pliocene of Kansas.
Amer. Mid. Nat., 18(3) :460-464, May, 1937.
1939. Notes on additional fauna of Edson Quarry of the Middle Pliocene
of Kansas. Trans. Kansas Acad. Sci., 42:457-462, 6 figs., 1939.
Howell, A. B.
1932. The saltatorial rodent Dipodomys: The functional and comparative
anatomy of its muscular and osseous systems. Proc. Amer. Acad.
Arts and Sci., 67(10) :377-536, 28 figs., December, 1932.
1944. Speed in animals, their specialization for running and leaping, pp.
xi -f 270, 55 figs., University of Chicago Press, Chicago, Ulinois,
1944.
Howell, A. B., and Gersh, I.
1935. Conservation of water by the rodent Dipodomys. Journ. Mamm.,
16:1-9, February 14, 1935.
Marshall, W. H.
1940. A survey of the mammals of the islands in Great Salt Lake, Utah.
Journ. Mamm., 21:144-159, May 14, 1940.
Mayr, E.
1942. Systematics and the origin of species, pp. x -f- 334, 29 figs., Colum-
bia University Press, New York, New York, 1942.
MlDGLEY, E. E.
1938. The visceral anatomy of the kangaroo rat. Journ. Mamm., 19:304-
317, 16 figs., August 18, 1938.
Miller, G. S., Jr.
1924. List of North American Recent mammals. 1923, U. S. Nat. Mus.
Bull., 128:xvi + 673, 1924.
RlDGWAY, R.
1912. Color standards and color nomenclature, pp. iii + 43, 53 pis., pub-
lished by the author, Washington, D. C, 1912.
Setzer: Subspeciation in Kangaroo Rat 573
Schultz, J. R.
1938. A late Quaternary mammal fauna from the tar seeps of McKittrick,
California. Carnegie Inst., Washington, 487:113-215, 17 pis., 12 figs.,
July 6, 1938.
Warren, E. R.
1942. The mammals of Colorado, their habits and distribution, pp. xviii
+ 330, 50 pis., University of Oklahoma Press, Norman, Oklahoma,
1942.
Wilson, R. W.
1939. Rodents and Lagomorphs of the late Tertiary Avawatz Fauna,
California. Carnegie Inst., Washington, 514:33-38, 1 pi., May 18,
1939.
Wood, A. E.
1935. Evolution and relationship of the heteromyid rodents with new
forms from the Tertiary of western North America. Annals Carnegie
Museum, 24:73-262, 157 figs., 3 tables, May 13, 1935.
Transmitted May 15, 1948.
□
22-6114
(Continued from inside of front cover.)
15. A new hylid frog from eastern Mexico. By Edward H. Taylor.
Pp. 257-264, 1 figure in text. August 16, 1948.
16. A new extinct emydid turtle from the Lower Pliocene of Oklahoma.
By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.
17. Pliocene and Pleistocene records of fossil turtles from western Kan-
sas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284, 1 figure
in text. August 16, 1948.
18. A new species of heteromyid rodent from the Middle Oligocene of
northeastern Colorado with remarks on the skull. By Edwin G.
Galbreath. Pp. 285-300, 2 plates. August 16, 1948.
19. Speciation in the Brazilian spiny rats (Genus Proechimys, Family
Echimyidae). By Joao Moojen. Pp. 301-406, 140 figures in text.
December 10, 1948.
20. Three new beavers from Utah. By Stephen D. Durrant. Pp. 407-
417, 7 figures in text. December 24, 1948.
21. Two new meadow mice from Michoacan, Mexico. By E. Ray-
mond Hall. Pp. 423-427, 2 figures in text. December 24, 1948.
22. An annotated check list of the mammals of Michoacan, Mexico.
By E. Raymond Hall and Bernardo Villa R. Pp. 431-472, 5 figures
in text. December 27, 1949.
23. Subspeciation in the kangaroo rat, Dipodomys ordii. By Henry
W. Setzer. Pp. 473^573, 27 figures in text, 7 tables. December 27,
1949.
Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp.
1-444, 140 figures in text. April 9, 1948.
S-lvA-L
Geographic Range of the Hooded Skunk,
Mephitis macroura, with Description of a
New Subspecies from Mexico
BY
E. RAYMOND HALL and WALTER W. DALQUEST
MRi 1DV
MAR ~8 19! I
UNIVERSITY
University of Kansas Publications
Museum of Natural History
Volume 1, No. 24, pp. 575-580, 1 figure in text
January 20, 1950
University of Kansas
LAWRENCE
1950
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Edward H. Taylor,
A. Byron Leonard, Robert W. Wilson
Volume 1, No. 24, pp. 575-580, 1 figure in text
January 20, 1950
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1950
23-1544
IHUt). UUftJI . LUUL.
LIBRARY
HAR -8 13' I
Geographic Range of the Hooded Skunk, Mephitis
macroura, with Description of a New Subspecies
from Mexico
By
E. RAYMOND HALL AND WALTER W. DALQUEST
The hooded skunk, Mephitis macroura Liechtenstein, can be dis-
tinguished from the only other species in the genus, Mephitis me-
phitis Schreber, by the larger tympanic bullae, in the white-backed
color phase by having some black hairs mixed with the white hairs
of the back, and in the black-backed phase by having the two white
stripes widely separated and on the sides of the animal instead of
narrowly separated and on the back of the animal. The starting
point for taxonomic work with Mephitis is A. H. Howell's "Revision
of the skunks of the genus Chincha (N. Amer. Fauna, 20, 1901)."
Of the species Mephitis macroura, Howell (op. cit.) recognized three
subspecies: M. m. macroura, M. m. milleri, and M. m. vittata.
The species M. macroura is restricted to the arid region made up
mostly of the Mexican Plateau. Also, wherever the species occurs
beyond this Plateau, as for example in Guatemala, at San Mateo
del Mar in Oaxaca, in the vicinity of Piedras Negras in Veracruz,
and in southern Arizona, aridity is marked. Whether the species
has a continuous distribution from the southern end of the Mexican
tableland southward to Duenas in Guatemala is not known but it
is unlikely that the lowland population at San Mateo del Mar on
the Pacific slope of Oaxaca has contact with M . m. macroura of the
Mexican Plateau and it is almost certain that the population, which
is here named M. m. eximius, from the arid coastal plain of eastern
Mexico in Veracruz, has no connection with the upland population,
M. m. macroura. The lowest elevation on the eastern slope of the
Plateau from which we have record of the occurrence of this species
is 4,500 feet at Jico. All along the eastern slope of the Plateau,
between the elevations of approximately 2,000 and 4,500 feet, the
belt of lush, dense vegetation of the upper humid division of the
Tropical Life-zone constitutes a barrier to Mephitis and tends to
exclude the hooded skunk from the arid territory below the humid
belt. Another kind of skunk, Conepatus tropicalis, lives in the
humid belt, at least on the eastern side of the Mexican tableland.
How the population of Mephitis, which was sampled by us from
(577)
578 University of Kansas Pubs., Mus. Nat. Hist.
west and west-northwest of Piedras Negras, arrived there is unknown
but we think that its geographic range is not now connected with
that of the population on the Plateau. The same can be said of the
lowland population at San Mateo del Mar in Oaxaca. There, on the
Pacific slope of the Mexican tableland, the lower humid division of
the Tropical Life-zone probably has tended to restrict the spread
southward and westward of Mephitis; however, on this Pacific slope
the humid belt is less humid and it is less continuous, we think, than
on the Atlantic slope.
Four subspecies of Mephitis macroura may be recognized. They
are as follows:
Mephitis macroura milleri Mearns
1897. Mephitis milleri Mearns, Proc. U. S. Nat. Mus., 20:467, 1897.
1901. Mephitis macroura milleri, Bull. Amer. Mus. Nat. Hist., 14:334,
November 12, 1901.
Type locality. — Fort Lowell, Pima County, Arizona.
Range. — Northwestern Mexico and southeastern Arizona. See figure 1.
Marginal occurrences (unless otherwise indicated, after Howell, N. Amer.
Fauna, 20:42, 43, 1901) are: Arizona: Santa Catalina Mountains; Tucson;
Fort Lowell. Chihuahua: Casas Grandes; Chihuahua (City). Coahuila:
La Ventura. Chihuahua: Guadalupe y Calvo (mountains near). Sonora:
Camoa; Hermosillo; Sierra Cubabi (Burt, Miscl. Publ., Mus. Zool., Univ.
Michigan, 39:30, 1938).
Characters. — Long skull ($Q0 mm, $56 mm) and large ml.
Mephitis macroura macroura Lichtenstein
1832. Mephitis macroura Lichtenstein, Darstellung Siiugethier, pi. 46,
with accompanying text, 1832.
Type locality. — Mountains northwest of the City of Mexico.
Range. — Southern half of Mexican Plateau and south to Guatemala. See
figure 1. Marginal occurrences (all from Howell, N. Amer. Fauna, 20:41, 42,
1901) are: Tamaulipas: Jaumave. Veracruz: Las Vigas; Jico; Orizaba.
Puebla: Tehuacan. Guatemala: Duehas (vicinity). Oaxaca: 15 mi. W
Oaxaca. Colima: Hacienda Magdalena. Jalisco: San Sebastian. Tepic:
Santa Teresa. Zacatecas: Valpariso.
Characters. — Skull of medium size (basal length, $ 56, 5 54) ; tail averag-
ing shorter than head and body.
Mephitis macroura vittata Lichtenstein
1832. Mephitis vittata Lichtenstein, Darstellung Siiugethier, pi. 47,
with accompanying text, 1832.
1901. Mephitis macroura vittata, Allen, Bull. Amer. Mus. Nat. Hist.,
14:334, November 12, 1901.
Type locality.— "San Matteo el Mar" [= San Mateo del Mar], Oaxaca.
Range.— Known only from the type locality. See figure 1.
Characters.— Skull short (5 54.6, $52.3); narrow across mastoid processes;
tail long; body short.
Geographic Range of Mephitis macroura
579
Mephitis macroura eximius new subspecies
Type.— Female, adult, skin with skull, No. 19272, Mus. Nat. Hist., Univ.
Kansas; 15 kilometers west of Piedras Negras, 300 feet elevation, Veracruz,
Mexico; 13 January 1947; obtained by J. Mazza and Walter W. Dalquest;
original No. 7017, W. W. Dalquest.
Range. — From the vicinity of the type locality on the arid coastal plain of
the lowlands of central Veracruz. See figure 1.
Diagnosis. — Size small (see measurements) ; tail long, ranging from 110 to
133 percent of length of head and body; color black, with white areas con-
taining a few black hairs, and in non-hooded phase with white lateral stripes
low on sides of body and in some specimens almost absent; skull small but
broad across mastoid processes.
"1
M.m.miHeri 2
M m. macroura 3. M m. vittata 4 M. m. eximius
Fig. 1. Map showing the geographic ranges of the four subspecies of the
species Mephitis macroura.
Comparisons. — From Mephitis macroura macroura of the southern part of
the Mexican Plateau, M. m. eximius differs in shorter head and body, relatively
(to body) longer tail, and smaller skull. From Mephitis macroura vittata of
the tropical lowlands of the Pacific slope of Oaxaca, M. m. eximius differs in
slightly larger average size throughout and relatively longer tail.
Remarks. — M. m. eximius is regarded as a subspecies of M. ma-
croura because there is some overlap in size between larger indi-
viduals of M. m. eximius and smaller individuals of M. m. macroura.
Actually, as indicated above, we doubt that the geographic ranges
580 University of Kansas Pubs., Mtjs. Nat. Hist.
of the two subspecies are continuous or that the geographic range
of M. m. eximius is continuous with the geographic range of M. to.
vittata. Small size and relatively long tail characterize both of the
lowland, tropical subspecies, eximius and vittata, whereas the two
upland subspecies of the temperate areas are larger and have rela-
tively shorter tails.
Habitat closely resembling that at the type locality extends from
the southern base of the first mountains north of Jalapa southward
as far as the north base of the Tuxtla Mountains — a distance of
approximately 110 miles from northwest to southeast along the gulf
coast. None of our 5 skins shows the hooded color-pattern so com-
mon on the Mexican Plateau and in vittata of Oaxaca. One of our
five specimens has well-developed lateral stripes ; three have greatly
reduced lateral stripes and one is black except for a white spot on
the right hip.
Measurements. — An adult male (University of Kansas Museum of Natural
History Catalogue Number, 17900), a subadult male (19273), and adult female
(19272, the holotype) and a subadult female (19902), measure, in millimeters,
respectively, as follows: Total length, — , 599, 578, 583; length of tail, — ,
319, 335, 305; length of hind foot, 58, 62, 58, 60; basal length of skull, 56.1,
55.0, 52.8, 53.1; basilar length of Hensel, 53.6, 52.6, 50.3, 51.2; greatest zygo-
matic breadth, 41.6, 38.0, 39.0, 37.0; greatest mastoid breadth, 34.6, 34.3, 33.3,
31.5; breadth across postorbital processes, 22.2, 20J2, 20.5, 21.0; least inter-
orbital breadth, 20.3, 18.2, 19.0, 18.5; palatal length, 242, 25.1, 24.2, 24.0; post-
palatal length, 31.5, 29.6, 28.8, 29.0; foramen magnum to plane of last molars,
30.8, 29.4, 27.5, 29.0.
Specimens examined. — Total number, 5, all from Veracruz, Mexico, as follows: Ri6
Blanco, 20 km. WNW Piedras Negras, 3; 15 km. W Piedras Negras, 300 ft., 2.
University of Kansas Museum of Natural History, Lawrence, Kansas.
Transmitted October 81, 1949.
□
23-1544
S-A/A-L-
Pipistrellus cinnamomeus Miller 1902
Referred to the Genus Myotis
BY
E. RAYMOND HALL and WALTER W. DALQUEST
r
LIBRARY
MAR
-8
if
50
1
University of Kansas Publications
Museum of Natural History
Volume 1, No. 25, pp. 581-590, 5 figures in text
January 20, 1950
University of Kansas
LAWRENCE
1950
University of Kansas Publications, Museum op Natural History
Editors: E. Raymond Hall, Chairman, Edward H. Taylor,
A. Byron Leonard, Robert W. Wilson
Volume 1, No. 25, pp. 581-590, 5 figures in text
January 20, 1950
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND. JR.. STATE PRINTER
TOPEKA. KANSAS
1950
23-1545
riAR -8 !•• I
■
Pipistrellijis cinnamomeiis Miller 1902
Referred to the Genus Myotis
By
E. RAYMOND HALL AND WALTER W. DALQUEST
Miller (Proc. Acad. Nat. Sci. Philadelphia, 1902, p. 390, Septem-
ber 3, 1902) based the name Pipistrellus cinnamomeus on a skin and
skull of a vespertilionid bat obtained on May 4, 1900, at Monte-
cristo, Tabasco, Mexico, by E. W. Nelson and E. A. Goldman. A
single specimen was available to Miller when he proposed the name
P. cinnamomeus. Dalquest and Hall (Jour. Mamm., 29:180, May
14, 1948) reported three additional specimens collected in 1946 by
W. W. Dalquest on the Rio Blanco, twenty kilometers west-north-
west of Piedras Negras, Veracruz, Mexico. No other published in-
formation concerning this species is known to us, although the name
has, of course, appeared in regional lists, for example in the "List
of North American Recent Mammals, 1923" (Bull. U. S. National
Museum, 128:75, April 29, 1924) by Gerrit S. Miller, Jr.
Additional specimens, nevertherless, are known. Two collected on
April 18 and 20, 1903, at Papayo, Guerrero, by Nelson and Goldman,
are in the Biological Surveys Collection in the United States Na-
tional Museum. A skin, probably of this species, for which the
skull cannot now be found, was taken on October 27, 1904, at Es-
quinapa, Sinaloa, by J. H. Batty and is in the American Museum of
Natural History. This is the skin referred by Miller and Allen
(Bull. U. S. Nat. Mus., 144:100, May 25, 1928) to Myotis occultus.
Three additional specimens, each a skin with skull, were collected
twenty kilometers east-northeast of Jesus Carranza, at 200 feet ele-
vation, Veracruz, by Walter W. Dalquest, two on April 13, 1949,
and one on May 16 of the same year. These are in the Museum of
Natural History of the University of Kansas, as also are the three
previously reported by Dalquest and Hall (loc. tit.). A total of ten
specimens, from five localities, all in Mexico, thus is accounted for.
On page 392 of the original description — which our study of the
holotype shows to be accurate — Miller wrote: "This bat differs so
widely from the other known American species of Pipistrellus as to
need no special comparisons. Superficially it has much the appear-
ance of an unusually red Myotis lucifugus, and only on examination
of the teeth do the animal's true relationships become apparent."
(583)
584 University of Kansas Pubs., Mus. Nat. Hist.
In referring to the teeth Miller almost certainly was thinking of the
premolars of which there are only two on each side of the upper
jaw and on each side of the lower jaw in Pipistrellus, including his
Pipistrellas cinnamomeus, whereas Myotis at that time was thought
always to have three premolars on each side of both the upper and
lower jaw, except in rare instances where one premolar might be
lacking on one side of one jaw or even more rarely on both sides of
the upper jaw. In his original description of P. cinnamomeus, Mil-
ler mentioned also that it had the "Inner upper incisor distinctly
smaller than the outer, not approximately equal to it as is the case
in P. subfiavus."
At this point it is well to make clear that each of the genera
Pipistrellus and Myotis contains a large number of species and that
the differences between the two genera are few. Our examination
of American specimens reveals only one differential character: In
Myotis the outer upper incisor is distinctly larger than the inner,
whereas the two incisors are of approximately equal size in Pipistrel-
lus. It may be noted that the outer upper incisor of several, but
not all, species of Myotis has a well-developed concave surface di-
rected toward the canine whereas this surface is flat or convex in
Pipistrellus. In both features, the type of Pipistrellus cinnamomeus
Miller agrees with Myotis and differs from Pipistrellus.
Five years after naming and describing Pipistrellus cinnamomeus,
Miller published his monumental work entitled "The families and
genera of bats" (Bull. U. S. Nat. Mus., 57, June 29, 1907) wherein
he points out the differences in the upper incisors between Pipistrel-
lus and Myotis (by a lapsus plumae ascribes subequal incisors to
Myotis and unequal incisors to Pipistrellus) but seemingly failed to
reexamine P. cinnamomeus in the light of this better understanding
of the two genera, or if he did examine P. cinnamomeus he possibly
was misled still by the absence of the third premolar on each side
of both the upper and lower j aw.
In 1928 when Miller and Allen published their account of "The
American bats of the Genera Myotis and Pizonyx" (Bull. U. S. Nat.
Mus., 144, May 25, 1928) they examined specimens of Myotis oc-
cultus which they implied (op. cit.: 99-100) had only two instead
of three premolars on each side of both the upper and lower jaws.
In preparing this taxonomic account of bats of the genus Myotis,
the specimens (type and two from Papayo) of Pipistrellus cinnamo-
meus seem not to have been examined. Indeed, it is almost certain
that they were not examined for the species was renamed; the new
PlPISTRELLUS CINNAMOMEUS MlLLER 585
name, Myotis lucifugus fortidens Miller and Allen (Bull. U. S. Nat.
Mus., 144:54, May 25, 1928), was based on a skull with the cor-
responding body in alcohol. The characters of this specimen are
almost exactly those of Pipistrellus cinnamomeus, named and de-
scribed by Miller 26 years earlier. The type locality (Teapa) of
M. I. fortidens is 80 miles westerly from the type locality of P. cin-
namomeus; both are in the state of Tabasco, and in the same life-
zone, at equivalent elevations (neither higher than 50 meters). Since
there are no characters of taxonomic worth to distinguish the two
named specimens, Myotis lucifugus fortidens Miller and Allen 1928
falls as a synonym of Pipistrellus cinnamomeus Miller 1902. But,
according to Miller and Allen (Bull. U. S. Nat. Mus., 144:19, 197),
Vespertilio cinnamomeus Wagner 1855 is a name based on Myotis
ruber (E. Geoffroy, 1806) from Paraguay and hence Myotis cin-
namomeus (Miller) 1902 is a homonym of Myotis cinnamomeus
(Wagner) 1855 and is unavailable for the animal from Montecristo
when it is transferred to the genus Myotis; the species of animal
concerned will take the next available name, which seems to be
Myotis lucifugus fortidens Miller and Allen 1928.
It may reasonably be asked if Myotis and Pipistrellus should be
retained as separate genera if the only constant difference between
the two is subequal versus unequal upper incisors. In our opinion
it would be worth-while for someone who had access to adequate
material from both the Old World and the New World to investigate
this question. We lack adequate material from the Old World.
When Miller and Allen named M. I. fortidens they had only two
specimens, the holotype from Teapa, Tabasco, and a referred speci-
men from Fort Hancock, El Paso County, Texas, approximately
1,200 miles north-northwest of Teapa. We have examined this
specimen from Texas (U. S. Nat. Mus., 21083/36121, skin and skull)
and regard it as Myotis lucifugus carissima Thomas. Furthermore,
we regard the holotype of Myotis lucifugus fortidens Miller and
Allen 1928 as specifically distinct from Myotis lucifugus of Miller
and Allen 1928. The Cinnamon Myotis, described below, therefore
may stand as:
586
University of Kansas Pubs., Mus. Nat. Hist.
Myotis fortidens Miller and Allen
Cinnamon Myotis
Pipistrellus cinnamomeus Miller, Proc. Acad. Nat. Sci. Philadelphia,
p. 390, September 3, 1902, type from Montecristo, Tabasco (preoccupied
by Vespertilio cinnamomeus Wagner, Schreber's Saugethiere, suppl,
5:755, 1855, a renaming of Vespertilio ruber E. Geoffroy Saint-Hilaire).
Myotis lucifugus fortidens Miller and Allen, Bull. U. S. Nat. Mus.,
144:54, May 25, 1928.
Type.—" Adult female (in alcohol) No. 88.8.8.18, British Museum (Natural
History). Collected at Teapa, Tabasco, Mexico, by H. H. Smith, January 5,
1888. Presented by Messrs. Salvin and Godman [after Miller and Allen, Bull.
U. S. Nat. Mus, 144:54, May 25, 1928]."
Range. — Known only from the lower part of the Tropical Life-zone of the
region of the Isthmus of Tehuantepec and east and west coasts of Mexico.
116
108 100
92
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0 50 100 200 300 MILES ~^^iltll§^^
I — 1 — 1 1 1 ^^§^j^^53
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lb
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108 100
92
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Fig. 1. Map showing localities from which Myotis fortidens has been re-
corded.
Diagnosis. — Among American species of the genus, over-all size medium
(total length 94 mm) ; body long (54) ; tail short (39) ; forearm of medium
length (37) ; tibia short (14.5) ; foot long (58 per cent of length of tibia) ; wing
membrane arising from side of foot at distal end of metatarsal; calcar simple
(not keeled) and 7 mm long; ears 15 to 16 mm long measured in the flesh
from the notch (posteroventral border of the meatus) ; tragus, measured from
same place, 7 to 8 mm high with posterobasal lobe; third metacarpal longest
and second metacarpal shortest; fifth shorter than fourth; ears brownish;
membranes of wing and tail blackish; uropatagium almost hairless, the few
hairs that are present being almost invisible; pelage of back 5 mm long with
some overhairs 8 to 9 mm long ; basal 3 mm of fur black, remainder Cinnamon-
PlPISTRELLUS CINNAMOMEUS MlLLER
587
Brown (capitalized color terms, after Ridgway, Color Standards and Color
Nomenclature, Washington, D. C, 1912) ; outline of skull viewed dorsally simi-
lar to that of Myotis lucifugus; sagittal crest well developed; distance across
upper canines equal to or slightly exceeding interorbital constriction; brain-
case low; two premolars on each side in upper jaw and also in lower jaw, the
one remaining small premolar in contact with both the canine and the fourth
premolar.
Remarks. — Myotis jortidens is known only from the Tropical
Life-zone. The skin, without a skull, from Esquinapa, Sinaloa,
agrees in color with the undoubted specimens of M. jortidens from
Papayo, Guerrero, but can be matched also by selected skins of
Myotis occultus from Blythe, Riverside County, California. With-
out the skull the reference of this specimen to M. jortidens is pro-
visional. Reason for referring it to jortidens rather than to M.
Figs. 2-5. Four views of the skull of Myotis jortidens. No. 32112, Uni-
versity of Kansas Museum of Natural History, $ , obtained 20 kilometers
east-northeast Jesus Carranza, 200 feet elevation, Veracruz, Mexico, on May 16,
1949, by Walter W. Dalquest; original no. 12869. X 2.
occultus is provided, however, by a series of eleven specimens of M.
occultus from Alamos, Sonora. These are Saccardo's Umber rather
than Cinnamon-Brown and they are geographically intermediate be-
tween the reddish M. occultus of California and the reddish M.
jortidens of Mexico. Furthermore, these specimens from Alamos
have large skulls of slightly different proportions than those of M.
jortidens or than those of M. occultus from California; possibly the
animals from Alamos are representative of the larger, duller-colored
variation for which Hollister proposed the name Myotis baileyi
(Proc. Biol. Soc. Washington, 22:44, March 10, 1909). This duller-
colored type of animal intervenes between the geographic ranges of
undoubted M. occultus and undoubted M. jortidens. The specimen
from Esquinapa, in the geographic sense, is on the jortidens side
rather than on the occultus side of the baileyi population. This geo-
graphic position is the basis on which the specimen from Esquinapa
is referred to M. jortidens. The third premolar is lacking from each
side of both the upper and the lower jaws of each individual of this
series from Alamos.
588 University of Kansas Pubs., Mus. Nat. Hist.
The specimens of M. fortidens are all distinguishable by their
color from other kinds of Myotis found in the same area. Occasional
individuals of Myotis velifer, as for example three from Las Vigas,
Veracruz, also are reddish but they are of brighter tone. In addi-
tion, the larger size and cranial features of these specimens of M.
velifer permit ready differentiation of them from specimens of M.
jortidens. One specimen (No. 32113) of M. fortidens from twenty
kilometers east-northeast of Jesus Carranza is lighter than the others,
being near (j) Cinnamon-Brown above and is lighter on the under-
pays than on the upper parts. Another individual (No. 32112) is
duller colored than the others, being Snuff Brown both above and
below. Otherwise the specimens of M . fortidens agree in color.
Among named kinds of Myotis, M. fortidens resembles Myotis
lucifugus and Myotis occultus. From the former, M. fortidens dif-
fers in possessing a strong sagittal crest and in lacking the third
premolar in both the upper jaw and the lower jaw. M. fortidens
lacks the glossy sheen found on the pelage of many individuals of
M. lucifugus. From M. occultus, M. fortidens differs in having the
rostrum (viewed from above) smaller in relation to the braincase.
This is true of specimens with the teeth showing much wear as well
as in specimens with the teeth unworn or only moderately worn.
Also, M. fortidens is longer bodied as may be seen by comparing the
measurements given here with those recorded for M . occultus by
Miller and Allen (Bull. U. S. Nat. Mus., 144:100, May 25, 1928).
We are agreed that M. fortidens is as closely related to M. occultus
as to any other named kind of Myotis, and that it is more closely
related to it than to most other species of the genus, but one of us
(Dalquest) thinks that M. fortidens is specifically distinct from M.
occultus, whereas the other author (Hall) inclines to the view that
additional specimens from localities intermediate between the known
geographic ranges of M. occultus and M. fortidens will reveal inter-
gradation between the two kinds. However that may be, there is
no proof at present of such intergradation and the binomial is there-
fore used for the Cinnamon Myotis.
Specimens examined. — Total number, 10, all from Mexico, each a skin with skull except
the skin-only from Sinaloa. Sinaloa: Esquinapa, 1 (Amer. Mus. Nat. Hist.). Guerrero:
Papayo, 2 (U. S. Biological Surveys Collection). Veracruz: 20 km. WNW Piedras Negras,
3 (Mus. Nat. Hist., Univ. Kansas); 20 km. ENE Jesus Carranza, 200 ft. elevation, 3 (Mus.
Nat. Hist., Univ. Kansas). Tabasco: Montecristo, 1 (U. S. Biological Surveys Collection).
Additional record. — Tabasco: Teapa, the holotype of Myotis lucifugus fortidens Miller
and Allen 1928.
University of Kansas Museum of Natural History, Lawrence, Kansas.
Transmitted October SI, 1949.
PlPISTRELLUS CINNAMOMEUS MlLLER
589
Cranial Measurements of Myotis fortidens
a
o
CO
£
A
a
o3
■s
•f-H
CI
O
a>
CO
c3
'3
*
o
1
Xi
+3
o3
03
o
i
Xi
O
o
60
u
o
t-.
o
CD
-4->
-u
to
c3
-Q
"3
o
+3
_Q
u
03
GO
X2
O
03
13
O
a
3
tf
03
c3
3
Sex
CD
>>
a
o
-(J
T3
F3
• ^H
T3
o
No. Age
Locality
CI
bC
a3
CD
a
e3
03
o3
a
03
a
O
>>
S3
a
i— i
PQ
§
£
S
2
&
25030 5
Esquinapa
?
126650 9
Papayo
15.0
14.2
9.7
3.9
7.1
11.5
5.5
5.6
6.0
6
126651 9
Do.
15.1
13.8
9.4
3.8
6.8
10.6
5.6
5.9
6.0
0
17834 5
P. Negras i
• • • •
4.1
10.6
5.6
5.7
6.0
0
17835 9
Do.
15.5
14.9
9.6
4.2
7.2
11.0
5.7
6.0
6.1
2
17S36 9
Do.
15.5
14.5
9.7
4.2
7.3
10.9
5.4
5.9
5.7
3
32112^
J.Carranza2
15.3
14.4
9.7
4.1
7.3
11.5
5.7
5.9
6.3
1
32113 4
Do.
15.0
14.0
9.5
4.2
7.2
10.9
5.5
5.9
5.9
1
32114 5
Do.
15.0
13.9
9.7
4.1
7.2
10.8
5.4
6.0
5.9
1
88.8.8.1893
Teapa
15.0
13.8
9.6
3.8
7.4
5.4
5.8
5.8
1
100231 9 4
Montecristo
15.0
14.1
9.0
4.0
7.2
11.4
5.S
6.0
0
Average
15.2
14.2
9.5
4.0
7.2
11.0
5.6
5.9
6.0
1. 20 km. WNW Piedras Negras.
2. 20 km. ENE Jesus Carranza, 200 ft.
3. Type of Myotis lucifugus fortidens; measurements after Miller and Allen, Bull. U. S.
Nat. Mus., 144 :100, 101, May 25, 1928.
4. Type of Pipistrellus cinnamomeus Miller 1902.
External Measurements of Myotis fortidens
>>
03
"c3
X\
x\
bD
T3
O
03
O
03
o3
O
o3
0
+3
O
CI
Sex
C
CD
T3
CI
03
T5
03
a
03
a
CD
a
CD
a
a
0
u
No. Age
Locality
-4J
o
o3
CD
03
-Q
o
o
O
3
'x\
03
H
w
h
H
Ph
fr
H
£
W
25030 5
Esquinapa
14.2
8.11
35.6
5.5
33.3
30.8
126650 9
Papayo
14.7
8.21
38.3
5.4
35.1
32.4
126651 9
Do.
14.8
7.91
35.6
5.7
32.7
31.1
17834 5
P. Negras 2
95
55
40
14.7
9.01
37.0
5.7
33.8
32.0
15
17835 9
Do.
93
55
38
15.6
9.41
37.5
6.0
35.4
32.2
15
17836 9
Do.
94
55
39
14.3
8.41
37.6
6.0
34.5
32.7
15
32112 5
J.Carranza3 94
53
41
14.5
8.91
38.2
5.0
35.1
33.8
16
32113 5
Do.
94
57
37
14.2
8.01
36.5
5.3
34.9
32.7
16
32114 5
Do.
90
53
37
• • • •
• • •
37.0
5.1
34.2
33.0
16
88.8.8.189 4
Teapa
46
39
15.6
8.0
38.6
6.2
34.8
33.0
100231 9 5
Montecristo 99
56
44
15.4
9.6
37.0
6.0
■ • ■ ■
• • • •
Average
94
53.8 39.414.8
8.6
37.2
5.6
34.4
32.4
15.5
1. Measured on the dried skin.
2. 20 km. WNW Piedras Negras.
3. 20 km. ENE Jesus Carranza.
4. Type of Myotis lucifugus fortidens; measurements after Miller and Allen, Bull. U. S.
Nat. Mus., 144:100, 101, May 25, 1928.
5. Type of Pipistrellus cinnamomeus Miller 1902.
□
23-1545
S-fVA-L
A Synopsis of the American Bats
of the Genus Pipistrellus
BY
E. RAYMOND HALL and WALTER W. DALQUEST
MUS. COMP. ZOOL
LIBRARY
MAR -8 1450
Ui
University of Kansas Publications
Museum of Natural History
Volume 1, No. 26, pp. 591-602, 1 figure in text
January 20, 1950
University of Kansas
LAWRENCE
1950
University of Kansas Publications, Museum of Natural History
Editors: E. Raymond Hall, Chairman, Edward H. Taylor,
A. Byron Leonard, Robert W. Wilson
Volume 1, No. 26, pp. 591-602, 1 figure in text
January 20, 1950
University of Kansas
Lawrence, Kansas
PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA. KANSAS
1950
23-1546
PS. CO^IP. ZOOL.
LIBRARY
HAR ~8 !" !
A Synopsis of the lAmbrican Bats
of the Genus Pipistrellus
By
E. RAYMOND HALL AND WALTER W. DALQUEST
Four nominal species of the genus Pipistrellus are currently recog-
nized in North America. They are Pipistrellus subjlavus (F. Cuvier)
of eastern North America, Pipistrellus hesperus (H. Allen) of west-
ern North America, Pipistrellus veracrucis (Ward) from Veracruz,
Mexico, and Pipistrellus cinnamomeus Miller from Tabasco, Mexico.
In the past three years, specimens have been obtained in Veracruz
(by Dalquest) of each of the southern species. One of these, P.
cinnamomeus, previously was known from a single specimen; the
other, P. veracrucis, was known only from six specimens which now
are lost or misplaced. The results of our study of these recently
acquired Mexican specimens constitute our principal contribution in
this paper; we have done little more with the material from the
United States and Canada than to codify the findings of other mam-
malogists with respect to the systematic status and geographic dis-
tribution.
Study of the available specimens reveals that there are only two
species, Pipistrellus hesperus and Pipistrellus subflavus; Pipistrellus
veracrucis proves to be only a subspecies (geographic race) of P.
subflavus, and Pipistrellus cinnamomeus proves to be a species of
another genus, Myotis (see Hall and Dalquest, page 583 of this
volume).
Genus Pipistrellus Kaup
1829. Pipistrellus Kaup, Skizzirte Entw.-Gesch. u. natiirl. Syst. europ.
Thierw., Vol. 1, p. 98, Type, Vespertilio pipistrellus Schreber (not seen
by us, after Miller, N. Amer. Fauna, 13:87, 1897).
Range in the New World. — In North America from southern Canada to
Honduras (47 degrees to 5 degrees North Latitude) and from the Atlantic to
the Pacific; not recorded from the West Indies or South America.
Characters. — Size small; tail approximately as long as outstretched leg;
ears well developed with prominent tragus; dental formula: i.f; c.|; p.f; m.| ;
two upper incisors subequal and outer one lacking a concavity on surface facing
canine; dentition otherwise essentially as in Myotis Kaup except that third
premolar is always, instead of rarely, absent.
Remarks. — There are two species in North America. Their geo-
graphic ranges, as now known, meet, but do not overlap. Certain
(593)
594
University of Kansas Pubs., Mus. Nat. Hist.
differences between the two species are listed in the parallel columns
below. Most of these differences in the skull and teeth are illustrated
in figures 22 and 23 on page 92 of Miller's "Revision of the North
American bats of the family Vespertilionidae (N. Amer. Fauna, 13,
1897)."
Structure
P. hesperus
P. subflavus
Color
Predominately gray
Predominately brown
Foot
Less than half as long
as tibia
More than half as long as
tibia
Thumb, length of
Less than 4.9 mm.
More than 4.9 mm.
Tragus
Blunt, terminal part
bent forward
Narrow, straight
Skull
(dorsal profile)
Nearly straight
Dish-faced
Braincase
(viewed from above)
Small
Large
Palate
Extending far behind
molars; spine short,
narrow at base
Extending short distance
behind molars; spine
long, wide at base
12
Unicuspidate
Bicuspidate
13
Accessory cusp present
on anterointernal
face
Accessory cusp absent on
anterointernal face
PI (occlusal view)
Less than a seventh as
large as canine
More than a seventh as
large as canine
PI (labial view)
Concealed by Cl and P4
Not concealed
P4
Touching canine
Not touching canine
i3
Touching i2 and cl
Separated by space from
i2 and cl
p3
Lower than anterior
cusp of canine
As high as anterior cusp
of canine
Distance from cl
to ml
Less than length of m2 ;
premolars crowded
More than length of m2;
premolars less crowded
Synopsis of Genus Pipistrellus 595
Pipistrellus hesperus
(Synonomy under subspecies)
Range. — Arid Sonoran life-zones of western North America from Washing-
ton southward to Jalisco.
Characters. — Smoke Gray to Buff Brown (Capitalized color terms after
Ridgway, Color Standards and Color Nomenclature, Washington, D. C, 1912)
dorsally; total length, 60 to 86; foot less than half as long as tibia; tragus
blunt with terminal part bent forward; skull nearly straight in dorsal profile;
inner upper incisor unicuspidate; outer upper incisor with accessory cusp on
anterointernal face; PI, viewed from occlusal face, less than a seventh of area
of canine, and from labial aspect concealed by canine and fourth premolar;
lower, third premolar lower than anterior cusp of canine; lower premolars
crowded, distance between canine and first molar less than length of second
lower molar.
Remarks. — In the United States and in the northern part of Mex-
ico, P. hesperus is the smallest bat found. Little is known about its
habits. It emerges earlier in the evening than other species of bats.
The frequency with which it is seen near cliffs suggests that it finds
concealment under rocks. In winter, in Nevada (Hall, Mammals of
Nevada, p. 150, 1946), P. hesperus has been found singly in crevices
in the roofs of mine tunnels.
In the United States National Museum in July, 1949, the specimen
providing the easternmost record station of occurrence was examined
by us. This is No. 23591, in alcohol, taken on August 24, 1890, by
William Lloyd, original No. 88, at the mouth of the Pecos River in
Texas. In the same collection there is a specimen of Pipistrellus
subflavus providing the westernmost record of occurrence of that
species. This specimen, a skin with skull, is No. 126729, $ , taken
on May 3, 1903, by Jas. H. Gaut, original No. 1271, at Comstock,
Texas. The two localities concerned are in the Valley of the Rio
Grande, and are only about five miles apart. Nevertheless, the two
specimens are clearly referable to their respective species and show
no tendency toward intergradation. Consequently, confidence is felt
in treating Pipistrellus hesperus and Pipistrellus subflavus as two
distinct species.
The most recent report upon geographic variation throughout the
entire species, Pipistrellus hesperus, was that by Hatfield (Jour.
Mamm., 17:257-262, August 14, 1936). Later, as explained below
in the account of P. h. australis, Burt (Miscl. Publ., Mus. Zool.,
Univ. Michigan, 39:25, February 15, 1938) examined specimens from
Sonora, Mexico, and for them and for specimens from southern Ari-
zona proposed a different nomenclatural arrangement.
596
University of Kansas Pubs., Mus. Nat. Hist.
Pipistrellus hesperus
Pipistrellus subflavus
Fig. 1. Map showing the geographic ranges of species and subspecies of
Pipistrellus.
1. Pipistrellus h. hesperus 5. Pipistrellus h. santarosae
2. Pipistrellus h. merriami 6. Pipistrellus s. subflavus
3. Pipistrellus h. australis 7. Pipistrellus s. obscurus
4. Pipistrellus h. maximus 8. Pipistrellus s. veracrucis
Pipistrellus hesperus hesperus (H. Allen)
Scotophilus hesperus H. Allen, Smithsonian, Miscl. Coll., No. 165,
Vol. 7 (art. 1) : p. 43, June, 1864.
Vesperugo hesperus True, Proc. U. S. Nat. Mus., 7:602, 1885.
Pipistrellus hesperus Miller, N. Amer. Fauna, 13:88, October 16, 1897.
Type locality.— Old Fort Yuma, Imperial County, California, on right bank
of Colorado River, opposite present town of Yuma, Arizona.
Range. — Intermontane region of the United States from south-central Wash-
ington south to Catavifia, Baja California, and from southeastern California
eastward to southeastern Utah. Marginal occurrences (unless otherwise indi-
cated, after Hatfield, Jour. Mamm., 17:258, 1936) are: Washington (Dalquest,
Univ. Kansas Publ., Mus. Nat. Hist., 2:165, 1948): Maryhill; Vantage; Al-
mota. Oregon: Watson. Idaho: 8 mi. W Rogerson (Davis, Mamms. Idaho,
p. 120, 1939). Nevada: Middle Stormy Spring (Hall, Mamms. Nevada, p.
Synopsis of Genus Pipistrellus 597
151, 1946). Utah: Goodridge. Arizona: 11 mi. NW Kayenta; Tinajas Altas.
Baja California: Catavifia; San Jose; Laguna Hanson. California: Dos
Palmos Spring; Banning; Victorville; 12 mi. below (down river) Bodfish;
Little Lake; 2 mi. S Benton Station. Nevada: 2 mi. NW Morgans Ranch;
Deephole. Oregon: Princeton.
Diagnosis. — Size medium for the species; total length, 71.8(66-74) ; tibia,
12.0(10.7-13.5) ; forearm, 29.4(27.8-31.8) ; greatest length of skull, 11.9(11.5-12.3) ;
breadth of braincase, 6.3(6.1-6.4). Color between Drab Gray and Smoke Gray,
dorsally; between Smoke Gray and Pale Smoke Gray, ventrally (after Hat-
field, Jour. Mamm, 17:257, 1936).
Pipistrellus hesperus merriami (Dobson)
Vesperugo merriami Dobson, Ann. and Mag. Nat. Hist., 18(ser. 5) :
124, August, 1896.
Pipistrellus hesperus merriami Grinnell, Proc. California Acad. Sci.,
3(ser. 4) :279, August 28, 1913.
Type locality .—Red Bluff, Tehama County, California.
Range.— California west of the Sierra Nevada; the Sacramento Valley, the
San Joaquin Valley, and the Coast Range from San Francisco Bay south to
San Diego County. Marginal occurrences (after Hatfield, Jour. Mamm., 17:260,
1936, unless otherwise noted) are: California: Dales on Paines Creek; Fyffe;
Yosemite Valley; Shaver Ranger Station; Springville; Fort Tejon; Painted
Gorge (P. H. Krutzsch, MS); Carrizo Creek; thence northward up the coast
probably to San Francisco Bay ; in the Sacramento Valley west to Rumsey .
Diagnosis. — Size medium for the species; total length, 71.3(66-78); tibia,
11.2(10.6-11.7) ; forearm, 28.9(27.5-30.8) ; greatest length of skull, 11.8(11.3-12.2) ;
breadth of braincase, 6.4(6.0-6.6). Color Buffy Brown to Army Brown, dorsally;
Wood Brown to Buffy Brown, ventrally (after Hatfield, op. cit.: 258, 260).
Pipistrellus hesperus australis Miller
Pipistrellus hesperus australis Miller, N. Amer. Fauna, 13:90, October
16, 1897.
Pipistrellus hesperus apus Elliot, Field Columb. Mus., pub. 90, zool.
ser., 3:269, March 8, 1904. Type from Providencia Mines, Sonora,
Mexico.
Type locality. — Barranca Ibarra, Jalisco, Mexico.
Range. — Central Arizona south to Jalisco and including the southern half
of Baja California. Marginal occurrences (after Hatfield, op. cit.: 261, unless
otherwise indicated) are: Arizona: Camp Verde; Fort Bowie. Sonora:
Pilares (Burt, Miscl. Publ., Mus. Zool., Univ. Michigan, 39:24, 1938). Jalisco:
Barranca Ibarra (Miller, orig. descr.). Baja California: Miraflores; San
Ignacio. Arizona: Bates Well.
Diagnosis. — Size small for the species; total length, 67.1(60-72); tibia,
11.3(10.1-12.3) ; forearm, 28.4(26.3-30.0) ; greatest length of skull, 11.7(11.3-12.0) ;
breadth of braincase, 6.1(5.9-6.3). Color: between Cinnamon Drab and Drab,
dorsally; Wood Brown to Light Drab, ventrally (after Hatfield, op. cit.:260).
Remarks. — Hatfield (op. cit.) examined no specimens from Mex-
ico (Baja California excepted) and Burt (op. cit.) who did examine
598 University of Kansas Pubs., Mus. Nat. Hist.
some specimens (from Sonora), referred one from northwestern
Sonora to P. h. hesperus and those from northeastern Sonora to P. h.
merriami. Since our treatment of subspecies of Pipistrellus (P. h.
veracrucis excepted) aims merely to reflect the latest systematic
treatment accorded the animals, we would follow Burt (op. tit.)
were it not for the fact that he shows the geographic range of P. h.
merriami separated by the range of P. h. hesperus into two parts.
This is inconsistent with the ordinarily accepted concept of sub-
species. Consequently, we have followed Hatfield (op.cit.). Clearly,
a critical study is needed of adequate material of Pipistrellus hes-
perus of Mexico.
Pipistrellus hesperus maximus Hatfield
Pipistrellus hesperus maximus Hatfield, Jour. Mamm, 17:261, August
14, 1936.
Type locality. — Dog Spring, Hidalgo County, New Mexico.
Range. — Southern New Mexico, western Texas and probably the adjoining
parts of Mexico. Marginal occurrences (after Hatfield [op. cit. :261] except
as otherwise indicated) are: New Mexico: Animas Valley; Florida Mountains;
Carlsbad Cave. Texas: Mouth of Pecos River (Bailey, N. Amer. Fauna,
25:210, 1905); Boquillas (Borell and Bryant, Univ. California Publ. Zool,
48:9, 1942); Glen Spring (Borell and Bryant, loc. cit.).
Diagnosis. — Size large for the species; total length, 80.3(78-83); tibia,
12.3(11.7-13.1) ; forearm, 32.9(31.8-33.3) ; greatest length of skull, 12.7(12.3-12.9) ;
breadth of braincase, 6.6(6.5-6.7). Color between Smoke Gray and Pale Drab
(after Hatfield, op. cit. -.261) .
Pipistrellus hesperus santarosae Hatfield
Pipistrellus hesperus santarosae Hatfield, Jour. Mamm., 17:261, August
14, 1936.
Type locality. — Santa Rosa, Guadalupe County, New Mexico.
Range. — New Mexico (excepting southern part) and western Colorado.
Marginal occurrences (after Hatfield, op. cit. :2G2) are: Colorado: Bedrock.
New Mexico: Santa Rosa; Socorro; Laguna.
Diagnosis. — Size large for the species; total length, 82.0(80-86) ; tibia,
12.4(11.9-13.0) ; forearm, 32.8(31.7-34.1) ; greatest length of skull, 12.7(12.3-13.1) ;
breadth of braincase, 6.6(6.3-6.8). Color between Buffy Brown and Wood
Brown (after Hatfield, op. cit. :261, 262).
Pipistrellus subflavus
(Synonomy under subspecies)
Range. — Canadian to Tropical life-zones of eastern North America from
Quebec southward to Honduras.
Characters. — Sayal Brown to darker than Mummy Brown, dorsally; total
length, 73-89; foot more than half as long as tibia; tragus tapering and straight;
Synopsis of Genus Pipistrellus 599
dorsal profile of skull convex in interorbital region; inner upper incisor bi-
cuspidate; outer upper incisor unicuspidate (lacking accessory cusp on antero-
internal face) ; PI viewed from occlusal face more than a seventh of area of
canine and visible from labial aspect; lower, third premolar as high as an-
terior cusp of canine; lower premolars less crowded than in P. hesperus and
distance between canine and first molar less than length of second lower molar.
Remarks. — In winter this species hibernates in caves in clusters
of fewer than fifty individuals, but in summer fewer of the bats live
there and at this season some have been captured as far as thirty
miles from any such retreat suggesting that the bats inhabit other
types of shelter. The wide range of this species in respect to life-
zones is noteworthy; it occurs in the Canadian Life-zone (Joliet,
Quebec), the Tropical Life-zone (30 km. SSE Jesus Carranza, Vera-
cruz) and in the intervening life-zones.
The longer thumb of this species, in comparison with that of
Pipistrellus hesperus, was verified by measuring the thumb includ-
ing its claw and the pad at the base of the thumb in 12 P. s. vera-
crucis and 10 P. h. maximus. In veracrucis the mean was 5.9 milli-
meters and the extremes were 5.5 and 6.4. In maximus the
corresponding figures were 3.9, 3.6 and 4.3.
Pipistrellus subfiavus subfiavus (F. Cuvier)
V[espert'ilio]. subfiavus F. Cuvier, Nouv. Ann. Mus. Hist. Nat. Paris,
1 : 17, 1832.
Vespertilio erythrodactylus Temminck, Monogr. de Mamm., II, 13me
monogr., p. 238, 1835-1841 (not seen — after Miller, N. Amer. Fauna,
13:90, October 16, 1897).
Scotophilus georgianus H. Allen, Smithsonian Miscl. Coll., No. 165,
Vol. 7 (art. 1), p. 35, June, 1864.
Vesperugo carolinensis H. Allen, U. S. Nat. Mus. Bull. 43:121, March
14, 1894.
Pipistrellus subfiavus Miller, N. Amer. Fauna, 13:90, figs. 22, 23, Oc-
tober 16, 1897.
Type locality. — Eastern United States, probably Georgia.
Range. — From approximately 40 degrees North Latitude in Pennsylvania
and Kansas southward to central Florida and at least to extreme southern
Texas; from the Atlantic Coast westward to south-central Kansas and Val
Verde County, Texas. Marginal occurrences are : Kansas (K. U. Collection) :
41/£ mi. SW Sun City; Ft. Leavenworth. Illinois (Necker and Hatfield, Bull.
Chicago Acad. Sci., 6(3) :45, 1941): Quincy; Urbana. hidiana (Lyon, Amer.
Midland Nat., 17:73, 1936): Monroe County; Franklin Co. Ohio (Bole
and Moulthrop, Sci. Pubis. Cleveland Mus. Nat. Hist., 5(6) :115, 1942: Ham-
ilton Co.; Smoky Creek. West Virginia (Kellogg, Proc. U. S. Nat. Mus.,
84:449, 1937): Charleston; Smoke Hole Cave. Pennsylvania (Rhoads,
Mamms. Pa. and N. J., p. 211, 1903): Carlisle; Germantown. New Jersey:
Haddonfield (Rhoads, Mamms. Pa. and N. J., p. 211, 1903). Florida: Tarpon
Springs (Sherman, Proc. Florida Acad. Sci., p. 107, 1936). Texas: Brownsville
600 University of Kansas Pubs., Mus. Nat. Hist.
(Bailey, N. Amer. Fauna, 25:211, 1905); Comstock (Bailey, loc. cit.); Kerr
Co. (Taylor and Davis, Game, Fish and Oyster Comm. Bull., 50:17, 1947).
Oklahoma: 10 mi. S and 2 mi. E Sulphur (Blair, Amer. Midland Nat., 22:100,
1939).
Diagnosis. — Size large; eight specimens from Barber and Butler counties,
Kansas, measure in total length, 84(77-89); tibia, 14.8(14.5-15); forearm,
33.5(31.8-35.3); greatest length of skull (exclusive of incisors), 12.8(12.3-13.1);
breadth of braincase immediately above roots of zygomatic arches, 6.5(6.4-6.7).
Color ranging from Snuff Brown to Sayal Brown.
Pipistrellus subflavus obscurus Miller
Pipistrellus subflavus obscurus Miller, N. Amer. Fauna, 13:93, Oc-
tober 16, 1897.
Type locality. — Lake George, Warren County, New York.
Range. — From southern Quebec and southern Ontario south to southern
Ohio and West Virginia; from the Atlantic Coast west into Wisconsin. Mar-
ginal occurrences are: Minnesota: St. Peter (Swanson and Evans, Jour.
Mamm., 17:39, 1936); Marine (Swanson, Tech. Bull. No. 2, Minnesota Dept.
Conservation, p. 60, 1945). Wisconsin: Hurley (Greeley and Beer, Jour.
Mamm., 30:198, 1949). Quebec: Joliet (Anderson, Nat. Mus. Canada, Biol.
ser. No. 31, Bull. 102:30, 1946). Vermont: Brandon (Osgood, Jour. Mamm.,
19:436, 1938). Maine: No locality more precise than the state (Allen, Occ.
Papers Boston Soc. Nat. Hist., 7(3) :35, June, 1904). New York: Hastings on
Hudson (Rowley, Abstr. of Proc. Linnean Soc. N. Y., for yr. ending March 11,
1902, p. 57). Pennsylvania: Beaver (Rhoads, Mamms. Pa. and N. J., 1903,
p. 211). West Virginia: Cornwall's Cave (Frum, Jour. Mamm., 25:195, 1944).
Ohio: Cat Run (Bole and Moulthrop, Sci. Pubis. Cleveland Mus. Nat. Hist.,
5(6):116, 1942); Symmes Creek (Bole and Moulthrop, loc. cit.); Dry Cave
(Bole and Moulthrop, loc. cit.) ; "Union County" (Rausch, Jour. Mamm.,
27:275, 1946). Wisconsin: Devils Lake (Jackson, Jour. Mamm., 1:38, 1919).
Diagnosis. — ". . . color duller and less yellow, and dark tips of shorter
hairs on back more conspicuous" than in P. subflavus subflavus according to
the original description.
Remarks. — No one, as far as we know, has carefully studied the
variation in Pipistrellus subflavus of the United States and Canada
since Miller named P. s. obscurus. With the more abundant ma-
terial now available, such an appraisal would be worth-while. The
occurrences cited above for Minnesota and Wisconsin were recorded
in the literature under the specific name without indication of sub-
specific affinity. The reference of specimens from these states to
the subspecies P. s. obscurus is an arbitrary assignment on our part;
we have not seen them. However, two specimens in the University
of Kansas Museum of Natural History from Potosi (Snake Cave)
Grant County, Wisconsin, are referable to P. s. obscurus. These
provide the southwesternmost record station of occurrence in Wis-
Synopsis of Genus Pipistrellus 601
consin but are not shown on the distribution map because the speci-
mens were received after figure 1 was prepared.
It is noteworthy that the species Pipistrellus subflavus has not
yet, as far as we can ascertain, been recorded from Michigan, north-
ern Indiana, northern Illinois, or Iowa. Probably the species occurs
in these areas.
Pipistrellus subflavus veracrucis (Ward)
Vesperugo veracrucis Ward, Amer. Nat., 25:745, August, 1891.
Pipistrellus veracrucis Miller, N. Amer. Fauna, 13:93, October 16,
1897.
Type locality. — Las Vigas, 8,500 ft., Veracruz.
Range. — Eastern Mexico, certainly from the type locality southward into
Honduras. Records of occurrence are: Veracruz: Las Vigas (13 specimens
from 4 km. E Las Vigas, 8,500 ft., K. U.) ; 30 km. SSE Jesus Carranza, 1
(K. U.). Honduras: Jilamo Farm, Tela District, 3 (Univ. Michigan).
Diagnosis. — Size small for the species; measurements of 13 near topotypes
are: total length, 78(73-85); tibia, 12.9(11.8-14.7); forearm, 31.8(29.5-33.1);
greatest length of skull (exclusive of incisors), 12.2(11.8-12.6) ; breadth of brain-
case immediately above roots of zygomatic arches, 6.3(6.0-6.7). Color darker
than Mummy Brown above and below.
Remarks. — The specimen from thirty kilometers south-southeast
of Jesus Carranza, Veracruz, and the three specimens from Honduras
agree in all respects with topotypes. The color of P. s. veracrucis
is much darker than that of P. s. obscurus and is between black and
the darkest brown in Ridgway's (op. cit.) color key. Rinker (Jour.
Mamm., 29:179-180, 1948) described the three specimens from Hon-
duras without assigning a specific name to them because he lacked
topotypes of P. s. veracrucis. We find nothing in his description to
correct, but can add that the upper tooth-rows in many, but not in
all, specimens of P. s. veracrucis are straighter than in P. s. sub-
flavus. Probably it was this feature to which Rinker referred when
he said that in veracrucis "The tooth rows tend to be more con-
vergent posteriorly." Rinker did not refer the three specimens from
Honduras to P. veracrucis because Ward's original description states
that veracrucis has evenly spaced lower incisors and a basal cusp
on the lower canine on only its forward edge. Rinker's specimens
from Honduras have the first incisors in contact with each other,
the second incisors in contact with the first incisors and the third
incisor on each side of the lower jaw separated by a space from the
second incisor and from the canine. The specimens from Honduras
have a basal cusp on the hinder edge of the lower canine. In these
two features they agree with the specimens from Veracruz and with
602 Univeksity of Kansas Pubs., Mus. Nat. Hist.
specimens of Pipistrellus subflavus from the United States and Can-
ada. It is clear that Ward (Amer. Nat., 25:747, 1891) was mistaken
in stating that the lower incisors of veracrucis were evenly spaced
and that the canine had a basal cusp on only the forward edge.
Ward (loc. cit.) was correct in regarding his Vesperugo veracrucis
as "most closely related to V. georgianus [r= Pipistrellus subflavus],"
but for want of actual specimens of P. subflavus to use in compari-
son was incorrect in supposing that P. subflavus had only two bands
of color on the fur, more hair on the legs, and a larger area of hair
on the interfemoral membrane. In these respects we perceive no
difference between specimens from Veracruz and the United States.
Vesperugo veracrucis Ward, therefore, proves to be only a sub-
species of Pipistrellus subflavus, but is well characterized by dark
color and small size.
University of Kansas Museum of Natural History, Lawrence,
Kansas. Transmitted October 31, 1949.
□
23-1546
Nfl-L
MUS. COWP. ZOOL
LIBRARY
JUN
INDEX TO VOLUME 1
' New systematic names are in boldface type
abspnM'THomomi-s bottae, 71
acdflltltf^«fyoliusJ204
-i^ nil nil, in hflaaainea r'"ti""'q, 211
acanthopus, Hoplopleura, 141
Accipiter, .
cooperi, 198
gentilis, 198
striatulus, 198
striatus, 198
velox, 198
Acris crepitans, 104
Actitis macularia, 201
acuta, Anas, 197
acutipennis, Chordeiles, 186
acutus, Liomys irroratus, 253, 455
adocetus, Citellus adocetus, 450
Aechmophorus occidentalis, 196
aedon, Troglodytes, 207
Aegolius,
acadicus, 204
funereus, 203
richardsoni, 203
aegypticus, Trionyx, 122
Aeronautes saxatalis saxatalis, 204
affinis, Aythya, 198
Agelaius phoeniceus, 168
agilis,
Dipodomys, 477
Dipodops, 477
Agkistrodon mokeson, 111
agrestis, Microtus, 142
Agropyron repens, 133
Aix sponsa, 197
alascensis, Calcarius lapponicus, 191,
212
Alaska,
hermit thrush, 208
longspur, 191, 212
three-toed woodpecker, 205
alba,
Melilotus, 133
Tyto, 263
albeola, Glaucionetta, 198
albescens, Myotis, 239
albicaudatus, Thomomys bottae, 37
albicollis, Zonotrichia, 212
albifrons,
Anser, 197
Petrochelidon pyrronota, 206
albipes, Sciurus, 450
albispinosus, Echimys, 390
albispinus,
Echimys, 388
Proechimys, 390
albociliatus, Phalacrocorax auritus,
196
albolarvatus, Dendrocopos, 205
Alcantar, Roberto, 435
Alcorn, J. R., 435
alcyon, Megaceryle, 204
alder flycatcher, 187
Alexander, Annie M., 435
alexandrae, Thomomys bottae, 72
alexandri, Archilochus, 204
alexandrinus,
Charadrius, 183
Rattus rattus, 468
alfalfa, 133
algodonera, rata,
leonada, 465
setosa, 465
alleni, Liomys irroratus, 253, 455
almae, Hylocichla ustulata, 188, 20S
alpestris, Eremophila, 205
alpine three-toed woodpecker, 205
alticola,
Melospiza lincolnii, 212
Vireo solitarius, 188
altipetens, Lagopus leucurus, 200
amatze, 446
Ambrosia,
artemisiifolia, 136
trifida, 134
Ambystoma,
mavortium, 103
texanum, 103
tigrinum, 103
American,
bittern, 196
coot, 200
golden-eye, 198
hawk owl, 203
magpie, 206
merganser, 198
pintail, 197
raven, 206
redstart, 210
rough-legged hawk, 199
americana,
Aythya, 197
Fuhca, 168, 200
Glaucionetta, 198
Mareca, 197
Recurvirostra, 201
— Univ. Kansas Publ., Mus. Nat. Hist., Vol. 1, 1946-1050.
606
University of Kansas Publs., Mus. Nat. Hist.
americanus,
Bufo, 103
Coccyzus, 202
Mergus merganser, 198
Numenius, 183, 201
Ammodramus,
pratensis, 191
savannarum, 168, 191
amoena,
Carphophis, 107
Passerina, 210
amoenissima, Polioptila caerulea, 208
amphichoricus, Proechimys, 344
Araphispiza,
belli, 212
nevadensis, 212
Amyda,
cartilageneus, 122
ferox, 120
mutica, 112, 119
sinensis, 119
spinifera, 112, 119
steindacheneri, 119
analogus,
Baiomys taylori, 460
Peromyscus taylori, 460
Anas,
acuta, 197
carolinensis, 197
cyanoptera, 197
discors, 168, 197
platyryhnchos, 197
strepera, 197
tzitzihoa, 197
anatum, Falco peregrinus, 199
Andropogon furcatus, 132
angustifrons, Spilogale angustifrons,
447
angustirostris,
Cratogeomys, 452
Platygeomys, 452
annectens, Cyanocitta stelleri, 206
annua, Poa, 133
Anser albifrons, 197
anthracinus, Eumeces, 87
Anthus,
pacificus, 188, 208
spinoletta, 188, 208
angulatus, Tayassu, 469
aparine, Galium, 133
apasr, 447
apatze, 446
apatzee, 447
Aphelocoma,
coerulescens, 206
woodhousei, 206
Apodemus sylvaticus, 142
apus, Pipistrellus hesperus, 597
aquaticus, Scalopus, 137
Aquila,
canadensis, 199
chrysaetos, 199
arabupu, Proechimys, 369
arborea, Spizella, 212
arboricola, ardilla, 450, 451
Archilochus alexandri, 204
arctic,
horned owl, 203
three-toed woodpecker, 205
towhee, 211
arcticus,
Picoides, 205
Pipilo maculatus, 211
Ardea,
herodias, 196
treganzai, 196
ardilla,
arboricola, 450, 451
de pedregal, 449
arescens, Proechimys, 366
argentatus,
Larus, 202
Myotis, 239
Aristada oligantha, 132
arizonae, Spizella passerina, 212
armadillo, 470
arnyi, Diadophis punctatus, 107
arrocera,
pigmea, rata, 465
tropical, rata, 464
artemisiae, Molothrus ater, 210
artemisiifolia, Ambrosia, 136
Articholaelaps,
glasgowi, 142
sigmodoni, 142
Artibeus,
hirsutus, 442
planirostris planirostris, 442
artichoke, Jerusalem, 135
arvensis, Sonchus, 136
Arvey, M. Dale, a check-list of the
birds of Idaho, 195
Ascoschongastia brevipes, 143
aserriensis, Chordeiles minor, 186
ash-throated flycatcher, 186, 205
Asio,
flammeus, 203
otus, 203
wilsonianus, 203
asio, Otus, 203
astutus, Bassariscus, 447
Asyndesmus lewis, 204
Atalapha mexicanus, 444
ater, Molothrus, 210
atrata, Leucosticte, 211
atratus, Sigmodon hispidus, 466
atricapillus, Parus, 206
attenuatus, Dipodomys ordii, 553
Audubon,
hermit thrush, 208
warbler, 209
Index to Volume 1
607
auduboni,
Dendroica, 209
Hylocichla guttata, 208
aura, Cathartes, 198
aureiventirs, Thomomys bottae, 28
aureus, Thomomys bottae, 61
auritus,
Colymbus, 196
Phalacrocorax, 198
aurocapillus, Seiurus, 189
australis, Pipistrellus hesperus, 597
avocet, 201
Aythya,
aftinis, 198
americana, 197
collaris, 197
marila, 198
valisineria, 197
baileyi, Myotis, 587
Baiomys,
analogus, 460
musculus, 460
taylori, 460
Baird sandpiper, 184
bairdii, Erolia, 184
Balantiopteryx plicata, 440
bald eagle, 199
baldpate. 197
baileyi, Castor canadensis, 410
banderanus, Peromyscus banderanus,
463
band-tailed pigeon, 202
bank swallow, 206
barn,
owl, 203
swallow, 206
Barrow golden-eye, 198
bassanus, Morus, 180
Bassariscus,
astutus, 447
consitus, 447
bat,
big brown, 444
big leaf-nosed, 442
brown, big, 444
cave, 443
free-tailed, Mexican, 445
hoary, 445
long-eared, 445
long-tongued, 440, 441
leaf-nosed, 441, 442
mastiff, 445
new from Mexico, 239
red, 444
sac-winged, 440
vampire, 443
Yuma, 443
Batchelder woodpecker, 205
Beals, Ralph L., 435
belli, Amphispiza, 212
bellii, Chrysemys picta, 112
Bendire crossbill, 211
bendirei,
Falco columbarius, 199
Loxia curvirostra, 211
Bent crossbill, 211
benti, Loxia curvirostra, 211
bernicla, Branta, 197
bicolor,
Iridoprocne, 206
Perisoreus canadensis, 206
big,
brown bat, 444
leaf-nosed bat, 442
birdseyi, Thomomys bottae, 63
bistincta, Hyla, 261
bittern, American, 196
black,
rat, 468
snake, 108
tern, 202
black-billed cuckoo, 202
black-bellied plover, 200
black-capped chickadee, 206
black-chinned hummingbird, 204
black-eared,
deermouse, 464
nuthatch, 207
black-headed,
jay, 206
snake, 110
black-necked stilt, 201
black-throated gray warbler, 209
blackbird,
Brewer, 210
yellow-headed, 210
blackish mouse, 463
Blarina,
brevicauda, 137
pergracilis, 440
blue,
grouse, 199
heron, 196
racer, 107
blue-winged teal, 168
bluebird,
mountain, 208
western, 208
bluegrass, 132
bluejoint, 132
bob-white, Texas, 200
bobcat, 449
bobolink, 210
Bohemian waxwing, 208
boimensis, Proechimys, 350
bolivianus, Proechimys, 316
bonafidei, Proechimys, 378
Bonaparte gull, 202
608
University of Kansas Publs., Mus. Nat. Hist.
bombifrons, Scaphiopus, 103
Bombycilla,
cedrorum, 208
garrulus, 208
pallidiceps, 208
Bonasa,
incanus, 200
phaia, 199
umbelloides, 199
umbellus, 199
bonnevillei, Thomomys bottae, 41
booby, red-footed, 180
Borbolla, Prof. Daniel Rubin F. de la,
435
borealis,
Lasiurus, 444
Nuttallomis, 205
Botaurus lentiginosus, 196
bottae, Thomomys, 28
box turtle. 111
boylei, Peromyscus, 461
boylii, Peromyscus, 461
brachycephala, Rana pipiens, 104
brachyrynchos, Corvus, 206
Branta,
bernicla, 197
canadensis, 196
canadensis hutchinsii, 182
hutchinsii, 182, 197
leucopareia, 197
moffitti, 196
nigricans, 197
occidentalis, 196
Brazilian spiny rats, 303
brenero, raton, 461
Breukeleman, John and Hobart M.
Smith, selected records of reptiles
and amphibians from Kansas, 103
brevicauda,
Blarina, 137
Proechimys, 349
breviceps, Geomys, 219
brevipes, Ascoschonagastia, 143
Brewer,
blackbird. 210
sparrow, 212
breweri, Spizella, 212
Brewster,
egret, 196
screech owl, 203
brewsteri,
Leucophoyx thula, 196
Otus asio, 203
broad-tailed hummingbird, 204
broad-winged hawk, 198
brome, smooth, 133
Bromus,
carina (us, 133
inermis, 133
bronzed grackle, 168
brooksi, Hesperiphona vespertina, 210
brown,
bat, big, 444
thrasher, 168
brunnescens, Lophortyx californica,
200
brush mouse, 461
Bubo,
lagophonus, 203
occidentalis, 203
pacificus, 166
virginianus, 159, 203
wapacuthu, 203
buccinator, Cygnus, 196
Buenrostro, Sr. Efrain, 435
buffle-head, 198
Bufo,
americanus, 103
cognatus, 95, 104
compactilis, 96
woodhousii, 104
bulbivorus, Thomomys talpoides, 3
bull snake, 108
bullfrog, 104
Bullock oriole, 210
bullocki, Icterus, 210
bunting,
eastern snow. 213
lark. 211
lazuli. 210
snow, 213
burrowing owl, 203
burrus, Proechimys, 316
bursarius, Geomys, 222
bush-tit, lead-colored, 207
Buteo,
calurus, 198
.iamaicensis, 198
lagopus, 199
lineatus texanus, 183
platypterus, 198
regalis, 199
s[anctil.-johannis, 199
swainsoni, 199
texanus, 183
cacomixtle, 447
caerulea, Polioptila, 208
Cafer,
cafer, 204
canescens, 204
collaris, 204
cafer, Colaptes, 204
cagottis,
Canis latrans, 449
Lyciscus, 449
cajennensis, Echinomys, 387
Calamospiza melanocorys, 211
Calaveras warbler, 209
Calcarius,
alascensis, 191, 212
lapponicus, 191, 212
Index to Volume 1
609
calendula, Regulus, 20S
calidior, Proechimys, 316
California,
cuckoo, 202
gull, 202
murcielago, 444
myotis, 444
pygmy owl, 203
shrike, 209
californica, Lophortyx, 200
califomicum, Glaucidium gnoma, 203
californicus,
Colymbus nipa-icolIis, 196
Geococcyx, 184
Larus, 202
Microtus, 142
Myotis, 444
calligaster, Lampropeltis, 108
calliope,
hummingbird, 204
Stellula, 204
callotis, Lepus, 469
calurus, Buteo jamaicensis, 198
Canachites franklinii, 199
Canada goose, 196
canadensis,
Aquila chrysaetos, 199
Branta, 182, 196
Castor, 409
Grus, 200
Perisoreus, 206
Sitta, 207
canescens,
Cafer cafer, 204
Didelphis (Micomrus), 439
Marmosa, 439
caniceps, Junco, 212
canicollis, Proechimys, 315
Canis,
cagottis, 449
latrans, 449
cantabrigensis, Rana sylvatica, 104
canvas-back, 197
canyon wren, northern, 207
caparoch, Surnia ulula, 203
Capella,
delicata, 201
gallinago, 201
Cardenas, General Lazaro, 435
carilargo, murcielargo, 441
carinatus, Bromus, 133
carissima, Myotis lucifugus, 585
Carolina, Porzana, 168, 200
carolinense, Solanum. 135
carolinensis,
Anas, 197
Dumetella, 207
Pandion haliaetus, 199
Sitta, 207
Vesper ugo, 599
Carpodacus,
cassinii, 210
mexicanus, 210
solitudinis, 210
Carphophis,
amoena, 107
vermis, 107
cartilagineus, Amyda, 122
casero, raton, 468
caspia, Hydroprogne, 202
Caspian tern, 202
Cassiar junco, 191
Cassidix,
mexicanus, 190
prosopidicola, 190
Cassin,
purple finch, 210
vino, 209
cassinii,
Carpodacus, 210
Vireo solitarius, 209
Castor,
baileyi, 410
canadensis, 409
concisor, 410
duchesnei, 413
frondator, 409
pallidus, 409
rostralis, 411
taylori, 410
repentinus, 409
cat, ring-tailed, 447
catbird, western, 207
catenatus, Sistrurus, 111
catenifer, Pituophis, 108
catesbeiana, Rana, 104
Cathartes aura teter, 19S
Catherpes,
griseus, 207
mexicanus, 207
Catoptrophorus,
inornatus, 201
semipalmatus, 201
caurina,
Certhia familiaris, 2-7
Megaceryle alcyon, 204
caurinus, Turdus migratorius, 208
cave bat, 443
cayennensis,
Echimys, 351
Proechimys, 314
cedar waxwing, 208
cedrorum, Bombycilla, 208
celata, Vermivora, 209
celeripes, Dipodomys ordii, 549
centralis,
Proechimys, 316
Thomomys bottae, 44
Centrocercus urophasianus, 200
Cercomys, 321
610
University of Kansas Publs., Mus. Nat. Hist.
Certhia,
caurina, 207
faniiliaris, 207
Chaetura, vauxi, 204
chaparral, raton de, 461
chapmani,
Cricetodipus, 536
Dipodomys ordii, 536
Perodipus, 536
Charadrius,
alexandrinus, 183
hiaticula, 183
nivosus, 183
semipalmatus, 183
tenuirostris, 183
vociferus, 200
vvilsonia, 184
chaf ,
long-tailed, 209
vellow-breasted, 189
Chelonia, 284
Chelydra, 284
serpentina, 111
Chen,
hyperborea, 197
rossi, 197
Cherrie nighthawk, 186
cherriei, Proechimys, 316
chestnut-backed chickadee, 207
chickadee,
black-capped, 206
chestnut-backed, 207
Columbian black-capped, 206
chipping sparrow, 212
chiriquinus, Proechimys, 316
Chlidonias,
nigra, 202
surinamensis, 202
Chlorura chlorura, 211
chlorura, Chlorura, 211
Choeronycteris mexicanus, 441
Chondestes,
grammacus, 191, 211
strigatus, 191, 211
Chordeiles,
acutipennis, 186
aserriensis, 186
hesperis, 204
howelli, 185
minor. 185, 186, 204
sennetti, 186
texensis, 186
chrysaeolus, Proechimys, 316
chrysaetos, Aquila, 199
Chrysemys,
bellii, 112
limnodytes, 269
picta, 112, 270
timida, 272
chrysopsis, Reithrodontomys, 459
cinarascens, Myiarchus, 186, 205
Cinclus,
mexicanus, 207
unicolor, 207
cinderensis, Dipodomys ordii, 540
cineraceus,
Dipodomys ordii, 550
Regulus calendula, 208
cinereoargenteus, Urocyon, 448
cinereus, Lasiurus cinereus, 445
cinnamomea, Tringa solitaria, 201
cinnamomeus,
Mus, 392
Myotis, 585
Pipistrellus, 593
Pipistrellus, referred to the genus
Myotis, 583
Vespertilio, 585
Circus,
cyaneus, 199
hudsonius, 199
cismontanus, Junco hyemalis, 191,212
Citellus,
adocetus, 450
tridecimlineatus, 143
varicgatus, 449
clangula, Glaucionetta, 198
Clark nutcracker, 206
cleavers, 133
Clethrionomys glareolus, 142
cliff swallow, 164, 206
clover, 132
clusius, Thomomys, 17
clypeata, Spatula, 197
Cnemidophorus sexlineatus, 106
coachwhip, 108
coati, 446
Coccyzus,
americanus, 202
erythropthalmus, 202
occidentalis, 202
coerulescens, Aphelocoma, 206
cognatus, Bufo, 95, 104
cola blanca, venado, 470
colchicus, Phasianus, 200
colimensis, Urocyon cinereoargenteus,
448
Colinus,
texanus, 200
virginianus, 200
colirugosa, murcielago, 444
collared,
lizard, 105
peccary, 469
collaris,
Aythya, 197
Cafer cafer, 204
Crotaphytus, 105
colombianus, Proechimys, 316
Coluber,
constrictor, 107
flaviventris, 107
Index to Volume 1
611
coludo, murcielago, 445
Columba,
fasciata, 202
livia, 168
columbarius, Falco, 199
Columbian,
black-capped chickadee, 206
sharp-tailed grouse, 200
columbiana, Nucifraga, 206
columbianus,
Cygnus columbianus, 196
Cricetodipus ordii, 544
Dipodomys ordii, 544
Pedioecetes phasianellus, 200
perodipus ordii, 544
Columbigallina passerina pallescens,
185
Colymbus,
auritus, 196
californicus, 196
grisegena, 196
holbollii, 196
nigricollis, 196
comadreja, 447
common,
redpoll, 211
rock wren, 207
tern, 202
comosa, Falcata, 135
compactilis, Bufo, 96
compactus,
Dipodomys ordii, 515
Cricetodops, 515
Perodipus, 515
concisor, Castor canadensis, 410
conejo,
de Florida, 468
Mexicano, 469
Conepatus,
mesoleucus, 448
nelsoni, 448
tropicalis, 577
confmis, Pooecetes gramineus, 211
consitus, Bassariscus astutus, 447
constrictor, Coluber, 107
Contopus richardsonii richardsonii,
205
contortrix, Heterodon, 107
contractus, Thomomys bottae, 50
convexus, Thomomys bottae, 47
Convolvulus sepium, 135
Cooper hawk, 198
cooperi, Synaptomys, 130
cooperii, Accipiter, 198
coot, 168
American, 200
copperhead, 111
corax, Corvus, 206
cormorant, Farallon, 196
cornutum, Phrynosoma, 106
coromandelicus, Trionyx, 122
Corrodopsylla hamiltoni, 140
Corvus,
brachyrynchos, 206
corax," 206
hesperis, 206
sinuatus, 206
Corynorhinus,
macrotis, 445
megalotis, 445
mexicanus, 445
pallescens, 445
rafinesquii, 445
cotton rat,
fulvous, 465
hispid, 465
cottontail,
Florida, 468
Mexican, 469
couesi, Oryzomys, 464
cowbird, Nevada, 210
coyote, 449
crabgrass, 132
Crane, Harold S., Stephen D. Durrant
and, Three new beavers from Utah,
409
crane, sandhill, 200
Cratogeomys, 219
angustirostris, 452
gymnurus, 252, 452
imparilis, 452
varius, 453
creeper, northwestern, 207
crepitans, Acris, 104
Cricetodipus,
chapmanij 536
columbianus, 544
compactus, 515
longipes. 556
ordii, 530
palmeri, 562
sennetti, 517
Crocethia, 201
alba, 201
crossbill,
Bendire, 211
Bent, 211
white-winged, 211
red, 211
Crotalus,
horridus, 111
viridus, 111
Crotaphytus collaris collaris, 105
crow, western, 206
crucifer, Hyla, 88
crusgalli, Eichinochloa, 134
Cryptotis,
parva, 137
pergracilis pergracilis. 440
Ctenophthalmus pseudagyrtes, 140
cuatralvo, raton, 461
Cuban snowy plover, 183
cuchjeramba, 469
612
University of Kansas Publs., Mus. Nat. Hist.
cuckoo,
black-billed, 202
California, 202
cuitziqui, 448
cumihuatz, 448
cumu, 451. 452, 453
cuiniqui, 450
cunicularia, Speotyto, 203
cupidineus, Dipodomys ordii, 561
Cupidinimus, 480
nebraskensis, 483
curlew,
long-billed, 183, 201
northern, 201
northern long-billed, 183
currucoides, Sialia, 208
curtatus, Pipilo maculatus. 211
curvirostra, Loxia, 211
eyaneus, Circus, 199
cyanocephalus,
Euphagus, 210
Gymnorhinus, 206
Cyanocitta,
annectens, 206
stelleri, 206
cyanoptera, Anas, 197
Cygnus,
buccinator, 196
columbianus, 196
Dalquest, Walter W., 259
Dalquest, Walter W., and E. Ray-
mon Hall, A new bat (genus Myo-
tis) from Mexico, 237
A synopsis of the American bats of
the genus Pipistrellus, 591
Geographic range of the hooded
skunk, Mephitis macroura, with
description of a new subspecies
from Mexico, 575
Pipistrellus cinnamomeus Miller
1902 referred to the genus Myo-
tis, 581
Tadarida femorosacca (Merriam)
in Tamaulipas, Mexico, 245
dandelion, 134
dasycnemus, Doratopsylla, 140
Dasypus,
novemcinctus, 470
mexicanus, 470
decumans, Proechimys, 316
deer, white-tailed, 470
deermouse, 461
delawarensis, Larus, 202
delicata, Capella gallinago, 201
Dendrocopos,
albolarvatus, 205
leucurus, 205
monticola, 204
pubcscens, 205
villosus, 204
Dendragapus,
obscurus, 199
pallidus, 199
richardsonii, 199
Dendroica,
auduboni, 209
petechia, 209
morcomi, 209
nigrescens, 209
townsendi, 209
denigratus, Proechimys, 381
densiflorum, Lepidium, 133
Dermacentor variabilis, 143
desert house finch, 210
deserti, Dipodomys deserti, 65
Desmodus,
murinus, 443
rotundus, 443
Diadophis,
arnyi, 107
punctatus, 107
Dichromanassa rufescens, 181
Didelphis,
canescens, 439
mesamericana, 439
pigra, 439
texensis, 439
virginiana, 439
Digitaria ischaemum, 132
dimidiatus,
Echimys, 371
Proechimys, 371
dioica, Gymnocladus, 134
Dipodomyinae, 480
Dipodomys,
agilis, 477
attenuatus, 553
celeripes, 549
chapmani, 536
cinaraceus, 550
cinderensis, 540
cleomophila, 556
columbianus, 544
compactus, 515
cupidineus, 561
deserti, 65, 496
elator, 496
elephantinus, 496
evexus, 518
extractus, 534
fetosus, 541
fremonti, 524
fuscus, 555
gidleyi, 482
heermanni, 496
idoneus, 546
inaquosus, 552
ingens, 496
insularis, 496
kansensis, 481
levipes, 489
Index to Volume 1
613
Dipodomys — Concluded
longipes, 556
luteolus, 533
margaritae, 496
marshalli, 551
medius, 519
merriami, 496
microps, 477
minor, 482
mitchelli, 496
mohavensis, 489
monoensis, 528
morroensis, 489, 496
nelsoni, 496
nexilis, 559
nitratoides, 496
obscurus, 521
oklahomae, 514
ordii, 530
ornatus, 496
pallidus, 558
palmeri, 562
panamintinus, 496
panguitchensis, 496
perotensis, 496
perplexus, 489
phillipsi, 496
platycephalus, 496
priscus, 547
richardsoni, 511
sanrafaeli, 526
santiluciae, 4S9
sennetti, 517
spectabilis, 496
stephensi, 496
terrosus, 523
utahensis, 543
venustus, 496
Dipodops,
agilis, 477
ordii, 530
dipper, 207
Diprionomys, 482
discors, Anas, 168, 197
dispersal of kangaroo rats, 498
dissimilis, Thomomys bottae, 60
Dolichonyx oryzivorus, 210
domesticus, Passer, 210
dominica,
Oxyura, 182
Pluvialis, 184. 200
dorado, raton, 459
Doratopsylla dasycnemus, 140
dorsalis, Picoides tridactylus, 205
dove,
ground, 185
Mexican ground, 185
mourning, 16S, 202
rock, 168
western mourning, 202
dowitcher, long-billed, 201
duchesnei, Castor canadensis, 413
duck,
lesser scaup, 198
greater scaup, 198
harlequin, 198
hawk, 199
masked, 182
ring-necked, 197
ruddy, 198
scaup, 198
western harlequin, 19S
wood, 197
Dumetella,
carolinensis, 207
ruficrissa, 207
Durrant, Stephen D.,
The pocket gophers (genus Thom-
omys) of Utah, 3
Three new beavers from Utah, 409
dusky horned lark, 205
dutcheri, Geomys breviceps, 224
dwarf wood rat, 467
eagle,
bald, 199
golden, 164. 199
northern bald, 199
eared grebe, 196
eastern,
kingbird, 205
nighthawk, 185
snow bunting, 213
sparrow hawk, 199
Echimyidae, 303
Echimys,
albispinosus, 390
albispinus, 369
brevicauda, 349
cayennensis, 351
dimidiatus, 371
elegans, 387
macrourus, 315
myosuros, 351
setosus, 385
trinitatis, 333
Echinochloa crusgalli, 134
Echinomys,
cajennensis, 387
fuliginosus, 385, 390
leptosoma, 392
myosuros, 392
Ectopistes migratoiius, 202
egret,
Brewster, 196
reddish, 181
Elaphe,
laeta, 108
obsoleta, 108
elasson, Gavia immer, 195
elegans,
Echimys, 387
Loncheres, 351
614 University of Kansas Publs., Mus. Nat. Hist.
elegans — Concluded
Proechiinys, 387
Pseudemys scripta, 112
Emory rat snake, 108
emoryi, Amyda, 119
Empidonax,
brewsteri, 205
flaviventris, 186
griseus, 205
harnmondi, 205
minimus, 187
traillii, 187
wrighti, 205
emydid turtle, extinct from lower
Pliocene of Oklahoma, 269
English spairow, 210
enucleator, Pinicola, 210
episcopa, Sonora, 109
Epitedia wenmanni, 140
Eptesicus,
fuscus, 444
miradorensis, 444
Ereunetes mauri, 201
Erolia,
bairdii, 184
melanotos, 201
minutilla, 201
eiythrodactylus, Vespertilio, 599
erythrogaster,
Hirundo rustica, 206
Natrix, 109
erythropthalmus, Coccyzus, 202
erythrorhynchos, Pelecanus, 196
escuinapae, Lynx rufus, 449
espinoso, raton, 454
Eulaelaps stabularis, 142
Eumeces,
anthracinus, 87
fasciatus, 106
obsoletus, 106
pluvialis, 87
septentrionalis, 106
Eumops,
underwoodi, 445
sonoriensis, 446
Euphagus cyanocephalus, 210
European partridge, 200
Eurycea,
longicauda, 88
lucifuga, 88
melanopleura, 88
evening grosbeak, 210
evexus, Dipodomys ordii, 518
evides. Peromyscus boylii, 461
excubitor, Lanius, 209
eximius, Mephitis macroura, 579
extractus, Dipodomys ordii, 534
Falcata comosa, 135
Falco,
anatum, 199
bendirei, 199
columbarius, 199
mexicanus, 199
peregrinus, 199
sparverius, 199
falcon, prairie, 199
fallax, Melospiza melodia, 212
familiaris, Certhia, 207
Farallon cormorant, 196
fasciata, Columba, 202
fasciatus,
Eumeces, 106
Nosopsyllus, 140
Picoi'des tridactylus, 205
fedoa, Limosa, 184, 201
femorosacca, Tadarida, range of, 247
ferox, Amyda, 120
ferruginea, Neotoma, 467
ferruginosa, rata, 467
ferruginous wood rat, 467
fetosus, Dipodomys ordii, 541
finch,
black rosy, 211
Cassin purple, 210
desert house, 210
gray-crowned rosy, 211
Hepburn rosy, 211
house, 210
purple, 210
rosy, 211
fisheri, Thomomys quadratus, 6
five-lined skink, 106
flagellum, Masticophis, 108
flammea, Acanthis, 211
flammeolus, Otus, 203
flammeus, Asio, 203
Flammulated screech owl, 203
flavescens, Kinosternum, 111
flavipes, Totanus, 201
flaviventris,
Coluber constrictor, 107
Empidonax, 186
flicker, red-shafted, 204
Florentiamys, 289
Florida,
conejo, 468
cottontail, 468
floridana, Pseudemys, 112
floridanus, Sylvilagus, 168, 468
frycatcher,
alder, 187
ash-throated, 186, 205
gray, 205
Hammond, 205
least, 187
little, 205
olive-sided, 205
scissor-tailed, 186
vermilion, 187
Wright, 205
yellow-bellied, 186
forficata, Muscivora, 186
formicorum, Mookomys, 481
Index to Volume 1
615
Forster tern, 202
forsteri, Sterna, 202
fortidens, Myotis, 585, 586
fortis, Agelaius phoeniceus, 210
fortuitus, Parus atricapillus, 206
fossor, Thomorays talpoides, 21
fox,
squirrel, 168
sparrow, slate-colored, 212
foxtail, 132
Franklin,
grouse, 199
gull, 202
franklinii, Canachites, 199
fraterna, Rectofrontia, 140
free-tailed bat, Mexican, 445
fremonti, Dipodomys ordii, 524
frenata, Mustek, 447
frenatus, Putorius, 447
frentuda, rata montera, 467
fringe-tailed myotis, 444
frog,
bull, 104
cricket, 104
leopard, 104
northern cricket, 104
wood, 104
frondator, Castor canadensis, 409
Fulica americana, 168, 200
fulicarius, Phalaropus, 202
fuliginosus,
Echinomys, 385, 390
Proechimys, 308
fulva, Pluvialis dominica, 200
fulvescens,
Oryzomys, 465
Reithrodontomys, 457
fulviventer, Microtus mexicanus, 425
fulvous,
cotton rat, 465
rice rat, 465
fulvus, Thomomys, 61
fundatus, Microtus mexicanus, 425,
467
funereus, Aegolius, 203
furcatus, Andropogon, 132
furvior, Seiurus aurocapillus, 189
fusca, Melanitta, 198
fuscescens, Hylocichla, 188, 208
fusco, murcielago, 444
fuscus, Dipodomys ordii, 555
fuscus, Eptesicus, 444
Eptesicus, 444
gadwall, 197
Galbreath, Edwin C,
A new extinct emydid turtle from
the lower Pliocene of Oklahoma,
269
A new species of heteromyid ro-
dent from the middle Oligocene
of northeast Colorado with re-
marks on the skull, 289
Pliocene and Pleistocene records of
fossil turtles from Western Kan-
sas and Oklahoma, 283
Galium aparinae, 133
gallinago, Capella, 201
Gambel sparrow, 212
gambeli,
Lanius ludovicianus, 209
Parus, 206
Zonotrichia leucophrys, 212
garmani, Sceloporus undulatus, 106
garrulus, Bombycilla, 208
garter snake, 110
garter snakes, hybridization, 99
gato del monte, 449
Gavia,
elasson, 195
immer, 195
stellata, 195
gentilis,
Accipiter, 198
Lampropeltis triangulum, 109
Geococcyx californicus, 184
Geomys,
breviceps, 219
bursarius, 222
dutcheri, 224
hylaeus, 234
illinoensis, 224
industrius, 226
jugossicularis, 226
llanensis, 234
levisagittalis, 234
lutescens, 222
major, 229
majusculus, 223
vinaceus, 234
georgianus, Scotophilus, 599
getulus, Lampropeltis, 109
giant ragweed, 134
glareolus, Clethrionomys, 142
glasgowi, Articholaelaps, 142
glass-snake lizard, 106
Glaucidium,
californicum, 203
gnoma, 203
Glaucionetta,
albeola, 198
americana, 198
clangula, 198
islandica, 198
Glossophaga,
leachii, 440
soricina, 440
glossy ibis, 181, 196
gnatcatcher, western, 208
gnoma, Glaucidium, 203
616
University of Kansas Publs., Mtjs. Nat. Hist.
godwit,
Hudsonian, 201
marbled, 184, 201
goeldi, Proechimys, 33S
golden,
eagle, 164, 199
plover, Pacific, 200
golden-crowned kinglet, 208
golden-eye,
American, 198
Barrow, 198
goldfinch, pale, 211
goldmani,
Nelsonia, 467
Proechimys, 316
goose,
cackling, 197
Canada, 196
Great Basin Canada, 196
Hutchins cackling, 182, 197
lesser Canada, 197
lesser snow, 197
Ross, 197
snow, lesser, 197
white-cheeked, 196
white-fronted, 197
gopher, pocket, 451, 452, 453
Gopherus, 184
gorgonae, Proechimys, 316
goshawk, western, 198
gracilis,
Tantilla, 110
Thomomys talpoides, 6
grackle,
bronzed, 168
Mesquite great-tailed, 190
grahami, Natrix, 109
gramineus, Pooecetes, 211
granosa, Trionys, 122'
grasshopper sparrow, 168, 191
gratiosus, Proechimys, 379
gratus, Peromyscus truei 462
gray,
fox, 448
titmouse, 207
oven-bird, 189
owl, 203
gray-headed junco, 212
Great Salt Lake horned lark. 206
great,
blue heron, 196
gray owl, 203
horned owl, 203
horned owl, postnatal development
of, 159
scaup duck, 198
gnat-tailed grackle, 190
greater yellow-legs, 201
grebe,
eared, 196
Holbocll, 196
horned, 196
pied-billed, 196
western, 196
green-tailed towhee, 211
gregorvi, Heliscomys, 481
Grinnell, 207
chickadee, 207
water-thrush, 209
grinnelli, Parus gambeli, 207
gris, zorro, 448
grisegena, Colymbus, 196
griseus,
Catherpes mexicanus, 207
Limnodromus, 201
Parus inornatus, 207
Grison, 437
grosbeak,
evening, 210
pine, 210
Rocky Mountain, 210
western evening, 210
ground snake, 109
grouse,
blue, 199
Columbian, 200
duskv, 199
Franklin, 199
gray ruffed, 199
hoary ruffed, 199
Idaho ruffed, 199
Oregon dusky, 199
Richardson, 199
ruffed, 199
sage, 200
sharp-tailed, 200
Grus,
canadensis, 200
tabida. 200
guairae, Proechimys, 316
gularis, Proechimys, 316
gull,
Bonaparte, 202
California, 202
Franklin, 202
ring-billed, 202
Thayer, 202
guttata, Hylocichla, 208
guyannensis, Proechimys, 355
Gymnocladus dioica, 134
Gymnorhinus cyanocephalus, 206
gymnurus,
Cratogeomys, 252, 452
Platygeomys, 452
haemastica, Limosa, 201
hairy woodpecker, 204
Haldea striatula, 110
Haliaeetus,
leucocephalus, 199
washingtoniensis, 199
haliaetus, Pandion, 199
Index to Volume 1
617
Hall, E. Raymond,
Two new meadow mice from
Michoacan, Mexico, 425
Hall, E. Raymond and Bernardo
Villa R.,
A new pocket gopher (Thomomys)
and a new spiny pocket mouse
(Liomys) from Michoacan, 249
An annotated check list of the
mammals of Michoacan, Mex-
ico, 431
Subspeciation in pocket gophers of
Kansas, 219
Hall, E. Raymond and Walter W.
Dalquest,
A new bat (genus Myotis) from
Mexico, 239
A synopsis of the American bats of
the genus Pipistrellus, 591
Geographic range of the hooded
skunk, Mephitis macroura, with
description of a new subspecies
from Mexico, 575
Pipistrellus cinnamomeus Miller
1902 referred to the genus Myo-
tis, 581
Tadarida femorosacca (Merriam)
in Tamaulipas, Mexico, 245
Hall,
Benjamin D., 435
E. Raymond, 425
Hubert H., 435
Mary F., 435
William Joel, 435
hamiltoni, Corrodopsylla, 140
harlequin duck, 198
Harris sparrow, 212
Hatt, Robert T, 436
hawk,
American rough-legged, 199
broad-winged, 198
Cooper, 198
duck, 199
eastern sparrow, 199
marsh, 199
owl, 203
pigeon, 199
red-tailed, 163, 198
rough-legged, 199
sharp-shinned, 198
sparrow, 163, 199
Swainson, 198
Texas red-shouldered, 183
western pigeon, 199
western red-tailed, 198
Helianthus, tuberosus, 135
Heliscomys, 481
gregoryi, 291, 481
hatcheri, 291
senex, 293
tenuiceps, 289
vetus, 293
Helmitheros vermivorus, 189
hendeei, Proechimys, 315
hermit thrush, 208
hernandezii, Procyon lotor, 446
herodias, Ardea, 196
heron,
black-crowned night, 196
blue, 196
great blue, 196
night, 196
hesperis, Corvus brachyrynchos, 206
hesperomydis, Hoplopleura, 141
Hesperiphona,
brooksi, 210
vespertina, 210
hesperus,
Pipistrellus, 596
Scotophilia, 596
Vesperugo, 596
Heterodon,
contortrix, 107
nasicus, 107
heteromyid rodent, new species from
Oligocene of Colorado, 289
Heteromyidae, defined, 480
Heteromys, 289
hiaticula, Charadrius, 183
hilda, Proechimys, 316
Himantopus mexicanus, 201
himantopus, Micropalama, 201
hirsuta, Hoplopleura, 141
hirsutus, Artibeus, 442
Hirundo,
erythrogaster, 206
rustica, 206
hirundo, Sterna, 202
hispid cotton rat, 465
hispidus, Sigmodon, 137, 138, 465
Histrionicus,
histrionicus, 198
pacificus, 198
histrionicus, Histrionicus, 198
hoactli, Nycticorax nycticorax, 196
hoary bat, 445
Hoffmeister, D. F. and Henry W.
Setzer,
The postnatal development of two
broods of great horned owls
(Bubo virginianus) , 157
hog-nosed snake, 107
Holboell grebe, 196
holbollii, Colymbus grisegena, 196
holbrooki, Lampropeltis getulus, 109
Holbrookia maculata maculata, 105
honeysuckle, 132
hooded skunk, 448
geographic range of, with descrip-
tion of a new subspecies, 577
hoplomyoides, Proechimys, 315
618
University of Kansas Publs., Mus. Nat. Hist.
Hoplopleura,
acanthopus, 141
hi speromydis, 141
hirsuta, 141
horned,
grebe, 196
lark, 205
lizard, 106
horridus, Crotalus, 111
horse nettle, 134
house,
finch, 210
mouse, 468
wren, western, 207
howardi, Orchopeas, 140
Howell nighthawk, 1S5
howelli,
Chordeiles minor, 185
Thomomys bottae, 57
hoyi, Pseudemys noridana, 112
hudsonia, Pica pica, 206
Hudsonian godwit, 201
hudsonius, Circus cyaneus, 199
humeralis, Tayassu angulatus, 469
hummingbird,
black-chinned, 204
broad-tailed, 204
calliope, 204
rufous, 204
Hutchins goose, 182
hutchinsii,
Branta, 197
Branta canadensis, 1S2
Hutton vireo, 209
Hydroprogne caspia, 202
hyemalis, Junco, 191, 212
Hyla,
bistincta, 261
crucifer, 104
proboscidea, 261
versicolor, 88
hylaeus, Geomys bursarius, 234
Hylatomus,
picinus, 204
pileatus, 204
hyleae, Proechimys, 361
hylocetes, Peromyscus, 462
Hylocichla,
almae, 188, 209
auduboni, 208
fuscescens, 188, 208
guttata, 208
salicicola, 188, 208
swainsoni, 188
ustulata, 188, 208
hyberborea, Chen, 197
hypugaea, Speotyto cunicularia, 203
ibis,
eastern glossy, 181
white-faced glossy, 181, 196
Icteria,
auricollis, 209
virens, 189, 209
Icterus bullockii, 210
Idaho,
a check-list of birds of, 195
jay, 206
idoneus, Dipodomys ordii, 546
inexspectatus, Reithrodontomys ful-
vescens, 458
ignotus, Proechimys, 316
iheringi, Proechimys, 373
iliaca, Passerella, 212
illinoensis, Geomys bursarius, 224
immer. Gavia, 195
imparilis,
Cratogeomys gymnurus, 452
Platygeomys gymnurus, 452
inaquosus, Dipodomys ordii, 552
incanus, Bonasa umbellus, 200
industrius, Geomys bursarius, 226
inornatus,
Catoptrophorus semipalmatus, 201
Parus, 207
inermis, Bromus, 133
invictus, Lanius excubitor, 209
Inyo, 207
chickadee, 207
nuthatch, 207
inyoensis, Parus gambeli, 207
Iridoprocne bicolor, 206
irroratus, Liomys, 253, 454
ischaemum, Digitaris, 132
isingu, 470
islandica, Glaucionetta, 198
Ixodes sculptus, 143
Ixoreus,
naevius, 208
meruloides, 208
jabalinde collar, 469
Jackson, H. H. T., 440
jaeger, Pomarine, 202
jaliscensis, Liomys irroratus, 253, 454
Jamaicensis,
Buteo, 198
Oxyura, 198
Jameson, E. W., Jr.,
Natural history of the prairie vole
(mammalian genus Microtus),
125
Japanese honeysuckle, 132
japonica, Lonicera, 132
jay,
black-headed, 206
Idaho, 206
pifion, 206
Woodhouse, 206
Jerusalem artichoke, 135
jeyaquihuira, 464
jihuatz, 449
Index to Volume 1
619
juateanapu, 4G9
jugossicularis, Geomys bursarius, 226
Junco,
caniceps, 212
cismontanus, 191
hyemalis, 191, 212
mearnsi, 212
montanus, 212
oreganus, 212
junco,
Cassiar, 191
gray-headed, 212
Montana. 212
Oregon, 212
pink-sided, 212
slate-colored, 212
kaibabensis, Thomomys talpoides, 23
kangaroo rat, subspeciation in Dip-
odomys ordii, 475
Kansas,
amphibians and reptiles from, 87
reptiles and amphibians, 103
Kentucky coffee tree, 134
kermiti, Proechimys, 345
killdeer, 200
king snake, 108
kingbird, eastern, 205
kingfisher,
belted, 204
western belted, 204
kinglet,
golden-crowned, 208
ruby-crowned, 208
Kinosternon, 283
flavescens. 111
kochi. Laelaps, 142
kuaraki, 449, 450
kyphosis, 119
labecula, Peromyscus maniculatus,
461
labradorius, Passerculus sandwichen-
sis, 190
Lactuca scariola, 133
Ladino clover, 132
laelaps kochi, 142
laeta, Elaphe, 108
lagophonus, Bubo virginianus, 203
Lagopus,
altipetens. 200
leucurus, 200
lagopus, Buteo, 199
lamprochroma, Eremophila alpestris,
205
Lampropeltis,
calligaster, 108
holbrooki, 109
gentilis, 109
getulus, 109
syspila, 109
triangulum, 109
Lanius,
i xcubitor, 209
gambeli, 209
invictus, 209
ludovicianus, 209
lapponicus, Calcarius, 191, 212
lark,
bunting, 211
dusky horned, 205
Great Salt Lake horned, 206
horned, 205
Oregon horned, 205
sparrow, 191, 211
Larus,
argentatus, 202
californicus, 202
delawarensis, 202
Philadelphia, 202
pipixcan, 202
thayeri, 202
Lasionycteris, noctivagans, 241
Lasiurus,
borealis, 444
cinereus, 445
mexicanus, 444
laterale, Leiolopisma, 106
latifrons, Neotoma, 467
latrans, Canis, 449
lazuli bunting, 210
leachii, Glossophaga soricina.. 440
lead-colored bush-tit, 207
leaf-nosed bat, 441
least,
flycatcher, 187
.sandpiper, 201
Leiolopisma laterale, 106
leioprimna, Proechimys, 364
lenis,
Oryzomys fulvescens, 465
Thomomys bottae, 53
lentiginosus, Botaurus, 196
leonada, rata algodonera, 465
lepida, Tachycineta thalassina, 206
Lepidium densiflorum, 133
Leptonycteris,
nivalis, 441
yerbabuenae, 441
leptosoma, Mus, 392
Lepus callotis, 469
lesser,
Canada goose, 197
loon, 195
scaup duck, 198
snow goose, 197
yellow-legs, 201
lettuce, wild, 133
leucocephalus, Haliaeetus, 199
leucomystax, Proechimys, 352
leucopareia, Branta canadensis, 197
620
University of Kansas Publs., Mus. Nat. Hist.
leucoparia,
Mustela frenata, 447
Putorius frenatus, 447
leucophrys, Zonotrichia, 212
Leucophoyx,
brewsteri, 196
thula, 196
leucoptera, Loxia, 211
leucopus,
Peromyscus, 137, 140
Orchopeas, 140
Leucosticte,
atrata, 211
lit (oralis, 211
1: phrocotis, 211
leucurus,
Dendrocopos pubescens, 205
Lagopus, 200
levidensis, Thomomys bottae, 54
Ievipes, Peromyscus bo3'lii, 462
levis, Thomomys talpoides, 24
levisagittalis, Geomys bursarius, 234
Lewis woodpecker, 204
lewis, Asyndesmus, 204
Liatrus, 135
liebre, 469
limicola, Rallus, 200
liminalis, Proechimys, 343
limnaeus, Seiurus noveboracensis, 189
Limnodromus,
griseus, 201
scolopaceus, 201
limnodytes, Chrysemys, 269
Limosa,
fedoa, 184, 201
haemastica. 201
Lincoln sparrow, 212
lincolnii, Melospiza, 212
lineatum, Tropidoclonion. 110
lined snake, 110
Liponyssus occidentalis, 142
Liomys,
acutus, 253, 455
alleni, 253, 455
irroratus, 253
jalicensis, 454
jaliscensis, 253, 455
parviceps, 454
p ictus, 289, 454
plant inarensis, 454
Lissemys, 122
Listrophorus, 143
little flycatcher, 205
littoralis. Leucosticte tephrocotis, 211
livia, Columba, 168
lizard,
collared, 105
earless, 105
horned, 106
northern plains, 106
plains, 106
Texas horned, 106
llanensis, Geomys bursarius, 234
Lobipes, lobatus, 202
lobatus, Lobipes, 202
Loncheres,
elegans, 387
myosuros, 351
long-billed,
curlew, 201
dowitcher, 201
marsh wren, 207
long-eared,
bat, 445
owl, 203
long-tailed, 206
chickadee, 206
long-tongued bat, 440, 441
longicauda, Euiycea, 88
longicaudatus, Proechimys, 351
longipes,
Cricetodipus, 556
Dipodomys ordii, 556
Dipodops, 556
Perodipus, 556
longspur, Alaskan, 191. 212
Lonicera japonica, 132
loon,
lesser, 195
red-throated, 195
Lophortyx,
brunnescens, 200
californica, 200
lot or, Procyon, 446
Louisiana, birds of, 179
louisianensis, Triturus viridescens, 103
Lowery, George H., Jr., Additions to
the list of the birds of Louisiana,
179
Loxia,
bendirei, 211
benti, 211
curvirostra, 211
leucoptera, 211
lucifuga, Eurycea, 88
lucifugus, Myotis, 583
ludoviciana, Piranga, 210
ludovicianus,
Lanius, 209
Microtus, 129
Luna, P., 440
luteolus,
Dipodomys ordii, 533
Perodipus ordii, 533
lutescens,
Geomys bursarius, 222
Setaria, 132
lutosus, Myotis yumanensis. 443
Lutra, 437
Index to Volume 1
621
Lynx,
escuinapae, 449
rufus, 449
ruffus, 449
M. Dale Arvey, A check list of the
birds of Idaho, 195
MacFarlane screech owl, 203
macfarlanei, Otus asio, 203
Macgillivray warbler, 209
macrotis, Corynorhinus, 445
macroura,
Mephitis, 577
Mephitis macroura, 448
Zenaidura, 168, 202
macularia, Actitis, 201
maculata, Holbrookia, 105
maculatus, Pipilo, 211
magpie, American, 206
mallard, 197
major, Geomys bursarius, 229
majusculus, Geomys bursarius, 223
manchado, zorillo, 447
maniculatus, Peromyscus, 137, 461
mapache, 446
marbled godwit, 184, 201
marchiana, Thamnophis, 99
morcomi, Dendroica petechia, 209
Mareca americana, 197
marginella, Zenaidura macroura, 202
marila, Aythya, 198
Marmosa canescens canescens, 439
marsh,
hawk, 199
mouse, 461
wren, western long-billed, 207
marshalli, Dipodomys ordii, 551
mascotensis, Sigmodon hispidus, 465
masked duck, 182
massasauga, 111
Masticophis,
flagellum, 108
testaceous, 108
mastin, murcielago, 445
mastiff bat, 445
mauri, Ereunetes, 201
mavortium, Ambystoma tigrinum, 103
maximus, Pipistrellus hesperus, 598
meadow mouse, 168
Mexican, 467
meadowlark, 158
western, 190, 210
mearnsi, Junco oreganus, 212
Medicago sativa, 133
mediogriseus, Passerculus sandwichen-
sis. 190
medius, Dipodomys ordii, 519
Megaceryle,
alcyon, 204
caurina, 204
megalotis,
Reithrodontonrys, 137, 455
Corynorhinus, 445
Melanitta,
fusca, 198
perspicillata, 198
melanocephalus, Pheucticus, 210
melanocorys, Calamospiza, 211
melanoleucus, Totanus, 201
melanophrys, Peromyscus, 463
melanopleura, Eurycea longicauda, 88
melanotis,
Sigmodon, 465
Sitta pygmaea, 207
Peromyscus, 464
melanotos, Erolia, 201
Melilotus alba, 133
melodia, Melospiza, 212
Melospiza,
alticola, 212
fallax, 212
lincolnii, 212
melodia, 212
merrilli, 212
Mephitis,
eximius, 579
macroura, 577
macroura macroura, 448
mephitis, 57S
milleri, 578
vittata, 578
mephitis, Mephitis, 577
merganser,
American, 198
Mergus, 198
red-breasted, 198
merriami,
Pipistrellus hesperus, 597
Vesperugo, 597
Merrill song sparrow, 212
merrilli,
Eremophila alpestris, 205
Melospiza melodia, 212
meruloides, Ixoreus naevius, 208
mesamericana, Didelphis, 438
mesoleucus, Conepatus, 448
mesquite great-tailed grackle, 190
metorito, 467
Mexican,
cottontail, 469
free-tailed bat, 445
meadow mouse, 467
mexicana,
Choeron3'cteris, 441
Tadarida, 445
Mexicano, conejo, 469
mexicanus,
Atalapha, 444
Carpodacus, 210
Catherpes, 207
622
University of Kansas Publs., Mus. Nat. Hist.
mexicanus — Concluded
Cassidix, 190
Cinclus, 207
Corynorhinus megalotis, 445
Corynorhinus rafinesquii, 445
Dasypus novemcinctus, 470
Falco, 199
Himantopus, 201
Lasiurus borealis, 444
Microtus, 425, 467
Molossus, 445
Myotis californicus, 444
Orvzomvs couesi, 465
Sialia, 208
Microdipodops, 480
Micropalama himantopus, 201
Microtus,
agrestis, 141
californicus, 142
fulviventer, 425
fundatus, 425, 467
ludovicianus, 129
mexicanus, 425, 467
ochrogaster, 128, 168
pennsylvanicus, 128
phaeus, 426, 468
salvus, 426, 467
Michoacan,
Mexico, an annotated check list of
the mammals of, 435
squirrel, 450
Microhyla olivacea, 105
migratorius,
Ectopistes, 202
Turdus, 168, 188, 208
milk snake, 109
milleri, Mephitis macroura, 578
millet, wild, 134
Mimus polyglottos, 168
mincae, Proechimys, 315
minimus,
Empidonax, 187
Psaltriparus, 207
Thomomys bottae, 32
minor, Chordeiles, 185, 186, 204
miniiscula, rata montera, 467
minutilla, Erolia, 201
miradorensis,
Eptesicus, fuscus, 444
Scotophilus, 444
misicpapu, 449
mockingbird, 168
mofcta rayada, 448
moffitti, Branta canadensis, 196
mohavensis, Dipodomys panaminti-
nus, 489
mokeson, Agkistrodon, 111
molaris, Nasua narica, 446
Molossus mexicanus, 445
Molothrus,
artemisiae, 210
ater, 210
monoensis,
Dipodomys ordii, 528
Perodipus, 528
Montana,
horned owl, 203
junco, 212
montane, Lincoln sparrow, 212
montana, Pinicola enucleator, 210
montanus,
Dipodomys ordii, 538
Junco oreganus, 212
Oreoscoptes, 208
Perodipus, 538
monticola, Dendrocopos villosus, 204
montera, rata,
frentuda, 467
miniiscula, 467
Moojen, Joao, Speciation in the Bra-
zilian spiny rats (genus Proechi-
mys, family Echimyidae), 301
Mookomys formicorum, 481
moorei, Thomomys talpoides, 19
Mountain vireo, 188
mountain, 206
bluebird, 208
chickadee, 206
song spaiTOw, 212
montanero, raton, 464
moreno, raton, 457
morning glory, wild, 135
Morus bassanus, 180
mourning dove, 168, 202
mouse,
black-eared, 464
blackish, 463
brush, 461
deer, 461
fulvous harvest, 457
harvest, 455
house, 468
marsh, 461
meadow, 168, 467
Mexican meadow, 467
northern pigmy, 460
northern spiny pocket, 454
pinon, 462
pygmy, 460
spiny pocket, 454
Tarascan, 463
tropical pygmy, 460
volcano harvest, 459
western harvest, 455
western spiny pocket, 454
woods, 462
mud turtle, 111
murcielago,
California, 444
carilargo, 441
colirugosa, 444
coludo, 445
fusco, 444
lenguilarga, 441
Index to Volume 1
623
murcielago — Concluded
mastin, 445
narigudo, 445
pai'do, 445
rojizo, 440
sacoptero, 440
siricotero, 440
vespertino, 443
Yuma, 444
zapotero, 442
murine opossum, 439
murinus, Desmodus rotundus, 443
Mus,
alexandrinus, 468
cinnamoneus, 392
leptosoma, 392
musculus, 468
musarana, 439
colicorta, 440
Muscivora forficata, 186
musculus,
Baiomys, 460
Mus, 468
Peromyscus, 460
Sitomys, 460
musk turtle, 111
Mustela,
frenata, 447
leucoparia, 447
mutica, Amyda, 112, 119
Myadestes townsendi, 208
Mviarchus cinexascens cinerascens,
186, 205
Myocoptes, 143
myosuros,
Echimys, 351
Echinomys, 392
Loncheres, 351
Proechimys, 392
Myotis,
albescens, 239
argentatus, 239
baileyi, 587
californicus, 444
carissima, 585
cinnamomeus, 585
fortidens, 585, 586
lucifugus, 583
lutosus, 443
mexicanus, 444
occultus, 587
ruber, 585
thysanodes, 444
velifer, 443, 588
yumanensis, 443
myotis,
California, 444
fringe-tailed, 444
Yuma, 443
naevius, Ixoreus, 208
narica, Nasua, 446
narigudo, murcielago, 445
nasicus, Heterodon, 107
Nasua,
molaris, 446
narica, 446
Natrix,
erythrogaster, 109
grahamii, 109
rhombifera, 109
sipedon, 109
transversa, 109
Navy, United States, 479
nebraskensis, Cupidinimus, 483
nebulosa, Strix, 203
neglecta, Sturnella, 190, 210
negra, rata, 468
nelsoni,
Conepatus mesoleucus, 448
Reithrodontomys fulvescens, 457
Nelsonia goldmani, 467
nemoralis, 450
Pitymys, 128
Neotoma,
ferruginea, 467
latifrons, 467
tenuicauda, 467
nesiotes, Proechimys, 363
nesophilus, Thomomys bottae, 34
nettle, horse, 134
Nevada,
cowbird, 210
red-wing, 210
savannah sparrow, 211
sparrow, 211
towhee, 211
nevadensis,
Agelaius phoeniceus, 210
Amphispiza belli, 212
Parus atricapillus, 206
Passerculus sandwichensis, 190, 211
Newfoundland,
oven-bird, 189
robin, 188
newt, 103
nexilis, Dipodomys ordii, 559
nicholi, Thomomys bottae, 66
niger. Sciurus, 168
nighthawk,
Cherrie, 186
eastern, 185
Howell, 185
Pacific, 204
Sennett, 186
nigra, Chlidonias, 202
nigrescens, Dendroica, 209
nigricans, Branta bernicla, 197
nigriceps, Tantilla, 110
nigricollis, Colymbus, 196
nigrideus, Turdus migratorius, 188
nigrita, Pseudacris, 104
nigrofulvus, Proechimys, 316
624 University of Kansas Publs., Mus. Nat. Hist.
nivalis,
Leptonycteris nivalis, 441
Plectrophenax, 213
nivosus, Charadrius alexandrinus, 183
noctivagans, LasionycterLs, 241
norteno, raton, 454
pigmeo, 460
northern,
bald eagle, 199
canyon wren. 207
cliff swallow, 206
curlew, 201
pileolated warbler, 210
phalarope, 202
pygmy mouse, 460
sage sparrow, 212
spiny pocket mouse, 454
varied thrush, 208
white-headed woodpecker, 205
northwestern,
creeper, 207
horned owl, 203
long-billed marsh wren, 207
marsh wren, 207
shrike, 209
norvegicus, Rattus, 168
Norway rat, 168
Nosopsyllus fasciatus, 140
notabilis, Seiurus noveboracensis, 209
noveboracensis, Seiurus, 189, 209
novemcinctus, Dasypus, 470
nuchalis, Sphyrapicus vai'ius, 2-4
Nucifraga Columbiana, 206
Numenius,
americanus, 183, 201
parvus, 183, 201
nutcracker, Clark, 206
nuthatch,
black-eared, 207
Inyo, 207
red-breasted, 207
Nuttall poorwill, 204
nuttallii, Phalaenoptilus, 204
Nuttallornis borealis, 205
\yctea scandiaca, 203
\ vH icorax,
hoactli, 196
nycticorax, 196
nycticorax, Nycticorax, 196
o'connelli, Proechimys, 316
obscuro, raton, 463
obscurus,
Dendragapus, 199
Dipodomys ordii, 521
Perodipus, 521
Pipistrellus subflavus, 600
obsoleta, Klaphe, 108
obsoletus,
Eumeces, 106
S'alpinctes, 207
occidental, raton espinoso, 454
occidentalis,
Aechmophorus, 196
Branta canadensis, 196
Bubo virginianus, 203
Coccvzus americanus. 202
Geothlypis, 189, 209
Liponyssus, 142
Sialia mexicanus, 208
occultus, 587
ocelot, 449
ochracea, Spizella arborea. 212
ochraceus, Proechimys, 316
ochrogaster, Microtus, 128, 168
ocius, Thomomys talpoides, 17
ocotero, raton, 462
Odocoileus,
sinaloae, 470
virginianus, 470
odoratus, Sternotherus, 111
officinale, Taraxacum, 134
Oklahoma, amphibians and reptiles
from, 87
oklahomae, Dipodomys ordii, 514
oligantha, Aristada, 132
olivacea, Microhyla, 105
olivaceus, Rcgulus satrapa, 208
olive-backed,
thrush, 188
thrush, western, 208
olive-sided flycatcher, 205
oquirrhensis, Thomomys talpoides, 11
Ophisaurus ventialis, 106
Oporornis, 209
opossum, 438
murine, 439
orange-crowned warbler, 209
Orchopeas,
howardi, 140
leucopus, 140
wickhami, 140
ordi, Perodipus, 530
ordii,
Cricetodipus, 530
Dipodomys ordii, 530
Dipodops, 530
Perodipus, 530
oreganus, Junco, 212
Uregon,
horned lark, 205
junco, 212
orejudo, raton, 455
Oreoscoptes montanus, 208
Oreortyx picta picta, 200
orestera, Vermivora celata, 209
oriole, Bullock, 210
oris, Proechimys, 365
ornata, Terrapene, 111
oryzivorus, Dolichonyx, 210
Oryzomys,
couesi, 484
fulvescens, 4G5
Index to Volume 1
625
Oryzomys — Concluded
lenis, 465
mexicanus, 465
rcgillus, 464
osgoodi, Thomomys bottae, 56
osprey, 199
Otus,
asio, 203
brewsteri, 203
flammeolus, 203
macfarlanei, 203
otus, Asio, 2u3
oven-bird.
gray. 189
Newfoundland, 189
owl,
American hawk, 203
Arctic horned, 203
barn, 163, 203
Brewster screech, 203
burrowing, 203
California pygmy, 203
Flammulated screech, 203
gray, 203
great horned, 159, 203
great gray, 203
hawk, 203
horned, great, 159, 203
long-eared, 164, 203
MacFarlane screech, 203
Montana horned, 203
northwestern "horned, 203
pygmy, 203
Richardson, 203
saw-whet, 204
screech, 164, 203
short-eared, 203
snowy, 203
western burrowing, 203
Oxyura.
dominica, 182
jamaicensis, 198
rubida, 198
pachita, Proechimys, 316
Pacific,
golden plover, 200
nighthawk, 204
pacificus,
Anthus spinoletta, 208
Histrionicus histrionicus, 198
Troglodytes troglodytes, 207
painted turtle, 112
pale goldfinch, 211
paleontology of kangaroo rats, 480
pallescens,
Columbigallina passerina, 185
Corynorhinus pallescens, 445
pallid black-capped, 206
chickadee, 206
pallidiceps, Bombycilla garrulus, 208
pallidus, Castor canadensis, 409
pallidus,
Dendragapus obscurus, 199
Dipodomys ordii, 558
Spinus tristis, 211
palmeri,
Cricctodipus ordii, 562
Dipodomys ordii, 562
Perodipus ordii, 562
palustris, Telmatodytes, 207
panamensis, Proechimys, 316
Pandion,
carolinensis, 199
haliaetus, 199
panema, Proechimys, 380
panguitchensis, Dipodomys ordii, 527
paratus, Proechimys, 382
pardo, murcielago, 445
parietalis, Thamnophis sirtalis, 110
parkmanii, Troglodytes aedon, 207
parowanensis, Thomomys talpoides,
22
partridge, European, 200
Parus,
atricapillus, 206
fortuitus, 206
gambeli, 206
grinnelli, 207
griseus, 207
inornatus, 207
inyoensis, 207
nevadensis, 206
rufescens, 207
septentrionalis, 206
parva, Cryptotis, 137
parviceps, Liomys pictus, 454
parvissima, Rana, 105
parvus, Numenius americanus, 183,
201
passenger pigeon, 202
Passer domesticus, 210
Passerculus,
labradorius, 190
mediogriseus, 190
nevadensis, 190, 211
sandwichensis, 190, 211
Passerella,
iliaca, 212
schistacea, 212
Passerina amoena, 210
passerina,
Columbigallina, 185
Spizella, 212
pavidus, Peromyscus, 462
peccary, collared, 469
pectoral sandpiper, 201
Pedioecetes,
columbianus, 200
phasianellus, 200
Pelecanus erythrorhynchos, 196
pelican, white, 196
pennsylvanicus, Microtus, 128
626
University of Kansas Publs., Mus. Nat. Hist.
peppergrass, 133
Perdix perdix perdix, 200
perdix, Perdix, 200
peregrinus, Falco, 199
perfulvus, Peromyscus, 461
pergracilis,
Blarina, 440
Cryptotis, 440
Perisoreus,
bicolor, 206
canadensis, 206
Perodipus,
chapmani, 536
columbianus, 544
compactus, 515
longipes, 525
luteolus, 533
monoensis, 528
montanus, 518
obscurus, 521
ordi, 530
ordii, 556
palmeri, 562
richardsoni, 518
sennetti, 517
utahensis, 543
Perognathinae. 480
Peromyscopsylla scotti, 140
Peromyscus.
banderanus, 463
boylei, 461
boylii, 461
evides, 461
gratus, 462
hylocetes, 462
labecula, 461
leucopus. 137
levipes, 462
maniculatus, 137, 461
melanophiys, 463
melanotis, 464
pavidus, 462
perfulvus, 13
sagax, 462
sonoriensis, 461
spicilegus, 461
truei, 462
vicinior, 463
zamorae, 463
zelotes, 462
perpallidus, Thomomys, 28
perplexis, Dipodomys agilis, 489
perspicillata, 198
petechia, Dendroica, 209
Petrochelidon,
albifrons, 206
pyrronota, 206
pewee, western wood, 205
pheasant, ring-necked, 200
phaeus, Microtus mexicanus, 425, 467
phaia, Bonasa umbellus, 199
Phalacrocorax,
auritus, 196
albociliatus, 196
Phalaenoptilus nuttallii nuttallii, 204
phalarope,
northern, 202
red, 202
Wilson, 184, 202
Phalaropus fulicarius, 202
phasianellus, Pedioecetes, 200
Phasianus colchicus, 200
Pheucticus melanocephalus, 210
Philadelphia, Larus, 202
phoebe, Say, 205
phoeniceus, Agelaius, 168, 210
Phrynosoma cornutum, 106
Phyllomys, 310
plrylogeny of kangaroo rats, 484
Pica,
hudsonica, 206
pica, 206
pica, Pica, 206
picinus, Hylatomus pileatus, 204
Picoi'des.
arcticus, 205
dorsalis, 205
fasciatus, 205
tridactylus, 205
picta,
Chrysemys, 112, 270
Oreortyx, 200
pictus, Liomys, 289, 454
pied-billed grebe, 196
pigeon,
band-tailed, 202
hawk, 199
passenger, 202
pigmea, rata arrocera, 465
pigmeo norteno, raton, 460
pigra. Didelphis virginiana, 439
pileated woodpecker, western, 204
pileatus, Hylatomus, 204
pileolata, 210
pileolated warbler, 210
pillus. Thomomys umbrinus, 251
pilot black snake, 108
pine,
grosbeak, Rocky Mountain, 210
siskin, western, 211
pinetorum, Pitymys, 12S
Pinicola,
enucleator, 210
montana, 210
pink-sided junco, 212
pifion mouse, 462
pinonero, raton, 462
pintail, American, 197
pinus, Spinus, 211
pipiens, Rana, 104
Index to Volume 1
G27
Pipilo.
arcticus, 211
curtatus, 211
maculatus, 211
Pipistrellus,
apus, 597
australis, 597
cinnamomeus, 583, 593
hesperus, 596
maximus, 598
merriami, 597
obscurus, 600
santarosae, 59S
subflavus, 584, 599
synopsis of the American bats of
the genus, 593
veracrucis, 601
pipit, western, 188, 208
pipixcan, Larus, 202
Piranga ludoviciana, 210
Pituophis,
catenifer, 108
sayi, 108
Pitymys,
nemoralis, 128
pinetorum, 128
pizote, 446
planirostre, Phyllostoma, 442
planirostris,
Artibeus, 442
Thomomys bottae, 66
plantinarensis, Liomys pictus, 454
platycephalus, Selasphorus, 204
Platygeomys,
angustirostris, 452
gymnurus, 452
imparilis, 452
tylorhinus, 452
varius, 453
Platypeltis, 121
platypterus, Buteo, 198
platyrhynchos, Anas, 197
Plectrophenax, 213
plegadis,
falcinellus, 181
mexicana, 181
plicata, Balantiopteryx, 440
plover,
American golden, 184
black-bellied, 200
Cuban snowy, 183
golden, 184, 200
Pacific, golden, 200
semipalmated, 183
snowy, 183
western snowy, 183
Wilson, 184
plumbeus, Psaltriparus minimus, 207
plumed quail, 200
Pluvialis,
dominica, 184, 200
fulva. 200
pluvialis, Eumeces, 85
Poa,
annua, 133
pratensis, 132
pocket gopher,
Michoacan, 453
new from Mexico, 251
plains, 452
southern, 451
pocket gophers of,
Kansas, 219
Utah, 3
pocket mouse,
northern spiny, 454
spiny, 454
western spiny, 454
podiceps, Podilymbus, 196
Podilymbus podiceps, 196
poliopus, 451
Proechimys, 316
Polioptila,
amoenissima, 208
caerulea, 208
polyglottos, Mimus, 168
Pomarine jaeger, 202
pomarinus, Stercorarius, 202
Pooecetes,
confinis, 211
gramineus, 211
poorwill, Nuttall, 204
Porzana Carolina, 168, 200
postnatal development, homed owl,
159
Potos, 437
prairie,
falcon, 199
threeawn, 132
pratincola, Tyto alba, 203
pratensis,
Ammodramus savannarum, 191
Poa, 132
priscus, Dipodomys ordii, 547
proboscidea, Hyla, 259
Procyon,
hernandezii, 446
lotor, 446
Prodipodomys, 480
kansensis, 481
minor, 484
Proechimys,
albispinus, 388
amphichoricus, 344
arabupu, 369
arescens, 366
boimensis, 350
bolivianus, 316
bonafidei, 378
628
University of Kansas Publs., Mus. Nat. Hist.
Proechimys — Concluded
brevicauda, 349
burriis, 316
calidior, 316
canicollis, 315
cayennensis, 314
centralis, 316
cherriei, 316
chiriquinus, 316
chrysaeolus, 316
colombianus, 316
decumanus, 316
denigratus, 381
dimidiatus, 371
elegans, 387
fuliginosus, 308
goeldi, 338
goldmani, 316
gorgonae, 316
gratiosus, 379
guairae, 316
gularis, 316
guyannensis, 355
hendeei, 315
hilda, 316
hoplomyoides, 315
hyleae,'361
ignotus, 316
iheringi, 373
kermiti, 345
leioprimna, 364
leucomystax, 352
liminalis, 343
longicaudatus, 351
mincae, 315
myosuros, 392
nesiotes, 363
nigrofulvus, 316
ochraceus, 316
o'connelli, 316
oris, 365
pachita, 316
panamensis, 316
panema, 380
paratus, 382
poliopus, 316
rattinus, 316
ribeiroi, 361
riparum, 367
roberti, 353
rosa, 316
rubellus, 316
securus, 316
semispinosus, 342
sertonius, 391
setosus, 385
simonsi, 316
steerei, 338
trinitatis, 316
urichi, 316
vacillator, 316
villicauda, 355
warreni, 316
propinquus, Turdus migratorius, 208
prosopidicola, Cassidix mexicanus, 190
proximus, Thamnophis sauritus, 110
Pseudacris,
nigrita, 104
triseriata, 104
pseudagyrtes, Ctenophthalmus, 140
Pseudemys, 270, 2S3
elegans, 112
floridana, 112
hoyi, 112
scripta, 112, 270
ptarmigan, white-tailed, 200
pubescens, Dendrocopos, 205
pullus, Thomomj's umbrinus, 251, 451
pullus. Troglodytes troglodytes, 188
pulverius, Telmatodytes palustris, 207
punctatus, Diadophis, 107
purple finch, Cassin, 210
pusilla, Wilsonia, 190, 210
pygmaeus, Thomomys talpoides, 14
pygmy owl, 203
pygmy mouse,
northern, 460
tropical, 460
Pyrocephalus,
mexicanus, 187
rubinus, 187
pyrronota, Petrochelidon, 206
quadratus, Thomomys, 6
quail,
plumed, 200
California, 200
querula, Zonotrichia, 212
Quiscalus versicolor, 168
rabbit,
jack, 469
white-sided jack, 469
raccoon, 446
racer, blue, 107
radix, Thamnophis, 99, 110
rafinesquii, Corynorhinus, 445
ragweed, giant, 134
rail, Virginia, 200
Rallus limicola, 200
Rana,
brachycephala, 104
cantabrigensis, 104
catesbeiana, 104
parvissima, 104
sylvatica, 104
rat.
black, 468
dwarf wood, 467
ferruginous wood, 467
fulvous cotton, 465
fulvous rice, 465
hispid cotton, 465
Index to Volume 1
629
rat — Concluded
kangaroo, subspeciation in. 475
Norway, 168
rice, 464
snake, 10S
tropical rice, 464
white-throated wood, 467
wood, 467
rata,
algodonera, 465
arrocera tropical, 464
leonada, algodonera, 465
minuscula, montera, 467
montera, 467
negra, 468
pigmea, arrocera, 465
setosa, algodonera, 465
tropical, arrocera, 464
raton
brefiero, 461
chaparral, de, 461
cuatralvo, 461
dorado, 459
montanero, 464
moreno, 457
norteno, 454
nortefio pigmeo, 460
obscuro, 463
ocotero, 462
occidental espinoso, 454
orejudo, 455
pigmeo, 460 .
pinonero, 462
Tarasco, 463
tlacuache, 439
tropical pigmeo, 460
volcanes, 466
rats, spiny, 303
rattinus, Proechimys, 316
rattlesnake, 111
prairie, 111
timber, 111
Rattus,
alexandrinus, 468
norvegicus, 168
rattus, 468
rattus, Rattus, 468
raven, American, 206
ravus, Thomomys talpoides, 15
Recurvirostra americana, 201
red, 211
bat, 444
phalarope, 202
red-breasted,
merganser, 198
nuthatch, 207
red-eyed vireo, 209
red-naped sapsucker, 204
red-shouldered hawk, 183
red-tailed hawk, 198
red-throated loon, 195
red-wing, 168, 210
Nevada, 210
thick-billed, 210
reddish egret, 181
redhead, 197
redpoll, common, 211
redstart, American, 210
regalis, Buteo, 199
regillus, Oryzomys couesi, 464
Regulus,
calendula, 208
cineraceus, 208
olivaceus, 208
satrapa, 208
Reithrodontomys,
chrysopsis chrysopsis, 459
fulvescens, 457
inexspectatus, 458
megalotis, 137, 455
nelsoni, 457
saturatus, 455
seclusus, 459
tenuis, 457
toltecus, 458
zacatecae, 456
repens, Agropyron, 133
repentinus, Castor canadensis, 409
restrictus, Sylvilagus floridanus, 469
rhombifera, Natrix, 109
ribbon snake, 110
ribeiroi, Proechimys, 361
rice rat,
fulvous, 465
tropical, 464
Richardson owl, 203
richardsoni,
Aegolius funereus, 203
Dipodomys ordii, 511
Perodipus montanus, 518
richardsonii,
Contopus, 205
Dendragapus obscurus, 199
ridgwayi, Vermivora ruficapilla, 209
ring-billed gull, 202
ring-necked
duck, 197
pheasant, 200
snake, 107
ring-tailed cat, 447
riggsi, Testudo, 284
Riparia riparia, 206
riparia, Riparia, 206
riparum, Proechimys, 367
road-runner, 184
roberti, Proechimys, 353
robin, 168, 188, 208
robustus, Thomomys bottae, 30
rock,
dove, 168
squirrel, 449
wren, 207
630
University of Kansas Publs., Mus. Nat. Hist.
Rocky Mountain,
grosbeak, 210
hairy woodpecker, 204
orange-crowned warbler, 209
pine grosbeak, 210
yellow warbler, 209
rojizo, murcielago, 444
rosa, Proechimys, 316
Ross goose, 197
rossi, Chen, 197
rostralis, Castor canadensis, 411
rosy finch, 211
black, 211
gray-crowned, 211
Hepburn, 211
rotundus, Desmodus, 443
rough-leg, ferruginous, 199
rough-legged hawk, 199
rough-winged swallow, 206
rubellus, Proechimys, 316
ruber, Myotis, 585
rubida, Oxyura jamaicensis, 198
ruby-crowned kinglet, 208
ruddy duck. 198
rufescens,
Dichromanassa, 1S1
Parus, 207
Thorn omys, 3
ruffed grouse, 199
ruffus, Lynx, 449
ruficapilla, Vermivora, 209
ruficollis, Stelgidopteryx, 206
ruficrissa, Dumetella carolinensis, 207
rufous hummingbird, 204
rufum, Toxostoma, 168
rufus,
Lynx, 449
Selasphorus, 204
rustica, Hirundo, 206
ruticilla, Setophaga, 210
sac-winged bat, 440
sacoptero, murcielago, 440
sagax, Peromyscus, 462
sage,
grouse, 200
sparrow, 212
thrasher, 208
salamander,
narrow-mouthed, 103
Tiger, 103
salicicola, Hylocichla fuscescens, 188
208
Salpinctes obsoletus obsoletus, 207
salvus, Microtus mexicanus, 426, 467
s[ancti].-johannis, Buteo lagopus 199
sanderling, 201
sandhill crane, 200
sandpiper,
Baird, 184
least, 201
pectoral, 201
solitary, 201
spotted, 201
stilt, 201
western, 201
western solitary, 201
sanctiluciae, Dipodomys venustus, 489
sandwichensis, Passerculus, 190, 211
sanrafaeli, Dipodomys ordii, 526
santarosae, Pipistrellus hesperus, 598
sapsucker,
red-naped, 204
Williamson, 204
sarisi, 464
sauritus, Thamnophis, 110
sativa, Medicago, 133
satrapa, Regulus, 208
saturatus, Reithrodontomys megalotis,
455
Saussure shrew, 439
saussurei, Sorex, 439
savannah sparrow, 190, 211
savannarum, Ammodramus, 168, 191
saw-whet owl, 204
saxatalis, Aeronautes, 204
Say phoebe, 205
say a, Sayornis, 205
sayi, Pituophis catenifer, 108
Sayornis saya, 205
scandiaca, Nyctea, 203
Scalopus aquaticus, 137
Scaphiopus, bombifrons, 103
scariola, Lactuca, 133
scaup,
greater, 198
lesser, 198
Sceloporus,
garmani, 106
undulatus, 106
Scirpus, 132
scissor-tailed flycatcher, 186
schistacea, Passerella iliaca, 212
Sciurus,
albipes, 450
nemoralis, 450
niger, 168
poliopus, 450
senex, 451
variegatus, 449
scolopaceus, Limnodromus griseus, 201
scoter,
surf, 198
white-winged, 198
Scotophilus,
georgianus, 599
hesperus, 596
miradorensis, 444
scotti, Peromyscopsylla, 140
screech owl, 203
scripta, Pseudemys, 112, 270
Index to Volume 1
631
sculptus, Ixodes, 143
seclusus, Reithrodontoinys chrysopsis,
459
securus, Proechimys, 316
Seiurus,
cinereus, 189
furvior, 189
limnaeus, 189
notabilis, 209
noveboracensis, 189, 209
Selasphorus,
platycephalus, 204
rufus, 204
semipalmated plover, 183
semipalmatus,
Catoptrophorus, 201
Charadrius hiaticula, 183
semispinosus, Proechimys, 342
senex, 451
Sennett nighthawk, 186
sennetti,
Chordeiles minor, 186
Cricetodipus, 517
Dipodomys ordii, 517
Perodipus, 517
sepium, Convolvulus, 135
septentrionalis,
Eumeces, 106
Parus atricapillus, 206
serpentina, Chelydra, 111
serrana, tuza, 451
serrator, Mergus, 198
serripennis, Stelgidopteryx ruficollis,
206
sertonius, Proechimys, 391
Setaria,
lutescens, 132
viridis, 132
Setophaga ruticilla, 210
setosa, rata algodonera, 465
setosus,
Echimys, 385
Proechimys, 385
Setzer, Henry W.,
Subspeciation in the kangaroo rat,
Dipodomys ordii, 475
The postnatal development of two
broods of great horned owls
(Bubo virginianus), 159
sevieri, Thomomys bottae, 45
sexlineatus, Cnemidophorus, 106
sharp-shinned hawk, 198
sharp-tailed grouse, 200
short-eared owl, 203
shoveller, 197
shrew,
Saussure, 439
short-tailed, 440
shrike,
California, 209
northwestern, 209
Sialia,
currucoides, 208
mexicanus, 208
occidentalis, 208
Sigmodon,
atratus, 466
hispidus, 137, 138, 465
mascotensis, 465
melanotis, 465
sigmodoni, Articholaelaps, 142
simonsi, Proechimys, 316
sinaloae, Odocoileus virginianus, 470
sinensis, Amyda, 119
sinuatus, Corvus corax, 206
sipedon, Natrix, 109
siricotero, murcielago, 440
sirtalis, Thamnophis, 110
siskin, western pine, 211
Sitta,
canadensis, 207
carolinensis, 207
caurina, 207
pygmaea, 207
melanotis, 207
tenuissimus, 207
skink,
common five-lined, 106
five-lined, 106
Sonoran, 106
skunk,
hooded, 448
spotted, 447
slate-colored,
fox sparrow, 212
junco, 212
Smith, Hobart M.,
Hybridization between two species
of garter snakes, 99
Kyphosis and other variations in
soft-shelled turtles, 119
The systematic status of Eumeces
pluvialis Cope, and noteworthy
records of other amphibians and
reptiles from Kansas and Okla-
homa, 87
The tadpoles of Bufo cognatus
Say, 95
Smith, Hobart M., John Breukelman
and, Selected records of reptiles and
amphibians from Kansas, 103
smooth brome, 133
snake,
black, 108
black-headed, 110
bull, 108
common bull, 108
common water, 109
diamond-backed water, 109
Emory rat, 108
garter, 110
Graham, 109
632
University of Kansas Publs., Mus. Nat. Hist.
.snake — Concluded
ground, 110
Great Plains black-headed, 110
hog-nosed, 107
king, 108
lined, 110
milk, 109
pilot black, 108
plains garter, 110
prairie ring-necked, 107
rat, 108
raffle, 111
red milk, 109
red-sided garter, 110
ribbon, 110
ring-necked, 107
pout hern ground, 110
speckled king, 109
water, 109
western milk, 109
western ribbon, 110
western worm, 107
worm, 107
yellow-bellied king, 108
yellow-bellied water, 109
snapping turtle. 111
snipe, Wilson, 201
snow bunting, eastern, 213
snowy,
owl, 203
plover, 183
soft-shelled turtle, 112, 119
Solanum carolinense, 135
solitaire, Townsend, 208
solitaria, Tringa, 201
solitarius, Vireo, 188, 209
solitary sandpiper, 201
solitudinis, Carpodacus mexicanus,
210
Sonchus arvensis, 136
Song sparrow, 212
Merrill, 212
Mountain, 212
Sonora episcopa, 109
Sonoran skink, 106
sonoriensis,
Eumops, 446
Peromyscus, 461
sora, 168, 200
Sorex sauasurei, 439
soricina, Glossophaga, 440
southern pocket gopher, 451
spadefoot, 103
sparrow,
English, 210
Brewer, 212
chipping, 212
fox. 212
Gambel, 212
grasshopper, 168
Harris, 212
lark, 211
Lincoln, 212
Merrill song, 212
montane Lincoln, 212
mountain song, 212
Nevada savannah, 211
northern sage, 212
sage, 212
savannah, 211
slate-colored fox, 212
song, 212
tree, 212
western chipping, 212
white-crowned, 212
white-throated, 212
vesper, 211
sparrow hawk, 199
sparrow hawks, 164
sparverius, Falco, 199
Spatula clypeata, 197
speargrass, 133
speckled king snake, 109
Speotyto,
cunicularia, 203
hypugaea, 203
Sphyrapicus,
nuchalis, 204
thyroideus, 204
varius, 204
spicilegus, Peromyscus, 461
spike-flower, 135
Spilogale,
angustifrons, 447
gracilis, 447
spinifera, Amyda, 112, 119
Spinus,
pallidus, 211
pinus vagrans, 211
tristis pallidus, 211
vagrans, 211
spinoletta, Anthus, 208
spiny pocket mouse, 454
Spizella,
arborea ochracea, 212
arizonae, 212
breweri, 212
ochracea, 212
passerina, 212
sponsa, Aix, 197
spotted,
sandpiper, 201
skunk, 447
towhee, 211
Squatarola squatarola, 200
squatarola, Squatarola, 200
squirrel,
fox, 168
ground, 450
lesser tropical ground, 450
rock, 449
tropical ground, 450
stabularis, Eulaelaps, 142
Index to Volume 1
633
stansburyi, Thomomys botiae, 36
starling, 168, 209
steerei, Proechimys, 316
Steganopus tricolor, 184, 202
stcindachneri, Amyda, 119
Steigidopteryx,
ruficollis, 206
serripennis, 206
stellata, Gavia, 195
stelleri, Cyanocitta, 208
Stercorarius pomarinus, 202
Sterna,
forsteri, 202
hirundo, 202
Sternotherus odoratus, 111
stilt, black-necked, 201
stilt sandpiper, 201
strepera, Anas, 197
striatula, Haldea, 110
striatulus, Accipiter gentilis, 198
striatus, Accipiter, 198
Strix nebulosa nebulosa, 203
Stumella, 168
neglecta, 190, 210
Sturnus vulgaris, 168, 209
subcinctus, Sylvilagus floridanus, 468
subfiavus,
Pipistrellus, 584
Pipistrellus hispidus, 599
vespertilio, 599
subspeciation,
in kangaroo rat, Dipodomvs ordii,
475
in pocket gophers of Kansas, 219
Sula sula, 180
sula, Sula, 180
supernus, Thomomys bottae, 251
surinamensis, Chlidonias nigra, 202
Surnia,
caparoch, 203
ulula, 203
Swainson hawk, 19S
swainsoni,
Buteo, 198
Hylocichla ustulata, 188
swallow,
bank, 206
barn, 206
cliff. 164, 206
northern cliff, 206
rough-winged, 206
tree, 206
violet-green, 206
swan,
trumpeter, 196
whistling, 196
sweet clover, 133
swift,
Vaux, 204
white-throated, 204
sylvatica, Rana, 104
sylvaticus, Apodemus, 142
Sylvilagus,
cunicularis. 469
floridanus, 168, 468
restricuts, 469
subcinctus, 468
Synaptomys cooperi, 130
syspila, Lampropeltis triangulum, 109
tabida, Grus canadensis, 200
Tachycineta,
lepida, 206
thalassina, 206
Tadarida,
femorosacca, range of, 247
mexicana, 445
tadpoles of Bufo cognatus, 95
talpoides, Thomomys, 6, 138
tanager, western, 210
Tantilla,
gracilis, 110
nigriceps, 110
tantilla, 110
Tarascan,
Empire, 435
mouse, 463
tarascensis, Zygogeomys trichopus,
453
Trasco, raton. 463
Taraxacum officinale, 134
Taxidea, 447
Tayassu,
angulatus, 469
humeralis, 469
Taylor, Edward H., A new hylid frog
from eastern Mexico, 259
taylori,
Baiomys, 460
Castor canadensis, 410
Peromyscus, 460
Tayra, 437
teal,
blue-winged, 168, 197
cinnamon, 197
green-winged, 197
Telmatodytes,
palustris, 207
pulverius, 207
tenuicauda, Neotoma ferruginea, 467
tenuiceps, Heliscomys, 289
tenuirostris, Charadrius alexandrinus,
183
tenuis, Reithrodontomys fulvescens,
457
tenuissima, Sitta carolinensis, 207
tephrocotis, Leucosticte, 211
tergeminus, Sistrurus catenatus, 111
tern,
black, 202
Caspian, 202
common, 202
Forster, 202
634
University of Kansas Publs., Mus. Nat. Hist.
Terrapene,
ornata, 111
triunguis, 112
terrosus, Dipodomys ordii, 523
testaceous, Masticophis flagellum, 108
teter, Cathartes aura, 198
Testudo riggsi, 284
texanum, Ambystoma, 103
texanus, Colinus virginianus, 200
Texas bob-white, 200
texensis,
Chordeiles acutipennis, 186
Didelphis mesamericana, 439
thalassina, Tachycineta, 206
Thamnophis,
marciana, 99
parietalis, 110
proximus, 110
radix, 99, 110
sauritus, 110
sirtalis, 110
Thayer gull. 202
thayeri, Larus argentatus, 202
thick-billed red-wing, 210
Thomomys,
absonus, 71
albicaudatus, 37
alexandrae, 72
aureus, 61
aureiventris, 28
birdseyei, 63
bonnevillei, 41
bottae, 28
bulbivorus, 3
centralis, 44
clusius, 17
contractus, 50
convexus, 47
dissimilis, 60
fisheri, 60
fossor, 21
fulvus, 61
gracilis, 6
howelli, 57
kaibabensis, 23
lenis, 53
levidensis, 54
levis, 24
minimus, 32
moorei, 19
nesophilus, 34
nicholi, 66
ocius, 17
of Utah, 3
osgoodi, 56
oquirrhensis, 11
parowanensis, 22
perpallidus, 28
planirostris, 66
pullus, 251, 451
pygmacus, 14
quadratus, 6
ravus, 15
robustus, 30
rufescens, 3
sevieri, 45
stansburyi, 36
supernus, 251
talpoides, 6, 138
tivius, 49
tolucae, 252
trumbullensis, 65
uinta, 13
umbrinus, 3, 251, 451
umbrinus, new from Mexico, 251
virgineus, 64
wahwahensis, 58
wasatchensis, 8
thrasher,
brown, 168
sage, 208
threeawn, 132
three-toed woodpecker,
Alaska, 205
alpine, 205
arctic, 205
thrush,
Alaska hermit, 208
Audubon hermit, 20S
hermit, 208
northern varied, 208
olive-backed, 208
varied, 208
western olive-backed, 208
willow, 188, 208
thula, Leucophoyx, 196
thyroideus, Sphyrapicus, 204
thysanodes, Myotis thysanodes, 444
tigrinum, Ambystoma, 103
timida, Chrysemys, 272
titmouse, gray, 207
tivius, Thomomys bottae, 49
tlacuache, 438
raton, 439
toad,
American, 103
common tree, 104
Great Plains, 104
narrow-mouthed, 105
Rocky Mountain, 104
tolmiei, Oporornis, 209
toltecus, Reithrodontomys fulvescens,
458
tolucae, Thomomys umbrinus, 252
toothed turtle, 112
towhee,
arctic, 211
green-tailed, 211
Nevada, 211
spotted, 211
townsendi,
Dendroica, 209
Mvadestes, 208
Index to Volume 1
635
Townsend,
solitaire, 208
warbler, 209
Toxostoma rufum, 168
traillii, Empidonax, 187, 205
transversa, Natrix erythrogaster, 109
tree sparrow, western, 212
tree swallow, 206
treganzai, Ardea herodias, 196
triangulum, Lampropeltis, 109
trichas, Geothlypis, 189, 209
trichopus, Zygogeomys, 453
tricolor, Steganopus, 184, 202
tridactylus, Picoi'des, 205
tridecimlineatus, Citellus, 143
trifida, Ambrosia, 134
Trifolium, 136
Trifolius, 132
trinitatis,
Eehimys, 333
Proechimys, 316
Tringa,
cinnamomea, 201
solitaria, 201
Trionyx, 122
triseriata, Pseudacris nigrita, 104
tristis, Spinus, 211
Triturus,
louisianensis, 103
viridescens, 103
triunguis,
Amyda, 122 .
Tenapene, 112
Troglodytes,
aedon, 207
pacificus, 207
parkmanii, 207
pullus, 188
troglodytes, 188, 207
troglodytes, Troglodytes, 188. 207
tropical,
arrocera, rata, 464
pigmeo raton, 460
pigmy mouse, 460
rice rat, 464
tropicalis, Conepatus, 577
Tromdoclonion lineatum, 110
truei, Peromyscus, 462
trumbullensis, Thomomys bottae, 65
tuberosus, Helianthus, 135
tularemia, 479
Turdus,
caurinus, 208
migratorius, 168, 188, 208
nigrideus, 188
propinquus, 208
turkey vulture, 198
turtle,
box, 111
Carolina, 112
common musk, 111
common snapping, 111
extinct emydid from Lower Plio-
cene of Oklahoma, 269
mud, 111
musk, 111
ornate box, 111
painted, 112
scribe, 112
snapping, 111
soft-shelled, 112, 119
spineless soft-shelled, 112
spiny soft-shelled, 112
toothed, 112
yellow mud, 111
turtles, fossil from Kansas and Okla-
homa, 283
tuza,
de Michoacan, 452
llanera, 452
serrana, 451
tylorhinus, Platygeomys, 452
Tyrannus tyrannus, 205
tyrannus, Tyrannus, 205
tzitzihoa. Anas acuta, 197
uakui, 450, 451
Uikiiri, 438
uinta, Thomomys talpoides, 13
ulula, Surnia, 203
umbelloides, Bonasa umbellus, 199
umbellus, Bonasa, 199
umbrinus,
Thomomys, 251, 451
Thomomys talpoides, 3
underwoodi, Eumops, 445
undulatus, Sceloporus, 106
unicolor. Cinclus mexicanus, 207
United States Navy, office of research
and invention, 479
urichi, Proechimys, 316
Urocj'on,
cinereoargenteus, 448
colimensis, 448
urophasianus, Centrocercus, 200
ustulata, Hylocichla, 188, 208
Utah, The pocket gophers (genus
Thomomys) of, 1
utahensis,
Dipodomys ordii, 525, 543
Eremophila alpestris, 206
Perodipus montanus, 53S
Perodipus ordii, 543
vacillator, Proechimys, 316
vagrans, Spinus pinus, 211
valisineria, Aythya, 197
vampire bat, 443
vampiro, 443
variabilis, Dermacentor, 143
varied thrush, northern, 208
variegatus,
Citellus variegatus, 449
Sciurus, 449
636
University of Kansas Publs., Mtjs. Nat. Hist.
varius,
Cartogeomys, 453
Platygeomys, 453
Sphyrapicus, 204
Vaux swift, 204
vauxi, Chaetura, 204
velifer, Myotis, 443, 588
velox. Accipiter striatus, 198
venado cola blanca, 470
ventralis, Ophisaurus, 106
veracrucis,
Pipistrellus subflavus, 601
Vesperugo, 601
vermis, Carphophis amoena, 107
Vermivora,
celata, 209
orestera, 209
ridgwayi, 209
ruficapilla, 209
vermivorus, Helmitheros, 189
versicolor,
Hyla, 104
Quiscalus, 168
vesper sparrow, 211
Vespertilio cinnamoraeus, 585
vespertina, Hesperiphona, 210
vespertino, murcielago, 443
Vesperugo,
carolinensis, 599
erythrodactylus, 599
hesperus, 596
merriami, 597
subflavus, 599
veracrucis, 601
vicinior, Peromyscus banderanus, 463
Villa R., Bernardo, 219, 251
Villa R., Bernardo and E. Raymond
Hall, 435
villicauda, Proechimys, 355
villosus, Dendrocopos, 204
vinaceus, Geomys bursarius, 234
violet-green swallow, 206
virens, Icteria, 189, 209
Vireo,
alticola, 188
cassinii, 209
gilvus, 209
huttoni, 209
olivaceus, 209
solitarius, 1S8, 209
swain son ii, 209
vireo,
I lassin, 209
Hutton, 209
mountain, 188
red-eyed, 209
warbling, 209
western warbling, 209
virgineus, Thomomys bottae, 64
Virginia rail, 200
virginiana, Didelphis, 439
virginianus,
Bubo, 159, 203
Colinus, 200
Odocoileus, 470
viridescens, Triturus, 103
viridis, Setaria, 132
viridus, Crotalus, 111
vittata, Mephitis macroura, 578
vociferus, Charadrius, 200
volcanes, raton los, 466
vulgaris, Sturnus, 168, 209
vulture,
turkey, 198
western turkey, 198
wahwahensis, Thomomys bottae, 58
wapacuthu, Bubo virginianus, 203
warbler,
Audubon, 209
black-throated gray, 209
Calaveras, 209
Macgillivray, 209
northern pileolated, 210
orange-crowned, 209
pileolated, 210
Rocky Mountain orange-crowned,
209
Townsend, 209
yellow, 209
warbling vireo, 209
warreni, Proechimys, 316
wasatchensis, Thomomys talpoides, 8
washingtoniensis, Haliaeetus
leucocephalus, 199
water snake, 109
water-thrush,
British Columbia, 189
Grinnell, 209
northern, 189
waxwing,
Bohemian, 208
cedar, 208
weasel, 447
western,
belted kingfisher, 204
bluebird, 208
burrowing owl, 203
catbird, 207
chipping sparrow, 212
crow, 206
gnatcatcher, 208
golden-crowned kinglet, 208
goshawk. 198
grebe, 196
grosbeak, 210
harlequin duck, 198
house wren, 207
lark sparrow, 211
Index to Volume 1
637
western — Concluded
meadowlark, 190, 210
mourning dove, 202
olive-backed thrush, 208
pigeon hawk. 199
pileated woodpecker, 204
pine siskin, 211
pipit, 188, 208
red-tailed hawk, 198
ruby-crowned kinglet, 208
sandpiper, 201
solitary, 201
spiny pocket mouse, 454
tanager, 210
tree sparrow, 212
turkey vulture, 198
vesper sparrow, 211
warbling vireo, 209
wenmanni, Epitedia, 140
white pelican, 196
white-cheeked goose, 196
white-crowned sparrow, 212
white-faced glossy ibis, 196
white-headed woodpecker. 205
white-sided jack rabbit, 469
white-tailed,
deer, 470
ptarmigan, 200
white-throated,
sparrow, 212
swift, 204
wood rat, 467 •
white-winged crossbill. 211
wild,
lettuce, 133
millet, 134
morning glory, 135
willet, western, 201
Williamson sapsucker, 204
willow thrush, 188, 208
Wilson,
phalarope, 184, 202
plover, 184
snipe, 201
warbler, 190
Wilsonia,
pileolata, 210
pusilla, 190, 210
wilsonia, Charadrius, 184
wilsonianus, Asio otus, 203
winter wren, 207
wood duck, 197
wood pewee, 205
wood rat,
dwarf, 467
ferruginous, 467
white-throated, 467
Woodhouse jay, 206
woodhousei, Aphelocoma coerulescens,
206
woodhousii. Bufo, 104
woodpecker,
Alaska three-toed, 205
alpine three-toed, 205
arctic three-toed, 205
Batchelder, 205
hairy, 204
Lewis, 204
northern white-headed, 205
pileated, 204
Rocky Mountain hairy, 204
three-toed. 205
western pileated, 204
white-headed, 204
wood mouse, 462
worm-eating warbler, 189
worm snake, 107
wren,
canyon, 207
common rock, 207
house, 207
long-billed, 207
northern canyon, 207
northwestern long-billed, 207
rock, 207
southern winter, 188
western house, 207
western winter, 207
winter, 188, 207
yellow warbler, 209
vellow-bellied,
flycatcher, 186
king snake, 108
yellow-breasted chat, 189
yellow-headed blackbird, 210
yellow-legs,
greater, 201
lesser, 201
yellow-throat, western, 189, 209
wrbabuenae, Lepton3'cteris nivalis,
441
Yuma,
murcielago, 443
myotis, 443
yumanensis. Myotis, 443
zacatecae, Reithiodontomys mega-
lotis, 456
zamorae, Peromvscus melanophrys,
463
zapotero, murcielago, 442
zelotes, Peromyscus, 462
Zenaidura,
macroura, 168, 202
marginella, 202
638
University of Kansas Publs., Mus. Nat. Hist.
Zonotrichia,
albicollis, 212
gambeli, 212
leucophrys, 212
querula, 212
zorrillo, 448
manchado, 447
real, 448
zorro gris, 448
Zygogeomys,
tarascensis, 453
trichopus, 453
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