Subspeciation in Pocket Gophers of Kansas

By
BERNARDO VILLA-R. and E. RAYMOND HALL

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
FRED VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
  1947

21-8188

Subspeciation in Pocket Gophers of Kansas

By

BERNARDO VILLA-R. AND E. RAYMOND HALL

[219]
Several full species of the genus Geomys have been recorded from
Kansas. The purpose of the study now reported upon was to determine
the present taxonomic status of these animals and the distribution
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 gophers
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° 30′.” He reported this animal as common
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) recognized
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 between
[220]
every pair of kinds having contiguous geographic ranges. The
characters previously thought by some writers constantly to differentiate,
say, Geomys lutescens of western Kansas from Geomys bursarius
of eastern Kansas, prove not to do so; instead, in areas geographically
intermediate between the geographic ranges of the two
kinds, the pocket gophers are intermediate in morphological characters
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 structural
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 restricted
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 different
kinds of pocket gophers to use only one age class of one season
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 magnum
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.

[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 border
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 after
Ridgway, Color Standards and Color Nomenclature, Washington, D. C.,
1912.

Fig. 1. Map showing the geographic distribution of the five
subspecies of
the Missipi Valley pocket gopher, Geomys bursarius, in Kansas, with insert
showing range of the species.

[222]

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 grateful
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 Somerset,
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, October
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. Zoöl., 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 Nebraska.

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 specimens;
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 subspecies
occurring in Kansas.

Remarks.—In his monographic revision of the pocket gophers,
Merriam (1895:129) recorded 3 “typical or nearly typical” specimens
from Trego County, and 18 “non typical” specimens as follows:
Garden Plain, Sedgwick County, 4; Belle Plain, Sumner[223]
County, 5; Cairo, Pratt County, 6; Kiowa, Barber 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 remarkably
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.

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 subspecies)
extend equally far posteriorly to nasals; diastema longer in
relation to basilar length; rostrum longer relative to its width; sagittal
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 coloration
and in two cranial characters as follows: Nasals straight-sided
instead of shaped like an hour-glass, and superficial canals on palatine
extending anteriorly beyond first molar, and from there anteriorly
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 exposure
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 premaxillae
do not extend so far posteriorly to nasals in either males
or females; interorbital constriction slightly narrower in adult females;
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 differentiated[225]
the animals from southeastern South Dakota, the eastern
parts of Nebraska and Kansas, and the western and southern parts
of Iowa, from G. bursarius bursarius to which he assigned a more
northern geographic range. In the absence of comparative materials
of the northern subspecies we cannot make an independent decision
on the validity of majusculus and recognize that if it is inseparable
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 measurements
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 maximum
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 Hamilton
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. majusculus
on the one hand and Geomys bursarius lutescens and Geomys
bursarius major on the other hand. In color they agree with majusculus,
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 anterodorsally,
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.

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. jugossicularis;
nevertheless, one specimen from Morton County has the occiput
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. Zoöl.,
Univ. Michigan, 420:1, June 28, 1940.

Type.—Male, adult, skin and skull, no. 14083 Museum of Natural History,
University of Kansas; from 1-1/2 miles north of Fowler, Meade County, Kansas;
obtained December 30, 1941, by H. H. Hildebrand, original number 16.

Distribution in Kansas.—Southwestern Kansas from Meade County eastward[227]
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.

Fig. 2 Three views of the skull of the type specimen of Geomys bursarius industrius.

A. Lateral view;

B. Dorsal view;

C. Ventral view.

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 anteriorly;
forefeet white; hind feet and approximately distal half of tail whitish.
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 border
of alveolus of first upper molar, occiput strongly inclined anterodorsally, temporal
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 border
of alveolus of first upper molar (not premolar); occiput less inclined anterodorsally[228]
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 upper
parts and underparts. Skull: Jugular part of zygomatic arch
more curved (convex dorsally) and occiput far more inclined anterodorsally;
lower part of mastoidal ridge more prominent.

For comparison with G. l. 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 counties
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 subspecies
with adjoining geographic ranges. The existence of this
unique (among adjoining subspecies) cranial character is the principal
reason for according subspecific status to this animal. Although
it has other characters which are fairly uniform over a
considerable geographic area, these other characters, namely, Cinnamon
Brown color of the upper parts and medium size of the body,
after all, are conditions intermediate between those in jugossicularis
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 indicative
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” subspecies
as compared with any one of the adjoining subspecies.

Hooper (1940:2) in naming as new Geomys lutescens jugossicularis
referred to his new subspecies a skin-only from Meade County[229]
State Park. Our more abundant material from there shows the
cranial conformation to be that of industrius to which we accordingly
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 indicates
intergradation between the two named subspecies. Specimens
from Pratt County are slightly darker than industrius thereby
indicating intergradation between industrius and major.

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 majusculus, major differs in slightly lighter[230]
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 flat, 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.[231]
majusculus; the underparts are only slightly paler than the upper
parts as in majusculus. Measurements of the skulls are intermediate
between the averages for G. b. majusculus and those for G. b.
major. These specimens from eight miles west of Rosalia are intermediate
structurally, and since they are intermediate geographically
between G. b. majusculus and G. b. major, they suggest intergradation
of the two subspecies. The specimens in question are referred
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 McPherson
County, Kansas, that have caused us to treat G. b. majusculus
as only subspecifically distinct from the more western subspecies,
major.

Measurements of Adult Males of Geomys

(In millimeters)
[232]

[233]

Measurements of Adult Females of Geomys

(In millimeters)

Footnotes

SUBSPECIES OF THE SPECIES GEOMYS BURSARIUS

If Geomys lutescens major Davis is correctly judged to intergrade
with Geomys bursarius 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 Upper
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 1-1/2 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.[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):l-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, pls. 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. Zoöl., 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. Mamm., 12:404-408,
1 pl., 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]

Merriam, C. H.

1890. Descriptions of twenty-six new species of North American mammals.
N. Amer. Fauna, 4: v + 60, 3 pls., 3 figs. in text. October 8, 1890.

1895. Monographic revision of the pocket gopher Family Geomyidae….
N. Amer. Fauna, 8:1-258, 19 pls. and frontispiece, 71 figs. in text, 4
maps. January 31, 1895.

Scheffer, T. H.

1910. The pocket gopher. Kansas State Agric. Coll. Ent. and Zoöl. Dept.,
Bull., 172:197-233, illustrated. September, 1910.

1931. Habits and economic status of the pocket gophers. U. S. Dept. Agric.,
Tech. Bull., 224:1-27, 8 pls., 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. Missouri
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 distribution
of the variants. Missouri Valley Fauna, 2:1-12. February 1,
1940.

Transmitted May 30, 1947.

PRINTED BY

FRED VOILAND, JR., STATE PRINTER

TOPEKA, KANSAS

1947

21-8188