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    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


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 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.

  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.

  [Illustration: 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.]

  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 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.

  _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 Elkader, 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, December 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 Zoöl. 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 averaging 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
   distance from alveolus of upper incisor to middle of lateral
   border of P^4 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 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
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.

  _Specimens examined._--Total number, 148, as follows: _Clay
  County_: 6 mi. SW Clay Center, 3. _Jackson County_: 10-1/2 mi.
  WSW Holton, 1; no locality more precise than county, 1.
  _Atchison County_: 3 mi. N Cummings, 2. _Jefferson County_:
  Oskaloosa, 1. _Leavenworth County_: Fort Leavenworth
  (Government Hill, 2; Engineer Hill, 1), 6; no locality more
  precise
  than county, 19. _Saline County_: Salina, Debold Farm, 4 (coll.
  of A. J. Kirn). _Morris County_: 1-1/2 mi. N Council Grove, 3.
  _Douglas County_: 1 mi. NW Midland, 2; 1 mi. N Lawrence, 1;
  2-1/2 mi. W Lawrence, 2; 1 mi. W K. U. Campus, 2; 1 mi. W
  Lawrence, 2; 1/2 mi. W Lawrence, 2; "W K. U. Campus," 2; K. U.
  Campus, 4; Lawrence, 23; South Lawrence, 1; 1/2 mi. SW K. U.
  Campus, 2; Southwest K. U. Campus, 1; Haskell Institute, 1;
  4-1/2 mi. S Lawrence, 1; 7 mi. SW Lawrence, 6; 7-1/2 mi. SW
  Lawrence, 1; 8 mi. SW Lawrence, 1; 10 mi. S Lawrence, 1; 11 mi.
  SW Lawrence, 3; no locality more precise than county, 15.
  _Marion County_: 1-1/2 mi. NE Lincolnville, 6; 4 mi. SE
  Lincolnville, 1; 6 mi. S Lincolnville, 1. _Greenwood County_:
  Hamilton, 1; 1/2 mi. S Hamilton, 4; 1 mi. S Hamilton, 4; 4 mi.
  S and 14 mi. W Hamilton, 6; 8 mi. SW Toronto, 1; 8-1/2 mi. SW
  Toronto, 5; no locality more precise than county, 6.


    =Geomys bursarius jugossicularis= Hooper

    _Geomys lutescens jugossicularis_ Hooper, Occas. Papers Mus.
    Zoöl., 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 certainly to Hamilton County and south certainly to
    Morton and Seward counties.

    _Description._--A yellowish-cinnamon colored animal, with body
    of medium size, zygomatic 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. 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 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.

  [Illustration: FIG. 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.]

  _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
  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 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_.

  _Specimens examined._--Total number, 58, distributed as
  follows: _Pawnee County_: Jct. Pawnee and Arkansas rivers,
  Larned, 6; 1 mi. S and 1 mi. E Larned, 7. _Edwards County_: 1
  mi. W and 3-1/2 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_: 3-1/2 mi. NE
  Fowler, 2; 2 mi. N Fowler, 2; 1-1/2 mi. N Fowler, 2; 1-1/4 mi.
  N and 3/4 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 lutescens 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 llanensis_, 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 Ellsworth County, westward certainly to Stafford
  and Barber counties and eastward 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
  defined 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 alveolus of incisor
  to middle of lateral border of P^4 at alveolar border. Length
  of auditory bulla (from anteroventral edge of paroccipital
  process of exoccipital 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 twice 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 majusculus_, _major_ differs in slightly lighter
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._
_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_.

  _Specimens examined._--Total number, 77, as follows: _Ellsworth
  County_: 2 mi. S Ellsworth, 1. _McPherson County_: Smoky Hill
  River, 1 mi. S and 1/2 mi. W Lindsborg, 5; 1/2 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 1/2 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_: 4-1/2
  mi. NE Danville, 8; 1 mi. N Harper, 11; 3 mi. S Harper, 1.
  _Cowley County_: 3 mi. SW Arkansas City, 4; 3 mi. SE Arkansas
  City, 9; 3 mi. S Arkansas City, 1.




    MEASUREMENTS OF ADULT MALES OF GEOMYS


    (In millimeters)

    Key for table headings in table on this page.

    N:  Number of individuals averaged or catalogue number
    L:  Total length
    T:  Length of tail
    H:  Length of hind foot
    B:  Basilar length
    Na: Length of nasals
    Z:  Zygomatic breadth
    M:  Mastoid breadth
    Rb: Breadth of rostrum
    I:  Interorbital constriction
    A:  Alveolar length of maxillary tooth row
    E:  Extension of premaxilla posterior to nasals
    S:  Depth of skull
    Rl: Length of rostrum

    ======+===+====+====+====+====+====+====+====+===+===+===+====+=====
      N   |L  | T  | H  | B  | Na | Z  | M  | Rb | I | A | E | S  | Rl
    ------+---+----+----+----+----+----+----+----+---+---+---+----+-----
                         _G. b. lutescens_; topotypes
    5 ave.|266|82.0|34.2|40.0|17.7|30.5|26.8|11.5|6.7|8.6|3.9|17.1|20.8
      min.|257|76.0|33.0|38.3|16.0|29.1|26.1|11.2|6.3|8.1|3.5|16.2|19.1
      max.|276|91.0|36.0|42.4|20.3|31.7|27.5|11.9|6.9|9.2|4.2|17.7|23.6

                     2 mi. NE Ludell, Rawlins Co., Kansas
     12088|272|92.0|35.0|43.2|19.1|32.3|27.7|11.3|6.6|8.4|2.8|18.0|22.1

                   _G. b. majusculus_; Douglas Co., Kansas
          |   |    |    |    |    |[A] |    |    |   |   |   |    |
  16 ave. |289|79.8|36.3|47.1|21.0|34.1|30.4|12.1|6.8|9.3|3.7|18.5|24.9
     min. |273|70.0|32.0|44.7|18.9|30.5|27.5|11.1|6.5|8.2|2.9|17.3|22.9
     max. |308|95.0|55.0|49.9|23.2|38.0|34.5|13.5|7.6|10.3|5.7|20.0|28.1

                  _G. b. jugossicularis_; Morton Co., Kansas
    4 ave.|265|82.0|34.2|40.7|16.9|30.0|27.9|10.7|6.0|8.6|5.2|17.3|21.2
      min.|250|68.0|30.0|38.5|16.1|29.0|27.5|10.5|5.5|8.2|4.7|16.4|20.2
      max.|285|92.0|37.0|42.4|17.4|31.1|28.4|11.0|6.2|9.2|5.5|17.9|22.0

                    _G. b. industrius_; Meade Co., Kansas
    8 ave.|265|82.0|35.0|40.9|18.1|30.0|28.0|11.0|6.2|8.8|4.3|17.7|21.8
      min.|247|70.0|33.0|37.9|15.5|28.2|26.5| 9.9|5.7|8.0|2.9|16.8|19.5
      max.|280|90.0|36.0|43.4|21.0|32.4|29.5|11.6|7.0|9.1|5.2|19.1|24.2

             _G. b. major_; Wells Ranch, Aetna, Barber Co., Kansas
     11724|256|66.0|34.0|41.0|18.3|31.6|28.2|10.6|6.1|9.0|4.0|17.0|21.3

                      1 mi. W Aetna, Barber Co., Kansas
     11153|240|75.0|32.0|36.7|15.7|26.9|24.6| 9.9|5.9|8.8|4.0|15.0|19.5
     11152|240|65.0|32.0|36.0|14.2|26.1|25.4|10.9|5.6|8.5|5.0|15.5|18.5

                  3 mi. SE Arkansas City, Cowley Co., Kansas
          |   |    |    |    |[E] |    |    |    |   |   |   |    |
     12870|246|76.0|32.0|42.1|16.0|33.7|29.7|11.5|6.3|9.4|4.5|17.6|21.3

                  3 mi. SW Arkansas City, Cowley Co., Kansas
     12892|282|84.0|33.0|41.7|17.3|....|27.7|10.8|6.4|8.9|4.2|17.2|21.5
    ------+---+----+----+----+----+----+----+----+---+---+---+----+-----




    MEASUREMENTS OF ADULT FEMALES OF GEOMYS


    (In millimeters)

    Key for table headings in table on this page.

    N:  Number of individuals averaged or catalogue number
    L:  Total length
    T:  Length of tail
    H:  Length of hind foot
    B:  Basilar length
    Na: Length of nasals
    Z:  Zygomatic breadth
    M:  Mastoid breadth
    Rb: Breadth of rostrum
    I:  Interorbital constriction
    A:  Alveolar length of maxillary tooth row
    E:  Extension of premaxilla posterior to nasals
    S:  Depth of skull
    Rl: Length of rostrum

    ======+===+====+====+====+====+====+====+====+===+===+===+====+=====
      N   | L | T  | H  | B  | Na | Z  | M  | Rb | I | A | E | S  | Rl
    ------+---+----+----+----+----+----+----+----+---+---+---+----+-----
                              _G. b. lutescens_; topotypes
    6 ave.|233|72.3|31.1|35.3|15.0|25.9|23.7|10.4|6.1|8.3|3.7|15.4|18.4
      min.|215|63.0|30.0|33.5|13.9|24.6|21.8|10.1|5.6|8.1|2.9|14.8|17.3
      max.|254|76.0|32.0|37.0|16.8|26.7|24.8|10.7|6.6|8.5|4.5|16.2|19.8

                          2 mi. NE Ludell, Rawlins Co., Kansas
     11733|230|63.0|31.0|35.3|15.1|26.5|24.1| 9.3|6.1|7.5|2.4|15.0|18.2
     12155|245|70.0|30.0|35.6|14.6|25.2|24.1|10.6|6.4|7.5|3.1|14.9|18.2

                        _G. b. majusculus_; Douglas Co., Kansas
          |   |    |    |[B] |[B] |[A] |    |    |   |   |   |    |
  17 ave. |265|78.6|32.8|40.6|17.2|28.6|26.4|10.9|6.5|9.1|3.6|16.6|21.0
     min. |222|59.0|30.0|37.1|15.9|26.7|24.9|10.0|5.9|8.5|2.0|15.2|18.8
     max. |304|92.0|35.0|47.0|20.1|33.4|29.1|12.3|7.3|10.0|5.9|19.1|24.1

                       _G. b. jugossicularis_; Morton Co., Kansas
      5012|244|72.0|30.0|36.2|16.4|25.4|25.0|10.0|5.9|8.0|4.2|16.0|19.3
      5395|230|72.0|30.0|34.6|13.9|24.7|24.8| 9.8|5.8|8.0|4.5|15.2|17.5

                         _G. b. industrius_; Meade Co., Kansas
          |   |[C] |    |[D] |    |    |[D] |
   7 ave. |238|73.0|31.3|36.4|14.9|26.3|24.8|10.0|6.0|8.4|4.1|16.2|18.6
     min. |231|65.0|30.0|35.4|14.0|25.8|24.5| 9.5|5.6|8.1|3.6|15.5|17.5
     max. |256|75.0|32.0|37.8|16.1|27.8|25.9|10.3|6.5|8.7|4.7|17.6|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|9.0|3.4|16.4|19.4

                               Aetna, Barber Co., Kansas
     10070|242|83.0|30.0|36.8|15.7|26.2|25.0|10.1|6.5|9.1|3.3|15.8|19.1

                         Wells Ranch, Aetna, Barber Co., Kansas
     12238|239|65.0|31.0|34.2|14.5|24.6|23.7| 9.6|6.0|8.0|3.6|15.2|17.7

                          1 mi. S.Sun City, Barber Co., Kansas
     11075|232|66.0|28.0|34.2|14.4|25.0|23.6| 9.9|5.9|8.0|3.4|15.0|17.0

                       3 mi. SW Arkansas City, Cowley Co., Kansas
     12872|242|66.0|30.0|38.1|15.0|28.0|26.2|10.3|6.3|7.8|4.5|16.1|19.1

                       3 mi. SE Arkansas City, Cowley Co., Kansas
     12894|230|82.0|30.0|38.5|15.5|28.0|25.6|10.0|6.7|8.7|4.0|16.6|19.5
     12893|246|83.0|32.0|36.5|14.2|25.6|24.8| 9.6|6.6|8.7|4.6|15.4|18.1
    ------+---+----+----+----+----+----+----+----+---+---+---+----+-----

[A] 15 averaged.

[B] 16 averaged.

[C] 6 averaged.

[D] 5 averaged.

[E] approximate.




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.




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.

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




Transcriber Notes:


Minor typographical errors were corrected without notice.

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