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Title: Comments on the Taxonomy and Geographic Distribution of Some North American Rodents
Author: Hall, E. Raymond (Eugene Raymond), 1902-1986, Kelson, Keith R.
Language: English
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Copyright Status: Not copyrighted in the United States. If you live elsewhere check the laws of your country before downloading this ebook. See comments about copyright issues at end of book.

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Comments on the Taxonomy and Geographic Distribution of Some North
American Rodents



University of Kansas Publications
Museum of Natural History

Volume 5, No. 26, pp. 343-371
December 15, 1952



Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Edward H. Taylor, Robert W. Wilson

Volume 5, No. 26, pp. 343-371
December 15, 1952

Lawrence, Kansas


[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies
words in bold. Words surrounded by underscores, like _this_, signifies
words in italics.]

Comments on the Taxonomy and Geographic Distribution of Some North
American Rodents



In preparing maps showing the geographic distribution of North American
mammals we have found in the literature conflicting statements
concerning the subspecific identity of several rodents. Wherever
possible, we have examined the pertinent specimens. Results of our
examination are given below.

Our studies have been aided by a contract (NR 161-791) between the
Office of Naval Research, Department of the Navy, and the University of
Kansas. Also, a grant from the Kansas University Endowment Association
has permitted field work that yielded some of the specimens used for
comparison. Grateful acknowledgment is made to the persons in charge of
the several collections of mammals that we have consulted in order to
satisfy ourselves concerning the subspecific status of specimens from
many localities.

~Marmota flaviventer luteola~ A. H. Howell

A. H. Howell (N. Amer. Fauna, 37:50, April 7, 1915) referred specimens
from Bridgers Pass, Wyoming, to _Marmota flaviventer dacota_, on the
basis of paler underparts because, according to the data of Howell (_op.
cit._), _M. f. dacota_ and _M. f. luteola_, the contiguous subspecies,
do not differ significantly in other ways. Casual comparison reveals to
us no additional differences between the two. We have examined the three
specimens available to Howell from Bridgers Pass (Nos. 18733/25527,
18734/25528, and 18735/25529 U. S. Biol. Surv. Coll.) and find the tone
of the underparts to be darker (more nearly russet) than in typical
_luteola_. The tone, however, varies considerably, both individually and
geographically, in _luteola_ and it is possible to match almost exactly
the ventral coloration of the specimens from Bridgers Pass with that of
specimens from within the geographic range of _luteola_; Nos. 160509,
from Bear Creek, 8 miles west of Eagle Peak, Wyoming, 18875 and
18731/25535, from the Laramie Mts., Wyoming, and No. 203744 from Sulphur
Springs, Grand County, Colorado, all in the United States Biological
Surveys Collection, are examples to the point. Being influenced by the
geography of the region, we therefore consider the three specimens from
Bridgers Pass best referred to the subspecies _Marmota flaviventer

~Spermophilus variegatus grammurus~ (Say)

A. H. Howell (N. Amer. Fauna, 56:147, May 18, 1938) accorded _Citellus_
[= _Spermophilus_] _variegatus utah_ Merriam a geographic range that
included the Kaibab Plateau of Arizona. Durrant (Univ. Kansas Publ. Mus.
Nat. Hist., 6:119, August 10, 1952) assigned to _S. v. grammurus_ a
geographic range that included southern Utah from the eastern to the
western border but in doing this did not mention the rock squirrel of
the Kaibab Plateau of Arizona that also might be expected to be
referable to _S. v. grammurus_. Howell (_loc. cit._) had two specimens
from the Kaibab Plateau. Of these we have examined the one from Big
Spring (161566 BS) and find that it lacks the darker (more tawny) head
and posterior back of _C. v. utah_ and agrees with _C. v. grammurus_. On
this basis we refer the rock squirrel of the Kaibab Plateau to the
subspecies _Spermophilus variegatus grammurus_ (Say).

~Tamias amoenus caurinus~ Merriam

This subspecies was named from the Olympic Peninsula of Washington. A.
H. Howell, in his "Revision of the American chipmunks" (N. Amer. Fauna,
52:77, and fig. 5, 1929) regarded the geographic range of _Eutamias_ [=
_Tamias_] _amoenus caurinus_ as the mountains of the Olympic Peninsula
and most of Mt. Rainier. The geographic range of the _amoenus_ chipmunk
on Mt. Rainier almost certainly is continuous with that of _T. a.
ludibundus_ in the Cascade Mountains of which Mt. Rainier is a
westward-projecting arm. There is no contact between the chipmunks of
Mt. Rainier and those of the Olympic Peninsula; those on the Peninsula
are geographically isolated from all others of the species and are
separated from those on Mt. Rainier by approximately eighty miles of
low-lying country, which is uninhabited by chipmunks of the species
_Tamias amoenus_. Therefore, Howell's (_loc. cit._) assignment of most
of the chipmunks on Mt. Rainier to _caurinus_ is open to question and
Dalquest, in the "Mammals of Washington" (Univ. Kansas Publ. Mus. Nat.
Hist., vol. 2, 1948) evidently thought that Howell had incorrectly
identified them. On page 256 Dalquest (_op. cit._) defined the
geographic range of _T. a. caurinus_ as restricted to the Olympic
Peninsula and showed (fig. 81) Mt. Rainier to be in the geographic range
of _T. a. ludibundus_. We would accept Dalquest's (_op. cit._)
arrangement without question and also would follow it because it is the
more recent one were it not for the fact that Dalquest gives no reason
for his changes. To allow us to decide the matter we have compared the
pertinent materials ourselves. Catalogue numbers below are of the United
States National Museum, Biological Surveys Collection, and each specimen
mentioned by catalogue number is an adult female which shows much wear
on the fourth upper premolar.

Of _T. a. caurinus_, Nos. 241902 and 241903 are from 2 mi. SW of Mount
Angeles; No. 241911 is from "near" head of Dosewallips River, 6000 ft.,
and No. 241915 is from Canyon Creek, 3 mi. S Soleduc River, 3550 ft. Of
_T. a. ludibundus_, Nos. 234776 and 235018 are from Barron, 5000 ft.,
and No. 230685 is from Suiattle River, 6500 ft. Of specimens in
question, from Mount Rainier, No. 90635 is from 6500 ft., west slope;
No. 232729 is from 4900 ft., Reflection Lakes, and No. 233114 is from
5300 ft., Indian Henrys.

In comparison with _T. a. ludibundus_, _T. a. caurinus_ is grayer on
most, or all, parts of the pelage, has less ochraceous on the sides, and
the dark stripes on the sides of the head are narrower and less reddish
(more grayish). The skull of _caurinus_ is larger in certain
measurements, as shown below:

Catalogue  Occipitonasal  Zygomatic  Cranial  Length of  Greatest width
number     length         breadth    breadth  nasals     across upper
_T. a. ludibundus_

234776        34.0        19.3        15.6        10.2        ...
235018        34.1        ....        ....        10.4        8.0
230685        33.5        18.8        15.5        10.4        7.9

Mt. Rainier

 90635        34.5        19.2        16.3        10.8        8.3
232729        ....        18.5        15.3        ....        8.2
233114        34.2        18.6        15.7        10.8        8.0

_T. a. caurinus_

241911        34.5        19.7        16.2        11.3        8.3
241915        34.2        ....        ....        10.3        8.3
241902        35.2        ....        16.8        11.1        8.1
241903        34.7        ....        16.0        10.8        8.4

Howell (_op. cit._:75) referred three specimens from Glacier Basin, on
the northeastern part of Mount Rainier, to _T. a. ludibundus_ as he did
also one specimen (_loc. cit._) from Reflection Lakes, on the southern
flank of the mountain. Our comparisons indicate the correctness of
Howell's identification of the specimens from Glacier Basin; they more
closely resemble _ludibundus_ than _caurinus_. The specimen from
Reflection Lakes, however, is only one of five or six from the same
place; the others were lumped by him among the 49 that he recorded from
Mount Rainier under the name _caurinus_. The series from Reflection
Lakes, so far as we can detect, is not unusually variable and the
differences that are apparent are within the normal range of variation
ascribable to season, age, and individualism. Also, the series from
Reflection Lakes, to us, is not appreciably different from the other
series, representing the following places on Mount Rainier: Indian
Henrys, 5300 ft.; W slope Mt. Rainier, 6600 ft.; St. Andrews Park, 5500
ft.; Spray Park, 5500 ft.; Paradise Park; Muddy Fork of Cowlitz River;
Sunset Park, 5000 ft.; ridge between St. Andrews Park and South Puyallup
River, 6000 ft.; and Owyhigh Lakes, 5350 ft.

Collectively, or individually, where there are as many as six specimens
from a place, the material from Mt. Rainier (Glacier Basin excepted) is
intermediate in color between _T. a. ludibundus_ and _T. a. caurinus_
and no more closely resembles one subspecies than the other. As may be
seen from the cranial measurements recorded above, specimens from Mt.
Rainier, although intermediate between the two subspecies just
mentioned, resemble _ludibundus_ in lesser zygomatic breadth and lesser
cranial breadth (and, it may be added, in lesser dorsolateral inflation
of the braincase), but resemble _caurinus_ in longer skull
(occipitonasal length), longer nasals and greater breadth across the
rows of upper molariform teeth.

In summary: The animals from Mount Rainier, in features of taxonomic
import, are almost exactly intermediate between _T. a. caurinus_ and _T.
a. ludibundus_. Being influenced by considerations of geographic
adjacency, we refer the animals on Mount Rainier to _Tamias amoenus
ludibundus_ (Hollister).

Dalquest's (_op. cit._: 85) explanation of the probable origin of
_Tamias amoenus caurinus_ is pertinent here. He writes: "The chipmunks
of the Olympic Mountains [_caurinus_] probably reached their present
range from the Cascades. Their probable path of emigration was westward
from Mt. Rainier, along the glacial outwash train of Nisqualli Glacier,
to the moraine and outwash apron of the Vashon Glacier and thence to the
Olympics. So similar are the chipmunks of Mt. Rainier and the Olympic
Mountains that Howell (1929) included Mt. Rainier in the range of

~Tamias townsendii cooperi~ Baird

Some uncertainty exists concerning the subspecific identity of the
Townsend Chipmunk in southern Washington because Dalquest (Univ. Kansas
Publ. Mus. Nat. Hist., 2:262, April 9, 1948) identified as _Tamias
townsendii cooperi_ specimens that he examined from Yocolt, a place well
within the geographic range of _T. t. townsendii_ as defined by A. H.
Howell (N. Amer. Fauna, 52: fig. 7, p. 107, November 30, 1929). Dalquest
(_op. cit._) referred other specimens, that he did not examine, from Mt.
St. Helens (90654, 231112 and 231114 BS) to _T. t. cooperi_ although
Howell (N. Amer. Fauna, 52:109, November 20, 1929) had previously
identified them as _E. t. townsendii_. By implication, and on his map,
Dalquest (_op. cit._, fig. 83, p. 261) assigned to _T. t. cooperi_ still
other specimens, that he had not examined, from: Government Springs, 15
mi. N Carson (230514, 230515, 230559, 230560, and 230563 BS); Stevenson
(230513 and 230517 BS); and Skamania (230518 BS). Earlier, Howell (op.
cit.) had listed the specimens from the three mentioned localities as
_Eutamias townsendii townsendii_.

Our examination of specimens in the Museum of Vertebrate Zoology from
1-1/2 mi. W Yocolt (94238 and 94239 MVZ) and from 3-1/2 mi. E and 5 mi.
N Yocolt (94240-94244 MVZ) reveals that the "average" of the coloration
is nearer to that of the paler _T. t. cooperi_ than to that of the
darker _T. t. townsendii_ and indicates why Dalquest, we think
correctly, identified specimens from Yocolt as _T. t. cooperi_. We have
examined also the specimens in the Biological Surveys Collection of the
United States National Museum (catalogue numbers given above) and have
compared them with specimens (comparable in age and seasonal condition
of pelage) of _T. t. townsendii_ (notably a series from Lake Quinalt,
Washington) and of _T. t. cooperi_ (including specimens from Bumping
Lake and Blewett Pass, Washington). In color, the specimens from Mt. St.
Helens are almost exactly intermediate between _T. t. cooperi_ and _T.
t. townsendii_. We choose to use for them the name _T. t. townsendii_ as
did Howell (_op. cit._:109). The specimens from 15 mi. N Carson, those
from Stevenson and the one from Skamania agree in nearly all features of
color with the relatively paler _T. t. cooperi_, as Dalquest (_op.
cit._) thought they would, and we, accordingly, use for them the name
_Tamias townsendii cooperi_.

In view of the findings resulting from our study of the above mentioned
specimens of the Townsend Chipmunk in Washington, it seemed worthwhile
to examine the material of the same species from Hood River, Oregon.
Howell (_op. cit._:109) listed one specimen from there as _E. t.
townsendii_, but (_op. cit.:_ fig. 7, p. 107) mapped the locality as
within the geographic range of _E. t. cooperi_. The specimen (89061 BS)
is a juvenile having external measurements of only 175, 80 and 31.
Although the color is intermediate between that of the two subspecies
concerned, greater resemblance is shown to _T. t. townsendii_. We have
not examined any other specimen of the species _Tamias townsendii_ so
young as No. 89061, but suspect that older specimens from the same place
would be paler by a slight degree. This suspicion, and more especially
the light color of an older specimen from nearby White Salmon,
Washington, and the light color of two older specimens from Parkdale,
Oregon, which seem to us to be referable to _T. t. cooperi_, influence
us to refer the specimen from Hood River to _Tamias townsendii cooperi_

~Tamias townsendii townsendii~ Bachman

A. H. Howell (N. Amer. Fauna, 52:111, November 30, 1929) referred
specimens of the Townsend Chipmunk from the lower elevations on the
Olympic Peninsula to _Eutamias townsendii townsendii_ but referred
specimens from the central mountains on that peninsula to _Eutamias
townsendii cooperi_. The subspecies _T. t. cooperi_ thus is represented
as having a geographic range of two separate parts: (1) The Cascade
Mountains from southern British Columbia into southern Oregon, and (2)
the area of the Olympic Mountains, the latter area being entirely
surrounded by the geographic range of _T. t. townsendii_. Dalquest
(Univ. Kansas Publ. Mus. Nat. Hist., 2:261 and 262, April 9, 1948)
employed Howell's arrangement.

We have examined the specimens, in the Biological Surveys Collection of
the United States National Museum, from the Olympic Peninsula and fail
to find significant differences in external measurements or in size or
shape of skulls between specimens from the mountains (alleged _T. t.
cooperi_) and those from other parts of the Peninsula (assigned to _T.
t. townsendii_). Nevertheless, the specimens from the higher parts of
the Olympic Mountains resemble _T. t. cooperi_ in being less ochraceous
than are specimens of _T. t. townsendii_ from elsewhere on the Olympic
Peninsula, and in this one respect, in series, they more closely
resemble _T. t. cooperi_. Even so, the upper parts of the specimens from
the mountains are darker than in _T. t. cooperi_ of the Cascades. In
dark color of the superciliary stripe the specimens in question are
referable to _T. t. townsendii_. The over-all gray tone, resembling that
of _T. t. cooperi_, upon close inspection is found to be in considerable
degree the result of wear, and the difference in grayness from _T. t.
townsendii_, when specimens in comparable pelage are compared, is
slight. This tendency to lighter color in specimens from higher
elevations is seen in other places in Washington within the geographic
range of _Tamias townsendii_. We feel, therefore, that the mentioned
resemblance in color between specimens from the Olympic Mountains and
those of _T. t. cooperi_ from the Cascade Mountains is not significant
taxonomically. To us, all of the animals of the species _Tamias
townsendii_ from the Olympic Peninsula seem best referred to the
subspecies _Tamias townsendii townsendii_ Bachman.

~Tamias striatus ohionensis~ Bole and Moulthrop

A. H. Howell (Jour. Mamm., 13:166, May 14, 1932) referred a specimen
(252979 USNM) from Athens, Ohio, to _Tamias striatus fisheri_.
Subsequently, Bole and Moulthrop (Sci. Publs. Cleveland Mus. Nat. Hist.,
5:83-181, September 11, 1942) named _Tamias striatus ohionensis_ and
_Tamias striatus rufescens_, both of which occur in Ohio. They (_op.
cit._: 137) also excluded _T. s. fisheri_ from the state list of mammals
of Ohio. The locality of Athens lies between the ranges of _T. s.
ohionensis_ and _T. s. rufescens_, as outlined by referred specimens,
and thus the identity of the specimen from that place was left in doubt.
We have examined the specimen and among named kinds find that it most
closely resembles _T. s. ohionensis_ in its less widely spreading
zygomata, slender incisors and dull-colored pelage. We prefer the
specimen to _T. s. ohionensis_.

The subspecific identity of specimen No. 174762 USNM, a skin only, from
Nobleville, Hamilton Co., Indiana, assigned by Howell (N. Amer. Fauna,
52:21, November 30, 1929) to _T. s. griseus_ and by Lyon (Amer. Mid.
Nat., 17(1):191, January, 1936) to _T. s. fisheri_, was left in doubt by
Bole and Moulthrop's (_op. cit._) assignment of specimens to _T. s.
ohionensis_. Although the specimen lacks a skull and tail, on the basis
of its dull-colored pelage and dark brown (anteriorly) median dorsal
stripe, we identify No. 174762 as _T. s. ohionensis_. For the same
reason, specimen No. 125445 USNM, from Bascom, Indiana, referred by
Howell (_op. cit._:16) to _T. s. striatus_, and by Lyon (_op. cit._:191)
to _T. s. fisheri_, required re-examination. The specimen appears to be
an intergrade between _T. s. striatus_ and _T. s. ohionensis_; it is
probably best referred to the latter subspecies which it resembles in
having short nasals. In color it is intermediate, but it does not
possess the narrowly spreading zygomata of _T. s. ohionensis_ and, in
this respect, more nearly approaches _T. s. striatus_.

Specimen No. 13815 USNM, an alcoholic, from Wheatland, Knox Co.,
Indiana, was assigned by Howell (_op. cit._, 1929:21) to _T. s. griseus_
and by Lyon (_loc. cit._) to _T. s. fisheri_. Although the specimen is
much faded and cannot be identified with certainty, we assign it to _T.
s. ohionensis_. Allowing for fading, it seems to resemble _ohionensis_
more in the lighter color of the anterior part of the median dorsal
stripe, than it does either _griseus_ or _fisheri_. We are also
influenced in making this allocation by Bole and Moulthrop's (_op.
cit._:137) finding intergradation between _T. s. ohionensis_ and _T. s.
striatus_ in a specimen obtained at New Harmony, Posey Co., Indiana.

Howell (Jour. Mamm., 13:166, August 9, 1932) referred two specimens from
Boone County, Indiana, to _T. s. fisheri_. We have examined a specimen
(5675 AMNH) from that place and think it is one of the two seen by
Howell. The specimen is a poorly made skin in worn winter pelage with
the skull inside. Because it differs from _T. s. fisheri_ and agrees
with _T. s. ohionensis_ in the color of both upper parts and underparts
(comparisons made with material of comparable stage of molt), we assign
it to the latter subspecies. Howell (_loc. cit._) referred specimens
from Overton (57394), Wooster (57398, 57399, and 57442), and Loudonville
(57391-57393), all from Ohio, in the Museum of Zoology of the University
of Michigan, to _Tamias striatus fisheri_. We have examined these
specimens and find them to be readily separable from _T. s. rufescens_
on the basis of darker coloration. The affinities of the specimens in
question are with _T. s. fisheri_ and _T. s. ohionensis_. As a standard
for comparison we have used specimens in the Museum of Zoology,
University of Michigan, in comparable pelage of _T. s. ohionensis_ from
Dearborn County, Indiana, taken in August and specimens of _T. s.
fisheri_ from "near" summit Butt Mtn. and Little Meadows, both places in
Giles County, Virginia, as well as two specimens from Allair, Monmouth
County, New Jersey. On the basis of buffy (instead of white) edging of
the tail, buffy (not white) light dorsal stripes, and buffy (not black)
anterior third of the median dark stripe, the specimens from Overton,
Wooster, and Loudonville are referred to _Tamias striatus ohionensis_.

~Tamias striatus pipilans~ Lowery

A. H. Howell (N. Amer. Fauna, 29:16, November 30, 1929) recorded six
specimens of _Tamias striatus striatus_ from Greensboro, Alabama.
Subsequently, Lowery (Occas. Papers Mus. Zool., Louisiana State Univ.,
13:235, November 22, 1943) named _T. s. pipilans_ and assigned to it
specimens from northeastern Alabama. Lowery did not, however, mention
the specimens from Greensboro and, thus, their subspecific identity was
placed in doubt. We have examined five of the six specimens mentioned by
Howell (_loc. cit._) (57034-57036, 57588, and 77037 BS) and because of
their brilliant color and large size, refer them to _Tamias striatus
pipilans_ Lowery.

~Tamias striatus rufescens~ Bole and Moulthrop

A. H. Howell (Jour. Mamm., 13:166, August 9, 1932) also referred a
specimen (13154), from La Porte, Indiana, in the Chicago Nat. History
Museum to _T. s. fisheri_. We find the specimen to be distinguishable
from _T. s. fisheri_ in darker, richer pelage, brown instead of blackish
anterior third of the median dorsal stripe, more buffy light dorsal
stripes, and more heavily constructed skull. The specimen most closely
resembles _T. s. rufescens_ in having, as compared to _T. s.
ohionensis_, brighter, more rufescent color, wider incisors,
proportionately narrower interorbital region, and more widely spreading
zygomatic arches. We refer it to that subspecies.

~Sciurus carolinensis pennsylvanicus~ Ord

When J. A. Allen considered what name to apply to the gray squirrel of
northeastern United States and adjacent parts of Canada, (Monogr. N.
Amer. Rodentia, p. 709, 1877) he selected the name _leucotis_ of Gapper
(Zool. Jour., 5:206, 1830) as applicable. Allen rejected Ord's
(Guthrie's Geog., 2nd Amer. Ed., Zool. App., 2:292, 1815) earlier name,
_Sciurus Pennsylvanica_, because (_loc. cit._) "it was given to
specimens from the Middle Atlantic States, and hence from a locality
bordering upon the habitat of the southern form, and consequently the
name is not strictly applicable to the northern type as developed in the
Northern and Northeastern States and the Canadas." It must be recalled
that Allen had not at that time seen a copy of Ord's exceedingly rare
work and was basing his comments on Baird's statements on Ord's
treatment of the squirrels.

Subsequently, Rhoads obtained a copy of the second edition of Guthrie's
Geography and had Ord's zoological appendix thereto reprinted. The
reprinted version (now known generally as Ord's Zoology by Rhoads, 1894)
contains (Appendix, p. 19) Rhoads' review of the _pennsylvanicus vs.
leucotis_ controversy. Rhoads concluded that _pennsylvanicus_ must apply
because it has priority and is available. The habitat was given by Ord
as "those parts of Pennsylvania which lie to the westward of the
Allegany ridge," not the "Middle Atlantic States" as Allen thought.

Notwithstanding Rhoads' comments, Bangs (Proc. Biol. Soc. Washington,
10:156, December 28, 1896), in his "Review of the Squirrels of Eastern
North America," employed _leucotis_ Gapper and rejected Ord's name
because it "is a _nomen nudum_" and of uncertain application. There
seems to have been no attempt subsequently to review the pertinent

We are of the opinion that Rhoads' (_loc. cit._) analysis and
conclusions are correct and as cogent today as then. We do not agree
with Bangs that _pennsylvanicus_ is a _nomen nudum_ for the following
reasons. The name was based on melanistic individuals and could
conceivably be applied to three species of squirrels, the red squirrel,
the fox squirrel, and the gray squirrel. Melanistic red squirrels,
_Tamiasciurus hudsonicus_, are everywhere rare and in any case appear as
individuals and not populations. Ord (_loc. cit._) reported that his
_Sciurus Pennsylvanica_ was _abundant_. Ord, we think, was not referring
to the fox squirrel, _Sciurus niger_, because he wrote that _S.
Pennsylvania_ "has always been confounded with... [_Sciurus niger_], but
it is a different species," and (_loc. cit._) described _S. niger_ as a
"Large Black Squirrel" and _Sciurus Pennsylvanica_ as a "Small Black
Squirrel." Therefore, _pennsylvanicus_ Ord can refer only to _Sciurus
carolinensis_. Further, melanistic gray squirrels then, as now, were
common in western Pennsylvania and exceedingly rare in eastern
Pennsylvania. Additionally, Ord described his animal, although
admittedly inadequately (small, black, not _S. niger_). The name
_Sciurus Pennsylvanica_ Ord is clearly not a _nomen nudum_ and must
replace _leucotis_ Gapper.

Allen's (_loc. cit._) argument that the specimens were not
representative of "_leucotis_" because they were from the Middle
Atlantic States is based on an initial misunderstanding of the locality.
Further, whether or not "topotypes" are representative of a subspecies
has no bearing on the availability of the name appended to them. The
name and synonomy of the northern gray squirrel are as follows:

     ~Sciurus carolinensis pennsylvanicus~ Ord

     1815. _Sciurus Pennsylvanica_ Ord, Guthrie's Geog., 2nd
     Amer. Ed., 2:292. Type locality, western Pennsylvania.

     1894. _Sciurus carolinensis pennsylvanicus_, Rhoads,
     Appendix of reprint of Ord (_supra_), p. 19.

     1792. _Sciurus cinereus_ Schreber, Säuget., 4:766. Type
     locality, eastern United States, probably New York State.
     (_Nec Sciurus cinereus_ Linnaeus.)

     1830. _Sciurus leucotis_ Gapper, Zool. Jour., 5:206. Type
     locality, region between York and Lake Simcoe, Canada.

     1849. _Sciurus migratorius_ Audubon and Bachman, Quad. N.
     Amer., 1:265 (based on _S. leucotis_ Gapper).

     1877. _Sciurus carolinensis_ var. _leucotis_, J. A. Allen,
     Monogr. N. Amer. Rodentia, Sciuridae, p. 700 (_et auct._).

~Sciurus niger rufiventer~ Geoffroy

Two specimens (36192/48550, a young male with unworn teeth, and
36193/48551, an adult male with much worn teeth, both in the United
States Biological Surveys Collection in the National Museum) were
recorded by Bailey (N. Amer. Fauna, 25:75, 1905) as _Sciurus
ludovicianus_ from Gainesville, Texas. Bailey (_loc. cit._) further
stated that if the name _Sciurus rufiventer_ Geoffroy proved usable it
would apply to the specimens from Gainesville. Since the name
_rufiventer_ was revived there would be no question concerning the
identity of these specimens had not Lowery and Davis (Occas. Papers,
Mus. Zool., Louisiana State Univ., 9:172, 1942) assigned three specimens
(not seen by us) to _Sciurus niger limitis_ Baird from a point only
thirteen miles northwesterly. Lowery and Davis (_loc. cit._) say that
their specimens are intergrades (presumably with _rufiventer_) and
Bailey (_loc. cit._) noted that his two specimens from Gainesville "are
in size and color nearer to _ludovicianus_ [= _rufiventer_] than to
typical _limitis_." Examination of the two specimens from Gainesville
convinces us that Bailey was correct and the specimens therefore are
referable to _Sciurus niger rufiventer_. More in detail, the color
agrees with that of _rufiventer_ and differs from that of _limitis_ and
from that of darker specimens of _Sciurus niger ludovicianus_ (in the
restricted sense used by Lowery and Davis, _op. cit._: 104). Also the
size is larger than in _limitis_ and as in _rufiventer_ or
_ludovicianus_. Selected measurements of Nos. 36192/48550 and
36193/48551 are, respectively, as follows: Total length, 505, 500;
length of tail, 237, 228; length of hind foot, 72, 70; basilar length of
Hensel, 48.5, 48.6; zygomatic breadth, 35.1, 36.0; length of nasals,
21.4, 22.3; alveolar length of maxillary tooth-row, 11.8, 11.1; width
across posterior tongues of premaxillae, 17.5, 18.4.

~Sciurus variegatoides rigidus~ Peters

Harris (Occas. Papers Mus. Zool., Univ. Michigan, 266:1, June 28, 1933)
named _Sciurus variegatoides austini_ with type locality at Las Agujas,
Province of Puntarenas, Costa Rica. Later, in his revision of the
species _Sciurus variegatoides_, he (Misc. Publs. Mus. Zool., Univ.
Michigan, 38:19, September 7, 1937) referred specimens from Chomes,
Costa Rica, to _S. v. austini_ and (_op. cit._:24) specimens from
Puntarenas, Province of Puntarenas, to _S. v. rigidus_, an inland
subspecies. The geographic arrangement of these referred specimens
seemed to warrant a reconsideration of the material. We have examined
specimens of _S. variegatoides_ in the Museum of Zoology, University of
Michigan, from the following localities in Costa Rica: Puntarenas
(62703-62706), Las Agujas (65118 [type of _S. v. austini_],
59847-59850), Río Las Agujas (65114-65117), Agua Caliente (66483),
Zarcéro (75757-75761, 75765), Cartago (67546, 67547), and Esparta
(75762-75764). The specimens listed by Harris (_op. cit._, 1937:19) as
from Chomes, in the Museum of Zoology of the University of Michigan, are
not now in that museum and we have not seen them.

Harris (_op. cit._:19) characterized _S. v. austini_ as differing from
_S. v. rigidus_ in having brightly rufous legs (Ochraceous-Orange) in
_S. v. rigidus_ and a dorsal coloration resulting from a mixture of
shiny black and silver (Ochraceous-Orange mixed with black in _S. v.
rigidus_). We find that in the color of the legs of the paratypes of _S.
v. austini_ there is considerable variation ranging from bright rufous
in No. 65116 to much darker and duller in No. 59849. In six of the ten
specimens of the type series, the color is rufous, but in the other four
the color of the legs approaches and overlaps that found in the referred
specimens of _S. v. rigidus_. The color of the dorsum of _S. v. austini_
is also variable. No. 59850, for example, is dark brown and closely
resembles No. 75762, from Esparta, which was referred to _S. v.
rigidus_. Further, some specimens referred to _S. v. rigidus_ (67546 and
67547) have the bright-colored legs of _S. v. austini_ and some (75759,
for example) have the black-and-silver back of _austini_. We recognize
differences of an average sort between the now-available specimens of
the two alleged subspecies, but because of the individual variation that
exists, we feel that recognition of two subspecies is not indicated.
There is also some variation that is the result of wear and molt and one
of us (Kelson) feels that some of the differences are explainable on
this basis. Accordingly, we prefer to adopt a more conservative
taxonomic arrangement than that of Harris for this group of the Costa
Rican squirrels and arrange _Sciurus variegatoides austini_ Harris,
1933, as a synonym of _Sciurus variegatoides rigidus_ Peters, 1863.

~Thomomys bottae alienus~ Goldman

Six specimens (21249-21253, 212706 BS) from Rice, Arizona, were referred
by Goldman (Proc. Biol. Soc. Washington, 46:76, April 27, 1933) to the
subspecies _Thomomys bottae mutabilis_ Goldman when he proposed that
name as new, but these six specimens were not mentioned by him when he
later named _Thomomys bottae alienus_ (Jour. Washington Acad. Sci.,
28:338, July 15, 1938), to which subspecies the specimens in question
might be expected to belong. Examination of the six specimens reveals
that they are intergrades between _T. b. mutabilis_ and _T. b. alienus_
but that the specimens more closely resemble the latter. More precisely,
slightly larger size of skull, greater ventral inflation of tympanic
bullae, and less depressed occipital region ally the specimens with
_Thomomys bottae alienus_, and we identify them as that subspecies. The
two subspecies concerned are not so distinct as are most subspecies of
_Thomomys bottae_.

~Thomomys bottae aphrastus~ Elliott

Bailey (N. Amer. Fauna, 39:58, November 15, 1915) referred three
specimens from San Antonio, Baja California, to _Thomomys bottae
nigricans_. These specimens have not, to our knowledge, been re-examined
subsequently, although the current taxonomic treatment of the pocket
gophers of Baja California by Huey (Trans. San Diego Soc. Nat. Hist.,
10(4):245-268, 1 map, August 31, 1945) excludes _T. b. nigricans_ from
the area of San Antonio. The pertinent specimens are probably Nos.
10810-10812 in the Chicago Natural History Museum. We have examined the
specimens and, using the comparative materials listed under the account
of _T. b. siccovallis_, find them to be intermediate in most characters
between _T. b. aphrastus_ and _T. b. martirensis_. Because they more
nearly resemble _T. b. aphrastus_ in the weakly-spreading zygomatic
arches, we refer the specimens from San Antonio to that subspecies.

~Thomomys bottae jojobae~ Huey

When Huey (Trans. San Diego Soc. Nat. Hist., 10:256, August 31, 1945)
named _Thomomys bottae jojobae_ from Sangre de Cristo, Baja California,
México, he made no mention of a specimen that Bailey (N. Amer. Fauna,
39:58, November 15, 1915) identified as _Thomomys bottae nigricans_ from
La Huerta, which place is approximately eight miles northwest of Sangre
de Cristo. From a geographic standpoint, it seemed unlikely that the
specimen from La Huerta would be referable to _T. b. nigricans_.
Examination of the specimen (138752 BS) proves it to differ from
topotypes of _T. b. nigricans_ and to agree with _T. b. jojobae_ in
richer, more rufescent color, especially ventrally, and smaller,
slenderer, more delicate skull. The specimen is therefore tentatively
referred to _Thomomys bottae jojobae_. We have not, however, compared it
with specimens of _Thomomys bottae juarezensis_, a subspecies the range
of which lies to the east on the summit of the Sierra Juárez.

~Thomomys bottae martirensis~ J. A. Allen

Bailey (N. Amer. Fauna, 39:58, November 15, 1915) referred pocket
gophers from Piñon on the west slope of the San Pedro Mártir Mountains,
Baja California, to the subspecies _Thomomys bottae nigricans_. The
subspecific identity of these animals has now been reinvestigated
subsequently, although the locality whence they were obtained is far
removed from what is now thought to be the geographic range of _T. b.
nigricans_; further, several other subspecies are known to occur in the
intervening area. We have examined the available material from Piñon
(13853-13855 BS) and find the specimens to agree with _Thomomys bottae
martirensis_ and to differ from _T. b. nigricans_ in lighter color,
larger, more ridged and angular skull; proportionately greater mastoidal
breadth; narrower occipital shelf; more ventrally produced alveolar
ramus of the maxillae; and deeply concave posterior border of the
temporal root of the zygomatic arch. These specimens thus constitute the
northernmost record of _T. b. martirensis_ known to us.

~Thomomys bottae mohavensis~ Grinnell

Bailey (N. Amer. Fauna, 39:73, November 15, 1915) assigned a series of 7
specimens from Lone Willow Spring, California, to the subspecies
_Thomomys bottae perpes_. This locality lies at the northern edge of the
Mohave Desert. Later, Grinnell (Univ. California Publ. Zool., 17:427,
April 25, 1918) named the pocket gophers from approximately the eastern
half of the Mohave Desert, _Thomomys perpallidus_ [= _bottae_]
_mohavensis_, but failed to mention the specimens recorded by Bailey,
and thus their subspecific identity is in doubt. We find that _T. b.
mohavensis_ differs from _T. b. perpes_ in more pallid color (light
yellowish as opposed to dark rufescent) larger size, larger and more
angular skull, angular (as opposed to more evenly bowed) zygomatic
arches, larger and deeper audital bullae, narrower interpterygoid space,
and proportionately greater mastoidal breadth. In external measurements,
size and angularity of skull, width of interpterygoid space and
angularity of the zygomatic arch, the specimens from Lone Willow Spring
seem to be intermediate between the two subspecies, but perhaps show
more resemblance to _T. b. mohavensis_. Otherwise, the specimens closely
resemble _T. b. mohavensis_ to which they are here referred. The
specimens provide a northern marginal record of occurrence for that

Other specimens recorded as _T. b. perpes_ by Bailey (_loc. cit._) from
Grapevine Ranch, California, have also not been mentioned in later
publications although, from a geographic standpoint, they might be
better referred to either _Thomomys bottae pascalis_ or _T. b.
mohavensis_. Comparison of specimens of _T. b. mohavensis_ and _T. b.
pascalis_ from various localities show _T. b. pascalis_ to be larger
(including the skull), darker, and to possess a more nearly vertical
occipital plane, wider-spread but less angular zygomatic arches, less
inflated tympanic bullae, wider braincase (which consequently appears to
be less inflated), proportionately longer and slenderer rostrum, and
broader nasals distally. Cranially, _T. b. pascalis_ differs from _T. b.
perpes_ in essentially the same ways, but to an event greater degree. In
color, _T. b. pascalis_ differs from _T. b. perpes_ in being duller,
less rufescent.

The series of four specimens, in the U. S. Biological Surveys
Collection, from Grapevine Ranch clearly are not referable to _T. b.
perpes_. They do, however, agree with _T. b. mohavensis_ in all
essential particulars except that in two of the four specimens the
braincase is wider and the nasals are wider distally. This width is
evidence of intergradation with _T. b. pascalis_. Seemingly, then, they
are best referred to _Thomomys bottae mohavensis_.

~Thomomys bottae muralis~ Goldman

When Goldman (Jour. Washington Acad. Sci., 26(3):112, March 15, 1936)
described and named this pocket gopher from Arizona, he arranged it as a
full species and stated that there is no evidence of intergradation with
other named kinds. We have examined the holotype and three topotypes
(202579-202582 BS) and compared them with specimens of other kinds of
pocket gophers occurring in northern and central Arizona. The _muralis_
gopher is a depauperate form clearly belonging to the _bottae_ group.
The characters which Goldman (_loc. cit._) set forth as distinguishing
_muralis_ from other named kinds are readily apparent and, like Goldman,
we see no evidence of intergradation. Nevertheless, the characters which
serve to identify the race are, in a general way, those commonly found
in populations of depauperate individuals of _Thomomys bottae_ and _T.
talpoides_. The small size, delicate structure, well-inflated braincase,
short premaxillary tongues, and strongly recurved upper incisors, often
appear in populations existing in inhospitable areas of shallow,
unstable soils. For this reason we feel that the relationships of this
population are best shown by arranging _muralis_ as a subspecies of
_Thomomys bottae_; the name should stand as _Thomomys bottae muralis_

As far as known, _T. b. muralis_ is completely isolated from other
populations of pocket gophers by uninhabitable eroding cliffs. The
animals have been found only on isolated terraces in the lower end of
Prospect Valley (itself a lateral pocket) within the Grand Canyon of the
Colorado River, Hualpai Indian Reservation, Arizona. Consequently it is
unlikely that intergradation with other populations could exist at the
present time.

In short, in arranging _muralis_ as a subspecies of _Thomomys bottae_,
we are influenced, not by the demonstration of intergradation, but by
the degree of morphological differentiation of the population and the
probable reasons therefor.

~Thomomys bottae mutabilis~ Goldman

Goldman (Jour. Washington Acad. Sci., 28:342, July 15, 1938) named the
subspecies _Thomomys bottae pinalensis_ on the basis of only one
specimen, an immature female (245709 BS) from Oak Flat, five miles east
of Superior, Pinal Mountains, Arizona. Examination shows it to be
indistinguishable in characters of taxonomic importance (coloration,
external measurements, shape of skull and size of skull) from specimens
of _T. b. mutabilis_ of comparable sex and age. No. 245709 is well
within the limits of individual variation of _T. b. mutabilis_ as is
shown by the several specimens (all in the U. S. Biological Surveys
Collection) as follow: Nos. 214118, 214670 (topotypes from Camp Verde,
Arizona), 212707 (Chiricahua Ranch, 20 mi. E Calva), 208635 (H-bar
Ranch, 20 mi. S Payson), and 215762 (Turkey Creek). Therefore, the name
_Thomomys bottae pinalensis_ is here arranged as a synonym of the
earlier name, _Thomomys bottae mutabilis_ Goldman (Proc. Biol. Soc.
Washington, 46:75, April 27, 1933), the type locality of which is Camp
Verde, Yavapai County, Arizona.

~Thomomys bottae patulus~ Goldman

When Goldman (Jour. Washington Acad. Sci., 26:113, March 15, 1936) named
the subspecies _Thomomys bottae desitus_, he assigned to it (_op.
cit._:114) 10 specimens obtained at Wickenburg, Maricopa County,
Arizona. He did not mention specimens from Wickenburg when he
subsequently named the subspecies _Thomomys bottae patulus_ (Jour.
Washington Acad. Sci., 28:341, July 15, 1938) and stated that _T. b.
patulus_ was known only from the type locality in the "bottomland along
[the] Hassayampa River, two miles below Wickenburg." Examination in 1950
of specimens referable to _T. b. patulus_ in the U. S. Biological
Surveys Collection shows all of them, including the holotype, to be
labeled "Wickenburg." The 10 specimens from Wickenburg reported by
Goldman in 1936 as _T. b. desitus_ were included by him among the 16
(actually 17, one being a skull only) upon which he based his
description of _T. b. patulus_ in 1938. Examination of the field
catalogues of 3 of the 4 collectors who obtained the specimens discloses
that only the 7 specimens obtained last were recorded as occurring in
the Hassayampa River bottoms; the first 10 were recorded only as from
"Wickenburg." Briefly, only one subspecies, _T. b. patulus_, is present
in the area, and Goldman in 1938 seems to have thought that the two
localities were actually the same, and that "2 miles below Wickenburg"
was the more precise designation.

~Thomomys bottae providentialis~ Grinnell

We have examined a specimen, No. 26120/33526, from 12-Mile Spring,
California, in the U. S. Biological Surveys Collection, which Bailey (N.
Amer. Fauna, 39:73, November 15, 1945) referred to the subspecies
_Thomomys perpallidus_ [= _aureus_] _perpes_. We find the specimen to be
referable to the later named _Thomomys bottae providentialis_ on the
basis of smaller ear, more massive, more ridged and angular skull,
greater interorbital breadth, deeper and thicker rostrum, less globular
bullae, and U-shaped rather than V-shaped interpterygoid space.
Therefore, 12-Mile Spring is the northernmost locality of occurrence of
the subspecies _T. b. providentialis_.

~Thomomys bottae sanctidiegi~ Huey

In his discussion of the pocket gophers of Baja California, Huey (Trans.
San Diego Soc. Nat. Hist., 10:245-268, map, August 31, 1945) made no
mention of specimens from Ensenada, Baja California, recorded by Bailey
(N. Amer. Fauna, 39:58, November 15, 1915) as _Thomomys bottae
nigricans_. We have examined the specimens from Ensenada available to
Bailey in the U. S. Biological Surveys Collection, Nos. 137724, 139890,
and 139891, subadult, immature, and adult, respectively. As compared
with _Thomomys bottae sanctidiegi_ from the mouth of the Tiajuana River
(No. 126028) and _T. b. nigricans_ (topotypes), the one adult specimen
from Ensenada agrees with _T. b. sanctidiegi_ and differs from _T. b.
nigricans_ in lighter color, larger and more angular skull, and more
inflated braincase. The specimens from Ensenada differ from the adjacent
subspecies to the south, _Thomomys bottae proximarinus_ [to judge from
Huey's (_op. cit._) characterization of that subspecies] in lighter
color, and larger, more robust skull. Accordingly, the specimens from
Ensenada are referred to _Thomomys bottae sanctidiegi_.

~Thomomys bottae siccovallis~ Huey

Bailey (N. Amer. Fauna, 39:58, November 15, 1915) listed a specimen from
Mattomi, Baja California, as _Thomomys bottae nigricans_. When Huey
(Trans. San Diego Soc. Nat. Hist., 10:259, August 31, 1945) revived the
name _Thomomys_ [_bottae_] _aphrastus_ Elliot, and named (_op.
cit._:258) _Thomomys bottae siccovallis_ he made no mention of the
specimen, from Mattomi, which, on geographic grounds, would be expected
to be _T. b. aphrastus_, _T. b. martirensis_ J. A. Allen, or _T. b.
siccovallis_. We have examined an adult male (10832 CNHM), probably the
specimen seen by Bailey (_loc. cit._), from Mattomi, and have compared
No. 10832 with six topotypes (10813-10816, 10819 and 10820 CNHM) of _T.
b. martirensis_, the type and one topotype (10798 CNHM) of _T. b.
aphrastus_ and with the original description of _T. b. siccovallis_. The
specimen from Mattomi seems to be unique in the large size of the
tympanic bullae. The specimen in question differs from _T. b.
martirensis_ also in shorter and wider skull, shorter and wider rostrum,
and longer and wider molariform teeth. In these features resemblance is
shown to the holotype of _T. b. aphrastus_ and even greater resemblance
is shown to _T. b. siccovallis_ to which the specimen from Mattomi is

~Thomomys monticola mazama~ Merriam

This subspecies of the Cascades of Oregon and _Thomomys monticola
nasicus_ of the territory immediately to the east of the Cascades, in
the same state, were originally described (Merriam, Proc. Biol. Soc.
Washington, 11:214 and 216, respectively, July 15, 1897) and redescribed
(Bailey, N. Amer. Fauna, 39:123 and 125, respectively, November 15,
1915) as distinguished from each other by paler color, smaller tympanic
bullae and longer nasals in _T. m. nasicus_. The holotypes do differ in
these respects. The assigned (by Bailey, _loc. cit._) specimens indicate
that the opposite condition obtains with respect to the size of bullae;
that is to say, the bullae are smaller in _T. m. mazama_. In these
referred specimens from Oregon the nasals are actually and relatively
longer in _T. m. nasicus_, which averages paler (less black and more
red). Certain specimens of the two subspecies that are comparable as to
sex, age and season, are indistinguishable in color.

This is the background against which Bailey (_op. cit._:125), contrary
to his statement of geographic ranges (_op. cit._:123, 125) and map
(_op. cit._:fig. 5, p. 23), assigned, in his list of specimens
examined, two specimens ([** Male] ad. 79817 and [** Female] ad. 79818
BS) from Pengra, west of the Cascades, to the subspecies _T. m.
nasicus_. In the specimens from Pengra the bullae are angular as in
referred specimens of _nasicus_ (unlike those of the holotype), the
rostra are intermediate in length between those of the two subspecies
concerned, and the color is light as in _T. m. nasicus_ but can be
matched by that of certain specimens of _T. m. mazama_, for example by
that of No. 79821 BS from Diamond Lake, Oregon. Consequently, on
morphological grounds, the two specimens from Pengra can be assigned to
_T. m. mazama_ almost as well as to _T. m. nasicus_. Having regard for
the geographic relations, we assign them to _T. m. mazama_.

In making this tentative identification we are aware that the
acquisition of more nearly adequate material from Oregon, and critical
study of such material, may bring a subspecific arrangement of the
populations of _Thomomys monticola_ different from the current one.

~Thomomys talpoides bullatus~ Bailey

Bailey (N. Amer. Fauna, 39:101, November 15, 1915) identified as
_Thomomys talpoides clusius_ two specimens (66465 and 66523 BS) from
Pass (= Parkman) and one specimen (66464 BS) from Dayton, in Wyoming. We
have examined these specimens and find that they lack the broad
braincase and narrow nasals of _clusius_ and in these and in other
features the three specimens resemble _T. t. caryi_ and _T. t. bullatus_
more than they resemble any other named kinds. Although structurally,
and in color, intermediate between the two subspecies named immediately
above, the specimens show greater resemblance (large size and narrow
braincase) to the latter and are referred by us to _Thomomys talpoides

~Thomomys talpoides clusius~ Coues

Bailey (N. Amer. Fauna, 39:102, November 15, 1915) identified as
_Thomomys talpoides bullatus_ an adult male (147347 BS) from the J. K.
Ranch, 5900 ft., on Meadow Creek, Wind River, Wyoming [= Wind River of
Bailey, _loc. cit._] and a young female (168666 BS) from Sage Creek, 8
mi. NW Fort Washakie, Wyoming. The rosaceous tone of these pale
individuals is more as in some populations of _T. t. ocius_ and _T. t.
clusius_ to the southward. Also, the skull of the male, although large,
is distinctly narrower than in _T. t. bullatus_ and we think shows the
influence of the _T. t. tenellus_ stock. All features considered, we
refer the specimens to _T. t. clusius_.

~Thomomys talpoides glacialis~ Dalquest and Scheffer

Vernon Bailey (N. Amer. Fauna, 39:119, November 15, 1915) listed 19
specimens from Roy, Washington, as _Thomomys douglasi yelmensis_
Merriam. Our examination of 26 specimens (205039-205051, 205072-205077,
and 206545-206551 BS) labeled as "Roy," and presumably including those
listed by Bailey (_loc. cit._), leads us to identify all 26 as _Thomomys
talpoides glacialis_ on the basis of widely spreading zygomatic arches
and decidedly ochraceous hue of underparts.

~Geomys bursarius jugossicularis~ Hooper

Seven skins with skulls (35104/47369-35110/47375 BS) from Las Animas,
Colorado, probably formed the basis for Cary's (N. Amer. Fauna, 33:129,
August 17, 1911) record of _Geomys lutescens_ from that locality.
Comparison of the material reveals that the animals are referable
instead to the later named subspecies, _Geomys lutescens jugossicularis_
Hooper (Occas. Papers Mus. Zool., Univ. Michigan, 420:1, June 28, 1940),
on the basis of (1) more reddish color, (2) deeper zygomatic plate, (3)
shorter jugal as expressed as a percentage of the length of the part of
the zygomatic arch anterior to the jugal, and (4) larger area of inner
face of jugal exposed when skull is viewed from directly above. Possibly
it is noteworthy that the specimens from Las Animas are larger than
Hooper's holotype and one topotype; this larger size is indicative of
intergradation with _G. b. lutescens_ as represented by the specimens
examined by us from Pueblo.

Our examination of an adult female, No. 128242 BS and a juvenal female,
No. 128243 BS, from 15 mi. E Texline, Texas, recorded by Bailey (N.
Amer. Fauna, 25:132, October 24, 1905) under the name _Geomys lutescens_
reveals that the specimens are referable to _Geomys bursarius
jugossicularis_ instead of to _Geomys bursarius major_ on the basis of
(1) mastoid part of tympanic bulla more inflated posteriorly, (2)
narrowness of frontals between posterior tongues of the premaxillae and,
(3) lighter color.

~Liomys irroratus irroratus~ Gray

When Hooper and Handley (Occas. Papers Mus. Zool., Univ. Michigan,
514:1-34, October 29, 1948) published a revised map (_op. cit._:3)
showing the geographic distribution of the subspecies of _Liomys
irroratus_ they did not mention a specimen from Agusinapa, Guerrero,
which inferentially from their map would be _L. i. irroratus_ although
it previously had been recorded as _L. i. torridus_ by Goldman (N.
Amer. Fauna, 34:55, September 7, 1911). We have examined the specimen
(70228 BS), which retains the upper deciduous premolar. Its long foot
(32 mm.) and broad cranium (13 mm.) are the bases for identifying the
specimen as _Liomys irroratus irroratus_ instead of _L. i. minor_, which
is smaller.

~Liomys irroratus minor~ Merriam

When Hooper and Handley (Occas. Papers Mus. Zool., Univ. Michigan,
514:1-34, October 29, 1948) published a revised map (_op. cit._:3)
showing the geographic distribution of the subspecies of _Liomys
irroratus_ they did not mention five specimens from Tlapa, Guerrero,
which inferentially from their map would be _L. i. irroratus_ although
these specimens previously had been recorded as _L. i. torridus_ by
Goldman (N. Amer. Fauna, 34:55, September 7, 1911). We have examined the
five specimens (70221-70225 BS), three of which retain the upper
deciduous premolars and two of which have the upper fourth premolar
unworn. The short, wide rostrum is unlike the long slender rostrum of
topotypes of _L. i. torridus_ of comparable age, and agrees with the
condition in topotypes of _L. i. minor_ of comparable age. It is on this
basis of wider rostrum that we refer the five specimens from Tlapa to
_Liomys i. minor_ which Hooper and Handley (_op. cit._:13) described as
differing from the geographically adjacent _L. i. irroratus_ in "short
and strongly tapered rostrum." We would add that we have not
independently verified this difference between _L. i. minor_ and _L. i.
irroratus_ for want of specimens of _L. i. irroratus_ comparable in age
to the five individuals from Tlapa.

The map of Hooper and Handley (_loc. cit._) inferentially excludes
Tlalixtaquilla, Guerrero, from the geographic range of _L. i. minor_
(and places Tlalixtaquilla within the range of _L. i. irroratus_)
although Goldman (_op. cit._:56) previously had identified specimens
from this place as _L. i. minor_. Our examination of the two immature
specimens (70227 and 70230 BS) from Tlalixtaquilla reveals that they
closely resemble the holotype of _L. i. minor_ and leads to the
conclusion that they are _Liomys irroratus minor_.

~Perognathus amplus pergracilis~ Goldman

When Bole (Sci. Publ. Cleveland Mus. Nat. Hist., 5(2):6, December 4,
1937) named and described _Perognathus longimembris salinensis_, he
listed as comparative material of _P. l. bangsi_, a specimen in the
Museum of Comparative Zoology from Parker, Yuma Co., Arizona. There was
some reason to doubt the identification of the specimen since it is the
only record of occurrence of the subspecies from east of the Colorado
River. There is no specimen of _Perognathus longimembris_ from Arizona
in the Museum of Comparative Zoology. There is one specimen of pocket
mouse (18213, a skin only) from 30 miles east of Parker. We think that
this is the specimen seen by Bole because at one time according to the
label, it had been identified as _Perognathus panamintinus_ [=
_longimembris_] _bangsi_. If the identification of this skin-only had
been made by means of Osgood's key (N. Amer. Fauna, 18:14-15, September
20, 1900), the animal would have "keyed out" to _P. longimembris_
because the total length is recorded on the label as 130. Seth B. Benson
has subsequently examined the specimen. The label now bears in
handwriting the name of _P. amplus pergracilis_ and is followed by
Benson's initials as the identifier. Although we lack adequate
comparative material, we consider the specimen to be _P. amplus
pergracilis_ Goldman, because the skin answers well to the description
of _P. a. pergracilis_ and because of the name currently on the label
with Benson's initials.

~Perognathus longimembris panamintinus~ Merriam

In the current literature, Californian specimens of the little pocket
mouse stand identified as _Perognathus longimembris nevadensis_ from
Oasis and vicinity of Benton Station (Grinnell, Univ. California Publ.
Zool., 40:147, September 26, 1933). When one of us (Hall, Mammals of
Nevada, p. 360, July 1, 1946) reported specimens from southwestern
Nevada as _Perognathus longimembris panamintinus_ he did so on the basis
of study of specimens which included those from Oasis (in the California
Museum of Vertebrate Zoology) that he at that time (in ms.) identified
as _P. l. panamintinus_. Those specimens from Oasis have the hair on the
underparts white all the way to the base as also do specimens from
Morans, 5000 ft. (29583/41638 BS), in contrast to the plumbeous
underparts of _P. l. nevadensis_. It is on this basis that we identify
specimens from the places mentioned above as _Perognathus longimembris
panamintinus_. "Vicinity of Benton Station" as given by Grinnell (_loc.
cit._) is interpreted to include Morans, Mono County.

~Dipodomys agilis martirensis~ Huey

Elliot (Field Columb. Mus., Zool. Ser., Publ. 79, 3(12):221, August 15,
1903) referred specimens from Rosarito and Rosarito Divide, San Pedro
Mártir Mts., Baja California, to _Perodipus_ [= _Dipodomys_] _agilis_.
According to the currently known distribution of _Dipodomys agilis_ in
Baja California (see Huey, Trans. San Diego Soc. Nat. Hist., 11:237,
April 30, 1951), the specimens seemed likely to belong to the subspecies
_D. a. martirensis_. An examination of the specimens (10644, 10690-10693
CMNH from Rosarito, and 10694 from Rosarito Divide) shows that, on the
basis of large ear and comparatively narrow braincase, they are in fact
referable to _D. a. martirensis_. Only No. 10693, with its broader
braincase, seems atypical. Comparative materials used are in the Chicago
Natural History Museum as follows: _D. a. martirensis_: Baja California:
San Matias Spring, 2. _D. a. simulans_: Baja California: Ensenada, 8.
California: Dulzura, 1 (topotype); San Luis del Rey, 3.

~Dipodomys agilis simulans~ (Merriam)

J. A. Allen (Bull. Amer. Mus. Nat. Hist., 5:184, August 18, 1893) listed
as _Perodipus agilis_ a specimen (6306/4941 AMNH) from Valladares, Baja
California. Subspecies of this species were subsequently named without
mentioning this specimen that, on geographic grounds, might be either
_D. a. martirensis_ or _D. a. simulans_. Certain measurements of the
specimen are as follows: Total length, 288; length of tail, 171; length
of hind foot (dry), 41.0; greatest length of skull, 39.5; width of
maxillary arch at middle, 4.5. The long tail and wide (4.5) maxillary
arch are characteristic of _Dipodomys agilis simulans_ and constitute
the basis for identifying the specimen as of that subspecies.

~Baiomys taylori analogus~ Osgood

The geographic range currently assigned to _Baiomys taylori paulus_ (J.
A. Allen) is separated in two parts by the geographic range assigned to
_B. t. analogus_. The southern, separated part of the range of _B. t.
paulus_ rests wholly on ten specimens from Colima, Colima, identified as
_B. t. paulus_ by Osgood in his "Revision of the mice of the American
genus Peromyscus" (N. Amer. Fauna, 28, April 17, 1909) where (p. 255) he
places as a synonym of _Peromyscus taylori paulus_ J. A. Allen, 1903,
_Peromyscus allex_ Osgood, 1904. The later name was based on these ten
specimens (33422/45445-33427/45450, 33429/45452, 33432/45455, and
33435/45458 BS) from Colima. Osgood had a choice of synonymizing _P.
allex_ under _P. paulus_ or _P. t. analogus_. According to Osgood's
concept, _analogus_ was blackish and large; _allex_ was grayish and
small; and _paulus_ was fawn colored and intermediate in size. The more
nearly equal size of _paulus_ and _allex_ probably influenced Osgood in
making his choice. After examining the original materials we think there
is more to recommend the alternate choice. For example, two topotypes
of equal age of the same sex of _allex_ (33424/45447) and _analogus_
(120264 BS) are of almost the same size and, respectively, measure as
follows: Total length, 107, 108; length of tail, 42, 45; length of hind
foot (measured dry), 13.1, 12.8; greatest length of skull, 17.6, 17.7;
zygomatic breadth, 9.3, 9.2. Although _analogus_ does average darker, a
topotype, No. 120267 BS, from Zamora, is indistinguishable from several
of the topotypes of _allex_. Consequently, we arrange _Peromyscus allex_
Osgood as a synonym of _Baiomys taylori analogus_ (Osgood) 1909 and
refer the specimens from Colima to the latter.

~Peromyscus eremicus eremicus~ (Baird)

Osgood (N. Amer. Fauna, 28:242, April 17, 1909) listed a specimen of
this subspecies from Sierra Encarnación, Nuevo Leon. A specimen, No.
79614 BS, of this species was obtained on July 31, 1896, at Sierra
Encarnación, Coahuila, by Nelson and Goldman. We know of no specimens of
this subspecies from Sierra Encarnación, Nuevo Leon, and assume that
Osgood referred to the Coahuilan specimen. Further support for this
assumption is Osgood's (_loc. cit._) note that the Sierra Encarnación
specimen is aberrant and, to our eye, so is No. 79614 from Coahuila.

~Peromyscus merriami merriami~ Mearns

Osgood (N. Amer. Fauna, 28:239, April 17, 1909) placed _P. merriami_ in
synonymy under _Peromyscus eremicus eremicus_ (Baird). Because Seth B.
Benson, and subsequently the late Wilfred H. Osgood, told one of us
(Hall) that _Peromyscus merriami_ was specifically distinct from
_Peromyscus eremicus eremicus_, we have examined the specimens from
Sonoyta, Sonora, and Quitobaquita, Arizona, referred by Mearns (Bull. U.
S. Nat. Mus., 56:434-435, and 444, April 13, 1907) to _P. e. eremicus_
and _P. merriami_, respectively. We perceive the differences that Mearns
(_loc. cit._) described and recognize _P. merriami_ as a species
separate from _P. eremicus_.

Also we have compared the type and one topotype of _Peromyscus goldmani_
Osgood with the holotype and referred specimens mentioned above, of _P.
merriami_, and feel that the two kinds are no more than subspecifically
distinct. Accordingly, _P. goldmani_ should stand as _Peromyscus
merriami goldmani_. This arrangement is made with the knowledge that
Burt (Misc. Publ. Mus. Zool., Univ. Michigan, 39:56, February 15, 1938)
arranged _P. goldmani_ as a synonym of _Peromyscus eremicus_.

~Peromyscus truei preblei~ Bailey

Osgood (N. Amer. Fauna, 28: 171, April 17, 1909) listed two specimens
from Crooked River, 25 miles southeast of Prineville, Oregon, as
_Peromyscus truei gilberti_ with the notation "approaching _truei_?"
Subsequently, Bailey (N. Amer. Fauna, 55: 188, August 29, 1936) named
_Peromyscus truei preblei_ with type locality at Crooked River, 20 miles
southeast of Prineville, a place from which Bailey had two specimens. We
think the specimens recorded by the two authors are the same, and,
according to the specimen labels, were placed correctly as to locality
by Bailey. Our reasons are as follows: (a) The specimens mentioned by
Bailey were presumably available to Osgood, but Osgood made no mention
of specimens from "20 miles southeast of Prineville," (b) we find no
specimens nor other records pertaining thereto, of _Peromyscus truei_
from the locality given by Osgood, (c) Osgood indicated that the
specimens he saw were not typical of _P. t. gilberti_ and (d) _P. m.
gilberti_, geographically the nearest subspecies, is recorded otherwise
no closer to Prineville than Grants Pass, approximately 175 miles
southwest in southwestern Oregon.

~Sigmodon hispidus cienegae~ A. B. Howell

J. A. Allen (Bull. Amer. Mus. Nat. Hist., 5:28, March 16, 1893) listed
as _Sigmodon hispidus arizonae_ Mearns one specimen from Granados,
Sonora, at a time when _S. h. cienegae_ had not been named. We have
examined the specimen (5389 AMNH) which has the skull inside and which
lacks external measurements. It was taken on November 16, 1890, and is
darker than specimens of _S. h. arizonae_ collected in September at Fort
Verde, Arizona. The color is essentially as in specimens of _S. h.
cienegae_ from Fairbank, Arizona (March-taken specimens). Because of
this agreement in color and because of the geographic origin of the
specimen from Granados, we refer the animal to _Sigmodon hispidus

~Sigmodon hispidus zanjonensis~ Goodwin

Goodwin (Bull. Amer. Mus. Nat. Hist., 79:169, May 29, 1942) listed four
specimens from Honduras (El Jaral, 2; and Las Ventanas, 2) as _Sigmodon
hispidus saturatus_ Bailey. Because these localities fall within the
geographic range of _S. h. zanjonensis_ we were lead to examine the
specimens. Three are young and one (126113 AMNH from Las Ventanas) is an
adult female. The underparts of the young are washed with rufous as in
_S. h. saturatus_. The adult lacks this rufous as do specimens of _S. h.
zanjonensis_ and some specimens of _S. h. saturatus_. In the adult the
color of the upper parts and size of the upper cheek-teeth are
intermediate between the dark-backed, small-toothed _S. h. saturatus_
and the paler-backed, large-toothed _S. h. zanjonensis_. The rostrum is
intermediate in width but definitely nearer the broad condition which
obtains in _S. h. saturatus_. The tail is long, actually and in relation
to the body (total length 275, tail 130), as in _S. h. zanjonensis_ to
which we refer the specimens in question.

~Oryzomys couesi couesi~ (Alston)

For alleged occurrence at Reforma in Oaxaca, México (Goldman, N. Amer.
Fauna, 43:31, September 23, 1918), see under _Oryzomys couesi mexicanus_

~Oryzomys couesi mexicanus~ J. A. Allen

Goldman (N. Amer. Fauna, 43, September 23, 1918) listed, as in the Field
Museum of Natural History [= Chicago Natural History Museum] one
specimen from Reforma, Oaxaca, under _O. c. mexicanus_ (p. 35) and one
specimen from the same place under _O. c. couesi_ (p. 31). In the
Chicago Natural History Museum we can find only one specimen. It is a
young male, skull with skin, in which the last molar has not yet
erupted, and bears the catalogue number 13654. It is, in our opinion,
referable to _O. c. mexicanus_. Because we suspect that Goldman (_op.
cit._) by error listed this one specimen twice (once under _O. c.
couesi_ and once under _O. c. mexicanus_) it seems best to exclude
Reforma, Oaxaca, from the geographic range of _O. c. couesi_.

~Oryzomys alfaroi saturatior~ Merriam

A series of _Oryzomys alfaroi_ in the U. S. Biological Surveys
Collection obtained at Tumbala, 5000 ft., Chiapas, México, the type
locality of _Oryzomys alfaroi saturatior_, contains individuals some of
which Goldman (N. Amer. Fauna, 43:66, September 23, 1918) referred to
the subspecies _O. a. saturatior_ and one which he referred to _O. a.
palatinus_. This latter specimen, to judge from the external
measurements given by Goldman (_loc. cit._), is No. 76328. In comparison
with the other material which Goldman saw, we find the specimen to agree
with _O. a. palatinus_ in pale color and posterior concavity of the
posterior border of the palate. In some other diagnostic cranial
characters, it is indistinguishable from specimens of _O. a. saturatior_
from the same locality, and in other characters, notably the slenderness
of the rostrum, it is intermediate between the two subspecies
concerned. In short, although we see the reasons for Goldman's
subspecific identification of this individual, we think, in view of the
structural intermediacy of the animal and the characters of the series
_en masse_, that it is best referred to _Oryzomys alfaroi saturatior_.

~Zapus princeps idahoensis~ Davis

Preble (N. Amer. Fauna, 15:23, August 8, 1899) referred two specimens
from Henry House and three from 15 miles south of Henry House, both
localities in Alberta, Canada, to the subspecies _Zapus princeps
princeps_. Subsequently, when _Z. p. kootenayensis_ (Anderson, Nat. Mus.
Canada, Ann. Rept. 1931, p. 108, November 24, 1932) and _Z. p.
idahoensis_ (Davis, Jour. Mamm., 15(3):221, August 10, 1934) were named,
no mention was made of these specimens although the ranges assigned to
_Z. p. kootenayensis_ and _Z. p. idahoensis_ seemed to isolate the Henry
House area from the remainder of the range (as recorded) of _Z. p.
princeps_. We have examined the pertinent specimens in the U. S.
Biological Surveys Collection (75452 and 75453 from Henry House;
81509-81510 from 15 mi. S Henry House). On the basis of paler color,
reduced lateral line, smaller skull, shorter palatal bridge and
zygomatic arches, they are, among named subspecies, best referred to
_Zapus princeps idahoensis_.

_Transmitted July 30, 1952._

*** End of this Doctrine Publishing Corporation Digital Book "Comments on the Taxonomy and Geographic Distribution of Some North American Rodents" ***

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