J. KNOX JONES, JR.

In his "Revision of the pocket mice of the genus Perognathus," Osgood

(1900:18-20) reviewed the distribution, as then known, of _Perognathus

fasciatus_ and recognized two geographic races--_Perognathus fasciatus_

[_fasciatus_] Wied-Neuwied in eastern Montana and Wyoming and adjacent

parts of North and South Dakota, and _Perognathus fasciatus infraluteus_

Thomas, known only from the type locality at Loveland, Larimer County,

Colorado. Later, Cary (1911:61) described _Perognathus fasciatus litus_

as a pale subspecies occurring in the lower Sweetwater Valley and

adjacent parts of the Red Desert of south-central Wyoming. After 1911 no

important taxonomic contributions dealing with _Perognathus fasciatus_

appeared.

In studying the kinds of pocket mice known from Nebraska, I examined

thirteen specimens of _P. fasciatus_ from the northwestern part of the

state which did not agree satisfactorily with the descriptions of any

known subspecies of _fasciatus_. This impelled me to examine material

from the entire range of _P. fasciatus_. This examination revealed that:

(1) _Perognathus callistus_ Osgood, heretofore considered to be a full

species, should be reduced to subspecific rank under _P. fasciatus_; and

(2) specimens from eastern Wyoming and adjacent parts of Montana, South

Dakota and Nebraska represent an heretofore unrecognized subspecies.

Further investigation, however, revealed that _Perognathus flavescens

olivaceogriseus_ Swenk, described from northwestern Nebraska (Swenk,

1940:6), is not _flavescens_ but actually _fasciatus_. Since the

specimens on which Swenk's description was based were taken within the

range of this newly recognized subspecies, and since my examination of

the holotype shows it to be of the species _Perognathus fasciatus_, the

subspecific name _olivaceogriseus_ is available.

The subspecies of _P. fasciatus_ are most easily distinguished by color

of pelage, in which there is a general cline from northeast (dark) to

southwest (pale). Cranially, the subspecies are less distinct. The

skulls of _P. f. callistus_ can be distinguished from those of the other

subspecies by several differences; however, among the other four

subspecies, only minor cranial differences are evident. Individual

variation was found to be greater than secondary sexual variation.

[Illustration: FIG. 1. Distribution of the five subspecies of

_Perognathus fasciatus_. Solid symbols represent specimens examined;

circled symbols represent type localities.

Guide to subspecies

1. _P. f. fasciatus_

2. _P. f. olivaceogriseus_

3. _P. f. infraluteus_

4. _P. f. litus_

5. _P. f. callistus_

]

In connection with this study, grateful acknowledgment is made to

the following persons for the loan of specimens in their care: Dr.

H. E. Anthony, American Museum of Natural History (AMNH); Mr. Colin

C. Sanborn, Chicago Natural History Museum (CNHM); Mr. Austin W.

Cameron, National Museum of Canada (NMC); Miss Viola S. Schantz,

Biological Surveys Collection, U. S. National Museum (USBS); Dr.

Seth B. Benson, University of California Museum of Vertebrate

Zoology (MVZ); Mr. J. E. Moore, Department of Zoology, University

of Alberta (UADZ); Dr. William H. Burt, University of Michigan

Museum of Zoology (MZ); Dr. Otis Wade, University of Nebraska

Department of Zoology (UNDZ); Dr. C. Bertrand Schultz, University

of Nebraska State Museum (NSM); and to those in charge of the

collections of the Nebraska Game, Forestation and Parks Commission

(NGFPC). Specimens from the Museum of Natural History, University

of Kansas (KU), also have been used.

Capitalized color terms are those of Ridgway, Color Standards and

Color Nomenclature, Washington, D. C., 1912. Assistance with field

work to obtain specimens for the University of Kansas Museum of

Natural History is acknowledged from the Kansas University

Endowment Association and the National Science Foundation.

=Perognathus fasciatus fasciatus= Wied-Neuwied

_Perognathus fasciatus_ Wied-Neuwied, Nova Acta Phys.-Med. Acad.

Caesar. Leop.-Carol., 19:369, 1839, type from Upper Missouri

River near its junction with the Yellowstone, near Buford,

Williams County, North Dakota.

_Geographic distribution._--Southwestern Manitoba, extreme

northeastern Montana and all of North Dakota except extreme eastern

part, south through central South Dakota to the northeastern part

of Cherry County, Nebraska (see Figure 1).

_Remarks._--For comparisons with _Perognathus fasciatus

olivaceogriseus_, geographically adjacent to the southwest, see account

of that subspecies.

Specimens from the eastern part of the range of _P. f. fasciatus_ are

somewhat smaller and slightly darker than topotypes from Buford, North

Dakota. Specimens from 9 mi. SE Bainville, Johnson Lake and 3 mi. S

Medicine Lake, Montana, and from Crosby, North Dakota, are also paler

than those from farther east. At the southern limit of the range of the

subspecies, specimens from the Rosebud Agency and Minichaduza River,

South Dakota, approach _olivaceogriseus_ in pale dorsal coloration.

_Specimens examined._--Total number, 83, as follows: MANITOBA:

Aweme, 3 (USBS 2, NMC 1). Junction of Antler and Souris rivers, 1

(NMC). Oak Lake, 2 (NMC). MONTANA: _Roosevelt County_: 9 mi. SE

Bainville, 4 (MZ); Johnson Lake, 2 (USBS). _Sheridan County_: 3 mi.

S Medicine Lake, 1880 ft., 1 (KU). NEBRASKA: _Cherry County_: Ft.

Niobrara Game Reserve, 1 (NSM); Sparks, 1 (MZ); Valentine, 2

(UNDZ). NORTH DAKOTA: _Benson County_: 2 mi. W Fort Totten, 1400

ft., 3 (KU). _Bottineau County_: Bottineau, 1 (CNHM). _Burleigh

County_: 9 mi. E Bismark, 7 (MZ). _Dickey County_: Oakes, 2 (CNHM

1, USBS 1). _Divide County_: Crosby, 1 (USBS). _Kidder County_:

Dawson, 1 (USBS); 6 mi. W Steele, 6 (MZ). _McHenry County_: 1/2 mi.

E Upham, 1 (USBS). _Morton County_: 12 mi. W Mandan, 2 (MZ).

_Oliver County_: Ft. Clark, 8 (USBS). _Sargent County_: 7-2/10 mi.

E, 1-2/10 mi. S Oakes, 1200 ft., 1 (KU). _Sioux County_: Cannon

Ball, 6 (USBS). _Stark County_: 1 mi. S Dickinson, 1 (MZ); 9 mi. W

Dickinson, 3 (MZ); 2 mi. W Taylor, 4 (MZ). _Stutsman County_: 7 mi.

N Jamestown, 1 (MZ); 14 mi. W Jamestown, 1 (MZ). _Wells County_:

Bowdon, 1 (USBS). _Williams County_: Buford, 9 (USBS). SOUTH

DAKOTA: _Todd County_: Minichaduza River, 2 (USBS); Rosebud Agency,

1 (USBS). _Tripp County_: Colome, 3 (MZ). _Walworth County_: Swan

Creek, 13 mi. S Shelby, 1600 ft., 1 (KU).

=Perognathus fasciatus olivaceogriseus= Swenk

_Perognathus flavescens olivaceogriseus_ Swenk, Missouri Valley

Fauna, 3:6, June 5, 1940, type from [Little Bordeaux Creek,

sec. 14, T. 33 N, R. 48 W, 3 mi. E] Chadron, Dawes County,

Nebraska.

_Geographic distribution._--Southeastern Alberta, southeast to

eastern Wyoming and adjacent parts of South Dakota and Nebraska.

_Diagnosis._--External measurements of the holotype and average and

extreme external measurements of six adults (five males and one

female) from several localities in eastern Wyoming are,

respectively, as follows: Total length, 124, 132 (125-140); length

of tail-vertebrae, 53, 63 (59-68); length of hind foot, 16.5, 17

(17-18); length of ear, 6.5, 6.6 (6.0-7.0). Color: Upper parts

Cream Buff, lined with black and giving a pale olivaceous

appearance; lateral line near (16'd) Light Ochraceous-Buff;

postauricular patches Cream Buff; subauricular patches and

underparts white; tail indistinctly bicolor, dusky above, whitish

below. Skull: Size medium for species (see Table 1); braincase and

auditory bullae moderately inflated; interorbital region narrow;

mastoidal region broad.

_Remarks._--From topotypes of _Perognathus fasciatus fasciatus_,

geographically adjacent to the northeast, _P. f. olivaceogriseus_

differs in: Upper parts paler, especially face which is heavily washed

with buff; hind foot smaller; skull averaging slightly smaller in all

measurements taken (especially least interorbital breadth and cranial

depth), except mastoidal breadth which is slightly more and

interparietal breadth, length of tympanic bulla and length of maxillary

tooth-row which are approximately the same. From topotypes of

_Perognathus fasciatus infraluteus_, geographically adjacent to the

south, _P. f. olivaceogriseus_ differs in: Upper parts paler and less

drab; lateral line brighter and more ochraceous; underparts never buffy;

skull averaging larger in all measurements taken except interparietal

breadth and length of maxillary tooth-row which are smaller. From

topotypes of _Perognathus fasciatus litus_, geographically adjacent to

the southwest, _P. f. olivaceogriseus_ differs in: Upper parts darker;

skull, when specimens of equal age are compared, averaging larger in

mastoidal breadth, interparietal breadth and basal length. From

topotypes of _Perognathus fasciatus callistus_, _P. f. olivaceogriseus_

differs in: Upper parts slightly darker, pelage not silky, coarser in

appearance; hind foot shorter; skull averaging smaller in all

measurements taken (especially mastoidal breadth), except interparietal

breadth which is more.

Swenk (_loc. cit._) originally described _P. f. olivaceogriseus_ as a

subspecies of _Perognathus flavescens_. I have examined the holotype,

kindly made available to me by Dr. Seth B. Benson of the Museum of

Vertebrate Zoology, University of California, and the paratype, in the

collection of the University of Nebraska State Museum. Neither is _P.

flavescens_, but both are clearly specimens of _P. Fasciatus_ on the

basis of olive-gray dorsal coloration, larger over-all dimensions of the

skull, and in that the tympanic bullae do not meet anteriorly. In

addition, a third specimen from Glen, Nebraska, purported by Swenk to be

intermediate between _flavescens_ and _olivaceogriseus_, has been

examined and found to be an immature _fasciatus_.

[Illustration: FIGS. 2-7. Skulls of two species of pocket mice. All x

1-1/2. In comparison with _P. flavescens_ note that _P. fasciatus_ has

the tympanic bullae separated anteriorly and that the skull is slightly

larger in over-all dimensions.

FIGS. 2-4. _Perognathus fasciatus fasciatus_. Buford, Williams County,

North Dakota, May 6, 1910, [** Male], No. 168599 USBS.

FIGS. 5-7. _Perognathus flavescens flavescens_, Kennedy, Cherry County,

Nebraska, July 9, 1935, [** Female], No. 66883 MVZ.]

In the original description Swenk indicated the type locality as

Chadron, Dawes County, Nebraska. The locality given on the specimen

label of the holotype, however, is 5 mi. E Chadron. In addition, Swenk

(_loc. cit._), in the paragraph preceding the description of

_olivaceogriseus_, states that the holotype was actually taken on Little

Bordeaux Creek, sec. 14, T. 33 N, R. 48 W, 3 mi. E Chadron, on the farm

of L. M. Gates, who obtained the holotype. It seems best to restrict the

type locality to the place last mentioned.

Four specimens from southeastern Alberta recently reported by Moore

(1953:143) have been examined and seem best referred, among known

subspecies of _fasciatus_, to _P. f. olivaceogriseus_. They differ from

the latter in several minor cranial features and more drab back, sides

and lateral line. When adequate material is available they may prove to

be subspecifically different from _olivaceogriseus_. Moore referred

these specimens to _P. f. fasciatus_.

Specimens from extreme northwestern Nebraska, including the holotype,

are slightly darker than specimens from eastern Wyoming. A specimen from

Ft. Steele, Wyoming, shows approach to _P. f. litus_ in pale dorsal

coloration.

_Specimens examined._--Total number, 59, as follows: ALBERTA:

Foremost, 2 (UADZ). Manyberries, 1 (Alberta Dept. Pub. Health).

Medicine Hat, 1 (Alberta Dept. Pub. Health). MONTANA: _Carter

County_: Little Missouri River, 8 mi. NE Albion, 1 (USBS). _Custer

County_: Calf Creek, 2 (AMNH); 13 mi. E Miles City, 1 (MZ).

_Garfield County_: Piney Buttes, 1 (USBS). _Powder River County_:

Powderville, 2 (USBS). _Yellowstone County_: Lake Basin, 1 (USBS).

_County undesignated_: Wolf's Creek, 1 (AMNH). NEBRASKA: _Cherry

County_: 10 mi. E Gordon, 1 (CNHM); 12 mi. ESE Gordon, 2 (CNHM).

_Dawes County_: Chadron State Park, 1 (NGFPC); 1 mi. SW Chadron, 1

(NSM); Little Bordeaux Creek, 3 mi. E Chadron, 1 (MVZ); 10 mi. S

Chadron, 3 (MZ); 3 mi. SW Crawford, 1 (KU). _Sheridan County_:

Mirage Township, 1 (MZ). _Sioux County_: Glen, 1 (NSM); Monroe

Canyon, 1 (NGFPC). SOUTH DAKOTA: _Custer County_: Cheyenne River, 1

(AMNH); Elk Mountain, 4800 ft., 4 (USBS). _Meade County_:

Smithville, 1 (USBS). _Shannon County_: Corral Draw, 3 (AMNH); Pine

Ridge, 3 (CNHM 1, USBS 2). _Washabaugh County_: White River flood

plain, 7 mi. S Kadoka, 1 (MZ). WYOMING: _Campbell County_: 1-1/4

mi. N, 1/2 mi. E Rockypoint, 3850 ft., 1 (KU). _Carbon County_: Ft.

Steele, 1 (USBS). _Fremont County_: 40 mi. E Dubois, 1 (MZ). _Hot

Springs County_: Kirby Creek, 5000 ft., 1 (USBS). _Johnson County_:

6-1/2 mi. W, 2 mi. S Buffalo, 5620 ft., 4 (KU); 1 mi. WSW Kaycee,

4700 ft., 1 (KU). _Natrona County_: Casper, 1 (USBS); 1 mi. NE

Casper, 5150 ft., 1 (KU). _Platte County_: 2-1/2 mi. S Chugwater, 1

(KU). _Sheridan County_: Arvada, 2 (USBS); 5 mi. NE Clearmont, 3900

ft., 1 (KU). _Weston County_: Newcastle, 1 (USBS); 23 mi. SW

Newcastle, 4500 ft., 4 (KU).

=Perognathus fasciatus infraluteus= Thomas

_Perognathus infraluteus_ Thomas, Ann. Mag. Nat. Hist., ser. 6,

11:406, May, 1893, type from Loveland, Larimer County,

Colorado.

_Perognathus fasciatus infraluteus_, Osgood, N. Amer. Fauna, 18:19,

September 20, 1900.

_Geographic distribution._--Known only from the type locality and

from 7 mi. N Ramah, Colorado. Probably in other parts of

northeastern Colorado (see Figure 1).

_Remarks._--For comparisons with _Perognathus fasciatus olivaceogriseus_

Swenk, geographically adjacent to the north, see the preceding account

of that subspecies on page 520.

Of the seven topotypes available to me, six show various degrees of

encroachment of the color of the lateral line upon the underparts,

giving the ventral surface a buffy appearance, the "distinguishing

character" of the subspecies according to Osgood (_op. cit._:20). When

additional specimens are available, this character may be found to be

one of individual variation, although no specimens from other parts of

the range of the species have been found to exhibit it. Of the

subspecies _P. f. infraluteus_, each of the eight specimens examined by

me possesses buffy subauricular patches.

The specimen from 7 mi. N Ramah, Colorado, is slightly darker and less

drab than the topotypes and does not possess buffy underparts. The skull

of the specimen is broken but on the basis of characters of the pelage

it seems to be intermediate between _infraluteus_ and _olivaceogriseus_

and is referred to the former on geographic grounds.

_Specimens examined._--Total number, 8, as follows: COLORADO:

_Elbert County_: 7 mi. N Ramah, 1 (MZ). _Larimer County_: Loveland,

7 (USBS).

=Perognathus fasciatus litus= Cary

_Perognathus fasciatus litus_ Cary, Proc. Biol. Soc. Wash., 24:61,

March 22, 1911, type from Sun, Sweetwater Valley, Natrona

County, Wyoming.

_Geographic distribution._--Lower Sweetwater Valley and adjacent

parts of Red Desert in Carbon, Fremont, Natrona and Sweetwater

counties, Wyoming (see Figure 1).

_Remarks._--For comparisons with _Perognathus fasciatus

olivaceogriseus_, geographically adjacent to the northeast, see account

of that subspecies. From _Perognathus fasciatus callistus_,

geographically adjacent to the southwest, _P. f. litus_ differs in:

Upper parts paler with no trace of olivaceous; hind foot shorter; skull,

when specimens of equal age are compared, averaging smaller in all

measurements taken (especially occipitonasal length, mastoidal breadth,

length of tympanic bulla and cranial depth), except interparietal

breadth which is more.

No fully adult specimens of _P. f. litus_ have been available to me for

this study. Two of the three specimens mentioned by Cary in the original

description (holotype not seen) have been examined and found to possess

adult pelage, but cranially they must be classed as young adults. These

specimens are paler than those of any other subspecies of _fasciatus_

and do not have the olivaceous dorsal coloration present in other

subspecies.

_P. f. litus_ is seemingly an endemic race in the lower Sweetwater

Valley and adjacent parts of the Red Desert, Wyoming. The type locality,

recorded by Cary in the original description as "Sun, Sweetwater Valley,

Wyoming," is here placed in Natrona County on the basis of the map

(frontispiece) in Cary's (1917) "Life Zone Investigations in Wyoming."

_Specimens examined._--Total number, 9, as follows: WYOMING:

_Carbon County_: 8 mi. SE Lost Soldier, 6700 ft., 1 (USBS).

_Fremont County_: Granite Mts., 1 (MZ). _Natrona County_: 5 mi. W

Independence Rock, 6000 ft., 4 (KU); Sun, 1 (USBS); 16 mi. S, 11

mi. W Waltman, 6950 ft., 1 (KU). _Sweetwater County_: 27 mi. N

Table Rock, 1 (MZ).

=Perognathus fasciatus callistus= Osgood

_Perognathus callistus_ Osgood, N. Amer. Fauna, 18:28, September

20, 1900, type from Kinney Ranch, near Bitter Creek, Sweetwater

County, Wyoming.

_Geographic distribution._--East of the Green River in central and

southern parts of Sweetwater County, Wyoming, and adjacent parts of

Moffat County, Colorado (see Figure 1).

_Remarks._--For comparisons with _Perognathus fasciatus litus_,

geographically adjacent to the north, see account of that subspecies.

When Osgood (_loc. cit._) described _P. f. callistus_ as a full species,

he characterized it as having " ... the attractive coloration of

_fasciatus_, but softer and more delicate. Its position is evidently

between _fasciatus_ and _apache_, and its nearest relations are clearly

with the latter." He further remarked that, "Its large size immediately

separates it from _fasciatus_ which it resembles externally, especially

before maturity."

Comparison of the average and extreme external measurements of ten adult

_P. f. callistus_ from Sweetwater County, Wyoming, with adult

individuals of other subspecies of _P. fasciatus_ shows that _callistus_

does not average larger than _P. f. olivaceogriseus_ except in length of

hind foot and that it averages slightly smaller in all external

measurements than topotypes of _P. f. fasciatus_. Moreover, judging from

the accounts of Goldman (1918:24) and Durrant (1952:235), _Perognathus

apache caryi_ (the subspecies of _P. apache_ nearest to the range of _P.

f. callistus_) is significantly larger externally and has no trace of

olivaceous in the dorsal pelage.

Comparison of the skulls of _callistus_ with a skull of _Perognathus

apache apache_ from Wingate, New Mexico (USBS 137388), reveals the

following differences: Interparietal bone wider in _callistus_,

averaging 4.5 (as opposed to 4.0) and more pentagonal; lacrimal bone

shorter and stouter in _callistus_; tympanic bullae more inflated in

_callistus_; interorbital foramina larger in _callistus_; lower premolar

approximately the same size as the last lower molar in _callistus_,

approximately half the size of the last lower molar in _apache_.

Conversely, comparison of skulls of _callistus_ with those of _P.

fasciatus_ as concerns the above mentioned features reveals that they

closely resemble each other. In view of this resemblance it seems best

to arrange _callistus_ as a subspecies of _P. fasciatus_.

_P. f. callistus_ is distinct cranially from all other subspecies of _P.

fasciatus_ in the narrowness of the interparietal, the greater length of

the tympanic bulla and the greater mastoidal breadth. The fine, silky

nature of the pelage is shared, to some extent, with _P. f. litus_.

A specimen from 27 mi. N, 37 mi. E Rock Springs, Wyoming, referred to

_callistus_ resembles _litus_ in pale dorsal coloration and slightly

wider interparietal. Four immature specimens from 25 mi. N, 38 mi. E

Rock Springs, also referred to _callistus_, have extremely pale juvenal

pelage and also are judged to be intergrades with _litus_.

_Specimens examined._--Total number, 23, as follows: WYOMING:

_Sweetwater County_: 18 mi. S Bitter Creek, 6800 ft., 3 (KU);

Kinney Ranch, 21 mi. S Bitter Creek, 6800 ft., 9 (KU 7, USBS 2); 30

mi. S Bitter Creek, 2 (KU); 33 mi. S Bitter Creek, 6900 ft., 2

(KU); 32 mi. S, 22 mi. E Rock Springs, 7025 ft., 2 (KU); 25 mi. N,

38 mi. E Rock Springs, 6700 ft., 4 (KU); 27 mi. N, 37 mi. E Rock

Springs, 6700 ft., 1 (KU).

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