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Title: Geographic Variation in the Pocket Gopher, Thomys bottae, in Colorado

Author: Phillip M. Youngman

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University of Kansas Publications
Museum of Natural History

Volume 9, No. 12, pp 363-384, 7 figs, in text, 1 table
February 21, 1958

Geographic Variation
in the Pocket Gopher, Thomomys bottae,
in Colorado

BY
PHILLIP M. YOUNGMAN

University of Kansas
Lawrence
1958

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 9, No. 12, pp. 363-384, 7 figs. in text, 1 table
Published February 21, 1958

University of Kansas
Lawrence, Kansas

PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1958

27-1765

Geographic Variation in the Pocket Gopher, Thomomys bottae, in Colorado

BY
PHILLIP M. YOUNGMAN

INTRODUCTION

Two species of pocket gophers of the genus Thomomys (Family Geomyidae) occur in Colorado, Thomomys bottae (see fig. 1) in the low valleys in the south-central and southwestern parts of the state and Thomomys talpoides mainly in the mountains and high valleys.

Thomomys bottae occurs primarily in the Piñon-juniper, Ponderosa Pine, and Short Grass zones of Daubenmire (1943) but in some localities is found in the Douglas Fir Zone. Thomomys talpoides occupies primarily the Douglas Fir Zone and Engelmann Spruce-Subalpine Spruce Zone but is found also in the Piñon-juniper and Short Grass zones in some localities.

The ranges of the two species do not overlap in the strict sense but interdigitate in a parapatric type of distribution.

Two other pocket gophers, Geomys bursarius and Cratogeomys castanops, also occur in Colorado—in the Upper Sonoran Life-Zone. Geomys bursarius occupies much of the Great Plains, whereas Cratogeomys castanops is found only on the plains in the southeastern part of the state.

The objectives of the study, reported on here, were to learn the geographic distribution of Thomomys bottae in Colorado, to find means for recognizing the different subspecies, and to describe individual and geographic variation.

I am indebted to Mr. Sydney Anderson and Professor E. Raymond Hall for many helpful suggestions and for their critical reading of the manuscript, to Dr. Richard S. Miller, who made the collection of many of the specimens possible, and to Dr. Richard M. Hansen for numerous suggestions. I wish to express my appreciation also to the following for the loan of specimens in their care: Alfred M. Bailey and A. A. Rogers, Colorado Museum of Natural History, Denver, Colorado; David H. Johnson, United States National Museum, Washington, D. C; Robert W. Lechleitner, Colorado State University, Fort Collins, Colorado; and Robert Z. Brown, Colorado College, Colorado Springs, Colorado.

METHODS

Adults of approximately equal age were compared in the study of geographic variation. Three criteria of adulthood are: (a) suture obliterated between supraoccipital and exoccipital, (b) suture at least partly obliterated between basisphenoid and basioccipital, (c) supraorbital crests not widely separated and almost parallel. In males the crests encroach on the lateral borders of the interparietal; in females the crests approach the lateral borders of the interparietal but are more widely separated than in males.

In studying geographic variation, greater emphasis was placed on females than on males. As noted by Grinnell (1931:4), males vary more than females, especially in length of rostrum and associated nasal measurements.

Color terms are those of Munsell (1954). Color measurements were standardized by the use of a single 100 watt General Electric blue daylight bulb in a 12 inch white reflector suspended 24 inches above the specimen. All other light was excluded. The individual hairs of Thomomys bottae are either bicolored or tricolored. The darkness of a specimen often may be attributed to the presence of dark-tipped hairs. The color given in the description is the basic reddish or yellowish color of the hairs. The presence of a grizzled effect or a dark dorsal stripe, or any other pattern resulting from dark hairs, is noted in the remarks.

[pg 367] Specimens examined are listed by counties in the following order:

• Mesa
• Montrose
• San Miguel
• Dolores
• Montezuma
• La Plata
• Archuleta
• Conejos
• Chaffee
• Fremont
• El Paso
• Pueblo
• Custer
• Huerfano
• Alamosa
• Las Animas

Localities are listed from north to south within a county. If two localities lie on the same line of latitude, the western precedes the eastern. Localities omitted on the map in order to prevent overlapping of symbols are in Italics. Unless otherwise indicated, specimens are in the University of Kansas, Museum of Natural History. The following initials are used to designate specimens in other collections:

The following measurements of the skull are listed in the tables:

Condylobasal length.—The shortest distance between the anteriormost projections of the premaxillaries and a line touching the posterior surfaces of the exoccipital condyles.

Length of nasals.—The distance from the most anterior projection of the nasal bones to the most posterior projection of a nasal bone.

Zygomatic breadth.—The greatest distance across the zygomatic arches, at right angles to the long axis of the skull.

Squamosal breadth.—The greatest distance between the mastoidal processes of the squamosal.

Length of rostrum.—The shortest distance from the shallow notch that lies lateral to the hamulus of the lacrymal bone, to the tip of the nasal on the same side of the skull.

Breadth of rostrum.—The greatest width of the rostrum, anterior to the maxillae, transverse to the long axis of the skull.

Alveolar length of upper maxillary tooth-row.—Distance between the anterior margin of the alveolus of the first cheek-tooth and the posterior margin of the alveolus of the last upper cheek-tooth, on one side of the skull.

Least interorbital breadth.—The least distance across the frontal bones at the interorbital constriction as seen in dorsal view.

PHYSIOGRAPHY

Thomomys bottae occurs in the Colorado Plateau Province (terminology of Fenneman, 1931), the Southern Rocky Mountain Province and a small part of the Great Plains Province.

The Colorado Plateau Province, in the southwestern part of the state, is mostly above 5000 feet and is characterized by the great number of canyons cut by rivers and streams in the nearly horizontal strata. Prominent features of the landscape are cuestas, such as Mesa Verde, and laccoliths, such as Ute Peak.

The Southern Rocky Mountain Province consists mainly of high granitic mountains running north and south, many of which extend [pg 368] to more than 14,000 feet above sea level. Included in this region are several large basins, such as North Park and South Park and the San Luis Valley. The San Juan Mountains, which separate the Colorado Plateau Province from the San Luis Valley, and the Sangre De Cristo and Wet mountains, which intervene between the San Luis Valley and the Great Plains, importantly influence the distribution of Thomomys bottae.

The Great Plains Province is a broad highland that slopes gradually eastward from the Rocky Mountains. Of importance to the present study are two subdivisions of the Great Plains, the Colorado Piedmont and the Raton Section.

The Colorado Piedmont is a much dissected fluviatile plain, roughly extending from the vicinity of the Arkansas River to the northern boundary of the state. In general the topography of the Colorado Piedmont is broadly rolling with greater relief than the high plains to the east; however, buttes and steep bluffs occur locally.

The Raton Section imperceptibly blends into the southern boundary of the Colorado Piedmont and extends south into New Mexico and Texas. A trenched peniplane of greater relief and altitude than the Colorado Piedmont, it is characterized by high mesas, extensive dissected lava-capped plateaus, deep canyons, and mountains of volcanic origin.

GEOGRAPHIC VARIATION

Six subspecies of Thomomys bottae occur in Colorado. T. b. aureus and T. b. howelli occupy the Colorado Plateau Province (see fig. 1) and are characterized by a yellowish color; nasals posteriorly truncate or rounded; posterior extensions of premaxillae long; basioccipital wide; and interpterygoid space U-shaped with a median spicule.

T. b. internatus, T. b. cultellus, and a new subspecies from the vicinity of Cañon City described on page 376, inhabit the Sangre De Cristo and Wet mountains in the Southern Rocky Mountain Province and adjacent parts of the Colorado Piedmont and Raton Section of the Great Plains Province (see fig. 1). This group of closely related subspecies is characterized by reddish color; posterior margins of nasals forming a V; posterior extensions of premaxillae short; basioccipital narrow; and interpterygoid space V-shaped, lacking a median spicule.

T. b. pervagus occupies part of the San Luis Valley to the west of the Rio Grande (see fig. 1). In Colorado T. b. pervagus is iso [pg 369]lated from T. b. internatus and T. b. cultellus by the Sangre De Cristo and Culebra ranges and is separated from T. b. aureus by the San Juan Mountains. T. b. pervagus occupies an area geographically intermediate between T. b. aureus to the west and T. b. internatus and T. b. cultellus to the east and has some characters in common with these subspecies. T. b. pervagus resembles T. b. aureus in having long posterior extensions of the premaxillae and in sometimes having rounded posterior margins of the nasals. T. b. pervagus resembles T. b. internatus and T. b. cultellus in color, the presence of a V-shaped interpterygoid space, and a narrow basioccipital. Kelson (1951:69) has pointed out that in New Mexico the separation of the ranges of T. b. pervagus and T. b. cultellus is probably complete, but probably incomplete between T. b. pervagus and T. b. aureus. Nevertheless, the similarities between T. b. pervagus and T. b. cultellus and T. b. internatus suggest that T. b. pervagus was originally derived from the more eastern stock.

T. b. aureus is a variable subspecies which, according to Durrant (1952:211), intergrades with T. b. howelli in Utah. Specimens of T. b. aureus showing the greatest amount of geographic variation cranially are from the ecotone between the Piñon-juniper and Douglas Fir zones at the edge of the range of the subspecies.

T. b. howelli is a markedly distinct subspecies that shows certain similarities to T. b. aureus, but the degree of cranial difference from T. b. aureus suggests an isolation of long duration, or a rapid evolution from the parent stock.

T. b. internatus and T. b. cultellus probably intergrade east of the Sangre De Cristo Range in the vicinity of the Colorado-New Mexico boundary. The amount of intergradation is obscured by the great amount of geographic variation occurring in T. bottae at the edge of the plains and by the lack of specimens from this area.

T. b. internatus is a widespread subspecies showing its greatest variation at the edge of the plains. This area is an ecotone between the coniferous forest and the grassland and is by nature an area of change owing to the alternation of wet and dry periods such as the pluvial, interpluvial, and postpluvial periods. This seems to support Durrant's observation (1952:496) that "the greatest range of morphological variation is in animals from the least stable environments."

Specimens from a small area north of the Arkansas River in the vicinity of Cañon City (see fig. 1) differ sufficiently from T. b. internatus to be given nominal recognition. High mountains and the Arkansas River isolate the new subspecies found at Cañon City from [pg 370] populations of T. b. internatus to the west and south; however there are no apparent geographic barriers between the newly named subspecies and populations of T. b. internatus twelve miles to the north or from the vicinity of Pueblo to the east. This new subspecies is the most extreme of the variants occurring in the unstable environment at the edge of the plains.

Thomomys bottae aureus Allen

Remarks.—T. b. aureus is a variable subspecies, which differs considerably from T. b. internatus, T. b. cultellus, and T. b. rubidus and includes several microgeographic races distinguishable to a taxonomist specializing in the group. These slightly varying populations are here not considered sufficiently distinct for nominal recognition.

Characters such as color of the pelage and conformation of the bullae and zygomatic arches vary with the locality, and to some extent vary among specimens from a single locality.

The name Thomomys bottae optabilis, given to specimens from [pg 371] Coventry by Goldman (1936:116), is here placed in synonymy under T. b. aureus Allen. The characters originally used to describe T. b. optabilis are of the type that vary between populations only a few miles apart, or often vary within a population. The skulls of specimens from Coventry are not lighter in structure than those of T. b. aureus. The premaxillae are not narrower, nor is the frontal region narrower or more constricted than in T. b. aureus.

The name Thomomys bottae apache, given to specimens from Lake La Jara, New Mexico, by Bailey (1910:79), and later applied to specimens from Colorado by Bailey (1915:75), is here also placed in synonymy under T. b. aureus. Specimens from Lake La Jara, New Mexico, and nearby localities in Colorado may be separated from topotypes of T. b. aureus on the basis of color only. The topotypes of T. b. aureus are mostly pale; some, however, are dark. The number of pale specimens in any given series decreases gradually in a clinal pattern from west to east. Since there is no noticeable step in the cline and since all specimens show close cranial similarity, it is felt that nominal recognition of the darker specimens does not present a realistic picture of the relationships of the relatively unisolated populations in the Colorado Plateau Province.

Since Thomomys bottae in the Colorado Plateau Province is especially plastic, varying from locality to locality, emphasis is here placed on similarities that unite specimens from different localities. The individual and microgeographic variations are outlined below.

Specimens from Bedrock have zygomatic arches that are heavy anteriorly. Specimens from Coventry are dorsally almost uniformly Strong Brown (7.5YR 5/6) and lack a strong dorsal stripe. The venters are Reddish Yellow (7.5YR 8/6). Specimens from 15 miles west of Cortez are the palest specimens of T. b. aureus from Colorado, and closely resemble topotypes. The basic color varies from Reddish Yellow (7.5YR 7/6 and 6/6) to Strong Brown (7.5YR 5/6). Specimens are marked with a narrow dark dorsal stripe. The venters are white. Specimens from Ute Peak and Cortez have Reddish Yellow (7.5YR 6/6) flanks and are slightly darker dorsally. Many specimens from Mesa Verde are indistinguishable from specimens from Coventry and from Cortez. Others have dark diffuse dorsal stripes. The venters are Pink (7.5YR 7/4) or Pinkish White (7.5YR 8/2). Some specimens from the Mancos River have wide dorsal stripes. Specimens from three miles west of Durango have especially wide-spreading zygomatic arches posteriorly and have wide black dorsal stripes. The venters are Pink (7.5YR 7/4). One specimen from Florida is dark and grizzled and has a dark dorsal [pg 372] stripe. Another specimen is pale and has only a small dorsal stripe. Specimens from 12 miles west of Pagosa Springs have thin rostra and diffuse dorsal stripes. Specimens from Bondad have a V-shaped interpterygoid space and in it a small median spicule. One specimen is uniformly grizzled and lacks a dorsal stripe, giving an overall effect of Dark Yellowish Brown (10YR 3/3). Another specimen has Strong Brown (7.5YR 5/6) flanks and is only slightly darker dorsally.

Thomomys bottae howelli Goldman

Remarks.—T. b. howelli most closely resembles T. b. aureus; however, since only one adult specimen of T. b. howelli is known, it is impossible to appraise adequately its characters. Durrant (1952:211) records intergradation between T. b. howelli and T. b. osgoodi, and between T. b. howelli and T. b. aureus in Utah.

[pg 373] An attempt to collect specimens of T. b. howelli, in March, 1957, by Richard S. Miller and the writer was unsuccessful.

Thomomys bottae pervagus Merriam

Remarks.—T. b. pervagus is a well-defined subspecies. There is little variation between the topotypes and specimens from Colorado.

Thomomys bottae internatus Goldman

Remarks.—The dividing line between T. b. internatus and T. b. cultellus is drawn arbitrarily since only one specimen has been collected between La Veta Pass and the border of New Mexico.

When Goldman (1936:115) named T. b. internatus he included specimens from Union and Colfax counties, New Mexico, and specimens from Gardner, Colorado (not Garfield as stated by Kelson, 1951:66). The specimens from New Mexico and a specimen from Fishers Peak, Colorado, were subsequently assigned to T. b. cultellus by Kelson (loc. cit.).

The specimen from Fishers Peak shows some characters that might be interpreted as intermediate between internatus and cultellus, but shows also some unique characters that can be understood only by further collecting in the regions north and northeast of the type locality of T. b. cultellus.

Variation is slight in the large series of topotypes of T. b. internatus. Specimens from other localities in the western part of the range differ little from the topotypes. Specimens from one mile west of Coaldale have slightly more inflated bullae that are more flattened ventrally. Specimens from five miles south of Cotopaxi also have the bullae more flattened ventrally.

Specimens from localities bordering the plains differ from the topotypes and near topotypes, and in general show greater variation from locality to locality. Specimens from 12 miles north of Cañon City are dark, resembling T. b. rubidus, but cranially agree with specimens from near Colorado Springs in being indistinguishable from specimens from Salida. Specimens from St. Charles Mesa and Bear Creek near Walsenburg differ from the topotypes in having wider rostra. The specimens from St. Charles Mesa have more inflated bullae.

Thomomys bottae cultellus Kelson

Remarks.—Kelson (1951:64) named T. b. cultellus on the basis of six dark specimens (Dark Reddish Brown 5YR 3/4 and 2/2). Nowhere else within the range of this subspecies, as defined by Kelson, do any specimens resemble the topotypes in color.

After comparing topotypes of T. b. cultellus with topotypes of T. b. internatus of approximately equal age, I disagree with Kelson (loc. cit.) on some of the characters which he used to separate cultellus from internatus. My findings indicate that T. b. cultellus is not smaller, that its skull is not smaller and not less angular, and that the tympanic bullae are not less pointed ventrally. Further collecting is needed better to limit and diagnose this subspecies.

Figs. 2-7. Dorsal views of skulls of Thomomys bottae. × 1.

Fig. 2. Thomomys b. aureus, 3 mi. W Durango, La Plata Co., Colorado. No. 72967, ♀.

Fig. 3. Thomomys b. howelli, holotype, Grand Junction, 4600 ft., Mesa Co., Colorado. No. 75684 USNM, ♀.

Fig. 4. Thomomys b. pervagus, Espanola, 5000 ft., Rio Arriba Co., New Mexico. No. 133614 USNM, ♀.

Fig. 5. Thomomys b. cultellus, Fishers Peak, 8000 ft., Las Animas Co., Colorado. No. 129285 USNM, ♀.

Fig. 6. Thomomys b. internatus, Salida, 7050 ft., Chaffee Co., Colorado. No. 2757 ERW, ♀.

Fig. 7. Thomomys b. rubidus, holotype, 2-9/10 mi. E Cañon City, Fremont Co., Colorado. No. 72954, ♀.

Thomomys bottae rubidus new subspecies

Remarks.—The range of T. b. rubidus is surrounded by the range of T. b. internatus; nevertheless, intergradation has not been found. For a discussion of the geographic relation of T. b. rubidus to T. b. internatus see page 374.

SUMMARY

A study of 249 specimens of Thomomys bottae from Colorado reveals six subspecies in the state. T. b. aureus and T. b. howelli occupy the Colorado Plateau Region in the western and southwestern parts of the state. T. b. internatus, T. b. cultellus, T. b. pervagus, and the newly named T. b. rubidus occupy part of the Southern Rocky Mountain Region and a narrow strip of the Great Plains.

The greatest amount of geographic variation, in Thomomys bottae in Colorado, occurs in the ecotone between the grassland and coniferous forest at the edge of the Great Plains, and in the ecotone between the Piñon, juniper, and sage of the Colorado Plateau and the Coniferous forest of the southern Rocky mountains.

Table 1. Measurements, in Millimeters, of Thomomys bottae

Unless otherwise noted, specimens are adults from Colorado

LITERATURE CITED

Bailey, V.

Daubenmire, R. F.

Durrant, S. D.

Fenneman, N. M.

Goldman, E. A.

Grinnell, J.

Kelson, K. R.

Munsell, A. H.

Transmitted November 14, 1957.

□
27-1765

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