Fig. 2. Probable distribution of the Subfamily Geomyinae in the early Pleistocene (late Blancan), depicting major areas of differentiation of the modern genera.

1. Thomomys
2. Geomys
3. Zygogeomys
4. Pappogeomys
5. Orthogeomys

Thomomys

The earliest Pleistocene records of Thomomys are mostly isolated teeth. Although they can be identified as genus Thomomys, most of the materials are too fragmentary to be identified to species. In Thomomys two distinct patterns of occlusal surfaces of the molars can be recognized: the generalized elliptical pattern in the subgenus Pleisothomomys, not unlike the pattern in other geomyids, and the pear-shaped pattern in the subgenus Thomomys, which results from constriction of the upper molars on the labial side and constriction of the lower molars on the lingual side. Some fossils assigned to Thomomys were not examined with this distinction in mind by the persons who made the assignments. Consequently some of the identifications now in the literature may be subject to change.

Three occurrences of Thomomys are from the early and middle Pleistocene, with a possible fourth (depending upon the age of the Hay Springs local fauna of Nebraska). The earliest Pleistocene record is from the Broadwater-Lisco beds along the North Platte River in Morrill County, western Nebraska. Possibly the specimen from there was misidentified. Those beds are Lower Pleistocene, and are regarded by Schultz and Stout (1948:560-561, 573) and by Hibbard (1958:11), as having been deposited mostly during the Aftonian interglacial. There is also some indication that some of the strata were deposited late in the Nebraskan glaciation. There are no other early Pleistocene records of Thomomys. Savage (1951:228) reported the genus from the Irvington local fauna, Alameda County, California. The specimens were not identified to species, although they were described as indistinguishable from Thomomys bottae. Paulson (1961:137) recorded specimens from the Cudahy local fauna, Meade County, Kansas. These fragmentary specimens are referable to the subgenus Thomomys, owing to the strong constriction of the molars, but have not been identified to species. The Cudahy is an Irvingtonian local fauna, and is considered to have been deposited during the late Kansan glaciation. The stratum containing the Cudahy local fauna immediately underlies the Pearlette Ash. The Cudahy material includes five isolated molars and a fragmentary ramus bearing only the premolar. The genus Thomomys has been recovered also from the Hay Springs local fauna in Sheridan County, northwestern Nebraska, by Shultz and Tanner (1957:71). The Hay Springs local fauna is considered to have been deposited in late Kansan glaciation or in early Yarmouth interglacial by Shultz and Tanner (op. cit.:69), or of Irvingtonian age; however, Hibbard (1958:25) regarded the beds containing this fauna as Illinoian (thus post-Irvingtonian in age), and equivalent in age to the Berends local fauna of Oklahoma and the Butler Springs and Mt. Scott local faunas of Kansas. The Thomomys from Hay Springs local fauna has not been referred to species.

The relative abundance of Geomys, and rarity of Thomomys, in Great Plains fossil beds of early and middle Pleistocene is probably due to allopatric distributions of the two genera. The Great Plains area was evidently the center of distribution and differentiation of Geomys. Perhaps Thomomys evolved earlier to the west, in the Great Basin and Pacific Coastal regions, and not on the Great Plains.

Upper Pleistocene records of Thomomys are more common. The genus was widespread in beds identified with the Illinoian and Sangamon and extended its range eastward to the Atlantic Coast. Stephens (1960:1961) reported Thomomys from the Doby Springs local fauna, Harper County, northwestern Oklahoma. The material (34 isolated teeth) was too fragmentary to permit assignment to species. The molars are constricted on one side, indicative of the subgenus Thomomys, like the Cudahy specimens reported by Paulson (see discussion above). Stephens erroneously mentioned that the enamel plate on the posterior face of the upper premolar is unique in Thomomys; this plate occurs also in Zygogeomys. The Doby Springs local fauna was recovered from beds that have been identified as Illinoian deposits, and it is correlated with the Berends local fauna in Beaver County, Oklahoma, and the Butler Springs local fauna in Meade County, Kansas (see Stephens, op. cit.: 1700).

Local faunas in Maryland and Florida of Rancholabrean age include Thomomys, in every instance referable to the subgenus Pleisothomomys on the basis of unconstricted molars. Thomomys potomacensis (Gidley and Gazin, 1933), from Cumberland Cave local fauna, Allegany County in western Maryland, is the type of the genus Pleisothomomys Gidley and Gazin (1933:354). Pleisothomomys is here regarded as a subgenus. The material used in the original description included four lower jaws, one with a complete dentition. Hibbard (1958:25) pointed out that the Cumberland Cave assemblage is a composite fauna including both glacial and interglacial forms. He placed the stratigraphic position of the fauna as definitely Upper Pleistocene, probably deposited in both Illinoian glaciation and during the Sangamon interglacial. T. potomacensis is significantly larger than T. orientalis Simpson (1928:6), from the Saber-tooth Cave local fauna, Citrus County, Florida. Simpson's material included a rostral fragment with an incisor, premolar, and first molar. The Saber-tooth Cave local fauna is regarded by Kurten (1965:219) as having been recovered from Sangamon deposits. Thomomys is unknown from Wisconsin deposits in the eastern United States, and today the genus does not occur east of the Great Plains.

Thomomys of Rancholabrean provincial age from the western United States and México is known only from Wisconsin beds.