Part 5

Zygogeomys

Remains found in the Curtis Ranch local fauna, Cochise County, in southeastern Arizona are regarded as of middle Pleistocene age. See Gazin (1942:481-484), Wilson (1937:39-40), Hibbard (1958:25), and Hibbard et al. (1965:510-511). Although some question as to the exact age of the Curtis Ranch local fauna still seems to exist, most authorities on the Pleistocene agree that the age is not Pliocene and that it is older than Rancholabrean. Gidley (1922:122) described the pocket gopher found in the Curtis Ranch beds as Geomys parvidens, which is preoccupied by Geomys parvidens Brown (1908:194), a name proposed for the pocket gopher from the Conard Fissure of Arkansas; therefore, Hay (1927:136) proposed the name Geomys persimilis for the Curtis Ranch species to replace Geomys parvidens Gidley. Geomys persimilis Hay became the type species of Gazin's genus Nerterogeomys (1942:507). In this paper, Nerterogeomys is considered to be a junior synonym of Zygogeomys.

Zygogeomys persimilis is represented by a rostral fragment bearing all the cheek teeth on the left side and the upper incisors. In addition, two lower jaws, one with the first three cheek teeth, are referred to the species (see Gazin, 1942:507). The fossils identified as Geomys from the Arroyo San Francisco, Cedazo fauna, in Aguascalientes, México, by Mooser (1959:413) may be referable instead to Zygogeomys. I have not seen the specimens and no figures are available; Mooser states that a cranium was recovered. If either the upper premolar or third molar is in place, generic identification could be made with reasonable certainty. No other fossils of Zygogeomys have been uncovered in late Pleistocene deposits and the significance of the absence of Zygogeomys has been discussed in an earlier paragraph of this section. Geomys has not been found so far south as Aguascalientes, but Zygogeomys occurs farther south now and presumably had a more extensive range on the plateau to the north in the Pleistocene.

Geomys

Geomys is common in Pleistocene deposits, especially on the Great Plains. Certainly the center of differentiation for Geomys was in this region, although at times, probably when conditions were favorable, Geomys expanded its range into adjacent areas, reaching the Pacific Coast in Irvingtonian times and the Atlantic Coast at the time of the Illinoian glaciation. The earliest Pleistocene records of the genus are from the Great Plains. McGrew (1944:49) described Geomys quinni from the Sand Draw local fauna, Brown County, Nebraska, considered by Hibbard (1958:11) to be Nebraskan in age. As mentioned in the account of Pliocene geomyids, Geomys quinni occurs also in the late Pliocene deposits of southwestern Kansas. Also, Geomys quinni occurs in the Broadwater-Lisco local fauna of Morrill and Garden counties, western Nebraska (Barbour and Schultz, 1937:3; Schultz and Stout, 1948:560-563; Schultz et al., 1951: table 1). The Broadwater-Lisco is currently regarded as Aftonian deposits (Schultz and Stout, loc. cit.; Hibbard, 1958:11). Hibbard (1956:174) identified Geomys quinni from the Deer Park local fauna, probably deposited during the early Aftonian interglacial, of Meade County, Kansas. Strain (1966:36) described Geomys paenebursarius on the basis of fossils obtained from early Pleistocene deposits of the Hudspeth local fauna from western Hudspeth County in the Trans-Pecos of Texas. The Hudspeth fossils were probably deposited during the Aftonian interglacial. From Kingman County, Kansas, Hibbard (op. cit.: 164) recovered isolated teeth of Geomys from the Dixon local fauna, regarded by him (op. cit.:153-154) as deposited during the latest Nebraskan glaciation, and correlated by him with the Sand Draw local fauna of Nebraska. Hibbard (1958:11) later regarded the Dixon as a transitional fauna between Nebraskan and Aftonian. The remains of Geomys from the Dixon are known only from isolated teeth. The teeth are small, and suggest that a smaller species of Geomys may have occurred along with the more common and larger G. quinni during the early Pleistocene (see discussion beyond of the Saunders Geomys). Geomys quinni was widespread and common throughout the central Great Plains from the late Pliocene (Rexroad fauna) through the early Pleistocene (Nebraskan and Aftonian deposits).

Hibbard (1956:179) referred the pocket gopher remains taken from the Saunders local fauna in Meade County, Kansas, to Geomys tobinensis, a small species having continuous enamel bands around the lower premolar in younger specimens. The Saunders local fauna was deposited in the late Aftonian and is younger than the Deer Park local fauna discussed above. Paulson (1961:138) later pointed out that the Saunders Geomys is distinct from Geomys tobinensis; hence, the small pocket gopher from the Saunders local fauna is probably an unnamed species, perhaps more closely allied to paenebursarius than to quinni. The small Geomys reported from the Aftonian Broadwater-Lisco local fauna of Nebraska (Schultz and Stout, 1948:563) may also be the same as the Saunders pocket gopher, but the smaller adult specimens occurring in the same bed with larger specimens probably are females and the larger specimens males. In all living Geomyini females have smaller skulls than males.

The Irvingtonian provincial age is currently regarded as Middle Pleistocene and includes the late Kansan glaciation (that part occurring after the glacial maximum) and the Yarmouthian interglacial (see Hibbard et al., 1965:512-514). The Irvingtonian provincial age, therefore, follows the late Blancan provincial age of the early Pleistocene and is succeeded by the Rancholabrean provincial age of the late Pleistocene. No specimen of an Irvingtonian Geomys is referable to any living species. Two Irvingtonian species have been described. Hibbard (1944:735) named Parageomys tobinensis [= Geomys tobinensis] from the Tobin local fauna of Russell County, Kansas. This species since has been reported from the Cudahy local fauna of Meade County, Kansas (Paulson, 1961:137). Hibbard (1956:183) also identified as Geomys tobinensis the pocket gopher recovered from the Saunders local fauna, a late Aftonian deposit of Meade County, Kansas, and reduced the technical name Parageomys from generic to subgeneric rank. Paulson (op. cit.:138) pointed out that the Saunders specimens differ from G. tobinensis, and he, therefore, restricted the name to the small Geomys of the Cudahy and Tobin local faunas of Irvingtonian provincial age. G. tobinensis is markedly smaller than the Blancan G. quinni. The Cudahy and Tobin local faunas are of approximately the same age, and presently both are included in one unit, the Cudahy fauna. The Cudahy fauna is considered to have been deposited in late Kansan as it occurs in strata immediately below the Pearlette ash.

Recently, White and Downs (1961:8) described a new Irvingtonian species, Geomys garbanii, from the middle Pleistocene Vallecito Creek local fauna of San Diego County, California. Many well preserved fossils of the new species were recovered. Geomys garbanii is of medium size (approximately the size of one of the larger subspecies of G. bursarius), and significantly larger than the Irvingtonian Geomys tobinensis of the Great Plains. The Vallecito Creek occurrence of Geomys is the first authenticated record from the Pacific Coast region. Matthew (1902:320) erroneously referred remains of Thomomys to the genus Geomys in his revised list of Cope's earlier report on the Fossil Lake (or Silver Lake) fauna (see discussion of Thomomys above).

A number of Irvingtonian fossil remains of Geomys have not been identified with particular species. Hibbard (1941a:206) found Geomys in the Borchers local fauna (deposited in the time of the Yarmouthian interglacial) of Meade County, Kansas. Also, Geomys has been reported from several sites in Nebraska. Schultz and Tanner (1957:67) reported Geomys from the Angus fossil quarry in Nuckolls County, south-central Nebraska. The Angus fossils were found in sediments of the Sappa Formation considered by Schultz and Tanner to be a Yarmouthian deposit. Fossil quarries (Hay Springs, Rushville, and Gordon) along the south side of the Niobrara River Valley in Sheridan County, Nebraska, have also provided records of geomyids. Both a large and small species of Geomys have been reported from the more recently excavated Rushville and Gordon sites (Schultz and Stout, 1948:562-567, and table 3). In view of the great disparity in size owing to sex, these may actually be males and females of the same species, as mentioned above. The name Hay Springs has been used in reference to all three sites. The ages of the Hay Springs sites are approximately the same, but their correlation is presently under debate. Schultz and Tanner (1957:68-71) maintain that the fossils are distinctly middle Pleistocene, and that they were deposited during late Kansan glaciation, or perhaps from early Yarmouthian into early Illinoian, with the largest concentration coming from the Sappa sands of pre-Illinoian (Yarmouth) age. Hibbard (1958:25), basing his opinion on the presence of Microtus pennsylvanicus, and the stage of evolution of other species in the assemblage, regards the Hay Springs sites as probably Illinoian deposits, but certainly no older than that.

Mooser (1959:413) identified as Geomys the pocket gopher from Irvingtonian deposits in Arroyo San Francisco (loc. no. 5) near the city of Aguascalientes, México. As suggested elsewhere in this account, these fossils may be referable to Zygogeomys rather than Geomys. The Irvingtonian provincial age of this fauna was established by Hibbard and Mooser (1963:245-250). Other alleged occurrences have recently been compiled by Alvarez (1965:19-20). Maldonado-Koerdell (1948:20) noted four fossil occurrences of the genus Geomys in México. Two of these from San Josecito Cave in Nuevo León have since been identified with the genera Orthogeomys and Pappogeomys (Russell, 1960:543-548); the third listed by Maldonado-Koerdell from "near Ameca, Jalisco," was based on Brown's (1912:167) mention of some bones supposedly of the family "Geomyidae," and the fourth refers to pocket gopher remains from the "Hochtals von Mexiko" listed as Geomys by Freudenberg (1921:139). His generic identification is doubtful and the specimens should be compared with Mexican genera of the Geomyinae.

Upper Pleistocene records of Geomys also are common. Upper Pleistocene is here understood to include late Illinoian, Sangamon and Wisconsin deposits; all are considered to be of Rancholabrean provincial age (see Hibbard et al., 1965:512-515) and post-Irvingtonian. The presence of remains of Bison and/or Microtus pennsylvanicus are currently considered mammalian index fossils of Rancholabrean faunas. In the Illinoian, Geomys extended its range to the Atlantic Coast in the southeastern United States. The eastern and western species-groups evidently were isolated throughout much of the late Pleistocene, and, therefore, evolved separately. Of the two, the eastern, or pinetis, species-group seems to have remained somewhat more generalized, and the western, or bursarius, species-group has become more specialized. The Rancholabrean Geomys from deposits in the southeastern United States are referable (see Ray, 1963:325) to Geomys pinetis.

Marsh (1871:121) described Geomys bisulcatus from the North Prong of the Loup River (near Camp Thomas), Nebraska. These beds are also termed the Loup Fork or Loup River fossil beds (see discussion on p. 485), and they lie along the upper reaches of the Middle Loup River in Thomas County (near Senea), Hooker County (near Mullen), and southeastern Cherry County (probably the North Prong beds northwest of Mullen). These beds were at first thought to be of Miocene age, but later were regarded as early Pliocene (see Schultz and Stout, 1948:562-566 for a historical account of expeditions to these fossil sites). Schultz and Tanner (1957:71-72) pointed out that the principal fossiliferous beds in the Middle Loup region are of middle to late Pleistocene age, with most of the fossils coming from the Crete sand and silt beds which are probably early Illinoian deposits, and, therefore, younger than the Hay Springs faunas. Some fossils may have come from the Sappa deposits dated by Schultz and Tanner (loc. cit.) as mostly Yarmouthian deposits. Geomys bisulcatus, judging from the original description and Hibbard's discussion of the cotypes (1954:357), does not differ significantly from Geomys bursarius. However, Geomys bisulcatus is tentatively retained as a valid species. Based on the evidence cited above it seems unlikely that Geomys bisulcatus occurred in pre-Irvingtonian times as often suggested in the literature.

The genus Geomys has been identified in several faunas of Illinoian age, all from the Great Plains. Stephens (1960:1961) reported the genus from the Doby Springs local fauna in Harper County, Oklahoma, and Starrett (1956:1188) reported it from the Berends local fauna in Beaver County, Oklahoma. Schultz (1965:249) assigned 21 isolated teeth, including six incisors, from Butler Springs local fauna (considered by him to be late Illinoian, following the glacial maximum) to Geomys cf. bursarius. Hibbard and Taylor (1960:167) reported a baculum tentatively identified as that of Geomys from the early Illinoian Butler Springs local fauna (including the Adams fauna) of Meade County, Kansas. Hibbard (1963:206) recorded the genus Geomys from the Mt. Scott local fauna (late Illinoian deposits) of Meade County, Kansas; the specimens probably are referable to the living species bursarius. From McPherson County, Kansas, Hibbard (1952:7) reported the genus Geomys from the Kentuck Assemblage, which he (1958:25) regarded as a composite of Illinoian and Sangamon species. Specific identification of the Illinoian pocket gophers is uncertain, primarily due to the fragmentary nature of the material. On the basis of dental characters alone most specimens could be referred to G. bursarius; however the taxonomic status of G. bisulcatus is in doubt, and more complete material may indicate that the Illinoian gophers are specifically distinct from the living species. Consequently, most authors, including myself, have made no attempt to refer these specimens to species. Nevertheless, the Illinoian Geomys from the Great Plains is more closely allied to the living species of Geomys than it is to the earlier Irvingtonian species.

Geomys bursarius has been collected from a number of Sangamon fossil sites on the Great Plains. Although specific identification of specimens of Geomys from Illinoian faunas is uncertain, the Great Plains Geomys from Sangamon and later deposits probably is referable to the living species as Hibbard and Taylor (1960:165) pointed out. They found no difference between Geomys recovered from the Cragin Quarry local fauna (early Sangamon) of Meade County, Kansas, and the living species Geomys bursarius. Isolated teeth of the same species were collected from the Jinglebob local fauna of Meade County, Kansas (Hibbard, 1955b:206), a fauna of the late Sangamon. Hibbard (1943:240) also recorded the genus Geomys (referable to G. bursarius) from the Rezabek local fauna of Lincoln County, Kansas. According to Schultz et al. (1951:6 and table 1) the genus Geomys occurs in buried or "fossil" soils of Sangamon age, lying just above the Loveland Loess, in Nebraska. No specific localities were given by them, nor were any particular specimens mentioned. Dalquest reported Geomys bursarius from two Sangamon faunas in northern Texas. The species is represented in the Ward Quarry local fauna of Cooke County, Texas (1962a:42), and the Good Creek local fauna of Foard County, Texas (1962b:575).

Geomys bursarius has been reported from Wisconsin fossil deposits of the Great Plains and adjacent areas as follows: Jones local fauna, Meade County, Kansas (Hibbard and Taylor, 1960:64-66); Two Creeks Forest beds of the third interstadial soils formed between Cary and Mankato glaciations, late Wisconsin (Schultz et al., 1951:8 and table 1); Cita Canyon local fauna in the northern part of the Panhandle of Texas (Johnson and Savage, 1955:39); Howard Ranch local fauna of Hardeman County in northwestern Texas (Dalquest, 1965:70); Quitaque local fauna of Motley County, Texas (Dalquest, 1964:501); Clear Creek local fauna of Denton County in north-central Texas (Slaughter and Ritchie, 1963:120); Ben Franklin local fauna, of late Wisconsin beds along the North Sulphur River in Delta County, NE Texas (Slaughter and Hoover, 1963:137); Bulverde Cave (Hay, 1920:140; 1924:247) and Friesenhahn Cave (Tamsitt, 1957:321), both in Bexar County, south-central Texas; Alton, Illinois (Hay, 1923:338-339); Wisconsin drift of Illinois, without mention of specific locality (Bader and Techter, 1959:172); Wisconsin drift of southwestern Wisconsin and northeastern Iowa (Hay, op. cit.:343); Wisconsin drift near Galena, Illinois, and mouth of Platte River in eastern Nebraska (Leidy, 1869:406).

Brown (1908:194) described Geomys parvidens from the Conard Fissure, in northern Arkansas. Hibbard (1958:25) concluded that the Conard Fissure fauna represents a glacial stage, probably the Illinoian, and Hibbard et al. (1965:510-511) regarded the fauna as a composite including both Irvingtonian and Rancholabrean elements. White and Downs (1961:21) considered G. parvidens to be a subspecies of Geomys bursarius.

The first Pleistocene occurrence of Geomys in the southeastern United States is from the Reddick I deposits reported by Gut and Ray (1963:325), who found the remains of Geomys pinetis among the fossils comprising the "rodent beds" of Marion County, Florida. Gut and Ray tentatively identified the beds as Illinoian, but Kurten (1965:219) regarded the Reddick I fauna as early Sangamon. Simpson (1928:2) reported Geomys floridanus [= pinetis] from Saber-tooth Cave deposits of Citrus County, Florida. The Saber-tooth Cave (or Lecanto Cave) local fauna is considered by Kurten (op. cit.:219) also to be a Sangamon deposit. Geomys floridanus [= pinetis] was reported from the Seminole Field deposits by Simpson (1929:563); both Simpson and Kurten (op. cit.:221) agreed that the Seminole Field fauna is mainly late Wisconsin, although sub-Recent fossils occur at the tops of the beds. Ray (1958:430) collected remains of Geomys pinetis from the Melbourne Bone Bed of Brevard County, Florida. The Melbourne local fauna is considered to be from Wisconsin deposits by Kurten (op. cit.:220). The eastern species of Geomys were probably derived from Great Plains stock that reached the southeastern Coastal Plains in early Rancholabrean (Illinoian) time. Presently there is no contact between the eastern and western populations of the genus, and it is assumed that disjunction occurred as a result of Wisconsin glaciation. It is interesting to note that the genus Thomomys occurred in this region at approximately the same time; both genera occur in Saber-tooth Cave deposits.

Pappogeomys

The genus Pappogeomys is not known from Pleistocene deposits older than the Wisconsin glaciation, but a pre-Pleistocene occurrence in the Benson beds of Arizona (see discussion of the Pliocene above) shows that Pappogeomys had been differentiated by late Pliocene time. The absence of Pappogeomys, beginning in the early Pleistocene and continuing well into the late Pleistocene, is attributed to the southern distribution of the genus, where its range probably was centered on the Central Plateau of México. The paucity of early and middle Pleistocene deposits from this critical region prevents any definite statements about phyletic development within the genus. All of the late Pleistocene records pertain to the subgenus Cratogeomys (long in use as a generic name but in the present paper reduced to subgeneric rank in the genus Pappogeomys). Schultz and Howard (1935:280) found Cratogeomys [= Pappogeomys] castanops in Burnett Cave in the Guadalupe Mountains of south-central New Mexico. The Burnett deposits are probably late Wisconsin (see Schultz and Tanner, 1957:75, for discussion of the age of these deposits based on carbon-14 tests). These writers (loc. cit.) also referred the mandible of a small pocket gopher to the genus Pappogeomys [= subgenus Pappogeomys]. However, neither genera nor subgenera of the tribe Geomyini can be distinguished on the basis of their inferior dentitions. Judging from the distribution of the modern geomyines, it seems unlikely that the subgenus Pappogeomys has occurred beyond its present range in the late Pleistocene; therefore the small mandible is most likely that of a young individual of Pappogeomys castanops. Russell (1960:543) referred specimens collected at San Josecito Cave in Nuevo León, México, to the group of small subspecies Cratogeomys [= Pappogeomys] castanops. Also, Russell (loc. cit.) identified a rostral fragment as of the genus Cratogeomys [= subgenus Cratogeomys] although the fragment had a combination of features different than in any named species of the genus; he did not name the fragment as a new species, preferring to wait for additional material that could clarify its taxonomic relationships.

Hibbard (1955a:52-53) identified Cratogeomys [= Pappogeomys] tylorhinus from the Becerra Superior deposits in the valley of Tequixquic in the northern part of the state of México. The Wisconsin age of these beds suggests an earlier Pleistocene derivation of the gymnurus-group of species.

Several specimens of the subgenus Cratogeomys have been reported from beds of latest Wisconsin (certainly after the glacial maximum) or post-Wisconsin age. Gilmore (1947:158) found fossil remains of Cratogeomys [= Pappogeomys] castanops commonly in Quaternary cave deposits on the mountain slopes in the vicinity of Cuatro Ciénegas, in central Coahuila. These deposits actually may be of post-Wisconsin origin (see discussion above). Alvarez (1964:8) obtained fragments of Cratogeomys [= Pappogeomys] tylorhinus from sub-Recent deposits of Capa III in the Cueva La Nopalera in southwestern Hidalgo, México. Pappogeomys merriami lives in the area today. Mayer-Oakes (1959:373) reported remains of Cratogeomys [= Pappogeomys] merriami from levels eight and eleven of the excavations at El Risco II, in the northern part of Mexico City. The ages of these deposits are unknown to me, but they probably are no older than late Wisconsin with most of the beds dating from the post-Wisconsin.

Orthogeomys

This genus is not known from the Pleistocene, except for its occurrence in the San Josecito cave deposits of southwestern Nuevo León, México (Russell, 1960:544). Although Orthogeomys does not occur in the immediate vicinity of the cave at the present time, the northern limits of its range is nearby in southern Tamaulipas. The Orthogeomys from San Josecito Cave differs from living species, and has been named Heterogeomys [= Orthogeomys] onerosus Russell (loc. cit.), and is evidently referable to the subgenus Heterogeomys. As mentioned before, the San Josecito Cave local fauna represents deposits of Wisconsin glaciation.

HISTORY OF CLASSIFICATION

The account of the Tucan or Indian mole by Hernandez (sometimes listed as Fernandez) in 1651 probably is the earliest published one of a geomyid (see Merriam, 1895:201; Coues, 1877:607-608). Linnaeus in 1758 did not mention geomyids. In 1772, Kerr described Hernandez's Tucan under the name Sorex mexicana on the basis of Hernandez's account without having seen any specimens. Lichtenstein in 1827 applied the technical name Ascomys mexicana to three specimens collected by Deppe from unknown localities on the tableland of México. Merriam (loc. cit.) pointed out that the name mexicanus of Lichtenstein in 1827 is a nomen nudum, and that it is preoccupied by mexicanus used by Kerr in 1792. The latter can not be technically identified with any particular species of geomyid.

Bartram in 1791 wrote of the pocket gopher of Florida, without formally describing it. The first available technical name is Mus bursarius of Shaw in 1800. Rafinesque in 1817 proposed the first generic names for the geomyids when he described Geomys and Diplostoma. In 1839, Waterhouse referred the genus Geomys to his family Arvicolidae, considered by him to be a subgroup of muroids. In 1841, he suggested that Geomys was related to Bathyergus and Spalax. Waterhouse in 1848 (p. 8) treated the pocket gophers as a subgroup of rodents under the group name Saccomyina, in which he included the genera Heteromys, Saccomys, Perognathus, and Dipodomys. Hence, Waterhouse was the first to recognize the relationship between the heteromyids and geomyids. In the next year Gervais erected the family Pseudostomidae for a group of specialized squirrels to include Geomys and Thomomys and the same genera (at least in part) of heteromyids that Waterhouse classified in the "family" Saccomyina.

In 1839 the name Thomomys was proposed by Maximilian (Wied-Neuwied). All of the generic names previously proposed for pocket gophers were considered by subsequent authors to be synonyms of Geomys.

A third family name, Sciurospalacoides, was proposed by Brandt (1855:188) who referred Geomys and Thomomys to that family. He placed his new family phylogenetically between the family Sciuridae and the family Spalacoides (a group in which Brandt included the genera Spalax, Sipheus, and Ellobius). Brandt took exception to the classification of Waterhouse (1848), who united the geomyids and heteromyids in one family. Brandt placed the heteromyid genera in other groups: Perognathus in the Muridae, and Macrocolus [= Dipodomys] in the Macrolini, a subfamily of the family Dipodoides.