Part 11

Of the molariform dentition only the lower premolar and first two lower molars are known. The enamel investment of p4 is complete, and would not be subject to interruption at any stage of wear; the two prisms are joined at their mid-points, and the isthmus of dentine is relatively broad (as in Pliosaccomys) when compared with modern pocket gophers of this tribe. Also, the re-entrant folds, rather than having parallel sides, diverge broadly to the sides. The divergence is especially noticeable in the labial fold. The lower deciduous premolar would have formed essentially the same enamel pattern with wear as observed in Nerterogeomys [= Zygogeomys] cf. minor (see Hibbard, 1954:fig. 5, A and B) and Pliosaccomys dubius (see Wilson, 1936; pl. 1, fig. 1). Each molar is a single column in the final stages of wear; pre-final stages are unknown. Anterior and posterior enamel plates are present on m1 and m2 (m3 has not been recovered). The dentine tracts of m1 are exposed over a relatively wide surface; therefore, the enamel plates are distinctly separated. The tracts of dentine of m2 are much narrower than in m1 and the enamel plates are barely separated at the anterolateral margin of the tooth. Possibly the enamel band of m2 was continuous in an earlier stage of wear.

The mandible is stout and its general construction not unlike that in modern geomyines. The capsule at the base of the angular process that receives the terminal end of the lower incisor is well developed. The base of the angular processes is preserved, and suggests that the process was short and decidedly smaller than in living examples of the tribe. The masseteric ridge is distinct but weakly developed, and not at all massive as in living pocket gophers. The mental foramen is immediately anterior, and slightly ventral, to the anterior extension of the crest. The basitemporal fossa is absent as such, but its position is marked by a slight depression.

Specimens examined.--Two rami; nos. 29147 (holotype) and 33446; several isolated teeth 30194 and 30195, including an upper incisor and a dp4 (deciduous lower premolar), all from Univ. Michigan Mus. Paleo.

Referred species.--One.

*Pliogeomys buisi Hibbard, 1954. Papers Michigan Acad. Sci., Arts, and Letters, 39:353. Type from Buis local fauna, latest middle Pliocene, Beaver County, Oklahoma.

Genus =Zygogeomys= Merriam

1895. Zygogeomys Merriam, N. Amer. Fauna, 8:195, January 31.

1942. Nerterogeomys Gazin, Proc. U. S. Nat. Mus., 92:507 (type, Geomys persimilis Hay, 1927).

Type.--Zygogeomys trichopus Merriam, 1895, from Nahuatzen, Michoacán.

Chronologic range.--Late Pliocene (Benson and Curtis Ranch local faunas, Arizona, and ?Rexroad Formation, Kansas) to Recent.

Description and discussion.--The size is small to medium for the subfamily Geomyinae. This genus is distinguished principally by the retention of primitive features. In the living species, the skull is generalized, rather than specialized toward either extreme dolichocephaly or platycephaly. The angular process is short, barely exceeding the lateral extensions of the mastoid process of the squamosal. The rostrum is remarkably narrow in relation to its length. The jugal is reduced and displaced ventrally, causing the maxillary arm of the zygomata to articulate with the squamosal arm of the zygomata along the dorsal border of the zygomatic arch (a feature observed also in Orthogeomys cherriei costaricensis).

The upper incisor, recovered in material from the late Pliocene and middle Pleistocene, is bisulcate as in the genus Geomys and the primitive genus Pliogeomys. The enamel plate across the posterior wall of P4 is either complete (late Pliocene to late Pleistocene) or restricted to the lingual half of the tooth (always restricted in living species). The Pliocene specimens of the Rexroad local fauna referred to Nerterogeomys cf. minor by Hibbard (1950:138-139) are exceptional. In these specimens the length and position of the posterior enamel plate is variable; however, all but one specimen had persistant enamel. Evidently, in approximately 43 per cent of the specimens, a complete enamel blade was present (see Paulson, 1961:139), and in the others (except the one without any enamel) the plate was restricted to a small area of the ventral surface, usually on the lingual side of the loph. Hibbard suggested that the decrease in size of the plate, and its restriction to the lingual side, may be a function of age. Hence, most adults would be characterized by the reduced posterior plate on the upper premolar. Although age may be the important factor, intragroup variation cannot be ruled out. It is of interest to note that in all specimens from the Benson (type series of P. minor) and Curtis Ranch local faunas, the former of late Pliocene age and the latter of middle Pleistocene age, the enamel plates are complete on the posterior face of the upper premolar. As mentioned before, the specimens from Kansas may actually represent the transitional stages of the early evolution of Geomys in which the posterior plate of P4 is entirely lost. The enamel pattern of p4 is like that in other members of the tribe (excepting the genus Pliogeomys). The re-entrant angles of P4 and p4 are widely open (obtuse) in the examples recovered from late Pliocene and middle Pleistocene deposits, representing retention of a trait that is primitive in the Geomyini (see account of phylogeny).

M1 and M2 are elliptical in cross-section and each has an enamel plate on both the anterior and posterior surface. In the living species (Z. trichopus), the posterior enamel plate fails to reach the labial margin of the tooth and is restricted to the lingual two-thirds of the posterior surface; however, the enamel plates are complete in the late Pliocene species (Z. minor) and the middle Pleistocene species (Z. persimilis), being only slightly separated from the anterior plate by narrow tracts of dentine on the ends of the tooth. M3 is partly biprismatic in the living species, the two incompletely divided lophs being separated by a distinct outer sulcus. The posterior loph is elongated and forms a conspicuous heel paralleling the evolution of this trait in the genus Orthogeomys; therefore, the crown is longer than wide. The posterior part of the tooth is protected by two lateral enamel plates; of the two, the lingual plate is especially long and extends to the end of the heel. M3 has not been recovered in the Pliocene species, but in the middle Pleistocene species (Z. persimilis) M3 is subtriangular, no longer than wide, and the lateral inflections are weakly developed. The trend towards elongation of M3 evidently occurred in late Pleistocene evolution of the genus. All three of the inferior molars are elliptical, and only the posterior enamel plate is present (as in all other genera of the tribe except Pliogeomys).

The masseteric ridge of the mandible is well developed. In the late Pliocene species Z. persimilis and Z. minor the mental foramen is directly beneath the anterior extension of the masseteric ridge, but in the living species, Z. trichopus, the foramen lies well anterior to the ridge. The basitemporal fossa in the living species is well developed and deep; in the Pliocene species it is usually distinct but shallow (late Pliocene specimens of Z. minor).

Referred species.--Three (two extinct and one living; the last has two subspecies):

*Zygogeomys minor (Gidley), 1922. U. S. Geol. Surv. Prof. Paper, 131:123, December 26. Type from Benson local fauna (late Pliocene), Cochise County, Arizona; also known from the Rexroad local fauna, Meade County, Kansas.

*Zygogeomys persimilis Hay, 1927. Carnegie Inst. Washington Publ., 136. Originally described by Gidley, 1922 (U. S. Geol. Surv. Prof. Papers, 131:123, December 26) as Geomys parvidens which was preoccupied by G. parvidens Brown, 1908. Type from Curtis Ranch local fauna (middle Pleistocene), Cochise County, Arizona.

Zygogeomys trichopus trichopus Merriam, 1895. N. Amer. Fauna, 8:196, January 31. Type from Nahuatzen, Michoacán.

Zygogeomys trichopus tarascensis Goldman, 1938. Proc. Biol. Soc. Washington, 51:211, December 23. Type from 6 mi. SE Pátzcuaro, 8,000 ft., Michoacán.

Genus =Geomys= Rafinesque

1817. Geomys Rafinesque, Amer. Monthly Mag., 2(1):45, November.

1817. Diplostoma Rafinesque, Amer. Monthly Mag., 2(1):44-45, November. Included species: Diplostoma fusca Rafinesque [= Mus bursarius Shaw] and Diplostoma alba Rafinesque [= Mus bursarius Shaw] from the Missouri River region.

1820. Saccophorus Kuhl, Beitr. Zool. und Vergl. Anat., pp. 65, 66. Type: Mus bursarius Shaw, from upper Mississippi Valley.

1823. Pseudostoma Say, Long's Expd. Rocky Mts., I, pp. 406. Type: Pseudostoma bursaria [= Mus bursarius Shaw], from upper Mississippi Valley.

1825. Ascomys Lichtenstein, Abh. K. Akad. Wiss. Berlin (1822), p. 20., fig. 2. Type: Ascomys canadensis Lichtenstein [= Mus bursarius Say], probably from upper Mississippi Valley.

1944. Parageomys Hibbard, Bull. Geol. Soc. Amer., 55:735, June. Type: Parageomys tobinensis Hibbard, from Pleistocene, Cudahy (Tobin) local fauna, Russell Co., Kansas.

Type.--Geomys pinetis Rafinesque, 1817, restricted to Screven County, Georgia, in region of the pines.

Chronologic range.--Late Pliocene faunas of Blancan age (Rexroad, Kansas, and Sand Draw, Nebraska, local faunas) to Recent. Reported from numerous Pleistocene deposits of all stratigraphic levels, especially from the Great Plains, where common today.