Part 5

Hibbard proposed the generic name _Parageomys_ (1944:55), but later regarded it as a subgenus of _Geomys_ (1956:182) that includes those species retaining continuous enamel bands until relatively late in ontogeny; no other differences have been noted. When the early phylogeny of _Geomys_ is better understood, _Parageomys_ may serve as a subgeneric taxon in which the primitive species of _Geomys_ can be grouped, but as of now _Parageomys_ is arranged as a synonym of _Geomys_.

_Pappogeomys_ and _Cratogeomys_ also form a natural group. Their close relationship is best reflected in formal taxonomy by including them in the same genus. Their dissimilarities are of the sort that separate a primitive ancestral lineage from a divergent and progressively more specialized assemblage. The fossil record is inadequate, and I can only speculate that _Cratogeomys_ diverged from primitive _Pappogeomys_-stock in the earlier Pleistocene, at least before the end of the Irvingtonian. _Cratogeomys_ probably originated on the Mexican Plateau and probably underwent its subsequent evolution there. The living species of the subgenus _Pappogeomys_ are evidently relics of the ancestral stock of the genus. Hooper (1946:397), I think correctly, considered _Platygeomys_ as congeneric with _Cratogeomys_, although the highest degree of specialization of the genus is attained in those species formerly classed in the genus _Platygeomys_. Even so, in my opinion, the differences are insufficient to warrant even subgeneric recognition.

CLASSIFICATION

Family GEOMYIDAE Gill, 1872

Rodents of the superfamily Geomyoidea specialized for completely fossorial life (early Pliocene to Recent); specialized earlier (late? Oligocene and early Miocene) for semi-fossorial habits; body thickset, fusiform without apparent neck (in modern geomyids); legs short; forelegs especially stout; eyes and ears small (pinna reduced to inconspicuous crest concealed beneath pelage); tail tactile, shorter than head and body; lips closing behind incisors; cheek pouches external, fur-lined; baculum rodlike, arched, having expanded quadriform platelike base; pelage long, soft without underfur, covering body in thick coat (in some species of _Orthogeomys_ scant, harsh or scattered bristles); color varying from pale tints of buffy (almost white) to metallic black.

Skull thick-walled, massive, angular, relatively broad, and flattened; distinctly murine form, but having zygomasseteric structure of advanced sciuromorphs, including small infraorbital canal (that transmits no part of masseter muscle) and well-developed, broad zygomatic plate; zygomata massive and widely flaring, jugals stout; rostrum robust, relatively broad and deep, and without evidence of transverse canal (as in Heteromyidae); anterior projection of nasals only slightly exceeding that of upper incisors; interorbital region usually constricted, narrower than rostrum; anterior opening of infraorbital canal far forward on side of rostrum, about half way between zygomatic plate and upper incisor and just behind premaxillary-maxillary suture, its opening countersunk in oblique sulcus (for protection from muscle contraction); postorbital process lacking, except for rudimentary knoblike projection in subgenus _Macrogeomys_; palate relatively narrow, its deeply sculptured surface sloping steeply downward posteriorly causing region supporting maxillary tooth-row to be markedly depressed; palatine bone reduced, forming, on two abruptly different levels, posterior margin of hard palate behind tooth-rows; parietals compressed and narrow, and most of cerebral cavity roofed by squamosals (in some species squamosals overlap lateral parts of parietals); tympanic bullae completely inferior in position and fully ossified, external meatus being developed laterally as elongated tube; mastoid not inflated, but broadly exposed at posterolateral margin of the skull; occiput large, its surface usually rugose, and paroccipital processes large and flangelike, at least in advanced groups (early Pliocene to Recent); ramus relatively short and stout, having distinct crest and ridges for muscle attachments; coronoid process well developed, erect; articular condyle prominent; angular process prominent, reflected laterally, and in modern groups lateral extension protruding from posterior border of ramus nearly at right angle; capsule for root of lower incisor, prominent between angular process and articular condyle.

Anterior surface of incisors broad and flat, always smooth on lower teeth, but either smooth or grooved on upper teeth depending on taxon; cheek teeth hypsodont, becoming progressively higher crowned in modern groups, rooted in primitive groups (late? Oligocene to middle Pliocene), rootless and ever-growing in modern groups (late Pliocene to Recent); upper and lower premolars persistently bicolumnar; upper and lower molars bicolumnar only in primitive groups (late? Oligocene and early Miocene), becoming progressively monocolumnar in advanced groups (early Pliocene to Recent), primitive bicolumnar pattern being retained on occlusal surface only in early stages of ontogeny and in third molar throughout life; enamel pattern of occlusal surface of cheek teeth based on sextituberculate prototype (see Wood and Wilson, 1936:388-391), having cusps arranged in two transverse rows of three cusps each, excepting three anterior cusps of premolars that are arranged in trefoil, especially on p4 (sometimes only one or two, rather than three, cusps develop in a particular set, especially in p4), conules absent; protostyle and endostyle in upper teeth and protostylid and hypostylid in lower teeth formed from cingulum; cusps of each row uniting with wear into transverse enamel lophs (or lophids), each tooth having two lophs, one on anterior column, protoloph and protolophid, and one on posterior column, hypoloph and hypolophid, that unite with additional wear forming continuous enamel band; enamel lacking on sides of each column in advanced lineages, thereby restricting enamel to anterior and posterior walls; with extreme reduction, posterior plates of upper teeth and, more commonly, anterior plates of lower molars, missing. Dental formula: 1/1, 0/0, 1/1, 3/3.

Key to the Subfamilies of Geomyidae

A Angular process of ramus mostly below alveolar level of mandibular tooth-row; pattern of premolar like that of molars, consisting of two subequal crests united at one or both margins of tooth; molars persistently bicolumnar; molariform teeth always rooted. Subfamily Entoptychinae p. 513

A´ Angular process of ramus mostly above level of mandibular tooth-row; pattern of permolar unlike that of molars, consisting of two prisms differing in size and united at their mid-points but never at either margin; molars progressively monocolumnar, except for early Miocene forms; molariform teeth rooted only in primitive genera (late? Oligocene to middle Pliocene), and rootless and ever-growing in later genera (late Pliocene to Recent). Subfamily Geomyinae p. 514

Subfamily ENTOPTYCHINAE Miller and Gidley, 1918

Anterior face of upper incisor usually smooth, sometimes bearing faint groove in center or near medial margin of tooth, at least in _Gregorymys_; cheek teeth hypsodont, medium to high crowned, and rooted in all but _Entoptychus_ (has rootless, ever-growing teeth); cheek teeth identical in form, premolars resembling molars and lower cheek teeth mirror images of upper teeth; crowns biprismatic, having two columns joined at edge of protomeres (for description of term, see discussion of primitive morphotype on page 537) and with persistent lateral fissure between them; lateral re-entrant fold deep, penetrating at least half width of crown, from external side in upper teeth and internal side in lower teeth (in specialized genus _Entoptychus_ lophs, upon additional wear, join also at edge of parameres, thus uniting columns at both ends and thereby enclosing interior part of lateral fissure as a transverse fossette in center of tooth); enamel investment of prisms usually complete, including inflection bordering re-entrant folds, occlusal pattern becoming interrupted with wear only in _Entoptychus_, where enamel disappears first from sides of crowns (following union of anterior and posterior columns at both sides) and later, in final stages of attrition, from anterior wall of lower molars and posterior wall of upper molars.

Maxillary bone without pronounced vertical depth in part supporting cheek teeth, its inferior border only slightly lower than inferior border of premaxillary and alveolar lips of molariform teeth consequently approximately level with, or slightly below, alveolar lip of upper incisor; squamosal without lateral expansion, therefore, meatal tube of auditory bulla separated from zygomatic process of squamosal by deep, well-developed postglenoid notch; angular part of mandible below alveolar level of mandibular cheek teeth; angular process only slightly reflected laterally; coronoid process low, tip only slightly above condyle.

For information concerning the structure and relationships of the known genera, and for accounts of species, see Wood (1936). A list of the named genera in order of specialization is as follows:

*_Pleurolicus_ Cope, 1878. Proc. Amer. Phil. Soc., 18:66.

*_Gregorymys_ Wood, 1936. Amer. Mus. Novit., 866:9.

*_Grangerimus_ Wood, 1936. Amer. Mus. Novit., 866:13.

*_Entoptychus_ Cope, 1878. Proc. Amer. Phil. Soc., 18:64.

Five new species have been described since Wood's (1936) revision. They are: _Pleurolicus clasoni_ MacDonald (1963:180); _Gregorymys kayi_ Wood (1950:335); _Gregorymys montanensis_ Hibbard and Keenmon (1950:198); _Grangerimus dakotensis_ MacDonald (1963:182); _Grangerimus sellardsi_ Hibbard and Wilson (1950:623).

Subfamily GEOMYINAE Baird, 1858

Anterior face of upper incisor primitively smooth, grooves consistently developed only in one modern lineage (Geomyini); cheek teeth hypsodont, primitively rooted and having crown of medium height (late Oligocene to middle Pliocene), being higher crowned, rootless and ever-growing in modern lineages (late Pliocene to Recent); primitively crowns of cheek teeth biprismatic, having two columns joined at mid-points by narrow isthmus and entire crown sheathed in continuous band of enamel; premolars retaining primitive biprismatic form, anterior and posterior columns never uniting at edge of protomeres or parameres, and with both lateral re-entrant folds persistent throughout life; primitive biprismatic pattern becoming decidedly modified in molars (except in M3), having two prisms progressively uniting into one column by reduction and loss of lateral inflections, primitive biprismatic patterns being retained only in early stages of ontogeny; third upper molars retaining, at least partially, primitive bicolumnar pattern (except in Thomomyini), with relatively broad isthmus and horizontally shallow re-entrant folds, lingual fold sometimes wanting; enamel pattern becoming discontinuous (late Pliocene to Recent) owing to loss of enamel from sides of each column; remaining enamel restricted to anterior and posterior plates, or cutting blades, and enamel bordering lateral inflections in premolars (considering both sides together, these plates constitute essentially two transverse cutting blades); enamel pattern of M3 varying, depending on taxon; with specialization, anterior plates of lower molars and posterior plates of upper premolar and molars may be reduced or lost; except in primitive species (early Miocene), no enamel fossettes retained in adult dentitions.

Maxillary bone having pronounced vertical depth in part supporting cheek teeth, inferior border arching downward well below inferior border of premaxillary; consequently, alveolar lips of molariform teeth decidedly below level of alveolar lip of upper incisor; squamosal with marked lateral expansion at expense of postglenoid notch; notch compressed and reduced between meatal tube of auditory bulla and zygomatic process of squamosal; angular part of mandible mostly above alveolar level of mandibular cheek teeth; angular process reflected laterally at right angles to axis of ramus and developed into heavy knoblike projection; coronoid process well developed, tip decidedly higher than condyle; fossorial specializations remarkably well developed in advanced lineages, degree of specialization of primitive Miocene species unknown but probably only semi-fossorial as in Entoptychinae.

Key to the Tribes of the Geomyinae

A Enamel investment complete and uninterrupted, even in final (adult) stages of wear; cheek teeth rooted, with crowns of medium height; third lower molar biprismatic, the two columns separated by inner and outer re-entrant folds as in lower premolar. Tribe Dikkomyini p. 515

A´ Enamel investment incomplete and discontinuous, reduced, at least in final (adult) stages of wear, to interrupted enamel plates; cheek teeth rootless and ever-growing (except in extinct genus _Pliogeomys_), crowns of maximum height; third lower molar monoprismatic, without trace of inner and outer re-entrant folds as in first and second lower molars.

B Upper incisors smooth, occasionally with a fine indistinct groove near inner margin of tooth; form of third upper molar same as M1 and M2, monoprismatic, anteroposteriorly compressed, and having transverse enamel plates on both anterior and posterior faces, and without suggestion of either labial or lingual re-entrant folds; basitemporal fossa absent (except for a shallow depression in one Recent species, _T. townsendii_); forefoot small and narrow with claws not elongated for digging. Tribe Thomomyini p. 518

B´ Upper incisors grooved, bearing either one or two sulci; form of third upper molar distinctly different from M1 and M2, fully or partially biprismatic (with a few exceptions discussed beyond), without marked anteroposterior compression (either subtriangular, elongated, suborbicular or quadriform in cross-section, but not elliptical as in M1 and M2), and having typical transverse anterior plate and two lateral plates (varying in their development, depending on taxa), but no posterior plate, and with lateral re-entrant folds usually developed, especially labial inflection (although sometimes minute in a few species, as described beyond); basitemporal fossa well-developed, although occasionally shallow or absent (primitive species of _Zygogeomys_); forefoot large and broad, with elongated claws for digging. Tribe Geomyini p. 521

Tribe DIKKOMYINI, new tribe

_Genotype._--_Dikkomys_ Wood, 1936.

_Chronologic and geographic range._--Early to Middle Pliocene (early Arikareean to mid-Hemphillian) in western United States. Known from Miocene fossil sites in Montana, South Dakota, and Nebraska and Pliocene sites in South Dakota, Oregon, Nevada, and southern California. For precise localities see accounts of _Dikkomys_ and _Pliosaccomys_ beyond.

_Diagnosis._--Small Geomyinae; lacking specializations of more advanced tribes; upper incisors smooth, at least in _Pliosaccomys_; molariform teeth always rooted and having crowns of medium height; enamel investment of cheek teeth complete and uninterrupted in all stages of wear; crowns of molars primitively biprismatic, having two columns united at mid-points, thus forming narrow isthmus separating lateral re-entrant folds as in premolars, and, with wear, also uniting secondarily at protomeres (with exception of third lower molars), consequently, isolating remnant of that inflection as shallow fossette (columns uniting first at protomeres in _Pliosaccomys_); anterior and posterior columns of first and second molars, both above and below, becoming progressively united into one column in advanced Dikkomyini (early and middle Pliocene), but m3 (M3 unknown) retaining primitive biprismatic pattern, with columns joined at centers but never at protomeres (for details of dentition see generic accounts); mandible stout, its angle mostly above mandibular tooth-row; masseteric ridge low; basitemporal fossa barely discernable in some fragments of _Pliosaccomys_; postcranial skeleton unknown.

Key to the Genera of the Tribe Dikkomyini

A Molars biprismatic throughout life; anterior and posterior lophs of first and second molars in pre-final stages of wear uniting first at their mid-points and later at edge of protomeres; anterior lophid of lower premolar having distinct anteroexternal inflection. Genus _Dikkomys_ p. 516

A' First and second molars becoming monoprismatic in final (adult?) stages of wear, biprismatic only in pre-final stages of wear; third molars persistently biprismatic; anterior and posterior lophs of first and second molars uniting first at edge of protomeres; anterior lophid of lower premolar lacking anteroexternal inflection. Genus _Pliosaccomys_ p. 517

Genus =Dikkomys= Wood

1936. _Dikkomys_ Wood, Amer. Mus. Novit., 866:26, July 2.

_Type._--_Dikkomys matthewi_ Wood, 1936, from Lower Harrison deposits near Agate, Sioux County, Nebraska.

_Chronologic range._--Early Miocene, from early Arikareean (Lower Harrison local fauna of Nebraska) to middle Miocene, late Hemingfordian (Upper Rosebud local fauna, South Dakota, and the Deep River Formation, Montana). According to MacDonald (1963:149-150), the Upper Rosebud is middle Miocene rather than early Miocene.

_Description._--Size small, about as in small kinds of _Thomomys_; known only from fragmentary mandible, including molariform dentition in place, and isolated cheek teeth, including M1 (see Wood, 1936:26-28 and fig. 32; Galbreath, 1948:316-317 and fig. 1; and Black, 1961:13-14 and fig. 58); upper incisors unknown; cheek teeth hyposodont, persistently rooted, and having crowns of medium height compared with Recent geomyids; enamel investment complete and uninterrupted in all molariform teeth in all stages of wear; P4 unknown, but probably formed like p4; p4 persistently biprismatic, two crowns joined at mid-points by relatively narrow isthmus separating lateral re-entrant folds; anterior lophid of p4 having distinct anteroexternal inflection; molars also biprismatic throughout life; two lophids of lower molars first uniting at mid-points as in p4, and, with additional wear, m1 and m2 secondarily uniting at edge of protomeres and forming isolated enamel fossette between point of connection (detailed description of stages of wear discussed in account of phylogeny of subfamily); m3 permanently joined at mid-point only, without lateral union at edge of protomeres; upper molars, judging by M1 (M2 and M3 unknown), having same pattern as lower molars, but first union of lophs decidedly on lingual side of center, consequently, lingual re-entrant fold small; M1 probably developing U-pattern in advanced stages of wear by union of protomeres, with minute lingual fossette developing in transition as lophs secondarily become united at lingual edge of columns; mandible stout and geomyidlike; masseteric ridge weakly developed; basitemporal fossa absent.

Evidently, _Dikkomys matthewi_ is more primitive than _Dikkomys woodi_. The modified H-pattern in m1 and m2, with the metalophid and hypolophid joined at both their mid-points and also at their protomeres (by union of the protostylid and hypostylid in the lower dentition), is persistent throughout life. Therefore, the enclosed enamel fossette is not eradicated with wear. In m1 and m2 of _Dikkomys woodi_, the fossette is shallower, and, at least in advanced stages of wear, it would disappear, therefore, forming a U-pattern on the occlusal surface, as in M1 and M2, but lateral inflection horizontally shallow rather than deep as in entoptychines.

Specimen (No. P 26284 FMNH) reported as _Dikkomys matthewi_ by Galbreath (1948:316) is referable to the recently described species _Dikkomys woodi_ Black, 1961.

_Specimens examined._--One, no. P 26284, Field Mus. Nat. Hist., from upper Rosebud, Shannon Co., South Dakota.

_Referred species._--two:

_Dikkomys matthewi_ Wood, 1936. Amer. Mus. Novit., 866:26, July. Type from early Arikareean Lower Harrison deposits (early Miocene) near Agate, Sioux County, Nebraska.

_Dikkomys woodi_ Black, 1961. Postilla, Yale Peabody Museum, 48:13, January 16. Type from Deep River Formation, late Hemingfordian (middle Miocene), Meagher County, Montana; also known from Upper Rosebud deposits (middle Miocene) near Wounded Knee, Shannon County, South Dakota.

Genus =Pliosaccomys= Wilson

1936. _Pliosaccomys_ Wilson, Carnegie Inst. Washington Publ., 473:20, May 21.

_Type._--_Pliosaccomys dubius_ Wilson, 1936, from Smiths Valley local fauna in Lyon County, Nevada.

_Chronologic range._--Early Pliocene, late Clarendonian (Wolf Creek local fauna, South Dakota, and Nettle Springs local fauna, California) to Middle Pliocene, middle part of Hemphillian (Smiths Valley local fauna, Nevada, and McKay Reservoir and Otis Basin local faunas, Oregon).

_Description._--Size small (alveolar length of mandibular tooth-row measuring 6.0 in holotype), about as in _Thomomys monticola_; upper incisor relatively broad and flat, having anterior face smooth, without trace of grooving; crowns of cheek teeth of medium height and rooted; enamel investment continuous and uninterrupted in all stages of wear; premolars permanently, biprismatic; P4 having anterior prism subtriangular and decidedly smaller that sub-crescentic posterior prism, and joined near centers by narrow, obliquely oriented isthmus; p4 having anterior prism subovate, posterior prism strongly compressed anteroposteriorly, and joined at mid-points by relatively broad and straight isthmus; first and second molars, both above and below, monoprismatic in final (?adult) stage of wear, derived ontogenetically from primitive bilophate pattern by coalescence of two columns into one; M1 and M2 mirror images of m1 and m2 in pre-final stages of wear, two columns first uniting at edge of protomeres forming U-pattern, and primitive H-pattern never developing in either series (for detailed description of stages of wear, see account of phylogeny, p. 546); m3 (M3 unknown, but probably with same form as in Geomyini, see p. 552) persistently biprismatic, two columns joined by relatively broad isthmus at centers, consequently, forming H-pattern of primitive ancestors; rostrum heavy and broad as in modern geomyids; palate narrow and strongly ribbed; mandible stout; masseteric ridge and fossa well developed; basitemporal fossa absent.

_Specimens examined._--Six, nos. 1796 (holotype)--1799, 1804 and 1806 (CIT) now in the Los Angeles County Museum, all from Smiths Valley local fauna, Middle Pliocene, Nevada.

_Referred species._--two:

*_Pliosaccomys dubius_ Wilson, 1936. Carnegie Inst. Washington Publ., 743:20, May 21. Known from early and middle Pliocene faunas including Wolf Creek local fauna (late Clarendonian), Shannon County, South Dakota; McKay Reservoir local fauna and Otis Basin local fauna (Hemphillian), Oregon; type from Smiths Valley local fauna (probably middle Hemphillian), Lyon County, Nevada.

*_Pliosaccomys wilsoni_ James, 1963. Univ. California Publ. Geol. Sci., 45:101, June 26. Type from Nettle Springs local fauna of late Clarendonian (early Pliocene), Ventura County, California.

Tribe THOMOMYINI, new tribe

_Type._--_Thomomys_ Wied-Neuwied, 1839.