Chapter IV: Part 4

In _Dikkomys_, the anterior and posterior column first unite near their mid-points in the first stages of wear thus producing a figure-8 shaped (H-shaped) occlusal pattern in the premolar and all three molars. Evidently in the first two upper molars, the columns unite closer to their lingual margins than their mid-points, but at any rate both outer and inner re-entrant folds are evident at this stage of wear. With continued attrition on m1 and m2 of _Dikkomys_, the anterior and posterior columns secondarily unite at the edge of their labial margins thus enclosing a fossette of enamel in the labial half of the tooth. The lateral coalescence at the ends of the protomeres occurs because of the shallow vertical depth of the labial re-entrant fold, and the fossette itself does not reach the base of the crown and with continued wear it too would disappear, but not until the last stages of wear, at least in _Dikkomys matthewi_. The lingual re-entrant fold is deep, and therefore, persistent through all stages of wear. Although the amount of wear required for its effacement would be great, the occlusal configuration of the first and second lower molars in _Dikkomys_ could be eventually ground down to a U-pattern as in the entoptychids. Only one upper molar of _Dikkomys_, the first, has been recovered (see Wood, 1936:23, fig. 32B). Although the tooth is in an early stage of wear, the lingual valley is minute. Less attrition than required in m1 and m2 would progressively reduce the lingual fold until it too would essentially form a U-pattern, perhaps retaining a slight lingual inflection. Hence, the first upper molar becomes a mirror image of the first lower molar, and the second upper molar probably had the same pattern as the first (at least it does so in _Pliosaccomys_). Both of the lateral re-entrant folds of the premolar are deep vertically, and consequently would not disappear with occlusal wear. Therefore, the H-pattern of the premolars is retained throughout life.

The m3 (M3 unknown for _Dikkomys_ or _Pliosaccomys_) also has deep lateral folds; hence, it too retains the H-pattern in all stages of attrition, although the isthmus between the two prisms may become wider in the final phases of wear (as it does in _Pliosaccomys_).

In _Pliosaccomys_, the stages of wear are essentially the same as those described for _Dikkomys_, except that the anterior and posterior loph of the first and second molars tend to unite closer to one side of the tooth, lingual side in upper molars and labial in lower. Only a slight inflection of the re-entrant fold is evident on the side of union, and the inflection disappears in the first phases of wear as the columns unite. Concomitant with the lateral shift in the initial point of coalescence of the transverse lophs, the occlusal penetration of the re-entrant fold from the opposite side increases in horizontal depth, and the fold extends medially more than half way across the occlusal surface, thus forming a pattern essentially like that of the entoptychids. The U-pattern in _Pliosaccomys_ appears in the initial stages of wear without going through an earlier H-pattern as is the case in its Miocene ancestors of the genus _Dikkomys_, unless the minute inflection is considered as indicative of that stage. The two columns of the premolar and m3 are joined near their mid-points as in _Dikkomys_; therefore, they retain their primitive H-pattern, a feature unique to the Geomyinae.

The evolutionary trend toward an ontogenetically earlier U-pattern in the first two molars in the primitive lineage of the Geomyinae suggests that the U-pattern characteristic of the Entoptychinae was simply an earlier tendency toward the same specialization that occurred later in the subfamily Geomyinae. If so, early entoptychines would have been characterized by an H-pattern in the first stages of attrition, like _Dikkomys_, and later developed union at the edge of the protomeres. However, in the entoptychines, all the molariform dentition, and not merely the first and second molar, became specialized; consequently the U-pattern was produced on the occlusal surfaces of each of the cheek teeth. As in _Pliosaccomys_, the transitional phase, in which the two columns were united at their mid-points, was eventually eliminated from the pattern of wear and only the U-pattern, that now appeared in the initial stages of wear, was retained. In the entoptychines of the early Miocene there is no suggestion of the H-pattern that characterizes the Geomyinae, except in the position of the cusps before wear in the lower molars of _Pleurolicus sulcifrons_, which, according to Wood (1936:6), suggests the H-pattern. In earlier unknown Oligocene stages of evolution, the prisms possibly united first at their mid-points, and the columns may have joined at the side of the tooth only in the terminal stages of wear. The U-pattern of pre-Miocene entoptychines, therefore, may have become the dominant occlusal pattern only in the later stages of phyletic development.

According to the recently expressed views of several paleontologists, the Entoptychinae constitute the primitive lineage of the family and the early Geomyinae constitute a specialized offshoot of the entoptychine ancestral assemblage. The structure of the Entoptychinae, especially of the less advanced genera, closely approximates that of the hypothetical primitive morphotype. But, according to my view, the subfamily Geomyinae constitutes the ancestral assemblage and its structure is essentially that of the primitive morphotype of the family. At any rate the structure of the early geomyines more closely approximates the structure of the ancestral stock than the more divergent entoptychines. Therefore, the genus _Dikkomys_ of the early Miocene, the first known geomyine, is considered to be a generalized geomyid, and, although it is a contemporary of the more specialized entoptychid assemblage, is considered to be more closely allied to the ancestral stock.

The entoptychines were the dominant and most highly differentiated geomyids of the early and middle Miocene. Nevertheless, they became extinct in the middle Miocene, and the geomyines of that time survived and later gave rise to the modern pocket gophers. Therefore, the early history of the family Geomyidae is characterized by an early radiation and trend toward specialization, followed by survival of the less specialized Geomyinae and extinction of the more specialized Entoptychinae.

Entoptychid Radiation

The most abundant geomyids of the early and middle Miocene, the Entoptychinae, consisted of at least 24 species (see Wood, 1936:4-25) classified in four genera: _Pleurolicus_, _Gregorymys_, _Grangerimus_, and _Entoptychus_. The genera were essentially contemporaneous (see Figure 3). Even so, the subfamily was morphologically varied, pointing to an earlier origin in the Oligocene (actually a part of the John Day Fauna, including _Pleurolicus_ may be correlated with late Oligocene Whitneyian age) followed by a relatively rapid radiation including all four genera in the early Miocene. Two genera, _Pleurolicus_ and _Gregorymys_, continued into the Middle Miocene (Hemingfordian). This divergence, specialization, and subsequent radiation suggest that the entoptychines evolved into a new major adaptive zone, in the sense described by Simpson (1945:199-206).

The radiation is correlated geographically and temporally with the southward retreat of the Neotropical flora of the Tertiary from the western United States and southward movement of the Arctic flora of the Tertiary (see Axlerod, 1950; Berry, 1937:31-46; Chaney, 1947:139-148; and Kendeigh, 1961:280-283). In the early Tertiary the Neotropical-tertiary geoflora occurred northward to at least 49° latitude in western North America, and the boreal Arctic-tertiary flora was restricted to a circumpolar zone. The southward and eastward shift of the Neotropical-tertiary flora, associated with the drying and chilling of the continent, began in the middle or late Oligocene and was concurrent with the divergence and radiation of the Entoptychinae. Beginning in late Oligocene and continuing at least into middle Miocene, most of the region in which the entoptychines occurred was occupied by the Arcto-tertiary geoflora of which the temperate forest division contributed the dominate plant associations. The maples, chestnuts, dogwoods, beeches, walnuts, oaks, elms, birches, and sycamores of that flora were the forerunners of today's eastern deciduous forest. It is my view that the entoptychines became adapted to the conditions of this paleoecological environment and radiated rapidly in the Arikareean when the major change occurred in climax vegetation. The ancestral stock of the Geomyinae was not so successful in the Arcto-tertiary climax, and most of it probably was displaced southward along with the tropical flora.

The skeleton in the entoptychines is not so strongly fossorial as in the modern geomyids (Wilson, 1949:117), and these early geomyids probably were semi-fossorial with somewhat the same burrowing habits as those of the living mountain beaver (_Aplodontia_). Inasmuch as the morphology and taxonomy of the entoptychines were discussed in detail by Cope (1884) and reviewed later by Wood (_loc. cit._), there is no need to recount the details here. According to Wood (_op. cit._, 27-28), _Pleurolicus_ occupied a central position in the entoptychid radiation and perhaps appeared slightly earlier than the other genera. Wilson (1949) suggested that the lower part of the John Day may actually be Upper Oligocene rather than Lower Miocene, and this arrangement is followed here. Also, _Pleurolicus_ is less specialized than the other genera and occurs in deposits of both the Great Plains and the Pacific Coast. _Gregorymys_, also little specialized, occurred only on the Great Plains. The more specialized genera, _Grangerimus_ and _Entoptycus_, evidently appeared somewhat later than _Pleurolicus_ and evolved from it. Except for a record from southern Texas reported recently by Hibbard and Wilson (1950:621-623) and the new species described by MacDonald (1963:182) from the Sharps Formation of South Dakota (early Arikareean), _Grangerimus_ is known only from the Pacific coast. _Entoptycus_ was restricted to the Pacific Coast (John Day fauna).

_Entoptycus_ is the most specialized of the known genera; it has pronounced fossorial adaptations, especially in the skull. Its molariform teeth are rootless and ever-growing as in the modern geomyines. Moreover, the continuous enamel bands on only moderately worn teeth become separated in the final stages of wear into anterior and posterior enamel plates by tracts of dentine that extend toward the crown on the sides of each tooth. This extension was made possible by the union of the two columns at both the lingual and labial margins of the tooth forming an O-pattern, and the crown is essentially monoprismatic save for the isolated enamel fossette in the center of the tooth. The fossette is all that remains of the lateral re-entrant fold that characterized the preceding U-pattern of the earlier stages of wear. Late in the sequence of wear, the anterior enamel plate is lost in the lower molars and the posterior plate in the upper molars. The U-pattern characterizes the final stages of attrition in the other genera of the Entoptychinae; none developed the dental specializations seen in _Entoptycus_. Rootless, ever-growing cheek teeth, discontinuous enamel patterns, and monoprismatic molars were not evolved in the subfamily Geomyinae until the late Pliocene.

Phyletic Trends in Subfamily Geomyinae

The subfamily Geomyinae is made up of three groups, recognized taxonomically for the first time in this account as tribes--Dikkomyini, Thomomyini, and Geomyini (for full discussion of classification, see previous account). The phylogeny proposed by me is illustrated in Figure 3. The tribe Dikkomyini is characterized by generalized and primitive features that together form the basic structural foundation of the subfamily. Evolution within the Dikkomyini resulted in the acquisition and perfection of fossorial adaptations. The Thomomyini and Geomyini are considerably more specialized than the ancestral Dikkomyini from which they evolved. The Geomyini are clearly more specialized than the Thomomyini, suggesting closer affinity between the Thomomyini and the Dikkomyini than between the Geomyini and the Dikkomyini. The specializations in the dentition and the associated changes in the skull of the Thomomyini and Geomyini permit more efficient mastication of fibrous vegetation. Along with these specializations, fossorial adaptations inherited from the Dikkomyini are retained without noteworthy modification.

_Dikkomys_, the earliest known genus of the tribe Dikkomyini, can be taken as a starting point of evolution for the subfamily Geomyinae. The Pliocene genus _Pliosaccomys_ is the only other known geomyine having primitive features closely resembling those of _Dikkomys_. The relatively close but previously unrecognized relationship between _Dikkomys_ and _Pliosaccomys_ can be understood when patterns of wear on the occlusal surfaces of the cheek teeth are taken into account. It appears that _Pliosaccomys_ descended from _Dikkomys_-like stock, if not _Dikkomys_ itself. Although _Dikkomys_ is towards the beginning of this phyletic sequence and _Pliosaccomys_ towards the end of the sequence, the primitive features shared by the two provide a generalized morphotype for the subfamily Geomyinae.

In the molariform dentition, an almost complete series of stages of wear in _Pliosaccomys_ has been preserved, and those of _Dikkomys_ can be reconstructed with reasonable accuracy from those that are known (see Fig. 4):

(1) In the initial stage of wear in _Dikkomys_ the anterior and posterior columns are separated by an intervening valley (Fig. 4A), and the occlusal surface of each column bears a loph of dentine surrounded by a ring of enamel: protoloph on the anterior column and metaloph on the posterior column of the upper teeth (protolophid and hypolophid in corresponding positions in the lower teeth). Actually this stage is not preserved in the known material of _Dikkomys_, but does occur in both geomyines and entoptychines in all stages of evolution, and it must have also occurred in _Dikkomys_ in order for the next two stages, which are preserved, to have developed.

(2) The occlusal surfaces are ground down to a level where the enamel loops of the two columns join at their mid-points, thus forming an H-shaped pattern (Fig. 4B), or more exactly a pattern resembling a figure 8. Probably this was the primitive pattern in the final stage of wear in the geomyid ancestor of the Oligocene.

(3) In the pre-final stage of wear, the anterior and posterior lophs of the first and second molars unite secondarily at the edge of their protomeres (labial side in the lower and lingual in the upper), thus enclosing an isolated enamel fossette (Fig. 4C). Lateral union occurs in the lower teeth because the vertical depth of the labial re-entrant angle is less than the depth of the lingual re-entrant fold. In the upper teeth the reverse is true. The re-entrant angle on one side of the premolar is as deep vertically as the angle on the other side of that tooth, and both reach the base of the crown; therefore, they do not disappear at any stage of attrition. The same pertains in the third lower molar.

(4) In the final stage of wear (Fig. 4D), the enamel fossette disappears as a result of continued attrition on the occlusal surface in the upper series. The fossette may vary somewhat in vertical depth in m1 and m2, but the amount of wear required for its effacement would be greater than in the upper teeth. Therefore, upon wear, the U-pattern would become characteristic of the final stage in M1 (and probably also M2), but the modified H-pattern described in Fig. 4C would prevail in m1 and m2. Perhaps, in extremely worn teeth, the labial fossette of m1 and m2 would disappear. If this advanced stage of effacement is obtained, then the two columns would be united across the entire surface of their protomeres from the center of the crown to its labial edge, and the occlusal pattern would be in the shape of a U.

The occlusal pattern, at least in M1 and M2, in the final stages of wear in _Dikkomys_ resembles that in the subfamily Entoptychinae, but the U-pattern develops on only the first and probably the second molar in _Dikkomys_ and not on all of the cheek teeth as it does in the entoptychines. Judging from the material that has been described, the U-pattern did not develop in the lower teeth of _Dikkomys_ until the Hemingfordian (_D. woodi_), upper Rosebud, and specimens of _D. matthewi_ from the earlier Arikareean, lower Harrison, suggest that the modified H-pattern, with secondary coalescence at the edge of the protomeres, persisted throughout life, without developing the U-pattern in the final stages of wear.

Essentially the same patterns of wear characterize the genus _Pliosaccomys_, except that the earlier stages were telescoped and the second stage was omitted while another (final) stage was added. The stages are reconstructed in sequence in figure 4, and all are based on preserved dentitions, as follows:

(1) The first phases of wear produced the pattern (Fig. 4E and I) described for _Dikkomys_ in the previous account (Fig 4A).

(2) A small additional amount of wear produced the 2nd stage (Fig. 4F and J) characterized by a U-pattern, formed by union of the anterior and posterior columns at the edge of the protomeres of the first and second molars, both above and below, without first forming an H-shaped pattern. Union at the mid-points thus was omitted from the sequence of wear in these two teeth. In the premolars and third molars the primitive H-pattern did form, as in _Dikkomys_. The pattern of wear in the first two molars is the same as in the entoptychines of the early Miocene. The trend of evolution through which the _Pliosaccomys_ lineage passed must have featured a progressively earlier union at the edge of the tooth until the lateral coalescence occurred simultaneously with the median union. At that stage, emphasis was shifted to the union at the edge of the tooth, and eventually the teeth failed to unite at their mid-points and the U-pattern developed directly. Therefore, the horizontally deep re-entrant fold that separates the two lophs of the U-pattern is equivalent to one fold plus the apex of the opposite fold.

(3) The horizontal re-entrant fold of the U-pattern was remarkably shallow vertically and disappeared with little additional wear. Thus the two parts of M1, and also of M2, are united into a single column except for a slight inflection on the labial side and this is true also of m1 and m2 except for a slight inflection on the lingual side (Fig. 4G and K). The inflection appears to have persisted in the upper teeth (Fig. 4H), but evidently with slight wear, disappeared in the lower teeth (Fig. 4L). The final monocolumnar pattern was attained early ontogenetically, evidently before the permanent premolar had fully erupted; hence, the earlier stages occurred only in transition, persisted for only a brief interval in the teeth of juveniles, and the final stage developed in the young animal and lasted throughout the rest of its life in _Pliosaccomys_. In _Dikkomys_ the two columns never united into a single column, and a bilophodont occlusal pattern persisted throughout life.

The early phyletic development of the subfamily Geomyinae took place in the tribe Dikkomyini from the early Miocene into the early Pliocene. Compared with the rapid evolution of the specializations that distinguish the Entoptychinae, the structural changes in the early Geomyinae occurred at a remarkably slow rate. In fact the lineage changed but little from _Dikkomys_ to _Pliosaccomys_, in parts of the animal that can be compared, as illustrated by the low-crowned and rooted cheek teeth, the continuous enamel bands, the lack of grooving of the upper incisor, the retention of the primitive H-pattern, both above and below, in the premolar and third lower molar, and the ridges and fossae of the mandible to which the muscles of mastication attach. The only major changes detected in the known fragments are in the pattern of wear and the final configuration of the first and second molars, as described above. The unification of the two lophs in each of these two teeth into a single column was a significant step in the evolution of the Geomyinae, and is a stage between the primitive bilophodont pattern of the early and middle Miocene geomyines having continuously bicolumnar teeth and the monolophodont pattern in the modern pocket gophers of both lineages in which these teeth consist of a single column in all but the initial stages of wear. The monocolumnar structure of the first and second molars in the final stages of wear, therefore, is closer to that in the lineage of _Thomomys_ than it is to that of _Dikkomys_. Other specializations in the dentition of _Pliosaccomys_, especially in m1 and m2 where the H-pattern has been completely eliminated from the sequence of wear, are too far advanced for _Pliosaccomys_ to have given rise to the tribe Geomyini. The teeth in the immediate ancestor of the Geomyini must have been less specialized in m1 and m2, perhaps about as in _Dikkomys_. In the m1 and m2 of the tribe Geomyini, the H-pattern is formed in the initial stages of wear; therefore, in the early Pliocene ancestor, presently unknown in the fossil record, the H-pattern probably was present. Even so, the ancestor of the Geomyini and that of _Pliosaccomys_ probably were closely allied otherwise, and both probably had attained the highly specialized fossorial adaptations characterizing all modern pocket gophers, before the divergence of _Pliosaccomys_ and the Geomyini took place.

The evidence points to a major divergence of the geomyines that lived in the latest Miocene or the early Pliocene (probably the latter) and that gave rise to the two modern lineages, Thomomyini and Geomyini (see Fig. 3). One, the most primitive of the two, gave rise to the Thomomyini lineage that eventually evolved into _Thomomys_. _Pliosaccomys_ is closely allied to the ancestry of this lineage, although it is probably not the actual ancestor, as mentioned previously. Aside from the aforementioned specializations of the first and second molars, the features of the Thomomyini are less advanced than in the other specialized lineage (tribe Geomyini). Primitive traits retained in the tribe Thomomyini (and also characteristic of the ancestral tribe Dikkomyini) are: (1) Small size, in general no larger than the ancestral morphotype; (2) lack of grooving on the upper incisor (although a slight rudimentary groove is developed rarely in some living species); (3) retention of anterior and posterior enamel plates in lower and upper cheek teeth; (4) premolars having widely open re-entrant folds; (5) smooth and generalized skull lacking marked angularity, regosity or cresting (neither the sagittal nor the lambdoidal crest are ordinarily well developed except in _Thomomys bulbivorus_); (6) forefoot small, less modified for digging than in the Geomyini.

A-D. _Dikkomys woodi_, right lower tooth-row, including p4-m3.
Patterns based on No. P26284 (FMNH) from Upper Rosebud (Middle
Miocene), Shannon Co., South Dakota (B above).

E-H. _Pliosaccomys dubius_, left upper tooth-row, including P4-M2
(M3 unknown). Patterns based on Nos. 1798 and 1799 (LAM) from
Smiths Valley (Middle Pliocene), Lyon Co., Nevada.

I-L. _Pliosaccomys dubius_, right lower tooth-row, including p4-m3.
Patterns based on Nos. 1796 (holotype), 1804, and 1806 (LAM)
from Smiths Valley (Middle Pliocene), Lyon Co., Nevada.
]

The lineage of the Thomomyini is essentially rectilinear and without the major branching seen in the tribe Geomyini. The one genus, _Thomomys_, appears first in the Upper Pliocene (early Blancan time), and the specializations characterizing the lineage had already developed by that time. Evidently, the early stages of divergence from the ancestral stock resulted in the development of rootless, ever-growing, more hypsodont cheek teeth, simplification of M3, and enlargement of the masseteric ridge on the mandible. The enamel investment on the sides of the molariform teeth is interrupted owing to intrusion of tracts of dentine on the sides of each column. Even so, complete anterior and posterior plates are retained on all of the cheek teeth (Fig. 5, K and L) and there is no trend toward additional loss of enamel as in the Geomyini. The enamel on the sides of the column has little functional value, and its elimination probably reduces friction during the anteroposterior movements of the lower jaw, thereby increasing the efficiency of the cutting blades on the anterior and posterior wall of the tooth. The simplification of M3 was achieved by union of the two columns of the primitive pattern into a single column and obliteration of both the labial and lingual re-entrant folds in the first stages of wear. The adult tooth (see Fig. 5L) is without trace of the bilophate pattern and is not elongated; therefore, its structure is essentially the same as that of the first and second upper molars.

In the Thomomyini, the two lophs of the unworn molars unite entirely across the width of their surfaces with the first traces of wear (see Fig. 5, I and J), owing to the shallow and uniform depth of the transverse valley. In the molars, the final pattern is acquired, therefore, before the deciduous premolar has been replaced by the permanent tooth. A relatively shallow re-entrant inflection between the ends of the parameres sometimes is retained, although it also will disappear with slight additional wear. Therefore, both lophs tend to unite completely with the first stages of wear in the Thomomyini, thus omitting both U and H patterns from the sequence of wear. This is the highest degree of specialization attained in the Geomyidae in regard to the patterns of wear, since a sequence of bilophodont patterns appear in both the Dikkomyini and Geomyini before the monoprismatic pattern is developed.

A and E. _Geomys bursarius majusculus_, No. 2948 (KU), Douglas Co.,
Kansas. Right upper (A) including DP4-M3; lower left (E)
including dp4-m3.

B and F. _Pappogeomys bulleri burti_, No. 100444 (KU), 10 mi. NNW
Barra de Navidad, Jalisco. Right upper (B) including P4-M3;
right lower (F) including p4-m3 (both P4 and p4 with unworn
enamel caps).

C and G. _Pappogeomys bulleri albinasus_, No. 31044 (KU), 10 mi. S
and 8 mi. W Guadalajara, Jalisco. Right upper (C) including
P4-M3; right lower (G) including p4-m3.

D and H. _Pappogeomys bulleri albinasus_, No. 31002 (KU), W side La
Venta, 13 mi. W and 4 mi. N Guadalajara, Jalisco. Right
upper (D) including P4-M3; right lower (H) including p4-m3.

I and J. _Thomomys talpoides bridgeri_, No. 6865 (KU), 2 mi. up Mink
Creek, Pocatella, Bannock Co., Idaho. Left upper (I),
DP4-M3; left lower (J), dp4-m3.

K and L. _Thomomys talpoides fossor_, No. 13205 (KU), Wasson Ranch,
3 mi. E Creede, Mineral Co., Colorado. Right lower (K),
p4-m3; left upper (L), P4-M3.
]

Relationship of the Geomyini with the ancestral Dikkomyini is most clearly demonstrated in the sequence of wear on the occlusal surfaces of the molars. As in all geomyids, the upper part of the crown is biprismatic in the newly erupted tooth, and the two columns are separated by an intervening valley. With slight attrition on the unworn enamel cap, the weakly developed cusps merge and form a transverse enamel loop on each of the two columns (see third molar in Fig. 5, A and E), each loop enclosing a core of dentine that had become exposed. The valley between the two columns is shallow, and upon further wear of the tooth, the two loops unite. The two columns become joined at different points in the upper and lower molars depending on the varying depth of the valley in different teeth. Therefore, upper and lower molars develop distinctly different occlusal configurations.

In the lower molars, the pattern characteristic of _Dikkomys_ (Fig. 4C) is preserved without significant modification, as illustrated in an immature specimen of _Geomys_ (see Fig. 5E). The H-pattern and modified H-pattern are developed in the same sequence of wear in the Geomyini. A juvenal female (not illustrated), KU 2931, provides an example of the intermediate H-pattern. In this specimen, the protolophid and hypolophid of the left m2 are united only at their mid-points, indicating that the pattern of wear occurs in the same sequence in the Geomyini as it did in the Miocene genus _Dikkomys_. After the two columns have become united at their mid-points, a secondary union is formed at the edge of their protomeres, thus enclosing the enamel fossette as illustrated in Figure 5E (this is the modified H-pattern mentioned above). However, the fossette itself is shallow and soon disappears with slight wear. At this stage, the occlusal configuration would be in a U-pattern (m1 in Fig. 5E). The lingual re-entrant fold is also shallow in vertical depth; therefore, it is obliterated by wear following the eradication of the labial fossette. Consequently, the two columns are united into one. In m3 (see Figs. 5E, F, and G), the two columns merge by progressive lateral expansion of the medial isthmus.

In the first and second upper molars, the two columns unite across the entire surface of their protomeres from near the lingual edge of the crown to near its center. A minute inner inflection may be temporarily retained in some teeth. At this stage (see Fig. 5B), the parameres are still separated by the labial fissure, and the occlusal pattern is in the shape of a U, resembling, but not exactly duplicating, the pre-final pattern of Ml and M2 in the genus _Pliosaccomys_ (see Fig. 4H). The labial fissure is shallow, and, with further wear, the inflection is worn away and the parameres also unite, thereby forming a monoprimatic crown in the final stage. In M3, the two lophs first become united near the edge of their protomeres (see Fig. 5B), therefore forming a U-pattern similar to that developed in Ml and M2 of _Pliosaccomys_. The connection of the two lophs is not directly at the end of the protomere; consequently a shallow lingual inflection remains. The lingual edge of the valley is also shallow, and, with continued wear a second union of the two lophs takes place near the ends of their parameres, and the deeper, interior part of the valley remains as an isolated enamel fossette (see Fig. 5C). The two primary lophs of the tooth are now joined near both sides, having shallow lingual and labial re-entrant angles on the sides and the enamel island in the center. With continued effacement of the occlusal surface, the fossette will be eradicated, and the pattern of the occlusal surface will become the partially biprismatic pattern of the final stages (adult) of wear (see Fig. 5D). M3's of _Dikkomys_ and _Pliosaccomys_ are not known; however, it seems reasonable to assume that the pattern of wear in the M3 of Dikkomyini was not essentially different from that of the Geomyini, except that it is likely that the U-pattern of the second stage of wear in the Geomyini was probably the final stage in the genus _Dikkomys_.

Judging from the pre-final stages of wear, the dentition of the Geomyini provides a curious combination of patterns that resemble in part the Miocene genus _Dikkomys_ and in part the early and middle Pliocene genus _Pliosaccomys_. There is no significant variation in the premolars or third molars (at least in the lower teeth) of the Geomyinae from the early Miocene to late Pliocene; therefore, deviations of major significance are in the character of the first and second molars. In the Geomyini, the patterns of wear of m1 and m2 are the same as those of _Dikkomys_, and are distinctly different from those of _Pliosaccomys_ where the two columns first unite at the edge of their protomeres to form a U-pattern, rather than at their mid-points to form an H-pattern. Even though the intermediate stages of ontogeny in m1 and m2 of _Pliosaccomys_ and the Geomyini are entirely different, the bicolumnar crowns of both eventually unite, upon wear, into a single column. On the other hand, the patterns of M1 and M2 in the Geomyini most closely resemble those of _Pliosaccomys_, rather than _Dikkomys_. In this regard it should be pointed out that the upper molars of _Dikkomys_ are presently represented by only one tooth, an M1 in an early stage of wear. As described already, the patterns of M1-2 evidently would be mirror images of m1-2 in corresponding stages of wear. However, the initial union of the two columns, in the M1 that is known, is somewhat to the lingual side of center and the relatively small lingual valley does not reach the base of the crown, indicating, that eventually with wear, the two columns of _Dikkomys_ might have become united across the entire surface of their protomeres as in _Pliosaccomys_. Even so, the two columns of M1 do initially join closer to their mid-points than they do in _Pliosaccomys_, and, if they did actually unite across their protomeres, the union would have occurred with subsequent wear. That is, the first occlusal pattern would be H-shaped (but with the connection closer to the lingual than the labial side), as in m1 and m2, and it would become U-shaped only after additional wear. This sequence of patterns of M1 and M2, as already pointed out, does not pertain in _Pliosaccomys_ or the Geomyini, since the U-pattern is formed with the first union of the two columns at the edge of their protomeres, and the primitive H-pattern is never developed, unless one counts the slight lingual inflection, that occasionally is formed just after the two columns unite, as being indicative of the primitive pattern. As in the lower teeth, the bicolumnar crowns of early ontogeny in both _Pliosaccomys_ and the Geomyini become eventually united, with wear, into a single column.

Based upon the foregoing evidence, it would seem likely that the Geomyini evolved from an early Pliocene (perhaps late Miocene) Dikkomyini ancestor that had evolved the specializations of M1 and M2 that characterize its relative, _Pliosaccomys_, but had not also evolved the specializations of m1 and m2 that distinguish _Pliosaccomys_. Therefore, the ancestor of the Geomyini differed from the _Pliosaccomys_-Thomomyini lineage in its retention, unmodified, of the primitive patterns in m1 and m2 that characterized the earliest known Geomyines (_Dikkomys_). The same patterns are preserved in m1 and m2 of its modern descendents, the living Geomyini. In the _Pliosaccomys_-Thomomyini lineage the pattern of m1 and m2 are entirely different, as described above.

The earliest record of the Geomyini is the extinct genus _Pliogeomys_ (see Fig. 6) in the latest Hemphillian (middle Pliocene) and earliest Blancan (late Pliocene). _Pliogeomys_ is more primitive than any modern genus of the Geomyini, seems to have been a late survivor of the primitive stock, but was itself probably a collateral lineage and not on the direct line of descent. The cheek teeth in _Pliogeomys_ are rooted and less hypsodont than in the late Pliocene examples of the modern genera, and the anterior enamel plate of the lower molars shows no indication of reduction, as would be expected if _Pliogeomys_ were in the direct line of evolution. Separation of _Pliogeomys_ from the main stem of the Geomyini probably occurred after several specializations had already been achieved by the Geomyini. Two inheritances might have been grooving on the upper incisors and some reduction in amount of enamel on the sides of the cheek teeth. The dentine tracts on the sides of the cheek teeth of _Pliogeomys_ are narrow (see Fig. 7A) and barely separate the enamel blades and there is no discernible reduction in the anterior enamel blades on its lower molars. Those blades evidently were lost in the main lineage before the Pleistocene radiation of the living genera took place. _Pliogeomys_ is in an intermediate stage in evolution, and was not so advanced as was the main lineage at the time _Pliogeomys_ died out. Its structure does provide clues as to phyletic development that took place in the main lineage.

Specialized trends in the early phylogeny of the Geomyini included: development of rootless, ever-growing cheek teeth and an increase in hypsodonty; loss of the bicolumnar structure of the first and second molars, and, consequently, the formation of a single elliptical column in the final stage of wear; interruption of the enamel investment of the molariform teeth and formation of anterior and posterior enamel plates; and enlargement of the masseteric ridge and fossa. Each of these trends occurred independently in the Thomomyini, and each is an example of parallelism in the phyletic evolution of the two lineages. Three additional specializations lacking in the Thomomyini are the grooving on upper incisors, loss of anterior enamel plate in lower molars, and development of a basitemporal fossa on the mandible. Evidently, two grooves evolved in the ancestral incisors in the same bisculcate pattern preserved in _Pliogeomys_, _Zygogeomys_ and _Geomys_. The innermost groove is weakly developed in _Pliogeomys_, suggesting that this character was in an intermediate stage of evolution in the ancestral lineage at the time that _Pliogeomys_ split off. Numerous other specializations in the Geomyini appeared later, but evolved in the different genera that diverged from the ancestral lineage and are discussed separately in the next account. Only two of the major features characterizing the Dikkomyini are retained in the Geomyini: the H-pattern on the occlusal surface of the m1 and m2 developed during the initial stages of wear, and the bicolumnar pattern of M3. Adaptive radiation produced the living genera of the Geomyini in the late Pliocene and early Pleistocene (see Fig. 6) and subsequent specialization of the ancestral morphology followed.

Parallelism in the molars of later geomyines and the Entoptychinae is illustrated by the lateral interruption of the enamel investment and loss of enamel plates and by the omission of the H-pattern stage in the first and second molars (in _Pliosaccomys_). Resemblance of dentitions in certain stages of wear in _Pliosaccomys_ and in entoptychines led some investigators, for instance, Hibbard (1953:357), to suggest that _Pliosaccomys_ descended from one of the less specialized entoptychines, possibly _Grangerimus_ but probably _Gregorymys_. Actually, the highly specialized upper and lower premolars and third molars of the entoptychines rule them out as ancestors of the later geomyines. The evolution of entoptychine-like features in _Pliosaccomys_ is regarded as an example of iteration, a pattern of parallelism (see Simpson, 1953:248-253) where an allochronic and independent lineage undergoes the same evolutionary trend that phyletically characterized an earlier lineage, usually after the latter has become extinct. In this case, the lineage giving rise to _Pliosaccomys_ passed through the same phyletic stages in its evolution in the early Pliocene (and possibly the late Miocene) as did the entoptychines in the late Oligocene and early Miocene.

Another parallelism by iteration, occurring in the middle and late Pliocene in both the Thomomyini and Geomyini, is the loss of enamel from the lateral surfaces of the cheek teeth, and, in the Geomyini only, the eventual loss of the anterior plate in the lower teeth and the posterior plate in the upper teeth. Both features were evolved more than an epoch earlier in the specialized entoptychid genus _Entoptychus_ of the lower Miocene. In _Entoptychus_, only the posterior plate of the lower molars and the anterior plate of the upper molars remained in the final stages of attrition, although a central enamel fossette, a remnant of the re-entrant fold, remained throughout life. Iteration is also expressed in the subfamily Geomyinae by the development of grooving on the upper incisor and the formation of the basitemporal fossa. A shallow but distinct basitemporal fossa occurs between the coronoid process and the third lower molar in the genus _Entoptychus_ and a sulcated upper incisor, a single shallow groove usually near the median border of the tooth, is found in the genus _Gregorymys_ of the subfamily Entoptychinae. Both features are regarded as advanced specializations in the tribe Geomyini, even though each was evolved in the entoptychines of the Lower Miocene.

The postcranial skeleton of living genera of pocket gophers, as befits animals that spend most of their life within underground burrows, are highly specialized for a fossorial life. Elements of the postcranial skeleton recovered from Lower Miocene deposits indicate that the entoptychines were only semi-fossorial (see Cope, 1884:857; Wood, 1936:4-5; Wilson, 1949:117-118). One of the basic trends of the entoptychines was towards greater fossorial adaptation; the skeleton of _Entoptychus_ shows a greater degree of fossorial adaptation than earlier genera of the subfamily. There is no reason to suppose that the geomyine genus _Dikkomys_, which lived at the same times as the entoptychines, had acquired any more advanced fossorial adaptations than had the entoptychines.

The most pronounced fossorial adaptations seem to have evolved only in the ancestral lineage of the modern geomyines, probably in the latter part of the Miocene and in the early Pliocene, before the modern Thomomyini and Geomyini diverged. Extreme fossorial adaptations in herbivorous rodents, such as those characteristic of the modern pocket gophers and their immediate ancestors, are thought to have evolved only in response to pronounced arid conditions. The Entoptychinae and evidently the early geomyines lived in environments that were either tropical or temperate, and under conditions more mesic than I would consider necessary to bring about selection pressure resulting in fossorial specializations. In late Oligocene and early Miocene, according to Axelroad (1958:433-509), arid conditions did not exist in the United States, and the only xerophytic environments in North America occurred on the Central Plateau of México. Moreover (Axelroad, _loc. cit._), arid conditions did not develop in the western United States until the early Pliocene. Geomyids evidently became extinct in this region at the close of the Middle Miocene, and none appear in fossil deposits in the western United States until the latest Lower Pliocene (Clarendonian). The reappearance of geomyids, _Pliosaccomys_, in the western United States coincides with a trend toward aridity and the northward movement of the Madro-tertiary geoflora into the Great Basin and Great Plains from its place of origin on the Central Plateau of México (Axelroad, _loc. cit._). Later, in the middle and later Pliocene, the Madro-tertiary geoflora gave rise to the modern xerophytic plants that now characterize the desert vegetation of North America.

The Madro-tertiary climax does not appear as a major flora until the Miocene, but probably originated earlier. According to Axelroad (_loc. cit._), this xerophytic flora evolved from elements of the Neotropical-tertiary geoflora that became adapted to arid conditions that developed in the rain shadow of the high mountains flanking the Central Plateau of México. Originally, the Madro-tertiary flora consisted of small trees, shrubs, and grasses. Although some elements of this flora moved northward in the late Miocene, the major part of it remained in México until the early Pliocene. In the western United States, mountain formation increased in intensity in the Pliocene and continued on into the early Pleistocene. As the mountains became more elevated, especially the Sierra Nevada and Cascade ranges, they blocked the prevailing winds from the Pacific Ocean and extensive aridity developed on their leeward side. As xeric conditions became widespread, the Madro-tertiary flora successfully occupied the drier regions of southern California, the Great Basin, and the western parts of the Great Plains.

While the Entoptychinae probably evolved in response to the Arcto-tertiary flora, the late Tertiary geomyines probably evolved in response to the Madro-tertiary geoflora on the Central Plateau of México. Some of these early geomyines, especially ancestors of the modern lineages, probably were pushed southward by competition with the more specialized entoptychines. Most geomyines were pushed out of the northern area of distribution, except for _Dikkomys_ that survived in association with the entoptychids throughout the early and middle Miocene. During this time, and probably continuing on into the late Miocene, the geomyines occurring to the south in México became adapted to the arid environments of the Madro-tertiary geoflora.

Of course, information is lacking about climates in several parts of the late Miocene and early Pliocene. When such information becomes available it conceivably could modify the hypothesis outlined immediately above.

The principal trend of evolution in these semi-fossorial rodents was toward more complete fossorial adaptation, and the pronounced fossorial features characteristic of the modern pocket gophers were perfected. This trend continued in response to the intense selection pressures in this arid environment. The principal structural characters effected were in the postcranial anatomy, especially in the skeletal and muscular systems. Consequently, it is not surprising that in skull and dentition, _Pliosaccomys_ differs but little from _Dikkomys_. Therefore, most of the basic structural specializations so far developed for subterranean existence probably had evolved by the time geomyines moved back north in the early Pliocene. Both modern lineages, the tribes Thomomyini and Geomyini, have essentially the same fossorial features, and it seems unlikely that these features were acquired independently in the relatively short period of time available to them after their divergence; probably they were inherited from a common ancestor. These probabilities indicate that the evolution of the fossorial specialization was in the later phyletic development of the tribe Dikkomyini.

Plio-Pleistocene radiation of Geomyini

Unlike the lineage of the Thomomyini that remained essentially rectilinear through out its history, the Geomyini in the late Pliocene and the early Pleistocene underwent adaptive radiation in a degree comparable to the earlier radiation of the Entoptychinae, and all of the later history of the tribe is dominated by the radiation--the resulting structural diversity. At least four lineages were produced by the Plio-Pleistocene radiation (see Fig. 6); each originated at essentially the same time (late Pliocene) presumably from the same ancestral stock. Each of these lineages within the Geomyini has given rise to one of the four modern genera: _Zygogeomys_, _Geomys_, _Orthogeomys_, and _Pappogeomys_.

_Morphotype_

The immediate, unknown, ancestor probably lived on the Central Plateau of México. After the radiation began the ancestors of _Geomys_ and _Zygogeomys_ extended their ranges northward.

Features of the hypothetical morphotype, that would permit derivation of the modern genera would include the following: (1) Skull generalized, neither excessively long and narrow or short and broad; (2) skull smoothly rounded, without pronounced angularity, rugosity or cresting (sagittal crest probably lacking, even in old individuals); (3) zygomata slender, without lateral platelike expansions; (4) rostrum moderately broad; (5) upper incisors bisulcate, two grooves in pattern found in _Pliogeomys_, _Zygogeomys_ and _Geomys_; (6) lateral re-entrant angles of premolars obtuse; (7) p4 having four enamel plates (one on anterior wall, one on posterior wall, and two lateral plates) and lower molars having one enamel plate on the posterior wall of tooth (anterior plate is lacking); (8) P4 having four enamel plates, in same pattern as described for p4, M1 having two enamel plates (one anterior and one posterior), M2 same as M1, M3 having three plates (one anterior, two lateral on sides of posterior loph, none posterior); (9) M3 subtriangular in cross-section, distinctly bicolumnar, two columns marked by shallow re-entrant folds and connected by broad isthmus; (10) masseteric ridge large, forming high crest bordering masseteric fossa; (11) basitemporal fossa shallow; (12) angular process of mandible short, its lateral projection barely exceeding that of zygomatic arch.

_Specializations in Genera_

In relation to the primitive morphotype, increase in size, simplification of dentition, and changes in shape of skull are regarded as specializations. Considerable parallelism between the four lineages is seen. But each lineage is distinguished by a combination of specialized features, and three by a few unique specializations.

Among trends resulting in simplification of the dentition, reduction of enamel on the posterior wall of the upper cheek teeth has occurred in various degrees in all lineages of the Geomyini even to loss of all enamel on the posterior wall of the premolars and molars in two genera. Loss of some enamel is more common on P4 than on M1-2, and has occurred in all genera (see Figs. 7 and 9.)

In evolutionary sequence loss of enamel from M1 and M2 usually occurs after, but never preceding, the reduction of enamel on P4. Loss of enamel plates from the posterior face of M1 and M2 is associated with the evolution of an efficient anterotransverse shearing action of the teeth.

On the anterior wall of those teeth no reduction of the cutting blade has been observed; a complete anterior plate is retained in all living Geomyini.

Presence of both the posterior and anterior plates decreases the efficiency of transverse shearing, by providing two upper plates (anterior plate of one tooth and posterior plate of the preceding tooth) over which the lower cutting blade _simultaneously_ must pass with each movement. The advantages of shearing over the more common mechanics of planing are largely lost unless the posterior plates are eliminated. Also, none of the living Geomyini have retained a definitive posterior enamel plate on M3, the last upper molar; but two well-developed lateral plates, that extend almost all of the way back to the posterior apex of M3, have been retained, and, together function as a posterior plate. Loss of either or both of the lateral plates of M3 is rare, and occurs only in old individuals. Their loss in the final stages of wear may represent the beginning of a new trend in those species where it occurs (the _castanops_-group of the subgenus _Cratogeomys_). In any case, reduction of enamel takes place by transverse shortening of the plate through the complete loss of enamel on one end, the diminution beginning first on the labial end and proceeding by progressive atrophy to the lingual end of the plate. Evidently, when enamel has been eliminated from the labial end of a plate, the rate of loss decreases markedly, and the last stages of evolution, terminating in complete loss of an enamel plate, occurs more slowly. Evolution may be arrested before complete loss has occurred, and that part of the enamel that remains forms a short, vestigial plate restricted to the lingual one-fourth or one-third of the wall. The enamel pattern of the lower dentition is the same in all of the diverging lineages, with no evidence of additional loss of enamel from that which had already occurred in their common ancestor (see Figs. 7 and 9). Reduction and loss of enamel plates began and was terminated in the lower dentition before reduction began in the upper dentition.

Other dental specializations have occurred in the shape of the third upper molar and in the pattern of grooving in the upper incisor. Unlike M3 of the Thomomyini, that of the Geomyini differs in shape from M2, and its enamel investment differs from that of M2. Primitively, M3 was probably subtriangular in cross-section, and the posterior loph evidently projected posteriorly as a short, rudimentary heel that formed the apex of the triangle. Other shapes of M3 are considered to be specializations that have been derived from the primitive form. In addition to the primitive subtriangular pattern, the M3 of living Geomyini may be suborbicular, quadriform, elongate, or obcordate in shape. Usually each lineage is characterized by only one pattern, but in one genus (_Pappogeomys_) all patterns occur. Of the different forms, the elongate and obcordate seem to be the most highly specialized deviations from the triangular-shaped tooth. The bicolumnar pattern is accentuated in the elongate type (Fig. 7D, F, H) by deep lateral re-entrant folds, on both the lingual and labial sides, and by the elongation of the posterior loph into a pronounced heel. Teeth having this pattern have been illustrated by Merriam (1895:76-82) in Figures 27 (6 and 7), 28 (c and d), 34 (7 through 15), and 35 (8).

A. _Pliogeomys buisi_, No. 29157 (UMMP), holotype, Buis Ranch
(Upper Middle Pliocene), Beaver Co., Oklahoma. Right lower,
p4-m2 (m3 unknown).

B and C. _Zygogeomys trichopus trichopus_, adult female, No. 51971
(FMNH), Mt. Tancítaro, 10,500 ft., Michoacán. Left upper
(B), P4-M3; right lower (C), p4-m3.

D and E. Subgenus _Orthogeomys_. _Orthogeomys grandis guerrerensis_,
adult female, No. 39807 (KU), 1/2 mi. E La Mira, 300 ft.,
Michoacán. Left upper (D), P4-M3; right lower (E), p4-m3.

F and G. Subgenus _Heterogeomys_. _Orthogeomys hispidus hispidus_,
adult female, No. 23975 (KU), 4 km. W Tlapacoyan, 700 ft.,
Veracruz. Left upper (F), P4-M3; right lower (G), p4-m3.

H and I. Subgenus _Macrogeomys_. _Orthogeomys heterodus
cartagoensis_, adult female, No. 60664 (KU), Rancho
Redando, Volcán Lrozá, Prov. San José, Costa Rica. Left
upper (H), P4-M3; right lower (I), p4-m3.
]

The subcordate form is characterized by pronounced anteroposterior compression, and retention of a distinct labial re-entrant fold. The posterior loph apparently has been rotated in such a way that what was previously its posterior border now lies on the outer margin of the tooth; therefore, the axis of the posterior loph is strongly oblique in relation to the anteroposterior bearing of the maxillary tooth-row, and the median enamel plate also has been rotated and so lies transversely across the posterior wall of the tooth. Owing to the rotation of the posterior loph, the apex of the obcordate tooth is at its lingual side. The subcordate type is illustrated by Merriam (_loc. cit._) in Figures 27 (3 and 4), 28 (a and b), 34 (3 and 4), and 35 (5, 6, and 7). The suborbicular and quadriform types are less specialized than the two described above. Both are characterized by reduction, often obliteration, of the bicolumnar pattern of the subtriangular ancestral form, especially marked by the decrease in depth of the lateral re-entrant folds and the decrease in length of the posterior projection of the posterior loph. With these changes, the tooth becomes essentially monocolumnar, its occlusal surface oval in outline in one and squarish in shape in the other. Occlusal views of the suborbicular form are presented by Merriam (_loc. cit._) in Figure 33 (1, 5, 6, 7, 11, and 12) and the quadriform tooth is depicted in Figure 29. Grooved upper incisors are characteristic of the living Geomyini, but variation occurs in the number of grooves, and, if only one groove is present, its position on the anterior face of the tooth varies. Except for the previously mentioned (p. 480) abnormal tooth having three grooves, incisors with no more than two grooves are found in these pocket gophers, and this number of grooves is taken to be primitive. Loss of one or the other of the two grooves of the bisulcate pattern, therefore, is regarded as specialization. However, complete loss of both grooves never occurs in the Geomyini. Each of the four major lineages is characterized by one of the three patterns of grooving, and the particular groove-pattern is remarkably stable in each group.