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AMERICAN MUSEUM Norntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

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AMERICAN MUSEUM Norntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2812, pp. 1-20, figs. 1-6 April 10, 1985 Skull Morphology and Relationships of Geomyoid Rodents JOHN H. WAHLERT1 ABSTRACT Analysis of cranial and mandibular morpholo- Geomyinae, share common ancestry. The Ento- gy, especially the foramina and structures relating ptychidae are tentatively placed as the sister group to the masticatory musculature, supports mo- of these two families; the Florentiamyidae appear nophyly of the living Heteromyidae. The family to be the earliest known branch of the geomyoid includes two major divisions, the Heteromyinae clade. The Geomyoidea, which includes these on the one hand, and the Perognathinae and Di- families, and the extinct Eomyoidea comprise the podomyinae on the other. Derived features of infraorder Geomorpha. A classification of the Schizodontomys suggest that the genus may be an Geomorpha that reflects these phylogenetic hy- early member of the Dipodomyinae. The Heter- potheses is presented. omyidae and Geomyidae, containing only the INTRODUCTION Two basic theories on the interrelation- nial structures associated with the muscles of ships of geomyoid rodents have been pro- mastication; they have not resolved this con- posed. In the traditional view the geomyids troversy. (pocket gophers), and the heteromyids (kan- I have attempted to find cranial characters garoo rats and pocket mice) are two mono- that can be added to the diagnoses of groups phyletic groups that are descended from a ofliving geomyoids. The new evidence agrees common ancestor. In the alternative view re- with common, current subdivisions. The cent geomyids and their supposed relatives, family Geomyidae, subfamily Geomyinae, which are extinct, are derived independently contains four genera- Geomys, Orthogeo- from heteromyids. Published studies have mys, Pappogeomys, and Thomomys. The stressed morphology of the teeth and of cra- family Heteromyidae contains three subfam- 1 Research Associate, Department of Vertebrate Paleontology, American Museum of Natural History; Assistant Professor of Biology, Department of Natural Sciences, Baruch College of the City University of New York. Copyright © American Museum of Natural History 1985 ISSN 0003-0082 / Price $2. 1 0 2 AMERICAN MUSEUM NOVITATES NO. 2812 ilies that each include two genera: Hetero- the twentieth century. Wood (1935) pre- myinae-Heteromys and Liomys; Pero- sented a detailed phylogeny of heteromyids gnathinae-Chaetodipus and Perognathus; that included the known extinct and living Dipodomyinae-Dipodomys and Micro- taxa, and he discussed their relationship to dipodops. I examined the published descnp- geomyids. The first challenge to the tradi- tions of skulls of extinct taxa to find char- tional view was made by Wilson (1936); he acters that indicated relationships to these suggested that the extinct and living geomyid groups. Entoptychus, Pleurolicus, and subfamilies had separate ancestry in the Het- Schizodontomys have heretofore been re- eromyidae. Both hypotheses have been fur- garded as constituting one or more subfam- ther elaborated by later authors. ilies within the Geomyidae. Cranial evidence Wood (1935) proposed that the Dipodo- of the Florentiamyidae (Wahlert, 1983) and myinae and Perognathinae are each other's the Eomyoidea (Wahlert, 1978) is included closest relatives and that together they share in the discussion of phylogenetic relation- with the Heteromyinae a common ancestry ships; complete data are available in the pre- in the Oligocene genus Heliscomys. The crown vious publications. Supposed cranial remains morphology of geomyid cheek teeth could ofHeliscomys have been described; since their also be derived from that of Heliscomys, but generic assignments are uncertain (Wahlert, Wood believed the morphologic gap between 1983, 1984) I have omitted them. this genus and the John Day geomyids to be Rodents of the superfamily Geomyoidea so great and the time interval so short as to and its sister taxon, the extinct Eomyoidea, preclude the possibility of ancestor-descen- have a sciuromorphous lateral masseter. The dant relationship. Recent studies of the Het- geomyoids are distinguished by bilophodont eromyidae by M. S. Hafner (1982) and Haf- cheek teeth and the presence offur-lined cheek ner and Hafner (1983) support Wood's pouches that open lateral to the mouth in hypothesis of relationship among the living forms. The concept of the Geomyo- subfamilies. idea, its subdivisions and their contents de- A new subfamily ofheteromyids, the Flor- veloped gradually. Important early contri- entiamyinae, was described by Wood (1936a). butions were summarized by Coues (1877), The skull of Florentiamys loomisi, on which Wood (1935), and Russell (1968). the subfamily is based, is heteromyid-like but The Geomyinae are burrowers that range has many primitive features. The tooth crown from western Canada to Panama. Their fossil pattern appears more primitive than that of record extends back to the early Miocene and Heliscomys. Wood concluded (ibid., p. 47) is restricted to North America with the pos- that this florentiamyine species could not be sible exception of a Miocene fossil from "a direct ancestor ofthe heteromyids, though Shantung, China (Li, 1974). The Hetero- it might be a structural ancestor." Wahlert myidae are small, scampering and jumping (1983) described new specimens of florentia- rodents that range from southwestern Can- myids and found a combination of cranial ada to northern South America. Their fossil characters indicating that the group is neither record goes back farther in time, to the early ancestral to nor derived from heteromyids. Oligocene, and is entirely North American. Wood (1936b) discussed the relationships The living heteromyids have certain cra- of the Entoptychinae, an extinct geomyid nial and dental characters that are regarded subfamily ofmid-Tertiary age, and remarked as primitive among geomyoid rodents. The that "the evidence of the skulls is at least as oldest supposed heteromyid, Heliscomys, is strong for geomyine affinities as the evidence more ancient than other undisputed geo- of the teeth is for heteromyid ones" (ibid., p. myoid fossils. For these reasons systematists 4). He described a new genus, Dikkomys, as expect that the ancestors of all geomyoids representative of an early stage in differen- were heteromyid-like rodents. The tradition- tiation ofthe Geomyinae. This new material al phylogenetic hypothesis, that geomyids and did not serve to link the Geomyinae un- heteromyids are two monophyletic groups equivocally to a particular branch of the En- descended from a common, heteromyid-like toptychinae, but Wood found some points of ancestor, developed in the initial decades of similarity to the genus Pleurolicus. 1985 WAHLERT: GEOMYOID RODENTS 3 Russell (1968) revised the systematics of p. 66, fn.) stated that assignment of the En- the Geomyinae. He agreed with Wood on the toptychinae to the Geomyidae may be arti- ancestral position of Dikkomys. He derived ficial and that the family is then only a grade. the geomyines and entoptychines from a hy- He pointed out (ibid., p. 156) that dental sim- pothetical ancestor. Rensberger (1971, 1973a) ilarities suggest independent origin of the divided the early geomyids into two subfam- modern Geomyinae from the subfamily Het- ilies, the Entoptychinae and Pleurolicinae, eromyinae. and he added (1973b) the Florentiamyinae In this paper I use aspects of cranial mor- to this lineage. In his phylogeny (ibid.) Hel- phology to investigate the interrelationships iscomys is shown as the common ancestor of of heteromyid and geomyid subfamilies. I geomyids and heteromyids. have grounded the study in recent taxa, be- In the alternative phylogenetic hypothesis, cause skulls are abundant, and the contents the Geomyinae are considered to be derived ofany foramen can be checked by dissection. from the Heteromyidae at a time later than This information is sufficient to test the mo- the origin of the Entoptychinae; the hetero- nophyly of the living heteromyids. The ex- myids, which include Heliscomys, are a para- tinct taxa are examined in the context of the phyletic stem group. Wilson (1936) described cranial evidence that defines living taxa and a late Miocene geomyine, Pliosaccomys; he the concepts ofcharacter polarity in the Geo- pointed out many heteromyid-like features morpha that were developed in previous ofthe dentition and suggested that the genus publications (Wahlert, 1978, 1983). may illustrate a morphological stage passed through by the modern Geomyinae. Since the earlier entoptychines are more specialized ACKNOWLEDGMENTS than this genus in certain respects, Wilson This research was funded by National Sci- proposed that they might represent an earlier ence Foundation grant no. DEB79-03286. branch from the central stock. The departments ofVertebrate Paleontology A second challenge to monophyly of the and Mammalogy at the American Museum Heteromyidae was posed by Shotwell (1967). of Natural History gave me free access to On the basis of new specimens, he described collections; together with the Department of the tooth crown patterns in Pliosaccomys as Biology at Franklin and Marshall College they a developmental step from that in Dikkomys. provided the facilities for the research. The Morphological similarities led him to con- Nikon dissecting microscope with camera lu- clude that the Geomyinae are closely related cida that I used for drawing
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