Part 2

of the Pocket Gophers of the

Subfamily Geomyinae

ROBERT J. RUSSELL

CONTENTS

PAGE

INTRODUCTION 477

MATERIALS AND ACKNOWLEDGMENTS 477

TAXONOMIC CHARACTERS 478 Prismatic character of molars 478 Character of enamel patterns 479 Grooving of incisors 480 Masseteric ridge and fossa 480 Basitemporal fossa 481 Specializations of skull 481

FOSSIL RECORD 484 Miocene 485 Pliocene 486 Pleistocene 490 Thomomys 492 Zygogeomys 496 Geomys 496 Pappogeomys 503 Orthogeomys 504

HISTORY OF CLASSIFICATIONS 505

CLASSIFICATION 512 Family Geomyidae 512 Subfamily *Entoptychinae 513 Genus *Pleurolicus 514 Genus *Gregorymys 514 Genus *Grangerimus 514 Genus *Entoptychus 514 Subfamily Geomyinae 514 Tribe *Dikkomyini 515 Genus *Dikkomys 516 Genus *Pliosaccomys 517 Tribe Thomomyini 518 Genus Thomomys 518 Subgenus *Pleisothomomys 519 Subgenus Thomomys 520 Tribe Geomyini 521 Genus *Pliogeomys 522 Genus Zygogeomys 523 Genus Geomys 525 Genus Orthogeomys 528 Subgenus Orthogeomys 529 Subgenus Heterogeomys 530 Subgenus Macrogeomys 531 Genus Pappogeomys 532 Subgenus Pappogeomys 534 Subgenus Cratogeomys 535

PHYLOGENY OF THE GEOMYIDAE 536 Primitive Morphotype 537 Entoptychid Radiation 540 Phyletic Trends in Subfamily Geomyinae 542 Plio-Pleistocene Radiation of Geomyini 558 Morphotype 559 Specializations in Genera 560 Zygogeomys 564 Geomys 565 Orthogeomys 568 Pappogeomys 569

LITERATURE CITED 572

INTRODUCTION

When C. Hart Merriam wrote his monograph of the subfamily Geomyinae in 1895, he had no opportunity to examine fossil specimens. No doubt his phylogenetic conclusions and classification would have been greatly influenced had he enjoyed that opportunity because study of fossil geomyids reveals the historic sequence of phyletic development, and this sequence provides a firm basis for distinguishing specialized from primitive characters. The history of the Geomyinae has been characterized by the evolution of specializations. These evolutionary trends begin, as we presently know them, with a generalized ancestral stock in the early Miocene. The direction, degree, and rate of change, beginning with the primitive morphotype of the subfamily, has not been the same in the various lineages. The classification within the subfamily is based upon the phyletic interpretations of available data and the relationships they disclose. In turn, a new, and I hope more realistic, phylogeny and classification is offered.

MATERIALS AND ACKNOWLEDGMENTS

Recent specimens were studied of all the known genera, subgenera and 29 of the 36 living species. Most of the species not studied are monotypic and have restricted geographic ranges. They are: Geomys colonus, G. fontanelus, and G. cumberlandius, Orthogeomys cuniculus and O. pygacanthus of the subgenus Orthogeomys, and O. dariensis and O. matagalpae of the subgenus Macrogeomys. Examination of these modern species would not radically change the estimation of the degree of phyletic development of the genera and subgenera involved. All of the major polytypic and widespread species were studied.

Specimens of the extinct genera Dikkomys, Pliosaccomys, Pliogeomys, Nerterogeomys, and Parageomys also were studied, as were examples of the extinct species Geomys quinni, Geomys tobinensis, and Orthogeomys onerosus. Considerable fossil material of living species, especially of the genera Geomys and Pappogeomys, was used.

Inasmuch as the present account concerns mainly structural changes in the subfamily Geomyinae at the level of subgenera and above, and the temporal sequence of those changes, no attempt is made in the present account to revise taxonomy below the level of subgenera. Considerable modification of the classification below that level (for species and subspecies) is to be expected in Orthogeomys and Pleistocene taxa of Geomys when available specimens are studied.

I thank Prof. Robert W. Wilson for his assistance in securing fossil geomyids for study, and those in charge of the paleontological collections at the California Institute of Technology, Prof. Bryan Patterson, formerly of the Field Museum of Natural History, and Prof. Claude W. Hibbard of the University of Michigan, Museum of Zoology. For their kindness in lending Recent species, I thank Mr. Hobart M. Van Duesen of the American Museum of Natural History, Dr. David H. Johnson of the U. S. National Museum, and Dr. Oliver P. Pearson of the California Museum of Vertebrate Zoology, the late Colin C. Sanborn of the Field Museum of Natural History, and Profs. Emmet T. Hooper and William H. Burt of the University of Michigan Museum of Zoology.

I am especially grateful to Prof. E. Raymond Hall for his guidance and helpful criticisms with the manuscript. For assistance with paleontological problems, I thank Drs. Robert W. Wilson and William A. Clemens. Several persons have offered helpful suggestions and encouragement in the course of my study. For assistance of various sorts I especially thank Drs. J. Knox Jones, Jr., Rollin H. Baker, A. Byron Leonard, Sydney Anderson, James S. Findley, Robert L. Packard, and Robert G. Anderson. Advice concerning the drawings of the dentitions was generously given by Mr. Victor Hogg, and the drawings were done by Mrs. Lorna Cordonnier under his direction and by Mr. Thomas H. Swearingen. For assistance with secretarial tasks I thank Valerie Stallings, Violet Gourd, Ann Machin, Toni Ward, Sheila Miller, and my wife, Danna Russell.

TAXONOMIC CHARACTERS

Morphological features of the fossils and their stratigraphic provenience provide the information upon which phylogenetic interpretations are based. Although the most critical sequences of the fossil record are lacking, and although the existing fossils are mostly fragmentary and therefore seldom furnish ideally suitable data for the interpretations that have been made, phylogenetic conclusions drawn from fossil materials are superior to those drawn on other bases. The especially relevant characters are those disclosing primary trends in the evolution of the modern assemblages. The higher systematic categories recognized in the following account are based primarily upon such characters.

The most important characters found are in the teeth, although several structural changes in the lower jaw, especially those associated with the insertion of cranial musculature, are almost as important.

Prismatic Character of Molars

In primitive geomyines the molar consisted of two columns united at their mid-points and forming a figure 8 or H-pattern (see Fig. 4B). Both labial and lingual re-entrant folds were formed between the two columns. The primitive pattern is retained in the premolars of all known Geomyinae. Therefore, in the earliest (Miocene) members of the subfamily, the pattern of the molars was essentially like that of the premolars.

In Pliocene Geomyinae the two columns of the molars tend to merge into one. This is evident on the worn occlusal surface of the teeth; the lateral re-entrant folds are shallow vertically and progressively recede laterally until only a slight inflection remains. In the final stages of attrition, the inflection disappears and the tooth is a simple elliptical column. In the Pleistocene the monoprismatic pattern appears at earlier stages of wear owing to the decrease in depth of the re-entrant folds, and in Geomyinae of Recent time the initial stages of wear on the enamel cap of infants erase the last vestiges of two columns in the molar teeth.

The general trend in evolution, therefore, has been from a bicolumnar to a monocolumnar pattern. The particular patterns of wear characterizing each genus are described in detail beyond.