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Palaeontologia Electronica Taxonomic Diversity of Late Palaeontologia Electronica http://palaeo-electronica.org Taxonomic diversity of Late Cenozoic Asian and North American ochotonids (an overview) Margarita A. Erbajeva, Jim I. Mead, Nadezhda V. Alexeeva, Chiara Angelone, Sandra L. Swift ABSTRACT Ochotonids (pikas) are an ancient group of mammals originating in the Oligocene of Asia and flourishing in the Miocene of the Old World. During the Pliocene they reduced both their diversity and abundance, Only the Pleistocene genus Ochotona survived to the present. Their current geographic diversity is concentrated in Asia, where 28 species exist today. Outside of Asia, ochotonids are represented by one liv- ing species in Europe and two living species in North America. There were likely three main immigrations to North America since the Miocene: (1) at the beginning of the Miocene represented by the appearance of Oreolagus; (2) at the Miocene- Pliocene boundary represented by the appearance of Ochotona span- glei; and (3) during the Early Pleistocene with the appearance of Ochotona whartoni, and small pikas close to the “Ochotona pusilla group”. Extant, endemic North American species appear in the Pleistocene. Plain-Language Summary Margarita A. Erbajeva, Geological Institute, Siberian Branch, Russian Academy of Sciences, Ulan-Ude, Russia 670047. [email protected] Jim I. Mead, Department of Geosciences, East Tennessee State University, PO Box 70267, Johnson City, Tennessee 37614, USA. [email protected] Nadezhda V. Alexeeva, Geological Institute, Siberian Branch, Russian Academy of Sciences, Ulan-Ude, Russia 670047. [email protected] Chiara Angelone, Universita di Roma Tre, 00146 Roma, Italy. [email protected] Sandra L. Swift, Department of Geosciences, East Tennessee State University, PO Box 70267, Johnson City, Tennessee 37614, USA. [email protected] Key Words: Ochotonid diversity; Eurasia; North America; Miocene-Pleistocene PE Article Number: 14.3.42A Copyright: Society of Vertebrate Paleontology November 2011 Submission: 15 June 2007. Acceptance: 19 September 2011 Erbajeva, Margarita A., Mead, Jim I., Alexeeva, Nadezhda V., Angelone, Chiara, and Swift, Sandra L. 2011. Taxonomic diversity of Late Cenozoic Asian and North American ochotonid an overview). Palaeontologia Electronica Vol. 14, Issue 3; 42A:9p; palaeo-electronica.org/2011_3/25_erbajeva/index.html ERBAJEVA ET AL.: TAXONOMIC DIVERSITY OF OCHOTONIDS INTRODUCTION CLASSIFICATION OF THE FAMILY OCHOTONIDAE The family Ochotonidae (Lagomorpha) includes 16 extinct genera and one extant genus Five families are recognized in the order Lag- (Ochotona). Studies indicate that the ochotonids omorpha: Mimolagidae Erbajeva, 1986, Leporidae show general evolutionary trends in the cheek Gray, 1821, Palaeolagidae Dice, 1929, Prolagidae teeth. These trends include a shift toward rootless- Gureev, 1960, and Ochotonidae Thomas, 1897. ness and hypsodonty, increasing complexity in the The family Ochotonidae includes two subfamilies, 3 enamel structure of the P and P3, increasing Sinolagomyinae Gureev, 1960 and Ochotoninae depth of the hypostria of P4-M2, and widening of Thomas, 1897. The chronologic range of Sinolago- the talonids of P -M from narrow to wide (Erba- myinae is Oligocene to Miocene. Ochotoninae is 4 2 known from the Miocene – Recent. The classifica- jeva 1994). tion of genera (after Erbajeva 1994, modified with Morphological examination of cranial and new data from Sen, 2003) is as follows: postcranial skeletons of both extinct and living taxa indicates that the main external features of ochoto- Ochotonidae Thomas, 1897 nids remained invariable for a long time (Gureev Sinolagomyinae Gureev, 1960 1964; Polyakova and Erbajeva 1974; Erbajeva Sinolagomys Bohlin, 1937 1994). Current data suggest that the principal Austrolagomys Stromer, 1926 ochotonid adaptations developed in the Oligocene Kenyalagomys MacInnes, 1953 and have mostly been retained. Based on similar Oreolagus Dice, 1917 dental structures in fossil and modern pikas, it is Bellatona Dawson, 1961 likely that fossil forms had similar ecologic require- Heterolagus Crusafont, Villata, and ments (e.g., open landscapes, grass diet) (Gureev Truyols, 1955 1964; Erbajeva 1988; Mead and Spaulding 1995). Bellatonoides Sen, 2003 Ochotoninae Thomas, 1897 MATERIALS Marcuinomys Groizet, 1839 Recent and fossil specimens examined for Lagopsis Schlosser, 1884 this study belong to: Geological Institute of the Albertona Lopez-Martinez, 1986 Siberian Branch RAS, Ulan-Ude; Institute of Plant Alloptox Dawson, 1961 and Animal Ecology USC, Yekaterinburg; Zoologi- Paludotona Dawson, 1959 cal Institute RAS, Saint Petersburg, Russia and Proochotona Chomenko, 1914 Institute of Geology SB RAS, Yakutsk (Russia); Ochotonoides Teilhard de Chardin and American Museum of Natural History, New York; Young, 1931 Smithsonian Institution, Washington, D.C.; Carne- Pliolagomys Erbajeva, 1983 in Agadja- gie Museum of Natural History, Pittsburgh; Denver nian and Erbajeva, 1983 Museum of Nature and Science; and the University Ochotonoma Sen, 1998 of California, Museum of Paleontology (USA). Ochotona Link, 1795 The systematic organization of fossil species presented below is based on the morphology of THE DIVERSITY OF OCHOTONIDS AND THEIR preserved skulls, mandibles and teeth, and on bio- EVOLUTIONARY DEVELOPMENT metric studies. Most living species, in addition to Subfamily Sinolagomyinae the mentioned characters, were diagnosed accord- ing to external features such as body ratio or pel- The earliest ochotonids referred to the genus age. The comparative analysis of living and extinct Sinolagomys are recognized from Oligocene species is based mainly on the tooth structures and deposits in China and Mongolia (Bohlin 1937, in part on preserved fragments of skull and mandi- 1942; Gureev 1964). The genus flourished in east- bles for fossil forms. ern Asia from the middle Oligocene through early We follow the standard Ochotona dental mor- Miocene (Gureev 1964; Li and Qiu 1980; Tong phology outlined by Erbajeva (1988). In a whole 1989). Oligocene taxa are characterized by having more than 2000 specimens of living pikas and reduced roots, cheek teeth with closed crown approximately 650 specimens of fossil Ochotona bases, with rather high crown, hypostria of the were examined from mentioned above Institutions. upper cheek teeth nearly half of the tooth width, and talonids of lower teeth being much narrower trigonids (Erbajeva 1994, figure 2). 2 PALAEO-ELECTRONICA.ORG At the beginning of the Miocene ochotonids the end of Late Miocene three genera, Ochoto- were widespread. A number of new genera of noides, Ochotonoma, and Ochotona, appeared in Sinolagomyinae appeared in Europe (Heterola- Asia, and Pliolagomys appeared in Eurasia to mid- gus), Africa (Kenyalagomys and Austrolagomys), dle Pliocene (Kretzoi 1959; Sen 1998; Topacewski North America (Oreolagus), and Asia (Bellatona and Scorik 1977; Erbajeva 1988). At the beginning and Bellatonoides). Sinolagomys, represented by of Pliocene the geographic range of ochotonids new Miocene species, persisted in Asia. Heterola- shrank, with all African forms becoming extinct. gus and Austrolagomys appeared at the end of Concurrently, Ochotona migrated to North America early Miocene. All Miocene taxa differ from the Oli- (Ochotona spanglei Shotwell 1956) and to Europe gocene in their lack of cheek tooth roots, higher (Ochotona sp. from Maritsa) (de Bruijn et al. 1970). crowns, nearly equal widths of trigonids and talo- During the Pliocene Ochotonoides (three spe- nids on P4-M2,, and deeper hypostria in the upper cies), Pliolagomys (three species) and Ochotona teeth. (36 species) flourished. They were represented by Sinolagomys, Kenyalagomys, and Oreolagus a number of species having wide distributions in became extinct the end of the Middle Miocene Eurasia, from Hungary, Romania and Moldavia in (Dawson 1965; Janvier and Muizon 1975; Li and the west, to North China in the east. They differed Qiu 1980). Bellatona and Bellatonoides survived much in body size and tooth morphology. Ochoto- until the Late Miocene. By the end of the Late Mio- noides and Pliolagomys are characterized by com- cene all genera in Sinolagomyinae became extinct. plicated structures of the P3 (Erbajeva 1988, figures 21-23). These genera became extinct in the Subfamily Ochotonodnae Late Pliocene, and only Ochotona survives to the The earliest record of subfamily Ochotoninae present. The latter had a wide distribution in Eur- is from the early Miocene, after which members of asia from the Late Miocene through Pleistocene both ochotonid subfamilies evolved in parallel and in North America from the Early Pleistocene throughout the Miocene (Erbajeva 1994). through Recent (Erbajeva 1988). This genus with It seems likely that Ochotoninae derived from simple structure of P3 was represented by a num- primitive Bellatona-like ancestors described by Qiu ber of species differing by the variations of teeth (1996). All Ochotoninae are characterized by root- size (corresponding to body size), from large to less cheek teeth except Marcuinomys, which has medium and small among ochotonids (Erbajeva teeth with closed crown bases similar to Sinolago- 1994, figure 6; Erbajeva and Zheng 2005). mys kansuensis. However, Marcuinomys differs from the latter by having more a complex structure OCHOTONID DEVELOPMENT AND HISTORY IN 3 of the most diagnostic teeth (P3 and P ). By the EURASIA end of Early Miocene all ochotonid cheek teeth are The genus Ochotona was represented in Eur- rootless and hypsodont. asia by several species differing in
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