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Fossil Imprint 3.2017.Indb FOSSIL IMPRINT • vol. 73 • 2017 • no. 3–4 • pp. 495–514 (formerly ACTA MUSEI NATIONALIS PRAGAE, Series B – Historia Naturalis) COMMENTS ON THE AGE AND DISPERSAL OF MICROTOSCOPTINI (RODENTIA: CRICETIDAE) We can know only that we know nothing. And that is the highest degree of human wisdom. Lev Nikolayevich Tolstoy, War and Peace LUTZ C. MAUL1,*, LEONID I. REKOVETS2, WOLF-DIETER HEINRICH3, ANGELA A. BRUCH4 1 Senckenberg Forschungsstation für Quartärpaläontologie, Am Jakobskirchhof 4, Weimar, Germany; e-mail: [email protected]. 2 Wrocław University of Environmental and Life Sciences, ul. C. K. Norwida 25, 50-375 Wrocław, Poland; e-mail: [email protected]. 3 Museum für Naturkunde – Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Germany; e-mail: [email protected]. 4 The Role of Culture in Early Expansions of Humans, Senckenberg Forschungsinstitut, Senckenberganlage 25, 60325 Frankfurt/M., Germany; e-mail: [email protected]. * corresponding author Maul, L. C., Rekovets, L. I., Heinrich, W.-D., Bruch, A. A. (2017): Comments on the age and dispersal of Microtoscoptini (Rodentia: Cricetidae). – Fossil Imprint, 73(3-4): 495–514, Praha. ISSN 2533-4050 (print), ISSN 2533-4069 (on-line). Abstract: The tribe Microtoscoptini, comprising the genera Microtoscoptes from Eurasia and Paramicrotoscoptes and Goniodontomys from North America, is an enigmatic group of microtoid cricetids, which was widespread during the Late Miocene. Although fossil remains have been reported from 33 localities, their evolutionary and dispersal history is still poorly understood. Here we give an overview of sites and records and discuss temporal ranges and some aspects of the dispersal history. The branch of cricetids that gave rise to the Microtoscoptini is still unknown. The currently oldest records are those of Microtoscoptes from Shala in China, considered 8 to 9 Ma and correlated with MN 10 or MN 11. All other remains from Eurasia are distinctly younger (MN 11 – MN 13). The earliest North American records of Paramicrotoscoptes and Goniodontomys are from the early Hemphillian beginning at 9.0 Ma (approximately MN 10 – MN 11 transition). Whether Microtoscoptes from Shala actually indicates the origin of Microtoscoptini in Asia and subsequent dispersal to North America is still unclear. The presence of Microtoscoptini in both Central Asia and North America during the Late Miocene does suggest dispersal through Beringia. The Microtoscoptini inhabited probably open grasslands locally differentiated by shrubs, tree patches and water bodies. It is far from clear why the Microtoscoptini became extinct even though they had developed an effi cient arvicoline-like cheek-tooth pattern. Key words: Microtoscoptes, Paramicrotoscoptes, Goniodontomys, Late Miocene records, dispersal Received: August 28, 2017 | Accepted: November 2, 2017 | Issued: December 31, 2017 Introduction There is a considerable similarity between the cheek- teeth of the Microtoscoptini and those of arvicolines (voles) Tolstoy’s phrase, the epigraph of this paper, previously owing to similar adaptations (for a summary, see Fejfar supposed to be taken from Socrates (who, however, probably et al. 2011). Also the molars of the Microtoscoptini are never said this), or from a thousand other wisemen, seems hypsodont, and they display a prismatic molar shape with particularly appropriate with regard to our knowledge of the Microtoscoptini, the enigmatic extinct tribe of so-called opposing or slightly alternating dental triangles. However, at microtoid cricetids or “vole-toothed hamsters” (Fahlbusch the fi rst lower molar the second and third triangles (counted and Mayr 1975, Fahlbusch 1996, Fejfar 1999, Fejfar et al. from distal side of the tooth) form a distinct rhombus – the 2011). Judging from the number of Microtscoptini-bearing “trademark” of the Microtoscoptini that is not seen in true localities (33), this group seems to be a well-known one. arvicolids or other branches of microtoid cricetids. However, considering our knowledge of its fossil history, Microtoscoptes, confi ned to Eurasia and the fi rst described there are still a lot question marks. genus of Microtoscoptini, comes from the Late Miocene DOI 10.1515/if-2017-0026 495 (Turolian) of Ertemte in northern China (Schaub 1934), datum, Hh – Hemphillian, LAD – last appearance datum, MN and it has been reported since then from several regions – Neogene Mammal Age Zone, NALMA – North American of Central Asia and Eastern Europe (e.g., Topachevskiy Land Mammal Ages, NMU – Neogene Mammal Unit. 1971, Zazhigin in Gromov and Polyakov 1977, Zazhigin in Pevzner et al. 1982, Savinov 1982, Zazhigin and Zykin 1984). Goniodontomys and Paramicrotoscoptes were Records of Microtoscoptini exclusively distributed in North America during the Late The records are listed in geographical order of the Miocene (Hemphillian) (e.g., Wilson 1937, Hibbard 1970, countries from east to west in Eurasia: China – Russia – Repenning 1987, Repenning et al. 1990). Mongolia – Kazakhstan – Ukraine and in North America. Because Microtoscoptes SCHAUB, 1934, Goniodontomys The numbers of the sites (loc. nr.) refer to those in Text-fi gs 1 WILSON, 1937, and Paramicrotoscoptes MARTIN, 1975 are and 4. In the morphological descriptions, lower case denotes morphologically rather similar, they seem to be closely lower teeth and upper case denotes upper teeth. The maps in related (Schaub 1940) and were grouped in one taxon: fi rst Text-fi gs 1 and 4 were created using ArcGIS® software by at subfamily level as Microtoscoptinae (Kretzoi 1955), then Esri. ArcGIS® and ArcMap™ are the intellectual property as a family Microtoscoptidae (Kretzoi 1969), and later as of Esri and are used herein under license. Copyright © tribe Microtoscoptini (Gromov and Polyakov 1977), the still Esri. All rights reserved. For more information about Esri® commonly accepted taxonomic rank (Fejfar et al. 2011). software, please visit http://www.esri.com. Although three genera with seven species have been named so far (Microtoscoptes praetermissus SCHAUB, China 1934, M. tjuvanensis ZAZHIGIN in GROMOV et POLYAKOV, 1977, M. pristinus SAVINOV, 1982, M. sibiricus ZAZHIGIN Ertemte 1 and 2 (loc. nr. 1) et ZYKIN, 1984, M. mongolicus ZAZHIGIN in PEVZNER et Location of the site: Northern China, Inner Mongolia (= al., 1982, Goniodontomys disjunctus WILSON, 1937 and Nei Mongol) Autonomous Region, Huade county, 4 km SE Paramicrotoscoptes hibbardi MARTIN, 1975), the fossil of the town of Huade, outskirts of village of Ertemte (Storch record of the Microtoscoptini is far from being satisfactorily and Zazhigin 1996). According to Fahlbusch et al. (1983b), investigated, nor even suffi ciently documented. Some of Ertemte 1 refers to the classical site excavated in the 1920s these species mentioned have not yet been adequately (Andersson 1923, Schlosser 1924, Schaub 1934), from described and illustrated. Therefore, many aspects of the where of Microtoscoptes was described (Schaub 1934), phylogeny, biostratigraphy and palaeobiogeography of these and Ertemte 2 refers to the site of the later rich material microtoid cricetids have been remained obscure until today. unearthed and investigated by Fahlbusch (Fahlbusch et al. For a long time the Microtoscoptini were to many 1983b, Fahlbusch 1987, Qiu Zhuding et al. 2013: 184). European paleontologists strange small mammals from the Fauna: Ertemte 1 (Schaub 1934, Fahlbusch et al. Far East. This view changed in the early 1970s, when a single 1983a, Qiu and Li 2016): Microtoscoptes praetermissus, tooth of Microtoscoptes was unearthed from Late Miocene associated with Erinaceomorpha: Erinaceidae: Erinaceus deposits exposed at the Ukrainian site Cherevychne 3 mongolicus, Soricomorpha: Talpidae: Yanshuella primaeva, (Topachevskiy 1971), which showed for the fi rst time that Soricidae: “Crocidura” kormosi, Lagomorpha: Leporidae: Microtoscoptini were distributed also in Eastern Europe. Alilepus annectens, Rodentia: Aplodontidae: Pseudaplodon Since then further sites have been discovered in the Ukraine asiaticus, Cricetidae: Nannocricetus mongolicus, that yielded numerous fi nds of Microtoscoptes that are Sinocricetus zdanskyi, Anatolomys teilhardi, Microtodon currently under investigation (Fejfar and the authors of the atavus, Dipodidae: Lophocricetus pusilllus, Lophocricetus present paper). There are also further recent discoveries grabaui, Paralactaga anderssoni, Brachyscirtites from Asia (e.g., locality Shala – Qiu et al. 1999, Li et al. wimani, Dipus fraudator, Spalacidae: Myospalacinae: 2003, Qiu Zhuding et al. 2013) that require modifi cations of Prosiphneus eriksoni, Muridae: Karnimatoides hipparionus, some previous conclusions about Microtoscoptini (Fejfar et Hansdebruijnia pusillus, Apodemus orientalis. al. 2011). Ertemte 2 (Qiu Zhuding et al. 2013, Qiu and Li The purpose of this paper is to summarise and discuss 2016): Microtoscoptes praetermissus, associated with current information on the available records and ages of Erinaceomorpha: Erinaceidae: Erinaceus mongolicus, the fi nds and the dispersal of Microtoscptini known so Soricomorpha: Talpidae: Quyania chowi, Yanshuella far. It is complementary to the studies on new material of primaeva, Soricidae: Sorex minutoides, Sorex ertemteensis, Microtoscoptes from the Ukraine, the description of which Sorex pseudoalpinus, Zelceina kormosi, Alloblarinella is in preparation. The present paper is devoted to Oldřich sinica, Paenepetenyia zhudingi, Cokia kowalskae, Fejfar and Rudolf Musil
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