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Eocene Pachynolophinae (Perissodactyla, Palaeotheriidae) from China, and Their Palaeobiogeographical Implications

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Eocene Pachynolophinae (Perissodactyla, Palaeotheriidae) from China, and Their Palaeobiogeographical Implications [Palaeontology, Vol. 60, Part 6, 2017, pp. 837–852] EOCENE PACHYNOLOPHINAE (PERISSODACTYLA, PALAEOTHERIIDAE) FROM CHINA, AND THEIR PALAEOBIOGEOGRAPHICAL IMPLICATIONS by BIN BAI1,2 1Key Laboratory of Vertebrate Evolution & Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology & Paleoanthropology, Chinese Academy of Sciences, Beijing, China 2State Key Laboratory of Palaeobiology & Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China; [email protected] Typescript received 29 March 2017; accepted in revised form 27 June 2017 Abstract: The Eocene perissodactyl family Palaeotheriidae China, in which the complete dentition is known. Qianohip- has traditionally been considered to be a nearly endemic pus is characterized by a molariform P2 and non-molariform European group within Equoidea, but a few palaeotheres P3–4; a relatively high degree of lophodonty; the absence of have been reported from Asia. Here, I reanalyse a maxilla mesostyles; an angular bending in the protoloph on P3-M3 containing M1–3 from the Lunan Basin, Yunnan Province, and the metaloph on M1–3 at the paraconule and meta- China. This element was initially assigned to a new tapiro- conule, respectively; and weakly developed ‘metastylid’ on morph species, Lophialetes yunnanensis, but is here placed in the lower cheek teeth. A cladistic analysis supports a close a new genus Lophiohippus within Pachynolophinae based relationship between Lophiohippus yunnanensis and Paran- mainly on the absence of mesostyles, the strongly oblique chilophus, and suggests that Qianohippus is closely related to metalophs, the strong development of lophodonty, parastyles some derived pachynolophs. The appearance of the pachyno- overlapping metastyles of preceding teeth and situated mesial lophins Lophiohippus and Qianohippus in China supports the to the paracone, and the fact that M3 is longer than wide existence of a biogeographical connection between Europe and has a large and buccally deflected metastyle. Lophiohip- and Asia in the Middle-Late Eocene, and the dispersal route pus differs from European Anchilophus and Paranchilophus in was probably along the Tethyan microcontinents in the that the parastyles are situated mesial or even slightly lingual, south. rather than mesiobuccal, to the paracones, and M3 is mark- edly relatively larger than M1. I further reanalyse Qianohip- Key words: Palaeotheriidae, Pachynolophinae, Lophiohip- pus magicus from the Shinao Basin of Guizhou Province, pus, Qianohippus, phylogeny, palaeobiogeography. P ALAEOTHERIIDAE was historically regarded as a group of Eocene Lingcha Formation, whereas the others are from equoids that flourished in Palaeogene Europe but also Middle–Upper Eocene strata. The Chinese palaeotheriid had a scattered distribution in China. Traditionally, material, in contrast with the numerous complete skulls Palaeotheriidae was thought to be the sister group to and skeletons from Messel in Germany and other locali- Equidae, the two clades collectively forming the super- ties in Europe (Franzen 2010), is mainly limited to tooth- family Equoidea (Franzen 1989; Froehlich 1999; Rose bearing maxillae and mandibles. 2006). Recent phylogenetic work, however, has suggested The holotype of ‘Lophialetes’ yunnanensis Huang & Qi, that equids are more closely related to tapiromorphs than 1982, a species of the well established genus Lophialetes to palaeotheriids, although palaeotheriids have been rep- (Matthew & Granger 1925; Radinsky 1965), is a maxilla resented in the recent analysis only by Pachynolophus with three molars (IVPP V 6508) that was only briefly eulaliensis (Bai et al. 2014). Known palaeotheriids from described when the species was first erected. The speci- China include Propalaeotherium sinense from Xintai Basin, men was collected from the Lumeiyi Basin, Yunnan Pro- Shandong Province (Zdansky 1930), Propachynolophus vince by an IVPP field crew in 1970. I am skeptical of the hengyangensis from the Hengyang Basin, Hunan Province referral of this maxilla to Lophialetes, and suggest that it (Young 1944; Li et al. 1979) and Qianohippus magicus represents a palaeotheriid rather than a tapiromorph, from the Shinao Basin, Guizhou Province (Miao 1982; based on reexamination of the type specimen. Accord- Fig. 1). Propachynolophus hengyangensis is from the Lower ingly, the present paper describes IVPP V 6508 in more © The Palaeontological Association doi: 10.1111/pala.12319 837 838 PALAEONTOLOGY, VOLUME 60 FIG. 1. E 90° 100° 110° 120° Distribution of Palaeotheriidae at Palaeogene locali- ties in China. 1, ‘Propachynolophus’ hengyangensis, Lower Eocene, Heng- yang Basin, Hunan Province; 2, Lophiohippus yunnanensis, Middle 40° Eocene, Lunan Basin, Yunnan Pro- Beijing vince; 3, ‘Propalaeotherium’ sinense, Middle Eocene, Xintai Basin, Shan- dong Province; 4, Qianohippus mag- 3 icus, Upper Eocene, Shinao Basin, Guizhou Province. 30° 4 1 N 20° 2 detail, and reanalyses its phylogenetic relationships. Here, The phylogenetic analyses presented in this paper were I also take the opportunity to redescribe and reanalyse conducted with a parsimony criteria in PAUP 4.0 (Swof- Qianohippus magicus, a palaeotheriid that has rarely been ford 2002) and TNT (Goloboff et al. 2008). The matrix mentioned by researchers since its original description by constructed for analysis was based on data from Danilo Miao (1982) even though the dentition is completely et al. (2013), with ‘Lophialetes’ yunnanensis and Qianohip- known in this species. Finally, the palaeobiogeography pus magicus added, and contained a total of 28 taxa and of palaeotheriids is discussed based on a reanalysis of 72 characters (Appendix 1; Bai 2017). All characters were Middle–Late Eocene palaeotheriids from China. unordered and equally weighted. The heuristic search algorithm was used with 1000 replications of random stepwise addition and tree-bisection-reconnection (TBR) MATERIAL AND METHOD branch swapping. The specimens of ‘Lophialetes’ yunnanensis (IVPP V 6508) Institutional abbreviations. AMNH FM, American Museum of and Qianohippus magicus (IVPP V 6519–6520) are housed Natural History, Fossil Mammal, New York; IVPP, Institute of in the Institute of Vertebrate Paleontology and Paleoan- Vertebrate Paleontology and Paleoanthropology, Beijing, China. thropology (IVPP), Chinese Academy of Sciences, Beijing, China. The upper dentition of the holotype (IVPP V 6519) of Qianohippus magicus is currently unavailable; the SYSTEMATIC PALAEONTOLOGY figure presented in this work is based on a cast (AMNH FM 144356) that was made from IVPP V 6519. Order PERISSODACTYLA Owen, 1848 Remy (2012) used the terms ‘crochet’ and ‘anticrochet’ Superfamily EQUOIDEA Hay, 1902 for the mesiobuccal end of the metaloph and a crista on Family PALAEOTHERIIDAE Bonaparte, 1850 the lingual side of the paracone, respectively. However, Subfamily PACHYNOLOPHINAE Pavlow, 1888 ‘crochet’ and ‘anticrochet’ have also been used for struc- Tribe ANCHILOPHINI Remy, 2012 tures on the metaloph and protoloph in rhinocerotoids. Genus LOPHIOHIPPUS nov. To avoid confusion and uncertainty regarding homolo- gies, I use the terms ‘postcrista’ and ‘crista’ as respective Type and only species. Lophiohippus yunnanensis (Huang replacements for ‘crochet’ and ‘anticrochet’ as used by & Qi, 1982). Remy (2012). Our usage is appropriate partly because the term ‘crista’ has previously been used for the structure on Derivation of name. Greek ‘lophia’ meaning crest, with the lingual side of the paracone in tapiroids and reference to the protolophs and metalophs of the upper rhinocerotoids (Radinsky 1965). molars lacking the paraconules and metaconules; and BAI: EOCENE PACHYNOLOPHINAE FROM CHINA 839 Greek ‘hippos’ meaning horse, a common used root in equid names. Type locality and horizon. As for the type and only spe- cies. Diagnosis. As for the type and only species. Lophiohippus yunnanensis (Huang & Qi, 1982) Figure 2 1982 Lophialetes yunnanensis Huang & Qi, p. 315, fig. 1, pl. 1, figs 4–5. LSID. urn:lsid:zoobank.org:act:83E11014-F0C4-474B-A779- A872888DDCB0 Holotype. IVPP V 6508, a left maxilla with M1–3. Type locality and horizon. Lumeiyi Formation (field local- ity number: 70002), Lunan Basin, Yunnan Province, China. Middle Eocene. Emended diagnosis (modified from Huang & Qi 1982). Upper molars highly lophodont, and gradually increasing in size from M1 to M3. Parastyles overlapping metastyles of preceding teeth, and situated mesial to the paracone. Paracone rib relatively narrow and hemicylin- drical. Metacone long and flat. Metaloph strongly oblique and joining paracone on moderately worn tooth. Metas- tyle of M3 large and deflected buccally. Differential diagnosis. Differs from Anchilophus and Paranchilophus in having the parastyles situated mesial or even slightly lingual, rather than mesiobuccal, to the para- cones, the parastyles overlapping the metastyles of the preceding teeth, the metastyle of M3 more buccally deflected, and M3 relatively pronouncedly larger than M1. FIG. 2. Left maxilla of Lophiohippus yunnanensis (IVPP V 6508) with M1–3. A, occlusal view. B, buccal view. C, Description lingual view. Scale bar represents 1 cm. M1–2. These teeth are generally similar in morphology (Fig. 2). However, M1 is roughly quadrate in outline and heavily worn, metacone is more lingually situated than the paracone and while M2 is larger, relatively mesiodistally longer on the buccal slightly lingually depressed, bearing a very weak rib on the buc- side and only moderately worn. The lingual side
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