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New Fossil Suid Specimens from the Terminal Miocene Hominoid Locality of Shuitangba, Zhaotong, Yunnan Province, China

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New Fossil Suid Specimens from the Terminal Miocene Hominoid Locality of Shuitangba, Zhaotong, Yunnan Province, China Journal of Mammalian Evolution (2019) 26:557–571 https://doi.org/10.1007/s10914-018-9431-3 ORIGINAL PAPER New Fossil Suid Specimens from the Terminal Miocene Hominoid Locality of Shuitangba, Zhaotong, Yunnan Province, China Sukuan Hou1,2,3 & Denise F. Su2 & Jay Kelley4 & Tao Deng1,3 & Nina G. Jablonski5 & Lawrence J. Flynn6 & Xueping Ji7,8 & Jiayong Cao9 & Xin Yang 10 Published online: 14 March 2018 # Springer Science+Business Media, LLC, part of Springer Nature 2018, corrected publication April/2018 Abstract Fossil suid specimens recovered from the latest Miocene site of Shuitangba, Zhaotong Basin, Yunnan Province, provide new information on the classification and relationship of Chinese Miocene Suinae. Most of the recovered specimens are referred to a relatively advanced and large species of Suinae, Propotamochoerus hyotherioides, based on dental dimensions and morphology. Detailed morphological comparisons were made between the Shuitangba Pr. hyotherioides and other Asian Miocene suines. From these comparisons, we suggest that Pr. hyotherioides from Shuitangba and northern China may be relatively derived compared to the specimens from Lufeng and Yuanmou, southern China and that Pr. hyotherioides and Pr. wui represent separate branches of the genus in China. Furthermore, Microstonyx differs from Pr. hyotherioides in p4/P4 and m3/M3 characters. Molarochoerus is suggested to represent a relatively derived taxon due to the uniquely molarized upper and lower fourth premolars. Miochoerus youngi is suggested to have a closer relationship to Sus and Microstonyx than to Propotamochoerus due to its small size and p4 morphology. Hippopotamodon ultimus, Potamochoerus chinhsienense, Dicoryphochoerus medius, and D. binxianensis exhibit complex morphologies that variously resemble Propotamochoerus, Microstonyx,andSus and are suggested to be possible transitional forms between Propotamochoerus, Microstonyx,andSus. However, the resolution of their classification requires further analysis when more material is recovered. Keywords Propotamochoerus . Suinae . Classification . Terminal Miocene . Shuitangba Abbreviations DTa Width of the First Lobe of a Cheek Tooth DAP Length of Premolar and Molar DTp Width of the Second Lobe of a Cheek Tooth DLL Labio-Lingual Diameter DTpp Width of the Third Lobe of the Third Molar and the DMD Mesio-Distal Diameter Deciduous Fourth Lower Premolar DT Maximum Width of a Cheek Tooth HHeight * Sukuan Hou 5 Department of Anthropology, The Pennsylvania State University, [email protected] University Park, PA 16802, USA 6 Peabody Museum of Archaeology and Ethnology, Harvard 1 Key Laboratory of Vertebrate Evolution and Human Origins of University, Cambridge, MA 02138, USA Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 7 Yunnan Institute of Cultural Relics and Archaeology, Beijing 100044, China Kunming, Yunnan 650118, China 2 Department of Paleobotany and Paleoecology, Cleveland Museum of 8 Natural History, Cleveland, OH 44106, USA Key Laboratory for Paleobiology, Yunnan University, Kunming, Yunnan 650091, China 3 Department of Earth Sciences, University of Chinese Academy of Sciences, Beijing 100049, China 9 Zhaotong Institute of Cultural Relics, Zhaotong, Yunnan 657000, 4 Institute of Human Origins and School of Human Evolution and China Social Change, Arizona State University, Tempe, AZ 85287-4101, USA 10 Zhaoyang Museum, Zhaotong, Yunnan 657000, China 558 J Mammal Evol (2019) 26:557–571 Introduction relationship of these enigmatic Miocene taxa from China. Here, we describe those fossil Suinae specimens Fossil Suidae are common in the late Miocene faunas of recovered from Shuitangba between 2007 and 2015 China. However, their classification and relationships are (Figs. 2, 3, 4, 5, 6, 7, 8,and9,Tables1, 2,and 3)and highly controversial. Among the late Miocene fossil suids, compare them with those from other Miocene sites in the medium- to large-sized Suinae, which have a China. Almost all the new fossils resemble Dicoryphochoerus-type p4, where the protoconid and the Propotamochoerus hyotherioides from Lufeng, Yunnan metaconid are separated, the metaconid is posterolingual to Province (van der Made and Han 1994), and northern the protoconid, and the talonid is lower than the main cen- China (Pearson 1928), except for one isolated M3 (ZT- tral cusps (Schlosser 1903;Pearson1928;Li1963;Chang 2009-03-511; Fig. 5i) and three incisors (LI1 [ZT-2007-02- 1974;Liuetal.1978;Tangetal.1985;Dong1987;Han 024; Fig. 2d], LDI1 [ZT-2015-2418; Fig. 2e], and RI3 [ZT- 1987; van der Made and Han 1994;Chen1997;Pickford 2015-1820; Fig. 2f]). The M3 (DAP = 22.3, DT = 16.8; see and Liu 2001; Zhang et al. 2002;LiuandPan2003;Liuand below Abstract for Abbreviations) has a very weak talon, Ji 2004;Liuetal.2004; Pan et al. 2006;DongandZhang which makes it resemble an M2. However, its hypoconule 2014), may be the most complicated and controversial is proportionally large compared to that of M2 and compa- group in terms of their classification. Schmidt-Kittler rable to the hypoconule of M3; the talon, though very weak, (1971) established Dicoryphochoerini in addition to the is still proportionally larger than those found in M2s, and is tribes Potamochoerini, Hippohyini, and Suini of Thenius more posteriorly extended than those in M2s. Thus, it may (1970),basedonp4charactersofKorynochoerus, represent a small- to medium-sized suid and bears an inflated Microstonyx, and Dicoryphochoerus. Pickford (1993) later main cusp and a very weak talon. The three incisors (ZT- erected Propotamochoerini for Propotamochoerus, 2007-02-024, I1, DMD > 10, DLL = 6.4; ZT-2015-2418, Hippopotamodon,andMicrostonyx, which is a synonym of DI1, DMD = 7, DLL = 5.1; ZT-2015-1820, I3, DMD = 8.2, Dicorypochoerini. DLL = 5.6) have mesio-distal lengths within the range of Recently, abundant large-sized Suinae specimens were Pr. wui (I1, DMD = 10.2–12.8, DLL = 5.5–6.3; DI1, discovered from Shuitangba, a terminal Miocene site (6.5– DMD = 6.5, DLL = 3.6; I3, DMD = 12.2, DLL = 4.2), but 6.0 Ma) located in northeastern Yunnan Province (Ji et al. are less labio-lingually compressed, and the I1 crown is low- 2013; Jablonski et al. 2014;Fig.1). Most of these new er than those of Pr. wui. As these specimens are only isolated Suinae remains show great similarity to many of the teeth and appear to be different from the other specimens Chinese late Miocene large-sized Suinae and provide an recovered from Shuitangba, they are referred to Suinae opportunity to better understand the classification and indet. and are not considered further in this paper. Fig. 1 Site location map J Mammal Evol (2019) 26:557–571 559 Materials and Methods Tooth measurements follow van der Made (1996); tooth width was measured at the base of the crown at its widest part. The Pr. hyotherioides specimens described in this paper were recovered from Shuitangba between 2007 and 2015 (Figs. 2, 3, 4, 5, 6, 7, 8,and9,Tables1, 2,and3), including six Data Availability Statement maxillary fragments, three mandiblular fragments, and several isolated teeth, which are all housed in the Zhaoyang Museum All data generated or analyzed during this study are included in Zhaotong city, Yunnan Province. in this published article. The Shuitangba Pr. hyotherioides specimens were com- pared with those of medium-large sized fossil Suinae from both northern and southern China and with those of Systematic Paleontology Hippopotamodon cf. H. hyotheroides from Thailand (Pickford et al. 2004b) to better understand the relation- Order Artiodactyla Owen, 1848 ships among these medium- to large sized suids bearing a Superfamily Suoidea Cope, 1887 Dicoryphochoerus-type p4 morphology. Family Suidae Gray, 1821 Tooth orientation and dental nomenclature of common fea- Subfamily Suinae Gray, 1821 tures on all tooth crowns (main cusps and accessory cusps) Genus Propotamochoerus Pilgrim, 1926 mostly follow Pickford (1986, 1988). The work of Boisserie et al. (2010) was also used for crest nomenclature on premolars. Propotamochoerus hyotherioides (Schlosser, 1903) Fig. 2 Incisors of Propotamochoerus hyotherioides (a-c, g) and Suinae indet. (d-f) from Shuitangba. a Left i1 (ZT- 2015-0240), (1) labial, (2) lingual, (3) mesial, (4) distal, (5) occlusal; b Right i1 (ZT-2007-01-149), (1) labial, (2) lingual, (3) mesial, (4) distal, (5) occlusal; c Left i1 (ZT- 2015-0366), (1) labial, (2) lingual, (3) mesial, (4) distal, (5) occlusal; d Left I1 (ZT-2007-02-024), (1) labial, (2) lingual, (3) mesial; e Left DI1 (ZT-2015-2418), (1) labial, (2) lingual, (3) mesial; f Right I3 (ZT-2015-1820), (1) labial, (2) lingual, (3) occlusal; g Right I2 (ZT-2015-1166), (1) labial, (2) lingual. Scale bar equals 5 cm 560 J Mammal Evol (2019) 26:557–571 Fig. 3 Upper cheek teeth of Propotamochoerus hyotherioides from Shuitangba (ZT-2015- 1062). a Left DP3-M2*, (1) occlusal, (2) lingual, (3) buccal; b Right DP3, (1) occlusal, (2) buccal, (3) lingual; c Right DP4, (1) occlusal, (2) buccal, (3) lingual; d Right P1, (1) occlusal, (2) buccal, (3) lingual; e Right P3, (1) occlusal, (2) buccal, (3) lingual; f Right P4, (1) occlusal, (2) buccal, (3) lingual; g Right M1, (1) occlusal, (2) buccal, (3) lingual; h Right M2, (1) occlusal, (2) buccal, (3) lingual. Scale bar equals 5 cm. * Note that in 3a, the DP3 is improperly positioned against the broken surface of the DP4. There should be a gap between the distal margin of the DP3 and the broken surface of the DP4 that would have accommodated the missing anterior half of the broken DP4 Description root is compressed mesio-distally and slightly curved. There is a shallow median groove on both the mesial and the distal Permanent Incisors sides of the root. Only one isolated upper second incisor (Fig. 2g, Table 1)is Permanent Premolars and Molars preserved. It is compressed labio-lingually and elongated mesio-distally. There may be a tiny facet on top of the mesial There is one isolated upper first premolar (Fig. 3d, Table 2), side of the crown, but it is not clearly visible.
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