旧大陆“Hippotherium”的存续:两种欧亚大陆三趾马的修订
收稿日期: 2024-04-19
网络出版日期: 2024-11-20
基金资助
国家重点研发计划(2023YFF0804501);中国科学院青年创新促进会项目(2021069);第二次青藏高原综合科学考察研究项目(2019QZKK0705)
Occurrence of “Hippotherium” in the Old World: a revision of two hipparion species in Eurasia
Received date: 2024-04-19
Online published: 2024-11-20
对一个具有争议的类群Hipparion plocodus进行了回顾。山西保德的头骨材料具有明确的鉴定特征,证实Hi. plocodus为一有效种。使用新的特征矩阵进行系统发育分析,表明Hi. plocodus和欧洲种Hippotherium malpassii构成一个单系群。事实上所谓的Hm. malpassii和Hippotherium属无较近的亲缘关系,而该属目前无晚中新世晚期分布记录。因此之前所定义的Hi. plocodus和Hm. malpassii在更好的材料发现之前暂时归入“Hipparion” 属当中。二者演化水平略有差异,时代相当,表明它们独立起源于某一更原始类群。在晚中新世晚期,亚洲夏季风的盛行加强了中国地区的湿润程度,森林和灌木林生境在这一背景下迅速扩张,最终令欧亚大陆某一喜好封闭环境的类群向中国扩散,在这片适应的土地上演化出了“Hi.” plocodus。
孙博阳 , 刘艳 , 王世骐 , 邓涛 . 旧大陆“Hippotherium”的存续:两种欧亚大陆三趾马的修订[J]. 古脊椎动物学报, 2025 , 63(1) : 57 -80 . DOI: 10.19615/j.cnki.2096-9899.241120
A controversial taxon, Hipparion plocodus, is reviewed in the present study. Hi. plocodus has been confirmed to be a valid species with definite diagnostic characteristics, represented by cranial specimens from Baode, Shanxi Province. The phylogenetic analysis performed in the present study, with a new matrix, shows that Hi. plocodus forms a monophyletic group with a European species, Hippotherium malpassii. Actually, no close relationship between so-called Hm. malpassii and the genus Hippotherium has been identified, and the record of stratigraphic range of this genus in late stage of Late Miocene is currently absent. Herein previously Hi. plocodus and Hm. malpassii have both attributed into “Hipparion” before the discovery of better material. Evolutionary stages and correlative absolute age showed that these two species should derive independently from some primitive clade. During the late stage of the Late Miocene, the development of the Asian summer monsoon enhanced the humidity of China, with forest and wood habitats expanding considerably under this setting. As the result, one Eurasian closed-habitat lineage thus extended its range into China, which had become very suited for it, give rise to “Hi.” plocodus.
Key words: Eurasian hipparion; systematic revision; phylogeny; paleozoogeography
[1] | Alberdi M T, 1989. A review of Old World hipparionine horses. In: Rrothero D R, Schoch R M eds. The Evolution of Perissodactyls. New York, Oxford: Oxford University Press. 234-261 |
[2] | Bernor R L, 1985. Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (Late Miocene, Turolian age). Palaeovertebrata, 15(4): 173-269 |
[3] | Bernor R L, Hussain S T, 1985. An assessment of the systematic, phylogenetic and biogeographic relationships of Siwalik hipparionine horses. J Vert Paleont, 5: 32-87 |
[4] | Bernor R L, White T D, 2009. Systematics and biogeography of “Cormohipparion” africanum, early Vallesian (MN 9, ca. 10.5 Ma) of Bou Hanifia, Algeria. In: Albright B ed. Papers on Geology, Vertebrate Paleontology, and Biostratigraphy in Honor of Michael O. Woodburne. Bull Mus North Arizona, 65: 635-658 |
[5] | Bernor R L, Qiu Z X, Tobien H, 1987. Phylogenetic and biogeographic bases for an Old World hipparionine horse geochronology. Proceedings of the VIIIth International Congress of the Regional Committee on Mediterranean Neogene Stratigraphy, Budapest. Ann Inst Geol Publ Hung, 70: 43-53 |
[6] | Bernor R L, Qiu Z X, Hayek L A C, 1990. Systematic revision of Chinese hipparion species desribed by Sefve, 1927. Am Mus Novit, 2984: 1-60 |
[7] | Bernor R L, Koufos G D, Woodburne M O et al., 1996. The evolutionary history and biochronology of European and southwestern Asian late Miocene and Pliocene hipparionine horses. In: Bernor R L, Fahlbusch V, Mittmann H W eds. The Evolution of Western Eurasian Neogene Mammal Faunas. New York: Columbia University Press. 307-338 |
[8] | Bernor R L, Tobien H, Hayek L A C et al., 1997. Hippotherium primigenium (Equidae, Mammalia) from the late Miocene of H?wenegg (Hegau, Germany). Andrias, 10: 1-230 |
[9] | Bernor R L, Kaiser T M, Kordos L et al., 1999. Stratigraphic context, systematic position and paleoecology of Hippotherium sumegense Kretzoi, 1984 from MN 10 (late Vallesian of the Pannonian Basin). Mitt Bayer Staatsslg Pal?ont Hist Geol, 39: 115-149 |
[10] | Bernor R L, Armour-Chelu M, Kaiser T M et al., 2003. An evaluation of the late MN9 (Late Miocene, Vallesian age), Hipparion assemblage from Rudabánya (Hungary): systematic background, functional anatomy and palaeoecology. Coloq Palaeontol, Ext, 1: 35-45 |
[11] | Bernor R L, Kaiser T M, Nelson S V et al., 2011. Systematics and paleobiology of Hippotherium malpassii n. sp. (Equidae, Mammalia) from the latest Miocene of Baccinello V3 (Tuscany, Italy). Boll Soc Paleontol Ital, 50(3): 175-208 |
[12] | Bernor R L, G?hlich U B, Harzhauser M et al., 2017. The Pannonian C hipparions from the Vienna Basin. Palaegeogr Palaeoclim Palaeoecol, 476: 28-41 |
[13] | Bernor R L, Wang S Q, Liu Y et al., 2018. Shanxihippus dermatorhinus (new gen.) with comparisons to Old World hipparions with specialized nasal apparati. Riv Ital Paleontol Stratigr, 124: 361-386 |
[14] | Bernor R L, Cirilli O, Mittmann H W, 2022. H?wenegg Hippotherium primigenium: geological context, cranial and postcranial morphology, palaeoecological and biogeographic importance. Hist Biol, 35(8): 1376-1390 |
[15] | Deng T, 2012. A skull of Hipparion (Proboscidipparion) sinense (Perissodactyla, Equidae) from Longdan, Dongxiang of northwestern China - addition to the Early Pleistocene Longdan mammalian fauna (3). Vert PalAsiat, 50: 74-84 |
[16] | Deng T, Wang W M, Yue L P et al., 2004. New advances in the establishment of the Neogene Baode Stage. J Stratigr, 28(1): 41-47 |
[17] | Deng T, Qiu Z X, Wang B Y et al., 2013. Chapter 9: Late Cenozoic Biostratigraphy of the Linxia Basin, Northwestern China. In: Wang X M, Flynn L J, Fortelius M eds. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York: Columbia University Press. 243-273 |
[18] | Deng T, Wang H J, Wang X M et al., 2016. The Late Miocene Hipparion (Equidae, Perissodactyla) fossils from Baogeda Ula, Inner Mongolia, China. Hist Biol, 28: 53-68 |
[19] | Eisenmann V, Alberdi M T, de Giuli C et al., 1988. Methodology. In: Woodburne M, Sondaar P eds. Studying Fossil Horses. Leiden: E J Brill. 1-71 |
[20] | Fang X, Wang J, Zhang W et al., 2016. Tectonosedimentary evolution model of an intracontinental flexural (foreland) basin for paleoclimatic research. Glob Planet Change, 145: 78-97 |
[21] | Forstén A M, 1968. Revision of the Palearctic Hipparion. Acta Zool Fenn, 119: 1-134 |
[22] | Forstén A M, 1985. Chinese Turolian Hipparion in the Lagrelius Collection. In: Lucas S G, Mateer N J eds. Studies in Chinese Fossil Vertebrates. Bull Geol Inst Univ Upps, New Ser, 11: 113-124 |
[23] | Fortelius M, Eronen J T, Kaya F et al., 2014. Evolution of Neogene mammals in Eurasia: environmental forcing and biotic interactions. Ann Rev Earth Planet Sci, 42: 579-604 |
[24] | Garcés M, Cabrera L, Agustí J et al., 1997. Old World first appearance Datum of “Hipparion” horses: Late Miocene large-mammal dispersal and global events. Geology, 25(1): 19-22 |
[25] | Goloboff P A, Farris J S, Nixon K C, 2008. TNT, a free program for phylogenetic analysis. Cladistics, 24: 774-786 |
[26] | Gromova V, 1952. Le genre Hipparion. Bur Rech Geol Min, 12: 1-288 |
[27] | Kaakinen A, Passey B H, Zhang Z Q et al., 2013. Chapter 7: Stratigraphy and paleoecology of the classical dragon bone localities of Baode County, Shanxi Province. In: Wang X M, Flynn L J, Fortelius M eds. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York: Columbia University Press. 203-217 |
[28] | Kaiser T M, Bernor R L, Scott R S et al., 2003. New interpretations of the systematics and palaeoecology of the Dorn-Dürkheim 1 hipparions (Late Miocene, Turolian Age [MN11]), Rheinhessen, Germany. Senckenbergiana lethaea, 83: 103-133 |
[29] | Li Y F, Sun B Y, Deng T et al., 2023. Late Miocene Hipparion (Equidae, Perissodactyla) fossils from Fugu, northern Shaanxi, China, and their stratigraphic significance. J Mamm Evol, 30: 657-671 |
[30] | Liu Y, 2013. Late Miocene hipparionine fossils from Lantian, Shaanxi Province and phylogenetic analysis on Chinese Hipparionines. Ph. D thesis. Beijng: University of Chinese Academy of Sciences. 1-130 |
[31] | MacFadden B J, Woodburne M O, 1982. Systematics of the Neogene Siwalik hipparions (Mammalia, Equidae) based on cranial and dental morphology. J Vert Paleont, 2: 185-218 |
[32] | Pang L B, 2015. Hipparion (Equidae, Perissodactyla) fossils from the Shilei Locality of the Linxia Basin, Gansu Provinve. Quat Sci, 35(3): 502-512 |
[33] | Qiu Z X, Huang W L, Guo Z H, 1987. The Chinese hipparion fossils. Palaeont Sin, New Ser C, 25: 1-250 |
[34] | Rook L, Bernor R L, 2013. Hippotherium malpassii (Equidae, Mammalia) from the latest Miocene (late Messinian; MN13) of Monticino gypsum quarry (Brisighella, Emilia-Romagna, Italy). Boll Soc Paleontol Ital, 52(2): 95-102 |
[35] | Rook L, Oms O, Benvenuti M et al., 2011. Magnetostratigraphy of the Late Miocene Baccinello-Cinigiano Basin (Tuscany, Italy) and the age of Oreopithecus bambolii faunal assemblages. Palaegeogr Palaeoclimat Palaeoecol, 305: 286-294 |
[36] | Sefve I, 1927. Die Hipparionen Nord-Chinas. Palaeont Sin, New Ser C, 4(2): 1-93 |
[37] | Sen S, 1989. Hipparion Datum and its chronologic evidence in the Mediterranean area. In: Lindsay E H, Fahlbusch V, Mein P eds. European Neogene Mammal Chronology. New York: Plenum Press. 495-506 |
[38] | Sisson S, 1953. The Anatomy of the Domestic Animals. Philadelphia: W. B. Saunders Company. 1-972 |
[39] | Sun B Y, 2024. Fossil Equidae in the Linxia Basin with biostratigraphic and paleozoogeographic significance. Acta Geol Sin-Engl, 98(1): 1-9 |
[40] | Sun B Y, Deng T, 2019. The Equus Datum and the Early Radiation of Equus in China. Front Ecol Evol, 7: 429 |
[41] | Sun B Y, Zhang X X, Liu Y et al., 2018. Sivalhippus ptychodus and Sivalhippus platyodus (Perissodactyla, Mammalia) from the late Miocene of China. Riv Ital Paleontol Stratigr, 124: 1-22 |
[42] | Sun B Y, Liu Y, Chen S et al., 2022. Hippotherium Datum implies Miocene palaeoecological pattern. Sci Rep, 12: 3605 |
[43] | Tong H W, 2012. Evolution of the non-Coelodonta dicerorhine lineage in China. C R Palevol, 11: 555-562 |
[44] | Wang M Y, Wang H Y, Zhu Z X et al., 2021. Late Miocene-Pliocene Asian summer monsoon variability linked to both tropical Pacific temperature and Walker Circulation. Earth Planet Sci Lett, 561: 116823 |
[45] | Woodburne M O, 2005. A new occurrence of Cormohipparion, with implications for the Old World Hippotherium Datum. J Vert Paleont, 25(1): 256-257 |
[46] | Woodburne M O, 2007. Phyletic diversification of the Cormohipparion occidentale complex (Mammalia, Perissodactyla, Equidae), Late Miocene, North America, and the origin of the Old World Hippotherium Datum. Bull Am Mus Nat Hist, 306: 1-180 |
[47] | Woodburne M O, Bernor R L, 1980. On superspecific groups of some Old World hipparionine horses. J Paleontol, 54(6): 1319-1348 |
[48] | Xue X X, Zhang Y X, Yue L P, 2006. Paleoenvironments indicated by the fossil mammalian assemblages from red clay-Loess sequence in the Chinese Loess Plateau since 8.0 Ma B.P. Sci China Ser D, 49(5): 518-530 |
[49] | Zdansky O, 1923. Fundorte der Hipparion-Fauna um Pao-Te-Hsien in NW-Shansi. Bull Geol Surv China, 5: 69-81 |
[50] | Zhang Y X, Xue X X, Yue L P, 1995. Age and division of Neogene “Red Bed” of Laogaochuan, Fugu County, Shaanxi. J Stratigr, 19: 214-219 |
[51] | Zhang Z Q, Kaakinen A, Liu L P et al., 2013. Chapter 6: Mammalian biochronology of the Late Miocene Bahe Formation. In: Wang X M, Flynn L J, Fortelius M eds. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York: Columbia University Press. 187-202 |
[52] | Zhegallo V I, 1971. Hipparions from the Neogene deposits of Western Mongolia and Tuva. Sovm Sovet-Mongol Nauch-Issled Geol Eksped Tr, 3: 98-119 |
[53] | Zhu Y M, Zhou L P, Mo D W et al., 2008. A new magnetostratigraphic framework for late Neogene Hipparion Red Clay in the eastern Loess Plateau of China. Palaegeogr Palaeoclimat Palaeoecol, 268: 47-57 |
[54] | Zouhri S, Bensalmia A, 2005. Révision systématique des Hipparion sensu lato (Perissodactyla, Equidae) de l’Ancien Monde. Estud Geol, 61: 61-99 |
/
〈 |
|
〉 |