歐亞大陸晚中新世“ 麝牛類” 牛科動物

The Late Miocene ‘ovibovin’ bovids in Eurasia

Abstract During the Late Miocene， numerous medium to large-sized herbivores， that resemblethe living Ovibos in skeletal morphology， dispersed throughout the Holarctic realm and comprisedeleven genera： Lantiantragus， Shaanxispira， Tsaidamotherium， Criotherium， Mesembriacerus，Hezhengia， Urmiatherium， Parurmiatherium， Plesiaddax， Sinotragus， and Prosinotragus.These genera are primarily found in eastern Asia， in which nine genera are found in northernChina， as well as in countries in eastern Europe and western Asia. They are distinguished byunique characteristics， including a perpendicular braincase， specialized horn cores， and a robustbasioccipital. Previous studies have often classified these ‘ovibovin’ bovids as part of theconventional subfamily/tribe Ovibovinae/Ovibovini， along with extant Ovibos. Nevertheless， anincreasing number of studies do not support the monophyly of the subfamily/tribe Ovibovinae/Ovibovini， nor is a close relationship likely between these Late Miocene ‘ovibovin’ bovids andextant Ovibos. Among the eleven genera of ‘ovibovin’ bovids， Plesiaddax， Hezhengia， andUrmiatherium are often considered to have a very close relationship and conventionally formthe tribe Urmiatheriini. However， previous phylogenetic analyses do not support the monophylyof Urmiatheriini. This paper presents a summary of the transmutation of the terms Ovibovidae/Ovibovinae/Ovibovini/Ovibovina， the temporal and spatial distribution of the Late Miocene‘ovibovin’ bovids in Eurasia， the principal characteristics of these taxa， and the previousphylogenetic analyses.

Key words Eurasia， Late Miocene， ‘ovibovin’ bovids， Bovidae， temporal and spatial distribution

Citation Ilyas M， Li Q， Shi Q Q et al.， 2024. The Late Miocene ‘ovibovin’ bovids in Eurasia.Vertebrata PalAsiatica， 62（4）： 262–290

1 Introduction

Bovids are ruminant artiodactyls that were the dominant mammalian herbivores in OldWorld terrestrial communities with diverse extant and extinct species in Africa， Eurasia， andNorth America （Simpson， 1945）. The extant bovids include approximately 45 genera and 140 species （Vrba， 1985; Kingdon， 1989; Grubb， 1993; Bibi et al.， 2009） in two subfamiliesand approximately 13 tribes （Bibi and Tyler， 2022）. Of all the living bovids， the NorthAmerican muskox （Ovibos moschatus） is a particularly unique animal due to its distinctivecranial features and high adaptability to the harsh environment of the Arctic tundra （Lent，1988）. Traditionally， the muskox was believed to be closely related to the takin （Budorcas）from southern China （Simpson， 1945）. Gray （1872） proposed the family Ovibovidae forOvibos and Budorcas. In the same year， Gill （1872） proposed the subfamily Ovibovinae，which included both living Ovibos and extinct Bootherium （Table 1）. Lydekker （1913）and Bohlin （1935a） also used the subfamily Ovibovinae. Sickenberg （1933） establishedthe subfamily Urmiatheriinae， which included two extinct genera： Parurmiatheriumand Urmiatherium. Bohlin （1935a） subsequently merged the Urmiatheriinae into theOvibovinae. Bohlin （1935a） even placed four additional extinct genera， Parurmiatherium，Plesiaddax， Pseudobos， and Urmiatherium， in the subfamily Ovibovinae. Bohlin （1937）reevaluated his earlier classification and agreed that the subfamily Urmiatheriinae couldbe legitimate due to the unique horn core growth processes between Urmiatherium andmodern Ovibos. Simpson （1945） concurred with the close relationship between Ovibos andBudorcas and grouped them together with nine extinct genera， namely Boopsis， Bootherium，Criotherium， Euceratherium， Parurmiatherium， Plesiaddax， Symbos， Tsaidamotherium，and Urmiatherium， in the tribe Ovibovini under the subfamily Caprinae. These generaspan a temporal range from the Late Miocene to the Pleistocene. Simpson’s classificationhas been widely accepted for a long time. McKenna and Bell （1997） adopted Simpson’s（1945） framework but expanded the Ovibovini tribe to include additional extinct generasuch as Damalavus， Lyrocerus， Makapania， Megalovis， Palaeoreas， Praeovibos， andSoergelia. Grubb （2001） classified the two genera， Ovibos and Budorcas， within the tribeOvibovini， which is part of the subfamily Caprinae. Chen and Zhang （2004） resurrectedthe subfamily Urmiatheriinae， proposed by Sickenberg （1933）， for some Late Miocenebovids in northern China. Their subfamily includes Turcocerus， Urmiatherium， Plesiaddax，Hezhengia， and Shaanxispira. Chen and Zhang （2009） divided the subfamily Urmiatheriinaeinto two tribes： Turcocerini and Urmiatheriini. The latter tribe includes Lantiantragus，Urmiatherium， Plesiaddax， Hezhengia， Tsaidamotherium， Shaanxispira， Sinotragus，and Prosinotragus. Nevertheless， a recent phylogenetic analysis did not corroborate themonophyly of the tribe Urmiatheriini （Shi and Deng， 2021）. In the Handbook of HoofedMammals， Groves and Leslie （2011） classified Ovibos and Budorcas in the tribe Capriniunder the subfamily Antilopinae， along with several other living genera， includingAmmotragus， Arabitragus， Capra， Carpricornis， Hemitragus， Naemorhedus， Nilgiritragus，Oreamnos， Ovis， Pantholops， Pseudois， and Rupicapra. However， molecular evidence，particularly mitochondrial DNA analysis， indicates that Ovibos is more closely related to Capricornis and Naemorhedus than to Budorcas （Lalueza-Fox et al.， 2005; Ropiquet andHassanin， 2005; Bibi et al.， 2012; Hassanin et al.， 2012）. Hassanin et al. （2012） proposed thesubtribe Ovibovina， which belongs to the tribe Caprini and includes only the living Ovibos，Capricornis， and Naemorhedus. For the fossil bovids that are similar to Ovibos， Lazaridis etal. （2017） used a general term， “ovibovine-like taxa”. In this text， it is proposed to use theterm ‘ovibovin’ bovids to refer all fossil taxa that are not closely related to Ovibos， followingKostopoulos et al. （2023）， as the conventional Ovibovinae/Ovibovini is no longer applicableto the living Ovibos and the “urmiatherines” are not monophyly. It is also worth noting thatthe positions of these ‘ovibovin’ fossil bovids remain unsolved.

Institutional abbreviations AMNH， American Museum of Natural History， New York，USA; BGR， Bundesanstalt für Geowissenschaften und Rohstoffe， Hannover， Germany; BMNH，Natural History Museum， London， UK; BSPM， Bayerische Staatssammlung für Pal?ontologieund Geologie， Pal?ontologisches Museum， Munich， Germany; ?O， ?ank?r? Museum， Turkey;DOE， Department of the Environment， Maragheh， Iran; HMV， Hezheng PaleozoologicalMuseum， Gansu， China; HUW， Howard University， Laboratory of Evolutionary Biology，Washington， DC， USA; IGPM， Institute für Geologie und Pal?ontologie， Universit?t Münster，Germany; IVPP， Institute of Vertebrate Paleontology and Paleoanthropology， ChineseAcademy of Sciences， Beijing， China; LGPUT， Laboratory of Geology and Palaeontology，Aristotle University of Thessaloniki， Thessaloniki， Greece; MMTT， fossils from Maraghehlocalities of LRE deposited either in Tehran or in Washington， DC， USA; MNHN， Muséumnational d’Histoire Naturelle， Paris， France; MTA， Maden Tetkik ve Arama Museum， Ankara，Turkey; MY?E PV， Natural History Museum of Ege University， Izmir， Turkey; NHML，Natural History Museum， London， UK; NHMUK， Natural History Museum， London， UK;NHMW， Naturhistorisches Museum， Vienna， Austria; NLB， Nieders?chsisches Landesamt fürBodenforschung， Hannover， Germany; NMNH， National Museum of Natural History， Sofia，Bulgaria; PGMA， Palaeontology and Geology Museum， University of Athens， Athens， Greece;PMU， Museum of Evolution in Uppsala University， Uppsala， Sweden.

2 Geographic and stratigraphic distribution of Late Miocene ‘ovibovin’ bovids in Eurasia

The Late Miocene ‘ovibovin’ bovids include Criotherium， Hezhengia， Lantiantragus，Mesembriacerus， Parurmiatherium， Plesiaddax， Shaanxispira， Sinotragus， Prosinotragus，Tsaidamotherium， and Urmiatherium. They are concentrated in eastern Europe and northernChina （Fig. 1）. Plesiaddax， Sinotragus， and Urmiatherium are widely distributed in bothnorthern China and eastern Europe （Shi and Deng， 2021）. In contrast， Criotherium andMesembriacerus are restricted to eastern Europe. Criotherium is distributed to the LateMiocene of Greece， Bulgaria， and Turkey （Forsyth-Major， 1891; Solounias， 1981; Bouvrain，1994; Geraads and Spassov， 2008; Kostopoulos et al.， 2023）， while Mesembriacerus is onlyfound in Greece （Bouvrain and Bonis， 1984; Spassov et al.， 2018）. Parurmiatherium has beendiscovered in Greece， Iraq， and southwestern Turkey （Sickenberg， 1932， 1933; Bouvrain etal.， 1995; Kostopoulos and Karakütük， 2013）. Nine genera occur in northern China; five ofthem are endemic， including Hezhengia， Lantiantragus， Shaanxispira， Prosinotragus， andTsaidamotherium. Hezhengia is primarily discovered in the poorly defined “Liushu Formation”（see Qiu et al.， 2023） in the Linxia Basin located at the northeastern edge of the TibetanPlateau （Qiu et al.， 2000; Deng et al.， 2013; Shi and Deng， 2021）. Lantiantragus is limited tothe Bahe Formation in the Lantian Basin， Shaanxi Province （Chen and Zhang， 2004， 2009）.Shaanxispira is found in the Bahe and Liushu formations in northwestern China （Liu et al.，1978; Zhang， 2003; Shi et al.， 2014）. Tsaidamotherium was discovered in the ShangyoushashanFormation in the Qaidam Basin， Qinghai， and the Liushu Formation in the Linxia Basin，Gansu， northwestern China （Bohlin， 1935b; Shi， 2014; Wang and Ye， 2023）. Prosinotragus isonly discovered in Qingyang， Gansu Province （Bohlin， 1935a）.

The majority of the ‘ovibovin’ bovids in Eurasia from the Late Miocene are from theearly Late Miocene， except for Urmiatherium intermedium， U. rugosifrons， Sinotraguswimani， and Prosinotragus tenuicornis， which appear in the late Late Miocene （Fig. 2）.Tsaidamotherium is the earliest member among all the Late Miocene ‘ovibovin’ bovidsabovementioned， appearing at the boundary between the Middle and the Late Miocene.This genus is endemic to northwestern China and comprises two species， the number ofspecimens of which is limited. T. hedini was primarily discovered in the ShangyoushashanFormation in the Quanshuiliang area of the eastern Qaidam Basin （Wang et al.， 2011），while T. brevirostrum was found in the “Liushu Formation” in the Linxia Basin （Shi，2014）. Both species of Tsaidamotherium belong to the Bahean Age of the Chinese LandMammal Age. Lantiantragus is a monotypic genus that includes only the type species L.longirostralis， which was discovered in loc. 12 of the lower part of the Bahe Formation inthe Lantian Basin， Shaanxi. This species is in the Bahean Age， with a magnetic dating ofapproximately 9.95 Ma （Chen and Zhang， 2004; Zhang et al.， 2013）. Hezhengia consistsof two species， H. bohlini and H. minor. The holotype of H. bohlini was discovered in thegreyish-yellow silts in the Linxia Basin. Qiu et al. （2000） considered that the fossils fromthis layer can be correlated with the Hipparion faunas in the Hezheng and Baode area，which are believed to date to the late Late Miocene. H. minor was originally referred to thegenus Plesiaddax by Bohlin （1935a） with some reservation and then was transferred to thegenus Hezhengia by Shi and Deng （2021）. This species was discovered in Fugu， Shanxi，and its age is approximately 8.0 Ma， according to palaeomagnetic data （Bohlin， 1935a;Xue et al.， 1995）. Urmiatherium has a wide distribution throughout Eurasia with threeknown species. The type species， U. polaki， was discovered in the Maragheh Formationof Karaj Abad， Maragheh， Iran， with an estimated age of ～8.7–7.5 Ma （Rodler， 1889; deMecquenem， 1925; Kostopoulos and Bernor， 2011）. U. intermedium was discovered in thered clays of Baode， Shanxi， and Qingyang， Gansu， as well as in the “Liushu Formation”in the Linxia Basin， Gansu， northwestern China. Its age is within the Baodean Age （lateLate Miocene）. U. kassandriensis is a recently established species based on the materialfrom the Antonios Formation in Fourka， Chalkidili， Greece， and its age is about 9.0–8.0Ma （Lazaridis et al.， 2017）. Parurmiatherium is a monotypic genus that includes onlythe type species P. rugosifrons. The species was discovered in the Mytilinii Formation ofSamos， eastern Greece， the Bakhtiari Formation of Injana， Iraq， and the Salihpa?alar and?erefk?y-2 localities in the Yata?an Formation of southwestern Turkey （Sickenberg， 1932，1933， Bouvrain et al.， 1995， Kostopoulos and Karakütük， 2013）. Plesiaddax is distributedfrom northern China to Turkey， with three species. Its type species is P. depereti， whichwas discovered from the Baode Formation in Shanxi， China. The latter two species，P. inundates and P. simplex， have recently been described as Hezhengia with some reservation （Kostopoulos et al.， 2023）. Criotherium has two species. The type species C.argalioides is known from the Mytilinii Formation in Samos， Greece， the Sinap Formationof Kemiklitepe-D， and the Tüg?lu Formation of ?orakyerler in Turkey （Forsyth-Major，1891; Solounias， 1981; Bouvrain， 1994; Geraads and Spassov， 2008; Kostopoulos et al.，2023）. C. nikolovi was discovered in the Kalimantsi Formation in Kalimantsi-1， Bulgaria.Shaanxispira is endemic to northern China， where it is represented by three species. Thefirst two species， S. chowi and S. baheensis， were discovered from the Bahe Formation inthe Lantian Basin， while S. linxiaensis was discovered from the “Liushu Formation” in theLinxia Basin （Liu et al.， 1978; Zhang， 2003; Shi et al. 2014）. Mesembriacerus melentisi isthe sole representative of its genus found in the Antonios Formation of the Axios Valley inGreece， with an age of approximately 9.3 Ma （Bouvrain and Bonis， 1984; Spassov et al.，2018）. Lantiantragus is based on an anterior part of the skull from the Bahe Formation inthe Lantian Basin， Shaanxi， which comprises only the type species L. orientalis. The speciesis restricted to the Bahe Formation of the Lantian Basin （Chen and Zhang， 2004， 2009）.Sinotragus was established based on cranial materials with two species. The type species，S. wimani， was discovered in the Baode Formation of Baode County， Shanxi， China. Thesecond species S. occidentalis was found from an unidentified locality within the Yata?anFormation in Turkey （Geraads et al.， 2002）. Prosinotragus was erected with its type species P.tenuicornis in the Liushu Formation of Qingyang area， Gansu with an age of approximately6.5 Ma （Bohlin， 1935a）. Sinotragus is considered as synonymous with Prosinotragus in fewstudies （Chen and Zhang， 2009）.

3 Systematic paleontology

Artiodactyla Owen， 1848

Bovidae Gray， 1821

Urmiatherium Rodler， 1889

Type species Urmiatherium polaki Rodler， 1889.

Distribution China， Iran， Greece.

Included species Urmiatherium intermedium Bohlin， 1935a; U. kassandriensis Lazaridis et al.， 2017.

Ages Urmiatherium polaki， ～8.2–7.4 Ma; U. intermedium， ～7.2–5.3 Ma; U.kassandriensis， 9.7–8.7 Ma （MN 10）; see Table 2.

Remarks Urmiatherium was named by Rodler （1889） based on the material fromMaragheh， Iran， and currently includes three species： U. polaki， U. intermedium， and U.kassandriensis. Urmiatherium is a Late Miocene bovid characterized by closely inserted andcaudally oriented horn cores （Fig. 3A）， a perpendicular cranial roof， a thickened basicranium，and hypsodont cheek teeth （Rodler， 1889; de Mecquenem， 1925; Bohlin， 1935a， Sickenberg，1932， 1933; Kostopoulos， 2009; Kostopoulos and Bernor， 2011; Jafarzadeh et al.， 2012; Kayaet al.， 2012; Kostopoulos and Karakütük， 2013; Shi et al.， 2016; Lazaridis et al.， 2017）. Amongthe Late Miocene ‘ovibovin’ bovids， Urmiatherium has the highest tooth crown. The earliestUrmiatherium is U. kassandriensis， whose horn cores are short and separated （Fig. 3E）. Thehorn core of Urmiatherium became more specialized in the two later lineages in both northernChina and western Asia， with the horn bases merging together and extending on the frontal andparietal crania （Fig. 3A， C）.

At the tribal level， the assignment of Urmiatherium remains a topic of debate. Apart fromits conventional classification as the Ovibovinae/Ovibovini （Bohlin， 1935a; Solounias， 1981），Kostopoulos （2009， 2014） referred to it as the Antilopini tribe and the Oiocerina subtribe.However， some other researchers followed Sickenberg’s （1932） original assignment andreferred it to the Urmiatheriini tribe （Chen and Zhang， 2009; Shi et al.， 2016; Shi and Deng，2021）.

Criotherium Forsyth-Major， 1891

Type species Criotherium argalioides Forsyth-Major， 1891.

Included species Criotherium nikolovi Geraads amp; Spassov， 2008.

Distribution Greece， Bulgaria， and Turkey.

Ages Criotherium argalioides， ～8.7–7.5 Ma; C. nikolovi， ～8.9–7.5 Ma; see Table 2.

Remarks Criotherium is the only ‘ovibovin’ bovid with heteronymously twisted horncores. As such， Criotherium is unlikely to be closely related to the other genera of ‘ovibovin’bovids， assuming the direction of twists evolved separately. Its cranium morphologies， suchas a short and broad braincase， low and broad occipital， strong posterior tuberosities， andsupplementary articulations medial to the paroccipital processes， are similar to other ‘ovibovin’bovids. Criotherium is common in Samos （Greece）， but it is quite rare in Kemiklitepe-D（Western Turkey）， Kalimantsi-1 （Bulgaria）， and ?orakyerler （Turkey） （Solounias， 1981;Bouvrain， 1994; Geraads and Spassov， 2008; Kostopoulos et al.， 2023）. C. nikolovi differsfrom the type species， C. argalioides， in its smaller size， less upright horn cores （Fig. 4D， H），and less lengthened and broader muzzle （Fig. 4A， E）， which are probably primitive （Geraads and Spassov， 2008）. According to Gentry （1971）， Criotherium was a large antilopine closelyrelated to Palaeoreas， evolving in parallel with Urmiatherium and sharing many similar traits.Morphologically， Criotherium is the most similar to Garkin’s ？Plesiaddax inundates， althoughit cannot be included in Plesiaddax due to the weak homonymous torsion of ？P. inundates（Bosscha-Erdbrink， 1978; Geraads and Spassov， 2008; Kostopoulos et al.， 2023）.

Plesiaddax Schlosser， 1903

（Fig. 5）

Type species Plesiaddax depereti Schlosser， 1903.

Included species Plesiaddax inundates Bosscha-Erdbrink， 1978; P. simplex K?hler，1987.

Distribution Western Turkey， northern China.

Ages Plesiaddax depereti， ～7.2–5.3 Ma; P. inundates， ～8.7–7.5 Ma （MN 11） and P.simplex， ～8.7–7.5 Ma （MN 11）; see Table 2.

Remarks Plesiaddax was established by Schlosser （1903） based on the materialfrom Hequ， Shanxi， China. It is an ‘ovibovin’ bovid with extremely short and laterallyextended horn cores， similar to Hezhengia but much shorter. Solounias （1981） suggested thatPlesiaddax is synonymous with Parurmiatherium， but Shi et al. （2016） treated them as twoindependent genera. Plesiaddax shares similarities with Hezhengia in terms of skull and horncore morphology. They may have a closer relationship than other Late Miocene ‘ovibovin’bovids， such as Urmiatherium， Tsaidamotherium， and Criotherium， as pointed out by Qiuet al. （2000）. Shi and Deng （2021） also suggested a close relationship among Plesiaddax，Hezhengia， and Urmiatherium. The type species of Plesiaddax is P. depereti， which ischaracterized by several beautifully preserved skulls and mandibles. The two includedspecies， namely P. simplex and P. inundates， have inadequate material. In contrast to P.depereti， the horn bases in P. simplex are not enlarged anteriorly by buttresses. Additionally，P. inundates is more similar to Criotherium or Hezhengia than P. depereti （Bouvrain et al.，1995; Geraads and Spassov， 2008; Kostopoulos et al.， 2023）. Kostopoulos et al. （2023） havequestioned the generic attribution of these two species and suggested that they should betransferred to the genus Hezhengia. （Fig. 5）

Parurmiatherium Sickenberg， 1932（Fig. 6）

Type species Parurmiatherium rugosifrons Sickenberg， 1932.

Distribution Greece， southwestern Turkey， and Iraq.

Age ～7.4–6.7 Ma; see Table 2.

Remarks Parurmiatherium was named by Sickenberg （1932） based on material fromSamos， E Greece. It includes only the species P. rugosifrons. This genus is characterizedby its diminutive size and the presence of a horn core that is very short， robust， mediallyunfused， strongly homonymously twisted， and grooved. It is also distinguished by the presenceof low buttresses that extend anteriorly from the frontals （Sickenberg， 1932; Gentry et al.，1999; Kostopoulos， 2009; 2014）. Gentry et al. （1999） considered this genus a synonym ofUrmiatherium. This classification was followed by Kostopoulos （2009， 2014） and Kostopouloset al. （2023）. Solounias （1981） synonymized Plesiaddax with Parurmiatherium， whereas Shiet al. （2016） treated Parurmiatherium as an independent genus. These debates reflect thatParurmiatherium exhibits mixed characteristics of both Urmiatherium and Plesiaddax. Forinstance， the backwards extending horn cores and the extremely strong posterior tuberosity aresimilar to Urmiatherium. In contrast， the wide opisthocranium and the separate horn bases aresimilar to Plesiaddax （Fig. 6A， B）.

Sinotragus Bohlin， 1935a（Fig. 7）

Type species Sinotragus wimani Bohlin， 1935a.

Included species S. occidentalis Geraads et al.， 2002.

Distribution China and Turkey.

Ages Sinotragus wimani， ～5.7 Ma; S. occidentalis 7.4–6.8 Ma （MN 12）; see Table 2.

Remarks Sinotragus and its type species S. wimani were named by Bohlin （1935a）based on cranial materials from Baode， Shanxi. The genus is characterized by prominent horncores and a relatively short skull. The horn cores are long， slightly homonymously twisted，and closely inserted with each other. Geraads et al. （2002） erected a new species of Sinotragus，S. occidentalis， based on the material from the Late Miocene of Mu?la， Turkey. However，Chen and Zhang （2009） noted that S. occidentalis differs from Sinotragus in China in severalcranial features， such as the position of the cranial curve not being located at the frontal boneanterior to the base of the horn core， the long length of the braincase （Fig. 7A， C）， and theuntwisted horn cores. They suggested that S. occidentalis may not belong to Sinotragus. For along time， Sinotragus has been considered a representative of Protoryx in the East Asia. Thisgroup was first included in the subfamily Hippotraginae but was later referred to as the tribeCaprini （Bohlin， 1935a; Gentry， 1971， 2000）. However， previous works have often classifiedSinotragus in the tribe Urmiatheriini due to its homonymously twisted horn cores， shortbraincase， wide and flat occiput， and strongly developed basioccipital tuberosities （Zhang，2003; Chen and Zhang， 2009）.

Prosinotragus Bohlin， 1935a（Fig. 8）

Type species Prosinotragus tenuicornis Bohlin， 1935a.

Distribution China.

Age P. Tenuicornis， ～6.5 Ma; see Table 2.

Remarks Prosinotragus was also named by Bohlin （1935a） based on cranial materialsfrom Qingyang， Gansu. The genus is characterized by its slender， distinctive anterior keels andweakly twisted horn cores. P. tenuicornis was originally the type species of the Prosinotragus，but its cranial morphology is identical to that of Sinotragus wimani. Chen and Zhang （2009）proposed that Prosinotragus is synonymous with Sinotragus because of its cranial morphology.However， P. tenuicornis can be distinguished from S. wimani by its slender， distinctive anteriorkeels and weakly twisted horn cores （Fig. 8A， B）. Further research will elucidate and unveilthe characteristics of this genus and species.

Tsaidamotherium Bohlin， 1935b（Fig. 9）

Type species Tsaidamotherium hedini Bohlin， 1935b.

Included species Tsaidamotherium brevirostrum Shi， 2014.

Distribution Northwestern China.

Ages Tsaidamotherium hedini， ～11.1–10.1 Ma （Tuosu Fauna）; T. brevirostrum， ～11.1–10.5 Ma （Guonigou Fauna; equivalent to MN 9）; see Table 2.

Remarks Bohlin （1935b） named Tsaidamotherium and its type species T. hedinibased on three incomplete skulls from the south limb of the Oulongbuluk Anticline in theQuanshuiliang， eastern Qaidam Basin （See Wang et al.， 2011： locality 447 and 457）. Despitethe lack of evidence for the anterior skull parts （Bohlin， 1935a， b）， T. hedini was placed in thesubfamily Ovibovinae by Bohlin （1935b） due to its cranial resemblance to Plesiaddax， Ovibos，and Urmiatherium. The second species of the genus T. brevirostrum was established by Shi（2014） based on an almost complete skull （HMV 1865） from the Linxia Basin in Gansu. Thisgenus is an endemic bovid found only in northwestern China and is characterized by its uniqueplate-like and asymmetric horn core apparatus （Fig. 9C）. T. hedini and T. brevirostrum aresimilar in size and have similar posterior horn core plate morphology （Fig. 9A， D）. However，the proportion of the real horn cores relative to the skull is smaller in T. brevirostrum than in T.hedini （Fig. 9C， F）. Wang and Ye （2023） reported the upper and lower dentitions of T. hedinifrom the Tuosu Lake area in the eastern Qaidam Basin. These dentitions are comparable tothose of T. brevirostrum， exhibiting an elongated M3， a flat labial wall， and a well-developedmesostyle in the upper molars.

The phylogenetic position of Tsaidamotherium is a topic of debate. According toconventional views， this genus was classified as either Ovibovinae/Ovibovini （Bohlin，1935b; Shi， 2014） or Urmiatheriini （K?hler， 1987）. Chen and Zhang （2009） placed thisgenus in the tribe Urmiatheriini under the subfamily Urmiatherinae， along with other LateMiocene “ovibovines”. Shi and Deng （2021） considered it a distantly related bovid tourmiatherins. The argument presented by Wang et al. （2003a， b） and Wang and Ye （2023） isthat Tsaidamotherium belongs to the stem taxon Discokerycinae and is a giraffoid rather thana bovid， due to differences in the median parietal headgear and bony labyrinth. However， Houet al. （2023） commented that Tsaidamotherium has mesodont teeth， a short premolar row， fullyfused cones in molars， and flat labial walls on the upper molars， which suggests that it belongsto the Bovidae rather than the Giraffidae.

Mesembriacerus Bouvrain， 1975

Type species Mesembriacerus melentisi Bouvrain， 1975.

Distribution Northern Macedonia， Greece.

Age ～9.3 Ma; see Table 2.

Remarks Bouvrain （1975） named Mesembriacerus based on material from Macedonia，Greece. This genus comprises only its type species， M. melentisi， which is characterized bylong， straight， and strongly inclined horn cores. The horn cores of Mesembriacerus are insertedfar behind the orbit and an accessory articular surface between the paroccipital process andthe occipital condyle. These characteristics are similar to those of Ovibos， which classifies Mesembriacerus as a member of Ovibovini according to conventional views （Bouvrain andBonis， 1984）. Additionally， the relatively small body size， slender horn cores， and primitivebraincase suggest a primitive evolutionary stage for ‘ovibovin’ bovids. Mesembriacerusis considered the most primitive member of the lineage that gave rise to Plesiaddax，Tsaidamotherium， and Urmiatherium （Bouvrain and Bonis， 1984）. It possibly originatedin Asia and arrived in the southern Balkans during the late Vallesian period （Lazaridis etal.， 2017）. However， its small parieto-occipital angle and postero-ventrally faced occiput（Fig. 10C） are distinct from all other Late Miocene ‘ovibovin’ bovids， implying a differentevolutionary lineage.

Shaanxispira Liu et al.， 1978

Type species Shaanxispira chowi Liu et al.， 1978.

Included species Shaanxispira baheensis Zhang， 2003; S. linxiaensis Shi et al.， 2014.

Distribution Northern China.

Ages Shaanxispira chowi， 8.21 Ma; S. baheensis， 8.07 Ma; S. linxiaensis， ～9.5–8 Ma;see Table 2.

Remarks Shaanxispira is an endemic genus found only in northern China. It ischaracterized by long， straight， and strongly homonymously twisted horn cores. Althoughthe horn core of Shaanxispira is not similar to that of Ovibos， its cranium is short and strong，making this genus also a phylogenetically close relative of urmiatherins （Chen and Zhang，2009）. All three known species of Shaanxispira are contemporaneous and of similar size. Themost notable difference between them lies in the morphology of their horn cores （Shi et al.，2014）. The type species， S. chowi， has an extra postero-lateral keel on the horn core （Fig. 11E，F(xiàn)）， distinguishing it from the other two species. S. baheensis （Fig. 11C， D）， on the other hand，has more inclined horn cores and weaker keels than S. linxiaensis （Fig. 11A， B） （Zhang， 2003;Shi et al.， 2014）.

Hezhengia Qiu et al.， 2000

Type species Hezhengia bohlini Qiu et al.， 2000.

Included species Hezhengia minor （Bohlin， 1935a）.

Distribution Northwestern China.

Ages Hezhengia bohlini， ～9.5–8 Ma （Dashengou Fauna）; H. minor， ～7.8 Ma （Lamagou Fauna）; see Table 2.

Remarks Hezhengia is a medium-sized ‘ovibovin’ bovid that is endemic to northernChina. It is characterized by short and robust horn cores that insert well behind the orbit andextend mainly laterally （Fig. 12A） （Qiu et al.， 2000; Shi and Deng， 2021）. Qiu et al. （2000）established the genus Hezhengia and its type species， H. bohlini， based on hundreds of wellpreservedskulls from the Linxia Basin. Shi and Deng （2021） acknowledged that the two upperjaws from Fugu County， Shaanxi， originally identified as Plesiaddax minor， share similartooth morphology， particularly that of the P3， with H. bohlini. They reclassified P. minor as H.minor. H. minor coexisted with Miotragocerus gregarius， Gazella gaudryi， Dorcadoryx sp.，and Urmiatherium sp.， forming the Lamagou Fauna with an estimated age of ～7.8 Ma （Xue etal.， 2006）. H. bohlini is part of the Dashengou Fauna， estimated from an earlier age of ～9.5–9.0 Ma （Deng et al.， 2019）. However， Shi （2023） correlated the Dashengou Fauna with theLamagou Fauna due to the high similarity of the bovid taxa， suggesting that these two faunasmay be contemporaneous.

Lantiantragus Chen amp; Zhang， 2004

Type species Lantiantragus longirostralis Chen amp; Zhang， 2004.

Distribution Northern China.

Age 9.95 Ma， the Bahean Age; see Table 2.

Remarks Lantiantragus was named by Chen and Zhang （2004） based on a skullfrom the Bahe Formation in the Lantian Basin， Shaanxi. It includes only its type species，L. longirostralis. Lantiantragus is the only genus established on the material without horncores among all the ‘ovibovin’ bovids from northern China. This large bovid genus hasan extremely long muzzle （Chen and Zhang， 2004）. Chen and Zhang （2009） classifiedLantiantragus in the tribe Urmiatheriini and suggested a close relationship amongLantiantragus， Urmiatherium， and Plesiaddax. Our observation indicates that the dentitionof Lantiantragus is similar to that of Shaanxispira， suggesting a close relationship betweenthese two genera. However， Lantiantragus differs from Ovibos， which has an extremelyshort and high muzzle.

4 Classification and phylogeny of the Late Miocene ‘ovibovin’ bovids

The high-ranking classification of Late Miocene ‘ovibovin’ bovids in Eurasia remainsunclear. Bohlin （1935a） classified the ‘ovibovin’ bovids into three groups based on horncore morphology： 1） the Ovibos-group， which includes Plesiaddax and Ovibos， whosehorn cores extend laterally and horn bases enlarge forwards， backwards， and mediallydirected， 2） the Urmiatherium-group， which includes Urmiatherium， Parurmiatheriumand Criotherium， whose horn cores are medially and posteriorly inserted， the cranial roofis raised， and the horn bases are enlarged mainly forwards， and 3） the Tsaidamotheriumgroup，which includes only the Tsaidamotherium， whose real horn cores are inserted abovethe orbit primitively， and the horn base extends backward and occupies the whole cranialroof. Chen and Zhang （2004， 2009） grouped all the Late Miocene ‘ovibovin’ bovids inChina， including Lantiantragus， Urmiatherium， Plesiaddax， Hezhengia， Tsaidamotherium，Shaanxispira， Sinotragus， and Prosinotragus， into the tribe Urmiatheriini， because of thelarge body size， short braincase， long face， specialized horn cores， and short premolartooth row. Recently， based on the phylogenetic analysis， Shi and Deng （2021） suggestednarrowing down the Urmiatheriini tribe and considering it paraphyletic. The Late Miocene‘ovibovin’ bovids in China （see Shi and Deng， 2021） have been classified into threegroups： the Hezhengia-Plesiaddax-Urmiatherium group， the Shaanxispira-Lantiantragusgroup， and the more distant clade of Tsaidamotherium group. Plesiaddax and Hezhengiashare similarities in the postero-laterally inserted horn cores. Urmiatherium are alsosimilar in having short horn cores， short braincase， robust occipital condyle， and accessoryarticular surface medial to the paroccipital process （de Mecquenem， 1925; Bohlin，1935a; Qiu et al.， 2000; Jafarzadeh et al.， 2012; Shi and Deng， 2021）. Lantiantragusand Shaanxispira have similar dentition and are likely to have a close relationship. InLantiantragus， the basal pillars are particularly pronounced in the lower molars， whichserves to distinguish it from the Hezhengia-Plesiaddax-Urmiatherium group （Shi andDeng， 2021）. The homonymously twisted horn core of Shaanxispira is comparable tothat of Oioceros， but the link between Shaanxispira and Oioceros cannot be determined（Shi and Deng， 2021）. The Tsaidamotherium group consists of only one genus，Tsaidamotherium. This genus is the smallest and most distinctive among all knownLate Miocene ‘ovibovin’ bovids in Eurasia （Shi， 2014）. A study on the second speciesof Tsaidamotherium from the Linxia Basin， T. brevirostrum， revealed significant facialand dental characters， such as a strongly shortened nasal， short and high muzzle， smallpremaxilla， very short premolar row， and elongated M3， suggesting that this genus may bea distantly related bovid clade （Shi and Deng， 2021）.

The phylogenetic relationships of these ‘ovibovin’ bovids are highly debated in differentphylogenetic cladograms using different taxa and characters （Fig. 13） （Bohlin， 1935a， 1937;Bouvrain and Bonis， 1984; K?hler， 1987; Chen and Zhang， 2004， 2009; Lalueza-Fox et al.，2005; Ropiquet and Hassanin， 2005; Geraads and Spassov， 2008; Bibi et al.， 2012; Hassaninet al.， 2009， 2012; Kostopoulos， 2014; Shi and Deng， 2021; Hou et al.， 2023， Wang et al.，2023a， b; Wang and Ye， 2023）. The conventional classifications of Ovibovinae/Ovibovini andUrmiatheriini have failed to form a monophyletic group. Most of the Late Miocene ‘ovibovin’bovids are probably not very closely related to the extant Ovibos， but rather belong to differentfossil bovid clades without extant descendants.

5 Discussion

The phylogenetic relationships of the ‘ovibovin’ bovids are difficult to solve becauseof their highly specialized horn core and skull morphology as well as their relatively simpleand similar tooth structure， which might have evolved convergently. Transitional fossiltaxa are also absent to help to bridge the large morphological gaps. In some monotypicgenera， the horn core is sometimes so distinct that it is difficult to find any close relatives.For example， as in Tsaidamotherium， the classification was highly controversial at thefamily/superfamily level because of its strange plate-like horn apparatus. In genera withmore species， such as Plesiaddax， the main problem is the attribution of the species， whichis partially due to the lack of key characters of the genus. The attribution could be highlydebatable when the materials are poorly preserved， such as in P. simplex. There are alsodebates about synonymy， such as the validity of Parurmiatherium， which is similar to bothUrmiatherium and Plesiaddax in cranium morphology. The swelling frontal above the orbitin Parurmiatherium is similar to Urmiatherium， and the former is considered a synonym ofthe latter in some studies （Gentry et al.， 1999; Kostopoulos， 2009， 2014; Kostopoulos et al.，2023）.

The most important issue regarding these ‘ovibovin’ bovids is the phylogeneticrelationships among the most common Late Miocene ‘ovibovin’ bovids： Urmiatherium，Plesiaddax， Shaanxispira， and Hezhengia. All these genera have plenty of skulls thatprovide enough morphological information for further research. Most importantly， theyare closely related and compose one of China’s main evolutionary lineages of bovids.Three questions about this group remain to be solved. First， is Plesiaddax more closelyrelated to Urmiatherium or Hezhengia？ In Shi and Deng’s （2021） cladogram， the BaodeanUrmiatherium and Plesiaddax are sister groups and are later members of the tribeUrmiatheriini. However， in the traditional view， Plesiaddax is more closely related toHezhengia because of its similar horn core position and direction （Qiu et al.， 2000）. Thesecond question is whether or not Shaanxispira is an early branch of Urmiatheriini or relatedto Oioceros. The long and strongly twisted horn core of Shaanxispira is different from allthe urmiatherins， but it is similar to that of the small bovid Oioceros. However， their skullshave many differences， and more studies are needed to confirm their relationships. Third，what is the relationship between urmiatherins and caprins？ All the urmiatherins becameextinct， and previous studies inferred that their closest extant relatives might be the caprins.However， a recent study proposed that the stem Caprini， Pantholops， might have evolvedfrom an early ancestor more than 11 million years ago （Tseng et al.， 2022）， which means thaturmiatherins and caprins have been separated for a very long time and are probably not veryclosely related.

Acknowledgements We are thankful to Prof. D. Geraads and Prof. D. S. Kostopoulos forproviding the photographic material and data at their disposal. We are especially thankfulto Prof. D. S. Kostopoulos for reviewing the manuscript and providing comments as wellas suggestions. We thank the reviewers Prof. Wang Shiqi from the IVPP and Prof. WangXiaoming from the Natural History Museums of Los Angeles County for their constructive comments and suggestions， which led to the improvement of this manuscript. The work wassupported by the Second Tibetan Plateau Scientific Expedition and Research Program （STEP）（Grant No. SQ2019QZKK0705-02）， the National Key Research and Development Program ofChina （Grant No. 2023YFF0804500）， and the Key Research Program of the Chinese Academyof Sciences （IVPP）. The first author is grateful to the University of the Chinese Academy ofSciences （UCAS） for providing financial support through the Belt amp; Road Scholarship for hisMaster’s program.

摘要：晚中新世時期出現(xiàn)了許多骨骼形態(tài)類似于現(xiàn)生麝牛（Ovibos）的中到大型食草動物，它們廣泛分布于全北界，包括11個屬：Lantiantragus， Shaanxispira， Tsaidamotherium，Criotherium， Mesembriacerus， Hezhengia， Urmiatherium， Parurmiatherium， Plesiaddax，Sinotragus和Prosinotragus。這些屬主要發(fā)現(xiàn)于東亞、東歐和西亞地區(qū)，其中9個屬在中國北方有分布。它們具有獨特的特征，包括垂直的腦顱、特化的角心、粗壯的基枕骨。以前的研究經(jīng)常將這些 “麝牛類” ?？苿游锱c現(xiàn)生的麝牛一起歸入傳統(tǒng)的麝牛亞科/麝牛族（Ovibovinae/Ovibovini）。然而，越來越多的研究表明麝牛亞科/麝牛族不是單系的，同時也不支持“麝牛類”動物與現(xiàn)生麝牛存在近的親緣關(guān)系。通常認(rèn)為在 “麝牛類” ?？苿游锏?1個屬中，近旋角羊（Plesiaddax）、和政羊（Hezhengia）和烏米獸（Urmiatherium）三屬具有更密切的關(guān)系，三者構(gòu)成烏米獸族（Urmiatheriini）。然而，先前的系統(tǒng)發(fā)育分析并不支持烏米獸族的單系性?？偨Y(jié)了Ovibovidae， Ovibovinae， Ovibovini， Ovibovina等術(shù)語的嬗變、歐亞大陸晚中新世 “麝牛類” ?？苿游锏臅r空分布、各類群的主要特征及前人的系統(tǒng)發(fā)育分析結(jié)果。

關(guān)鍵詞：歐亞大陸，晚中新世，牛科，“麝牛類” 動物，時空分布

中圖法分類號：Q915.876 文獻(xiàn)標(biāo)識碼：A 文章編號：2096–9899（2024）04–0262–29

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第二次青藏高原綜合科學(xué)考察研究項目（編號：SQ2019QZKK0705-02）、國家重點研發(fā)計劃（編號：2023YFF0804500）、中國科學(xué)院古脊椎動物與古人類研究所重點部署項目和中國科學(xué)院 “一帶一路” 碩士生獎學(xué)金資助。