欧亚大陆晚中新世 “麝牛类” 牛科动物

doi:10.19615/j.cnki.2096-9899.240902

摘要/Abstract

摘要：

晚中新世时期出现了许多骨骼形态类似于现生麝牛(Ovibos)的中到大型食草动物，它们广泛分布于全北界，包括11个属：Lantiantragus, Shaanxispira, Tsaidamotherium, Criotherium, Mesembriacerus, Hezhengia, Urmiatherium, Parurmiatherium, Plesiaddax, Sinotragus和Prosinotragus。这些属主要发现于东亚、东欧和西亚地区，其中9个属在中国北方有分布。它们具有独特的特征，包括垂直的脑颅、特化的角心、粗壮的基枕骨。以前的研究经常将这些 “麝牛类” 牛科动物与现生的麝牛一起归入传统的麝牛亚科/麝牛族(Ovibovinae/Ovibovini)。然而，越来越多的研究表明麝牛亚科/麝牛族不是单系的，同时也不支持“麝牛类”动物与现生麝牛存在近的亲缘关系。通常认为在 “麝牛类” 牛科动物的11个属中，近旋角羊(Plesiaddax)、和政羊(Hezhengia)和乌米兽(Urmiatherium)三属具有更密切的关系，三者构成乌米兽族(Urmiatheriini)。然而，先前的系统发育分析并不支持乌米兽族的单系性。总结了Ovibovidae, Ovibovinae, Ovibovini, Ovibovina等术语的嬗变、欧亚大陆晚中新世 “麝牛类” 牛科动物的时空分布、各类群的主要特征及前人的系统发育分析结果。

关键词: 欧亚大陆, 晚中新世, 牛科, “麝牛类” 动物, 时空分布

Abstract:

During the Late Miocene, numerous medium to large-sized herbivores, that resemble the living Ovibos in skeletal morphology, dispersed throughout the Holarctic realm and comprised eleven 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 northern China, as well as in countries in eastern Europe and western Asia. They are distinguished by unique characteristics, including a perpendicular braincase, specialized horn cores, and a robust basioccipital. Previous studies have often classified these ‘ovibovin’ bovids as part of the conventional subfamily/tribe Ovibovinae/Ovibovini, along with extant Ovibos. Nevertheless, an increasing 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 and extant Ovibos. Among the eleven genera of ‘ovibovin’ bovids, Plesiaddax, Hezhengia, and Urmiatherium are often considered to have a very close relationship and conventionally form the tribe Urmiatheriini. However, previous phylogenetic analyses do not support the monophyly of 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 previous phylogenetic analyses.

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

中图分类号:

Q915.876

穆罕默德·伊利亚斯, 李强, 史勤勤, 倪喜军. 欧亚大陆晚中新世 “麝牛类” 牛科动物. 古脊椎动物学报, 2024, 62(4): 262-290.

Muhammad ILYAS, LI Qiang, SHI Qin-Qin, NI Xi-Jun. The Late Miocene ‘ovibovin’ bovids in Eurasia. Vertebrata Palasiatica, 2024, 62(4): 262-290.

图/表 15

Table 1 Various concepts of scientists about the groups of fossil ‘ovibovin’ bovids

Gray	Gill	Lydekker	Bohlin	Simpson	McKenna and Bell	Grubb	Groves and Leslie	Hassanin et al.
(1872)	(1872)	(1913)	(1935a)	(1945)	(1997)	(2001)	(2011)	(2012)
Family	Family	Family	Subfamily	Superfamily	Superfamily	Family	Family	Family
Ovibovidae	Bovidae	Bovidae	Ovibovinae	Bovoidea	Bovoidea	Bovidae	Bovidae	Bovidae
	Subfamily	Subfamily		Family	Family	Subfamily	Subfamily	Subfamily
	Ovibovinae	Ovibovinae		Bovidae	Bovidae	Caprinae	Caprinae	Antilopinae
				Subfamily	Subfamily	Tribe	Tribe	Tribe
				Caprinae	Caprinae	Ovibovini	Caprini	Caprini
				Tribe	Tribe			Subtribe
				Ovibovini	Ovibovini			Ovibovina
Ovibos	Ovibos	Ovibos	†Parurmiatherium	Ovibos	Ovibos	Ovibos	Ovibos	Ovibos
Budorcas	†Bootherium		†Plesiaddax	Budorcas	Budorcas	Budorcas	Carpricornis	Capricornis
			†Pseudobos	†Boopsis	†Boopsis		Naemorhedus	Naemorhedus
			†Urmiatherium	†Bootherium	†Bootherium		Pantholops
				†Criotherium	†Criotherium		Oreamnos
				†Euceratherium	†Damalavus		Budorcas
				†Parurmiatherium	†Euceratherium		Ammotragus
				†Plesiaddax	†Lyrocerus		Arabitragus
				†Symbos	†Makapania		Hemitragus
				†Tsaidamotherium	†Megalovis		Pseudois
				†Urmiatherium	†Palaeoreas		Capra
					†Parurmiatherium		Nilgiritragus
					(including †Plesiaddax)		Ovis
					†Praeovibos		Rupicapra
					†Soergelia
					†Symbos
					†Tsaidamotherium
					†Urmiatherium

Fig. 1 Map showing the geographic distribution of fossil ‘ovibovin’ bovids in Eurasia Red dots represent the localities of the Late Miocene ‘ovibovin’ bovids. Localities 1-8 are in China, 9 in Iran, 10 in Iraq, 11-14 in Turkey, 15-17 in Greece, 18 in Bulgaria. Plesiaddax (1 Siziwangqi, 2 Hequ, 4 Baode; 12 Kayadibi, 13 Garkin), Hezhengia (3 Fugu, 7 Linxia), Urmiatherium (4 Baode, 6 Qingyang, 9 Maragheh, 14 Muğla [Salihpaşalar and Şerefköy-2], 15 Samos, 16 Fourka), Parurmiatherium (10 Injana, 14 Muğla, 15 Samos), Sinotragus (4 Baode, 14 Muğla), Prosinotragus (6 Qingyang), Shaanxispira (5 Lantian; 7 Linxia), Lantiantragus (5 Lantian), Tsaidamotherium (7 Linxia, 8 Qaidam), Criotherium (11 Çorakyerler, 15 Samos, 18 Kalimantsi), and Mesembriacerus (17 Ravin de la Pluie, N Macedonia).Map Approval Number: GS(2023)2752

Fig. 2 Biochronological distribution of the Late Miocene ‘ovibovin’ bovids in Eurasia

Table 2 List of fossil ‘ovibovin’ bovids in the Late Miocene with age and localities described (modified from Shi, 2012)

Species	Material	Locality	Country	Housed	Age (Ma)	Reference
Urmiatherium polaki	A snout and partial cranium along with horn cores	Karaj Abad site, Maragheh	Iran	DOE MNHN NHML	~8.2-7.4	Rodler, 1889; de Mecquenem, 1925; Kostopoulos and Bernor, 2011
U. intermedium	Plenty of skulls, teeth, and bone fragments	Baode (Locs. 30, 43, 44, 49 and 108) and Qingyang (Locs. 115, 116)	China	IVPP	7.2-5.3	Schlosser, 1903; Bohlin, 1925, 1935a, Shi et al., 2016
U. kassandriensis	A partial cranium and an opisthocranium with horn cores	Fourka locality, Chalkidiki	Greece	LGPUT PGMA	9.7-8.7	Lazaridis et al., 2017
Parurmiatherium rugosifrons	A partial opisthocranium, frontlets, axis, and cervical vertebra	Samos [unknown levels], Injana, Salihpaşalar and Şerefköy-2	E Greece, Iraq and SW Turkey	MYŞE NHMW BSPM	~7.4-6.7	Sickenberg, 1932, 1933; Bouvrain et al., 1995; Kostopoulos and Karakütük, 2013
Tsaidamotherium hedini	Two broken posterior parts of the skull and two cervical vertebrae	Qaidam Basin	China	IVPP	Tuosu Fauna: ~11.1-10.1	Bohlin, 1935b; Xue et al., 2006
T. brevirostrum	A complete skull, cranial part of skull with horn core apparatus, and an upper jaw	Yancaiping	China	HMV IVPP	Guonigou Fauna: ~11.1-10.5	Shi, 2014
Hezhengia bohlini	More than 100 skulls and few mandibles	Guanghe County	China	HMV IVPP	Dashenggou Fauna: ~ 9.5-8	Qiu et al., 2000; Shi and Deng, 2021
H. minor	Two fragmentary skulls and upper jaws	Fugu County	China	IVPP PMU	Lamagou Fauna: 7.8	Bohlin, 1935a
Plesiaddax depereti	Plenty of skulls, few snouts, mandibles, and isolated teeth	Baode, Hequ and Siziwang Banner	China	IVPP PMU	~7.2-5.3	Schlosser, 1903; Bohlin, 1935a; Deng et al., 2011
P. inundates	Skulls, mandibles, upper jaws and post-cranials	Garkin	Turkey	NLB BGR	~8.7-7.5	Bosscha-Erdbrink, 1978
Plesiaddax cf. P. inundates	Isolated teeth	Mahmutgazi	Turkey	BGR	~8.7-7.5	Köhler, 1987
P. simplex	Partial cranium, few mandibles and an upper jaw	Kayadibi	Turkey	BGR	~8.7-7.5	Köhler, 1987
Criotherium argalioides	Plenty of skulls, jaws and man-dibles	Samos, Mytilinii Basin (precise locality unknown) and Kemiklitepe-D, Çorakyerler	Greece and Turkey	ÇO, NHMW, NHML, BSPM, IGPM AMNH	~8.7-7.5	Solounias, 1981; Bouvrain, 1994; Kostopoulos et al., 2023
C. nikolovi	Cranium	Kalimantsi-1	Bulgaria	NMNH, Sofia	8.9-7.5	Geraads and Spassov, 2008
Shaanxispira chowi	A fractured skull with horn cores, jaw, and mandible	Lantian	China	IVPP	8.21	Liu et al., 1978
S. baheensis	A partial skull, frontlet with horn cores, and mandible	Lantian	China	IVPP	8.07	Zhang, 2003
S. linxiaensis	An almost complete and un-deformed skull	Panyang, Linxia	China	IVPP	~9.5-8	Shi et al., 2014
Mesembriacerus melentisi	Cranium	Axios valley	Greece	LGPUT	~9.3	Bouvrain and Bonis, 1984 Spassov et al., 2018
Lantiantragus longirostralis	A snout and mandible	Loc. 12, Lantian	China	IVPP	9.95	Chen and Zhang, 2004
Sinotragus wimani	Three skulls, a frontlet, and three upper jaws	Loc.30, Baode	China	PMU	~5.7	Bohlin, 1935a
S. occidentalis	A relatively complete skull without muzzle and a frontlet with cranial base	Precise locality is unknown Akgedik-Bayir, Muğla (Salihpaşalar [Kemikalan], Örucebağ [Ankara])	Turkey	MTA Museum, Ankara Izmir Museum	~7.4-6.8	Geraads et al., 2002
Prosinotragus tenuicornis	A broken cranium with horn cores	Qingyang locs. 116, 115	China	PMU	~6.5	Bohlin, 1935a

Table 2 List of fossil ‘ovibovin’ bovids in the Late Miocene with age and localities described (modified from Shi, 2012)

Species	Material	Locality	Country	Housed	Age (Ma)	Reference
Urmiatherium polaki	A snout and partial cranium along with horn cores	Karaj Abad site, Maragheh	Iran	DOE MNHN NHML	~8.2-7.4	Rodler, 1889; de Mecquenem, 1925; Kostopoulos and Bernor, 2011
U. intermedium	Plenty of skulls, teeth, and bone fragments	Baode (Locs. 30, 43, 44, 49 and 108) and Qingyang (Locs. 115, 116)	China	IVPP	7.2-5.3	Schlosser, 1903; Bohlin, 1925, 1935a, Shi et al., 2016
U. kassandriensis	A partial cranium and an opisthocranium with horn cores	Fourka locality, Chalkidiki	Greece	LGPUT PGMA	9.7-8.7	Lazaridis et al., 2017
Parurmiatherium rugosifrons	A partial opisthocranium, frontlets, axis, and cervical vertebra	Samos [unknown levels], Injana, Salihpaşalar and Şerefköy-2	E Greece, Iraq and SW Turkey	MYŞE NHMW BSPM	~7.4-6.7	Sickenberg, 1932, 1933; Bouvrain et al., 1995; Kostopoulos and Karakütük, 2013
Tsaidamotherium hedini	Two broken posterior parts of the skull and two cervical vertebrae	Qaidam Basin	China	IVPP	Tuosu Fauna: ~11.1-10.1	Bohlin, 1935b; Xue et al., 2006
T. brevirostrum	A complete skull, cranial part of skull with horn core apparatus, and an upper jaw	Yancaiping	China	HMV IVPP	Guonigou Fauna: ~11.1-10.5	Shi, 2014
Hezhengia bohlini	More than 100 skulls and few mandibles	Guanghe County	China	HMV IVPP	Dashenggou Fauna: ~ 9.5-8	Qiu et al., 2000; Shi and Deng, 2021
H. minor	Two fragmentary skulls and upper jaws	Fugu County	China	IVPP PMU	Lamagou Fauna: 7.8	Bohlin, 1935a
Plesiaddax depereti	Plenty of skulls, few snouts, mandibles, and isolated teeth	Baode, Hequ and Siziwang Banner	China	IVPP PMU	~7.2-5.3	Schlosser, 1903; Bohlin, 1935a; Deng et al., 2011
P. inundates	Skulls, mandibles, upper jaws and post-cranials	Garkin	Turkey	NLB BGR	~8.7-7.5	Bosscha-Erdbrink, 1978
Plesiaddax cf. P. inundates	Isolated teeth	Mahmutgazi	Turkey	BGR	~8.7-7.5	Köhler, 1987
P. simplex	Partial cranium, few mandibles and an upper jaw	Kayadibi	Turkey	BGR	~8.7-7.5	Köhler, 1987
Criotherium argalioides	Plenty of skulls, jaws and man-dibles	Samos, Mytilinii Basin (precise locality unknown) and Kemiklitepe-D, Çorakyerler	Greece and Turkey	ÇO, NHMW, NHML, BSPM, IGPM AMNH	~8.7-7.5	Solounias, 1981; Bouvrain, 1994; Kostopoulos et al., 2023
C. nikolovi	Cranium	Kalimantsi-1	Bulgaria	NMNH, Sofia	8.9-7.5	Geraads and Spassov, 2008
Shaanxispira chowi	A fractured skull with horn cores, jaw, and mandible	Lantian	China	IVPP	8.21	Liu et al., 1978
S. baheensis	A partial skull, frontlet with horn cores, and mandible	Lantian	China	IVPP	8.07	Zhang, 2003
S. linxiaensis	An almost complete and un-deformed skull	Panyang, Linxia	China	IVPP	~9.5-8	Shi et al., 2014
Mesembriacerus melentisi	Cranium	Axios valley	Greece	LGPUT	~9.3	Bouvrain and Bonis, 1984 Spassov et al., 2018
Lantiantragus longirostralis	A snout and mandible	Loc. 12, Lantian	China	IVPP	9.95	Chen and Zhang, 2004
Sinotragus wimani	Three skulls, a frontlet, and three upper jaws	Loc.30, Baode	China	PMU	~5.7	Bohlin, 1935a
S. occidentalis	A relatively complete skull without muzzle and a frontlet with cranial base	Precise locality is unknown Akgedik-Bayir, Muğla (Salihpaşalar [Kemikalan], Örucebağ [Ankara])	Turkey	MTA Museum, Ankara Izmir Museum	~7.4-6.8	Geraads et al., 2002
Prosinotragus tenuicornis	A broken cranium with horn cores	Qingyang locs. 116, 115	China	PMU	~6.5	Bohlin, 1935a

Fig. 3 Selected crania of Urmiatherium A, B. skulls of Urmiatherium intermedium, PMU M823 (A), PMU M3620 (B), from Baode, China;C, D. skull of U. polaki, MNHN MAR 3215, from Maragheh, Iran; E, F. partial craniums of U. kassandriensis, LGPUT FRK19 (E) and LGPUT AMPG-103 (F) from Fourka locality, GreeceThe arrows show the specific peculiarity of horn core and basioccipital in each species A, C, E. posterior views; B, D, F. ventral views Scale bars = 5 cm. C-F, courtesy of D S Kostopoulos

Fig. 4 Selected crania of Criotherium A-D. Criotherium nikolovi, NMNH FM-1740 (holotype) from Kalimantsi-1, Bulgaria;E-H. C. argalioides, ÇO 1045 from Çorakyerler, Turkey. The arrows show the specific peculiarity of parietal, nasal, basioccipital and horn core twist and direction in each species A, E. dorsal views; B, F. posterior views; C, G. ventral views; D, H. left lateral views Scale bars = 5 cm. A-D, courtesy of D Geraads; E-H, courtesy of D S Kostopoulos

Fig. 5 Selected crania of Plesiaddax A, B, C. Plesiaddax inundates, NLB SaG 69 from Garkin, Turkey;D, E, F. P. depereti, PMU M1258 from Hequ, ChinaThe arrows show the specific peculiarity of frontal, frontal sinuses and basioccipital in each species A, D. dorsal views; B, E. lateral views; C, F. ventral views Scale bars = 5 cm. A-C, courtesy of D S Kostopoulos

Fig. 6 Cranium of Parurmiatherium rugosifrons, NHMW A4758 (lectotype), from Samos, Greece The arrows show the characteristics of basioccipital and horn core direction A. posterior view; B. ventral view. Scale bars = 5 cm. A and B, courtesy of D S Kostopoulos

Fig. 7 Selected crania of Sinotragus A, B. Sinotragus occidentalis, MTA 1894 (holotype, Geraads et al., 2002: fig 2), from Yatağan Formation, Muğla, SW Turkey; C, D. S. wimani, PMU M1297 (C) and PMU M3930 (D), from Baode, ChinaThe arrows show the parietal and horn core characters in each species A, C. lateral views; B, D. latero-dorsal views. Scale bars = 5 cm. A and B, courtesy of D Geraads

Fig. 8 Skull of Prosinotragus tenuicornis, PMU M3942 from Qingyang, China The arrows show the parietal and horn core characters. A. lateral view; B. latero-dorsal view Scale bars = 5 cm

Fig. 9 Selected crania of Tsaidamotherium A, B, C. Tsaidamotherium brevirostrum, IVPP V17925 (holotype), from Linxia Basin, China;D, E, F. T. hedini, IVPP RV35052, from Qaidam Basin, ChinaThe arrows show the specific peculiarity of horn core and basioccipital in each species A, D. posterior views; B, E. ventral views; C, F. dorsal views. Scale bars = 5 cm

Fig. 10 Cranium with horn cores of Mesembriacerus melentisi, LGPUT RP l43 (holotype) from Ravin de la Pluie, Macédoine, Greece Modified from Bouvrain and Bonis, 1984. The arrows show the frontal, occipital and horn core direction A. dorsal view; B. posterior view; C. lateral view. Scale bars = 5 cm

Fig. 11 Selected crania with horn cores of Shaanxispira and upper dentition of Lantiantragus longirostralis and Shaanxispira linxiaensis A, B, H. skull (A, B) and upper dentition (H) of Shaanxispira linxiaensis, HMV 1922 (holotype), from Linxia Basin, China; C, D. skull of S. baheensis, IVPP V13626 (holotype), from Lantian County, Shaanxi Province, China; E, F. horn cores of S. chowi, IVPP V3124 (holotype) from Lantian County, China; G. upper dentition of Lantiantragus longirostralis, IVPP V13908 (holotype), from Lantian County, ChinaThe arrows show the peculiarity of the horn core twist and direction in each species, as well as the teeth similarities between Shaanxispira and Lantiantragus A. anterodorsal view; C, E. dorsal views; B, D, F. lateral views. G, H. occlusal views. Scale bars = 5 cm

Fig. 12 Skull of Hezhengia bohlini, IVPP V12349, from Linxia Basin, China The arrows show the characters of basiocciptal, frontal and horn core direction A. dorsal view; B. lateral view; C. ventral view. Scale bar = 5 cm

Fig. 13 Different author’s concepts about the lineage of ‘ovibovin’ bovids based on morphological and molecular analysis

参考文献 75

[1]	Bibi F, Tyler J, 2022. Evolution of the bovid cranium: morphological diversification under allometric constraint. Commun Biol, 5: 69 DOI PMID
[2]	Bibi F, Bukhsianidze M, Gentry A W et al., 2009. The fossil record and evolution of Bovidae: state of the field. Palaeont Electron, 12: 1-11
[3]	Bibi F, Vrba E, Fack F, 2012. A new African fossil caprin and a combined molecular and morphological bayesian phylogenetic analysis of caprini (Mammalia: Bovidae). J Evol Biol, 25: 1843-1854
[4]	Bohlin B, 1925. Urmiatherium intermedium (Schlosser). Bull Geol China, 7: 111-113
[5]	Bohlin B, 1935a. Cavicornier der Hipparion-Fauna Nord-China. Palaeont Sin Ser C, 9: 1-166
[6]	Bohlin B, 1935b. Tsaidamotherium hedini, n. g., n. sp. Geogr Ann, 17: 66-74
[7]	Bohlin B, 1937. Einige Bemerkungen über die Hörner der Ovibovinae. Bull Geol Instit Uppsala, 27: 42-47
[8]	Bosscha P, Erdbrink D, 1978. Fossil ovibovines from Garkin near Afyon, Turkey. Proc K Ned Akad Wet, Ser B, 81: 145-185
[9]	Bouvrain G, 1975. Un nouveau bovidé du Vallésien de Macédoine, Grèce. C R Acad Sci Paris, 280: 1357-1359
[10]	Bouvrain G, 1994. The late Miocene mammal localities of Kemiklitepe, Turkey: 9. Bovidae. Bull Mus Natl Hist Nat, Sect C: Sci Terre: Paleontol Geol Mineral, 16: 175-209
[11]	Bouvrain G, Bonis L de, 1984. Le genre Mesembriacerus (Bovidae, Artiodactyla, Mammalia): un ovibovin primitif du Vallésian (Miocène supérieur) de Macédoine (Grèce). Palaeovertebrata, 14: 201-223
[12]	Bouvrain G, Sen S, Thomas H, 1995. Parurmiatherium rugosifrons Sickenberg, 1932, un Ovibovinae (Bovidae) du Miocène supèrieur d’Injana (Djebel Hamrin, Irak). Geobios, 28: 719-726
[13]	Chen G F, Zhang Z Q, 2004. Lantiantragus gen. nov. (Urmiatherinae, Bovidae, Artiodactyla) from the Bahe Formation, Lantian, China. Vert PalAsiat, 42: 205-215
[14]	Chen G F, Zhang Z Q, 2009. Taxonomy and evolutionary process of Neogene Bovidae from China. Vert PalAsiat, 47: 265-281
[15]	de Mecquenem R, 1925. Contribution à l’étude des fossils de Maragha. Ann Paléontol, 14: 1-36
[16]	Deng T, Liang Z, Wang S Q et al., 2011. Discovery of a Late Miocene mammalian fauna from Siziwang Banner, Inner Mongolia, and its paleozoogeographical significance. Chinese Sci Bull, 56: 526-534
[17]	Deng T, Qiu Z X, Wang B Y et al., 2013. 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, Hou S K, Wang S Q, 2019. Neogene integrative stratigraphy and timescale of China. Sci China Earth Sci, 62: 310-323 DOI
[19]	Forsyth-Major C I F, 1891. Considérations nouvelles sur la faune des Vertébrés du Miocène supérieur dans l’ île de Samos. C R Acad Sci, 108: 608-610
[20]	Gentry A W, 1971. The earliest goats and other antelopes from the Samos Hipparion fauna. Bull Br Mus Nat Hist Geol, 20: 229-296
[21]	Gentry A W, 2000. Caprinae and Hippotragini (Bovidae, Mammalia) in the Upper Miocene. In: Vrba E, Schaller G eds. Antelopes, Deer and Relatives. New Haven: Yale University Press. 65-83
[22]	Gentry A W, Rössner G E, Heizmann E P J, 1999. Suborder Ruminantia. In: Rössner G E, Heissig K eds. The Miocene Land Mammals of Europe. München: Friedrich Pfeil. 225-258
[23]	Geraads D, Spassov N, 2008. A new species of Criotherium (Bovidae, Mammalia) from the late Miocene of Bulgaria. Hell J Geosci, 43: 21-27
[24]	Geraads D, Güleç E, Kaya T, 2002. Sinotragus (Bovidae, Mammalia) from Turkey and the Late Miocene Asiatic Province. Neues Jahrb Geol Paläont Monatsh, 8: 477-489
[25]	Gill T, 1872. Arrangement of the families of mammals with analytical tables. Smithson Misc Collect, 11: 1-98
[26]	Gray J E, 1821. On the natural arrangement of vertebrose animals. London Med Reposit, 15: 296-310
[27]	Gray J E, 1872. Catalogue of the ruminant Mammalia (Pecora, Linnaeus) in the British Museum. London: Trustees of the British Museum. 1-102
[28]	Groves C P, Leslie D M, 2011. Family Bovidae. In: Wilson D E, Mittermeier R A eds. Handbook of the Mammals of the World. 2. Hoofed Mammals. Barcelona: Lynx Edicions. 444-779
[29]	Grubb P, 1993. Family Bovidae. In: Wilson D E, Reeder D A M ed. Mammal Species of the World. A Taxonomic and Geographic Reference, 2nd ed. , Washington DC: Smithsonian Institution Press. 393-414
[30]	Grubb P, 2001. Review of family-group names of living bovids. J Mammal, 82: 374-388
[31]	Hassanin A, Ropiquet A, Couloux A et al., 2009. Evolution of the mitochondrial genome in mammals living at high altitude: new insights from a study of the tribe Caprini (Bovidae, Antilopinae). J Mol Evol, 68(4): 293-310 DOI PMID
[32]	Hassanin A, Delsuc F, Ropiquet A et al., 2012. Pattern and timing of diversification of Cetartiodactyla (Mammalia, Laurasiatheria), as revealed by a comprehensive analysis of mitochondrial genomes. C R Biol, 335: 32-50
[33]	Hou S K, Shi Q Q, Benton M J et al., 2023. Comment on “Sexual selection promotes giraffoid head-neck evolution and ecological adaptation”. Science, 379: eadd9559
[34]	Jafarzadeh R, Kostopoulos D, Daneshian J, 2012. Skull reconstruction and ecology of Urmiatherium polaki (Bovidae, Mammalia) from the upper Miocene deposits of Maragheh, Iran. PalZ, 86: 103-111
[35]	Kaya T, Mayda S, Kostopoulos D S et al., 2012. Şerefköy-2, a new Late Miocene mammal locality from the Yatağan Formation, Muğla, SW Turkey. C R Palevol, 11: 5-12
[36]	Kingdon J, 1989. East African Mammals: an Atlas of Evolution in Africa. Vol. IIIC and D. Bovids. Chicago: University of Chicago Press. 1-746
[37]	Köhler M, 1987. Boviden des türkischen Miozäns (Känozoikum and Braunkohlen der Türkei). Paleont Evol, 21: 133-246
[38]	Kostopoulos D S, 2009. The late Miocene mammal faunas of the Mytilinii basin, Samos Island, Greece: new collection. 14. Bovidae. Beitr Paläontol, 31: 345-389
[39]	Kostopoulos D S, 2014. Taxonomic re-assessment and phylogenetic relationships of Miocene homonymously spiral-horned antelopes. Acta Palaeontol Pol, 59: 9-29
[40]	Kostopoulos D S, Bernor R L, 2011. The Maragheh bovids (Mammalia, Artiodactyla): systematic revision and biostratigraphic-zoogeographic interpretation. Geodiversitas, 33: 649-708
[41]	Kostopoulos D S, Karakütük S, 2013. Late Miocene bovids from Şerefköy-2 (SW Turkey) and their position within the subParatethyan biogeographic province. Acta Palaeontol Pol, 60: 49-66
[42]	Kostopoulos D S, Erol A S, Mayda S, 2023. Late Miocene ‘οvibovin’ bovids (Mammalia, Bovidae) from Çorakyerler, Turkey. J Vert Paleont, 43: e2232850
[43]	Lalueza-Fox C, Castresana J, Sampietro L et al., 2005. Molecular dating of caprines using ancient DNA sequences of Myotragus balearicus, an extinct endemic Balearic mammal. BMC Evol Biol, 5: 70 PMID
[44]	Lazaridis G, Kostopoulos D S, Lyras G et al., 2017. A new Late Miocene ovibovine-like bovid (Bovidae, Mammalia) from the Kassandra Peninsula (Chalkidiki, northern Greece) and implications to the phylogeography of the group. PalZ, 91: 427-437
[45]	Lent P C, 1988. Ovibos moschatus. Mamm Spec, 302: 1-9
[46]	Liu T S, Li C K, Zhai R J, 1978. Pliocene vertebrates of Lantian, Shensi. Prof Pap Stratigr Palaeontol, 7: 149-200
[47]	Lydekker R, 1913. Catalogue of the ungulate mammals in the British Museum (Natural History). Vol. 1. London: Trustees of the British Museum, 1-249
[48]	McKenna M C, Bell S K, 1997. Classification of Mammals above the Species Level. New York: Columbia University Press. 1-631
[49]	Owen R, 1848. Description of teeth and portions of jaw of two extinct anthracotherioid quadrupeds (Hyopotamus vectianus and Hyopbovinus) discovered by the Marchioness of Hastings in the Eocene deposits on the NW coast of the Isle of Wight: with an attempt to develop Cuvier’s idea of the classification of pachyderms by the number of their toes. Quart J Geol Soc London, 4: 103-141
[50]	Qiu Z X, Wang B Y, Xie G P, 2000. Preliminary report on a new genus of Ovibovinae from Hezheng District, Gansu, China. Vert PalAsiat, 38: 128-134
[51]	Qiu Z X, Qiu Z D, Zheng Y et al., 2023. Stratigraphic context of Oligocene to Pliocene mammal-bearing deposits in Linxia Basin, Gansu Province, China: a historical review and a discussion of ongoing controversies. Palaeogeogr Palaeoclimatol Palaeoecol, 628: 111773
[52]	Rodler A, 1889. Über Urmiatherium polaki n. g. n. sp., einen neuen Sivatheriiden aus dem Knochenfelde von Maragha. Denkschriften Kaiserlichen Akad Wiss Math-Naturwiss Kl, 56: 315-322
[53]	Ropiquet A, Hassanin A, 2005. Molecular phylogeny of caprines (Bovidae, Antilopinae): the question of their origin and diversification during the Miocene. J Zool Syst Evol Res, 43: 49-60
[54]	Schlosser M, 1903. Die fossilen Saugethiere Chinas nebst einer Odontographie der recenten Antilopen. Abh K Buyer AM Wiss, 22: 1-221
[55]	Shi Q Q, 2012. Two late Miocene ovibovines from Houshan locality, Linxia Basin, Gansu Province. Ph. D thesis. Beijing: University of Chinese Academy of Sciences. 1-146
[56]	Shi Q Q, 2014. New species of Tsaidamotherium (Bovidae, Artiodactyla) from China sheds new light on the skull morphology and systematics of the genus. Sci China Earth Sci, 57: 258-266
[57]	Shi Q Q, 2023. Fossil bovids from the Linxia Basin of Gansu Province, China, and their implications for regional biostratigraphy, palaeogeography and palaeoecology. Palaeogeogr Palaeoclimatol Palaeoecol, 619: 111543
[58]	Shi Q Q, Deng T, 2021. Redescription of the skull of Hezhengia bohlini (Artiodactyla, Mammalia) and a reassessment of the systematics of the Chinese late Miocene ‘ovibovines’. J Syst Palaeontol, 18: 2059-2074
[59]	Shi Q Q, He W, Chen S Q, 2014. A new species of Shaanxispira (Bovidae, Artiodactyla) from the upper Miocene of China. Zootaxa, 3794: 501-513
[60]	Shi Q Q, Wang S Q, Chen S K et al., 2016. The first discovery of Urmiatherium (Bovidae, Artiodactyla) from Liushu Formation, Linxia Basin. Vert PalAsiat, 54: 319-331
[61]	Sickenberg O, 1932. Eine neue Antilope, Parurmiatherium rugosifrons nov. gen. nov. sp., aus dem Unterpliozan von Samos. Anz Osterreichischen Akad Wiss, Math-Naturwiss, 1: 10-11
[62]	Sickenberg O, 1933. Parurmiatherium rugosifrons ein neuer Bovide aus dem Unterpliozän von Samos. Palaeobiologica, 5: 81-102
[63]	Simpson G G, 1945. The principles of classification and a classification of mammals. Bull Am Mus Nat Hist, 85: 1-350
[64]	Solounias N, 1981. The Turolian fauna from the island of Samos, Greece, with special emphasis on the hyaenids and the bovids. Contrib Vert Evol, 6: 1-232
[65]	Spassov N, Geraads D, Hristova L et al., 2018. The late Miocene mammal faunas of the Republic of Macedonia (FYROM). Palaeontogr Abt A, 311: 1-85
[66]	Tseng Z J, Wang X, Li Q et al., 2022. Qurliqnoria (Mammalia: Bovidae) fossils from Qaidam Basin, Tibetan Plateau and deep-time endemism of the Tibetan antelope lineage. Zool J Linn Soc, 196(3): 990-1012
[67]	Vrba E S, 1985. African Bovidae: evolutionary event since the Miocene. S Afr J Sci, 81: 263-266
[68]	Wang S Q, Ye J, 2023. The enigmatic spherical-horned Tsaidamotherium reveals an unsuccessful grazing adaptation of early giraffoids. Innov Life, 1: 100035-1
[69]	Wang S Q, Meng J, Mennecart B et al., 2023a. Response to comment on “Sexual selection promotes giraffoid head-neck evolution and ecological adaptation”. Science, 379: 3392
[70]	Wang S Q, Ye J, Meng J et al., 2023b. Sexual selection promotes giraffoid headneck evolution and ecological adaptation. Science, 376: eabl8316
[71]	Wang X M, Xie G P, Li Q et al., 2011. Early explorations of Qaidam Basin (Tibetan Plateau) by Birger Bohlin - reconciling classic vertebrate fossil localities with modern biostratigraphy. Vert PalAsiat, 49: 285-310
[72]	Xue X X, Zhang Y X, Yue L P, 1995. Discovery and chronological division of the Hipparion fauna in Laogaochuan Village, Fugu County, Shaanxi. China Sci Bull, 40: 926-929
[73]	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-Earth Sci, 49: 518-530
[74]	Zhang Z Q, 2003. A new species of Shaanxispira (Bovidae, Artiodactyla, Mammalia) from the Bahe Formation, Lantian, China. Vert PalAsiat, 41: 230-239
[75]	Zhang Z Q, Kaakinen A, Liu L P et al., 2013. 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