A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China

doi:10.19615/j.cnki.2096-9899.211025

Vertebrata Palasiatica ›› 2022, Vol. 60 ›› Issue (2): 81-116.DOI: 10.19615/j.cnki.2096-9899.211025

A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China

Henry GALIANO¹, Z. Jack TSENG²^,³^,⁴(), Nikos SOLOUNIAS³^,⁵, WANG Xiao-Ming³^,⁴^,⁶, QIU Zhan-Xiang⁶^,⁷, Stuart C. WHITE⁸

1 Maxilla and Mandible, Ltd 105 West 86th Street Suite # 117, New York 10024, USA
2 Department of Integrative Biology and Museum of Paleontology, University of California Berkeley 94720, USA
3 Division of Paleontology, American Museum of Natural History New York 10024, USA
4 Department of Vertebrate Paleontology, Natural History Museum of Los Angeles County Los Angeles 90007, USA
5 Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine Old Westbury, New York 11568, USA
6 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing 100044, China
7 University of Chinese Academy of Sciences Beijing 100049, China
8 School of Dentistry, University of California Los Angeles 90095, USA

Received:2021-05-25 Online:2022-04-20 Published:2021-10-26
Contact: * zjt@berkeley.edu

甘肃临夏盆地中新世鬣狗科一新属新种

Henry GALIANO¹, 曾志杰²^,³^,⁴(), Nikos SOLOUNIAS³^,⁵, 王晓鸣³^,⁴^,⁶, 邱占祥⁶^,⁷, Stuart C. WHITE⁸

1 美国Maxilla and Mandible有限公司纽约 10024
2 美国加州大学伯克利分校,加州大学古生物博物馆伯克利 94720
3 美国纽约自然历史博物馆纽约 10024
4 美国洛杉矶自然历史博物馆洛杉矶 90007
5 美国纽约理工大学医学院纽约 11568
6 中国科学院古脊椎动物与古人类研究所北京 100044
7 中国科学院大学北京 100049
8 美国加州大学洛杉矶分校牙医学院洛杉矶 90095

通讯作者: * zjt@berkeley.edu
基金资助:
美国科学基金会(DEB-1257572);美国纽约自然历史博物馆Frick博士后项目2013-2016;美国怀俄明恐龙国际有限责任公司和美国纽约Maxilla and Mandible有限公司资助

1. .zip(20KB)

Abstract

Abstract:

The aardwolf Proteles cristatus is the only known hyaenid, living or extinct, to exhibit an extremely reduced dentition related to its termite-specializing diet. The fossil record of extant aardwolves extends to 2 to 4 million years ago, but records that inform its evolutionary origins are essentially nonexistent. Such circumstance renders it difficult to place this unusual hyena in the broader evolutionary context of small-bodied hyaenid species in Eurasian Neogene deposits. Here we describe a new genus and species of a small-bodied hyaenid, Gansuyaena megalotis, representing the closest morphological link to aardwolves to date. This new fossil hyena is based on a skull with associated mandible, a rostrum preserving several teeth, and several referred specimens. The new specimens were discovered in Neogene deposits in Linxia Basin, Gansu Province, China. Phylogenetic analysis indicates that among early hyaenids, G. megalotis is most closely related, but unlikely ancestral, to the living aardwolf. Also recognized in this new species are the fossils previously referred to “Protictitherium” aff. P. gaillardi from Pasalar, Turkey. Additionally, “Plioviverrops” guerini from Los Mansuetos, Spain is interpreted to represent a second Gansuyaena species. In addition to the living aardwolf, Proteles cristatus, our analyses suggest that the proteline lineage includes the extinct genera Gansuyaena, Mesoviverrops, and Plioviverrops. Although the precise timing and geographic location of evolutionary divergence between the aardwolf and Gansuyaena remain elusive, critical new morphological information provided by Gansuyaena specimens reinforce findings from recent genomic analyses that the aardwolf lineage has an ancient origin from small-bodied stem hyaenids prior to the appearance of large and robust bone-cracking hyaenines.

Key words: Linxia Basin, Gansu, Middle and Late Miocene, Hyaenidae, Protelinae

摘要：

非洲土狼(Proteles cristatus)是鬣狗科中牙齿最为退化的一个种类,与其他化石及现生鬣狗物种粗壮的碎骨型牙齿形态截然不同。它的化石记录可以追溯到上新世,但尚缺少中间过渡形态的化石记录,以致生物学家对这一罕见的食蚁性食肉目动物的起源演化缺乏清楚的认知。描述了鬣狗科化石一新属甘肃鬣狗(Gansuyaena), 并讨论其对于土狼起源的重要指示。所记述的标本为从海外收回的甘肃临夏盆地新近纪材料,包括两件破损头骨和下颌。基于甘肃鬣狗的形态特征,进行了一系列的系统发育分析,结果显示Proteles, Mesoviverrops, Plioviverrops和Gansuyaena组成一个相对原始的鬣狗类群,即土狼亚科(Protelinae)。而且,新的形态证据支持土狼亚科在形态粗壮的鬣狗亚科出现之前就已形成一个独立的支系。

关键词: 甘肃临夏盆地, 中晚中新世, 鬣狗科, 土狼亚科

CLC Number:

Q915.874

Henry GALIANO, Z. Jack TSENG, Nikos SOLOUNIAS, WANG Xiao-Ming, QIU Zhan-Xiang, Stuart C. WHITE. A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China. Vertebrata Palasiatica, 2022, 60(2): 81-116.

Henry GALIANO, 曾志杰, Nikos SOLOUNIAS, 王晓鸣, 邱占祥, Stuart C. WHITE. 2022, 60(2): 81-116, 甘肃临夏盆地中新世鬣狗科一新属新种. 古脊椎动物学报.

Figures/Tables 11

Fig. 1 Gansuyaena megalotis gen. et sp. nov., IVPP V 13507-1 Holotype cranium (A-D) and line drawing with key anatomical features labeled (E-H) in right lateral (A, E), left lateral (B, F), dorsal (C, G), and ventral (D, H) views. For abbreviations see text

Fig. 2 Gansuyaena megalotis gen. et sp. nov., IVPP V 13507, holotype dentition A, B. V 13507-1, upper dentition: A. occlusal surface, B. lateral view; C-F. V 13507-2, lower posterior dentition: C, E. right lower dentition, D, F. left lower dentition, C, D. lateral views, E, F. occlusal views

Fig. 3 Gansuyaena megalotis gen. et sp. nov., IVPP V 13507-1, holotype, partially segmented CT scan images A. medial view of right basicranial area with a cross sectional (along rostrocaudal axis) slice of right bulla in CT scan images, indicating location of slice in B; B. segmentation of bony elements are assigned cross-hatched color patterns (dorsal is up); C. dorsolateral view of right bulla as digitally reconstructed, with the external bulla made semi-transparent to show internal structures; D. ventral view of right bulla as in C Color codes in segmentation: light blue, basioccipital and basisphenoid; deep blue, petrosal and ventral promontorial process; yellow (solid in A or semi-transparent in C and D), external bulla (presumably mostly ectotympanic); red, dorsal laminar of bullar septum (ectotympanic); pink, ventral laminar of bullar septum (entotympanic). Colors seen through semi-transparent bulla has a paler tone

Fig. 4 Gansuyaena megalotis gen. et sp. nov., IVPP V 13507-2, holotype mandibular fragments A, B, E, F. right hemimandibular fragment; C, D, G, H. left hemimandibular fragment;A, D, E, H. lateral views; B, C, F, G. medial views; A-D. photos;E-H. line drawing with key dental morphology labeled. For abbreviations see text

Fig. 5 Gansuyaena megalotis gen. et sp. nov., reconstruction of the skull based on IVPP V 13507-1, V 13507-2, and V 13508 Illustration by Mick Ellison. Reprinted from Mick Ellison under a CC BY license, with permission from Mick Ellison, original copyright 2020

Fig. 6 Gansuyaena megalotis gen. et sp. nov., IVPP V 13508, fragment of rostrum A. dorsal view; B. ventral view; C. left lateral view

Table 1 Skull measurements of Gansuyaena and Plioviverrops (mm)

Skull and mandible	IVPP V 13507-1, V 13507-2	IVPP V 13508	P. orbignyi Gaudry 1862
Length from ant end of premaxilla to end of occipital condyle	90.6 (R) 102.0 (L)	-	~105
Length from ant end of premaxilla to cleft of occipital condyle	94.9	-	(100.9)
Length from ant end of premaxilla to end of glenoid process	72.5	-	(77.4)
Postorbital width	18.3	-
Depth of braincase from basioccipital to top of parietal	25.8	-
Diameter of orbit	17.7	-
Transverse width inside nasal opening	9	6.6
Width across canines	16.4	15.9
Bulla length × width	20.5×11.9	-
Diameter of external auditory meatus	7	-
Width across mastoid processes	29.5	-
Transverse width occipital condyles	15.5	-
Width across P4s	32.8	-
Depth of skull at P4/M1 junction	26.1	-	(23.4)
Width of incisor series across from outer edges of I3	11.1	10
Length of mandibular symphysis to mandibular condyle	66.5	-	~80
Width across canines	9.9	-
Depth below p1&2	8.5	-	(10.7)
Depth below p2&3	8.9	-	(12.5)
Depth below m1	11.2	-	(12.5)
Thickness below m1	4.6	-

Table 2 Upper dentition measurements of Gansuyaena and Plioviverrops (mm)

Upper dentition	IVPP V 13507-1	V 13508	BSP-1967-VI 743	P. orbignyi Gaudry 1862
I1 width	1.5	-
I2 width	1.5	-
I3 width	1.9	1.8
Canine length × width	3.9×2.9	4.0×2.5
tooth row I1-M2	52.4 (approx.)	50.4		~54
P1 length × width	2.5×1.6	3.2×1.9		(4.4)
P2 length × width	6.2×2.1	5.3×2.4		(6.8)
P3 length × width	7.4×3.3	6.4×3.1 (approx.)		(6.3)
P4 length from parastyle to end of metacone	11.1	10.5	13.6	(10.4)
P4 length from protocone to end of metacone	12.7	12
P4 parastyle length from anterior edge to notch	1.7	1.4
P4 length from parastyle notch to carnassial notch	4.4	4
P4 length of metacone blade	3.4	3.3
P4 protocone basin anteroposterior width	2.5	2.9
P4 parastyle labial height	3.1	3.2
P4 paracone labial height	4	4.4
P4 metacone labial height	3	3.1	BSP-1967-VI 747
M1 length × width	4.6×7.4	4.5×9.0	5.9×11.6
M2 length × width	3.5×4.5	3.2×4.5	4.1×6.3

Table 3 Lower dentition measurements of Gansuyaena and Plioviverrops (mm)

Lower dentition	IVPP V 13507-2		G. guerini Los Mansuetos	P. orbignyi Gaudry 1862
Length of tooth row from canine to end of m2	40.8			~53
Length from anterior end of p4 to posterior end of m1	17.2
i1 width	1.1
i2 width	1.1
i3 width	1.5
Canine anteroposterior width	3.5
p1 length × width	1.5×?
p2 length × width	5.1×?		5.5×2.5	(6.5)×-
p3 length × width	?	BSP-1967-VI 746	6.9×3.0	(7.1)×-
p4 length × width	8.1×3.1	9.0×4.2	8.9×4.0	(7.5)×-
p4 height of principle cusp from posterior base of crown	5	BSP-1967-VI 745
m1 length × width	8.7×3.9	11.3×5.1	11.4×5.5	(8.4)×-
m1 height of protoconid from anterior crown base	4.4
m1 trigonid length × width	5.6×3.9
m1 talonid length × width	3.1×3.1		3.1×-
m2 length × width	3.9×2.5

Fig. 7 Comparison of the upper and lower dentitions of early fossil hyaenids A-C. lower premolar 4 and molars in lingual (A), labial (B), and occlusal (C) views; D. upper premolar 4 and molars in occlusal views

Fig. 8 Hyaenid phylogeny Single most parsimonious tree from parsimony analysis of 44 morphological characters in 11 taxa using the implicit enumeration method in TNT. Dark notches indicate clade synapomorphies and taxon autapomorphies, with character number followed by character state in parentheses. Labeled nodes correspond to synapomorphies listed in Table S2. The full data matrix is available in MorphoBank (http://dx.doi.org/10.7934/P1088)

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A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China

甘肃临夏盆地中新世鬣狗科一新属新种

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[1]	WANG Ban-Yue, QIU Zhan-Xiang. Micromammal fossils from the basal part of the Jiaozigou Formation in Yagou area, Linxia Basin, Gansu Province . Vertebrata Palasiatica, 2023, 61(4): 284-316.
[2]	WANG Ban-Yue, QIU Zhan-Xiang, WANG Shi-Qi. Mansancun Micromammal Fauna from the upper part of the Jiaozigou Formation in Linxia Basin, Gansu Province . Vertebrata Palasiatica, 2023, 61(2): 123-141.
[3]	WANG Ban-Yue. A new species of Pararhizomys (Tachyoryctoidinae, Muroidea) from Linxia Basin of Gansu Province . Vertebrata Palasiatica, 2022, 60(4): 271-277.
[4]	ZHANG Xiao-Xiao, SUN Dan-Hui. A cuboid bone of a large Late Miocene elasmothere from Qingyang, Gansu, and its morphological significance . Vertebrata Palasiatica, 2022, 60(1): 29-41.
[5]	LI Zhi-Heng, Alida M. BAILLEUL, Thomas A. STIDHAM, WANG Min, DENG Tao. Exceptional preservation of an extinct ostrich from the Late Miocene Linxia Basin of China . Vertebrata Palasiatica, 2021, 59(3): 229-244.
[6]	WANG Ban-Yue, QIU Zhan-Xiang. New Hystrix (Hystricidae, Rodentia) from the Neogene of Linxia Basin, Gansu, China . Vertebrata Palasiatica, 2020, 58(3): 204-220.
[7]	WANG Ban-Yue, QIU Zhan-Xiang, LI Lü-Zhou. A Late Miocene Huerzelerimys (Rodentia: Muridae) skull from Hezheng, Gansu, China . Vertebrata Palasiatica, 2020, 58(2): 120-133.
[8]	XIONG Wu-Yang. Basicranial morphology of Late Miocene Dinocrocuta gigantea (Carnivora: Hyaenidae) from Fugu, Shaanxi . Vertebrata Palasiatica, 2019, 57(4): 274-307.
[9]	HOU Su-Kuan, DENG Tao. A new species of Kubanochoerus (Suidae, Artiodactyla) from the Linxia Basin, Gansu Province, China . Vertebrata Palasiatica, 2019, 57(2): 155-172.
[10]	CHEN He, WANG Shi-Qi, TAO Da-Wei, XIA Xiu-Min, CHEN Shan-Qin,WU Yan. Implications for Late Miocene diet from Diceros gansuensis: starch granules in tooth calculus . Vertebrata Palasiatica, 2018, 56(4): 343-353.
[11]	SUN Dan-Hui, LI Yu, DENG Tao. A new species of Chilotherium (Perissodactyla, Rhinocerotidae) from the Late Miocene of Qingyang, Gansu, China . Vertebrata Palasiatica, 2018, 56(3): 216-228.
[12]	Dana BIASATTI, WANG Yang, DENG Tao. Paleoecology of Cenozoic rhinos from northwest China: a stable isotope perspective . Vertebrata Palasiatica, 2018, 56(1): 45-68.
[13]	YANG Xiang-Wen, LI Yu , WANG Shi-Qi . Cranial and dental material of Gomphotherium wimani (Gomphotheriidae, Proboscidea) from the Middle Miocene of the Linxia Basin, northwestern China . Vertebrata Palasiatica, 2017, 55(4): 331-346.
[14]	DENG Tao, ZHANG Yun-Xiang, Zhijie J. TSENG, HOU Su-Kuan. A skull of Machairodus horribilis and new evidence for gigantism as a mode of mosaic evolution in machairodonts (Felidae, Carnivora) . Vertebrata Palasiatica, 2016, 54(4): 302-318.
[15]	SHI Qin-Qin, WANG Shi-Qi, CHEN Shao-Kun, LI Yi-Kun . The first discovery of Urmiatherium (Bovidae, Artiodactyla) from Liushu Formation, Linxia Basin . Vertebrata Palasiatica, 2016, 54(4): 319-331.