A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China
Received date: 2021-05-25
Online published: 2021-10-26
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
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[J]. Vertebrata Palasiatica, 2022 , 60(2) : 81 -116 . DOI: 10.19615/j.cnki.2096-9899.211025
| [1] | Bernor R L, Tobien H, 1990. The mammalian geochronology and biogeography of Paşalar (Middle Miocene, Turkey). J Hum Evol, 19: 551-568 |
| [2] | Colbert E H, 1939. Carnivora of the Tung Gur Formation of Mongolia. Bull Am Mus Nat Hist, 76: 47-81 |
| [3] | Crusafont-Pairó M, Petter G, 1969. Contribution a l’etude des Hyaenidae. La sous-famille des Ictitheriinae. Ann Paleontol Vertebr, 55: 87-127 |
| [4] | de Beaumont G, 1968. Observations sur les Herpestinae (Viverridae, Carnivora) de l’oligocene superieur avec quelques remarques sur des Hyaenidae du neogene. Arch Sci, 20: 79-107 |
| [5] | de Beaumont G, 1969. Breves remarques sur Plioviverrops Kretzoi (Carnivora). Bull Soc Vaudoise Sci Nat, 70(6): 247-253 |
| [6] | de Beaumont G, Mein P, 1972. Recherches sur le genre Plioviverrops Kretzoi (Carnivora, ?Hyaenidae). Arch Sci Geneve, 25: 383-394 |
| [7] | de Vries J L, Pirk C W W, Bateman P W, et al. 2011. Extension of the diet of an extreme foraging specialist, the aardwolf (Proteles cristata). Afr Zool, 46: 194-196 |
| [8] | 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 |
| [9] | Depéret C, 1892. La fauna de mammiféres miocénes de La Grive-Saint-Alban (Isere). Arch Mus Hist Nat Lyon, 5: 1-89 |
| [10] | Fang X M, Wang J Y, Zhang W L, et al. 2016. Tectonosedimentary evolution model of an intracontinental flexural (foreland) basin for paleoclimatic research. Glob Planet Change, 145: 78-97 |
| [11] | Filhol H, 1883. Note sur quelques Mammifères fossiles de l’époque miocène. Publ Mus Confluences, 3: 1-99 |
| [12] | Forsyth-Major C I, 1903. III.-New Carnivoka from the Middle Miocene of La Grive-Saint-Alban, Isère, France. Geol Mag, 10: 534-538 |
| [13] | Gaillard C, 1899. Mammifères miocènes nouveaux ou peu connus de la Grive-Saint-Alban (Isère). Publ Mus Confluences, 7: 1-11 |
| [14] | Galiano H, Frailey D, 1977. Chasmaporthetes kani, new species from China, with remarks on phylogenetic relationships of genera within the Hyaenidae (Mammalia, Carnivora). Am Mus Novit, 2632: 1-16 |
| [15] | Gaudry A, 1862. Animaux fossiles et géologie de l’Attique: d’après les recherches faites en 1855-56 et en 1860 sous les auspices de l’Académie des Sciences. Paris: F. Savy. 1-474 |
| [16] | Granger D E, Gibbon R J, Kuman K, et al. 2015. New cosmogenic burial ages for Sterkfontein Member 2 Australopithecus and Member 5 Oldowan. Nature, 522: 85-88 |
| [17] | Gregory W K, 1910. The orders of mammals. Bull Am Mus Nat Hist, 27: 1-524 |
| [18] | Gregory W K, Hellman M, 1939. On the evolution and major classification of the civets (Viverridae) and allied fossil and recent Carnivora: a phylogenetic study of the skull and dentition. Proc Am Philos Soc, 81: 309-392 |
| [19] | Hendey Q B, 1974. New fossil carnivores from the Swartkrans australopithecine site (Mammalia: Carnivora). Ann Transvaal Mus, 29: 27-47 |
| [20] | Hough M J, 1953. Auditory region in North American fossil Felidae; its significance in phylogeny. US Geol Surv Prof Pap, 243-G: 95-115 |
| [21] | Hunt Jr R M, 1974. The auditory bulla in Carnivora: an anatomical basis for reappraisal of carnivore evolution. J Morphol, 143: 21-75 |
| [22] | Hunt Jr R M, 1987. Evolution of the aeluroid Carnivora: significance of auditory structure in the nimravid cat Dinictis. Am Mus Novit, 2886: 1-74 |
| [23] | Hunt Jr R M, 1989. Evolution of the aeluroid Carnivora: significance of the ventral promontorial process of the petrosal, and the origin of basicranial patterns in the living families. Am Mus Novit, 2930: 1-32 |
| [24] | Hunt Jr R M, 1991. Evolution of the aeluroid Carnivora: viverrid affinities of the Miocene carnivoran Herpestides. Am Mus Novit, 3023: 1-34 |
| [25] | Hunt Jr R M, 1998. Evolution of the aeluroid Carnivora: diversity of the earliest aeluroids from Eurasia (Quercy, Hsanda-Gol) and the origin of felids. Am Mus Novit, 3252: 1-65 |
| [26] | Hunt Jr R M, 2001. Basicranial anatomy of the living linsangs Prionodon and Poiana (Mammalia, Carnivora, Viverridae), with comments on the early evolution of aeluroid carnivorans. Am Mus Novit, 3330: 1-24 |
| [27] | Hunt Jr R M, Solounias N, 1991. Evolution of the aeluroid Carnivora: hyaenid affinities of the Miocene carnivoran Tungurictis spocki from Inner Mongolia. Am Mus Novit, 3030: 1-25 |
| [28] | Hunt Jr R M, Tedford R H, 1993. Phylogenetic relationships within the aeluroid Carnivora and implications of their temporal and geographic distribution. In: Szalay F, Novacek M, McKenna M eds. Mammal Phylogeny: Placentals. New York: Springer-Verlag. 53-73 |
| [29] | ICZN, 1999. International Code of Zoological Nomenclature. 4th ed [available online at http://www.iczn.org/iczn/index.jsp] |
| [30] | Jenks S, Werdelin L, 1998. Chapter 2:taxonomy and systematics of living hyaenas (Family Hyaenidae). In: Mills G, Hofer H eds. IUCN, Hyaenas: Status Survey and Conservation Action Plan. Oxford: Information Press. 8-17 |
| [31] | Jin Z D, Li F C, Cao J J, et al. 2006. Geochemistry of Daihai Lake sediments, Inner Mongolia, north China: implications for provenance, sedimentary sorting, and catchment weathering. Geomorphology, 80: 147-163 |
| [32] | Joeckel R M, 1998. Unique frontal sinuses in fossil and living Hyaenidae (Mammalia, Carnivora): description and interpretation. J Vert Paleont, 18: 627-639 |
| [33] | Koehler C E, Richardson P R K, 1990. Proteles cristatus. Mamm Species, 363: 1-6 |
| [34] | Koepfli K-P, Jenks S M, Eizirik E, et al. 2006. Molecular systematics of the Hyaenidae: relationships of a relictual lineage resolved by a molecular supermatrix. Mol Phylogenet Evol, 38: 603-620 |
| [35] | Kretzoi M, 1938. Die Raubtiere von Gombaszög nebst einer übersicht der Gesamtfauna (Ein beitrag zur stratigraphie des Altquartaers). Ann Mus Natl Hung, 31: 88-157 |
| [36] | Kretzoi M, 1945. Bemerkungen über das Raubtiersystem. Ann Mus Natl Hung, 38: 59-83 |
| [37] | Kruuk H, Sands W A, 1972. The aardwolf (Proteles cristata Sparrman) 1783 as predator of termites. Afr J Ecol, 10: 211-227 |
| [38] | Kurtén B, 1976. Fossil Carnivora from the Late Tertiary of Bled Douarah and Cherichira, Tunisia. Notes Serv Geol, 42: 177-214 |
| [39] | Merriam J C, Stock C, 1932. The Felidae of Rancho La Brea. Carnegie Inst Wash Publ, 422: 1-436 |
| [40] | Pei W C, 1934. On the Carnivora from locality 1 of Choukoutien. Palaeont Sin Ser C, 8: 1-217 |
| [41] | Pilgrim G E, Hopwood A T, 1931. Catalogue of the Pontian Carnivora of Europe in the Department of Geology. London:Printed by order of the Trustees of the British Museum. 1-174 |
| [42] | Qiu Z X, Ye J, Cao J X, 1988. A new species of Percrocuta from Tongxin, Ningxia. Vert PalAsiat, 26: 116-127 |
| [43] | Qiu Z X, Deng T, Wang B Y, 2004. Early Pleistocene mammalian fauna from Longdan, Dongxiang, Gansu, China. Palaeont Sin, New Ser C, 27: 1-198 |
| [44] | Radinsky L, 1975. Viverrid neuroanatomy: phylogenetic and behavioral implications. J Mammal, 56: 130-150 |
| [45] | Radinsky L B, 1982. Evolution of skull shape in carnivores. 3. The origin and early radiation of the modern carnivore families. Paleobiology, 8: 177-195 |
| [46] | Rao H Y, Yang Y M, Liu J Y, et al. 2020. Palaeoproteomic analysis of Pleistocene cave hyenas from east Asia. Sci Rep, 10: 16674 |
| [47] | Schmidt-Kittler N, 1976. Raubtiere aus dem Jungtertiär Kleinasiens. Palaeontogr Abt A, 155: 1-131 |
| [48] | Sheng G L, Soubrier J, Liu J Y, et al. 2014. Pleistocene Chinese cave hyenas and the recent Eurasian history of the spotted hyena, Crocuta crocuta. Mol Ecol, 23: 522-533 |
| [49] | Solounias N, 1981. The Turolian Fauna from the Island of Samos, Greece. Basel: S. Karger AG. 1-232 |
| [50] | Thenius E, 1966. Zur stammesgeschichte der hyanen (Carnivora, Mammalia). Z Saugertierkd, 31: 293-300 |
| [51] | Torre D, 1989. Plioviverrops faventinus n. sp., a new carnivore of late Messinian age. Boll Della Soc Paleont Ital, 28: 323-327 |
| [52] | Turner A, 1993. New fossil carnivore remains. In: Brain C K ed. Swartkrans: a Cave’s Chronicle of Early Man. Transvaal Mus Monogr, 8: 151-165 |
| [53] | Turner A, 1997. New fossil carnivore remains from Sterkfontein hominid site (Mammalia: Carnivora). Ann Transvaal Mus, 34: 319-347 |
| [54] | Turner A, Antón M, Werdelin L, 2008. Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe. Geobios, 41: 677-687 |
| [55] | Villalta Comella J F de, Crusafont-Pairó M, 1943. Los vertebrados del mioceno continental de la cuenca Valles-Panades (provincia de Barcelona), I, Insectivoros; II, Carnivoros. Bol Inst Geol Min Esp, 56: 145-336 |
| [56] | Villalta Comella J F de, Crusafont-Pairó M, 1945. Nuevas aportaciones al conocimiento de los carnivoros pontienses del Valles-Penedes. Barcelona: Inst Geolo?gico, Pub VII (Misc Almer Part 1). 81-121 |
| [57] | Viret J, 1951. Catalogue critique de la faune des mammiferes miocenes de la Grive Saint-Alban (Isere); premiere partie, chiropteres, carnivores, edentes pholidotes. Publ Mus Confluences, 3: 1-104 |
| [58] | Wang X M, 2004. New materials of Tungurictis (Hyaenidae, Carnivora) from Tunggur Formation, Nei Mongol. Vert PalAsiat, 42: 144-153 |
| [59] | Wang X M, Flynn L J, Fortelius M, 2013. Toward a continental Asian biostratigraphic and geochronologic framework. In: Wang X M, Flynn L J, Fortelius M eds. Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York: Columbia University Press. 1-25 |
| [60] | Wang X M, Tseng Z J, Wu W Y, et al. 2020. A new species of Tungurictis Colbert, 1939 (Carnivora, Hyaenidae) from the Middle Miocene of Junggar Basin, northwestern China and the early divergence of basal hyaenids in East Asia. Geodiversitas, 42: 29-45 |
| [61] | Werdelin L, 1988. Studies of fossil hyaenas: the genera Thalassictis Gervais ex Nordmann, Palhyaena Gervais, Hyaenictitherium Kretzoi, Lycyaena Hensel and Palinhyaena Qiu, Huang & Guo. Zool J Linn Soc, 92: 211-265 |
| [62] | Werdelin L, Solounias N, 1991. The Hyaenidae: taxonomy, systematics and evolution. Fossils Strata, 30: 1-104 |
| [63] | Westbury M V, De Cahsan B, Dalerum F, et al. 2019. Aardwolf population diversity and phylogenetic positioning inferred using complete mitochondrial genomes. Afr J Wildl Res, 49: 27-33 |
| [64] | Westbury M V, Le Duc D, Duchêne D A, et al. 2021. Ecological specialization and evolutionary reticulation in extant Hyaenidae. Mol Biol Evol, 1-14, doi: 10.1093/molbev/msab055 |
| [65] | Winge H, Deichmann E, Allen G M, et al. 1941. The interrelationships of the mammalian genera. Vol. 2, Rodentia, Carnivora, Primates. Copenhagen: C.A. Reitzel. 1-418 |
/
| 〈 |
|
〉 |