The European wild ass (Equus hydruntinus or Equus hemionus hydruntinus) or hydruntine is an extinct equine from the Middle Pleistocene to Late Holocene of Europe and West Asia, and possibly North Africa. It is a member of the subgenus Asinus, and closely related to the living Asiatic wild ass. The specific epithet, hydruntinus, means from Otranto (Hydruntum in Latin).
| European wild ass Temporal range: Middle Pleistocene - Holocene
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| Life restoration | |
Scientific classification 
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Perissodactyla |
| Family: | Equidae |
| Genus: | Equus |
| Subgenus: | Asinus |
| Species: | †E. hydruntinus
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| Binomial name | |
| †Equus hydruntinus Regalia, 1907
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Description
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In comparison to the Asiatic wild ass (Equus hemionus), the muzzle region of the skull is much shorter and somewhat proportionally wider, the palate is elongate, and the nasal notch is shorter. The teeth are relatively small compared to skull size, but are very hypsodont (high crowned). The shafts of the metacarpal and metatarsal bones are also more robust.[1]
Evolutionary history
editEquus hydruntinus appeared first in the fossil record around 600,000 years ago during the Middle Pleistocene. In the Late Pleistocene it was widespread throughout much of western Eurasia from the Middle East to Europe, especially along the Mediterranean, with fossil reports from Italy, Turkey, Spain, France and Portugal. In the east the range apparently stretched at least to the Volga and to Iran. In the north it reached almost to the North Sea in Germany and the British Isles. Some authors suggest that it may have reached North Africa.[1] While it was historically suggested that the species survived in the southern Iberian Peninsula into the Chalcolithic based on a phalange found at Cabezo Juré in Huelva, Spain, later research found it was more likely that the phalange came from a mule, and thus there are no unambigious specimens of E. hydruntinus in the Iberian Peninsula that date later than the end of the Late Pleistocene (also corresponding to the end of the Paleolithic).[2] Its range fragmented after the Last Glacial Maximum, surviving into the Holocene, its range contracted further, persisting in small regions of southern Europe, including the Danube river valley, the southern Italian Peninsula and southern France, probably surviving latest in Europe around the Danube until around 4000-3000 BC.[3] It likely survived later in West Asia, with reported dates in that region ranging until 1500–500 BC.[3]
The exact systematic position was formerly unclear but recent genetic and morphological analysis suggested that it is closely related to the Asiatic wild ass.[4][5] A 2017 genetic study based on a partial mitochondrial genome suggested that it was a subspecies of Asiatic wild ass, closer to the Khur than the Persian onager.[6] However, study of the full mitochondral and nuclear genomes of specimens from Çatalhöyük and Çadır Höyük in Anatolia (present day Turkey) dating to the early-mid 1st millennium BC, which represent the youngest known remains of the species (with the youngest specimen dated to around 2698–2356 cal years Before Present, or around 748–406 cal years BC) , suggest that all modern Asiatic wild ass lineages (sensu lato, including the kiang) are more closely related to each other than to E. hydruntinus, with the split between hydruntines and Asiatic wild asses estimated at around 800-600 thousand years ago. Analysis of the nuclear genome suggested that there had been gene flow during the Holocene from the hydruntine lineage into Middle Eastern Asiatic wild asses,[7] with analysis of a genome of Pleistocene specimens from Eastern Europe also suggesting gene flow with Asiatic wild asses in Western Russia.[8]
Cladogram based on whole nuclear genomes after Özkan et al. 2024.[7]
| Asinus (asses) |
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Ecology
editThe evidence shows that the European ass favoured semi-arid, steppic conditions and showed a preference for temperate climates, although it was also found in cool or cold conditions.[1] It may have retreated to warmer locales during the coldest periods[6] although the relatively short muzzle indicates an adaptation for cold conditions. It preferred open biotopes, between shrubland (favoured by true horses) and grassland (favoured by bovids). It is believed to have shared this habitat with species such as the woolly rhinoceros.[1] It is considered an ecologically important part of the ecosystem known as mammoth steppe where it filled a niche equivalent to that provided by the African wild ass or Zebra in the African savanna.[9] Dental wear analysis of specimens from the Iberian Peninsula suggests a primarily grazing diet, though they appear to have been flexible feeders, having seasonally consumed browse.[10][11]
Relationship with humans
editThe hydruntine is depicted in Palaeolithic cave paintings and engravings from France, as well as on Neolithic pottery from Anatolia.[6] Remains found with cut marks and/or in archaeological sites spanning from the Paleolithic to the youngest known remains of the species in the Iron Age across the species range, including Crimea, Italy, the Iberian Peninsula and Anatolia indicated that it was hunted by people, including both modern humans and Neanderthals.[12][11][13][14]
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Cave painting from Lascaux cave, France -
Engraving in the cave Les Trois Frères, France -
Engraving on a pendant from Putois cave, France -
Hunting scene on Neolithic pottery from Turkey
See also
editReferences
edit- ^ a b c d Boulbes, Nicolas; van Asperen, Eline N. (2019). "Biostratigraphy and Palaeoecology of European Equus". Frontiers in Ecology and Evolution. 7: 301. doi:10.3389/fevo.2019.00301. ISSN 2296-701X.
- ^ Bernáldez-Sánchez, Eloísa; García-Viñas, Esteban; Sanguino, Fernando; Villalón, David; Leonard, Jennifer A. (February 2024). "Equids (Equus sp.) in southern Spain from the Palaeolithic to the Bronze Age". Journal of Quaternary Science. 39 (2): 261–276. Bibcode:2024JQS....39..261B. doi:10.1002/jqs.3580. hdl:10261/362568. ISSN 0267-8179.
- ^ a b Crees, Jennifer J.; Turvey, Samuel T. (May 2014). "Holocene extinction dynamics of Equus hydruntinus, a late-surviving European megafaunal mammal". Quaternary Science Reviews. 91: 16–29. Bibcode:2014QSRv...91...16C. doi:10.1016/j.quascirev.2014.03.003. ISSN 0277-3791.
- ^ Burke, Ariane; Eisenmann, Vera; Ambler, Graeme K. (May 2003). "The systematic position of Equus hydruntinus, an extinct species of Pleistocene equid". Quaternary Research. 59 (3): 459–469. Bibcode:2003QuRes..59..459B. doi:10.1016/S0033-5894(03)00059-0. S2CID 84330920.
- ^ Orlando, Ludovic; Machkour, Maryam; Burke, Ariane (July 2006). "Geographic distribution of an extinct equid (Equus hydruntinus: Mammalia, Equidae) revealed by morphological and genetical analyses of fossils". Molecular Ecology. 15 (8): 2083–2093. Bibcode:2006MolEc..15.2083O. doi:10.1111/j.1365-294X.2006.02922.x. PMID 16780426. S2CID 9698728.
- ^ a b c Bennett, E. Andrew; Champlot, Sophie; Peters, Joris; Arbuckle, Benjamin S.; Guimaraes, Silvia; Pruvost, Mélanie; Bar-David, Shirli; Davis, Simon J. M.; Gautier, Mathieu; Kaczensky, Petra; et al. (2017-04-19). Janke, Axel (ed.). "Taming the late Quaternary phylogeography of the Eurasiatic wild ass through ancient and modern DNA". PLOS ONE. 12 (4): e0174216. Bibcode:2017PLoSO..1274216B. doi:10.1371/journal.pone.0174216. ISSN 1932-6203. PMC 5396879. PMID 28422966.
- ^ a b Özkan, Mustafa; Gürün, Kanat; Yüncü, Eren; Vural, Kıvılcım Başak; Atağ, Gözde; Akbaba, Ali; Fidan, Fatma Rabia; Sağlıcan, Ekin; Altınışık, Ezgi N.; Koptekin, Dilek; Pawłowska, Kamilla; Hodder, Ian; Adcock, Sarah E.; Arbuckle, Benjamin S.; Steadman, Sharon R. (July 2024). "The first complete genome of the extinct European wild ass ( Equus hemionus hydruntinus )". Molecular Ecology. 33 (14): e17440. Bibcode:2024MolEc..33E7440O. doi:10.1111/mec.17440. ISSN 0962-1083. PMID 38946459.
- ^ Pan, Jianfei; Liu, Xuexue; Baca, Mateusz; Calvière-Tonasso, Laure; Schiavinato, Stéphanie; Chauvey, Loreleï; Tressières, Gaétan; Perdereau, Aude; Aury, Jean-Marc; Oliveira, Pedro H.; Wincker, Patrick; Abdykanova, Aida; Arsuaga, Juan Luis; Bayarsaikhan, Jamsranjav; Belinskiy, Andrey B. (October 2024). "Genome-wide population affinities and signatures of adaptation in hydruntines, sussemiones and Asian wild asses". Molecular Ecology. 33 (19). Bibcode:2024MolEc..33E7527P. doi:10.1111/mec.17527. ISSN 0962-1083.
- ^ Vereshchagin, N. K.; Baryshnikov, G. F. (1991). "The Ecological Structure of the 'Mammoth Fauna' in Eurasia". Annales Zoologici Fennici. 28 (3/4): 253–259. JSTOR 23735450.
- ^ Strani, Flavia; DeMiguel, Daniel (June 2023). "The role of climate change in the extinction of the last wild equids of Europe: Palaeoecology of Equus ferus and Equus hydruntinus during the Last Glacial Period". Palaeogeography, Palaeoclimatology, Palaeoecology. 620: 111564. Bibcode:2023PPP...62011564S. doi:10.1016/j.palaeo.2023.111564. hdl:11573/1678265.
- ^ a b Sánchez-Hernández, Carlos; Gourichon, Lionel; Soler, Joaquim; Soler, Narcís; Blasco, Ruth; Rosell, Jordi; Rivals, Florent (August 2020). "Dietary traits of ungulates in northeastern Iberian Peninsula: Did these Neanderthal preys show adaptive behaviour to local habitats during the Middle Palaeolithic?". Quaternary International. 557: 47–62. Bibcode:2020QuInt.557...47S. doi:10.1016/j.quaint.2020.01.008.
- ^ Özkan, Mustafa; Gürün, Kanat; Yüncü, Eren; Vural, Kıvılcım Başak; Atağ, Gözde; Akbaba, Ali; Fidan, Fatma Rabia; Sağlıcan, Ekin; Altınışık, N. Ezgi (2023-06-07). The first complete genome of the extinct European wild ass ( Equus hemionus hydruntinus ) (Report). Genomics. doi:10.1101/2023.06.05.543765.
- ^ Cassoli, P. F., Fiore, I. & Tagliacozzo, A. Butchering and exploitation of large mammals in the Epigravettian levels of Grotta Romanelli (Apulia, Italy). Anthropozoologica25–26, 309–318 (1997).
- ^ Bataille, Guido (April 2017). "Neanderthals of Crimea – Creative generalists of the late Middle Paleolithic. Contextualizing the leaf point industry Buran-Kaya III, Level C". Quaternary International. 435: 211–236. Bibcode:2017QuInt.435..211B. doi:10.1016/j.quaint.2015.12.071.