{{Short description|Extinct species of mammal}} {{Speciesbox | fossil_range = Middle Pleistocene - Holocene {{fossilrange|0.6|0.0025}} | image = Drawing of a hydruntine.jpg | image_caption = Life restoration | extinct = 500 BC | genus = Equus | parent = Asinus | species = hydruntinus | authority = Regalia, 1907 }}
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).
== Description == left|thumb|Complete skull from Crimea 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.<ref name="BPEE" />
== Evolutionary history == ''Equus 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.<ref name="BPEE">{{Cite journal |title=Biostratigraphy and Palaeoecology of European Equus |journal=Frontiers in Ecology and Evolution |last1=Boulbes |first1=Nicolas |date=2019 |volume=7 |article-number=301 |last2=van Asperen |first2=Eline N. |doi=10.3389/fevo.2019.00301 |bibcode=2019FrEEv...7..301B |issn=2296-701X |doi-access=free}}</ref>
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.<ref name=":1">{{Cite journal |last1=Crees |first1=Jennifer J. |last2=Turvey |first2=Samuel T. |date=May 2014 |title=Holocene extinction dynamics of Equus hydruntinus, a late-surviving European megafaunal mammal |journal=Quaternary Science Reviews |volume=91 |pages=16–29 |bibcode=2014QSRv...91...16C |doi=10.1016/j.quascirev.2014.03.003 |issn=0277-3791}}</ref>
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 unambiguous 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).<ref>{{Cite journal |last1=Bernáldez-Sánchez |first1=Eloísa |last2=García-Viñas |first2=Esteban |last3=Sanguino |first3=Fernando |last4=Villalón |first4=David |last5=Leonard |first5=Jennifer A. |date=February 2024 |title=Equids (Equus sp.) in southern Spain from the Palaeolithic to the Bronze Age |journal=Journal of Quaternary Science |language=en |volume=39 |issue=2 |pages=261–276 |bibcode=2024JQS....39..261B |doi=10.1002/jqs.3580 |issn=0267-8179 |doi-access=free |hdl-access=free |hdl=10261/362568}}</ref> Suggestions that the hydruntine survived into the historical period to be described in written records as the ''zebro'', while popular "despite a total dearth of osteological finds ... during the Holocene", are unfounded.<ref>{{cite journal |last1=Nores |first1=Carlos |last2=Muñiz |first2=Arturo Morales |last3=Rodríguez |first3=Laura Llorente |last4=Bennett |first4=Andrew |last5=Geigl |first5=Eva-María |title=The Iberian zebro: what kind of a beast was it? |journal=Anthropozoologica |date=26 June 2015 |volume=50 |issue=1 |pages=21–32 |doi=10.5252/az2015n1a2 |url=https://zenodo.org/records/4313384 |access-date=20 March 2025|hdl=10651/34413 |hdl-access=free}}</ref>
It likely survived latest in West Asia, with reported dates in that region ranging until 1500–500 BC.<ref name=":1" />
The exact systematic position was formerly unclear but recent genetic and morphological analysis suggested that it is closely related to the Asiatic wild ass.<ref>{{cite journal |last1=Burke |first1=Ariane |last2=Eisenmann |first2=Vera |last3=Ambler |first3=Graeme K. |date=May 2003 |title=The systematic position of ''Equus hydruntinus'', an extinct species of Pleistocene equid |journal=Quaternary Research |volume=59 |issue=3 |pages=459–469 |doi=10.1016/S0033-5894(03)00059-0 |bibcode=2003QuRes..59..459B |s2cid=84330920}}</ref><ref>{{cite journal |last1=Orlando |first1=Ludovic |last2=Machkour |first2=Maryam |last3=Burke |first3=Ariane |date=July 2006 |title= Geographic distribution of an extinct equid (''Equus hydruntinus'': Mammalia, Equidae) revealed by morphological and genetical analyses of fossils |journal=Molecular Ecology |volume=15 |issue=8 |pages=2083–2093 |doi=10.1111/j.1365-294X.2006.02922.x |pmid=16780426 |bibcode=2006MolEc..15.2083O |s2cid=9698728}}</ref> 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.<ref name=":0">{{Cite journal |last1=Bennett |first1=E. Andrew |last2=Champlot |first2=Sophie |last3=Peters |first3=Joris |last4=Arbuckle |first4=Benjamin S. |last5=Guimaraes |first5=Silvia |last6=Pruvost |first6=Mélanie |last7=Bar-David |first7=Shirli |last8=Davis |first8=Simon J. M. |last9=Gautier |first9=Mathieu |last10=Kaczensky |first10=Petra |last11=Kuehn |first11=Ralph |display-authors=10 |date=2017-04-19 |editor-last=Janke |editor-first=Axel |title=Taming the late Quaternary phylogeography of the Eurasiatic wild ass through ancient and modern DNA |journal=PLOS ONE |language=en |volume=12 |issue=4 |article-number=e0174216 |doi=10.1371/journal.pone.0174216 |issn=1932-6203 |pmc=5396879 |pmid=28422966 |bibcode=2017PLoSO..1274216B |doi-access=free}}</ref> 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,<ref name=":4">{{Cite journal |last1=Özkan |first1=Mustafa |last2=Gürün |first2=Kanat |last3=Yüncü |first3=Eren |last4=Vural |first4=Kıvılcım Başak |last5=Atağ |first5=Gözde |last6=Akbaba |first6=Ali |last7=Fidan |first7=Fatma Rabia |last8=Sağlıcan |first8=Ekin |last9=Altınışık |first9=Ezgi N. |last10=Koptekin |first10=Dilek |last11=Pawłowska |first11=Kamilla |last12=Hodder |first12=Ian |last13=Adcock |first13=Sarah E. |last14=Arbuckle |first14=Benjamin S. |last15=Steadman |first15=Sharon R. |date=July 2024 |title=The first complete genome of the extinct European wild ass ( Equus hemionus hydruntinus ) |journal=Molecular Ecology |language=en |volume=33 |issue=14 |article-number=e17440 |doi=10.1111/mec.17440 |issn=0962-1083 |doi-access=free |pmid=38946459 |bibcode=2024MolEc..33E7440O |hdl=11511/110286 |hdl-access=free}}</ref> with analysis of a genome of Pleistocene specimens from Eastern Europe also suggesting gene flow with Asiatic wild asses in Western Russia.<ref>{{Cite journal |last1=Pan |first1=Jianfei |last2=Liu |first2=Xuexue |last3=Baca |first3=Mateusz |last4=Calvière-Tonasso |first4=Laure |last5=Schiavinato |first5=Stéphanie |last6=Chauvey |first6=Loreleï |last7=Tressières |first7=Gaétan |last8=Perdereau |first8=Aude |last9=Aury |first9=Jean-Marc |last10=Oliveira |first10=Pedro H. |last11=Wincker |first11=Patrick |last12=Abdykanova |first12=Aida |last13=Arsuaga |first13=Juan Luis |last14=Bayarsaikhan |first14=Jamsranjav |last15=Belinskiy |first15=Andrey B. |date=October 2024 |title=Genome-wide population affinities and signatures of adaptation in hydruntines, sussemiones and Asian wild asses |journal=Molecular Ecology |language=en |volume=33 |issue=19 |article-number=e17527 |doi=10.1111/mec.17527 |pmid=39279684 |bibcode=2024MolEc..33E7527P |issn=0962-1083 |doi-access=free}}</ref>
Cladogram based on whole nuclear genomes after Özkan et al. 2024.<ref name=":4" />
{{clade |label1=''Asinus'' (asses) |1={{clade |1=''Equus africanus'' + ''Equus asinus'' African wild ass + domestic donkeys |2={{clade |1={{extinct}}'''''Equus hydruntinus''''' (European wild ass/hydruntine) |2={{clade |1={{clade |1={{extinct}}''Equus hemionus hemippus'' (Syrian wild ass) |2=''Equus hemionus onager'' (Persian onager) }} |2={{clade |1=''Equus hemionus hemionus'' (Mongolian wild ass) + ''Equus (hemionus) kiang'' (kiang) }}}}}}}}}}
== Ecology == The 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.<ref name="BPEE" /> It may have retreated to warmer locales during the coldest periods<ref name=":0" /> 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.<ref name="BPEE" /> 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.<ref>{{Cite journal |title=The Ecological Structure of the 'Mammoth Fauna' in Eurasia. |journal=Annales Zoologici Fennici |last1=Vereshchagin |first1=N. K. |date=1991 |volume=28 |pages=253–259 |last2=Baryshnikov |first2=G. F. |issue=3/4 |jstor=23735450}}</ref> 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.<ref>{{Cite journal |last1=Strani |first1=Flavia |last2=DeMiguel |first2=Daniel |date=June 2023 |title=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 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=620 |article-number=111564 |doi=10.1016/j.palaeo.2023.111564 |doi-access=free |bibcode=2023PPP...62011564S |hdl=11573/1678265 |hdl-access=free}}</ref><ref name=":3">{{Cite journal |last1=Sánchez-Hernández |first1=Carlos |last2=Gourichon |first2=Lionel |last3=Soler |first3=Joaquim |last4=Soler |first4=Narcís |last5=Blasco |first5=Ruth |last6=Rosell |first6=Jordi |last7=Rivals |first7=Florent |date=August 2020 |title=Dietary traits of ungulates in northeastern Iberian Peninsula: Did these Neanderthal preys show adaptive behaviour to local habitats during the Middle Palaeolithic? |journal=Quaternary International |language=en |volume=557 |pages=47–62 |doi=10.1016/j.quaint.2020.01.008 |doi-access=free |bibcode=2020QuInt.557...47S}}</ref> Dental microwear evidence from Late Pleistocene specimens from the Crimean Peninsula likewise reveal it had a diet mainly composed of abrasive grasses.<ref>{{Cite journal |last1=Ramírez-Pedraza |first1=Iván |last2=Rivals |first2=Florent |last3=Uthmeier |first3=Thorsten |last4=Chabai |first4=Victor |date=27 October 2020 |title=Palaeoenvironmental and seasonal context of the Late Middle and Early Upper Palaeolithic occupations in Crimea: an approach using dental wear patterns in ungulates |url=https://link.springer.com/article/10.1007/s12520-020-01217-9 |journal=Archaeological and Anthropological Sciences |language=en |volume=12 |issue=11 |article-number=268 |doi=10.1007/s12520-020-01217-9 |bibcode=2020ArAnS..12..268R |issn=1866-9557 |access-date=29 December 2024 |via=Springer Nature Link |url-access=subscription}}</ref> Remains of European wild ass have been found in cave hyena dens, suggesting that they were likely predated upon by them.<ref name="diedrich&zak">Diedrich, C.G. & ŽÁK, K. 2006. [http://www.geology.cz/bulletin/fulltext/bullgeosci200604237.pdf Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic)]. ''Bulletin of Geosciences'' 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.</ref>
== Relationship with humans == The hydruntine is depicted in Palaeolithic cave paintings and engravings from France, as well as on Neolithic pottery from Anatolia.<ref name=":0" /> 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.<ref name=":2">{{Cite journal |last1=Özkan |first1=Mustafa |last2=Gürün |first2=Kanat |biorxiv=10.1101/2023.06.05.543765 |language=en |last3=Yüncü |first3=Eren |last4=Vural |first4=Kıvılcım Başak |last5=Atağ |first5=Gözde |last6=Akbaba |first6=Ali |last7=Fidan |first7=Fatma Rabia |last8=Sağlıcan |first8=Ekin |last9=Altınışık |first9=N. Ezgi |title=The first complete genome of the extinct European wild ass ( ''Equus hemionus hydruntinus'' ) |journal=Molecular Ecology |date=2024 |volume=33 |issue=14 |article-number=e17440 |doi=10.1111/mec.17440 |bibcode=2024MolEc..33E7440O |hdl=11511/110286 |hdl-access=free}}</ref><ref name=":3" /><ref>Cassoli, P. F., Fiore, I. & Tagliacozzo, A. Butchering and exploitation of large mammals in the Epigravettian levels of Grotta Romanelli (Apulia, Italy). ''Anthropozoologica''25–26, 309–318 (1997).</ref><ref>{{Cite journal |last=Bataille |first=Guido |date=April 2017 |title=Neanderthals of Crimea – Creative generalists of the late Middle Paleolithic. Contextualizing the leaf point industry Buran-Kaya III, Level C |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215014664 |journal=Quaternary International |language=en |volume=435 |pages=211–236 |doi=10.1016/j.quaint.2015.12.071|bibcode=2017QuInt.435..211B |url-access=subscription}}</ref><ref>{{cite book |last1=Anthony |first1=David W. |title=The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World |date=26 July 2010 |publisher=Princeton University Press |location=Princeton, NJ |isbn=978-1-4008-3110-4 |pages=136, 148, 198–199 |url=https://books.google.com/books?id=0FDqf415wqgC&pg=PA148 |access-date=20 March 2025 |language=en}}</ref>
{{gallery|Panel of the hemione.jpg|Cave painting from Lascaux cave, France|{{Not a typo|Engraving of a presumed hydruntine in the cave “Les Trois Frères”.jpg}}<!-- Do not change the spelling of the file! -->|Engraving in the cave Les Trois Frères, France|Engraving of a presumed hydruntine on a pendant.jpg|Engraving on a pendant from Putois cave, France|Vessel with a hunt scene of presumable hydruntines.jpg|Hunting scene on Neolithic pottery from Turkey|||||||||width=200|height=180|lines=|align=center}}
==See also== * List of extinct animals of Europe
==References== {{Reflist}}
{{Equus}} {{Taxonbar|from=Q631645}}
Category:Onager Category:Pleistocene mammals of Europe Category:Extinct mammals of Europe Category:Holocene extinctions Category:Middle Pleistocene Category:Late Pleistocene Category:Extinct mammals Category:Mammals described in 1907 Category:Çatalhöyük