Tolai hare

{{Short description|Species of mammal}} {{good article}} {{Speciesbox | image = Толай.jpg | image_alt = A hare sitting among brambles | image_caption = Altyn-Emel National Park, Kazakhstan | status = LC | status_system = IUCN3.1 | status_ref = <ref name="iucn status 19 November 2021">{{cite iucn |author=Smith, A.T. |author2=Johnston, C.H. |date=2019 |title=''Lepus tolai'' |volume=2019 |article-number=e.T41308A45193447 |doi=10.2305/IUCN.UK.2019-1.RLTS.T41308A45193447.en |access-date=19 November 2021}}</ref> | genus = Lepus | species = tolai | authority = Pallas, 1778 | range_map = Tolai Hare area-edit.png | range_map_caption = Tolai hare range | synonyms_ref = <ref name=mdd>{{Cite mdd|access-date=6 February 2026|id=1001107 |title=''Lepus tolai'' Pallas, 1778 }}</ref> | synonyms = {{Collapsible list| *''Lepus aralensis'' Severtzov, 1861 *''Lepus lehmanni'' Severtzov, 1873 *''Lepus butlerowi'' Bogdanov, 1882 *''Lepus kessleri'' Bogdanov, 1882 *''Lepus swinhoei'' O. Thomas, 1894 *''Lepus centrasiaticus'' Satunin, 1907 *''Lepus gansuicus'' Satunin, 1907 *''Lepus gobicus'' Satunin, 1907 *''Lepus filchneri'' Matschie, 1908 *''Lepus stegmanni'' Matschie, 1908 *''Lepus swinhoei subluteus'' O. Thomas, 1908 *''Lepus swinhoei brevinasus'' J. A. Allen, 1909 *''Lepus aurigineus'' Hollister, 1912 *''Lepus swinhoei sowerbyae'' Hollister, 1912 *''Lepus tolai bucharensis'' Ognev, 1922 *''Lepus wongi'' Yang Zhongjian, 1927 *''Lepus tolai desertorum'' Ognev & Heptner, 1928 *''Lepus europaeus turcomanus'' Heptner, 1934 *''Lepus europaeus cinnamomeus'' Shamel, 1940 *''Lepus capensis cheybani'' Baloutch, 1979 *''Lepus capensis habibi'' Baloutch, 1979 *''Lepus capensis petteri'' Baloutch, 1979 *''Lepus capensis huangshuiensis'' Luo Zexun, 1982 }} }}

The '''tolai hare''' ('''''Lepus tolai''''') is a species of hare native to Central Asia, including much of Mongolia, eastern Iran, Afghanistan, southern Kazakhstan, Turkmenistan, Uzbekistan, Kyrgyzstan, and North and Central China. It inhabits semi-desert, steppes, rocky habitats, grasslands, and river valleys. A yellow, brown or grey-furred hare with long, black tipped ears, the adult tolai hare weighs between {{Convert|1.7 and 2.7|kg|abbr=off}} and measures between {{convert|40|and|59|cm|in|0|abbr=off}}. It is mainly active at dusk and night but is occasionally active during the day to forage for seeds, roots, and other plant matter. Young hares are often more active in daylight hours.

A widespread, fairly common species, the tolai hare has a stable population. It breeds often, two to three times per year, and is hunted for its meat, fur, and for use in traditional medicine. Hustai National Park is one of several protected areas where the hare can be found. Authorities in China and Mongolia, as well as the International Union for Conservation of Nature, consider it a least-concern species.

==Taxonomy and etymology== thumb|left|alt=Drawing of a brown hare on a rock|Engraving of the tolai hare by Johann Nufsbiegel The tolai hare was first described by the German naturalist Peter Simon Pallas in 1778. He gave it the name {{gloss|tolai}} as this was the common name for {{Gloss|hare}} among the Mongols and Kalmyks.<ref>{{Cite book |last1=Pallas |first1=P.S. |author-link1=Peter Simon Pallas |url=https://www.biodiversitylibrary.org/page/15494826#page/33/mode/1up |title=Novae Species Quadrupedum E Glirium Ordine Cum Illustrationibus Variis Complurium Ex Hoc Ordine Animalium |date=1778 |publisher=Wolfgang Walther, Erlangen |volume=Fasciculus I |pages=17-28 |via=Biodiversity Heritage Library |lang=la}}</ref> This word is attested to in later works as {{Lang|mn|tōlai}},<ref>{{Cite book |last=Brill |first=E. J. |url=https://books.google.com/books?id=0ScVAAAAIAAJ |title=Histoire des Campagnes de Gengis Khan |date=1951 |publisher=Leiden |volume=1 |page=330 |language=fr |quote=le lièvre étant appelé ''tōlai'' en mongol}}</ref> ''dolai'' or ''taulai'' in the Mongolian language.<ref>{{Cite journal |last=Yang |first=Lien-sheng |date=1955 |title=Notes on Maspero's "Les documents chinois de la troisième expédition de Sir Aurel Stein en Asie centrale" |url=https://www.jstor.org/stable/2718415 |journal=Harvard Journal of Asiatic Studies |volume=18 |issue=1/2 |pages=142–158 |doi=10.2307/2718415 |issn=0073-0548 |url-access=subscription}}</ref> Its type locality was described as a rocky or sandy plain near the Selenga river; Ellerman and Morrison-Scott wrote in 1951 that a more likely description of the locality was at Adinscholo Mountain on the river Borzya, which is some {{Convert|700|km|abbr=off}} east of the Selenga.<ref>{{MSW3 Lagomorpha|id=13500237|heading=''Lepus (Proeulagus) tolai''}}</ref>

The phylogenetics of the tolai hare and its subspecies have been contested. It was formerly included with the cape hare (''L. capensis''),<ref name="pmid6972951">{{cite journal |last1=Lado |first1=S |last2=Alves |first2=PC |last3=Islam |first3=MZ |last4=Brito |first4=JC |last5=Melo-Ferreira |first5=J |date=November 2019 |title=The evolutionary history of the Cape hare (''Lepus capensis'' sensu lato): insights for systematics and biogeography |journal=Heredity |volume=123 |issue=5 |pages=634–646 |bibcode=2019Hered.123..634L |doi=10.1038/s41437-019-0229-8 |pmc=6972951 |pmid=31073237}}</ref> as well as the European hare (''L. europaeus'') and the desert hare (''L. tibetanus'').<ref name="iucn status 19 November 2021" /> Even after the tolai hare was separated into a distinct species, there remained confusion among its subspecies. The subspecies ''L. tolai centrasiaticus'' has since been reassigned to the desert hare, but genetic analysis has made the position of ''L. t. centrasiaticus'' less certain, indicating that it may be closer to the tolai hare than the desert hare. Furthermore, the subspecies ''L. tolai swinhoei'' has been proposed as representing a distinct species, and ''L. przewalskii'' has been reassigned as a synonym of the woolly hare (''L. oiostolus'').<ref name=Smith2018>{{Lagomorphs2018|last=Smith|first=Andrew T.|chapter=''Lepus tolai'' Pallas, 1778 Tolai hare|pages=216-217}}</ref>

Ten subspecies of the tolai hare are known, with eight accepted in the third edition of Mammal Species of the World and two more (''centrasiaticus'' and ''huangshuiensis'') proposed since its publication:<ref name="iucn status 19 November 2021" /><ref name=Smith2018 /> {{Columns-list|colwidth=30em|1= *''L. t. tolai'', nominate subspecies found in Inner Mongolia and Gansu *''L. t. aurigineus'', found in Jiangxi, Anhui, Hubei, Shaanxi, Sichuan, and Guizhou *''L. t. buchariensis'', unknown range, may extend into Iran and beyond *''L. t. centrasiaticus'', found in Xinjiang, Gansu, and Inner Mongolia *''L. t. cheybani'', unknown range, may extend into Iran and beyond *''L. t. cinnamomeus'', found in Sichuan and Yunnan *''L. t. filchneri'', unknown range, may extend into Iran and beyond *''L. t. huangshuiensis'', found in Qinghai *''L. t. lehmanni'', found in Xinjiang *''L. t. swinhoei'', found in Heilongjiang, Jilin, Liaoning, Inner Mongolia, Hebei, Beijing, Henan, Shaanxi, Shanxi, and Shandong }} According to molecular genetic analysis performed in 2024 by Leandro Iraçabal and colleagues, the sister clade of the tolai hare is a group that includes nine species of widely distributed hares:<ref name="Iraçabal-2024">{{Cite journal |last=Iraçabal |first=Leandro |last2=Barbosa |first2=Matheus R. |last3=Selvatti |first3=Alexandre Pedro |last4=Russo |first4=Claudia Augusta de Moraes |date=2024 |title=Molecular time estimates for the Lagomorpha diversification |journal=PLOS One |volume=19 |issue=9 |article-number=e0307380 |doi=10.1371/journal.pone.0307380 |doi-access=free|issn=1932-6203 |pmc=11379240 |pmid=39241029}}</ref> {{cladogram|title=The tolai hare, its sister clade, and other hares|align=left| {{clade |1={{clade |2={{clade |2={{clade |2={{clade |2={{clade |2={{clade |2=''Lepus castroviejoi'' |1=''Lepus corsicanus'' }} |1=''Lepus arcticus'' }} |1=''Lepus othus'' }} |1={{clade |2=''Lepus timidus'' |1=''Lepus townsendii'' }} }} |1={{clade |2={{clade |2=''Lepus mandshuricus'' |1=''Lepus coreanus'' }} |1=''Lepus capensis'' }} }} |1='''''Lepus tolai''''' }}|2=Other hares}}}}{{clear}}

==Description== [[File:Lepus tolai 68614347 (cropped).jpg|thumb|left|A tolai hare seen in the Altai Republic, Russia]] The tolai hare grows to a head-and-body length of {{convert|40|to|59|cm|in|0|abbr=off}} with a tail of {{convert|72|to|110|mm|in|1|abbr=out}}. The ears range from {{Convert|80 to 120|mm|abbr=on}} in length, and the hind feet measure {{Convert|110 to 127|mm|abbr=on}}. Adult hares weigh from {{Convert|1.7 to 2.7|kg|abbr=off}}. The tolai hare is variable in colouration across its range. Its upper parts range from dusty yellow to pale brown, sandy yellow or sandy grey and may have dark brownish or reddish stripes.<ref name=":0" /> The hip region is sometimes ochre or grey. The head has a pale, bare, greyish or ochraceous patch of skin surrounding the eye and extending forwards to near the muzzle and backwards to the base of the long ears, which have black tips. The underparts and flanks are pure white. The tail has a broad black or brownish-black stripe on the top.<ref name=Smith/> Like other leporids, it has a dental formula of {{DentalFormula|upper=2.0.3.3|lower=1.0.2.3|total=28}}—two pairs of upper and one pair of lower incisors, no canines, three upper and two lower premolars on each side, and three upper and lower molars on either side of the jaw.<ref>{{Cite web |last=Pintus |first=Kathryn |title=''Lepus tolai'' - Tolai hare |url=http://wildpro.twycrosszoo.org/S/0MLagomorph/Leporidae/lepus/Lepus_tolai.htm |archive-url=https://web.archive.org/web/20161026174012/http://wildpro.twycrosszoo.org/S/0MLagomorph/Leporidae/lepus/Lepus_tolai.htm |archive-date=October 26, 2016 |website=Wildpro |publisher=Twycross Zoo}}</ref>

There are few features that can be used to differentiate the subspecies of the tolai hare, as their original descriptions were based largely on external characteristics and had small sample sizes. The distinctions between them remain unclear due to the species' gradual shifts in colouration across its range.<ref name=":0" />

==Distribution and habitat== [[File:Lepus tolai 529211477.jpg|thumb|alt=A hare with its head to the ground eating grass|Feeding in Rashaant, Bulgan, Mongolia]] The tolai hare is native to central and eastern Asia. Its range extends from the eastern side of the Caspian Sea through eastern Iran, Afghanistan, southern Kazakhstan, Turkmenistan, Uzbekistan and Kyrgyzstan, through southern Siberia and Mongolia to western, central and north-eastern China. Its elevation range is generally between {{cvt|600|and|900|m}}, but a single individual has been recorded much higher, at an elevation of {{Convert|4900|m|abbr=on}}.<ref name="iucn status 19 November 2021" /> One specimen found in Jammu and Kashmir may indicate a southern extension of the species' distribution.<ref name=":0" />

The hare is found across various habitats, from arid sand dunes to river valleys with tall vegetation. The wetter regions are preferred, with populations being described as "abundant" in the Hailar river valley. It avoids steppes where low-lying vegetation grows.<ref name=":1">{{Cite journal |last=Loukashkin |first=Anatole S. |date=1943 |title=On the hares of northern Manchuria |url=https://www.jstor.org/stable/1374782 |journal=Journal of Mammalogy |volume=24 |issue=1 |pages=73–81 |doi=10.2307/1374782 |issn=0022-2372}}</ref> Other habitats with reported tolai hare populations include grasslands, deserts, semi-deserts, rocky habitats, and forest steppes, though it is not found in heavily forested areas.<ref name="Smith2018" /><ref name=":1" />

==Ecology and behaviour== [[File:Lepus tolai young 218774125.jpg|thumb|left|180px|A young tolai hare in Kegeti, Kyrgyzstan]] The tolai hare is a nocturnal species and feeds on grasses, sedges, wood and bark, seeds, bulbs, shoots and roots. It occasionally deviates from its nocturnal habits to forage during daylight; young hares are generally more active during the day.<ref name="Smith2018" /> It does not dig a burrow except when it is breeding, but scrapes out a depression in the ground (known as a form) with its front paws in which to rest during the day.<ref name=":0" /> This scoop is shallow in hot weather but is deeper in colder conditions. Breeding takes place two or three times a year, with litters of two to six young being produced each time.<ref name=Smith>{{cite book|author1=Smith, Andrew T.|author2=Xie, Yan|author3=Hoffmann, Robert S.|author4=Lunde, Darrin|author5=MacKinnon, John|author6=Wilson, Don E.|author7=Wozencraft, W. Chris|title=A Guide to the Mammals of China|url=https://books.google.com/books?id=ka-9f68nPT4C&pg=PA291 |year=2010 |publisher=Princeton University Press |isbn=978-1-4008-3411-2 |pages=291–292}}</ref> Subsequent litters are usually smaller than those that precede it in a season. The breeding season generally starts in late February and extends to March, but may start earlier or last for longer depending on location. In some regions, the breeding season lasts for up to five months. The species has a diploid chromosome number of 48.<ref name="Smith2018" />

Several parasites in the genus ''Eimeria'' are known to use the tolai hare as a host, namely ''E. leporis'' and ''E. gobiensis''.<ref>{{Cite journal |last=Gardner |first=Scott L. |last2=Seggerman |first2=Nathan A. |last3=Batsaikhan |first3=Nyamsuren |last4=Ganzorig |first4=Sumiya |last5=Tinnin |first5=David S. |last6=Duszynski |first6=Donald W. |date=2009 |title=Coccidia (Apicomplexa: Eimeriidae) from the lagomorph ''Lepus tolai'' in Mongolia |url=http://www.bioone.org/doi/abs/10.1645/GE-2137.1 |journal=Journal of Parasitology |language=en |volume=95 |issue=6 |pages=1451–1454 |doi=10.1645/GE-2137.1 |issn=0022-3395 |url-access=subscription}}</ref> 22 species of ticks have been found on tolai hares, including those from the genera ''Dermacentor'' and ''Rhipicephalus''. The spring and summer months typically bring the greatest numbers of mites. Fleas, trematodes, cestodes and nematodes are also found on or in tolai hares. The hares also carry diseases such as tularemia, plague, and the brucellosis pathogen.<ref name="Smith2018" />

==Interaction with humans== [[File:Lepus tolai 197486393.jpg|thumb|alt=A hare running in a row of crops|A tolai hare among crops in Zhongmu County, China]] Tolai hares are hunted for food and furs, though this latter use has declined, with fur harvests decreasing more than tenfold in southern Uzbekistan from 1959 to 1977. The hare is also used in traditional medicine.<ref name=":0">{{HMW Volume 6|chapter=''Lepus tolai'' Pallas, 1778|chapter-url=https://www.gbif.org/species/195832024}}</ref> In eastern Mongolia, a consistent harvest of about 1,000 hares is reported annually.<ref name="Smith2018" /> In the Neolithic Yangjiesha site of Loess Plateau, signs of commensal behavior (taming) between tolai hares and humans have been found.<ref>{{cite journal |last1=Sheng |first1=Pengfei |last2=Hu |first2=Yaowu |last3=Sun |first3=Zhouyong |last4=Yang |first4=Liping |last5=Hu |first5=Songmei |last6=Fuller |first6=Benjamin T. |last7=Shang |first7=Xue |date=June 2020 |title=Early commensal interaction between humans and hares in Neolithic northern China |journal=Antiquity |volume=94 |issue=375 |pages=622–636 |doi=10.15184/aqy.2020.36 |doi-access=free}}</ref>

=== Conservation status === The tolai hare has a wide range, a large population, and is generally a common species. Growing human populations and displacement of hares from their habitat poses a potential threat, though it has not been studied extensively. Protected areas, such as parks like Hustai National Park, cover roughly 12% of the species' range. The International Union for Conservation of Nature has assessed its conservation status as being of least concern, and this same status is used by regional conservation lists in China and Mongolia.<ref name="iucn status 19 November 2021" />

==References== {{Commons category|Lepus tolai}} {{Reflist}}

Category:Hares Category:Mammals of Pakistan Category:Mammals described in 1778 Category:Taxa named by Peter Simon Pallas