{{Short description|Genus of rodents}} {{Italic title}} {{Use dmy dates|date=September 2020}} {{Automatic taxobox | name = ''Spermophilus'' | fossil_range = Late [[Miocene]] - Recent | image = Asia Minor Ground Squirrel.jpg | image_caption = ''[[Spermophilus xanthoprymnus]]'' | taxon = Spermophilus | authority = [[Frederic Cuvier|F. Cuvier]], 1825 | type_species = ''[[European ground squirrel|Mus citellus]]'' | type_species_authority = [[Linnaeus]], [[12th edition of Systema Naturae|1766]] | subdivision_ranks = [[Species]] | subdivision = See text.<!-- A list of sensu stricto species will be added after genus articles are started --> }} '''''Spermophilus''''', also known as the '''Old World ground squirrels''', '''sousliks''' or, sometimes, '''spermophiles''', is an [[Old World]] [[genus]] of [[ground squirrel]]s in the squirrel family ([[Sciuridae]]).<ref>{{MSW3|id=12400978}}</ref> Formerly, the genus was more species rich, but it has since been reduced to contain only species from Eurasia, with many former species having been moved to other genera. The 18 known species are distributed across the [[Eurasian Steppe|Eurasian steppe belt]] and adjacent regions between [[Central Europe]] and [[Manchuria]]. They inhabit steppe and comparable short grassland habitats, including airports, as well semi-deserts and, in some cases, agricultural fields. Their diet is dominated by seeds, grasses, forbs, roots and tubers, but may also include small invertebrates. ''Spermophilus'' live in colonial burrows, and spend much of the year in [[hibernation]] and, in dry regions, [[aestivation]].
==Etymology== Some species are called ''susliks'' (or sousliks), which comes from Russian суслик, ''[[wiktionary:suslik|suslik]]''. In some languages, a variation of the name is in common usage, for example ''[[Wiktionary:suseł|suseł]]'' in [[Polish language|Polish]], or ''[[Wiktionary:Ziesel|Ziesel]]'' in [[German language|German]], all of which probably ultimately descend from a common [[Proto-Slavic]] root. The scientific name of this genus means "seed-lovers" (gr. σπέρμα ''sperma'', genitive σπέρματος ''spermatos'' – seed; φίλος ''philos'' – friend, lover).<ref>{{cite journal |last1= Palmer |first1= T.S. |year= 1904|title= Index Generum Mammalium: a List of the Genera and Families of Mammals|url= https://www.biodiversitylibrary.org/item/88553#page/647/mode/1up|journal= North American Fauna|volume= 23 |pages= 639|doi= 10.3996/nafa.23.0001 | access-date= 2018-02-09|doi-access= free}}</ref>
==Ecology and behavior== As typical ground squirrels, ''Spermophilus'' live in open habitats like grasslands, meadows, [[steppe]] and semideserts throughout the [[Eurasian Steppe|Eurasian steppe]] and adjacent regions, feed on the low plants, and use burrows as nests and refuge.<ref name=Krystufek2012>{{cite journal | author1=Kryštufek, B. | author2=B. Vohralík | year=2012 | title=Taxonomic revision of the Palaearctic rodents (Rodentia). Part 1 (Eutamias and Spermophilus) | journal=Lynx, N. S. (Praha) | volume=43 | pages=17–111 }}</ref><ref name=":0" /> They are [[Diurnality|diurnal]] and mostly live in colonies, although some species also can occur singly.<ref name=Smith2008>{{cite book | editor1=Smith, A.T. | editor2= Y. Xie | year=2008 | title=A Guide to the Mammals of China | publisher=Princeton University Press | pages=193–196 | isbn=978-0-691-09984-2 }}</ref> They are found in both lowlands to highlands, [[hibernate]] during the colder months (up to {{circa}} 8{{1/2}} months each year in some species) and in arid regions they may also [[aestivate]] during the summer or fall.<ref name=Krystufek2012/> The distributions of the various species are mostly separated, often by large rivers or mountain ranges, although there are regions inhabited by as many as three species and rarely two species may even form mixed colonies.<ref name=Krystufek2012/> Generally, however, [[interspecific competition]] is intense and thus, sufficient differences in size (as per [[Hutchinson's rule]]) seem to be a necessary precondition for [[sympatry]].<ref name=":1" /> A few species are known to [[Hybrid (biology)|hybridize]] where their ranges come into contact.<ref name=Krystufek2012/>
In contrast to most other, smaller grassland rodents, such as [[Vole|voles]], [[Gerbillinae|gerbils]] or the [[steppe pika]], but like other ground squirrels, ''Spermophilus'' are mainly active during the day, and rely on their good vision and their agility to detect and evade predators. Therefore, ''Spermophilus'' require open and low-growing habitats, and can cope well even with overgrazing.<ref>{{Cite journal |last1=Cao |first1=Chan |last2=Shuai |first2=Ling-Ying |last3=Xin |first3=Xiao-Ping |last4=Liu |first4=Zhi-Tao |last5=Song |first5=Yan-Ling |last6=Zeng |first6=Zhi-Gao |date=2016 |title=Effects of cattle grazing on small mammal communities in the Hulunber meadow steppe |journal=PeerJ |volume=4 |article-number=e2349 |doi=10.7717/peerj.2349 |doi-access=free |issn=2167-8359 |pmc=5012293 |pmid=27635323}}</ref> On the contrary, dense vegetation makes them vulnerable to predation by their many predators, including the [[Steppe eagle|steppe]] and [[Eastern imperial eagle|eastern imperial eagles]], and [[Steppe polecat|steppe]] and [[marbled polecat|marbled polecats]], often rendering them dependent on the presence of large grazing animals.<ref name=":1" /><ref>{{Cite journal |last1=Koshkina |first1=Alyona |last2=Freitag |first2=Martin |last3=Grigoryeva |first3=Irina |last4=Hölzel |first4=Norbert |last5=Stirnemann |first5=Ingrid |last6=Velbert |first6=Frederike |last7=Kamp |first7=Johannes |date=2023 |title=Post-Soviet fire and grazing regimes govern the abundance of a key ecosystem engineer on the Eurasian steppe, the yellow ground squirrel Spermophilus fulvus |journal=Diversity and Distributions |language=en |volume=29 |issue=3 |pages=395–408 |doi=10.1111/ddi.13668 |bibcode=2023DivDi..29..395K |issn=1472-4642 |doi-access=free}}</ref><ref>{{Cite journal |last1=Surkova |first1=Elena |last2=Popov |first2=Sergey |last3=Tchabovsky |first3=Andrey |date=2019 |title=Rodent burrow network dynamics under human-induced landscape transformation from desert to steppe in Kalmykian rangelands |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/1749-4877.12392 |journal=Integrative Zoology |language=en |volume=14 |issue=4 |pages=410–420 |doi=10.1111/1749-4877.12392 |pmid=30983144 |issn=1749-4877 |url-access=subscription}}</ref> Like other colonial ground squirrels, ''Spermophilus'' use high-pitched calls to communicate and warn colony members of predators.<ref>{{Cite journal |last1=Matrosova |first1=Vera A. |last2=Schneiderová |first2=Irena |last3=Volodin |first3=Ilya A. |last4=Volodina |first4=Elena V. |date=2012-01-01 |title=Species-specific and shared features in vocal repertoires of three Eurasian ground squirrels (genus Spermophilus) |url=https://doi.org/10.1007/s13364-011-0046-9 |journal=Acta Theriologica |language=en |volume=57 |issue=1 |pages=65–78 |doi=10.1007/s13364-011-0046-9 |issn=2190-3743|url-access=subscription }}</ref>
==Appearance== ''Spermophilus'' are overall yellowish, light orangish, light brownish or greyish. Although many are inconspicuously mottled or spotted, or have orange markings on the head, overall they lack strong patterns, except in ''[[Speckled ground squirrel|S. suslicus]]'' and ''[[Spermophilus odessanus|S. odessanus]]'', which commonly have brown upperparts with clear white spotting.<ref name=Krystufek2012/> Size varies with species and they have a head-and-body length of {{circa}} {{cvt|17-40|cm|in}}. Before hibernation the largest ''[[Yellow ground squirrel|S. fulvus]]'' may weigh up to {{cvt|2|kg|lb}} and the largest ''[[Russet ground squirrel|S. major]]'' up to almost {{cvt|1.4|kg|lb}}, but they always weigh much less earlier in the year and other species are considerably smaller, mostly less than {{cvt|0.5|kg|lb}} even in peak condition before hibernation.<ref name=Krystufek2012/> All have a fairly short tail that—depending on exact species—is around 10–45% of the length of the head-and-body.<ref name=Krystufek2012/>
==Taxonomy== As traditionally defined the genus was very species-rich, ranging through Europe, Asia and North America, but this arrangement was found to be [[paraphyletic]] to the certainly distinct prairie dogs (''[[Prairie dog|Cynomys]]''), marmots (''[[Marmot|Marmota]]''), and antelope squirrels (''[[Antelope squirrel|Ammospermophilus]]''). As a consequence, all the former ''Spermophilus'' species of North America have been moved to other genera, leaving the European and Asian species as true ''Spermophilus'' (the only exceptions being the two Asian ''[[Urocitellus]]'' species).<ref>{{cite journal |author=Helgen Kristofer M |display-authors=etal |year=2009 |title=Generic revision in the Holarctic ground squirrel genus Spermophilus |journal=Journal of Mammalogy |volume=90 |issue=2 |pages=270–305 |doi=10.1644/07-mamm-a-309.1 |doi-access=free}}</ref> The exact relations between these clades are slightly unclear.<ref>{{cite journal |author1=Helgen, Kristofer M. |author2=Cole, F. Russel |author3=Helgen, Lauren E. |author4=Wilson, Don E |name-list-style=amp |year=2009 |title=Generic Revision in the Holarctic Ground Squirrel Genus ''Spermophilus'' |journal=Journal of Mammalogy |volume=90 |issue=2 |pages=270–305 |doi=10.1644/07-MAMM-A-309.1 |doi-access=free}}</ref> According to Simonov et al. (2024), ''Spermophilus'' consists of 18 species. Also according to this study, the genus can be divided into four major clades that diverged during the [[Late Miocene]].<ref name=":0">{{Cite journal |last1=Simonov |first1=Evgeniy |last2=Lopatina |first2=Natalia V. |last3=Titov |first3=Sergey V. |last4=Ivanova |first4=Anastasiya D. |last5=Brandler |first5=Oleg V. |last6=Surin |first6=Vadim L. |last7=Matrosova |first7=Vera A. |last8=Dvilis |first8=Alisa E. |last9=Oreshkova |first9=Nataliya V. |last10=Kapustina |first10=Svetlana Yu. |last11=Golenishchev |first11=Fedor N. |last12=Ermakov |first12=Oleg A. |date=2024-06-01 |title=Traditional multilocus phylogeny fails to fully resolve Palearctic ground squirrels (Spermophilus) relationships but reveals a new species endemic to West Siberia |url=https://www.sciencedirect.com/science/article/pii/S1055790324000496 |journal=Molecular Phylogenetics and Evolution |volume=195 |article-number=108057 |doi=10.1016/j.ympev.2024.108057 |pmid=38471598 |bibcode=2024MolPE.19508057S |issn=1055-7903|url-access=subscription }}</ref> {{Sidebar | title= Internal phylogeny after Simonov et al. 2024<ref name=":0">{{Cite journal |last1=Simonov |first1=Evgeniy |last2=Lopatina |first2=Natalia V. |last3=Titov |first3=Sergey V. |last4=Ivanova |first4=Anastasiya D. |last5=Brandler |first5=Oleg V. |last6=Surin |first6=Vadim L. |last7=Matrosova |first7=Vera A. |last8=Dvilis |first8=Alisa E. |last9=Oreshkova |first9=Nataliya V. |last10=Kapustina |first10=Svetlana Yu. |last11=Golenishchev |first11=Fedor N. |last12=Ermakov |first12=Oleg A. |date=2024-06-01 |title=Traditional multilocus phylogeny fails to fully resolve Palearctic ground squirrels (Spermophilus) relationships but reveals a new species endemic to West Siberia |url=https://www.sciencedirect.com/science/article/pii/S1055790324000496 |journal=Molecular Phylogenetics and Evolution |volume=195 |article-number=108057 |doi=10.1016/j.ympev.2024.108057 |pmid=38471598 |bibcode=2024MolPE.19508057S |issn=1055-7903|url-access=subscription }}</ref> |content1= {{clade |label1=''Spermophilus'' |1={{clade |2={{clade |1=[[Daurian ground squirrel]] (''S. dauricus'') |2=[[Alashan ground squirrel]] (''S. alashanicus'') }} |1={{clade |2={{clade |1=[[Caucasian Mountain ground squirrel]] (''S. musicus'') |2=[[Little ground squirrel]] (''S. pygmaeus'') }} |1={{clade |2={{clade |2=[[Yellow ground squirrel]] (''S. fulvus'') |1={{clade |2={{clade |1=[[Tian Shan ground squirrel]] (''S.nilkaensis'') |2=[[Relict ground squirrel]] (''S. relictus'') }} |1={{clade |2=[[Pallid ground squirrel]] (''S. pallidicauda'') |1={{clade |2={{clade |1=[[Red-cheeked ground squirrel]] (''S. erythrogenys'') |2=''[[Spermophilus vorontsovi]]'' }} |1={{clade |2=[[Brandt's ground squirrel]] (''S. brevicauda'') |1={{clade |1=''[[Spermophilus selevini]]'' |2=[[Russet ground squirrel]] (''S. major'') }} }} }} }} }} }} |1={{clade |2={{clade |1=[[Speckled ground squirrel]] (''S. suslicus'') |2=''[[Spermophilus odessanus]]'' }} |1={{clade |2=[[Asia Minor ground squirrel]] (''S. xanthoprymnus'') |1={{clade |2=[[Taurus ground squirrel]] (''S. taurensis'') |1=[[European ground squirrel]] (''S. citellus'') }} }} }} }} }} }} }} }} {| class="wikitable mw-collapsible" |- !Clade !Image !Name !Common name !Distribution |- ! rowspan="2" style="text-align:center;"| East Asian clade |[[File:Spermophilus alashanicus 381342585.jpg|120px]] |''[[Alashan ground squirrel|Spermophilus alashanicus]]'' |Alashan ground squirrel |[[Northern and southern China|Northern China]] west of the [[Yellow River]]. |- | |''[[Daurian ground squirrel|Spermophilus dauricus]]'' |Daurian ground squirrel |Northern China and [[Manchuria]] between the [[Yellow River]] and the [[Amur|Amur River]]. |- ! rowspan="2" style="text-align:center;"| ''Pygmaeus'' clade |[[File:Суслик в поле.jpg|120px]] |''[[Caucasian Mountain ground squirrel|Spermophilus musicus]]'' |Caucasus Mountain ground squirrel |Southern [[Pontic–Caspian steppe|Pontic-Caspian steppe]] between the [[Dnieper]] and [[Volga]] rivers; [[Crimea]]. |- |[[File:Spermophilus pygmaeus 58814806.jpg|120px]] |''[[Little ground squirrel|Spermophilus pygmaeus]]'' |Little ground squirrel |Northern [[Central Asia]] east of the Volga. |- ! rowspan="5" style="text-align:center;"| Europe/Asia Minor clade |[[File:European souslik (Spermophilus citellus) Obrovisko 3.jpg|120px]] |''[[European ground squirrel|Spermophilus citellus]]'' |European ground squirrel |[[Balkans|Balkan Peninsula]] to the [[Czech Republic]] and [[Poland]], [[Moldova]] and [[western Ukraine]]. |- |[[File:Spotted souslik (Spermophilus odessanus).jpg|120px]] |''[[Spermophilus odessanus]]'' |Polonian ground squirrel |[[Eastern Europe]] between the [[Carpathian Mountains]] and the Dnieper River. |- |[[File:Spermophilus suslicus 384911573.jpg|120px]] |''[[Speckled ground squirrel|Spermophilus suslicus]]'' |Speckled ground squirrel |Northern [[Pontic–Caspian steppe|Pontic steppe]] between the [[Dnieper]] and [[Volga]] rivers. |- |[[File:Spermophilus taurensis.jpg|120px]] |[[Taurus ground squirrel|''Spermophilus taurensis'']] |Taurus ground squirrel |South-central Anatolia. |- |[[File:Asia Minor Ground Squirrel.jpg|120px]] |[[Asia Minor ground squirrel|''Spermophilus xanthoprymnus'']] |Anatolian ground squirrel |[[Lycia|South-western]], [[Central Anatolia region|central]] and [[Eastern Anatolia region|eastern Anatolia]]; [[Armenian highlands]]. |- ! rowspan="9" style="text-align:center;"| ''Colobotis'' clade | |[[Spermophilus brevicauda|''Spermophilus brevicauda'']] |Brandt's ground squirrel |Border region between [[Kazakhstan]] and [[China]] along the [[Tian Shan]]. |- |[[File:Spermophilus erythrogenys 392429402.jpg|120px]] |[[Red-cheeked ground squirrel|''Spermophilus erythrogenys'']] |Red-cheeked ground squirrel |Western Siberia between the [[Irtysh]] and [[Ob]] rivers. |- |[[File:Spermophilus fulvus Baikonur 02.jpg|120px]] |[[Yellow ground squirrel|''Spermophilus fulvus'']] |Yellow ground squirrel |Central Asia and the northern [[Iranian plateau|Iranian Plateau]]. |- |[[File:Spermophilus major 203118180.jpg|120px]] |[[Russet ground squirrel|''Spermophilus major'']] |Russet ground squirrel |[[Ural Mountains]] between the Volga and [[Tobol]] rivers. |- | |[[Pallid ground squirrel|''Spermophilus pallicauda'']] |Pale-bellied ground squirrel |Mongolia and [[Gobi Desert]] [[Altai Mountains]]. |- | |[[Tian Shan ground squirrel|''Spermophilus nilkaensis'']] |Tian Shan ground squirrel |Endemic to the Tian Shan. |- |[[File:Spermophilus relictus in Ertosh.jpg|120px]] |[[Relict ground squirrel|''Spermophilus relictus'']] |Relict ground squirrel |Central Asia between the Tian Shan and the [[Pamir Mountains]]. |- | |[[Spermophilus seleveni|''Spermophilus seleveni'']] | |Central Asia between the Tobol and Irtysh rivers, [[Lake Balkhash]] and the [[Tian Shan]]. |- |[[File:Spermophilus erythrogenys 121075441.jpg|120px]] |[[Spermophilus vorontsovi|''Spermophilus vorontsovi'']] | |[[Western Siberia]] between the Ob and the [[Kuznetsk Alatau|Kuznetsk Alatau Mountains]]. |} ===Extinct species=== Discovery and examination of one of the best preserved Eurasian ground squirrel fossils yet recovered allowed the study of many previously unknown aspects of ground squirrel cranial anatomy, and prompted a critical reassessment of their phylogenetic position.<ref>{{Cite journal|author1=Maxim V. Sinitsa |author2=Natalia V. Pogodina |author3=Lyudmila Y. Кryuchkova |year=2019 |title=The skull of ''Spermophilus nogaici'' (Rodentia: Sciuridae: Xerinae) and the affinities of the earliest Old World ground squirrels |journal=Zoological Journal of the Linnean Society |volume=186 |issue=3 |pages=826–864 |doi=10.1093/zoolinnean/zly092 }}</ref> As a result, three [[Pleistocene]] species previously considered members of the ''[[Urocitellus]]'' genus were moved to ''Spermophilus'':{{cn|date=January 2025}}
:* [[Extinction|†]]''[[Spermophilus nogaici]]'' :* [[Extinction|†]]''[[Spermophilus polonicus]]'' :* [[Extinction|†]]''[[Spermophilus primigenius]]'' In addition to the recent species, three now-extinct species are known from the Pleistocene of Europe:
''[[Spermophilus citelloides]]'' is known from the Middle Pleistocene to early Holocene of central Europe. It appears to be most closely related to the living ''S. suslicus''.<ref>{{Cite journal|last1=Sinitsa|first1=Maxim V.|last2=Virág|first2=Attila|last3=Pazonyi|first3=Piroska|last4=Knitlová|first4=Markéta|date=2019-10-29|title=Redescription and phylogenetic relationships of Spermophilus citelloides (Rodentia: Sciuridae: Xerinae), a ground squirrel from the Middle Pleistocene – Holocene of Central Europe|url=https://www.tandfonline.com/doi/full/10.1080/08912963.2019.1677640|journal=Historical Biology|volume=33|pages=19–39|doi=10.1080/08912963.2019.1677640|bibcode=2021HBio...33...19S |s2cid=208578857|issn=0891-2963|url-access=subscription}}</ref>
''[[Spermophilus severskensis]]'' is known from the late Pleistocene ([[Weichselian glaciation|Weichselian]]) of the [[Desna (river)|Desna]] area, Ukraine. It appears to have been a highly specialised grazer and close relative of the living ''S. pygmaeus''.<ref name=":1">{{Cite journal |last1=Popova |first1=L. V. |last2=Maul |first2=L. C. |last3=Zagorodniuk |first3=I. V. |last4=Veklych |first4=Yu. M. |last5=Shydlovskiy |first5=P. S. |last6=Pogodina |first6=N. V. |last7=Bondar |first7=K. M. |last8=Strukova |first8=T. V. |last9=Parfitt |first9=S. A. |date=2019-03-10 |title='Good fences make good neighbours': Concepts and records of range dynamics in ground squirrels and geographical barriers in the Pleistocene of the Circum-Black Sea area |url=https://www.sciencedirect.com/science/article/abs/pii/S1040618217307267 |journal=Quaternary International |series=Bridging Europe and Asia: Quaternary stratigraphy and paleolithic human occupation |volume=509 |pages=103–120 |bibcode=2019QuInt.509..103P |doi=10.1016/j.quaint.2018.03.023 |issn=1040-6182 |url-access=subscription}}</ref>
''[[Spermophilus superciliosus]]'' is known from the Middle Pleistocene to reportedly the early 20th century, with a vast range across much of Europe, from southern England to the [[Volga]] and the [[Ural Mountains]]. It was similar in size to the recent ''S. major'', and a probable ancestor of ''S. fulvus''.<ref>{{Cite journal |last=Popova |first=L. V. |date=2016-10-28 |title=Evolutionary lineage of Spermophilus superciliosus – S. fulvus (Rodentia, Sciuridae) in the quaternary of the Dnieper area: An ability of a biostratigraphical implication |journal=Quaternary International |series=The Quaternary of the Urals: Global trends and Pan-European Quaternary records |volume=420 |pages=319–328 |doi=10.1016/j.quaint.2015.10.104 |bibcode=2016QuInt.420..319P |issn=1040-6182|doi-access=free }}</ref>
==Relationship with humans== Ground squirrels may carry fleas that transmit diseases to humans (see [[Black Death]]), and have been destructive in tunneling underneath human habitation.<ref>[http://www.britannica.com/EBchecked/topic/246960/ground-squirrel Encyclopædia Britannica]</ref>
== References == {{Reflist}}
== External links == {{Commons|Spermophilus}} {{Wikispecies|Spermophilus}}
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[[Category:Spermophilus| ]] [[Category:Rodents of Europe]] [[Category:Mammals of North America]] [[Category:Mammals of Asia]] [[Category:Rodent genera]] [[Category:Taxa named by Frédéric Cuvier]]