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Lacerta agilis LINNAEUS, 1758

IUCN Red List - Lacerta agilis - Least Concern, LC
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Higher TaxaLacertidae, Lacertinae, Sauria, Lacertoidea, Squamata (lizards)
SubspeciesLacerta agilis agilis LINNAEUS 1758
Lacerta agilis argus (LAURENTI 1768)
Lacerta agilis boemica SUCHOW 1929
Lacerta agilis bosnica SCHREIBER 1912
Lacerta agilis brevicaudata PETERS 1958
Lacerta agilis chersonensis ANDRZEJOWSKI 1832
Lacerta agilis exigua EICHWALD 1831
Lacerta agilis grusinica PETERS 1960
Lacerta agilis ioriensis PETERS & MUSKHELISCHWILI 1968
Lacerta agilis mzymtensis TUNIYEV & TUNIYEV 2008
Lacerta agilis tauridica SUCHOW 1926 
Common NamesE: Sand Lizard
G: Zauneidechse
NL: Zandhagedis
Croatian: livadna gušterica
Russian: Прыткая ящерица
Chinese: 捷蜥蜴
G: Grusinische Zauneidechse [grusinica]
Italian: lucertola agile 
SynonymLacerta agilis LINNAEUS 1758: 203
Ameiva agilis — MEYER 1795: 28
Ameiva agilis —  LINK 1807: 61
Lacerta stirpium — MILNE-EDWARDS 1829: 83
Lacerta stirpium — MÉNÉTRIES 1832: 61
Lacerta stirpium — COCTEAU 1835
Lacerta agilis — COCTEAU 1835
Lacerta stirpium — DUMÉRIL & BIBRON 1839: 196
Lacerta stirpium — MÜLLER 1878: 412
Lacerta paradoxa BEDRIAGA 1886
Lacerta agilis — ANDERSSON 1900: 8
Lacerta agilis spinalis WERNER 1910
Lacerta exigua paradoxa — SUCHOW 1948
Lacerta agilis agilis — BISCHOFF 1988
Lacerta agilis — ENGELMANN et al 1993
Lacerta agilis — SZCZERBAK 2003
Lacerta agilis grusoni — KWET & TRAPP 2015 (in error)
Lacerta agilis agilis — BISCHOFF 2020

Lacerta agilis garzoni PALACIOS & CASTROVIEJO 1975
Lacerta agilis garzoni — BISCHOFF 2020

Lacerta agilis argus (LAURENTI 1768)
Lacerta agilis argus — BISCHOFF 1988
Lacerta agilis argus — SCHLÜTER 2004
Lacerta agilis argus — KÜHNEL 2013
Lacerta agilis argus — BISCHOFF 2020

Lacerta agilis boemica SUCHOW 1929
Lacerta boemica SUCHOV 1929
Lacerta exigua boemica — SUCHOW 1948
Lacerta agilis boemica — BORKIN & DAREVSKY 1987
Lacerta agilis boemica — BISCHOFF 1988
Lacerta agilis boemica — ROITBERG et al. 2000
Lacerta agilis boemica — BISCHOFF 2005
Lacerta agilis boemica — TUNIYEV & TUNIYEV 2008
Lacerta boemica — ANDRES et al. 2014
Lacerta agilis boemica — BISCHOFF 2020

Lacerta agilis bosnica SCHREIBER 1912
Lacerta agilis bosnica — BISCHOFF 1988
Lacerta agilis bosnica — NESSING 1989
Lacerta agilis bosnica — TRAPP 2005
Lacerta agilis bosnica — TRAPP 2011
Lacerta agilis bosnica — DIECKMANN 2017
Lacerta agilis bosnica — BISCHOFF 2020
Lacerta agilis bosnica — GEBHART 2020
Lacerta agilis bosnica — BAMANN et al. 2021

Lacerta agilis brevicaudata PETERS 1958
Lacerta agilis brevicaudata — BORKIN & DAREVSKY 1987
Lacerta agilis brevicaudata — BISCHOFF 1988
Lacerta agilis brevicaudata — BAUER & GÜNTHER 1995
Lacerta agilis brevicaudata — BISCHOFF 2007
Lacerta agilis brevicaudata — TUNIYEV & TUNIYEV 2008
Lacerta agilis brevicaudata — TUNIYEV et al. 2014
Lacerta agilis brevicaudata — BISCHOFF 2020
Lacerta agilis brevicaudata — DORONIN & DORONINA 2022

Lacerta agilis chersonensis ANDRZEJOWSKI 1832
Lacerta agilis L. var. Dzieduszyckii BAYGER 1909
Lacerta agilis chersonensis — SUCHOW 1948
Lacerta agilis euxinica FUHN & VANCEA 1964 (fide BISCHOFF 1984)
Lacerta agilis chersonensis — BORKIN & DAREVSKY 1987
Lacerta agilis euxinica — BORKIN & DAREVSKY 1987
Lacerta agilis chersonensis — BISCHOFF 1988
Lacerta agilis chersonensis — SCHLÜTER 2005
Lacerta agilis euxinica — GHERGHEL & STRUGARIU 2009
Lacerta agilis chersonensis — GROZDANOV & TZАNKOV 2014
Lacerta agilis chersonensis — BISCHOFF 2020

Lacerta agilis exigua EICHWALD 1831
Lacerta agilis var. dilepis LICHTENSTEIN 1856
Lacerta agilis var. orientalis KESSLER 1878 (fide DORONIN & DORONINA 2022)
Lacerta exigua — SUCHOW 1948
Lacerta agilis var. kurtuana KASHTCHENKO 1909: 125 (fide ZHAO & ADLER 1993: 205)
Lacerta agilis exigua — BISCHOFF 1988
Lacerta agilis exigua — BAUER & GÜNTHER 1995
Lacerta agilis exigua — TUNIYEV & TUNIYEV 2008
Lacerta agilis exigua — CHIRIKOVA et al. 2016
Lacerta agilis exigua — EPLANOVA 2016
Lacerta agilis exigua — DAVLETBAKOV et al. 2016
Lacerta agilis exigua — BISCHOFF 2020
Lacerta agilis exigua — DORONIN & DORONINA 2022

Lacerta agilis grusinica PETERS 1960
Lacerta agilis grusinica — BORKIN & DAREVSKY 1987
Lacerta agilis grusinica — BISCHOFF 1988
Lacerta agilis grusinica — TUNIYEV 1990
Lacerta agilis grusinica — BISCHOFF 2003
Lacerta agilis grusinica — TUNIYEV & TUNIYEV 2008
Lacerta agilis grusinica — TUNIYEV et al. 2014
Lacerta agilis grusinica — BISCHOFF 2020
Lacerta agilis grusinica — DORONIN & DORONINA 2022

Lacerta agilis ioriensis PETERS & MUSKHELISCHWILI 1968
Lacerta agilis ioriensis — BORKIN & DAREVSKY 1987
Lacerta agilis ioriensis — BISCHOFF 1988
Lacerta agilis ioriensis — BAUER & GÜNTHER 1995
Lacerta agilis ioriensis — BISCHOFF 2003
Lacerta agilis iorensis — TUNIYEV & TUNIYEV 2008 (in error)
Lacerta agilis ioriensis — NESSING 2011
Lacerta agilis ioriensis — DORONIN et al. 2019
Lacerta agilis ioriensis — BISCHOFF 2020

Lacerta agilis mzymtensis TUNIYEV & TUNIYEV 2008
Lacerta agilis mzymtensis — MAYER 2015
Lacerta agilis mzymtensis — KUKUSHKIN et al. 2020
Lacerta agilis mzymtensis — BISCHOFF 2020

Lacerta agilis tauridica SUCHOW 1926
Lacerta punctata L. — HABLIZL 1785:195 (part.)
Lacerta europaea — PALLAS 1813:29 (part.)
Lacerta paradoxa BEDRIAGA 1886:170 (part.)
Lacerta agilis var. concolor SCHREIBER 1912:482
Lacerta agilis tauridica SUCHOW 1927:331 (part.)
Lacerta exigua exigua — SUCHOW 1948:111 (part.)
Lacerta agilis exigua — NIKOLSKY 1915:298 (part.)
Lacerta agilis exigua — TERENTYEV & CHERNOV 1949:181 (part.)
Lacerta agilis exigua — TARATSHUK 1959:179 (part.)
Lacerta agilis exigua — MERTENS & WERMUTH 1960:102
Lacerta agilis exigua — SZCZERBAK 1966:97 (part.)
Lacerta agilis exigua — DAREVSKY et al. 1976 (part.)
Lacerta agilis exigua — BISCHOFF 1984:53 (part.)
Lacerta agilis exigua — BISCHOFF 1988:13 (part.)
Lacerta agilis tauridica — KALYABINA-HAUF et al. 2004
Lacerta agilis tauridica — BISCHOFF 2020 
DistributionAustria, Switzerland, Liechtenstein, Germany, France, Andorra, Denmark, Sweden, SE Norway,
Czech Republic (formerly Czechoslovakia), Slovakia, Hungary, Bulgaria, Greece, Albania, N Balkan,
Netherlands, Belgium, Luxembourg, S England, NE Italy,
Croatia, Bosnia-Hercegovina, Serbia, Macedonia, Montenegro,
Bulgaria, N Greece, Romania, E Poland, Belorussia, Belarus,
W Russia (in the north up to S Karelia and SE Finland, NE Caucasus),
Russia (north of the Caucasus Mts., east up to Lake Baikal),
Ukraina (east of the Dnjepr River and W Ukraina),
Armenia, NE Turkey,
Kazakhstan, Kirgistan (south up to Issyk Kul),
NW China (W Xinjiang)
Caucasian coast of the Black Sea in Russia near Sochi,
Republic of Georgia (coastal region and upper Iori River in the Caucasus Mts.),
Moldova, Latvia, Estonia, Lithuania, Azerbaijan, NW Mongolia

boemica: S Russia (Dagestan), NE Caucasus; Type locality: Russia, North Ossetia, the neighborhood of Vladikavkaz (43°03′18.3′′N 44°40′52.4′′E)

bosnica: Bulgaria, Greece, Croatia, Bosnia-Hercegovina; Type locality: “Bosnien”

brevicaudata: NE Turkey, Armenia, Republic of Georgia; elevation up to 1900 m.

chersonensis: Carpathian Mountains eastward, Romania.

dzieduszyckii: Ukraine; Type locality: Lwiw-Holosko, Ukraine.

exigua: Caucasus, Kyrgyzstan; Type locality: Russia, Ural Mountains

garzoni: Spain (Gerona), Andorra; Type locality: “Falda oeste del Puig de Basa, Tossas, Pirineos, provincia de Gerona, España. 1850 msnm”.

grusinica: Republic of Georgia, NE Turkey, Caucasus

ioriensis: Republic of Georgia; type locality: near Tianeti by the Iori river in E Republic of Georgia.

mzymtensis: Russia (Western Caucasus, high-mountains of the upper basing of Mzymta River; vicinity of Sochi); Type locality: “mountain Aishkha-2, Adler Rayon of Sochi City, Krasnodar Kray, Russia”

orientalis: Type locality: Pyatigorsk.

tauridica: Ukraine (Crimea): Type locality: near Simferopol, Chumakarka settl., Ukraine, Crimea (44°54′12′′N 34°06′16.1′′E)

Type locality: South Sweden  
Reproductionoviparous; Near the northern limit of their distribution (in Sweden), Lacerta agilis shifts tactics seasonally, such that the eggs in late clutches complete development more quickly (when incubated at a standard temperature) than do those of early clutches. That acceleration is achieved by a reduction in egg size, and by an increase in the duration of uterine retention of eggs (especially, after cool weather) (Shine et al. 2017). 
TypesHolotype: NRM (RMS, Royal Museum Stockholm)
Holotype: MNCN 44271.1, Falda oeste del Puig de Basa, Tossas, Pirineos, provincia de Gerona, España, 07.08.1972, Coll. F. Palacios and J. López. [argus]
Lectotype: ZISP 30363, designated by Doronin & Doronina 2020;
paralectotypes: ZISP, BMNH 1960.1.4.26–30, BMNH 1965.337–342 etc. (93); Coll. G.F. Suchow 1928. Some syntypes are not cataloged fide Doronin et al., 2018 [boemica]
Holotype: ZISP 12900, Armenia, Stepanawan, 26.06.1920, Coll. A.B. Shelkovnikov; paratypes: ZFMK, ZIL, ZMB, MM [brevicaudata]
Holotype: unlocated, Ukraine, Kherson, probably the author’s collection (fide I. Doronin, pers. comm.) [chersonensis]
Holotype: SNHM (National Museum of Natural History Lwiw = Lviv, Ukraine) [dzieduszyckii]
Holotype: unlocated fide I. Doronin (pers. comm., Nov 2018)[exigua]
Holotype: MNCN 44271, female [garzoni]
Lectotype: BMNH 1946.8.6.53-54 (?), Abkhazia, Sukhumi. It requires revision of the type specimens [grusinica]
Holotype: ZIT (Institute of Zoology of the Academy of Sciences of Georgia, now the Institute of Zoology Ilia State University), has not been cataloged, Georgia, Tianeti, Iori Valley, 19-23.07.1966, Coll. T.A. Muskhelischwili.. In May 2018, Doronin et al. did not find it, may be lost. Paratypes: ZFMK 8766-67 (ex-ZMB 39276), male and female, ZIT, ZMB, MM [ioriensis]
Holotype. ZISP 24648, adult female; paratypes: ZISP [mzymtensis]
Lectotype: ZISP 22902 [orientalis]
Lectotype: ZISP 12620, adult male, designated by Doronin & Doronina 2020 Leg.: V. Kuznetsov, 2.06.1924 [tauridica] 
DiagnosisDiagnosis (genus): Sharing with Timon the following combination of features that mainly occur elsewhere only in a minority of other Lacertini and are sometimes restricted to the two genera: large body size, head and body deep, nasal process of premaxilla often broad, nine or more premaxillary teeth in adults, usually 27 presacral vertebrae in males; often seven posterior presacral vertebrae with short ribs; occipital scale frequently relatively broad, collar strongly serrated, ventral scales with sloping sides and strong posterior overlap, preanal scale often surrounded by more than one semicircle of smaller scales, frequently green above and underside without bright contrasting colour except sometimes on throat, no blue spots on outer ventral scales, bodenaponeurosis without a lateral septum, retractor lateralis anterior muscle inserting in front of vent laterally away from midline, hatchlings ‘embryonic’ with very large head and short extremities compared to those of other Lacertini. Other more widely distributed features include: supraocular osteoderms complete in adults, inscriptional ribs present, and tail not brightly coloured in hatchlings.
Differs from Timon in smaller adult body sizes (70 to 175 mm from snout to vent); maxillary-jugal suture often stepped, dorsal body scales always keeled; 3–5 narrow light stripes in many juveniles and some females, no well defined blue ocelli on flanks of adults, hemipenial microornamentation usually of crown-shaped tubercles but with longer spines on sides of lobes (latter also present in Timon princeps). Also lacks distinctive karyology of Timon and is not associated with it by albumin immunology (Lutz & Mayer 1984) (from Arnold et al. 2007: 48). 
CommentSynonymy: The status of Lacerta agilis argus is doubtful [RAHMEL 1988]. Synonymy partly after KHALIKOV & ANANJEVA (pers. comm.). ANDRES et al. (2014) found that only the subspecies L. a. chersonensis, L. a. tauridica, L. a. garzoni, and L. a. boemica were genetically well supported. L. a. exigua, L. a. brevicaudata and L. a. grusinica are genetically similar and could not be separated from each other. PEREZ-MELLADO in SALVADOR 1998 considered L. a. garzoni as a synonym of L. a. agilis.

Subspecies: According to morphology (Bischoff, 1988), populations from the western Alps are L. a. agilis, while those of the eastern Alps are L. a. argus (Laurenti 1768). However, cytochrome-b trees did not recover the subspecies L. a. agilis and L. a. argus as separate evolutionary lineages (Kalyabina et al., 2001; Andres et al., 2014).

Hybridization: RYKENA & HENKE (1978) report hybrids between L. viridis and L. agilis.

Distribution: for a map of subspecies see Bischoff 1988, Andres et al. 2014.

Type Species: Lacerta agilis Linnaeus, 1758 is the type species of the genus Lacerta LINNAEUS 1758. Note that many species of the genus have been allocated to other genera by ARNOLD et al. (2007) and other authors. Lacerta is also the type genus of the tribus Lacertini, the subfamily Lacertinae (in this database synonymous with Lacertini), and the family Lacertidae.

Phylogenetics: for a phylogenetic analysis of the genus Lacerta see Amata-Orriols & Escoriza 2023.

Barcoding: Genetic identification has been successful at the species level across multiple species of Lacerta. However, in the composition of L. agilis within the Caucasus, Doronina et al. 2023 can diagnose only the subspecies L. a. boemica and L. a. exigua, while L. a. brevicaudata, L. a. grusinica, and L. a. mzymtensis do not differ in this marker from L. a. exigua and L. a. ioriensis from L. a. boemica. 
EtymologyNamed after Latin “agilis” = mobile, agile, swift.

Lacerta agilis mzymtensis is named after the Mzymta River, which valley is its area of distribution.

The genus is named from Latin “Lacerta” = lizard. 
References
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