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Zootoca vivipara (LICHTENSTEIN, 1823)

IUCN Red List - Zootoca vivipara - Least Concern, LC
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Higher TaxaLacertidae, Lacertinae, Sauria, Lacertoidea, Squamata (lizards)
SubspeciesZootoca vivipara louislantzi ARRIBAS 2009
Zootoca vivipara pannonica (LAC & KLUCH 1968)
Zootoca vivipara vivipara (LICHTENSTEIN 1823) 
Common NamesE: Viviparous Lizard, European common lizard
G: Waldeidechse
Croatian: živorodna gušterica
Russian: Живородящая ящерица
Chinese: 胎蜥
Mongolian: Zulzagalagch gurvel 
SynonymSeps? atra SCHRANK in SCHRANK & MOLL 1785
Lacerta vivipara VON JACQUIN 1787: 33 (nom. nud.)
Lacertus cinereus LACÉPÈDE 1788
Lacerta oedura SHEPPARD 1804
Lacerta montana MIKAN in STURM 1805
Lacerta nigra WOLF in STURM 1805
Lacerta crocea WOLF in STURM 1805
Lacerta fragilis PALMSTRUCH & SWARTZ 1808
Lac. unicolor KUHL 1820
Lac. ptychodes KUHL 1820
Lacerta pyrrhogaster MERREM 1820
Lacerta vivipara LICHTENSTEIN 1823
Lacerta schreibersiana MILNE-EDWARDS 1829: 83
Lacerta Schreibersiana var. a. fusca GACHET 1832
Lacerta Schreibersiana var. b. lutea GACHET 1832
Lacerta chrysogastra ANDRZEJOWSKI 1832
Lacerta crocea — EVERSMANN 1834: 347
Lacerta de jacquin COCTEAU 1835
Lacerta de guérin COCTEAU 1835
Lacerta vivipara — COCTEAU 1835
Lacerta crocea — COCTEAU 1835
Lacerta montana — COCTEAU 1835
Lacerta pyrrhogaster — COCTEAU 1835
Lacerta unicolor — COCTEAU 1835
Zootoca montana TSCHUDI 1837
Zootoca alpina TSCHUDI 1837
Zootica [sic] vivpara — STORK 1838: 413
Lacerta vivipara — DUMÉRIL & BIBRON 1839: 204
“Lacerta isidori Geoffr.” (sic!) SCHINZ 1840
Zootoca vivipara — WELLS 1846: 449
Zootoca vivipara var. nigra — GRAY 1857: 107
Lacerta vivipara var. pallida FATIO 1872
Lacerta vivipara — BOULENGER 1887: 23
Lacerta vivipara var. carniolica WERNER 1897 (nomen nudum)
Lacerta (Zootoca ) vivipara var. melanogastra PRAZÁK 1898
Lacerta vivipara var. barabensis KASHENKO 1902
Lacerta vivipara — STEJNEGER 1907: 251
Lacerta vivipara var. gedulyi FEJERVÁRYI 1923
Lacerta vivipara sachalinensis PERELJESHIN & TERENTJEV 1963 (nom. nud.)
Lacerta vivipara — DELY & BÖHME 1984: 362
Lacerta herseyi AUSTIN 1992: 35
Lacerta vivipara — ENGELMANN et al 1993
Zootoca vivipara — MAYER & BISCHOFF 1996
Zootoca vivipara — NECAS et al. 1997
Lacerta vivipara — SZCZERBAK 2003
Zootoca vivipara — BISCHOFF 2005
Zootoca vivipara — ARNOLD et al. 2007
Lacerta (Zootoca) vivipara sachalinensis — SINDACO & JEREMČENKO 2008
Zootoca vivipara vivipara — CORNETTI et al. 2014

Zootoca vivipara louislantzi ARRIBAS 2009
Zootoca vivipara louislantzi ARRIBAS 2009
Lacerta vivipara vasconiae OSENEGG 1995
Zootoca vivipara louislantzi — CORNETTI et al. 2014
Zootoca vivipara louislantzi — CHABAUD et al. 2022

Zootoca vivipara pannonica (LAC & KLUCH 1968)
Lacerta vivipara pannonica LAC & KLUCH 1968
Lacerta (Zootoca) vivipara pannonica — SINDACO & JEREMČENKO 2008 
DistributionNorway, Sweden, Finland
Switzerland, Liechtenstein, Germany, France, Austria, Denmark,
Poland, Czech Republic, Hungary,
Croatia, Slovenia, Bosnia and Hercegovina, Albania,
Monte Negro, Macedonia, Serbia, Romania, Bulgaria, Greece,
Belgium, Netherlands, Luxembourg,
England, Ireland, N Spain
In the north beyond the Arctic Circle, in the south up to N Italy, Montenegro,
Russia (E Siberia, Sakhalin Island), Estonia, Latvia, Lithuania, Ukraine, Kazakhstan, Mongolia,
Japan (Hokkaido Island)
China (Xinjiang [Altai region] and Heilongjiang)

Type locality: Schneeberg near Vienna, Austria

louislantzi: N Spain, Andorra, SW France (Pyrenees); type locality: Pla de Beret (Vall d’Aràn, Lleida). Spain.

pannonica: W/S Balkan peninsula; Type locality: Boťany, Kapu-sansky Forest in Eastern Slovakia). Mayer 2015 suggested to apply the name only to populations around the type locality.

sachalinensis: Poland, Belarus (Carpathians), Russia east through (Sakhalin island), Japan (n Hokkaido); Type locality: Sakhalin island.  
ReproductionViviparous and oviparous. Lacerta vivipara is ovovivparous throughout most of its range, but oviparous in the extreme southwest portion (Heulin et al., 1989). However, although lizards from the oviparous and ovovivparous populations are similar in many respects, they produce significant numbers of embryonic malformations when hybridized in the laboratory (Heulin et al., 1989). Zootoca vivipara louislantzi and carniolica are oviparous. Crosses between oviparous and ovovivparous populations of Zootoca (Lacerta) vivipara produce an F1 generation with an ‘‘intermediate’’ phenotype (Arrayago et al. 1996, Murphy & Thompson 2011), in which females retain embryos in utero for longer than oviparous females and embryos are surrounded by a thin, transparent shell. However, Cornetti et al. (2015) show that oviparous and ovovivparous populations in a contact zone in the Italian Alps do not hybridize and have been reproductively isolated for a long time, as shown by genetic differences. Hence, Cornetti et al. 2015 suggest to recognize Z. v. vivipara and Z. v. carniolica as two separate species.<br /><br />Recknagel et al. 2018 found that a single origin of viviparity and a subsequent reversal to oviparity in one clade.<br /><br />Z. v. pannonica is a ovovivparous lineage.<br />Z. v. louislantzi and carniolica are oviparous. 
TypesHolotype: unlocated
Holotype: MNCN 44290, Paratypes in ZFMK, MNCN, NMW and Coll. O. Arribas, holotype in MNCN [louislantzi]
Holotype: private collection of Ján Lác (№ 2) [pannonica] 
DiagnosisDiagnosis (genus): The only genus of Lacertini with a hemipenis that has a fully developed armature and complexly folded lobes, in which fully-formed young are produced over most of the geographic distribution, and which is also characterised by small short head, short limbs and thick short tail. Also possessing the following features found only in a minority of other Lacertini: nasal process of premaxilla often broad, postorbital and postfrontal bones fused throughout life, posterior presacral vertebrae with short ribs frequently five, row of supraciliary granules reduced or even absent, outer edge of parietal scale reaching the lateral border of parietal table both posteriorly and anteriorly, single postnasal scale, frequent contact between supranasal and anterior loreal scales above nostril, few dorsal body scales (25–37 across mid-back), serrated collar, imbricate ventral scales, no blue spots on outer ventral scales, insertion of retractor lateralis anterior muscle in front of vent thick, 36 macrochromosomes in males and often 35 in females, no microchromosomes, usually Z1Z2W-type sex chromosomes. Other more widely distributed features include head and body not strongly depressed and supraocular osteoderms complete in adults, usually seven premaxillary teeth in adults, usual number of presacral vertebrae 26 in males, inscriptional ribs sometimes absent, tail with bright colouring in juveniles of some populations (although largely masked by darker pigment), hemipenial microornamentation of crownshaped tubercles (from Arnold et al. 2007: 62). 
CommentNomenclature: Zootoca has been considered as a subgenus of Lacerta, but more recently as valid genus. VON JACQUIN 1787 did not use the name Lacerta vivipara in order to describe a new species but to report the fact that the species is viviparous. Hence, VON JACQUIN 1787 is not the author of Lacerta vivipara as indicated by much of the older literature but rather LICHTENSTEIN 1823 who provided the first description of the species (SCHMIDTLER & BÖHME 2011).

Synonymy: partly after SCHMIDTLER & BÖHME 2011. The validity of Z. v. sachalinensis has been questioned by GUILLAUME et al. (2006). The description of “Lacerta herseyi” is certainly not sufficient to establish a new species. His description seems to be based on an aberrant L. vivipara that displays an unusual behavior (“leaping lizard”). Z. v. sachalinensis is a nomen nudum fide Mayer & Böhme 2000, Mayer 2015.

In Z. vivipara, male mortality and emigration are not higher under male-biased adult sex ratio. Rather, an excess of adult males begets aggression toward adult females, whose survival and fecundity drop, along with their emigration rate (GALLIARD et al. 2005).

Karyotype: Z. vivipara includes populations with varying karyotypes (Kupriyanova et al. 2014, 2017). The genome has been sequenced by Yurchenko et al. 2019. Based on chromosomal and molecular studies Kupriyanova & Safronova (2020) concluded that Z. vivipara represents a group of cryptic taxa.

Subspecies: Z. v. pannonica was implicitly included in Surget-Groba et al. 2006 and both animals belonging to their clade E ("Western Viviparous") and animals belonging to the clade C (Central viviparous I) belong to that subspecies, hence it is not assignable to a any specific clade.

Distribution: Z. v. carniolica and Z. v. vivipara are parapatric in the Alpine chain (Cornetti et al. 2015). Only two areas may have potentially hybridizing oviparous and viviparous populations (Carinthia, Austria; Lindtke et al., 2010, and one in the central Italian Alps, Cornetti et al. 2015).

Conservation: Reptile of the year 2006. Zootoca vivipara has been called the “most successful reptile species of the world”, based on its distribution and the range of climates it inhabits (GUILLAUME et al. 2006).

Type species: Lacerta vivipara LICHTENSTEIN 1823 is the type species of the genus Zootoca WAGLER 1830.

References: see SCHMIDTLER & BÖHME 2011 for an extended bibliography. 
EtymologyNamed after Greek zoon (= animal) and tokos (= birth, child), and Latin viva (= alive) or vivum, -i (= living things), referring to the viviparous mode of reproduction.

Z. v. louislantzi has been named after Louis A. Lantz (1886-1953), French naturalist (see Ineich & Doronin 2017, Ineich et al. 2019).

WERNER'S name carniolica is a Latin feminine adjective meaning "belonging to the Carnian region" ("Krain" in German) i.e., the area where this new taxon seems to have its geographic and historical (réfugial) center. 
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