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Zootoca carniolica (MAYER, BÖHME, TIEDEMANN & BISCHOFF, 2000)

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Higher TaxaLacertidae, Lacertinae, Sauria, Lacertoidea, Squamata (lizards)
Common Names 
SynonymLacerta vivipara carniolica MAYER, BÖHME, TIEDEMANN & BISCHOFF 2000
Zootoca vivipara carniolica — BÖHME & RÖDDER 2006
Zootoca vivipara carniolica — BISCHOFF 2007
Lacerta (Zootoca) vivipara carniolica — SINDACO & JEREMČENKO 2008
Zootoca vivipara carniolica — CORNETTI et al. 2014
Zootoca carniolica — CORNETTI et al. 2015
Zootoca vivipara carniolica — CORNETTI et al. 2018
Zootoca carniolica — SPEYBROECK et al. 2020
Zootoca carniolica — SINDACO & RAZZETTI 2021 
DistributionSlovenia, NE Italy, S Austria (Kärnten: Doberbach Tal, Krainer Alpen), NW Croatia.

Type locality: Mt. Sneznik: 8 km SE Masun village, 1250 m elevation, Slovenia.  
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: ZFMK 68438, paratypes in NMW and MCSNT 
DiagnosisDiagnosis (carniolica): A subspecies of Zootoca vivipara which is distinguished from the nominotypic form by genetic properties (sequence differences between mitochondrial haplotypes) rather than by morphological peculiarities. As a main characteristic, it retains the oviparous reproductive mode, as it is the case in the populations of Spain and south-western France, which are, however, genetically only distantly related (from Mayer et al. 2000). Rodriguez-Prieto et al. 2017 found that males with a ratio of mean ventral scale rows / mean number of femoral pores (MVS/MFP) < 2.20 are unequivocally attributed to Z. carniolica, while males with MVS/MFP higher than 2.27 are unequivocally attributed to Z. v. vivipara; females with a ratio MVS/MFP lower than 2.45 are unequivocally attributed to Z. carniolica, while females with MVS/MFP higher than 2.48 are unequivocally attributed to Z. v. vivipara (”mean number” = average of left and right side per animal; based on 325 specimens). 
CommentSynonymy: partly after SCHMIDTLER & BÖHME 2011. Note that Lacerta (= Zootoca) vivipara carniolica MAYER et al. 2000 is not identical to Lacerta vivipara var. carniolica WERNER, 1897, which is a nomen nudum.

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).

See also Z. vivipara. 
EtymologyWERNER'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. 
  • Bischoff, W. 2007. Auf der Suche nach Gebirgseidechsen in drei Erdteilen. Draco 7 (27): 54-73 - get paper here
  • Böhme, W. & Rödder, D. 2006. Reptil des Jahres 2006: Die eurasische Wald- oder Bergeidechse (Zootoca vivipara), das erfolgreichste terrestrische Reptil der Welt. Reptilia (Münster) 11 (59): 55-60 - get paper here
  • Cornetti, L; GF Ficetola, S Hoban, C Vernesi 2015. Genetic and ecological data reveal species boundaries between viviparous and oviparous lizard lineages. Heredity; doi: 10.1038/hdy.2015.54 - get paper here
  • Cornetti, Luca; Francesco Belluardo, Samuele Ghielmi, Giovanni Giovine, Gentile F. Ficetola, Giorgio Bertorelle, Cristiano Vernesi and Heidi C. Hauffe 2015. Reproductive isolation between oviparous and viviparous lineages of the Eurasian common lizard Zootoca vivipara in a contact zone. Biological Journal of the Linnean Society, doi: 10.1111/bij.12478 - get paper here
  • Cornetti, Luca; Michele Menegon, Giovanni Giovine, Benoit Heulin, Cristiano Vernesi 2014. Mitochondrial and Nuclear DNA Survey of Zootoca vivipara across the Eastern Italian Alps: Evolutionary Relationships, Historical Demography and Conservation Implications. PLoS One 9 (1): e85912. doi:10.1371/journal.pone.0085912 - get paper here
  • Cornetti, Luca; Oliver W Griffith, Andrea Benazzo, Alex Panziera, Camilla M Whittington, Michael B Thompson, Cristiano Vernesi, Giorgio Bertorelle 2017. Candidate genes involved in the evolution of viviparity: a RAD sequencing experiment in the lizard Zootoca vivipara (Squamata: Lacertidae). Zoological Journal of the Linnean Society, zlx069 - get paper here
  • Mayer, W. , W. Böhme, F. Tiedemann & W. Bischoff 2000. On viviparous populations of Zootoca vivipara (Jacquin, 1787) in south-eastern Central Europe and there phylogenetic relationship to neighbouring viviparous and South-west European oviparous populations (Squamata: Sauria: Lacertidae). Herpetozoa 13 (1/2): 59-69 - get paper here
  • SINDACO, ROBERTO & EDOARDO RAZZETTI. 2021. An updated check-list of Italian amphibians and reptiles. Natural History Sciences 8(2): 35–46. - get paper here
  • Speybroeck, Jeroen; Wouter Beukema, Christophe Dufresnes, Uwe Fritz, Daniel Jablonski, Petros Lymberakis, Iñigo Martínez-Solano, Edoardo Razzetti, Melita Vamberger, Miguel Vences, Judit Vörös0, Pierre-André Crochet 2020. Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica.. Amphibia-Reptilia 41: 139-189 - get paper here
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