Amphibia-Reptilia 41 (2020): 139-189 brill.com/amre

Review

Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica

Jeroen Speybroeck1,∗, Wouter Beukema2, Christophe Dufresnes3, Uwe Fritz4, Daniel Jablonski5, Petros Lymberakis6, Iñigo Martínez-Solano7, Edoardo Razzetti8, Melita Vamberger4, Miguel Vences9, Judit Vörös10, Pierre-André Crochet11

Abstract. The last species list of the European herpetofauna was published by Speybroeck, Beukema and Crochet (2010). In the meantime, ongoing research led to numerous taxonomic changes, including the discovery of new species-level lineages as well as reclassifications at genus level, requiring significant changes to this list. As of 2019, a new Taxonomic Committee was established as an official entity within the European Herpetological Society, Societas Europaea Herpetologica (SEH). Twelve members from nine European countries reviewed, discussed and voted on recent taxonomic research on a case-by-case basis. Accepted changes led to critical compilation of a new species list, which is hereby presented and discussed. According to our list, 301 species (95 amphibians, 15 chelonians, including six species of sea turtles, and 191 squamates) occur within our expanded geographical definition of Europe. The list includes 14 non-native species (three amphibians, one chelonian, and ten squamates).

Keywords: Amphibia, amphibians, Europe, reptiles, Reptilia, taxonomy, updated species list.

Introduction

1 - Research Institute for Nature and Forest, Havenlaan 88 Speybroeck, Beukema and Crochet (2010) bus 73, 1000 Brussel, Belgium (SBC2010, hereafter) provided an annotated 2 - Wildlife Health Ghent, Department of Pathology, Bacteriology and Avian Diseases, Ghent University, species list for the European amphibians and Salisburylaan 133, 9820 Merelbeke, Belgium non-avian reptiles. A decade later, a sizable 3 - LASER, College of Biology and the Environment, amount of new research has been produced, Nanjing Forestry University, Nanjing, China 4 - Museum of Zoology, Senckenberg Dresden, A.B. fuelling the need for a contemporary update. Meyer Building, Königsbrücker Landstraße 159, Within the European Herpetological Society 01109 Dresden, Germany (Societas Europaea Herpetologica; SEH) and by 5 - Department of Zoology, Comenius University in Bratislava, Ilkovicovaˇ 6, Mlynská dolina, 842 15 invitation of the SEH Council, a newly com- Bratislava, Slovakia posed Taxonomic Committee (SEH TC, or fur- 6 - Natural History Museum of Crete, University of Crete, ther TC) was formed in early 2019, and its Knossou Ave. 71409, Crete, Irakleio, Greece 7 - Museo Nacional de Ciencias Naturales (MNCN- chair was approved by SEH membership during CSIC), c/ José Gutiérrez Abascal, 2, 28006 Madrid, the Ordinary General Meeting held in Milan, Spain 8 - Kosmos – Museo di Storia Naturale, Università di Pavia, Piazza Botta 9, 27100 Pavia, Italy 11 - CEFE, Université Montpellier, CNRS, EPHE, IRD, 9 - Division of Evolutionary Biology, Zoological In- stitute, Braunschweig University of Technology, Université Paul Valéry Montpellier 3, Montpellier, Mendelssohnstr. 4, 38106 Braunschweig, Germany France ∗ 10 - Department of Zoology, Hungarian Natural History Corresponding author; Museum, 1088 Budapest, Baross u. 13, Hungary e-mail: [email protected]

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September 5th 2019 (SEH News, Amphibia- Venchi and Grieco (2013). Together, these four Reptilia 40: 551-559). sources led to a starting point species list. In the We did not define our own limits for the geo- following, we only discuss taxonomic changes graphical area considered here, but adopted the which deviate from this species list. We decided limits defined by previous projects. Our goal against using online databases as starting points, was to provide a taxonomic reference for the fu- as they are changing constantly, with historical ture mapping projects of the SEH. Therefore, versions not remaining reliably and easily avail- we included all areas that were part of pre- able. vious European atlas projects (cf. Gasc et al., The taxonomic decisions adopted here are 1997; Sillero et al., 2014). We also aimed at in- not necessarily supported by all authors of this forming the taxonomic backbone of the Fauna work. According to TC guidelines, a change to Europaea initiative (https://fauna-eu.org), and the starting list will only be adopted if widely therefore our geographical area also includes all supported among its members, specifically by territories that are covered by Fauna Europaea. a >75% majority. When a change is recom- As a result, we enlarged the geographical area mended by a large majority of the TC mem- considered by SBC2010 to encompass all areas bers, but different members favour different covered by both Gasc et al. (1997) and Fauna outcomes, the adopted solution may be sup- Europaea. Areas included by Gasc et al. (1997), ported by only a simple majority (>50%). Note but not by Speybroeck, Crochet and Beukema that this process favours taxonomic stability, (2010), include the northern versant of the Cau- with changes requiring large support among TC casus (including north-eastern Azerbaijan), all members to become accepted. areas west of the Ural River (including western- TC members do not necessarily adhere to most Kazakhstan) and west of the Ural Moun- the same species concept. While many agree tains, and the Yekaterinburg Region. Areas in- with the General Lineage Concept (GLC) of De cluded in Fauna Europaea, but not by Gasc et Queiroz (2007), some prefer the general frame- al. (1997) or SBC2010, are Macaronesia (with- work of the Biological Species Concept. How- out Cape Verde), the Greek Islands off the west- ever, all agree on using reproductive isolation as ern Anatolian shore, and Cyprus. As such, our the primary operational criterion for the delimi- area exceeds that of the most recent European tation of species. The majority of TC members atlas (Sillero et al., 2014) by including Mac- is of the opinion that, while every species is a aronesia, all Greek islands, and parts of Azer- lineage, not every lineage is a species. The com- baijan and Kazakhstan (fig. 1). A Google Earth mon approach can be defined as either following .kml file with the limits of the area is provided the Biological Species Concept framework, or in the supplementary material. For the rationale as applying a Biological Species Criterion under of these limits, we refer to Gasc et al. (1997) and the GLC. More specifically, TC members ad- https://fauna-eu.org/data-handling). here to a “soft” version of the reproductive iso- Upon enlarging the scope area and prior to lation criterion. As such, we allow extensive in- discussing any taxonomic changes, a broader trogression between recognised species, as long baseline list had to be set for species occur- as there are intrinsic barriers to gene flow that ring outside the area considered by SBC2010. prevent wide-reaching introgression beyond the As such, in addition to SBC2010, we followed contact zones. Even in the absence of geograph- the taxonomy of Gasc et al. (1997, 2004, includ- ical barriers, a sufficient level of reproductive ing the changes adopted by Dubois and Cro- isolation has to exist in order to ensure long- chet in the 2004 reprint), and for species outside term persistence of the diverged lineages. Taxa Europe (including Macaronesia and Cyprus), connected by bimodal or trimodal hybrid zones Sindaco and Jeremcenkoˇ (2008) and Sindaco, (Gay et al., 2008) were unanimously treated as Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 141 lero et al. (2014). Extent of geographic area (Mollweide projection). Herpetofauna species within this area are dealt with. Shading indicates areas not included by Sil Figure 1.

Downloaded from Brill.com10/05/2021 05:01:43PM via free access 142 J. Speybroeck et al. valid species, but opinions often differed re- reptiles. For each case, we provide the rationale garding how much introgression was “allowed” underlying the respective decision of the TC, across unimodal hybrid zones, reflecting differ- and conclude by providing an updated species ent opinions relative to where to cut the grey list of the European herpetofauna. zone of speciation, how much reproductive iso- lation is necessary and when to treat incipient species as valid species. For allopatric taxa, or Amphibia when contact zones were not studied, lineages Caudata/Urodela that had divergence levels similar to closely re- lated, unambiguously distinct species were ac- A series of phylogenetic studies on mitochon- cepted as species. As an auxiliary criterion, we drial DNA, allozymes, and nuclear DNA se- sometimes use monophyly, even though we do quences of members of the family Salamandri- not consider it as a necessary requirement for dae (Litvinchuk et al., 2005; Weisrock et al., species status. 2006; Zhang et al., 2008; Kieren et al., 2018; For supraspecific classification, the TC Veith et al., 2018) confirmed that the newt genus agreed to accept only monophyletic units. Triturus sensu lato, as traditionally recognised, This causes issues regarding the class Rep- is not monophyletic. Litvinchuk et al. (2005) tilia, which in its traditional definition is pa- proposed the separation of Triturus into four raphyletic through the exclusion of birds. All genera, among which the new genus Omma- current hypotheses on the evolution of verte- totriton contains (the former) Triturus vitta- brates agree that a group including squamates, tus. SBC2010 accepted this new arrangement, turtles, Sphenodon and crocodiles, but not birds but did not explicitly acknowledge the need to is paraphyletic (see e.g. Chiari et al., 2012a; recognise Ommatotriton, as the new genus does Hasegawa, 2017). As a consequence, most cur- not occur in the area they considered. As we rent classifications of Vertebrata include Aves in herein consider a wider area, we formally ac- the class Reptilia (see e.g. Modesto and Ander- cept Ommatotriton as a separate genus. son, 2004; Ruggiero et al., 2015). To avoid con- Litvinchuk et al. (2005) also showed that fusion, we adopt the term ‘non-avian reptiles’ to morphology (number of trunk vertebrae, colour refer to the components of the European fauna pattern), genome size, and allozymes (Nei’s assigned to Testudines/Chelonii and Squamata. genetic distances of 0.44-0.83) strongly dif- For nomenclatural decisions, including fer between populations in the two widely dis- spelling, we followed the International Code joint areas inhabited by Ommatotriton vittatus, of Zoological Nomenclature (the Code here- namely south-eastern Anatolia and the Levant after, ICZN (1999 and subsequent changes), (O. v. vittatus) versus the southern and eastern see https://www.iczn.org/). Such decisions were Black Sea and western Caucasus regions (O. v. generally not submitted to voting, but they could ophryticus). This led them to elevate ophryticus be discussed, as many parts of the Code can be to species level. A more comprehensive study subject to interpretation, and many actual cases also demonstrated restricted introgression be- can be open to different decisions, even under tween an eastern and a western taxon along the the rules of the Code. Black Sea coast of Turkey (van Riemsdijk et al., In the following, we review taxonomic and 2017). Consequently, we recommend to recog- nomenclatural changes proposed since the pub- nise three species in the genus Ommatotriton: lication of the four literature sources that we O. vittatus (extralimital) in south-eastern Anato- used to build our starting point, as well as other lia and the Levant, O. ophryticus in Russia, the relevant new information pertaining the taxon- Caucasus and northern Anatolia west to the re- omy of European amphibians and non-avian gion of Samsun, and O. nesterovi (extralimital) Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 143 in Anatolia from Samsun to the Sea of Marmara. result of ancient admixture between T. mace- The populations inhabiting the native Omma- donicus and T. ivanbureschi. We do not entirely totriton range in the European part of Russia be- agree with their interpretation of the Code. As long to O. ophryticus, while the introduced (and they rightfully state, nomina based on geneti- persisting anno 2019; Speybroeck, pers. obs.) cally admixed individuals derived through sev- population in north-eastern Spain is of mixed eral generations of backcrossing are indeed left ancestry, with genetic contribution from both O. in limbo in the Code. Yet, we do not agree that ophryticus and O. nesterovi (van Riemsdijk et the Code should be interpreted as extending pro- al., 2018). visions of Art. 23.8 to nomina based on genet- In addition to mtDNA sequences, Vences et ically admixed individuals, at least not without al. (2014) used sequences from thirteen nuclear clear guidelines of what constitutes admixture loci to improve our understanding of the his- in the sense of the Code. We do, however, agree tory of the genus Salamandra. Their nuclear with Wielstra and Arntzen (2014) that arntzeni, data suggest that Salamandra salamandra con- based on individuals that carry less than 50% tains several deeply divergent lineages whose of alleles derived from the karelinii-complex monophyly relative to S. algira received weak taxon, should not be used as the valid name for support, although this requires confirmation. that taxon. We thus accept T. ivanbureschi as Based on mitochondrial data the subspecies lon- the valid nomen of the taxon of the karelinii- girostris is sister to all other S. salamandra lin- complex that occurs in Europe in the Balkans. eages. However, nuclear genes place it with the With the extension of the geographic range of subspecies morenica. The discordance between SBC2010, T. karelinii sensu stricto also occurs mtDNA and nuclear genes may result from past in our area in Russia, the Crimean Peninsula and introgression and admixture processes. Further- the Caucasus region. more, a clade comprising the subspecies fas- Using mitochondrial and nuclear DNA se- tuosa, bernardezi (including populations at- quences, Sotiropoulos et al. (2007) and Recuero tributed to the subspecies alfredschmidti, whose et al. (2014) investigated the phylogenetic struc- validity was rejected, as it is phylogenetically ture of Ichthyosaura alpestris. They identified nested within several subclades of bernardezi; several deeply divergent lineages that largely Beukema et al., 2016) and gigliolii is recov- correspond to currently recognised subspecies, ered with strong support. While additional data except for the subspecies alpestris, which is are clearly needed, this suggests a high amount further divided into a western and an eastern of evolutionary divergence within S. salaman- main lineage. Interestingly, nuclear sequences dra, and the existence of more than one species from two loci show a nearly complete lack of within S. salamandra cannot yet be fully ruled allele sharing between the eastern and west- out. ern clades, suggesting reproductive isolation be- SBC2010 recognised two species within the tween them. However, no samples were col- former Triturus karelinii: T. arntzeni from the lected close to their contact zones. As the east- Balkan Peninsula, and T. karelinii. Wielstra et ern and western clades should meet in the al. (2013) suggested that the type specimens of Balkans and the southern Carpathian Moun- T. arntzeni are in fact T. macedonicus, which led tains, we follow the recommendation of Re- them to place T. arntzeni in the synonymy of cuero et al. (2014) and await more data on T. macedonicus, and to create the name ivanbu- their level of reproductive isolation before mak- reschi for the taxon of the karelinii complex that ing any taxonomic change. Thus, we do not occurs in the Balkans and in Western Anato- follow Raffaëlli (2018) who (based on the ge- lia. Later, Wielstra and Arntzen (2014) demon- netic data presented in the aforementioned stud- strated that the types of T. arntzeni are in fact the ies) elevated the subspecies apuana, reiseri and Downloaded from Brill.com10/05/2021 05:01:43PM via free access 144 J. Speybroeck et al. veluchiensis to species level, and we note that schmidtleri, its range limits and contact zones such changes would make I. alpestris poly- with other European lineages remain poorly phyletic. We thus retain I. alpestris as a single known. Although data on the contact zone with species for now. L. v. kosswigi in Anatolia support a species-level Pabijan et al. (2017) reconstructed phyloge- divergence between L. v. schmidtleri and the netic relationships within the Lissotriton vul- graecus-kosswigi clade, areas of 100-300 km garis species complex and inferred patterns of inhabited by L. vulgaris s.l. of unknown iden- (historical) gene flow, using 74 nuclear DNA tity separate genotyped L. v. schmidtleri popu- markers from one individual from each of 127 lations from those of L. v. vulgaris and L. grae- locations. Five highly divergent lineages were cus (Pabijan et al., 2017; Wielstra et al., 2018). identified within our focal area: Lissotriton The TC thus feels that more information on con- montandoni and four lineages corresponding tact zones between L. v. schmidtleri and L. v. to the Lissotriton vulgaris subspecies graecus, vulgaris is warranted before the species rank of lantzi, schmidtleri and vulgaris. In spite of clear schmidtleri can be accepted. Finally, L. v. lantzi evidence of past historical introgression, their is endemic to the Caucasus and shows an al- contemporary gene flow is restricted. Therefore, lopatric distribution (Wielstra et al., 2018). Mi- the authors proposed to treat the three former tochondrial data suggests that L. v. lantzi was subspecies as species (Pabijan et al., 2017). Be- the first to diverge from other lineages around tween L. v. vulgaris and the morphologically 3.39 (1.42-5.37) Mya (Pabijan et al., 2015). Yet, diverged L. v. meridionalis regular episodes of nuclear data place L. v. lantzi as sister to L. v. gene flow were identified, thus meridionalis schmidtleri and other eastern and central Eu- was retained at subspecies level. While L. v. ropean Lissotriton, while suggesting that diver- graecus is easily distinguished from other Eu- gence within this group initiated with the split ropean populations by male nuptial character- of the graecus-kosswigi clade (Pabijan et al., istics, both L. v. lantzi and L. v. schmidtleri 2017). In conclusion, the TC recommends to are morphologically cryptic in respect to L. v. treat Lissotriton graecus as a valid species, but, vulgaris (Raxworthy, 1990). Rather than be- as awarding species status to lantzi but not to ing confined to Anatolia, Pabijan et al. (2015, schmidtleri may render L. vulgaris paraphyletic, 2017) showed that L. v. schmidtleri also occurs we prefer to maintain the other taxa, including in Greek and Turkish Thrace and on a number L. v. schmidtleri and L. v. lantzi, as subspecies of Greek islands. Because the sampling gaps be- of L. vulgaris for the time being. tween several of the lineages remained wide, A phylogeographic study based on two the TC has been reluctant to accept all syste- mtDNA markers by Martínez-Solano et al. matic conclusions of Pabijan et al. (2017). The (2006) revealed two major mitochondrial lin- taxon graecus comes into close contact with L. eages of Miocene origin in Lissotriton boscai. v. vulgaris along the northern and eastern bor- One of them is restricted to central and south- ders of its distribution. Although no dense sam- western coastal Portugal, while the other oc- pling has been performed to delineate the con- cupies the remainder of the species range, in- tact zone in detail, and mitochondrial introgres- cluding the type locality of L. boscai. Martínez- sion from southern L. v. vulgaris into graecus Solano et al. (2006) acknowledged that the occurs at its northern range border (Pabijan et two lineages might represent cryptic species, al., 2017), nuclear gene pools of these two taxa but called for additional morphological and appear to remain reciprocally distinct in rela- molecular studies, including data on variation tive close geographical proximity (Pabijan et al., in nuclear DNA markers. Dubois and Raffaelli 2017; Wielstra et al., 2018). We therefore accept (2009) resurrected the nomen Triton maltzani Lissotriton graecus as a valid species. For L. v. Boettger, 1879 for the southwestern lineage in Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 145 the new combination Lissotriton maltzani.Ac- Vörös, Ursenbacher and Jelic´ (2019) used 10 cording to these authors, L. maltzani can be dis- microsatellite loci to investigate patterns of dif- tinguished from L. boscai by its smaller size ferentiation between four Croatian cave popu- and by its paler dorsal coloration, especially in lations of Proteus anguinus. They uncovered females, with less distinct dark spots. Teixeira long-lasting isolation between caves belong- et al. (2015) used DNA sequences of one nu- ing to different hydrogeographic systems, with clear gene and found overall congruence with the most ancient divergence being older than 7 mtDNA genes in terms of sequence divergence Mya. This suggests that some of the evolution- and geographic structure. They, however, also ary lineages within this species might constitute revealed wide areas of admixture and evidence cryptic taxa, possibly of species rank (Vörös, for recombination, suggesting a lack of com- Ursenbacher and Jelic,´ 2019). plete reproductive isolation and the presence of incomplete speciation. In a recent study, Se- Anura queira et al. (2020) used ten microsatellites, one Alytes obstetricans is composed of four sub- mtDNA gene and two single copy nuclear DNA species: A. o. obstetricans, A. o. pertinax, A. o. markers in a cline analysis framework to inves- boscai and A. o. almogavarii. The latter taxon tigate one of the hybrid zones between L. boscai is endemic to Catalonia and adjacent areas in and L. maltzani. The results show evidence for north-eastern Spain and southern France, and partial reproductive isolation between L. boscai it is highly differentiated in allozyme (Arntzen and L. maltzani, with narrow clines (3-28 km) and García-París, 1995; García-París, 1995), consistent with selection against hybrids. We mitochondrial (Gonçalves et al., 2007, 2015) thus recognise L. maltzani as a separate species. and microsatellite markers (Maia-Carvalho et Wake (2012) addressed the taxonomy of al., 2018), and features peculiar osteological the Plethodontidae, advocating to treat Aty- (Martínez-Solano et al., 2004) and bioacous- lodes (comprising Speleomantes genei) and tics characters (Márquez and Bosch, 1995). Pos- Speleomantes (comprising the other European sibly due to ancestral hybridisation or incom- plethodontid species) as subgenera of a single, plete lineage sorting (Gonçalves et al., 2007; cross-Atlantic genus Hydromantes. Addressing Maia-Carvalho et al., 2014), the nuclear phy- the genus name confusion, Wake (2013) com- logeny, based on intron sequences, is not well pared five potential arrangements, missing how- resolved. Using microsatellites, Maia-Carvalho ever the arrangement of SBC2010 (i.e. two et al. (2018) identified a distinct cluster cor- genera without subgenera). Comparing with responding to all A. o. almogavarii popula- other plethodontid genera, he argued in favour tions, suggesting that the lack of monophyly in of an arrangement of a single genus with mtDNA data is due to cyto-nuclear discordance. three subgenera. However, the arguments of This cluster extends as far west as the southern SBC2010 still stand: the position of Atylodes slopes of the Pyrenees in the north-westernmost remains unresolved, and the European species parts of Aragon. These authors also found re- form a well-defined monophyletic group with a stricted genetic admixture between A. o. almo- large genetic distance from the five Californian gavarii and neighbouring subspecies (A. o. per- species Hydromantes brunus, H. platycephalus, tinax, A. o. obstetricans). Following up on this H. samweli, H. shastae, and H. wintu (Nascetti study, Dufresnes and Martínez-Solano (2020) et al., 1996; Pyron and Wiens, 2011; Chiari targeted the hybrid zone between A. o. almo- et al., 2012b; Bingham, Papenfuss and Wake, gavarii and A. o. pertinax with genomic analy- 2018). Thus, no change seems in order, and we ses using RADseq-derived markers along a fine- maintain the European species for the time be- scale transect in Catalonia. They documented ing in a single taxon, the genus Speleomantes. a very narrow mitochondrial (cline width ca. Downloaded from Brill.com10/05/2021 05:01:43PM via free access 146 J. Speybroeck et al.

13 km) and nuclear (cline width ca. 16 km) et al., 2019a, b), we accept the specific rank of transition, and detected portions of the genome Pelobates vespertinus. that were completely impermeable to gene flow. Dufresnes et al. (2019a) also demonstrated Given the absence of barriers to dispersal in the complete reproductive isolation between Pelo- contact zone, the authors concluded that these bates syriacus syriacus and P. s. balcanicus, lineages exhibit substantial (even if incomplete) warranting elevation of European populations, reproductive isolation. We adopt their recom- except those from south-eastern Bulgaria, parts mendation and treat Alytes almogavarii as a dis- of European Turkey and a number of eastern tinct species. Greek islands (including Limnos and Lesbos), Pabijan et al. (2012) found a lack of recipro- to species level as Pelobates balcanicus, with an cal monophyly in nuclear data of Iberian painted estimated Mio-Pliocene divergence (>5 Mya) frogs (Discoglossus spp.), while Dufresnes et between both taxa. Deep intraspecific diver- al. (2020a) found numerous introgressed indi- gence within each species further leads to recog- viduals in their RAD datasets and a broad hy- nise the subspecies P. balcanicus chloeae (Pelo- brid zone (average cline width of nuclear mark- ponnese) and P.syriacus boettgeri (all European ers >136 km). These results indicate weak (or parts of the distribution of Pelobates syriacus). no) restriction to gene flow and confirm that The complex pattern of genetic variation D. g. galganoi and D. g. jeanneae are better within Iberian Pelodytes has been known for treated as conspecific, as previously advocated several years (e.g., van de Vliet et al., 2012). It by SBC2010. was taxonomically formalised by Díaz- Borkin et al. (2001) reported differences in Rodríguez et al. (2017), who described two genome size between the morphologically sim- new species: Pelodytes atlanticus from Por- ilar eastern and western populations of Pelo- tugal and P. hespericus from central eastern bates fuscus, with a transition between both Spain, in addition to the previously established groups in north-eastern Ukraine and adjacent species, P. ibericus and P. punctatus.Thetwo parts of Russia. The name Rana vespertina Pal- new lineages show no consistent morphological las, 1771 is available for the eastern taxon. Sub- or bioacoustics differences, are only weakly dif- sequently, analyses using mtDNA data across ferentiated in mtDNA (<2% in the 16S rRNA the entire species range confirmed the existence gene, <7% in COI), but show limited sharing of two major mitochondrial lineages (Crottini et of nuclear alleles. Díaz-Rodríguez et al. (2017) al., 2007). Litvinchuk et al. (2013) confirmed recognised that this was a disputable case, in that these two lineages differ in their nuclear which, for the time being, “species status for genomes, with Nei’s genetic distance based on all four western lineages of Pelodytes [was] the allozymes being similar to the divergence be- hypothesis best fitting the available data”. A tween Pelobates syriacus syriacus and P. s. bal- new study by Dufresnes et al. (2020a), based on canicus (see below), and with a restricted intro- RADseq-derived loci from spatially dense sam- gression zone. Based on its independent evolu- pling, found that P. atlanticus and P. ibericus tionary history, they thus proposed to treat ves- are separated by a narrow hybrid zone, featur- pertinus as a species. More recently, based on ing little introgression, whereas the contact zone RADseq data, Dufresnes et al. (2019a) reported of P. hespericus and P. punctatus has a consid- narrow clines between fuscus and vespertinus erably more flattened cline, with an introgres- (average nuclear cline width: 16 km). On the ba- sion zone possibly extending up to 200-300 km. sis of the narrow hybrid zone, indicating strong There is no known contact zone between P. a t - (albeit incomplete) reproductive isolation, and lanticus and P. hespericus. While the position in spite of the relatively recent divergence of of P. atlanticus (sister to ibericus or to hesperi- this taxon (between 2 and 3 Mya, see Dufresnes cus) remains unresolved, its genetic divergence Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 147 from hespericus and punctatus is quite large. and most French populations of Bufo bufo (for We thus follow the suggestion of Dufresnes which the name Bufo spinosus is available), et al. (2020a) to accept P. atlanticus as sepa- and a clade composed of two lineages, repre- rate species, while retaining hespericus at sub- senting ii) verrucosissimus from the Caucasus species level as P. punctatus hespericus. and iii) bufo from northern France to Russia, Despite growing support for recognition of while populations from Greece, southern Italy various clades in the family Bufonidae as sepa- and Sicily and most of Anatolia carried bufo rate genera (Stöck et al., 2006; Van Bocxlaer et mtDNA but grouped with verrucosissimus in al., 2009), several authors advocated retaining nuclear DNA. Estimations of divergence times Bufo for all species from the Western Palaearc- indicated a long evolutionary history of the tic and Central Asia, at least as an ad interim group, starting with the split from eichwaldi at solution (Dubois and Bour, 2010; SBC2010). about 12 Mya, and the divergence of spinosus Formal taxonomic action was further hampered taking place around 6 Mya. The deep level of by nomenclatural confusion, such as the use genetic divergence observed between the west- of either Pseudepidalea or Bufotes as the valid generic nomen for the green toad group (Frost ern and eastern groups of common toads indi- et al., 2006; Dubois and Bour, 2010). Dubois cated that these groups may be different species. and Bour (2010) demonstrated that Pseudep- However, Garcia-Porta et al. (2012) found ex- idalea is a junior objective synonym of Bu- tensive admixture of mitochondrial lineages be- fotes, thereby giving priority to the latter. How- tween the eastern and western clades in the ever, based on the presence of hybridisation Languedoc area of southern France (as also re- between representatives of these clades, and ported by Arntzen et al., 2017 for the Provence the restricted sampling size of previous stud- area of southern France), and detected signs of ies, Dubois and Bour (2010) listed these nom- ancient introgression of bufo allozyme alleles ina as subgenera, rather than genera. Accord- into spinosus. They thus suggested to treat B. b. ing to Van Bocxlaer et al. (2010), Pyron and spinosus and B. b. bufo as conspecific pending Wiens (2011), and Beukema et al. (2013), which studies of the contact zones. Detailed analysis of together provide a dense sampling of bufonid the amount of reproductive isolation in two geo- species, most genera currently recognised by graphically distant contact zones on the basis of Frost (2019) represent well-supported mono- mtDNA, morphology and nuclear markers have phyletic units. Moreover, time-calibrated analy- subsequently been published by Arntzen et al. ses showed Bufo, Bufotes and Epidalea to be of (2016, 2017). Both studies documented an ex- similar age as or older than most other recog- tensive amount of hybridisation and introgres- nised bufonid genera (Beukema et al., 2013). sion in the contact zone, but also the presence Considering these results, we accept Bufo, Bu- of narrow and concordant clines for most nu- fotes and Epidalea at genus level. clear markers, resulting in a unimodal, yet nar- Recuero et al. (2012) published a multilocus mitochondrial and nuclear DNA sequence data row hybrid zone of ca. 30 km wide, and indi- set for Bufo bufo and associated species, cover- cating intrinsic barriers to gene flow in spite of ing the entire documented range and providing incomplete reproductive isolation. Karyological extensive genetic data. The study yields a fully analysis (albeit based on only four males and resolved phylogeny, with the recently described a single female) identified heteromorphic sex Bufo eichwaldi from the Talysh Mountains of chromosomes in spinosus, but not in bufo (Sko- southern Azerbaijan and northern Iran as the rinov et al., 2018). The combined available ev- sister taxon of a clade including three deeply idence justifies treating Bufo spinosus as a dis- diverged lineages: i) north African, Iberian, tinct species. Downloaded from Brill.com10/05/2021 05:01:43PM via free access 148 J. Speybroeck et al.

Based on differences in morphology and Stöck et al., 2006) and demonstrated strong re- karyotype and a study suggesting lowered fertil- productive isolation between them. As these ity of F1 hybrids (summary in Kuzmin, 1999), two taxa belong to the two most divergent lin- the Caucasian populations of the complex are eages (North African and Eurasian), a two- sometimes also treated as a distinct species as species split of the green toad complex is clearly Bufo verrucosissimus. Recent genetic studies warranted. The Sicilian taxon siculus is closely paint a more complex picture. Firstly, based on related to the North African taxon boulengeri, nuclear markers, the Anatolian populations, tra- which has priority over siculus. Subsequently, ditionally excluded from B. b. verrucosissimus nuclear data were added to the picture (mi- on the basis of morphology, are in fact geneti- crosatellites – Dufresnes et al., 2018a; Gerchen, cally closer to this group than to B. b. bufo or B. Dufresnes and Stöck, 2018, and RADseq mark- spinosus, even if they carry B. b. bufo mtDNA ers – Dufresnes et al., 2019c). Although cyto- (Garcia-Porta et al., 2012; Arntzen et al., 2013). nuclear discordance was present in large areas Secondly, mtDNA divergence between Cau- of Europe where populations of viridis carry casian (B. b. verrucosissimus) and Eastern Eu- the variabilis mtDNA lineage, these studies ropean (B. b. bufo) populations is much lower confirmed that overall the previously identified than between B. b. bufo and B. spinosus.In- mtDNA lineages correspond to distinct evolu- tionary units. Dufresnes et al. (2019c) also dis- deed, divergence in mtDNA and allozymes be- covered that the type locality of variabilis is in- tween the Caucasian and Eastern European lin- habited by the western lineage, rendering vari- eages is even lower than between the subclades abilis a junior synonym of viridis, and that the of B. spinosus (Garcia-Porta et al., 2012). Most valid name for the Anatolian lineage is sitibun- importantly, based on frequencies of allozyme dus. These two lineages (B. v. sitibundus and B. alleles, populations of B. b. bufo from Greece v. viridis) widely admix over a very large geo- and north-western Turkey appear intermediate graphic area in Anatolia and Russia (Dufresnes between Caucasian and European populations et al., 2019c), suggesting that they are best (Garcia-Porta et al., 2012) or group with Cau- treated as subspecies. The contact zone between casian populations in nuclear trees (Recuero et B. v. viridis and B. v. balearicus is narrower al., 2012), suggesting extensive introgression (63 km), but still extensive if compared with between B. b. verrucosissimus and B. b. bufo in other contact zones between taxa that are here north-western Turkey and the southern Balkans treated as different species (Gerchen, Dufresnes (but see Arntzen et al., 2013). In our opinion, and Stöck, 2018). In addition, raising B. v. the specific status of B. b. verrucosissimus is balearicus but not B. v. sitibundus to species currently insufficiently supported by the avail- rank would make B. v. viridis paraphyletic (see able evidence, and we maintain it as conspecific Dufresnes et al., 2019c). The TC therefore pre- with Bufo bufo for the time being. ferred to treat B. v. sitibundus and B. v. baleari- Stöck et al. (2006) elevated a number of mito- cus (and the extralimital B. v. perrini) as sub- chondrial lineages of green toads (now Bufotes) species of B. viridis. to species level (for Europe: B. variabilis and Dufresnes et al.’s (2019c) range-wide sam- B. balearicus, in addition to B. viridis) and sub- pling of all known taxa of the Bufotes viridis sequently described a new species from Sicily complex identified a total of eight diploid ge- based on the same lines of evidence. SBC2010 netic clusters, including a new lineage endemic argued against accepting these species on the to Cyprus, which they describe as Bufotes cy- basis of mitochondrial DNA alone. Colliard et priensis. Estimated to be of Messinian origin al. (2010) documented a contact zone in Sicily (5.3 Mya), this insular taxon is as old as or between the taxa siculus and balearicus (sensu older than many species of anuran amphibians Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 149 from the Western Palearctic. Yet, all ten indi- of other supporting characters (weak acous- viduals from four localities carry mtDNA hap- tic divergence: see Schneider, 1974; no well- lotypes of B. v. sitibundus. Secondary contact supported morphological characters), SBC2010 and hybridisation with the Anatolian mainland refrained from formally accepting this syste- lineage during the Pleistocene would explain matic treatment, as it would have rested solely this mitochondrial capture, as well as the rem- on mitochondrial DNA data. Several more re- nants of nuclear introgression that were detected cent studies offer a better understanding of as well. In addition to this deep genomic di- the phylogeny of this group and of the pat- vergence, the genome of B. cypriensis is also terns of gene flow across contact zones. Firstly, significantly larger than that of all other West- Dufresnes et al. (2015, 2016) investigated the ern Palearctic Bufotes taxa. Morphologically, arborea-orientalis contact zones in Poland and the Cyprus green toads are smaller than those in the Balkans with microsatellite markers. In from mainland populations, as previously re- Poland, they found evidence of admixture over a ported by Stugren and Tassoula (1987). The 200 km wide zone, with mosaic contacts and in- combination of an old divergence and a larger terspersed hybrid populations, but with strongly genome size led the TC to accept the species sta- restricted introgression at sex-linked loci and tus of B. cypriensis. Adriatic populations could many populations of seemingly pure ancestry in close contact to each other (Dufresnes et al., represent a valid subspecies of B. viridis (for 2016). In the Balkans, Dufresnes et al. (2015) which the nomen longipes Fitzinger in Bona- found narrow clines (30 and 32 km of aver- parte, 1840 might be available), while green age cline width in Serbia and Greece respec- toads from Naxos (Cyclades) and Crete dif- tively). In line with our treatment of taxa con- fer genetically from other populations and de- nected by unimodal hybrid zones with narrow serve genomic investigations (Dufresnes et al., clines, we recommend affording species status 2019c). In summary, for Europe, we accept (1) to Hyla orientalis. Secondly, the phylogenomic Bufotes viridis, including the widespread B. v. tree in Dufresnes et al. (2018b) unambiguously viridis, B. v. balearicus (Italian Peninsula, Cor- groups orientalis and molleri as sister taxa, with sica, Sardinia, Sicily, Balearic islands) and B. v. arborea splitting from a node basal to the mol- sitibundus (within Europe restricted to a num- leri – orientalis divergence. Therefore, we ac- ber of eastern Greek islands), (2) the Sicilian cept molleri at species rank as well. Although Bufotes boulengeri siculus and (3) Bufotes cy- the close relationships between molleri and ori- priensis. We note that the availability of the entalis could justify to treat them as conspecific, nomen siculus remains unclear, given that the we refrain for the time being from this arrange- corresponding description (Stöck et al., 2008) ment. We thus treat Hyla molleri and Hyla ori- was only issued electronically and prior to 2011 entalis as valid species. (see Article 8.5 of the Code): whether “numer- Supported by mtDNA and genomic data, two ous identical and durable copies” (Article 8.1) major cryptic lineages reside within Hyla inter- were registered by the authors in parallel is yet media (Canestrelli, Verardi and Nascetti, 2007; to be addressed (see also Dubois et al., 2013). Stöck et al., 2008; Dufresnes et al., 2018b). Based on a high level of divergence in mi- They are not known to differ in any diagnos- tochondrial DNA sequences of Hyla tree frogs, tic morphological or acoustic character, even Stöck et al. (2008) have suggested to recog- if they differ in the averages of one acoustic nise the Iberian taxon molleri and the eastern and some morphometric traits. The northern lin- European taxon orientalis as species distinct eage occupies the Po Plain and adjacent re- from Hyla arborea. However, given the lack gions (including Ticino in Switzerland), with Downloaded from Brill.com10/05/2021 05:01:43PM via free access 150 J. Speybroeck et al. the northern Apennines acting as the biogeo- 2013b, 2017). In a phylogeographic survey us- graphical barrier separating it from the south- ing genome-wide data, Dufresnes et al. (2020b) ern lineage. As the name intermedia applies to found one of these mitochondrial groups to be a the southern lineage, Dufresnes et al. (2018b, “ghost lineage”, not differentiated in the nuclear c) coined a new name, Hyla perrini,forthe genome. The two main lineages are strongly dif- northern lineage. Note that, as its registration ferentiated and are estimated to have diverged in the Official Register of Zoological Nomen- around 4 Mya. Despite the absence of geo- clature (ZooBank) occurred later than the pub- graphic or ecological barriers to dispersal, they lication of Dufresnes et al. (2018b), the name form a narrow hybrid zone (25 km) in the east- perrini is not made available by Dufresnes et al. ern Cantabrian Mountains. Because partial re- (2018b), but by Dufresnes et al. (2018c), who productive isolation is thus likely to prevent met all conditions of nomenclatural availabil- these two taxa from merging, we follow the ity. Detailed analyses of the contact zone based recommendation of Dufresnes et al. (2020b) to on genomic (RADseq) data by Dufresnes et al. treat R. parvipalmata as a distinct species. (2018b) revealed broad clines (96 km of average cline width) for nuclear markers and detectable Using allozymes and mtDNA, a detailed admixture over approximately 130 km. Despite analysis of gene flow patterns across the con- a relatively high cyt b distance of around 9%, tact zone of the northern Anatolian lineage and and even though Dufresnes et al. (2018b) ar- the Balkan lineage of the Pelophylax ridibun- gued that the extent of the contact zone would dus – bedriagae complex in northern Greece re- be even larger without some form of selection vealed the existence of a wide hybrid zone, with against hybrids, the TC felt that the observed ex- introgression detectable over more than 200 km tent of introgression and thus the lack of strong (Hotz et al., 2013). This suggests that the Eu- reproductive barriers did not unambiguously al- ropean and Anatolian lineages are conspecific low treatment of perrini at species rank. We thus (contra SBC2010). Since the European lineage recommend to treat the northern lineage as Hyla splits from a node basal to the diversification of intermedia perrini. Anatolian lineages with respect to mtDNA ac- The authorship of the family Ranidae has cording to Plötner et al. (2012), it further sug- been clarified by Dubois and Bour (2011), at- gests that most lineages of this complex are tributing it to Batsch, 1796. conspecific, with the possible exception of the Based on allozyme data (Arano, Esteban and populations from Syria and Jordan (possibly the Herrero, 1993; Veith et al., 2002, 2012), two ge- ‘true’ bedriagae, see Plötner et al., 2012). On netically distinct groups have been recognised the other hand, Plötner et al. (2010) suggested within Rana temporaria populations of northern hybrid breakdown between some of these lin- Spain. The first was assigned to the subspecies eages, indicating incipient speciation. While un- R. t. parvipalmata and is restricted to the west- doubtedly the last word on this subject has not ernmost edges of the Iberian distribution (Gali- cia and Asturias). Frogs from western Galicia been written, the TC felt that it was prema- feature reduced feet webbing, smaller size and ture to suggest taxonomic changes, especially lower number of pulses per call (Vences, 1992). as a large taxonomic paper based on genomic The second group corresponds to the nominal variation patterns is in preparation (G. Mazepa, taxon R. t. temporaria, which extends through- pers. comm.). The same applies to the newly out Europe. Mitochondrial data suggested a proposed P. cypriensis, which was found to be more complex picture, with four deeply differ- placed between the European and the Anato- entiated lineages in the same area (Vences et al., lian lineages in mtDNA, but grouped with P. Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 151 cretensis (albeit with very low support) in nu- cannot be explained by a geographic barrier ef- clear DNA (Plötner et al., 2012). For the mo- fect of this narrow sea strait, because in E. or- ment, we thus suggest no change to the taxon- bicularis recent gene flow occurred across the omy adopted in SBC2010 for European Pelo- Strait of Gibraltar (Velo-Antón et al., 2015), phylax. and possibly across the Adriatic Sea between southern Italy and the Balkans (Vamberger et al., 2015). In contrast, Pereira, Teixeira and Reptilia Velo-Antón (2018), using seven microsatellite loci, concluded that the Strait of Gibraltar cur- Testudines/Chelonii rently impedes gene flow between Iberian and North African pond turtles. However, they show Based on nuclear genomic ISSR fingerprints that the break between the two clusters corre- and mtDNA sequences, Fritz et al. (2005) de- sponds to the Central Iberian Mountains and scribed Emys trinacris as a distinct species en- not the sea strait. Clines of microsatellites for demic to Sicily and Calabria, albeit the record the trinacris – orbicularis contact zone across from Calabria was later questioned (Vamberger theStraitofMessinawereinferredtobevery et al., 2015). A phylogenetic study using seven wide (247 km) in Vamberger et al. (2015), al- nuclear genes (Spinks and Shaffer, 2009) found though the significance of this estimate is ham- E. orbicularis to be paraphyletic with respect to pered by a substantial (150 km) sampling gap trinacris: E. o. hellenica was sister to a clade between the two taxa. The TC takes therefore containing trinacris and the remaining lineages a conservative stance and treats trinacris as a of E. orbicularis. This led SBC2010 to conclude subspecies of E. orbicularis, waiting for further that trinacris should not be recognised as a studies resolving the complicated relationships species, which has been criticised by Vamberger of this complex. In this context, we note that and Fritz (2018). While trinacris is the most Pöschel et al. (2018) reported a sharp transition basal lineage in mtDNA trees (Fritz et al., 2005, between occidentalis and orbicularis + gal- 2007), this position is weakly supported, and the loitalica in north-eastern Spain, matching that level of mtDNA divergence between trinacris of distinct species recognised here, suggesting and the other lineages is not clearly larger than that the E. orbicularis complex could comprise that between some lineages of E. orbicularis. several species. In the meantime, we recom- A first combined analysis of genetic differen- mend to maintain the Sicilian lineage as Emys tiation at eight microsatellite loci and mtDNA orbicularis trinacris. sequences found concordant patterns for both markers: no evidence for admixture between Squamata Sicilian pond terrapins and the remaining lin- The family Agamidae is sometimes credited to eages was found, while extensive admixture was Fitzinger, 1826 or Gray, 1827 (see e.g. Melville found between some other lineages of E. orbic- and Smith, 1987). In fact the nomen was first ularis (Pedall et al., 2011). Using 15 microsatel- published by Spix in 1825 as “Familia Aga- lite loci and mtDNA sequences, Vamberger et mae”. Even if Spix did not use the correct suf- al. (2015) found evidence for limited admixture fix ‘-idae’, Article 11.7.1.3 of the Code states between the two taxa in one Sicilian population. that “a family-group name of which the family- In addition, Vamberger et al. (2015) reported group name suffix is incorrect is available with concordant clines for both markers, with cline its original authorship and date, but with a cor- centres matching the Strait of Messina. Vam- rected suffix”. Other conditions of availability berger et al. (2015) argue that the coincidence all seem to be fulfilled in Spix (1825), who is of the cline centres with the Strait of Messina thus the author of the nomen. Downloaded from Brill.com10/05/2021 05:01:43PM via free access 152 J. Speybroeck et al.

Macey et al. (2000a) sequenced a 1685-1778 spelling is not an available name. If one fol- bp long segment of mtDNA (including the ND1, lows this and Art. 11.7.1.1 (“a family-group ND2 and COI genes) to assess phylogenetic name when first published must [...] be a noun relationships of acrodont lizards. Their results in the nominative plural formed from the stem suggested that the agamid genus Laudakia is of an available generic name”), a family-group paraphyletic, yet, low bootstrap support pre- name based on an incorrect subsequent spelling vented definite conclusions. Subsequent stud- of an available genus name is not available. ies based on mitochondrial and nuclear genes Yet, the Code also states that “a family-group (ND2, RAG1) by Melville et al. (2009) and name is an incorrect original spelling and must Edwards and Melville (2011) recovered Lau- be corrected if it is formed from an incorrect dakia as monophyletic with high support. Baig subsequent spelling of a generic name” (Art. et al. (2012) summarised the results of the afore- 32.5.3.3) and that “a family-group name based mentioned studies in a morphology-based revi- upon [...] an incorrect spelling of the name of sion of Laudakia. Despite failing to find distinct the type genus must be corrected” (Art. 35.4.1). morphological variation within the genus, and It is thus clear that the Code has no inten- acknowledging that Melville et al. (2009) and tion to make a family-group name unavailable Edwards and Melville (2011) recovered Lau- based on an incorrect spelling of the name of the type genus (see also Dubois, 2010). In con- dakia as monophyletic, Baig et al. (2012) par- clusion, the family-group name Camaelonia is titioned Laudakia into three genera acknowl- available as published by Rafinesque (1815), but edging its potential paraphyly (Macey et al., its spelling needs to be corrected. The nowadays 2000a). This taxonomic act was subsequently prevailing spelling Chamaeleonidae should be criticised by Pyron, Burbrink and Wiens (2013), preserved (Art. 29.5), with its authorship at- who confirmed the monophyly of Laudakia us- tributed to Rafinesque (1815) instead of Gray ing a supermatrix approach. Within our focal (1825). area, the classification proposed by Baig et al. SBC2010 adopted the inclusion of Cyrtopo- (2012) would affect Laudakia stellio and L. cau- dion kotschyi in the well-supported mono- casia, as these authors placed the former species phyletic genus Mediodactylus, as proposed by in the newly erected genus Stellagama, and the Macey et al. (2000b), Cervenka,ˇ Frynta and latter into Paralaudakia. While these genera Kratochvíl (2008) and Bauer et al. (2013). were rapidly adopted by the wider herpetolog- By enlarging the focal area of SBC2010, we ical community, we do not follow the split of hereby also include the former C. russowi, Laudakia, pending substantial evidence to reject now Mediodactylus russowii. Another species its monophyly, and therefore retain L. stellio and occurring in the area considered by Gasc L. caucasia in Laudakia. et al. (1997) but not in that of SBC2010 The family Chamaeleonidae is sometimes is the former Cyrtopodion caspium.Wefol- credited to Gray, 1825 (e.g. Glaw, 2015). How- low Bauer et al. (2013) in recognising the ever, Rafinesque (1815) published the name well-resolved genus Tenuidactylus and list this “Famille CAMÆLONIA”, based on the genus species as Tenuidactylus caspius. Furthermore, “Camæleo Daud.”, which is an incorrect subse- an introduced, well-established population of quent spelling of the available genus Chamaeleo Tenuidactylus fedtschenkoi has recently been re- Laurenti, 1768. The Code is somewhat am- ported from the city of Odessa (Ukraine). The biguous regarding the availability of family- native range of T. fedtschenkoi lies in central group nomina based on an incorrect subsequent Asia (western Pamiro-Altay mountains), and it spelling of an available genus. Article 19.1 of was probably transported passively to Odessa the Code states that an incorrect subsequent (Duz’, Kukushkin and Nazarov, 2012). Not long Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 153 after the record from Odessa, a revision of delimitation of the ranges and contact zones of the genus Tenuidactylus led to the description these new species requires further investigation of a new species from Uzbekistan and Turk- (e.g. M. danilewskii and M. orientalis), we ac- menistan, which was split from T. fedtschenkoi cept these species here. The following species and named T. bogdanovi (Nazarov and Po- are thus recognised in our area: M. kotschyi yarkov, 2013). Krasylenko and Kukushkin (mainland Balkans, most of Aegean islands, and (2017) provided an update on the status of the Italy (Apulia)), M. bartoni (Crete and nearby non-native Odessa population and assigned it to islets), M. danilewskii (Black Sea region and T. bogdanovi. south-western Anatolia), M. oertzeni (southern Within Mediodactylus kotschyi, many sub- Dodecanese Islands), and M. orientalis (Levant, species have traditionally been recognised. As Cyprus, southern Anatolia, and south-eastern indicated by Kasapidis et al. (2005), the ge- Aegean islands). netic substructure of Mediodactylus kotschyi The endemic gecko from the Selvagens Is- shows a high degree of divergence, suggest- lands has variously been treated as the valid ing that M. kotschyi represents a species com- species Tarentola bischoffi (e.g. Rebelo, 2008; plex. Nearly range-wide data (with limited ar- Sindaco and Jeremcenko,ˇ 2008; Uetz, Freed and eas in eastern and northern Anatolia excluded) Hošek, 2019), or as a subspecies of the Ca- of three mtDNA and three nuclear DNA frag- narian T. boettgeri as T. boettgeri bischoffi (e.g. ments allowed unravelling the evolutionary his- Carranza et al., 2000; Pleguezuelos, Márquez tory of Mediodactylus kotschyi (Kotsakiozi et and Lizana, 2002; Gübitz, Thorpe and Malho- al., 2018). Divergence dates back to 15 Mya, tra, 2005; Sá-Sousa et al., 2009). Its phyloge- and several of the main lineages show overlap- netic affinities were examined by Carranza et ping distribution areas. Divergence in mtDNA al. (2000, 2002) and Gübitz, Thorpe and Mal- is known to be often quite large in geckos (e.g. hotra (2005). The genetic data available (al- Nagy et al., 2012). However, distances between beit based only on mtDNA) show that the mi- the main lineages are particularly large in this tochondrial lineages of bischoffi are closely re- case (>10% 16S and >15% cyt b and COI lated to those of T. boettgeri, especially to the p-distances). In addition, nuclear data groups subspecies T. b. hierrensis from El Hierro, and specimens in concordance with their mtDNA lineage, and not with their geographical ori- are in fact nested inside the mitochondrial di- gin, which suggests a high level of repro- versity of T. b. boettgeri from Gran Canaria, ductive isolation between them. The proposed suggesting that the Selvagens Islands were re- splits also largely agree with morphological cently colonised from El Hierro or Gran Canaria data (Štepánek,ˇ 1937, 1939; Szczerbak and Gol- (Carranza et al., 2000, 2002; Gübitz, Thorpe ubev, 1996). The narrow contact between M. and Malhotra, 2005). Treating bischoffi as a orientalis and M. danilewskii in the Western valid species, while retaining hierrensis as a Taurus Mountains, identified by Kotsakiozi et subspecies of boettgeri (as done by the above al. (2018), corresponds to a lack of morpho- authors who rank bischoffi as a species), is logical intergrades (despite the narrow gap be- thus likely to render Tarentola boettgeri para- tween the subspecies danilewskii and cilicien- phyletic. The amount of mitochondrial diver- sis in the same area, see Rösler, Schmidtler and gence between hierrensis and bischoffi is also Moravec, 2012), while these new species are smaller than within the populations of T. de- very close to the M. kotschyi groups defined by lalandii from Tenerife or between the popula- Beutler (1981). Five species were recognised, tions of T. angustimentalis from Fuerteventura for all of which previously existing names are and Lanzarote. Based on the available data, we available (Kotsakiozi et al., 2018). While the thus recommend to treat the geckos from the Downloaded from Brill.com10/05/2021 05:01:43PM via free access 154 J. Speybroeck et al.

Selvagens Islands as conspecific with the popu- distribution, indicate that the three lineages cor- lations from El Hierro and Gran Canaria, as Tar- respond to three independent species. The name entola boettgeri bischoffi. of the eastern species is often spelled P. ed- Sánchez-Vialas et al. (2018) re-examined the wardsianus. As shown by Crochet (2015), this description of Algyroides hidalgoi Boscá, 1916, is an incorrect subsequent emendation that does a nomen for which the holotype has been lost, not meet the requirements of the Code. As a to settle its position in the synonymy of the consequence, the valid spelling of the eastern genus Algyroides. They argued that the charac- species is Psammodromus edwarsianus. Thus, ters of the holotype, said to originate from the we add Psammodromus edwarsianus and P. oc- Sierra de Guadarrama and described by Boscá, cidentalis to the list of the European herpeto- fall within the morphological variability of Al- fauna species. gyroides marchi Valverde, 1958, and designated A 9 Mya split marks the divergence of Timon a specimen of A. marchi as neotype of A. hi- lepidus nevadensis from the nominal subspecies dalgoi. This would make A. marchi a junior T. l. lepidus (Miraldo et al., 2013). After studies subjective synonym of A. hidalgoi. Because the on genetic (Paulo et al., 2008; Miraldo et al., conditions for automated reversal of precedence 2011, 2013) and morphological differentiation are not met (Art. 23.9.1 of the Code), A. hi- (Mateo and Castroviejo, 1990; Mateo, López- dalgoi would become the valid nomen of the Jurado and Guillaume, 1996), mtDNA and mi- Spanish Algyroides. Several TC members felt, crosatellites were used to investigate gene flow however, that the interpretation of the descrip- patterns in a zone of secondary contact (Miraldo tion of Boscá (1916) by Sánchez-Vialas et al. et al., 2013). While hybridisation and introgres- (2018) left room for doubt, and that Boscá may sion were observed, gene flow was shown to be not have described a specimen of Spanish Algy- restricted. The cline width for nuclear markers roides when he created the name Algyroides hi- was estimated at around 10 km (although with dalgoi. If so, this could affect the validity of the a sampling gap of around 20 km, it may actu- neotype designation, respective to Art. 75.3.5 of ally be less). Furthermore, mostly pure popu- the Code, and result in the unnecessary change lations are present on either side of the sam- of the well-established and widely used name A. pling gap. Considering this together with the marchi. Although opinions in the TC were di- aforementioned old genetic divergence, we ac- vided about this, we recommend to maintain for cept the proposal of Miraldo et al. (2013) and the time being the use of A. marchi, in anticipa- treat T. nevadensis as a valid species. tion of an upcoming application to the Commis- Lacking range-wide sampling and adequate sion, which would maintain marchi in use until molecular analysis, the taxonomy of the Lac- the Commission has ruled on the case. erta trilineata-pamphylica complex has re- Morphological and molecular (mtDNA and mained unresolved, until mitochondrial phylo- nuclear) data support the split of Psammod- genies showed the eastern Anatolian species L. romus hispanicus into three distinct lineages pamphylica to be nested within trilineata (God- (Fitze et al., 2011, 2012; Mendes et al., 2017). inho et al., 2005; Ahmadzadeh et al., 2013; While not all areas of potential contact have Sagonas et al., 2014). Remarkably, Sagonas et been sampled (despite a recent distribution up- al. (2014) found evidence that central Aegean date by Molina et al., in press), the available populations (L. t. citrovittata) are closely related data seem to warrant accepting these lineages to L. pamphylica. Yet, this was poorly supported as distinct species. Age estimates, lineage al- across analyses and the biogeographically sur- lopatry, the lack of mitochondrial and nuclear prising relationship between L. t. citrovittata haplotype sharing between lineages, bioclimatic and L. pamphylica could reflect a methodologi- niche divergence, and the current biogeographic cal artefact (long branch attraction). Thus, the Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 155 monophyly of L. trilineata could not be re- phylogeography of the Lacerta viridis com- jected. More recently, analyses of SNPs and mi- plex. Four main mtDNA lineages, whose phy- tochondrial sequences by Kornilios et al. (2019, logenetic relationships were weakly resolved, 2020) found a sister-group relationship between emerged from the data: (i) Lacerta bilineata, (ii) pamphylica and eastern Aegean populations of Lacerta viridis s.s., (iii) an Adriatic and west- trilineata. This led to the identification of four ern Balkan lineage, and (iv) a newly discovered species-level units: L. trilineata (eastern Adri- lineage from the south-eastern Balkans as well atic Coast, Greece except northeast and includ- as the Turkish Black Sea Coast. A previously ing Milos archipelago and Crete, south-eastern supposed (Amann et al., 1997) contact zone be- Bulgaria and North Macedonia), L. pamphylica tween L. bilineata and L. viridis in north-eastern (southern Anatolia east of Antalya), L. citrovit- Italy was found to be in fact a contact zone be- tata (central Aegean islands, including among tween the Adriatic and western Balkan lineage others Naxos, Tinos, Andros, Syros, Mykonos, and L. bilineata. Another contact zone between Paros), and L. diplochondrodes (north-eastern the Adriatic and western Balkan lineage and L. Greece, Romania, Bulgaria except southeast, viridis s.s. lies further east in Slovenia and ad- Turkey, eastern Aegean Greek islands). All ev- jacent Croatia. As a consequence, there is no idence (morphology, blood biochemistry, im- known geographical contact zone between L. munoserology, congruence between highly di- viridis s.s. and L. bilineata. Several other con- vergent mtDNA and nuclear genomics, biocli- tact zones between mitochondrial lineages were matic niche differences, no signs of admixture) identified across the distribution range. How- suggests that L. pamphylica deserves species ever, the nuclear data were insufficient to exam- status. As the eastern Aegean populations of the ine gene flow among lineages. Consequently, no complex are the sister taxon of L. pamphylica, species-level changes were suggested. The pre- maintaining monophyly implies also treating viously reported occurrence of L. bilineata on them as a species, L. diplochondrodes. The lat- Cres, Croatia (Brückner et al., 2001), was con- ter taxon does not show a lot of contact, nor firmed. However, this record is isolated from signs of introgression with any of the other three the remaining distribution range and lies among clades. Concerning the population west of the Aegean Barrier, the conclusion may seem less surrounding records of the Adriatic and western straightforward. However, genomic analyses of- Balkan lineage (Marzahn et al., 2016), sugges- fer stronger support for the species status of L. tive of an introduced population. citrovittata than for that of L. pamphylica.The Speybroeck and Crochet (2007) and former equally corresponds to one of the ma- SBC2010 adopted the split of the former genus jor mtDNA lineages of Kornilios et al. (2019). Lacerta s.l., accepting all genus names recog- Given its allopatry, there is no evidence of ad- nised by Arnold, Arribas and Carranza (2007), mixture. While the (potential) contact zone be- as far as they pertained to taxa occurring within tween L. diplochondrodes and L. trilineata in their considered area. As we have adopted a north-eastern Greece and its wider surroundings broader area definition here, we additionally requires more comprehensive sampling, we sug- formally accept the genera Anatololacerta and gest to accept the new four species arrangement Phoenicolacerta. The latter only has a single and to recognise Lacerta trilineata, L. diplo- representative within our area, the Cypriot P. chondrodes, L. citrovittata,aswellastheex- troodica. Within our area, the genus Anatololac- tralimital L. pamphylica. erta can be found on several Greek islands, in- Using one mitochondrial and one nuclear cluding Ikaria, Samos, Rhodes, Pentanisos and gene, Marzahn et al. (2016) investigated the Kastellorizo. Based on morphology, Eiselt and Downloaded from Brill.com10/05/2021 05:01:43PM via free access 156 J. Speybroeck et al.

Schmidtler (1986) treated A. anatolica (includ- the next available nomen is Iberolacerta Ar- ing the subspecies aegaea from Samos), A. dan- ribas, 1999. However, because Darevskia Ar- fordi and A. oertzeni (including the subspecies ribas, 1999 is a junior synonym of Caucasi- A. o. pelasgiana on Rhodes, Symi and Pen- lacerta Harris, Arnold and Thomas, 1998, Bu- tanisos and the nominotypical subspecies on sack et al. (2016) conclude that Caucasilac- Ikaria) as valid species. However, because the erta must replace Darevskia as the valid nomen biochemical differentiation (based on albumins) for the genus of the Caucasian Rock Lizards. between them was deemed to be too small, The conclusions drawn by Busack et al. (2016) Sindaco and Jermencenkoˇ (2008) maintained prompted a number of responses (Arribas, 2016; Arribas et al., 2017), advocating that Cau- all populations within Anatololacerta danfordi, casilacerta is a nomen nudum and hence is un- following Mayer and Lutz (1989). Bellati et al. available. Subsequently, Arribas et al. (2018) (2015) combined multi-locus species tree ap- appealed to the ICZN (under Articles 78.1 and proaches with species delimitation methods to 81 of the Code) to accept Arribas (1997) as pub- suggest the recognition of four distinct species lished in the sense of the Code and preserve in this group. Some of these species are closely Darevskia Arribas, 1997 and Iberolacerta Ar- related and were inferred to have diverged as ribas, 1997. While we regard Caucasilacerta as recently as 500,000 years ago, i.e. a remark- available, we support the current application to ably young age for speciation events within preserve Darevskia. We note that, while the case Lacertidae. Yet, there seems to be evidence of is under consideration by the Commission, the reproductive isolation between some of these prevailing usage of Darevskia and Iberolacerta species, as shown by the patterns of allele shar- with their authorship as Arribas, 1997 is to be ing in some nuclear markers. While more de- maintained (Article 82). tailed analyses of patterns of gene flow in con- Based on morphological variation, Stugren tact zones would be desirable, we tentatively ac- (1961) advocated the recognition of three sub- cept the four-species taxonomy (A. anatolica, A. species of Darevskia praticola: the nomino- budaki, A. danfordi and A. pelasgiana)ofBel- typical subspecies in the east of the distribu- lati et al. (2015). In our area, A. anatolica is tion (central and eastern Caucasus, northern present on Ikaria and Samos, and A. pelasgiana and south-eastern Georgia, northern Armenia, on Symi, Pentanisos and Rhodes. In addition, A. southern Azerbaijan, and north-eastern Iran), pelasgiana was also found (although presumed the subspecies pontica in the west of the Cauca- sus and in western Georgia, and D. p. hungarica introduced in both cases) on Kasos (Kornilios in Europe (north-eastern Serbia, southern Ro- and Thanou, 2016) and Kastellorizo (Kalaentzis mania, north-eastern Greece, eastern and west- et al., 2018), while A. budaki was found on the ern Bulgaria). Later, he recommended to treat (nearby) islet Psomi (Kalaentzis et al., 2018). D. p. hungarica as a synonym of D. p. pontica, As pointed out by Busack et al. (2016), resulting in a two-subspecies classification (Stu- the microfiche publication of Arribas’s (1997) gren, 1984). On the basis of multivariate anal- thesis does not meet the criteria of publica- ysis of morphological, meristic and qualitative tion for valid zoological nomenclature, nei- characters of Balkan and Caucasus populations, ther under the current version of the Code Ljubisavljevic et al. (2006) confirmed the valid- nor under the previous (and then relevant) ity of the two subspecies D. p. praticola and version. As a consequence, the names Ibero- D. p. pontica. Tuniyev et al. (2011) presented lacerta and Darevskia were not made avail- a detailed morphological study for the popula- able in Arribas (1997). For Iberolacerta,this tions of the Caucasian Isthmus. They confirmed has no major nomenclatural consequences, as two morphological clusters in the Caucasus and Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 157

Transcaucasia (western Caucasus: D. p. pon- was part of the Kingdom of Hungary, includ- tica; eastern Caucasus + Transcaucasia: D. p. ing parts of today’s Romania), should be applied praticola), and described a new subspecies from to the Balkan lineage. Given the incomplete ge- the Talysh Mountains (the extralimital D. p. hyr- ographic coverage of Freitas et al. (2016) and canica). On the basis of morphological differen- the somewhat borderline level of genetic diver- tiation and unpublished genetic data, they also gence between the Balkans and Caucasian + advocated species status for D. p. pontica. Later, Transcaucasian lineages, we refrain from rec- Freitas et al. (2016) provided a phylogeographic ommending species status for the time being, study of the species, based on comprehensive and thus suggest to treat the Balkan popula- sampling covering most of the range (although tions as Darevskia praticola hungarica.Wefur- the central and eastern Caucasus parts of the ther disagree with the conclusion of Freitas et range were missing). They recovered two clades al. (2016) to treat D. p. pontica as a synonym in the Caucasus and in Transcaucasia, one in the of D. p. praticola, as all their samples corre- western Caucasus (corresponding with the Cau- spond with the range of D. p. pontica according casian range of the D. p. pontica subspecies) to Tuniyev et al. (2011), and they have there- and one in Transcaucasia (Armenia) and the fore not analysed any sample from the range Talysh Mountains, corresponding with the ex- of D. p. praticola. Conclusively, we maintain tralimital subspecies D. p. hyrcanica and Tu- the morphologically well-supported subspecies niyev’s et al. (2013) D. p. loriensis. Samples D. p. pontica and D. p. praticola for the Cau- of the nominotypical subspecies were not in- casian populations occurring in our area, while cluded, although their morphology suggests that the Balkan populations are referred to as the they would presumably group with the Tran- subspecies D. p. hungarica. scaucasian samples (see Tuniyev et al., 2011, Genetic relationships based on a mtDNA (cyt 2013). Freitas et al.’s (2016) results thus seem b) fragment suggest that Darevskia brauneri to support the divergence of two lineages in the szczerbaki is sister to D. saxicola rather than Caucasus, corresponding to Caucasian popula- grouping with other lineages of D. brauneri tions of pontica on the one hand, and popula- (Doronin, Tuniyev and Kukushkin, 2013; tions of the praticola phenotype (including lo- Tarkhnishvili et al., 2016). Based on this and riensis and hyrcanica) on the other hand. How- on analysis of morphological variation cou- ever, genetic analyses of samples of praticola pled with differences in ecology, Tuniyev and from the eastern Caucasus would be needed to Tuniyev (2012) and Doronin, Tuniyev and confirm this. More surprisingly, Freitas et al. Kukushkin (2013) suggested to treat szczerbaki (2016) also uncovered a deep split within popu- as a valid, monotypic species. However, this hy- lations traditionally classified as pontica, with pothesis is in conflict with the results of Mac- the European samples from the Balkans forming Culloch et al. (2000), who found a lack of fixed a deeply divergent lineage in both mtDNA and differences in 15 polymorphic allozyme loci be- nuclear DNA that splits from a node basal to tween D. b. brauneri, D. b. darevskii and D. the clade grouping the Caucasian and Transcau- b. szczerbaki, and concluded that these three casian samples. They estimated the divergence taxa are conspecific. The high incidence of hy- between the European and Caucasian + Tran- bridisation within Caucasian Darevskia makes scaucasian clades at around 2.5 Mya, with a mtDNA relationships potentially unreliable. In much younger split of the Transcaucasian lin- addition, because of the low level of cyt b di- eage around 650,000 year ago. As pointed out vergence (2.4% between D. saxicola and D. b. by Ljubisavljevic et al. (2006), the name D. p. szczerbaki, and 4.8-5.6% of both D. saxicola hungarica (Sobolevsky, 1930), with type local- and D. b. szczerbaki towards other D. brauneri ity in the Transylvanian Alps (which at the time sequences) and the conflict with allozyme data, Downloaded from Brill.com10/05/2021 05:01:43PM via free access 158 J. Speybroeck et al. we consider this species complex to be cur- in northern Italy, and did not detect any hybrid rently too poorly studied to recognise additional or admixed genotype out of around 30 individ- species without extensive evidence from nuclear uals (genotyped with 13 microsatellites). Cor- genes and/or strong morphological differentia- netti et al. (2015b) genotyped many more in- tion in sympatry or close parapatry. As both of dividuals from several regions in the northern these lines of evidence are still lacking in this Italian Alps where the two taxa come in close case, we maintain D. b. szczerbaki at subspecies parapatry, and found a similar lack of admix- rank for the time being. ture. Thus, carniolica and vivipara appear to Two parthenogenetic Darevskia species represent two different species, and we accept which are not native to our area have been intro- Zootoca carniolica at species level. duced near Zhytomyr, Ukraine (Nekrasova and After being placed into the genus Lacerta, Kostiushyn, 2016). Darevskia armeniaca oc- Teira dugesii and Scelarcis perspicillata were curs naturally in north-western Armenia, west- transferred to the subgenus Teira and the genus ern Azerbaijan, southern Georgia and north- Podarcis, respectively, based on morphology eastern Turkey, and is believed to have been introduced in 1963, whereas D. dahli, known (Richter, 1980). Subsequently, Teira was ele- from northern Armenia and southern Georgia, vated to genus level by Mayer and Bischoff was discovered more recently (1980) within (1996). Yet, the generic allocation of S. per- the introduced armeniaca population. A third spicillata and T. dugesii remained a matter of species, D. mixta, was also introduced in 1968, confusion, with several authors attributing both but its contemporary presence could not be con- species to the genus Lacerta until quite recently firmed (Nekrasova and Kostiushyn, 2016). (e.g. Brehm et al., 2003; Perera et al., 2007). Mayer et al. (2000) described the oviparous Even though Pavlicev and Mayer (2009) sug- Zootoca vivipara populations of Carinthia (Aus- gested treating Scelarcis as a junior synonym tria), Slovenia, Friuli and isolated sites across of Teira and placing both species into the genus the Po Plain (Italy) as the distinct and allopatric Teira, SBC2010 adopted the genus arrangement subspecies Z. v. carniolica. Additional molecu- of Arnold, Arribas and Carranza (2007) and as- lar analysis confirmed its validity (Surget-Groba signed the Moroccan Rock Lizard to Scelar- et al., 2001, 2002). More recently, this taxon cis as Scelarcis perspicillata. However, a recent was shown to inhabit a wider range than previ- study using data from anchored phylogenomics ously known, not only occupying parts of south- and a fossil-dated time tree by Garcia-Porta et ern Austria, Slovenia, northern Croatia and al. (2019) confirmed that Teira and Scelarcis north-eastern Italy, but actually extending west are sister taxa and that their divergence is at a as far as the central southern Italian Alps (Cor- similar level or even lower than among species netti et al., 2015a). Lindtke, Mayer and Böhme of many well-established genera. According to (2010) documented a narrow contact zone in this study, Teira and Scelarcis diverged from Carinthia (Austria), and detected two natural hybrids (identified by clutch features) in a sam- each other in the mid-Miocene, coinciding with ple of 36 specimens. As they did not use genetic or even post-dating intrageneric splits in many markers, they could not investigate the level other lacertid genera such as Algyroides, Gallo- of introgression. More recently, two studies by tia, Iberolacerta, Lacerta, Podarcis, Psammod- Cornetti et al. (2015a, b) demonstrated complete romus, Takydromus, and Timon. Mendes et al. reproductive isolation between carniolica and (2016) argued to maintain Scelarcis mainly be- vivipara, in spite of widespread close parapatry cause of nomenclatural stability. We follow this between these two taxa. Cornetti et al. (2015a) approach and maintain Teira and Scelarcis as focused on the contact zone with local syntopy valid genera, awaiting additional evidence. 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Psonis et al. (2017, 2018) examined species populations, currently classified as several sub- delimitation in the Podarcis tauricus group (in- species of P. siculus, to species rank as Podar- cluding P. tauricus, P. milensis, P. gaigeae cis latastei,aslatastei is the oldest name avail- and P. melisellensis) using mitochondrial and able for this lineage. At this stage we are re- nuclear DNA sequences, microsatellites and luctant to follow this decision for two reasons. RADseq data. Their most complete phyloge- Firstly, the amount of divergence between the nomic analysis offered a fully resolved species Western Pontine populations and the rest of the level phylogeny, and confirmed the existence P. siculus complex in Senczuk et al. (2019) is of extensive genetic structuring within P. tauri- only presented as a combined mtDNA and nu- cus. Two main clades correspond with the west- clear DNA tree, not allowing to check if nuclear ern and eastern part of the distribution of the data independently supports this species-level species on either side of the Pindos Mountains. divergence or not. As multiple examples of deep The level of divergence between them is com- mtDNA divergence that are not reflected in ge- parable with their divergence from the well- nomic divergence are known, we feel that com- established species P. melisellensis, P. milen- pelling evidence of nuclear DNA divergence is sis and P. gaigeae, and extensive admixture is needed before adopting this new species. Sec- not apparent from the available data. They dif- ondly, Senczuk et al. (2019) presented a simpli- fied picture of the patterns of genetic divergence fer quite obviously in coloration of adult males. in P. siculus. More detailed results by Senczuk Even if no information on the nature of the con- et al. (2017, 2018) display a much more com- tact zones is available, the data at hand support plex situation, where the Western Pontine lin- the split of P. tauricus into an eastern species eage appears as one of several deeply diver- (retaining the name P. tauricus and including gent lineages in P. siculus, and latastei and sicu- the subspecies P. t. thasopulae) and a western lus are not reciprocally monophyletic. While species, for which the oldest available name is Senczuk et al. (2018) indicated differentiation P. ionicus (Psonis et al., 2017, 2018). Several in nuclear DNA between at least some of the of the subclades within P. ionicus, supported by main mtDNA lineages in P. siculus, no infor- mitochondrial and genomic data, are as diver- mation on their level of reproductive isolation is gent as other species pairs in the genus Podar- yet available. As a consequence, while the TC cis and probably warrant species status as well, acknowledges that P. siculus as currently under- but formal recognition as separate species is still stood is possibly made up of several species, lacking. On the basis of published information, one of them corresponding to the Western Pon- we accept the split of Podarcis tauricus into two tine islands populations, we consider adopting species: P. tauricus and P. ionicus. any formal taxonomic change premature. Using published data on three mitochondrial Michels and Bauer (2004) provided a list of and three nuclear genes, Senczuk et al. (2019) what they argued to be justified emendations examined the phylogeny of the genus Podarcis (citing Article 31.1.2 of the Code) of non-avian to address the systematic status of the Podar- reptile and amphibian scientific names which cis siculus populations from the Western Pon- the authors identified as inappropriate origi- tine Islands, off the Italian west coast. They nal constructions of patronyms or matronyms. estimated that the Western Pontine lineage di- Among the listed species was Podarcis raffonei verged from all other P. siculus lineages around (as Lacerta sicula raffonei Mertens, 1952). Be- 4 Mya, and showed that this genetic diver- cause this taxon was named in honour of the gence is similar to or greater than that between collectors’ wife, Michels and Bauer (2004) con- many other species pairs of Podarcis lizards. cluded that raffonei should be changed into raf- Hence, they formally raised the Western Pontine foneae. However, Article 31.1.2 simply states Downloaded from Brill.com10/05/2021 05:01:43PM via free access 160 J. Speybroeck et al. how species-group names should originally be (2014) provided additional morphological anal- formed. The article does not provide grounds yses, and formally named them as Podarcis for subsequent changes to the name. Further- guadarramae (former type 1) and P. virescens more, justified emendations can only be made (former type 2). Morphological and molecular to ‘spellings that must be corrected’ (Article differentiation within Podarcis guadarramae re- 32.5, see also 33.2.2), in which these cases are sulted in the description of the subspecies P. g. not included. Even though Michels and Bauer lusitanicus from northern Portugal and north- (2004) are linguistically correct (Arribas, 2017), western Spain. We accept P. guadarramae and amending raffonei to raffoneae is unjustified P. virescens as valid species, while the status of (Article 33.2.3), making the latter an unjustified lusitanicus will be the focus of a forthcoming emendation and hence a junior objective syn- study (G. Dias and C. Pinho, pers. comm.). onym of the former (see also Dubois, 2007). Although sometimes credited to Gray, 1825 We thus recommend to maintain the original (e.g. Sindaco and Jeremcenko,ˇ 2008), the fam- spelling P. raffonei. As a side note, we highlight ily name Scincidae must be credited to Oppel that the results of Senczuk et al. (2019) also cast (1811), as adopted by Hedges (2014) and oth- serious doubts on the validity of the species sta- ers. Oppel (1811) created the name as “Familia. tus of P. raffonei, but we refrain here from ad- Scincoides”. Article 11.7.3 of the Code (see vocating any change for now. family Agamidae above) makes it clear that the Investigating the species group comprising nomen is made available by its publication in Podarcis cretensis, P. levendis and P. pelo- Oppel (1811), even if the suffix must be cor- ponnesiacus, Spilani et al. (2019) found well- rected. supported differentiation within the latter A split of the Scincidae into seven families species. Based on 17 microsatellites, the east- would affect the systematics of the European ern populations diverged 1.86 Mya. Most note- genera, with Ablepharus being placed into Eu- worthy, the eastern and western lineages occur gongylidae, Heremites into Mabuyidae and the in actual or near syntopy without any apparent remaining genera Chalcides, Ophiomorus and sign of admixture. The observed level of diver- Eumeces remaining in Scincidae s.s. (Hedges gence, together with evidence of reproductive and Conn, 2012; Hedges, 2014). This was nei- isolation, warrant a species-level split within P. ther adopted in the large-scale squamate phy- peloponnesiacus. However, Spilani et al. (2019) logeny of Zheng and Wiens (2016), nor in refrain from making any taxonomic change. The the online Reptile Database (Uetz, Freed and TC follows this position, as we await a forth- Hošek, 2019). As most skinks are readily recog- coming morphological study, prior to proposing nisable as belonging to this group, and the a formal change. monophyly of Scincidae sensu lato is well es- From a long line of studies on the Iberian tablished, we follow the argument of the ‘phe- Podarcis species, the acceptance of Podarcis li- notypic diagnosability’ taxon naming criterion olepis and the redefinition of Podarcis hispan- of Vences et al. (2013a) for the time being, keep- icus sensu stricto were the most recent steps ing all species (even if there are many) within a that were adopted by SBC2010. At the time, single family. Distinction of subfamilies, how- two additional species were sufficiently sub- ever, certainly makes sense as it provides more stantiated, genetically as well as morphologi- manageable units for taxonomic revision and cally. Awaiting formal naming, they were listed other purposes. as Podarcis hispanicus type 1 (north-western For a long time, the skink genus Mabuya in- Iberian Peninsula and central Iberian moun- cluded numerous species from South America, tain chains) and type 2 (parts of central and Africa and Asia. A series of phylogenetic works south-western Iberian Peninsula). Geniez et al. (e.g. Mausfeld et al., 2000, 2002; Carranza and Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 161

Arnold, 2003; Mausfeld and Schmitz, 2003; (three nuclear genes) for single-locus and mul- Whiting et al., 2006) confirmed the monophyly tilocus coalescent-based species delimitation. of the genus, but also uncovered deep evolu- The established COI p-distance is substantial tionary divergence between its main radiations, (12.7%). No allele sharing was found, and the which also differ consistently in several mor- Asian and European lineages appeared recipro- phological and life-history traits. The initial di- cally monophyletic for all three nuclear genes, vergence within the group is estimated to date confirming their long-term isolation. Even if back to the Eocene, between 40 and 50 million suboptimal preservation of specimens and/or years ago (Karin et al., 2016). Mausfeld et al. morphological conservatism in these fossorial (2002) were the first to suggest a split of the species did not allow identifying any morpholo- genus Mabuya to reflect the long evolutionary gical differentiation between the two allopatric divergence of its main lineages, a proposal that Ophiomorus lineages, the authors advocated has been universally adopted since. The Middle recognition of the eastern lineage from Ana- Eastern Mabuya species (M. aurata, M. septem- tolia as a new species, which they described taeniata and M. vittata) form a well-supported as O. kardesi. The status of several mitochon- monophyletic group in all phylogenies. They drial lineages recovered within O. punctatis- have been classified with the African species simus awaits further study. Based on the diver- in the genus Trachylepis for several years but, gence time and level of genetic divergence be- even if their phylogenetic position within the tween these two cryptic species, we accept the radiation of the former Mabuya is still poorly species status of O. kardesi. In our area, the new resolved, they are not part of the radiation of species occurs on the Greek island of Kastel- African species in the genus Trachylepis (Karin lorizo (Kornilios et al., 2018). et al., 2016). Their origin is quite ancient (esti- Phylogenetic analysis based on mitochon- mated around the late Oligocene, more than 30 drial and nuclear markers (Skourtanioti et al., million years ago) and their clade is thus now 2016) showed strong divergence, supporting afforded genus rank by all recent works on the the independent history of a clade of snake- subject, a position that we follow here. The valid eyed skinks (Ablepharus) from Kastellorizo and genus-group nomen of the Middle-East clade is southwestern Turkey. The samples of this clade Heremites Gray, 1845 (masculine, see Karin et originate from an area where the only taxon al., 2016). We thus recommend calling the Mid- known to occur according to Schmidtler (1997) dle East species Heremites auratus, H. vittatus is Ablepharus budaki anatolicus. As a conse- and the extralimital H. septemtaeniatus. quence, Skourtanioti et al. (2016) suggested The phylogeography of Ophiomorus punc- that the name anatolicus is available for this tatissimus was examined by Poulakakis et al. new species-level lineage, which splits from a (2008) using three mitochondrial fragments. more basal node than all other but one (A. pan- On the basis of divergence rates of mitochon- nonicus from Iran) species of the genus, and drial genes calibrated in other groups of squa- whose divergence was estimated to date back to mates, they uncovered a deep divergence be- the mid-Miocene. As the genetic differentiation tween the Asian (Anatolian) and European and deep phylogenetic divergence of the new populations of the species, estimated to have species was confirmed by independent analy- occurred around 10 Mya. This date fits nicely ses of both mitochondrial and nuclear genes, we with the opening of the mid-Aegean trench, agree that the genetic data strongly support the estimated from geological information to have recognition of an additional Ablepharus species started around 12 Mya and being completed from SW Turkey (including Kastellorizo and around 9 Mya. More recently, Kornilios et al. hence entering our area). However, we refrain (2018) used mtDNA and nuclear DNA data from formally accepting a new species here and Downloaded from Brill.com10/05/2021 05:01:43PM via free access 162 J. Speybroeck et al. prefer to wait until genetic data confirm that ani- and La Gomera and El Hierro (C. v. coeruleop- mals from the type locality of anatolicus indeed unctatus) are not each other’s closest relative, represent this new lineage. with coeruleopunctatus being more closely re- Sindaco and Jeremcenkoˇ (2008), who pro- lated to the taxa from Gran Canaria (sexlinea- vided the starting point of our taxonomy for tus and bistriatus) than to viridanus (Brown and areas not covered by SBC2010 or Gasc et Pestano, 1998; Carranza et al., 2008). These re- al. (1997), follow Pasteur, Keymar and Perret lationships rule out maintaining coeruleopunc- (1988) in treating the Gran Canaria skinks as tatus as a subspecies of viridanus, as was tra- two species, Chalcides bistriatus in the north- ditionally the case based on their morpholo- east of the island and C. sexlineatus in the south- gical similarity. In addition, the amount of di- west. This classification was based on a sharp vergence between C. sexlineatus, C. viridanus geographic transition in morphology and (es- and C. coeruleopunctatus is typical of interspe- pecially) colour pattern between the skinks in- cific divergences in the genus Chalcides (Car- habiting the mesic and xeric habitats on Gran ranza et al., 2008). We thus accept the conclu- Canaria (see Brown and Thorpe, 1991a, b, al- sion of Carranza et al. (2008) and treat the an- though these authors rejected the hypothesis of imals from La Gomera and El Hierro as Chal- speciation between these morphotypes). Studies cides coeruleopunctatus. on the evolutionary history of these populations The publication dates of the works of Daudin using allozymes (Mayer and Tiedemann, 1991) have been worked out by Harper (1940). The or a combination of mtDNA and microsatel- volume 4 of “Histoire Naturelle des Reptiles”, lites (Suárez, Pestano and Brown, 2014) estab- containing the description of Eumeces schnei- lished that this differentiation has a genetic ba- derii, was published in August 1802. The spe- sis, originating from allopatric diversification cific epithet is often spelled schneideri,even within Gran Canaria. This divergence is linked if both schneideri and schneiderii areinuse. to volcanic activity 1.5-3 Mya and is currently The original spelling is schneiderii (see Daudin, maintained by strong selection on coloration, 1802), with schneideri as an incorrect subse- in spite of considerable introgression and gene quent spelling. The only reason to use this flow between the two morphs. While the deepest spelling rather than the original spelling would divergence between the mtDNA clades within Gran Canaria is dated to 1.5 Mya, the diver- be if the incorrect subsequent spelling schnei- gence between the two morphs was dated to deri was in prevailing usage (see Art. 33.3.1 260,000 years ago. This discrepancy could re- of the Code). Prevailing usage is defined by flect real differences between coalescence time the Code as “that usage of the name which is of mtDNA lineages versus divergence time of adopted by at least a substantial majority of the populations, or analytical issues with one of the most recent authors concerned with the rele- methods. Either way, the available data suggest vant taxon”. We checked the number of publica- a young divergence of C. s. sexlineatus and C. tions using either spelling after 2000 using the s. bistriatus, together with a lack of reproduc- electronic version of the Zoological Record and tive isolation. We thus recommend to treat these found that 17 works published after 2000 used taxa as conspecific as C. s. sexlineatus and C. s. schneiderii while 28 used schneideri. We inter- bistriatus, as adopted by Mayer and Tiedemann pret this as a lack of clear support for a prevail- (1991), Carranza et al. (2008) and Salvador and ing usage of the incorrect subsequent spelling Brown (2015). and thus adopt the original spelling E. schnei- Mitochondrial DNA data of the Canarian derii. skinks establish with a very high support that Considerable divergence within the genus the Chalcides from Tenerife (C. v. viridanus) Anguis has been revealed by means of genetic Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 163 and morphological analyses (Cabela and Gril- Iberian Blanus (SBC2010). These issues were litsch, 1989; Gvoždík et al., 2010). This led to tackled by Ceríaco and Bauer (2018), who ar- the recognition of A. graeca and A. colchica,re- gued that the type locality of Amphisbaena sulting in the acceptance of at least four Anguis cinerea Vandelli, 1797 falls within the range of species in Europe by Gvoždík et al. (2010) and, populations attributed to B. mariae by Albert subsequently, SBC2010. Although no Italian and Fernández (2009). To further strengthen specimens were incorporated in previous anal- their interpretation, they made a valid neotype yses, Anguis fragilis was hitherto presumed to designation for B. cinereus, selecting a neo- inhabit much of western, central and eastern Eu- type belonging to the south-western Iberian rope, including the Italian Peninsula (Gvoždík species. This effectively places B. mariae in et al., 2010). More recently, Gvoždík et al. the synonymy of B. cinereus, which becomes (2013) showed Italian Anguis to represent a the valid name for the south-western Iberian deeply differentiated mtDNA clade, which pre- Blanus species. In agreement with Sampaio et sumably diverged during or shortly after the al. (2015), who confirmed that the two Iberian basal radiation within the genus. Genetic dis- Blanus clades show high levels of mitochon- tances between the Italian clade and all recog- drial differentiation and no signs of nuclear nised species are larger than the interspecific haplotype sharing, Ceríaco and Bauer (2018) distances among A. fragilis, A. graeca and A. accepted the existence of two Iberian Blanus colchica. Hence, Gvoždík et al. (2013) proposed species. However, they deemed no name to to recognise the populations from Italy and ad- be valid for the Central Iberian species. Yet, jacent extreme south-eastern France as Anguis two older names, Amphisbaena oxyura Wagler, veronensis. Two specimens (one from NE Italy 1824 and Amphisbaena rufa Hemprich, 1820, and one from Slovenia) were heterozygous for are available and relate to Blanus from Spain. the most common A. fragilis and A. veronen- The type specimen of A. oxyura is lost, while sis PRLR haplotypes and carried A. fragilis its type locality (‘Spain’, according to Wagler, mtDNA, indicating hybridisation. While mor- 1830) makes it impossible to link this name phological differentiation between A. veronen- to either of the two Spanish Blanus species. sis and A. fragilis is significant, the ranges of all The name A. rufa was based on a holotype that studied characters overlap. In light of the earlier is still present in the Zoological Museum in accepted splits among the European Anguis by Berlin (ZMB), but the catalogue entry for this SBC2010, the divergence during the basal ra- specimen shows that it originates from south- diation of the genus, and the genetic and mor- ern Spain, where both Blanus clades occur. As phological evidence presented by Gvoždík et morphological identification of the two Span- al. (2013), we recognise the Italian and south- ish Blanus species is currently impossible, ro- eastern French populations as A. veronensis. bust attribution of A. rufa to either clade re- Albert and Fernández (2009) partitioned the quires molecular data. Yet, this is problematic Iberian Worm Lizard Blanus cinereus into two with the currently available methods (Ceríaco species, attributing southwestern Iberian popu- and Bauer, 2018). Ceríaco and Bauer (2018) lations to the cryptic Blanus mariae and desig- thus treated these two names as nomina du- nating a lectotype for B. cinereus. Their lecto- bia and created the new nomen Blanus vandel- type designation was, however, found to be in- lii. SBC2010 had already recognised the two valid, due to the lack of explicitly stated tax- Iberian Blanus clades at species level. We fol- onomic purpose, while they did not establish low this arrangement, and accept that B. mariae whether the original type series consists of one is a junior synonym of B. cinereus, with the holotype or several syntypes, nor examined the latter being the valid name for the southwest- status of other, older nomina in the synonymy of ern Blanus clade (Ceríaco and Bauer, 2018). Downloaded from Brill.com10/05/2021 05:01:43PM via free access 164 J. Speybroeck et al.

However, instead of introducing a new name in proposed to treat H. viridiflavus and H. car- the form of B. vandellii, we promote the use bonarius as different species on the basis of of the oldest available name, Blanus rufus,as (i) different morphology of the W sex chro- the valid name for the central clade until in- mosome, (ii) reciprocal monophyly in mtDNA formation on the identity of the A. rufa holo- with a genetic divergence of 4% in both cyt type can be obtained. Rather than discarding b and ND4, and (iii) morphological differen- the old name until proven to apply, we advocate tiation consistent with mtDNA differentiation. that when an available name exists, it should be However, Rato et al. (2009) found a lack of used until proven not to apply to the relevant general agreement between colour pattern and taxon. We thus list the two Iberian species as mtDNA clade in Italy, and no differences in B. cinereus (south-western species) and B. ru- the nuclear intron beta-fibrinogen intron 7 be- fus (north-eastern species). tween the two lineages. The amount of diver- Hedges et al. (2014) present molecular and gence in mtDNA between H. v. carbonarius and morphological data to clarify the phylogeny of H. v. viridiflavus is much lower than between the Typhlopidae and split the large genus Ty- H. gemonensis and H. viridiflavus, while the phlops into a number of well-defined new gene- amount of divergence in PRLR between them ra. As a consequence, the European blind snake resembles the divergence within H. gemonensis. species is now referred to as Xerotyphlops ver- The sample size for the chromosomes is quite micularis. Additionally, according to Hedges et low as well. The split of H. v. carbonarius from al. (2014), the non-native Flowerpot Snake, re- H. v. viridiflavus did not reach broad support in cently introduced to Italy and Spain (Zamora- the TC, as several members felt the evidence Camacho, 2017; Paolino, Scotti and Grano, is still insufficient, and they prefer to wait for 2019), is now called Indotyphlops braminus. information on genetic variation near the con- The name of Hierophis viridiflavus has been tact zones. We thus recommend to maintain Hi- involved in nomenclatural debate over the last erophis viridiflavus carbonarius as a subspecies few decades, but are now solved (ICZN Opin- for the time being. ions 1463 and 1686). In 1833, Charles-Lucien Jablonski et al. (2019) applied phylogenetic Bonaparte (the nephew of Napoleon) described and morphological analyses to range-wide sam- Coluber viridi-flavus carbonarius for the east- ern Italian populations that are completely or pled data, aiming to better understand the in- almost completely black. The subspecies H. v. traspecific relationships and biogeography of carbonarius was considered valid, until Schätti Elaphe sauromates, and subsequently revised and Vanni (1986) placed it in the synonymy of its taxonomy. Known from the Balkans, Ana- the nominal form. A series of phylogeographic tolia, the Caucasus, the Ponto-Caspian steppes, studies based on mtDNA and nuclear DNA con- and the Levant, this species has been suspected firmed that viridiflavus and carbonarius are dis- to be composed of two or more genetically tinct evolutionary lineages which form recip- diverse populations (Lenk, Joger and Wink, rocally monophyletic clades in mtDNA (Nagy 2001). Sequences from 63 specimens and mor- et al., 2001; Rato et al., 2009; Mezzasalma et phological data from 95 specimens were anal- al., 2015; Avella, Castiglia and Senczuk, 2017). ysed. The authors found two distinct evolution- These two lineages differ in the morphology of ary lineages, one of which represents a new their W chromosomes (submetacentric in the E species, Elaphe urartica. The new species is clade and telocentric in the W clade – Mezza- distributed in eastern Turkey, Georgia, Armenia, salma et al., 2015). As a consequence, they have Azerbaijan, Nagorno-Karabakh, Iran, and Rus- been treated as valid subspecies since Nagy et sia (Dagestan). The mtDNA genetic distances al. (2001). In 2015, Mezzasalma et al. (2015) between E. sauromates and E. urartica (6-8% Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 165 in COI and ND4) were interpreted as indicat- naming criterion of Vences et al. (2013a) and as ing a split at the Miocene-Pliocene boundary a named genus of uncontroversial monophyly is (5-8 Mya; see Kornilios et al., 2014), thus they obtained, we here follow this. presumably separated not very long after their The California Kingsnake Lampropeltis common ancestors separated from E. quatuor- getula californiae has been introduced to Gran lineata. Two out of four analysed nuclear genes Canaria, where it turned into a problematic in- were also clearly differentiated, though closely vasive species (Monzón-Arguëllo et al., 2015; related, between E. urartica and E. sauromates, Fisher et al., 2019). Based on mtDNA data and while the remaining two showed signs of in- morphological variation, Pyron and Burbrink complete lineage sorting. Both lineages are also (2009) elevated L. g. californiae and four other moderately morphologically differentiated and, subspecies to species status. While the basal di- while none of the characters are exclusively di- vision in their phylogeny has L. g. getula and L. agnostic, their combination can be used for con- g. nigra in the one main branch and L. g. hol- fident lineage identification. Conclusively, we brooki, L. g. splendida and L. g. californiae in accept Elaphe urartica as a separate species. the other, the division between L. g. splendida The ladder snake Rhinechis scalaris is char- and L. g. californiae is retrieved as the most re- acterised by a backward-oriented rostral scale cent one. Pending the publication of a multilo- that is wedged between the internasal scales. cus analysis or a detailed study of the various Apart from this striking feature, external mor- contact zones, we maintain L. g. californiae as phology of R. scalaris has long been acknowl- a subspecies of the Common Kingsnake Lam- edged to closely match that of Zamenis rat propeltis getula for the time being. snakes (Salvi et al., 2018). Multiple recent The groundwork of contemporary elucidation phylogenetic assessments have revealed that of the Natrix natrix complex was initiated by Rhinechis (with the sole member R. scalaris) the mitochondrial phylogeography of Kindler and Zamenis are closely related (Burbrink and et al. (2013), leading to rejection of Natrix Lawson, 2007; Pyron et al., 2011; Pyron, Bur- megalocephala and the identification of multi- brink and Wiens, 2013; Zheng and Wiens, ple contact zones, three of which were further 2016). Some of these studies found Zamenis to investigated more recently (see below). Subse- be paraphyletic in respect to Rhinechis, while quently, a virtual lack of gene flow between others placed Rhinechis as sister to all Zamenis N. natrix helvetica and N. natrix astreptophora species, albeit with low support (reviewed by and the differentiation between both taxa es- Salvi et al., 2018). The latter authors argue that tablished on multiple sources of evidence, such this phylogenetic uncertainty reflects an evo- as microsatellites, mtDNA, osteology and exter- lutionary scenario in which early cladogenetic nal morphology, led to suggesting species status events took place at the same time or within a for Natrix astreptophora (Pokrant et al., 2016). short time frame, making it hard to disentangle Within central Europe, Kindler et al. (2017) in- which (if any) lineage branched off first. Us- vestigated two macro-scale transects across Na- ing two mitochondrial and five nuclear genes, trix transition zones. The eastern contact zone Salvi et al. (2018) then inferred phylogenetic between two lineages identified by Kindler et relationships between Zamenis and Rhinechis. al. (2013) as running from Central Europe to They recovered a strongly supported clade con- the southern Balkans displayed wide-reaching sisting of Rhinechis scalaris and Zamenis, while introgression. In contrast, the western contact the monophyly of Zamenis without scalaris re- zone between the traditional subspecies N. n. mained poorly supported. As a result, these au- natrix and N. n. helvetica in western Germany thors proposed to include scalaris in Zamenis. was shown to be narrow (microsatellite cline In agreement with the ‘clade stability’ taxon width of about 40 km), with a deficit of admixed Downloaded from Brill.com10/05/2021 05:01:43PM via free access 166 J. Speybroeck et al. individuals generating a bimodal hybrid zone, V. r. lotievi and V. r. shemakhensis as conspecific indicative of strongly restricted gene flow. We with V. renardi. therefore accept N. astreptophora and N. hel- Tuniyev and Ostrovskikh (2001) described vetica as valid species in addition to N. natrix. Vipera orlovi and V. magnifica, two viper Ferchaud et al. (2012) provided a well- species from the Russian Caucasus that are re- resolved mitochondrial phylogeny of the Vipera lated to Vipera kaznakovi. Vipera orlovi inhabits ursinii complex, which indicates that the re- both slopes of the northwesternmost Caucasus, nardi clade, the ursinii clade and the graeca while V. magnifica is described from the Malaja clade diverged approximately 3-4 Mya. The di- Laba river valley and restricted to Krasnodar vergence between these three clades is of the territory. As no molecular data was provided, same magnitude as between the two widely ac- and it was later shown that both taxa share cepted species V. berus and V. seoanei.There- mtDNA haplotypes with V. kaznakovi (Zinenko nardi and ursinii clades do not seem to share et al., 2015), further enquiry was needed. Zi- haplotypes, although they inhabit similar habi- nenko et al. (2016) used genomic methods to tats along the north-western Black Sea coast demonstrate the hybrid origin of these taxa, with where Vipera ursinii moldavica occurs. Given parapatric V. kaznakovi and V. renardi as likely the level of genetic divergence, lack of haplo- parental species. Low admixture and limited ge- type sharing, marked morphological divergence netic differentiation led these authors to treat V. and lack of well supported morphological in- magnifica as a synonym of V. kaznakovi. The au- tergradation, it seems more adequate to treat V. thors build a case for not revoking the species renardi as specifically distinct from V. ursinii. status of V. orlovi by advocating caution with re- Given its well-supported position at the base gards to its conservation status. While we do not of a renardi + ursinii clade and although we reject species of hybrid origin per se, the 2016 verified a very low genetic distance between data does not suggest reproductive isolation or ursinii and graeca in cyt b (3.2%), this arrange- strong genetic differentiation of either taxon in ment makes it necessary to treat V. graeca as a relation to V. kaznakovi. Thus, we do not treat distinct species as well. Additional morpholo- V. magnifica or V. orlovi as valid species. As gical and mtDNA and nuclear DNA data sup- they are more closely related to V. kaznakovi port the distinct position of the graeca clade than to V. renardi, we treat both as conspecific (Mizsei et al., 2017). Several other taxa from with V. kaznakovi. In addition, we disagree with the Caucasus and the Middle East have been Zinenko et al. (2016) that the Code does not al- put forward as valid species, two of which oc- low the recognition of taxa with hybrid ancestry. cur in our area: the Caucasian lotievi and the re- The Code rules nomenclature, not systematics, cently described shemakhensis (Tuniyev et al., and nothing in the Code precludes treating taxa 2013) from north-eastern Azerbaijan. A recent of admixed ancestry as valid. As both orlovi and phylogenomic (RADseq data) analysis of mul- magnifica were described on the base of dis- tiple specimens of renardi and lotievi by Zi- tinct phenotypic features, we tentatively main- nenko et al. (2016) showed a lack of genetic dif- tain them as subspecies of V. kaznakovi (V. k. ferentiation at the genome scale between spec- orlovi and V. k. magnifica) until a detailed exam- imens collected in the lowlands (renardi) and ination of morphological variation might show Caucasian uplands (lotievi). There is no genetic them not to be morphologically distinct. information for shemakhensis, which is, how- A new viper species, named Vipera walser, ever, very similar to lotievi morphologically. In was described from northern Piedmont, Italy conclusion, we consider three European species (Ghielmi et al., 2016) based on mtDNA, nuclear within the Vipera ursinii complex: V. graeca, V. DNA and morphological discriminant analy- renardi and V. ursinii, and recommend treating ses. Remarkably, while morphologically very Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 167 similar to north Italian V. berus, mtDNA data were based on populations resulting from in- showed the new taxon to be nested with geo- tergradation between berus and nikolskii and graphically distant taxa of the ursinii-kaznakovi cannot apply to the later taxon, leaving nikol- complex. For two protein-coding nuclear genes skii as the valid nomen for this taxon. While (BACH2 and RAG1), V. walser does not share it was first described at species rank, nikolskii haplotypes with V. berus. However, only single is now variously treated as a full species (e.g. individuals of both species have so far been se- Bakiev, Böhme and Joger, 2005; Uetz, Freed quenced. Moreover, for RAG1, the new species and Hošek, 2019) or as a subspecies of Vipera is similar to V. berus, while no sequences of the berus (e.g. SBC2010; Zinenko,Turcanu ¸ and ursinii complex exist for this gene. Therefore, Strugariu, 2010; Mizsei et al., 2017; Crnobrnja the existence of cytonuclear discordance can- Isailovic et al., 2019 by implication). Depend- not be discarded. Until additional nuclear DNA ing on their geographical origin, specimens of data confirms the distinct nature of the walser nikolskii either carry berus mtDNA haplotypes, population, we consider acceptance of the new or haplotypes grouping (within the diversity of species to be premature, and tentatively regard the berus group) close to V. b. barani (Joger it as a population of V. berus with distinct, per- et al., 1997; Kalyabina-Hauf et al., 2004). A haps introgressed, mtDNA. Morphologically, V. multilocus phylogeny places nikolskii within V. berus (Mizsei et al., 2017). Detailed morpholo- walser does not differ much from the V. berus gical studies show extensive introgression in the populations inhabiting the Italian Alps (the Ital- contact zones with V. b. berus (Zinenko, 2004; ian clade of V. berus in Ursenbacher et al., 2006) Milto and Zinenko, 2005; Zinenko,Turcanu ¸ and but it is easily distinguishable from the Cen- Strugariu, 2010). We thus maintain the treat- tral and Northern European populations of V. ment of SBC2010 and consider nikolskii as con- berus. Thus, the Italian Alps populations of V. specific with V. berus, supporting its recognition berus could be treated as a subspecies under the as V. b. nikolskii to acknowledge its morpholo- name V. b. walser. However, we refrain from gical and ecological peculiarities. making formal recommendations here, pending The correct spelling of the specific epithet of a detailed analysis of morphological variation the Lataste’s viper has been a matter of long between populations of V. berus from the Italian lasting debates because Boscá (1878) used the Alps (including the populations described as V. spelling “Vipera Latasti” in the text and “Vipera walser) and the rest of the V. berus populations. latastei” in the caption for the accompanying The populations of mostly black adders in- plate, with both spellings in current use in the habiting the forest-steppe zone of the south literature (see Alonso-Zarazaga, 2013 and Sal- and south-eastern parts of the East European vador et al., 2014 for summaries). Following an Plain have been described as Vipera nikol- application to the International Commission on skii Vedmederya, Grubant and Rudaeva, 1986. Zoological Nomenclature (ICZN) by Salvador The distribution and morphological characters et al. (2014), the ICZN (2017) confirmed that of nikolskii compared to berus have been re- Vipera latastei Boscá, 1878 is the correct origi- viewed by Bakiev, Böhme and Joger (2005), nal spelling of the specific name. Milto and Zinenko (2005) and Zinenko,Tur- ¸ Frétey (2019) provided compelling argumen- canu and Strugariu (2010, who showed that tation of why the capitalised epithet Lebetinus populations of nikolskii from the western parts in Coluber Lebetinus Linnaeus, 1758 is a noun of its range can be mostly composed of non- in apposition and must be treated as invari- melanistic specimens). According to Milto and able. We agree with his interpretation, and ac- Zinenko (2005), several older names created by cept that the Blunt-nosed Viper should be called Pallas for black adders from southern Russia Macrovipera lebetinus. Downloaded from Brill.com10/05/2021 05:01:43PM via free access 168 J. Speybroeck et al.

Based on morphology, Nilson and Andrén a collected specimen. The possible occurrence (1988) elevated Macrovipera lebetinus schweiz- of Gloydius within Europe is based on several eri to species rank. Stümpel (2012), however, ancient sources that report the genus between found that the mtDNA haplotypes found in the Volga and Ural rivers in the dry steppes or M. schweizeri are not limited to the Milos deserts east of Astrakhan (see Conant, 1982 and archipelago, but are also present near Mersin on Bour, 1993 for details). They all seem to orig- the Turkish mainland. This author thus pointed inate from just two original sources: Gmelin out that treating M. schweizeri as a valid species (1789), who mentions the arid deserts of As- might render M. lebetinus paraphyletic. Further- trakhan for Coluber halys, and Pallas (1799), more, they showed that in their mtDNA trees who, in his book devoted to his second journey schweizeri is neither basal to the rest of the to the southern areas of Russia, mentions (on p. M. lebetinus haplotypes, nor highly divergent. 112) the species “Berus” and “Halys” were seen Yet, while Stümpel’s dissertation is available, in an area called “Saltan-Murat” and located and a (brief) sentence in Stümpel and Joger east of the Volga River, in an area of dry steppes (2009) refers to these results, these findings close to Astrakhan (more precisely north-east have not yet been properly published in a peer- of present-day Krasnyy Yar, see Conant, 1982 reviewed journal, and the relevant Macrovipera and Bour, 1993). All subsequent references are sequences are not yet available in GenBank based on these two sources. It is even unclear if either. As a consequence, many TC members the mention of the deserts of Astrakhan in J.F. felt that a formal taxonomic decision on this Gmelin’s book is based on original information case would be premature, and, while it is likely he received from his uncle J.G. Gmelin, who that schweizeri is best treated as a subspecies, also travelled widely in Russia including the the decision can only be taken after the evi- Caspian Sea area, from Pallas himself, or from dence has been properly published, or the data any other source. J.F. Gmelin never visited these has been made publicly available. Conclusively, areas himself. We may never know how reli- we maintain Macrovipera schweizeri at species able these old records are. However, the other level for now. venomous species mentioned by Pallas (1799) The occurrence of the genus Gloydius in- from the same area, Vipera berus, is currently side the area covered by this list is unclear. The only European area where the occurrence absent from the dry areas between the Ural and of the genus has been reported is north of the Volga Rivers. This casts doubt on the reliability Caspian Sea, between the Volga and Ural val- of these observations. In summary, the presence leys. As explained by Conant (1982) and Dirk- of the genus Gloydius west of the Ural River sen and Böhme (2005), all records supported entirely rests on unsubstantiated records that by specimens in museum collections are either cannot be verified. We thus believe the species on the east side of the Ural River, hence out- should be formally removed from the list of Eu- side our area (Induski Hills = Indersky Moun- ropean species. In addition, we note that the tains), or cannot be precisely localised (one systematics and nomenclature of the Gloydius specimen from “Gur’jev”, now Atyrau, which halys complex (for a brief review, see Wagner et lies at the mouth of the Ural River on both sides al., 2016) remain problematical. One of the con- of the river). Attempts to locate Gloydius west tentious issues is the type locality of Coluber of the Ural River have failed (Conant, 1982; halys Pallas 1776, which determines the valid Bakiev, Ratnikov and Zinenko, 2007; Sarayev name of the Gloydius taxon occurring close to and Pestov, 2010; Simonov, pers. comm.). Thus, Europe in western Kazakhstan, but this question no modern records of the presence of the genus might need to be settled by an application to the in Europe exist, and no record is supported by Commission. Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 169

Species list Euproctus Gené, 1839 Euproctus montanus (Savi, 1838) – Corsi- Within the area considered for our species list, can Brook Newt a number of non-native species have been intro- Euproctus platycephalus duced and/or have become established, and are (Gravenhorst, reproducing. As far as they were not yet listed in 1829) – Sardinian Brook Newt the starting point species lists mentioned in the Ichthyosaura Sonnini and Latreille, 1801 introduction, we add these naturalised species Ichthyosaura alpestris (Laurenti, 1768) – here. More precisely, these are Tenuidactylus Alpine Newt bogdanovi (see above), Darevskia dahli (see Lissotriton Bell, 1839 above), Indotyphlops braminus (see above), Lissotriton boscai (Lataste in Tourneville, Elaphe schrenckii (introduced and persisting in 1879) – Bosca’s Newt the Netherlands, see Van de Koppel et al., 2012), Lissotriton graecus (Wolterstorff, and Lampropeltis getula (see above). 1906) – Greek Smooth Newt While no objective standards exist to choose Lissotriton helveticus (Razoumowsky, vernacular names, to improve user-friendliness 1789) – Palmate Newt of the list, we added English species names, Lissotriton italicus (Peracca, 1898) – Ital- most of which were adopted from Speybroeck ian Newt et al. (2016). Others were added to harmonise Lissotriton maltzani (Boettger, 1879) – as much as possible with those of Speybroeck Maltzan’s Newt et al. (2016) and Arnold and Ovenden (2002), Lissotriton montandoni (Boulenger, or with suggestions in the respective species 1880) – Montandon’s Newt description, or they are newly proposed here. Lissotriton vulgaris (Linnaeus, 1758) – Changes, as discussed in the text, are marked Smooth Newt in bold. Non-native species are marked with an asterisk. Taxa above genus level are listed in tra- Lyciasalamandra Veith and Steinfartz, 2004 ditional order, genera and species arranged al- Lyciasalamandra helverseni (Pieper, phabetically. For the Class Reptilia, we restrict 1963) – Karpathos Salamander ourselves to non-avian reptiles and disregard the Lyciasalamandra luschani (Steindachner, Subclass Aves. 1891) – Luschan’s Salamander Class Amphibia Linnaeus, 1758 Pleurodeles Michahelles, 1830 Pleurodeles waltl Michahelles, 1830 – Order Caudata Scopoli, 1777 or Urodela Sharp-ribbed Newt Duméril, 1805 Salamandra Garsault, 1764 Family Hynobiidae Cope, 1859 (1856) Salamandra atra (Laurenti, 1768) – Alpine Salamandrella Dybowski, 1870 Salamander Salamandrella keyserlingii Dybowski, Salamandra corsica (Savi, 1838) – Corsi- 1870 – Siberian Salamander can Fire Salamander Family Salamandridae Goldfuss, 1820 Calotriton Gray, 1858 Salamandra lanzai (Nascetti, Andreone, Calotriton arnoldi Carranza and Amat, Capula and Bullini, 1988) – Lanza’s 2005 – Montseny Brook Newt Salamander Calotriton asper (Dugès, 1852) – Pyre- Salamandra salamandra (Linnaeus, nean Brook Newt 1758) – Fire Salamander Chioglossa Bocage, 1864 Salamandrina Fitzinger, 1826 Chioglossa lusitanica Bocage, 1864 – Salamandrina perspicillata (Savi, 1821) – Golden-striped Salamander Northern Spectacled Salamander Downloaded from Brill.com10/05/2021 05:01:43PM via free access 170 J. Speybroeck et al.

Salamandrina terdigitata (Bonnaterre, Proteus Laurenti, 1768 1789) – Southern Spectacled Salaman- Proteus anguinus Laurenti, 1768 – Olm der Order Anura Duméril, 1805 Triturus Rafinesque, 1815 Family Alytidae Fitzinger, 1843 Triturus carnifex (Laurenti, 1768) – Italian Alytes Wagler, 1829 Crested Newt Alytes almogavarii Arntzen and García- Triturus cristatus (Laurenti, 1768) – Great París, 1995 – Catalonian Midwife Crested Newt Toad Triturus dobrogicus (Kiritzescu, 1903) – Alytes cisternasii Boscá, 1879 – Iberian Danube Crested Newt Midwife Toad Triturus ivanbureschi Arntzen and Wiel- Alytes dickhilleni Arntzen and García- stra, 2013 – Buresch’s Crested Newt París, 1995 – Betic Midwife Toad Triturus karelinii (Strauch, 1870) – Kare- Alytes muletensis (Sanchiz and Adrover, lin’s Crested Newt Triturus macedonicus (Karaman, 1922) – 1979) – Mallorcan Midwife Toad Macedonian Crested Newt Alytes obstetricans (Laurenti, 1768) – Triturus marmoratus (Latreille, 1800) – Common Midwife Toad Marbled Newt Discoglossus Otth, 1837 Triturus pygmaeus (Wolterstorff, 1905) – Discoglossus galganoi Capula, Nascetti, Southern Marbled Newt Lanza, Bullini and Crespo, 1985 – Ommatotriton Gray, 1850 Iberian Painted Frog Ommatotriton ophryticus (Litvinchuk, Discoglossus montalentii Lanza, Nascetti, Zuiderwijk, Borkin and Rosanov, Capula and Bullini, 1984 – Corsican 2005) – Northern Banded Newt Painted Frog Family Plethodontidae Gray, 1850 Discoglossus pictus Otth, 1837 – Painted Speleomantes Dubois, 1984 Frog Speleomantes ambrosii (Lanza, 1955) – Discoglossus sardus Tschudi in Otth, Ambrosi’s Cave Salamander 1837 – Tyrrhenian Painted Frog Speleomantes flavus (Stefani, 1969) – Family Bombinatoridae Gray, 1825 Monte Albo Cave Salamander Bombina Oken, 1816 Speleomantes genei (Temminck and Bombina bombina (Linnaeus, 1761) – Schlegel, 1838) – Gené’s Cave Salaman- Fire-bellied Toad der Bombina variegata (Linnaeus, 1758) – Speleomantes imperialis (Stefani, 1969) – Yellow-bellied Toad Imperial Cave Salamander Family Pipidae Gray, 1825 Speleomantes italicus (Dunn, 1923) – Ital- Xenopus Wagler, 1827 ian Cave Salamander *Xenopus laevis (Daudin, 1802) – African Speleomantes sarrabusensis Lanza, Leo, Clawed Toad Forti, Cimmaruta, Caputo and Nascetti, Family Pelobatidae Bonaparte, 1850 2001 – Sette Fratelli Cave Salamander Pelobates Wagler, 1830 Speleomantes strinatii (Aellen, 1958) – Pelobates balcanicus Karaman, 1928 – Strinati’s Cave Salamander Balkan Spadefoot Toad Speleomantes supramontis (Lanza, Pelobates cultripes (Cuvier, 1829) – West- Nascetti and Bullini, 1986) – Supra- ern Spadefoot Toad monte Cave Salamander Pelobates fuscus (Laurenti, 1768) – Com- Family Proteidae Gray, 1825 mon Spadefoot Toad Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 171

Pelobates syriacus Boettger, 1889 – East- Hyla orientalis Bedriaga, 1890 – Eastern ern Spadefoot Toad Tree Frog Pelobates vespertinus (Pallas, 1771) – Hyla sarda (de Betta, 1857) – Tyrrhenian Pallas’s Spadefoot Toad Tree Frog Family Pelodytidae Bonaparte, 1850 Hyla savignyi Audouin, 1827 – Savigny’s Pelodytes Bonaparte, 1838 Tree Frog Pelodytes atlanticus Díaz-Rodríguez, Family Ranidae Batsch, 1796 Gehara, Márquez, Vences, Gonçalves, Pelophylax Fitzinger, 1843 Sequeira, Martínez-Solano and Pelophylax bedriagae (Camerano, 1882) – Tejedo, 2017 – Portuguese Parsley Levant Water Frog Frog Pelophylax cretensis (Beerli, Hotz, Tunner, Pelodytes caucasicus Boulenger, 1896 – Heppich and Uzzell, 1994) – Cretan Wa- Caucasian Parsley Frog ter Frog Pelodytes ibericus Sánchez-Herráiz, Bar- Pelophylax epeiroticus (Schneider, Sofi- badillo, Machordom and Sanchiz, 2000 – anidou and Kyriakopoulou-Sklavounou, Iberian Parsley Frog 1984) – Epirus Water Frog Pelodytes punctatus (Daudin, 1802) – Pars- Pelophylax kl. esculentus (Linnaeus, ley Frog 1758) – Edible Frog Family Bufonidae Gray, 1825 Pelophylax kl. grafi (Crochet, Dubois, Bufo Garsault, 1764 Ohler and Tunner, 1995) – Graf’s Hy- Bufo bufo (Linnaeus, 1758) – Common brid Frog Toad Pelophylax lessonae (Camerano, 1882) – Bufo spinosus Daudin, 1803 – Spiny Pool Frog Toad Pelophylax perezi (Seoane, 1885) – Iberian Bufotes Rafinesque, 1815 Water Frog Bufotes boulengeri (Lataste, 1879) – Pelophylax ridibundus (Pallas, 1771) – African Green Toad Marsh Frog Bufotes cypriensis Litvinchuk, Mazepa, Pelophylax shqipericus (Hotz, Uzzell, Jablonski and Dufresnes in Dufresnes Günther, Tunner and Heppich, 1987) – et al., 2019 – Cyprus Green Toad Albanian Pool Frog Bufotes viridis (Laurenti, 1768) – Green Lithobates Fitzinger, 1843 Toad *Lithobates catesbeianus (Shaw, 1802) – Epidalea Cope, 1864 American Bullfrog Epidalea calamita (Laurenti, 1768) – Rana Linnaeus, 1758 Natterjack Toad Rana arvalis Nilsson, 1842 – Moor Frog Family Hylidae Rafinesque, 1815 Rana dalmatina Fitzinger in Bonaparte, Hyla Laurenti, 1768 1838 – Agile Frog Hyla arborea (Linnaeus, 1758) – Common Rana graeca Boulenger, 1891 – Greek Tree Frog Stream Frog Hyla intermedia Boulenger, 1882 – Italian Rana iberica Boulenger, 1879 – Iberian Tree Frog Stream Frog *Hyla meridionalis Boettger, 1874 – Rana italica Dubois, 1987 – Italian Stream Stripeless Tree Frog Frog Hyla molleri Bedriaga, 1889 – Iberian Tree Rana latastei Boulenger, 1879 – Italian Frog Agile Frog Downloaded from Brill.com10/05/2021 05:01:43PM via free access 172 J. Speybroeck et al.

Rana macrocnemis Boulenger, 1885 – Mauremys rivulata (Valenciennes, 1833) – Caucasian Frog Balkan Terrapin Rana parvipalmata Seoane, 1885 – Gali- Family Emydidae Rafinesque, 1815 cian Common Frog Emys Duméril, 1805 Rana pyrenaica Serra-Cobo, 1993 – Pyre- Emys orbicularis (Linnaeus, 1758) – Euro- nean Stream Frog pean Pond Terrapin Rana temporaria Linnaeus, 1758 – Com- Trachemys Agassiz, 1857 mon Frog *Trachemys scripta (Thunberg in Schoepff, 1792) – Pond Slider (Red- Class Reptilia Laurenti, 1768 eared Slider for ssp. elegans) Order Testudines Linnaeus, 1758 or Chelonii Family Trionychidae Fitzinger, 1826 Brongniart, 1800 Trionyx Geoffroy, 1827 Trionyx triunguis (Forskål, 1775) – Nile Family Cheloniidae Oppel, 1811 Soft-shelled Turtle Caretta Rafinesque-Schmaltz, 1814 Caretta caretta (Linnaeus, 1758) – Log- Order Squamata Oppel, 1811 gerhead Turtle Family Agamidae Spix, 1825 Chelonia Brongniart, 1800 Laudakia Gray, 1845 Chelonia mydas (Linnaeus, 1758) – Green Laudakia caucasia (Eichwald, 1831) – Turtle Caucasian Agama Eretmochelys Fitzinger, 1843 Laudakia stellio (Linnaeus, 1758) – Eretmochelys imbricata (Linnaeus, 1766) – Starred Agama Hawksbill Turtle Phrynocephalus Kaup, 1825 Lepidochelys Fitzinger, 1843 Phrynocephalus guttatus (Gmelin, 1789) – Lepidochelys kempii (Garman, 1880) – Toad-headed Agama Phrynocephalus helioscopus (Pallas, Kemp’s Ridley 1771) – Sunwatcher Toad-headed Lepidochelys olivacea (Eschscholtz, Agama 1829) – Olive Ridley Phrynocephalus mystaceus (Pallas, Family Dermochelyidae Fitzinger, 1843 (1825) 1776) – Secretive Toad-headed Agama Dermochelys de Blainville, 1816 Trapelus Cuvier, 1817 Dermochelys coriacea (Vandelli, 1761) – Trapelus agilis (Olivier, 1804) – Brilliant Leatherback Turtle Ground Agama Family Testudinidae Batsch, 1788 Family Chamaeleonidae Rafinesque, 1815 Testudo Linnaeus, 1758 Chamaeleo Laurenti, 1768 Testudo graeca Linnaeus, 1758 – Spur- *Chamaeleo africanus Laurenti, thighed Tortoise 1768 – African Chameleon Testudo hermanni Gmelin, 1789 – Her- Chamaeleo chamaeleon (Linnaeus, mann’s Tortoise 1758) – Mediterranean Chameleon Testudo marginata Schoepff, 1792 – Family Sphaerodactylidae Underwood, 1954 Marginated Tortoise Euleptes Fitzinger, 1843 Family Geoemydidae Theobald, 1868 Euleptes europaea (Gené, 1839) – Euro- Mauremys Gray, 1869 pean Leaf-toed Gecko Mauremys caspica (Gmelin, 1774) – Family Gekkonidae Oppel, 1811 Caspian Terrapin Alsophylax Fitzinger, 1843 Mauremys leprosa (Schweigger, 1812) – Alsophylax pipiens (Pallas, 1827) – Spanish Terrapin Caspian Even-fingered Gecko Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 173

Hemidactylus Oken, 1817 Algyroides marchi Valverde, 1958 – Span- Hemidactylus turcicus (Linnaeus, 1758) – ish Algyroides Turkish Gecko Algyroides moreoticus Bibron and Bory de Mediodactylus Szczerbak and Golubev, 1977 Saint-Vincent, 1833 – Greek Algyroides Mediodactylus bartoni (Štepánek,ˇ Algyroides nigropunctatus (Duméril and 1934) – Cretan Bent-toed Gecko Bibron, 1839) – Dalmatian Algyroides Mediodactylus danilewskii (Strauch, Anatololacerta Arnold, Arribas and Car- 1887) – Bulgarian Bent-toed Gecko ranza, 2007 Mediodactylus kotschyi (Steindachner, Anatololacerta anatolica (Werner, 1870) – Kotschy’s Bent-toed Gecko 1900) – Anatolian Rock Lizard Mediodactylus oertzeni (Boettger, 1888) – Anatololacerta budaki (Eiselt and Oertzen’s Bent-toed Gecko Schmidtler, 1986) – Budak’s Rock Mediodactylus orientalis (Štepánek,ˇ Lizard 1937) – Eastern Bent-toed Gecko Anatololacerta pelasgiana (Mertens, Mediodactylus russowii (Strauch, 1887) – 1959) – Pelasgian Rock Lizard Russow’s Bent-toed Gecko Archaeolacerta Mertens, 1921 Tenuidactylus Szczerbak and Golubev, Archaeolacerta bedriagae (Camerano, 1984 1885) – Tyrrhenian Rock Lizard *Tenuidactylus bogdanovi Nazarov and Dalmatolacerta Arnold, Arribas and Car- Poyarkov, 2013 – Bogdanov’s Bent- ranza, 2007 toed Gecko Dalmatolacerta oxycephala Duméril and Tenuidactylus caspius (Eichwald, 1831) – Bibron, 1839 – Sharp-snouted Rock Caspian Bent-toed Gecko Lizard Family Phyllodactylidae Gamble, Bauer, Darevskia Arribas, 1997 Greenbaum and Jackman, 2008 Darevskia alpina (Darevsky, 1967) – Tarentola Gray, 1825 Alpine Rock Lizard Tarentola angustimentalis Steindachner, *Darevskia armeniaca (Méhely, 1909) – 1891 – East Canary Gecko Armenian Rock Lizard Tarentola boettgeri Steindachner, 1891 – Darevskia brauneri (Méhely, Gran Canaria Gecko 1909) – Brauner’s Rock Lizard Tarentola delalandii Duméril and Bibron, Darevskia caucasica (Méhely, 1909) – 1836 – Tenerife Gecko Caucasus Rock Lizard Tarentola gomerensis Joger and Bischoff, Darevskia daghestanica (Darevsky, 1983 – La Gomera Gecko 1967) – Dagestan Rock Lizard Tarentola mauritanica (Linnaeus, 1758) – *Darevskia dahli (Darevsky, 1957) – Moorish Gecko Dahl’s Rock Lizard Family Lacertidae Batsch, 1788 Darevskia derjugini (Nikolsky, 1898) – Acanthodactylus Wiegmann, 1834 Derjugin’s Rock Lizard Acanthodactylus erythrurus (Schinz, Darevskia lindholmi (Lantz and Cyrén, 1833) – Spiny-footed Lizard 1936) – Crimean Rock Lizard Acanthodactylus schreiberi Boulenger, Darevskia praticola (Eversmann, 1834) – 1918 – Schreiber’s Spiny-footed Lizard Meadow Lizard Algyroides Bibron and Bory de Saint-Vincent, Darevskia rudis (Bedriaga, 1886) – 1833 Rough-tailed Rock Lizard Algyroides fitzingeri (Wiegmann, 1834) – Darevskia saxicola (Eversmann, 1834) – Pygmy Algyroides North-western Caucasus Rock Lizard Downloaded from Brill.com10/05/2021 05:01:43PM via free access 174 J. Speybroeck et al.

Dinarolacerta Arnold, Arribas and Carranza, Iberolacerta martinezricai (Arribas, 2007 1996) – Peña de Francia Rock Lizard Dinarolacerta montenegrina Ljubisavljevic,´ Iberolacerta monticola (Boulenger, Arribas, Džukic´ and Carranza, 2007 – 1905) – West Iberian Rock Lizard Prokletije Rock Lizard Lacerta Linnaeus, 1758 Dinarolacerta mosorensis Kolombatovic,´ Lacerta agilis Linnaeus, 1758 – Sand 1886 – Mosor Rock Lizard Lizard Eremias Fitzinger in Wiegmann, 1834 Lacerta bilineata Daudin, 1802 – Western Eremias arguta (Pallas, 1773) – Steppe Green Lizard Runner Lacerta citrovittata Werner, 1935 – Cy- Eremias velox (Pallas, 1771) – Rapid clades Green Lizard Steppe Runner Lacerta diplochondrodes Wettstein, Gallotia Boulenger, 1916 1952 – Eastern Balkan Green Lizard Gallotia atlantica (Peters and Doria, Lacerta media (Lantz and Cyrén, 1920) – 1882) – Atlantic Lizard Middle East Green Lizard Gallotia bravoana Hutterer, 1985 – La Lacerta schreiberi Bedriaga, 1878 – Gomera Giant Lizard Schreiber’s Green Lizard Gallotia caesaris (Lehrs, 1914) – Lacerta strigata (Eichwald, 1831) – Boettger’s Lizard Caspian Green Lizard Gallotia galloti (Oudart in Webb and Lacerta trilineata Bedriaga, 1886 – Balkan Berthelor, 1939) – Tenerife Lizard Green Lizard Gallotia intermedia Hernández, Nogales Lacerta viridis (Laurenti, 1768) – Eastern and Martín, 2000 – Tenerife Speckled Green Lizard Lizard Ophisops Ménétries, 1832 Gallotia simonyi (Steindachner, 1889) – El Ophisops elegans Ménétries, 1832 – Hierro Giant Lizard Snake-eyed Lacertid Gallotia stehlini (Schenkel, 1901) – Gran Parvilacerta Arnold, Arribas and Carranza, Canaria Giant Lizard 2007 Hellenolacerta Arnold, Arribas and Car- Parvilacerta parva (Boulenger, 1887) – ranza, 2007 Dwarf Lizard Hellenolacerta graeca Bedriaga, 1886 – Phoenicolacerta Arnold, Arribas and Car- Greek Rock Lizard ranza, 2007 Iberolacerta Arribas, 1997 Phoenicolacerta troodica (Werner, 1936) – Iberolacerta aranica (Arribas, 1993) – Troodos Rock Lizard Aran Rock Lizard Podarcis Wagler, 1830 Iberolacerta aurelioi (Arribas, 1994) – Au- Podarcis bocagei (Seoane, 1884) – relio’s Rock Lizard Bocage’s Wall Lizard Iberolacerta bonnali (Lantz, 1927) – Pyre- Podarcis carbonelli Pérez-Mellado, 1981 – nean Rock Lizard Carbonell’s Wall Lizard Iberolacerta cyreni (Müller and Hellmich, Podarcis cretensis (Wettstein, 1952) – Cre- 1937) – Cyren’s Rock Lizard tan Wall Lizard Iberolacerta galani Arribas, Carranza and Podarcis erhardii (Bedriaga, 1876) – Er- Odierna, 2006 – Galan’s Rock Lizard hard’s Wall Lizard Iberolacerta horvathi (Méhely, 1904) – Podarcis filfolensis (Bedriaga, 1876) – Horvath’s Rock Lizard Maltese Wall Lizard Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 175

Podarcis gaigeae (Werner, 1930) – Skyros Psammodromus edwarsianus (Dugès, Wall Lizard 1829) – Edwards’s Psammodromus Podarcis guadarramae (Boscá, 1916) – Psammodromus occidentalis Fitze, Guadarrama Wall Lizard Gonzalez-Jimena, San-Jose, San Podarcis hispanicus (Steindachner, Mauro and Zardoya, 2012 – Western 1870) – Spanish Wall Lizard Psammodromus Podarcis ionicus (Lehrs, 1902) – Ionian Scelarcis Fitzinger, 1843 Wall Lizard Scelarcis perspicillata (Duméril and Podarcis levendis Lymberakis, Poulakakis, Bibron, 1839) – Moroccan Rock Lizard Kaliontzopoulou, Valakos and Mylonas, Teira Gray, 1838 2008 – Pori Wall Lizard Teira dugesii (Milne-Edwards, 1829) – Podarcis lilfordi (Günther, 1874) – Lil- Madeira Rock Lizard ford’s Wall Lizard Timon Tschudi, 1836 Podarcis liolepis (Boulenger, 1905) – Cat- Timon lepidus (Daudin, 1802) – Ocellated alonian Wall Lizard Lizard Podarcis melisellensis (Braun, 1877) – Timon nevadensis (Buchholz, 1963) – Dalmatian Wall Lizard Sierra Nevada Ocellated Lizard Podarcis milensis (Bedriaga, 1882) – Mi- Zootoca Wagler, 1830 los Wall Lizard Zootoca carniolica Mayer, Böhme, Podarcis muralis (Laurenti, 1768) – Com- Tiedemann and Bischoff, 2000 – mon Wall Lizard Carniolian Lizard Podarcis peloponnesiacus (Bibron and Zootoca vivipara (Jacquin, 1787) – Bory de Saint-Vincent, 1833) – Pelo- Viviparous Lizard ponnese Wall Lizard Family Scincidae Oppel, 1811 Podarcis pityusensis (Boscá, 1883) – Ibiza Ablepharus Fitzinger in Eversmann, 1823 Wall Lizard Ablepharus kitaibelii Bibron and Bory de Podarcis raffonei (Mertens, 1952) – Aeo- Saint-Vincent, 1833 – Snake-eyed Skink lian Wall Lizard Ablepharus budaki Göcmen, Kumlutas and Podarcis siculus (Rafinesque-Schmaltz, 1810) – Italian Wall Lizard Tosunoglu, 1996 – Budak’s Snake-eyed Podarcis tauricus (Pallas, 1814) – Balkan Skink Wall Lizard Chalcides Laurenti, 1768 Podarcis tiliguerta (Gmelin, 1789) – Chalcides bedriagai (Boscá, 1880) – Tyrrhenian Wall Lizard Bedriaga’s Skink Podarcis vaucheri (Boulenger, 1905) – Chalcides chalcides (Linnaeus, 1758) – Vaucher’s Wall Lizard Italian Three-toed Skink Podarcis virescens Geniez, Sá-Sousa, Chalcides coeruleopunctatus Salvador, Guillaume, Cluchier and Crochet, 1975 – La Gomera Skink 2014 – Geniez’s Wall Lizard Chalcides ocellatus (Forskål, 1775) – Podarcis waglerianus Gistel, 1868 – Sicil- Ocellated Skink ian Wall Lizard Chalcides sexlineatus Steindachner, Psammodromus Fitzinger, 1826 1891 – Gran Canaria Skink Psammodromus algirus (Linnaeus, 1758) – Chalcides simonyi Steindachner, 1891 – Large Psammodromus East Canary Skink Psammodromus hispanicus Fitzinger, Chalcides striatus (Cuvier, 1829) – Iberian 1826 – Spanish Psammodromus Three-toed Skink Downloaded from Brill.com10/05/2021 05:01:43PM via free access 176 J. Speybroeck et al.

Chalcides viridanus (Gravenhorst, 1851) – Xerotyphlops Hedges, Marion, Lipp, Marin West Canary Skink and Vidal, 2014 Eumeces Wiegmann, 1834 Xerotyphlops vermicularis (Merrem, Eumeces schneiderii (Daudin, 1802) – 1820) – Worm Snake Schneider’s Skink Family Erycidae Bonaparte, 1840 Heremites Gray, 1845 Eryx Daudin, 1803 Heremites auratus (Linnaeus, 1758) – Eryx jaculus (Linnaeus, 1758) – Sand Boa Levant Skink Eryx miliaris (Pallas, 1773) – Dwarf Sand Heremites vittatus (Olivier, 1804) – Bri- Boa dled Skink Family Psammophiidae Boie, 1827 Ophiomorus Duméril and Bibron, 1839 Malpolon Fitzinger, 1826 Ophiomorus kardesi Kornilios, Malpolon insignitus (Geoffroy Saint- Kumluta¸s, Lymberakis and Ilgaz, Hilaire, 1827) – Eastern Montpellier 2017 – Anatolian Limbless Skink Snake Ophiomorus punctatissimus (Bibron and Malpolon monspessulanus (Hermann, Bory de Saint-Vincent, 1833) – Limb- 1804) – Western Montpellier Snake less Skink Family Natricidae Bonaparte, 1840 Natrix Laurenti, 1768 Family Anguidae Gray, 1825 Natrix astreptophora (Seoane, 1884) – Anguis Linnaeus, 1758 Iberian Grass Snake Anguis cephallonica Werner, 1894 – Pelo- Natrix helvetica (Lacépède, 1789) – ponnese Slow Worm Barred Grass Snake Anguis colchica (Nordmann, 1840) – East- Natrix maura (Linnaeus, 1758) – Viperine ern Slow Worm Snake Anguis fragilis Linnaeus, 1758 – Slow Natrix natrix (Linnaeus, 1758) – Grass Worm Snake Anguis graeca Bedriaga, 1881 – Greek Natrix tessellata (Laurenti, 1768) – Dice Slow Worm Snake Anguis veronensis Pollini, 1818 – Italian Family Colubridae Oppel, 1811 Slow Worm Coronella Laurenti, 1768 Pseudopus Merrem, 1820 Coronella austriaca Laurenti, 1768 – Pseudopus apodus (Pallas, 1775) – Glass Smooth Snake Lizard Coronella girondica (Daudin, 1803) – Family Blanidae Kearney, 2003 Southern Smooth Snake Blanus Wagler, 1830 Dolichophis Gistel, 1868 Blanus cinereus (Vandelli, 1797) – Iberian Dolichophis caspius (Gmelin, 1789) – Worm Lizard Caspian Whip Snake Blanus rufus (Hemprich, 1820) – Cen- Dolichophis jugularis (Linnaeus, 1758) – tral Iberian Worm Lizard Black Whip Snake Blanus strauchi (Bedriaga, 1884) – Anato- Dolichophis schmidti (Nikolsky, 1909) – lian Worm Lizard Schmidt’s Whip Snake Family Typhlopidae Merrem, 1820 Eirenis Jan, 1863 Indotyphlops Hedges, Marion, Lipp, Marin Eirenis collaris (Ménétries, 1832) – Col- and Vidal, 2014 lared Dwarf Snake *Indotyphlops braminus (Daudin, 1803) – Eirenis levantinus Schmidtler, 1993 – Lev- Flowerpot Snake ant Dwarf Snake Downloaded from Brill.com10/05/2021 05:01:43PM via free access Species list of European herpetofauna 177

Eirenis modestus (Martin, 1838) – Masked Telescopus fallax (Fleischmann, 1831) – Dwarf Snake Cat Snake Elaphe Fitzinger, 1833 Zamenis Wagler, 1830 Elaphe dione (Pallas, 1773) – Steppe Zamenis hohenackeri (Strauch, 1873) – Snake Transcaucasian Rat Snake Elaphe quatuorlineata (Bonnaterre, Zamenis lineatus (Camerano, 1891) – Ital- 1790) – Four-lined Snake ian Aesculapian Snake Elaphe sauromates (Pallas, 1814) – Zamenis longissimus (Laurenti, 1768) – Blotched Snake Aesculapian Snake *Elaphe schrenckii Strauch, 1876 – Amur Zamenis scalaris (Schinz, 1822) – Lad- Rat Snake der Snake Elaphe urartica Jablonski, Kukushkin, Zamenis situla (Linnaeus, 1758) – Leopard Avcı, Bunyatova, Ilgaz, Tuniyev and Snake Jandzik in Jablonski et al., 2019 Family Viperidae Oppel, 1811 Macrovipera Reuss, 1927 Hemorrhois Boie, 1826 Macrovipera lebetinus (Linnaeus, 1758) – *Hemorrhois algirus (Jan, 1863) – Alge- Blunt-nosed Viper rian Whip Snake Macrovipera schweizeri (Werner, 1935) – Hemorrhois hippocrepis (Linnaeus, Milos Viper 1758) – Horseshoe Whip Snake Montivipera Nilson, Tuniyev, Andrén, Orlov, Hemorrhois nummifer (Reuss, 1834) – Joger and Herrmann, 1999 Coin-marked Snake Montivipera xanthina (Gray, 1849) – Ot- Hemorrhois ravergieri (Ménétries, 1832) – toman Viper Spotted Whip Snake Vipera Garsault, 1764 Hierophis Fitzinger in Bonaparte, 1834 Vipera ammodytes (Linnaeus, 1758) – Hierophis gemonensis (Laurenti, 1768) – Nose-horned Viper Balkan Whip Snake Vipera aspis (Linnaeus, 1758) – Asp Viper Hierophis viridiflavus (Lacépède, 1789) – Vipera berus (Linnaeus, 1758) – Adder Western Whip Snake Vipera dinniki (Nikolsky, 1913) – Dinnik’s Hierophis cypriensis (Schätti, 1985) – Viper Cyprus Whip Snake Vipera graeca Nilson and Andrén, 1988 – Lampropeltis Fitzinger, 1843 Greek Meadow Viper *Lampropeltis getula (Linnaeus, 1766) – Vipera kaznakovi (Nikolsky, 1909) – Cau- Common Kingsnake casus Viper Macroprotodon Guichenot, 1850 Vipera latastei (Boscá, 1878) – Lataste’s Macroprotodon brevis (Günther, 1862) – Viper Iberian False Smooth Snake Vipera renardi (Christoph, 1861) – Steppe *Macroprotodon cucullatus (Geoffroy Viper Saint-Hilaire, 1827) – Algerian False Vipera seoanei (Lataste, 1879) – Seoane’s Smooth Snake Viper Platyceps Blyth, 1860 Vipera ursinii (Bonaparte, 1835) – Platyceps collaris (Müller, 1878) – Red- Meadow Viper dish Whip Snake Platyceps najadum (Eichwald, 1831) – Acknowledgements. We thank the editor Salvador Car- Dahl’s Whip Snake ranza, as well as Shai Meiri and an anonymous reviewer for Telescopus Wagler, 1830 their constructive contributions to improve our manuscript. Downloaded from Brill.com10/05/2021 05:01:43PM via free access 178 J. Speybroeck et al.

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