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Review Species List of the European Herpetofauna 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] Downloaded from Brill.com03/08/2021 01:42:59PM © Speybroeck et al., 2020. DOI:10.1163/15685381-bja10010via Museo Nacional de Ciencias Naturales This is an open access article distributed under the terms of the CC-BY 4.0 License. 140 J. Speybroeck et al. 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.com03/08/2021 01:42:59PM via Museo Nacional de Ciencias Naturales 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.com03/08/2021 01:42:59PM via Museo Nacional de Ciencias Naturales 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
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