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

May 2020
Amphibia-Reptilia 41(2):139-189

DOI:10.1163/15685381-bja10010

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CC BY 4.0

Authors:

Jeroen Speybroeck

Research Institute for Nature and Forest

Wouter Beukema

Ravon

Christophe Dufresnes

Nanjing Forestry University

Uwe Fritz

Senckenberg Research Institute

Show all 12 authorsHide

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).

Extent of geographic area (Mollweide projection). Herpetofauna species within this area are dealt with. Shading indicates areas not included by Sillero et al. (2014).

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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.

1 - Research Institute for Nature and Forest, Havenlaan 88

bus 73, 1000 Brussel, Belgium

2 - Wildlife Health Ghent, Department of Pathology,

Bacteriology and Avian Diseases, Ghent University,

Salisburylaan 133, 9820 Merelbeke, Belgium

3 - LASER, College of Biology and the Environment,

Nanjing Forestry University, Nanjing, China

4 - Museum of Zoology, Senckenberg Dresden, A.B.

Meyer Building, Königsbrücker Landstraße 159,

01109 Dresden, Germany

5 - Department of Zoology, Comenius University in

Bratislava, Ilkoviˇ

cova 6, Mlynská dolina, 842 15

Bratislava, Slovakia

6 - Natural History Museum of Crete, University of Crete,

Knossou Ave. 71409, Crete, Irakleio, Greece

7 - Museo Nacional de Ciencias Naturales (MNCN-

CSIC), c/ José Gutiérrez Abascal, 2, 28006 Madrid,

Spain

8 - Kosmos – Museo di Storia Naturale, Università di

Pavia, Piazza Botta 9, 27100 Pavia, Italy

9 - Division of Evolutionary Biology, Zoological In-

stitute, Braunschweig University of Technology,

Mendelssohnstr. 4, 38106 Braunschweig, Germany

10 - Department of Zoology, Hungarian Natural History

Museum, 1088 Budapest, Baross u. 13, Hungary

Introduction

Speybroeck, Beukema and Crochet (2010)

(SBC2010, hereafter) provided an annotated

species list for the European amphibians and

non-avian reptiles. A decade later, a sizable

amount of new research has been produced,

fuelling the need for a contemporary update.

Within the European Herpetological Society

(Societas Europaea Herpetologica; SEH) and by

invitation of the SEH Council, a newly com-

posed Taxonomic Committee (SEH TC, or fur-

ther TC) was formed in early 2019, and its

chair was approved by SEH membership during

the Ordinary General Meeting held in Milan,

11 - CEFE, Université Montpellier, CNRS, EPHE, IRD,

Université Paul Valéry Montpellier 3, Montpellier,

France

∗Corresponding author;

e-mail: jeroen.speybroeck@inbo.be

©Speybroeck et al., 2020. DOI:10.1163/15685381-bja10010

This is an open access article distributed under the terms of the CC-BY 4.0 License.

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140 J. Speybroeck et al.

September 5th 2019 (SEH News, Amphibia-

Reptilia 40: 551-559).

We did not deﬁne our own limits for the geo-

graphical area considered here, but adopted the

limits deﬁned by previous projects. Our goal

was to provide a taxonomic reference for the fu-

ture mapping projects of the SEH. Therefore,

we included all areas that were part of pre-

vious European atlas projects (cf. Gasc et al.,

1997; Sillero et al., 2014). We also aimed at in-

forming the taxonomic backbone of the Fauna

Europaea initiative (https://fauna-eu.org), and

therefore our geographical area also includes all

territories that are covered by Fauna Europaea.

As a result, we enlarged the geographical area

considered by SBC2010 to encompass all areas

covered by both Gasc et al. (1997) and Fauna

Europaea. Areas included by Gasc et al. (1997),

but not by Speybroeck, Crochet and Beukema

(2010), include the northern versant of the Cau-

casus (including north-eastern Azerbaijan), all

areas west of the Ural River (including western-

most Kazakhstan) and west of the Ural Moun-

tains, and the Yekaterinburg Region. Areas in-

cluded in Fauna Europaea, but not by Gasc et

al. (1997) or SBC2010, are Macaronesia (with-

out Cape Verde), the Greek Islands off the west-

ern Anatolian shore, and Cyprus. As such, our

area exceeds that of the most recent European

atlas (Sillero et al., 2014) by including Mac-

aronesia, all Greek islands, and parts of Azer-

baijan and Kazakhstan (ﬁg. 1). A Google Earth

.kml ﬁle with the limits of the area is provided

in the supplementary material. For the rationale

of these limits, we refer to Gasc et al. (1997) and

https://fauna-eu.org/data-handling).

Upon enlarging the scope area and prior to

discussing any taxonomic changes, a broader

baseline list had to be set for species occur-

ring outside the area considered by SBC2010.

As such, in addition to SBC2010, we followed

the taxonomy of Gasc et al. (1997, 2004, includ-

ing the changes adopted by Dubois and Cro-

chet in the 2004 reprint), and for species outside

Europe (including Macaronesia and Cyprus),

Sindaco and Jeremˇ

cenko (2008) and Sindaco,

Venchi and Grieco (2013). Together, these four

sources led to a starting point species list. In the

following, we only discuss taxonomic changes

which deviate from this species list. We decided

against using online databases as starting points,

as they are changing constantly, with historical

versions not remaining reliably and easily avail-

able.

The taxonomic decisions adopted here are

not necessarily supported by all authors of this

work. According to TC guidelines, a change to

the starting list will only be adopted if widely

supported among its members, speciﬁcally by

a>75% majority. When a change is recom-

mended by a large majority of the TC mem-

bers, but different members favour different

outcomes, the adopted solution may be sup-

ported by only a simple majority (>50%). Note

that this process favours taxonomic stability,

with changes requiring large support among TC

members to become accepted.

TC members do not necessarily adhere to

the same species concept. While many agree

with the General Lineage Concept (GLC) of De

Queiroz (2007), some prefer the general frame-

work of the Biological Species Concept. How-

ever, all agree on using reproductive isolation as

the primary operational criterion for the delimi-

tation of species. The majority of TC members

is of the opinion that, while every species is a

lineage, not every lineage is a species. The com-

mon approach can be deﬁned as either following

the Biological Species Concept framework, or

as applying a Biological Species Criterion under

the GLC. More speciﬁcally, TC members ad-

here to a “soft” version of the reproductive iso-

lation criterion. As such, we allow extensive in-

trogression between recognised species, as long

as there are intrinsic barriers to gene ﬂow that

prevent wide-reaching introgression beyond the

contact zones. Even in the absence of geograph-

ical barriers, a sufﬁcient level of reproductive

isolation has to exist in order to ensure long-

term persistence of the diverged lineages. Taxa

connected by bimodal or trimodal hybrid zones

(Gay et al., 2008) were unanimously treated as

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Species list of European herpetofauna 141

Figure 1. Extent of geographic area (Mollweide projection). Herpetofauna species within this area are dealt with. Shading indicates areas not included by Sillero et al. (2014).

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142 J. Speybroeck et al.

valid species, but opinions often differed re-

garding how much introgression was “allowed”

across unimodal hybrid zones, reﬂecting differ-

ent opinions relative to where to cut the grey

zone of speciation, how much reproductive iso-

lation is necessary and when to treat incipient

species as valid species. For allopatric taxa, or

when contact zones were not studied, lineages

that had divergence levels similar to closely re-

lated, unambiguously distinct species were ac-

cepted as species. As an auxiliary criterion, we

sometimes use monophyly, even though we do

not consider it as a necessary requirement for

species status.

For supraspeciﬁc classiﬁcation, the TC

agreed to accept only monophyletic units.

This causes issues regarding the class Rep-

tilia, which in its traditional deﬁnition is pa-

raphyletic through the exclusion of birds. All

current hypotheses on the evolution of verte-

brates agree that a group including squamates,

turtles, Sphenodon and crocodiles, but not birds

is paraphyletic (see e.g. Chiari et al., 2012a;

Hasegawa, 2017). As a consequence, most cur-

rent classiﬁcations of Vertebrata include Aves in

the class Reptilia (see e.g. Modesto and Ander-

son, 2004; Ruggiero et al., 2015). To avoid con-

fusion, we adopt the term ‘non-avian reptiles’ to

refer to the components of the European fauna

assigned to Testudines/Chelonii and Squamata.

For nomenclatural decisions, including

spelling, we followed the International Code

of Zoological Nomenclature (the Code here-

after, ICZN (1999 and subsequent changes),

see https://www.iczn.org/). Such decisions were

generally not submitted to voting, but they could

be discussed, as many parts of the Code can be

subject to interpretation, and many actual cases

can be open to different decisions, even under

the rules of the Code.

In the following, we review taxonomic and

nomenclatural changes proposed since the pub-

lication of the four literature sources that we

used to build our starting point, as well as other

relevant new information pertaining the taxon-

omy of European amphibians and non-avian

reptiles. For each case, we provide the rationale

underlying the respective decision of the TC,

and conclude by providing an updated species

list of the European herpetofauna.

Amphibia

Caudata/Urodela

A series of phylogenetic studies on mitochon-

drial DNA, allozymes, and nuclear DNA se-

quences of members of the family Salamandri-

dae (Litvinchuk et al., 2005; Weisrock et al.,

2006; Zhang et al., 2008; Kieren et al., 2018;

Veith et al., 2018) conﬁrmed that the newt genus

Triturus sensu lato, as traditionally recognised,

is not monophyletic. Litvinchuk et al. (2005)

proposed the separation of Triturus into four

genera, among which the new genus Omma-

totriton contains (the former) Triturus vitta-

tus. SBC2010 accepted this new arrangement,

but did not explicitly acknowledge the need to

recognise Ommatotriton, as the new genus does

not occur in the area they considered. As we

herein consider a wider area, we formally ac-

cept Ommatotriton as a separate genus.

Litvinchuk et al. (2005) also showed that

morphology (number of trunk vertebrae, colour

pattern), genome size, and allozymes (Nei’s

genetic distances of 0.44-0.83) strongly dif-

fer between populations in the two widely dis-

joint areas inhabited by Ommatotriton vittatus,

namely south-eastern Anatolia and the Levant

(O. v. vittatus) versus the southern and eastern

Black Sea and western Caucasus regions (O. v.

ophryticus). This led them to elevate ophryticus

to species level. A more comprehensive study

also demonstrated restricted introgression be-

tween an eastern and a western taxon along the

Black Sea coast of Turkey (van Riemsdijk et al.,

2017). Consequently, we recommend to recog-

nise three species in the genus Ommatotriton:

O. vittatus (extralimital) in south-eastern Anato-

lia and the Levant, O. ophryticus in Russia, the

Caucasus and northern Anatolia west to the re-

gion of Samsun, and O. nesterovi (extralimital)

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Species list of European herpetofauna 143

in Anatolia from Samsun to the Sea of Marmara.

The populations inhabiting the native Omma-

totriton range in the European part of Russia be-

long to O. ophryticus, while the introduced (and

persisting anno 2019; Speybroeck, pers. obs.)

population in north-eastern Spain is of mixed

ancestry, with genetic contribution from both O.

ophryticus and O. nesterovi (van Riemsdijk et

al., 2018).

In addition to mtDNA sequences, Vences et

al. (2014) used sequences from thirteen nuclear

loci to improve our understanding of the his-

tory of the genus Salamandra. Their nuclear

data suggest that Salamandra salamandra con-

tains several deeply divergent lineages whose

monophyly relative to S. algira received weak

support, although this requires conﬁrmation.

Based on mitochondrial data the subspecies lon-

girostris is sister to all other S. salamandra lin-

eages. However, nuclear genes place it with the

subspecies morenica. The discordance between

mtDNA and nuclear genes may result from past

introgression and admixture processes. Further-

more, a clade comprising the subspecies fas-

tuosa,bernardezi (including populations at-

tributed to the subspecies alfredschmidti, whose

validity was rejected, as it is phylogenetically

nested within several subclades of bernardezi;

Beukema et al., 2016) and gigliolii is recov-

ered with strong support. While additional data

are clearly needed, this suggests a high amount

of evolutionary divergence within S. salaman-

dra, and the existence of more than one species

within S. salamandra cannot yet be fully ruled

out.

SBC2010 recognised two species within the

former Triturus karelinii:T. arntzeni from the

Balkan Peninsula, and T. karelinii. Wielstra et

al. (2013) suggested that the type specimens of

T. arntzeni are in fact T. macedonicus, which led

them to place T. arntzeni in the synonymy of

T. macedonicus, and to create the name ivanbu-

reschi for the taxon of the karelinii complex that

occurs in the Balkans and in Western Anato-

lia. Later, Wielstra and Arntzen (2014) demon-

strated that the types of T. arntzeni are in fact the

result of ancient admixture between T. mace-

donicus and T. ivanbureschi. We do not entirely

agree with their interpretation of the Code. As

they rightfully state, nomina based on geneti-

cally admixed individuals derived through sev-

eral generations of backcrossing are indeed left

in limbo in the Code. Yet, we do not agree that

the Code should be interpreted as extending pro-

visions of Art. 23.8 to nomina based on genet-

ically admixed individuals, at least not without

clear guidelines of what constitutes admixture

in the sense of the Code. We do, however, agree

with Wielstra and Arntzen (2014) that arntzeni,

based on individuals that carry less than 50%

of alleles derived from the karelinii-complex

taxon, should not be used as the valid name for

that taxon. We thus accept T. ivanbureschi as

the valid nomen of the taxon of the karelinii-

complex that occurs in Europe in the Balkans.

With the extension of the geographic range of

SBC2010, T. karelinii sensu stricto also occurs

in our area in Russia, the Crimean Peninsula and

the Caucasus region.

Using mitochondrial and nuclear DNA se-

quences, Sotiropoulos et al. (2007) and Recuero

et al. (2014) investigated the phylogenetic struc-

ture of Ichthyosaura alpestris. They identiﬁed

several deeply divergent lineages that largely

correspond to currently recognised subspecies,

except for the subspecies alpestris, which is

further divided into a western and an eastern

main lineage. Interestingly, nuclear sequences

from two loci show a nearly complete lack of

allele sharing between the eastern and west-

ern clades, suggesting reproductive isolation be-

tween them. However, no samples were col-

lected close to their contact zones. As the east-

ern and western clades should meet in the

Balkans and the southern Carpathian Moun-

tains, we follow the recommendation of Re-

cuero et al. (2014) and await more data on

their level of reproductive isolation before mak-

ing any taxonomic change. Thus, we do not

follow Raffaëlli (2018) who (based on the ge-

netic data presented in the aforementioned stud-

ies) elevated the subspecies apuana,reiseri and

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144 J. Speybroeck et al.

veluchiensis to species level, and we note that

such changes would make I. alpestris poly-

phyletic. We thus retain I. alpestris as a single

species for now.

Pabijan et al. (2017) reconstructed phyloge-

netic relationships within the Lissotriton vul-

garis species complex and inferred patterns of

(historical) gene ﬂow, using 74 nuclear DNA

markers from one individual from each of 127

locations. Five highly divergent lineages were

identiﬁed within our focal area: Lissotriton

montandoni and four lineages corresponding

to the Lissotriton vulgaris subspecies graecus,

lantzi,schmidtleri and vulgaris. In spite of clear

evidence of past historical introgression, their

contemporary gene ﬂow is restricted. Therefore,

the authors proposed to treat the three former

subspecies as species (Pabijan et al., 2017). Be-

tween L. v. vulgaris and the morphologically

diverged L. v. meridionalis regular episodes of

gene ﬂow were identiﬁed, thus meridionalis

was retained at subspecies level. While L. v.

graecus is easily distinguished from other Eu-

ropean populations by male nuptial character-

istics, both L. v. lantzi and L. v. schmidtleri

are morphologically cryptic in respect to L. v.

vulgaris (Raxworthy, 1990). Rather than be-

ing conﬁned to Anatolia, Pabijan et al. (2015,

2017) showed that L. v. schmidtleri also occurs

in Greek and Turkish Thrace and on a number

of Greek islands. Because the sampling gaps be-

tween several of the lineages remained wide,

the TC has been reluctant to accept all syste-

matic conclusions of Pabijan et al. (2017). The

taxon graecus comes into close contact with L.

v. vulgaris along the northern and eastern bor-

ders of its distribution. Although no dense sam-

pling has been performed to delineate the con-

tact zone in detail, and mitochondrial introgres-

sion from southern L. v. vulgaris into graecus

occurs at its northern range border (Pabijan et

al., 2017), nuclear gene pools of these two taxa

appear to remain reciprocally distinct in rela-

tive close geographical proximity (Pabijan et al.,

2017; Wielstra et al., 2018). We therefore accept

Lissotriton graecus as a valid species. For L. v.

schmidtleri, its range limits and contact zones

with other European lineages remain poorly

known. Although data on the contact zone with

L. v. kosswigi in Anatolia support a species-level

divergence between L. v. schmidtleri and the

graecus-kosswigi clade, areas of 100-300 km

inhabited by L. vulgaris s.l. of unknown iden-

tity separate genotyped L. v. schmidtleri popu-

lations from those of L. v. vulgaris and L. grae-

cus (Pabijan et al., 2017; Wielstra et al., 2018).

The TC thus feels that more information on con-

tact zones between L. v. schmidtleri and L. v.

vulgaris is warranted before the species rank of

schmidtleri can be accepted. Finally, L. v. lantzi

is endemic to the Caucasus and shows an al-

lopatric distribution (Wielstra et al., 2018). Mi-

tochondrial data suggests that L. v. lantzi was

the ﬁrst to diverge from other lineages around

3.39 (1.42-5.37) Mya (Pabijan et al., 2015). Yet,

nuclear data place L. v. lantzi as sister to L. v.

schmidtleri and other eastern and central Eu-

ropean Lissotriton, while suggesting that diver-

gence within this group initiated with the split

of the graecus-kosswigi clade (Pabijan et al.,

2017). In conclusion, the TC recommends to

treat Lissotriton graecus as a valid species, but,

as awarding species status to lantzi but not to

schmidtleri may render L. vulgaris paraphyletic,

we prefer to maintain the other taxa, including

L. v. schmidtleri and L. v. lantzi, as subspecies

of L. vulgaris for the time being.

A phylogeographic study based on two

mtDNA markers by Martínez-Solano et al.

(2006) revealed two major mitochondrial lin-

eages of Miocene origin in Lissotriton boscai.

One of them is restricted to central and south-

western coastal Portugal, while the other oc-

cupies the remainder of the species range, in-

cluding the type locality of L. boscai. Martínez-

Solano et al. (2006) acknowledged that the

two lineages might represent cryptic species,

but called for additional morphological and

molecular studies, including data on variation

in nuclear DNA markers. Dubois and Raffaelli

(2009) resurrected the nomen Triton maltzani

Boettger, 1879 for the southwestern lineage in

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Species list of European herpetofauna 145

the new combination Lissotriton maltzani.Ac-

cording to these authors, L. maltzani can be dis-

tinguished from L. boscai by its smaller size

and by its paler dorsal coloration, especially in

females, with less distinct dark spots. Teixeira

et al. (2015) used DNA sequences of one nu-

clear gene and found overall congruence with

mtDNA genes in terms of sequence divergence

and geographic structure. They, however, also

revealed wide areas of admixture and evidence

for recombination, suggesting a lack of com-

plete reproductive isolation and the presence of

incomplete speciation. In a recent study, Se-

queira et al. (2020) used ten microsatellites, one

mtDNA gene and two single copy nuclear DNA

markers in a cline analysis framework to inves-

tigate one of the hybrid zones between L. boscai

and L. maltzani. The results show evidence for

partial reproductive isolation between L. boscai

and L. maltzani, with narrow clines (3-28 km)

consistent with selection against hybrids. We

thus recognise L. maltzani as a separate species.

Wake (2012) addressed the taxonomy of

the Plethodontidae, advocating to treat Aty-

lodes (comprising Speleomantes genei) and

Speleomantes (comprising the other European

plethodontid species) as subgenera of a single,

cross-Atlantic genus Hydromantes. Addressing

the genus name confusion, Wake (2013) com-

pared ﬁve potential arrangements, missing how-

ever the arrangement of SBC2010 (i.e. two

genera without subgenera). Comparing with

other plethodontid genera, he argued in favour

of an arrangement of a single genus with

three subgenera. However, the arguments of

SBC2010 still stand: the position of Atylodes

remains unresolved, and the European species

form a well-deﬁned monophyletic group with a

large genetic distance from the ﬁve Californian

species Hydromantes brunus,H. platycephalus,

H. samweli,H. shastae, and H. wintu (Nascetti

et al., 1996; Pyron and Wiens, 2011; Chiari

et al., 2012b; Bingham, Papenfuss and Wake,

2018). Thus, no change seems in order, and we

maintain the European species for the time be-

ing in a single taxon, the genus Speleomantes.

Vörös, Ursenbacher and Jeli´

c (2019) used 10

microsatellite loci to investigate patterns of dif-

ferentiation between four Croatian cave popu-

lations of Proteus anguinus. They uncovered

long-lasting isolation between caves belong-

ing to different hydrogeographic systems, with

the most ancient divergence being older than 7

Mya. This suggests that some of the evolution-

ary lineages within this species might constitute

cryptic taxa, possibly of species rank (Vörös,

Ursenbacher and Jeli´

c, 2019).

Anura

Alytes obstetricans is composed of four sub-

species: A. o. obstetricans,A. o. pertinax,A. o.

boscai and A. o. almogavarii. The latter taxon

is endemic to Catalonia and adjacent areas in

north-eastern Spain and southern France, and

it is highly differentiated in allozyme (Arntzen

and García-París, 1995; García-París, 1995),

mitochondrial (Gonçalves et al., 2007, 2015)

and microsatellite markers (Maia-Carvalho et

al., 2018), and features peculiar osteological

(Martínez-Solano et al., 2004) and bioacous-

tics characters (Márquez and Bosch, 1995). Pos-

sibly due to ancestral hybridisation or incom-

plete lineage sorting (Gonçalves et al., 2007;

Maia-Carvalho et al., 2014), the nuclear phy-

logeny, based on intron sequences, is not well

resolved. Using microsatellites, Maia-Carvalho

et al. (2018) identiﬁed a distinct cluster cor-

responding to all A. o. almogavarii popula-

tions, suggesting that the lack of monophyly in

mtDNA data is due to cyto-nuclear discordance.

This cluster extends as far west as the southern

slopes of the Pyrenees in the north-westernmost

parts of Aragon. These authors also found re-

stricted genetic admixture between A. o. almo-

gavarii and neighbouring subspecies (A. o. per-

tinax,A. o. obstetricans). Following up on this

study, Dufresnes and Martínez-Solano (2020)

targeted the hybrid zone between A. o. almo-

gavarii and A. o. pertinax with genomic analy-

ses using RADseq-derived markers along a ﬁne-

scale transect in Catalonia. They documented

a very narrow mitochondrial (cline width ca.

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146 J. Speybroeck et al.

13 km) and nuclear (cline width ca. 16 km)

transition, and detected portions of the genome

that were completely impermeable to gene ﬂow.

Given the absence of barriers to dispersal in the

contact zone, the authors concluded that these

lineages exhibit substantial (even if incomplete)

reproductive isolation. We adopt their recom-

mendation and treat Alytes almogavarii as a dis-

tinct species.

Pabijan et al. (2012) found a lack of recipro-

cal monophyly in nuclear data of Iberian painted

frogs (Discoglossus spp.), while Dufresnes et

al. (2020a) found numerous introgressed indi-

viduals in their RAD datasets and a broad hy-

brid zone (average cline width of nuclear mark-

ers >136 km). These results indicate weak (or

no) restriction to gene ﬂow and conﬁrm that

D. g. galganoi and D. g. jeanneae are better

treated as conspeciﬁc, as previously advocated

by SBC2010.

Borkin et al. (2001) reported differences in

genome size between the morphologically sim-

ilar eastern and western populations of Pel o -

bates fuscus, with a transition between both

groups in north-eastern Ukraine and adjacent

parts of Russia. The name Rana vespertina Pal-

las, 1771 is available for the eastern taxon. Sub-

sequently, analyses using mtDNA data across

the entire species range conﬁrmed the existence

of two major mitochondrial lineages (Crottini et

al., 2007). Litvinchuk et al. (2013) conﬁrmed

that these two lineages differ in their nuclear

genomes, with Nei’s genetic distance based on

allozymes being similar to the divergence be-

tween Pelobates syriacus syriacus and P. s. bal-

canicus (see below), and with a restricted intro-

gression zone. Based on its independent evolu-

tionary history, they thus proposed to treat ves-

pertinus as a species. More recently, based on

RADseq data, Dufresnes et al. (2019a) reported

narrow clines between fuscus and vespertinus

(average nuclear cline width: 16 km). On the ba-

sis of the narrow hybrid zone, indicating strong

(albeit incomplete) reproductive isolation, and

in spite of the relatively recent divergence of

this taxon (between 2 and 3 Mya, see Dufresnes

et al., 2019a, b), we accept the speciﬁc rank of

Pelobates vespertinus.

Dufresnes et al. (2019a) also demonstrated

complete reproductive isolation between Pelo-

bates syriacus syriacus and P. s. balcanicus,

warranting elevation of European populations,

except those from south-eastern Bulgaria, parts

of European Turkey and a number of eastern

Greek islands (including Limnos and Lesbos),

to species level as Pelobates balcanicus, with an

estimated Mio-Pliocene divergence (>5 Mya)

between both taxa. Deep intraspeciﬁc diver-

gence within each species further leads to recog-

nise the subspecies P. balcanicus chloeae (Pelo-

ponnese) and P. syriacus boettgeri (all European

parts of the distribution of Pelobates syriacus).

The complex pattern of genetic variation

within Iberian Pelodytes has been known for

several years (e.g., van de Vliet et al., 2012). It

was taxonomically formalised by Díaz-

Rodríguez et al. (2017), who described two

new species: Pelodytes atlanticus from Por-

tugal and P. hespericus from central eastern

Spain, in addition to the previously established

species, P. ibericus and P. punctatus.Thetwo

new lineages show no consistent morphological

or bioacoustics differences, are only weakly dif-

ferentiated in mtDNA (<2% in the 16S rRNA

gene, <7% in COI), but show limited sharing

of nuclear alleles. Díaz-Rodríguez et al. (2017)

recognised that this was a disputable case, in

which, for the time being, “species status for

all four western lineages of Pelodytes [was] the

hypothesis best ﬁtting the available data”. A

new study by Dufresnes et al. (2020a), based on

RADseq-derived loci from spatially dense sam-

pling, found that P. atlanticus and P. ibericus

are separated by a narrow hybrid zone, featur-

ing little introgression, whereas the contact zone

of P. hespericus and P. punctatus has a consid-

erably more ﬂattened cline, with an introgres-

sion zone possibly extending up to 200-300 km.

There is no known contact zone between P. at -

lanticus and P. hespericus. While the position

of P. atlanticus (sister to ibericus or to hesperi-

cus) remains unresolved, its genetic divergence

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Species list of European herpetofauna 147

from hespericus and punctatus is quite large.

We thus follow the suggestion of Dufresnes

et al. (2020a) to accept P. atlanticus as sepa-

rate species, while retaining hespericus at sub-

species level as P. punctatus hespericus.

Despite growing support for recognition of

various clades in the family Bufonidae as sepa-

rate genera (Stöck et al., 2006; Van Bocxlaer et

al., 2009), several authors advocated retaining

Bufo for all species from the Western Palaearc-

tic and Central Asia, at least as an ad interim

solution (Dubois and Bour, 2010; SBC2010).

Formal taxonomic action was further hampered

by nomenclatural confusion, such as the use

of either Pseudepidalea or Bufotes as the valid

generic nomen for the green toad group (Frost

et al., 2006; Dubois and Bour, 2010). Dubois

and Bour (2010) demonstrated that Pseudep-

idalea is a junior objective synonym of Bu-

fotes, thereby giving priority to the latter. How-

ever, based on the presence of hybridisation

between representatives of these clades, and

the restricted sampling size of previous stud-

ies, Dubois and Bour (2010) listed these nom-

ina as subgenera, rather than genera. Accord-

ing to Van Bocxlaer et al. (2010), Pyron and

Wiens (2011), and Beukema et al. (2013), which

together provide a dense sampling of bufonid

species, most genera currently recognised by

Frost (2019) represent well-supported mono-

phyletic units. Moreover, time-calibrated analy-

ses showed Bufo,Bufotes and Epidalea to be of

similar age as or older than most other recog-

nised bufonid genera (Beukema et al., 2013).

Considering these results, we accept Bufo,Bu-

fotes and Epidalea at genus level.

Recuero et al. (2012) published a multilocus

mitochondrial and nuclear DNA sequence data

set for Bufo bufo and associated species, cover-

ing the entire documented range and providing

extensive genetic data. The study yields a fully

resolved phylogeny, with the recently described

Bufo eichwaldi from the Talysh Mountains of

southern Azerbaijan and northern Iran as the

sister taxon of a clade including three deeply

diverged lineages: i) north African, Iberian,

and most French populations of Bufo bufo (for

which the name Bufo spinosus is available),

and a clade composed of two lineages, repre-

senting ii) verrucosissimus from the Caucasus

and iii) bufo from northern France to Russia,

while populations from Greece, southern Italy

and Sicily and most of Anatolia carried bufo

mtDNA but grouped with verrucosissimus in

nuclear DNA. Estimations of divergence times

indicated a long evolutionary history of the

group, starting with the split from eichwaldi at

about 12 Mya, and the divergence of spinosus

taking place around 6 Mya. The deep level of

genetic divergence observed between the west-

ern and eastern groups of common toads indi-

cated that these groups may be different species.

However, Garcia-Porta et al. (2012) found ex-

tensive admixture of mitochondrial lineages be-

tween the eastern and western clades in the

Languedoc area of southern France (as also re-

ported by Arntzen et al., 2017 for the Provence

area of southern France), and detected signs of

ancient introgression of bufo allozyme alleles

into spinosus. They thus suggested to treat B. b.

spinosus and B. b. bufo as conspeciﬁc pending

studies of the contact zones. Detailed analysis of

the amount of reproductive isolation in two geo-

graphically distant contact zones on the basis of

mtDNA, morphology and nuclear markers have

subsequently been published by Arntzen et al.

(2016, 2017). Both studies documented an ex-

tensive amount of hybridisation and introgres-

sion in the contact zone, but also the presence

of narrow and concordant clines for most nu-

clear markers, resulting in a unimodal, yet nar-

row hybrid zone of ca. 30 km wide, and indi-

cating intrinsic barriers to gene ﬂow in spite of

incomplete reproductive isolation. Karyological

analysis (albeit based on only four males and

a single female) identiﬁed heteromorphic sex

chromosomes in spinosus, but not in bufo (Sko-

rinov et al., 2018). The combined available ev-

idence justiﬁes treating Bufo spinosus as a dis-

tinct species.

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148 J. Speybroeck et al.

Based on differences in morphology and

karyotype and a study suggesting lowered fertil-

ity of F1 hybrids (summary in Kuzmin, 1999),

the Caucasian populations of the complex are

sometimes also treated as a distinct species as

Bufo verrucosissimus. Recent genetic studies

paint a more complex picture. Firstly, based on

nuclear markers, the Anatolian populations, tra-

ditionally excluded from B. b. verrucosissimus

on the basis of morphology, are in fact geneti-

cally closer to this group than to B. b. bufo or B.

spinosus, even if they carry B. b. bufo mtDNA

(Garcia-Porta et al., 2012; Arntzen et al., 2013).

Secondly, mtDNA divergence between Cau-

casian (B. b. verrucosissimus) and Eastern Eu-

ropean (B. b. bufo) populations is much lower

than between B. b. bufo and B. spinosus.In-

deed, divergence in mtDNA and allozymes be-

tween the Caucasian and Eastern European lin-

eages is even lower than between the subclades

of B. spinosus (Garcia-Porta et al., 2012). Most

importantly, based on frequencies of allozyme

alleles, populations of B. b. bufo from Greece

and north-western Turkey appear intermediate

between Caucasian and European populations

(Garcia-Porta et al., 2012) or group with Cau-

casian populations in nuclear trees (Recuero et

al., 2012), suggesting extensive introgression

between B. b. verrucosissimus and B. b. bufo in

north-western Turkey and the southern Balkans

(but see Arntzen et al., 2013). In our opinion,

the speciﬁc status of B. b. verrucosissimus is

currently insufﬁciently supported by the avail-

able evidence, and we maintain it as conspeciﬁc

with Bufo bufo for the time being.

Stöck et al. (2006) elevated a number of mito-

chondrial lineages of green toads (now Bufotes)

to species level (for Europe: B. variabilis and

B. balearicus, in addition to B. viridis) and sub-

sequently described a new species from Sicily

based on the same lines of evidence. SBC2010

argued against accepting these species on the

basis of mitochondrial DNA alone. Colliard et

al. (2010) documented a contact zone in Sicily

between the taxa siculus and balearicus (sensu

Stöck et al., 2006) and demonstrated strong re-

productive isolation between them. As these

two taxa belong to the two most divergent lin-

eages (North African and Eurasian), a two-

species split of the green toad complex is clearly

warranted. The Sicilian taxon siculus is closely

related to the North African taxon boulengeri,

which has priority over siculus. Subsequently,

nuclear data were added to the picture (mi-

crosatellites – Dufresnes et al., 2018a; Gerchen,

Dufresnes and Stöck, 2018, and RADseq mark-

ers – Dufresnes et al., 2019c). Although cyto-

nuclear discordance was present in large areas

of Europe where populations of viridis carry

the variabilis mtDNA lineage, these studies

conﬁrmed that overall the previously identiﬁed

mtDNA lineages correspond to distinct evolu-

tionary units. Dufresnes et al. (2019c) also dis-

covered that the type locality of variabilis is in-

habited by the western lineage, rendering vari-

abilis a junior synonym of viridis, and that the

valid name for the Anatolian lineage is sitibun-

dus. These two lineages (B. v. sitibundus and B.

v. viridis) widely admix over a very large geo-

graphic area in Anatolia and Russia (Dufresnes

et al., 2019c), suggesting that they are best

treated as subspecies. The contact zone between

B. v. viridis and B. v. balearicus is narrower

(63 km), but still extensive if compared with

other contact zones between taxa that are here

treated as different species (Gerchen, Dufresnes

and Stöck, 2018). In addition, raising B. v.

balearicus but not B. v. sitibundus to species

rank would make B. v. viridis paraphyletic (see

Dufresnes et al., 2019c). The TC therefore pre-

ferred to treat B. v. sitibundus and B. v. baleari-

cus (and the extralimital B. v. perrini) as sub-

species of B. viridis.

Dufresnes et al.’s (2019c) range-wide sam-

pling of all known taxa of the Bufotes viridis

complex identiﬁed a total of eight diploid ge-

netic clusters, including a new lineage endemic

to Cyprus, which they describe as Bufotes cy-

priensis. Estimated to be of Messinian origin

(5.3 Mya), this insular taxon is as old as or

older than many species of anuran amphibians

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Species list of European herpetofauna 149

from the Western Palearctic. Yet, all ten indi-

viduals from four localities carry mtDNA hap-

lotypes of B. v. sitibundus. Secondary contact

and hybridisation with the Anatolian mainland

lineage during the Pleistocene would explain

this mitochondrial capture, as well as the rem-

nants of nuclear introgression that were detected

as well. In addition to this deep genomic di-

vergence, the genome of B. cypriensis is also

signiﬁcantly larger than that of all other West-

ern Palearctic Bufotes taxa. Morphologically,

the Cyprus green toads are smaller than those

from mainland populations, as previously re-

ported by Stugren and Tassoula (1987). The

combination of an old divergence and a larger

genome size led the TC to accept the species sta-

tus of B. cypriensis. Adriatic populations could

represent a valid subspecies of B. viridis (for

which the nomen longipes Fitzinger in Bona-

parte, 1840 might be available), while green

toads from Naxos (Cyclades) and Crete dif-

fer genetically from other populations and de-

serve genomic investigations (Dufresnes et al.,

2019c). In summary, for Europe, we accept (1)

Bufotes viridis, including the widespread B. v.

viridis,B. v. balearicus (Italian Peninsula, Cor-

sica, Sardinia, Sicily, Balearic islands) and B. v.

sitibundus (within Europe restricted to a num-

ber of eastern Greek islands), (2) the Sicilian

Bufotes boulengeri siculus and (3) Bufotes cy-

priensis. We note that the availability of the

nomen siculus remains unclear, given that the

corresponding description (Stöck et al., 2008)

was only issued electronically and prior to 2011

(see Article 8.5 of the Code): whether “numer-

ous identical and durable copies” (Article 8.1)

were registered by the authors in parallel is yet

to be addressed (see also Dubois et al., 2013).

Based on a high level of divergence in mi-

tochondrial DNA sequences of Hyla tree frogs,

Stöck et al. (2008) have suggested to recog-

nise the Iberian taxon molleri and the eastern

European taxon orientalis as species distinct

from Hyla arborea. However, given the lack

of other supporting characters (weak acous-

tic divergence: see Schneider, 1974; no well-

supported morphological characters), SBC2010

refrained from formally accepting this syste-

matic treatment, as it would have rested solely

on mitochondrial DNA data. Several more re-

cent studies offer a better understanding of

the phylogeny of this group and of the pat-

terns of gene ﬂow across contact zones. Firstly,

Dufresnes et al. (2015, 2016) investigated the

arborea-orientalis contact zones in Poland and

in the Balkans with microsatellite markers. In

Poland, they found evidence of admixture over a

200 km wide zone, with mosaic contacts and in-

terspersed hybrid populations, but with strongly

restricted introgression at sex-linked loci and

many populations of seemingly pure ancestry

in close contact to each other (Dufresnes et al.,

2016). In the Balkans, Dufresnes et al. (2015)

found narrow clines (30 and 32 km of aver-

age cline width in Serbia and Greece respec-

tively). In line with our treatment of taxa con-

nected by unimodal hybrid zones with narrow

clines, we recommend affording species status

to Hyla orientalis. Secondly, the phylogenomic

tree in Dufresnes et al. (2018b) unambiguously

groups orientalis and molleri as sister taxa, with

arborea splitting from a node basal to the mol-

leri –orientalis divergence. Therefore, we ac-

cept molleri at species rank as well. Although

the close relationships between molleri and ori-

entalis could justify to treat them as conspeciﬁc,

we refrain for the time being from this arrange-

ment. We thus treat Hyla molleri and Hyla ori-

entalis as valid species.

Supported by mtDNA and genomic data, two

major cryptic lineages reside within Hyla inter-

media (Canestrelli, Verardi and Nascetti, 2007;

Stöck et al., 2008; Dufresnes et al., 2018b).

They are not known to differ in any diagnos-

tic morphological or acoustic character, even

if they differ in the averages of one acoustic

and some morphometric traits. The northern lin-

eage occupies the Po Plain and adjacent re-

gions (including Ticino in Switzerland), with

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150 J. Speybroeck et al.

the northern Apennines acting as the biogeo-

graphical barrier separating it from the south-

ern lineage. As the name intermedia applies to

the southern lineage, Dufresnes et al. (2018b,

c) coined a new name, Hyla perrini,forthe

northern lineage. Note that, as its registration

in the Ofﬁcial Register of Zoological Nomen-

clature (ZooBank) occurred later than the pub-

lication of Dufresnes et al. (2018b), the name

perrini is not made available by Dufresnes et al.

(2018b), but by Dufresnes et al. (2018c), who

met all conditions of nomenclatural availabil-

ity. Detailed analyses of the contact zone based

on genomic (RADseq) data by Dufresnes et al.

(2018b) revealed broad clines (96 km of average

cline width) for nuclear markers and detectable

admixture over approximately 130 km. Despite

a relatively high cyt bdistance of around 9%,

and even though Dufresnes et al. (2018b) ar-

gued that the extent of the contact zone would

be even larger without some form of selection

against hybrids, the TC felt that the observed ex-

tent of introgression and thus the lack of strong

reproductive barriers did not unambiguously al-

low treatment of perrini at species rank. We thus

recommend to treat the northern lineage as Hyla

intermedia perrini.

The authorship of the family Ranidae has

been clariﬁed by Dubois and Bour (2011), at-

tributing it to Batsch, 1796.

Based on allozyme data (Arano, Esteban and

Herrero, 1993; Veith et al., 2002, 2012), two ge-

netically distinct groups have been recognised

within Rana temporaria populations of northern

Spain. The ﬁrst was assigned to the subspecies

R. t. parvipalmata and is restricted to the west-

ernmost edges of the Iberian distribution (Gali-

cia and Asturias). Frogs from western Galicia

feature reduced feet webbing, smaller size and

lower number of pulses per call (Vences, 1992).

The second group corresponds to the nominal

taxon R. t. temporaria, which extends through-

out Europe. Mitochondrial data suggested a

more complex picture, with four deeply differ-

entiated lineages in the same area (Vences et al.,

2013b, 2017). In a phylogeographic survey us-

ing genome-wide data, Dufresnes et al. (2020b)

found one of these mitochondrial groups to be a

“ghost lineage”, not differentiated in the nuclear

genome. The two main lineages are strongly dif-

ferentiated and are estimated to have diverged

around 4 Mya. Despite the absence of geo-

graphic or ecological barriers to dispersal, they

form a narrow hybrid zone (25 km) in the east-

ern Cantabrian Mountains. Because partial re-

productive isolation is thus likely to prevent

these two taxa from merging, we follow the

recommendation of Dufresnes et al. (2020b) to

treat R. parvipalmata as a distinct species.

Using allozymes and mtDNA, a detailed

analysis of gene ﬂow patterns across the con-

tact zone of the northern Anatolian lineage and

the Balkan lineage of the Pelophylax ridibun-

dus –bedriagae complex in northern Greece re-

vealed the existence of a wide hybrid zone, with

introgression detectable over more than 200 km

(Hotz et al., 2013). This suggests that the Eu-

ropean and Anatolian lineages are conspeciﬁc

(contra SBC2010). Since the European lineage

splits from a node basal to the diversiﬁcation of

Anatolian lineages with respect to mtDNA ac-

cording to Plötner et al. (2012), it further sug-

gests that most lineages of this complex are

conspeciﬁc, with the possible exception of the

populations from Syria and Jordan (possibly the

‘true’ bedriagae, see Plötner et al., 2012). On

the other hand, Plötner et al. (2010) suggested

hybrid breakdown between some of these lin-

eages, indicating incipient speciation. While un-

doubtedly the last word on this subject has not

been written, the TC felt that it was prema-

ture to suggest taxonomic changes, especially

as a large taxonomic paper based on genomic

variation patterns is in preparation (G. Mazepa,

pers. comm.). The same applies to the newly

proposed P. cypriensis, which was found to be

placed between the European and the Anato-

lian lineages in mtDNA, but grouped with P.

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Species list of European herpetofauna 151

cretensis (albeit with very low support) in nu-

clear DNA (Plötner et al., 2012). For the mo-

ment, we thus suggest no change to the taxon-

omy adopted in SBC2010 for European Pelo -

phylax.

Reptilia

Testudines/Chelonii

Based on nuclear genomic ISSR ﬁngerprints

and mtDNA sequences, Fritz et al. (2005) de-

scribed Emys trinacris as a distinct species en-

demic to Sicily and Calabria, albeit the record

from Calabria was later questioned (Vamberger

et al., 2015). A phylogenetic study using seven

nuclear genes (Spinks and Shaffer, 2009) found

E. orbicularis to be paraphyletic with respect to

trinacris:E. o. hellenica was sister to a clade

containing trinacris and the remaining lineages

of E. orbicularis. This led SBC2010 to conclude

that trinacris should not be recognised as a

species, which has been criticised by Vamberger

and Fritz (2018). While trinacris is the most

basal lineage in mtDNA trees (Fritz et al., 2005,

2007), this position is weakly supported, and the

level of mtDNA divergence between trinacris

and the other lineages is not clearly larger than

that between some lineages of E. orbicularis.

A ﬁrst combined analysis of genetic differen-

tiation at eight microsatellite loci and mtDNA

sequences found concordant patterns for both

markers: no evidence for admixture between

Sicilian pond terrapins and the remaining lin-

eages was found, while extensive admixture was

found between some other lineages of E. orbic-

ularis (Pedall et al., 2011). Using 15 microsatel-

lite loci and mtDNA sequences, Vamberger et

al. (2015) found evidence for limited admixture

between the two taxa in one Sicilian population.

In addition, Vamberger et al. (2015) reported

concordant clines for both markers, with cline

centres matching the Strait of Messina. Vam-

berger et al. (2015) argue that the coincidence

of the cline centres with the Strait of Messina

cannot be explained by a geographic barrier ef-

fect of this narrow sea strait, because in E. or-

bicularis recent gene ﬂow occurred across the

Strait of Gibraltar (Velo-Antón et al., 2015),

and possibly across the Adriatic Sea between

southern Italy and the Balkans (Vamberger et

al., 2015). In contrast, Pereira, Teixeira and

Velo-Antón (2018), using seven microsatellite

loci, concluded that the Strait of Gibraltar cur-

rently impedes gene ﬂow between Iberian and

North African pond turtles. However, they show

that the break between the two clusters corre-

sponds to the Central Iberian Mountains and

not the sea strait. Clines of microsatellites for

the trinacris –orbicularis contact zone across

theStraitofMessinawereinferredtobevery

wide (247 km) in Vamberger et al. (2015), al-

though the signiﬁcance of this estimate is ham-

pered by a substantial (150 km) sampling gap

between the two taxa. The TC takes therefore

a conservative stance and treats trinacris as a

subspecies of E. orbicularis, waiting for further

studies resolving the complicated relationships

of this complex. In this context, we note that

Pöschel et al. (2018) reported a sharp transition

between occidentalis and orbicularis +gal-

loitalica in north-eastern Spain, matching that

of distinct species recognised here, suggesting

that the E. orbicularis complex could comprise

several species. In the meantime, we recom-

mend to maintain the Sicilian lineage as Emys

orbicularis trinacris.

Squamata

The family Agamidae is sometimes credited to

Fitzinger, 1826 or Gray, 1827 (see e.g. Melville

and Smith, 1987). In fact the nomen was ﬁrst

published by Spix in 1825 as “Familia Aga-

mae”. Even if Spix did not use the correct suf-

ﬁx ‘-idae’, Article 11.7.1.3 of the Code states

that “a family-group name of which the family-

group name sufﬁx is incorrect is available with

its original authorship and date, but with a cor-

rected sufﬁx”. Other conditions of availability

all seem to be fulﬁlled in Spix (1825), who is

thus the author of the nomen.

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152 J. Speybroeck et al.

Macey et al. (2000a) sequenced a 1685-1778

bp long segment of mtDNA (including the ND1,

ND2 and COI genes) to assess phylogenetic

relationships of acrodont lizards. Their results

suggested that the agamid genus Laudakia is

paraphyletic, yet, low bootstrap support pre-

vented deﬁnite conclusions. Subsequent stud-

ies based on mitochondrial and nuclear genes

(ND2, RAG1) by Melville et al. (2009) and

Edwards and Melville (2011) recovered Lau-

dakia as monophyletic with high support. Baig

et al. (2012) summarised the results of the afore-

mentioned studies in a morphology-based revi-

sion of Laudakia. Despite failing to ﬁnd distinct

morphological variation within the genus, and

acknowledging that Melville et al. (2009) and

Edwards and Melville (2011) recovered Lau-

dakia as monophyletic, Baig et al. (2012) par-

titioned Laudakia into three genera acknowl-

edging its potential paraphyly (Macey et al.,

2000a). This taxonomic act was subsequently

criticised by Pyron, Burbrink and Wiens (2013),

who conﬁrmed the monophyly of Laudakia us-

ing a supermatrix approach. Within our focal

area, the classiﬁcation proposed by Baig et al.

(2012) would affect Laudakia stellio and L. cau-

casia, as these authors placed the former species

in the newly erected genus Stellagama, and the

latter into Paralaudakia. While these genera

were rapidly adopted by the wider herpetolog-

ical community, we do not follow the split of

Laudakia, pending substantial evidence to reject

its monophyly, and therefore retain L. stellio and

L. caucasia in Laudakia.

The family Chamaeleonidae is sometimes

credited to Gray, 1825 (e.g. Glaw, 2015). How-

ever, Raﬁnesque (1815) published the name

“Famille CAMÆLONIA”, based on the genus

“Camæleo Daud.”, which is an incorrect subse-

quent spelling of the available genus Chamaeleo

Laurenti, 1768. The Code is somewhat am-

biguous regarding the availability of family-

group nomina based on an incorrect subsequent

spelling of an available genus. Article 19.1 of

the Code states that an incorrect subsequent

spelling is not an available name. If one fol-

lows this and Art. 11.7.1.1 (“a family-group

name when ﬁrst published must [...] be a noun

in the nominative plural formed from the stem

of an available generic name”), a family-group

name based on an incorrect subsequent spelling

of an available genus name is not available.

Yet, the Code also states that “a family-group

name is an incorrect original spelling and must

be corrected if it is formed from an incorrect

subsequent spelling of a generic name” (Art.

32.5.3.3) and that “a family-group name based

upon [...] an incorrect spelling of the name of

the type genus must be corrected” (Art. 35.4.1).

It is thus clear that the Code has no inten-

tion to make a family-group name unavailable

based on an incorrect spelling of the name of

the type genus (see also Dubois, 2010). In con-

clusion, the family-group name Camaelonia is

available as published by Raﬁnesque (1815), but

its spelling needs to be corrected. The nowadays

prevailing spelling Chamaeleonidae should be

preserved (Art. 29.5), with its authorship at-

tributed to Raﬁnesque (1815) instead of Gray

(1825).

SBC2010 adopted the inclusion of Cyrtopo-

dion kotschyi in the well-supported mono-

phyletic genus Mediodactylus, as proposed by

Macey et al. (2000b), ˇ

Cervenka, Frynta and

Kratochvíl (2008) and Bauer et al. (2013).

By enlarging the focal area of SBC2010, we

hereby also include the former C. russowi,

now Mediodactylus russowii. Another species

occurring in the area considered by Gasc

et al. (1997) but not in that of SBC2010

is the former Cyrtopodion caspium.Wefol-

low Bauer et al. (2013) in recognising the

well-resolved genus Tenuidactylus and list this

species as Tenuidactylus caspius. Furthermore,

an introduced, well-established population of

Tenuidactylus fedtschenkoi has recently been re-

ported from the city of Odessa (Ukraine). The

native range of T. fedtschenkoi lies in central

Asia (western Pamiro-Altay mountains), and it

was probably transported passively to Odessa

(Duz’, Kukushkin and Nazarov, 2012). Not long

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Species list of European herpetofauna 153

after the record from Odessa, a revision of

the genus Tenuidactylus led to the description

of a new species from Uzbekistan and Turk-

menistan, which was split from T. fedtschenkoi

and named T. bogdanovi (Nazarov and Po-

yarkov, 2013). Krasylenko and Kukushkin

(2017) provided an update on the status of the

non-native Odessa population and assigned it to

T. bogdanovi.

Within Mediodactylus kotschyi, many sub-

species have traditionally been recognised. As

indicated by Kasapidis et al. (2005), the ge-

netic substructure of Mediodactylus kotschyi

shows a high degree of divergence, suggest-

ing that M. kotschyi represents a species com-

plex. Nearly range-wide data (with limited ar-

eas in eastern and northern Anatolia excluded)

of three mtDNA and three nuclear DNA frag-

ments allowed unravelling the evolutionary his-

tory of Mediodactylus kotschyi (Kotsakiozi et

al., 2018). Divergence dates back to 15 Mya,

and several of the main lineages show overlap-

ping distribution areas. Divergence in mtDNA

is known to be often quite large in geckos (e.g.

Nagy et al., 2012). However, distances between

the main lineages are particularly large in this

case (>10% 16S and >15% cyt band COI

p-distances). In addition, nuclear data groups

specimens in concordance with their mtDNA

lineage, and not with their geographical ori-

gin, which suggests a high level of repro-

ductive isolation between them. The proposed

splits also largely agree with morphological

data (Štˇ

epánek, 1937, 1939; Szczerbak and Gol-

ubev, 1996). The narrow contact between M.

orientalis and M. danilewskii in the Western

Taurus Mountains, identiﬁed by Kotsakiozi et

al. (2018), corresponds to a lack of morpho-

logical intergrades (despite the narrow gap be-

tween the subspecies danilewskii and cilicien-

sis in the same area, see Rösler, Schmidtler and

Moravec, 2012), while these new species are

very close to the M. kotschyi groups deﬁned by

Beutler (1981). Five species were recognised,

for all of which previously existing names are

available (Kotsakiozi et al., 2018). While the

delimitation of the ranges and contact zones of

these new species requires further investigation

(e.g. M. danilewskii and M. orientalis), we ac-

cept these species here. The following species

are thus recognised in our area: M. kotschyi

(mainland Balkans, most of Aegean islands, and

Italy (Apulia)), M. bartoni (Crete and nearby

islets), M. danilewskii (Black Sea region and

south-western Anatolia), M. oertzeni (southern

Dodecanese Islands), and M. orientalis (Levant,

Cyprus, southern Anatolia, and south-eastern

Aegean islands).

The endemic gecko from the Selvagens Is-

lands has variously been treated as the valid

species Tarentola bischofﬁ (e.g. Rebelo, 2008;

Sindaco and Jeremˇ

cenko, 2008; Uetz, Freed and

Hošek, 2019), or as a subspecies of the Ca-

narian T. boettgeri as T. boettgeri bischofﬁ (e.g.

Carranza et al., 2000; Pleguezuelos, Márquez

and Lizana, 2002; Gübitz, Thorpe and Malho-

tra, 2005; Sá-Sousa et al., 2009). Its phyloge-

netic afﬁnities were examined by Carranza et

al. (2000, 2002) and Gübitz, Thorpe and Mal-

hotra (2005). The genetic data available (al-

beit based only on mtDNA) show that the mi-

tochondrial lineages of bischofﬁ are closely re-

lated to those of T. boettgeri, especially to the

subspecies T. b. hierrensis from El Hierro, and

are in fact nested inside the mitochondrial di-

versity of T. b. boettgeri from Gran Canaria,

suggesting that the Selvagens Islands were re-

cently colonised from El Hierro or Gran Canaria

(Carranza et al., 2000, 2002; Gübitz, Thorpe

and Malhotra, 2005). Treating bischofﬁ as a

valid species, while retaining hierrensis as a

subspecies of boettgeri (as done by the above

authors who rank bischofﬁ as a species), is

thus likely to render Tarentola boettgeri para-

phyletic. The amount of mitochondrial diver-

gence between hierrensis and bischofﬁ is also

smaller than within the populations of T. de-

lalandii from Tenerife or between the popula-

tions of T. angustimentalis from Fuerteventura

and Lanzarote. Based on the available data, we

thus recommend to treat the geckos from the

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154 J. Speybroeck et al.

Selvagens Islands as conspeciﬁc with the popu-

lations from El Hierro and Gran Canaria, as Tar-

entola boettgeri bischofﬁ.

Sánchez-Vialas et al. (2018) re-examined the

description of Algyroides hidalgoi Boscá, 1916,

a nomen for which the holotype has been lost,

to settle its position in the synonymy of the

genus Algyroides. They argued that the charac-

ters of the holotype, said to originate from the

Sierra de Guadarrama and described by Boscá,

fall within the morphological variability of Al-

gyroides marchi Valverde, 1958, and designated

a specimen of A. marchi as neotype of A. hi-

dalgoi. This would make A. marchi a junior

subjective synonym of A. hidalgoi. Because the

conditions for automated reversal of precedence

are not met (Art. 23.9.1 of the Code), A. hi-

dalgoi would become the valid nomen of the

Spanish Algyroides. Several TC members felt,

however, that the interpretation of the descrip-

tion of Boscá (1916) by Sánchez-Vialas et al.

(2018) left room for doubt, and that Boscá may

not have described a specimen of Spanish Algy-

roides when he created the name Algyroides hi-

dalgoi. If so, this could affect the validity of the

neotype designation, respective to Art. 75.3.5 of

the Code, and result in the unnecessary change

of the well-established and widely used name A.

marchi. Although opinions in the TC were di-

vided about this, we recommend to maintain for

the time being the use of A. marchi, in anticipa-

tion of an upcoming application to the Commis-

sion, which would maintain marchi in use until

the Commission has ruled on the case.

Morphological and molecular (mtDNA and

nuclear) data support the split of Psammod-

romus hispanicus into three distinct lineages

(Fitze et al., 2011, 2012; Mendes et al., 2017).

While not all areas of potential contact have

been sampled (despite a recent distribution up-

date by Molina et al., in press), the available

data seem to warrant accepting these lineages

as distinct species. Age estimates, lineage al-

lopatry, the lack of mitochondrial and nuclear

haplotype sharing between lineages, bioclimatic

niche divergence, and the current biogeographic

distribution, indicate that the three lineages cor-

respond to three independent species. The name

of the eastern species is often spelled P. ed-

wardsianus. As shown by Crochet (2015), this

is an incorrect subsequent emendation that does

not meet the requirements of the Code. As a

consequence, the valid spelling of the eastern

species is Psammodromus edwarsianus. Thus,

we add Psammodromus edwarsianus and P. oc-

cidentalis to the list of the European herpeto-

fauna species.

A 9 Mya split marks the divergence of Timon

lepidus nevadensis from the nominal subspecies

T. l. lepidus (Miraldo et al., 2013). After studies

on genetic (Paulo et al., 2008; Miraldo et al.,

2011, 2013) and morphological differentiation

(Mateo and Castroviejo, 1990; Mateo, López-

Jurado and Guillaume, 1996), mtDNA and mi-

crosatellites were used to investigate gene ﬂow

patterns in a zone of secondary contact (Miraldo

et al., 2013). While hybridisation and introgres-

sion were observed, gene ﬂow was shown to be

restricted. The cline width for nuclear markers

was estimated at around 10 km (although with

a sampling gap of around 20 km, it may actu-

ally be less). Furthermore, mostly pure popu-

lations are present on either side of the sam-

pling gap. Considering this together with the

aforementioned old genetic divergence, we ac-

cept the proposal of Miraldo et al. (2013) and

treat T. nevadensis as a valid species.

Lacking range-wide sampling and adequate

molecular analysis, the taxonomy of the Lac-

erta trilineata-pamphylica complex has re-

mained unresolved, until mitochondrial phylo-

genies showed the eastern Anatolian species L.

pamphylica to be nested within trilineata (God-

inho et al., 2005; Ahmadzadeh et al., 2013;

Sagonas et al., 2014). Remarkably, Sagonas et

al. (2014) found evidence that central Aegean

populations (L. t. citrovittata) are closely related

to L. pamphylica. Yet, this was poorly supported

across analyses and the biogeographically sur-

prising relationship between L. t. citrovittata

and L. pamphylica could reﬂect a methodologi-

cal artefact (long branch attraction). Thus, the

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Species list of European herpetofauna 155

monophyly of L. trilineata could not be re-

jected. More recently, analyses of SNPs and mi-

tochondrial sequences by Kornilios et al. (2019,

2020) found a sister-group relationship between

pamphylica and eastern Aegean populations of

trilineata. This led to the identiﬁcation of four

species-level units: L. trilineata (eastern Adri-

atic Coast, Greece except northeast and includ-

ing Milos archipelago and Crete, south-eastern

Bulgaria and North Macedonia), L. pamphylica

(southern Anatolia east of Antalya), L. citrovit-

tata (central Aegean islands, including among

others Naxos, Tinos, Andros, Syros, Mykonos,

Paros), and L. diplochondrodes (north-eastern

Greece, Romania, Bulgaria except southeast,

Turkey, eastern Aegean Greek islands). All ev-

idence (morphology, blood biochemistry, im-

munoserology, congruence between highly di-

vergent mtDNA and nuclear genomics, biocli-

matic niche differences, no signs of admixture)

suggests that L. pamphylica deserves species

status. As the eastern Aegean populations of the

complex are the sister taxon of L. pamphylica,

maintaining monophyly implies also treating

them as a species, L. diplochondrodes. The lat-

ter taxon does not show a lot of contact, nor

signs of introgression with any of the other three

clades. Concerning the population west of the

Aegean Barrier, the conclusion may seem less

straightforward. However, genomic analyses of-

fer stronger support for the species status of L.

citrovittata than for that of L. pamphylica.The

former equally corresponds to one of the ma-

jor mtDNA lineages of Kornilios et al. (2019).

Given its allopatry, there is no evidence of ad-

mixture. While the (potential) contact zone be-

tween L. diplochondrodes and L. trilineata in

north-eastern Greece and its wider surroundings

requires more comprehensive sampling, we sug-

gest to accept the new four species arrangement

and to recognise Lacerta trilineata,L. diplo-

chondrodes,L. citrovittata,aswellastheex-

tralimital L. pamphylica.

Using one mitochondrial and one nuclear

gene, Marzahn et al. (2016) investigated the

phylogeography of the Lacerta viridis com-

plex. Four main mtDNA lineages, whose phy-

logenetic relationships were weakly resolved,

emerged from the data: (i) Lacerta bilineata, (ii)

Lacerta viridis s.s., (iii) an Adriatic and west-

ern Balkan lineage, and (iv) a newly discovered

lineage from the south-eastern Balkans as well

as the Turkish Black Sea Coast. A previously

supposed (Amann et al., 1997) contact zone be-

tween L. bilineata and L. viridis in north-eastern

Italy was found to be in fact a contact zone be-

tween the Adriatic and western Balkan lineage

and L. bilineata. Another contact zone between

the Adriatic and western Balkan lineage and L.

viridis s.s. lies further east in Slovenia and ad-

jacent Croatia. As a consequence, there is no

known geographical contact zone between L.

viridis s.s. and L. bilineata. Several other con-

tact zones between mitochondrial lineages were

identiﬁed across the distribution range. How-

ever, the nuclear data were insufﬁcient to exam-

ine gene ﬂow among lineages. Consequently, no

species-level changes were suggested. The pre-

viously reported occurrence of L. bilineata on

Cres, Croatia (Brückner et al., 2001), was con-

ﬁrmed. However, this record is isolated from

the remaining distribution range and lies among

surrounding records of the Adriatic and western

Balkan lineage (Marzahn et al., 2016), sugges-

tive of an introduced population.

Speybroeck and Crochet (2007) and

SBC2010 adopted the split of the former genus

Lacerta s.l., accepting all genus names recog-

nised by Arnold, Arribas and Carranza (2007),

as far as they pertained to taxa occurring within

their considered area. As we have adopted a

broader area deﬁnition here, we additionally

formally accept the genera Anatololacerta and

Phoenicolacerta. The latter only has a single

representative within our area, the Cypriot P.

troodica. Within our area, the genus Anatololac-

erta can be found on several Greek islands, in-

cluding Ikaria, Samos, Rhodes, Pentanisos and

Kastellorizo. Based on morphology, Eiselt and

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156 J. Speybroeck et al.

Schmidtler (1986) treated A. anatolica (includ-

ing the subspecies aegaea from Samos), A. dan-

fordi and A. oertzeni (including the subspecies

A. o. pelasgiana on Rhodes, Symi and Pen-

tanisos and the nominotypical subspecies on

Ikaria) as valid species. However, because the

biochemical differentiation (based on albumins)

between them was deemed to be too small,

Sindaco and Jermenˇ

cenko (2008) maintained

all populations within Anatololacerta danfordi,

following Mayer and Lutz (1989). Bellati et al.

(2015) combined multi-locus species tree ap-

proaches with species delimitation methods to

suggest the recognition of four distinct species

in this group. Some of these species are closely

related and were inferred to have diverged as

recently as 500,000 years ago, i.e. a remark-

ably young age for speciation events within

Lacertidae. Yet, there seems to be evidence of

reproductive isolation between some of these

species, as shown by the patterns of allele shar-

ing in some nuclear markers. While more de-

tailed analyses of patterns of gene ﬂow in con-

tact zones would be desirable, we tentatively ac-

cept the four-species taxonomy (A. anatolica,A.

budaki,A. danfordi and A. pelasgiana)ofBel-

lati et al. (2015). In our area, A. anatolica is

present on Ikaria and Samos, and A. pelasgiana

on Symi, Pentanisos and Rhodes. In addition, A.

pelasgiana was also found (although presumed

introduced in both cases) on Kasos (Kornilios

and Thanou, 2016) and Kastellorizo (Kalaentzis

et al., 2018), while A. budaki was found on the

(nearby) islet Psomi (Kalaentzis et al., 2018).

As pointed out by Busack et al. (2016),

the microﬁche publication of Arribas’s (1997)

thesis does not meet the criteria of publica-

tion for valid zoological nomenclature, nei-

ther under the current version of the Code

nor under the previous (and then relevant)

version. As a consequence, the names Ibero-

lacerta and Darevskia were not made avail-

able in Arribas (1997). For Iberolacerta,this

has no major nomenclatural consequences, as

the next available nomen is Iberolacerta Ar-

ribas, 1999. However, because Darevskia Ar-

ribas, 1999 is a junior synonym of Caucasi-

lacerta Harris, Arnold and Thomas, 1998, Bu-

sack et al. (2016) conclude that Caucasilac-

erta must replace Darevskia as the valid nomen

for the genus of the Caucasian Rock Lizards.

The conclusions drawn by Busack et al. (2016)

prompted a number of responses (Arribas,

2016; Arribas et al., 2017), advocating that Cau-

casilacerta is a nomen nudum and hence is un-

available. Subsequently, Arribas et al. (2018)

appealed to the ICZN (under Articles 78.1 and

81 of the Code) to accept Arribas (1997) as pub-

lished in the sense of the Code and preserve

Darevskia Arribas, 1997 and Iberolacerta Ar-

ribas, 1997. While we regard Caucasilacerta as

available, we support the current application to

preserve Darevskia. We note that, while the case

is under consideration by the Commission, the

prevailing usage of Darevskia and Iberolacerta

with their authorship as Arribas, 1997 is to be

maintained (Article 82).

Based on morphological variation, Stugren

(1961) advocated the recognition of three sub-

species of Darevskia praticola: the nomino-

typical subspecies in the east of the distribu-

tion (central and eastern Caucasus, northern

and south-eastern Georgia, northern Armenia,

southern Azerbaijan, and north-eastern Iran),

the subspecies pontica in the west of the Cauca-

sus and in western Georgia, and D. p. hungarica

in Europe (north-eastern Serbia, southern Ro-

mania, north-eastern Greece, eastern and west-

ern Bulgaria). Later, he recommended to treat

D. p. hungarica as a synonym of D. p. pontica,

resulting in a two-subspecies classiﬁcation (Stu-

gren, 1984). On the basis of multivariate anal-

ysis of morphological, meristic and qualitative

characters of Balkan and Caucasus populations,

Ljubisavljevic et al. (2006) conﬁrmed the valid-

ity of the two subspecies D. p. praticola and

D. p. pontica. Tuniyev et al. (2011) presented

a detailed morphological study for the popula-

tions of the Caucasian Isthmus. They conﬁrmed

two morphological clusters in the Caucasus and

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Species list of European herpetofauna 157

Transcaucasia (western Caucasus: D. p. pon-

tica; eastern Caucasus +Transcaucasia: D. p.

praticola), and described a new subspecies from

the Talysh Mountains (the extralimital D. p. hyr-

canica). On the basis of morphological differen-

tiation and unpublished genetic data, they also

advocated species status for D. p. pontica. Later,

Freitas et al. (2016) provided a phylogeographic

study of the species, based on comprehensive

sampling covering most of the range (although

the central and eastern Caucasus parts of the

range were missing). They recovered two clades

in the Caucasus and in Transcaucasia, one in the

western Caucasus (corresponding with the Cau-

casian range of the D. p. pontica subspecies)

and one in Transcaucasia (Armenia) and the

Talysh Mountains, corresponding with the ex-

tralimital subspecies D. p. hyrcanica and Tu-

niyev’s et al. (2013) D. p. loriensis. Samples

of the nominotypical subspecies were not in-

cluded, although their morphology suggests that

they would presumably group with the Tran-

scaucasian samples (see Tuniyev et al., 2011,

2013). Freitas et al.’s (2016) results thus seem

to support the divergence of two lineages in the

Caucasus, corresponding to Caucasian popula-

tions of pontica on the one hand, and popula-

tions of the praticola phenotype (including lo-

riensis and hyrcanica) on the other hand. How-

ever, genetic analyses of samples of praticola

from the eastern Caucasus would be needed to

conﬁrm this. More surprisingly, Freitas et al.

(2016) also uncovered a deep split within popu-

lations traditionally classiﬁed as pontica, with

the European samples from the Balkans forming

a deeply divergent lineage in both mtDNA and

nuclear DNA that splits from a node basal to

the clade grouping the Caucasian and Transcau-

casian samples. They estimated the divergence

between the European and Caucasian +Tran-

scaucasian clades at around 2.5 Mya, with a

much younger split of the Transcaucasian lin-

eage around 650,000 year ago. As pointed out

by Ljubisavljevic et al. (2006), the name D. p.

hungarica (Sobolevsky, 1930), with type local-

ity in the Transylvanian Alps (which at the time

was part of the Kingdom of Hungary, includ-

ing parts of today’s Romania), should be applied

to the Balkan lineage. Given the incomplete ge-

ographic coverage of Freitas et al. (2016) and

the somewhat borderline level of genetic diver-

gence between the Balkans and Caucasian +

Transcaucasian lineages, we refrain from rec-

ommending species status for the time being,

and thus suggest to treat the Balkan popula-

tions as Darevskia praticola hungarica.Wefur-

ther disagree with the conclusion of Freitas et

al. (2016) to treat D. p. pontica as a synonym

of D. p. praticola, as all their samples corre-

spond with the range of D. p. pontica according

to Tuniyev et al. (2011), and they have there-

fore not analysed any sample from the range

of D. p. praticola. Conclusively, we maintain

the morphologically well-supported subspecies

D. p. pontica and D. p. praticola for the Cau-

casian populations occurring in our area, while

the Balkan populations are referred to as the

subspecies D. p. hungarica.

Genetic relationships based on a mtDNA (cyt

b) fragment suggest that Darevskia brauneri

szczerbaki is sister to D. saxicola rather than

grouping with other lineages of D. brauneri

(Doronin, Tuniyev and Kukushkin, 2013;

Tarkhnishvili et al., 2016). Based on this and

on analysis of morphological variation cou-

pled with differences in ecology, Tuniyev and

Tuniyev (2012) and Doronin, Tuniyev and

Kukushkin (2013) suggested to treat szczerbaki

as a valid, monotypic species. However, this hy-

pothesis is in conﬂict with the results of Mac-

Culloch et al. (2000), who found a lack of ﬁxed

differences in 15 polymorphic allozyme loci be-

tween D. b. brauneri,D. b. darevskii and D.

b. szczerbaki, and concluded that these three

taxa are conspeciﬁc. The high incidence of hy-

bridisation within Caucasian Darevskia makes

mtDNA relationships potentially unreliable. In

addition, because of the low level of cyt bdi-

vergence (2.4% between D. saxicola and D. b.

szczerbaki, and 4.8-5.6% of both D. saxicola

and D. b. szczerbaki towards other D. brauneri

sequences) and the conﬂict with allozyme data,

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158 J. Speybroeck et al.

we consider this species complex to be cur-

rently too poorly studied to recognise additional

species without extensive evidence from nuclear

genes and/or strong morphological differentia-

tion in sympatry or close parapatry. As both of

these lines of evidence are still lacking in this

case, we maintain D. b. szczerbaki at subspecies

rank for the time being.

Two parthenogenetic Darevskia species

which are not native to our area have been intro-

duced near Zhytomyr, Ukraine (Nekrasova and

Kostiushyn, 2016). Darevskia armeniaca oc-

curs naturally in north-western Armenia, west-

ern Azerbaijan, southern Georgia and north-

eastern Turkey, and is believed to have been

introduced in 1963, whereas D. dahli, known

from northern Armenia and southern Georgia,

was discovered more recently (1980) within

the introduced armeniaca population. A third

species, D. mixta, was also introduced in 1968,

but its contemporary presence could not be con-

ﬁrmed (Nekrasova and Kostiushyn, 2016).

Mayer et al. (2000) described the oviparous

Zootoca vivipara populations of Carinthia (Aus-

tria), Slovenia, Friuli and isolated sites across

the Po Plain (Italy) as the distinct and allopatric

subspecies Z. v. carniolica. Additional molecu-

lar analysis conﬁrmed its validity (Surget-Groba

et al., 2001, 2002). More recently, this taxon

was shown to inhabit a wider range than previ-

ously known, not only occupying parts of south-

ern Austria, Slovenia, northern Croatia and

north-eastern Italy, but actually extending west

as far as the central southern Italian Alps (Cor-

netti et al., 2015a). Lindtke, Mayer and Böhme

(2010) documented a narrow contact zone in

Carinthia (Austria), and detected two natural

hybrids (identiﬁed by clutch features) in a sam-

ple of 36 specimens. As they did not use genetic

markers, they could not investigate the level

of introgression. More recently, two studies by

Cornetti et al. (2015a, b) demonstrated complete

reproductive isolation between carniolica and

vivipara, in spite of widespread close parapatry

between these two taxa. Cornetti et al. (2015a)

focused on the contact zone with local syntopy

in northern Italy, and did not detect any hybrid

or admixed genotype out of around 30 individ-

uals (genotyped with 13 microsatellites). Cor-

netti et al. (2015b) genotyped many more in-

dividuals from several regions in the northern

Italian Alps where the two taxa come in close

parapatry, and found a similar lack of admix-

ture. Thus, carniolica and vivipara appear to

represent two different species, and we accept

Zootoca carniolica at species level.

After being placed into the genus Lacerta,

Teira dugesii and Scelarcis perspicillata were

transferred to the subgenus Teira and the genus

Podarcis, respectively, based on morphology

(Richter, 1980). Subsequently, Teira was ele-

vated to genus level by Mayer and Bischoff

(1996). Yet, the generic allocation of S. per-

spicillata and T. dugesii remained a matter of

confusion, with several authors attributing both

species to the genus Lacerta until quite recently

(e.g. Brehm et al., 2003; Perera et al., 2007).

Even though Pavlicev and Mayer (2009) sug-

gested treating Scelarcis as a junior synonym

of Teira and placing both species into the genus

Teira , SBC2010 adopted the genus arrangement

of Arnold, Arribas and Carranza (2007) and as-

signed the Moroccan Rock Lizard to Scelar-

cis as Scelarcis perspicillata. However, a recent

study using data from anchored phylogenomics

and a fossil-dated time tree by Garcia-Porta et

al. (2019) conﬁrmed that Teira and Scelarcis

are sister taxa and that their divergence is at a

similar level or even lower than among species

of many well-established genera. According to

this study, Teira and Scelarcis diverged from

each other in the mid-Miocene, coinciding with

or even post-dating intrageneric splits in many

other lacertid genera such as Algyroides,Gallo-

tia,Iberolacerta,Lacerta,Podarcis,Psammod-

romus,Takydromus, and Timon. Mendes et al.

(2016) argued to maintain Scelarcis mainly be-

cause of nomenclatural stability. We follow this

approach and maintain Teira and Scelarcis as

valid genera, awaiting additional evidence.

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Species list of European herpetofauna 159

Psonis et al. (2017, 2018) examined species

delimitation in the Podarcis tauricus group (in-

cluding P. tauricus,P. milensis,P. gaigeae

and P. melisellensis) using mitochondrial and

nuclear DNA sequences, microsatellites and

RADseq data. Their most complete phyloge-

nomic analysis offered a fully resolved species

level phylogeny, and conﬁrmed the existence

of extensive genetic structuring within P. tauri-

cus. Two main clades correspond with the west-

ern and eastern part of the distribution of the

species on either side of the Pindos Mountains.

The level of divergence between them is com-

parable with their divergence from the well-

established species P. melisellensis,P. milen-

sis and P. gaigeae, and extensive admixture is

not apparent from the available data. They dif-

fer quite obviously in coloration of adult males.

Even if no information on the nature of the con-

tact zones is available, the data at hand support

the split of P. tauricus into an eastern species

(retaining the name P. tauricus and including

the subspecies P. t. thasopulae) and a western

species, for which the oldest available name is

P. ionicus (Psonis et al., 2017, 2018). Several

of the subclades within P. ionicus, supported by

mitochondrial and genomic data, are as diver-

gent as other species pairs in the genus Podar-

cis and probably warrant species status as well,

but formal recognition as separate species is still

lacking. On the basis of published information,

we accept the split of Podarcis tauricus into two

species: P. tauricus and P. ionicus.

Using published data on three mitochondrial

and three nuclear genes, Senczuk et al. (2019)

examined the phylogeny of the genus Podarcis

to address the systematic status of the Podar-

cis siculus populations from the Western Pon-

tine Islands, off the Italian west coast. They

estimated that the Western Pontine lineage di-

verged from all other P. siculus lineages around

4 Mya, and showed that this genetic diver-

gence is similar to or greater than that between

many other species pairs of Podarcis lizards.

Hence, they formally raised the Western Pontine

populations, currently classiﬁed as several sub-

species of P. siculus, to species rank as Podar-

cis latastei,aslatastei is the oldest name avail-

able for this lineage. At this stage we are re-

luctant to follow this decision for two reasons.

Firstly, the amount of divergence between the

Western Pontine populations and the rest of the

P. siculus complex in Senczuk et al. (2019) is

only presented as a combined mtDNA and nu-

clear DNA tree, not allowing to check if nuclear

data independently supports this species-level

divergence or not. As multiple examples of deep

mtDNA divergence that are not reﬂected in ge-

nomic divergence are known, we feel that com-

pelling evidence of nuclear DNA divergence is

needed before adopting this new species. Sec-

ondly, Senczuk et al. (2019) presented a simpli-

ﬁed picture of the patterns of genetic divergence

in P. siculus. More detailed results by Senczuk

et al. (2017, 2018) display a much more com-

plex situation, where the Western Pontine lin-

eage appears as one of several deeply diver-

gent lineages in P. siculus, and latastei and sicu-

lus are not reciprocally monophyletic. While

Senczuk et al. (2018) indicated differentiation

in nuclear DNA between at least some of the

main mtDNA lineages in P. siculus, no infor-

mation on their level of reproductive isolation is

yet available. As a consequence, while the TC

acknowledges that P. siculus as currently under-

stood is possibly made up of several species,

one of them corresponding to the Western Pon-

tine islands populations, we consider adopting

any formal taxonomic change premature.

Michels and Bauer (2004) provided a list of

what they argued to be justiﬁed emendations

(citing Article 31.1.2 of the Code) of non-avian

reptile and amphibian scientiﬁc names which

the authors identiﬁed as inappropriate origi-

nal constructions of patronyms or matronyms.

Among the listed species was Podarcis raffonei

(as Lacerta sicula raffonei Mertens, 1952). Be-

cause this taxon was named in honour of the

collectors’ wife, Michels and Bauer (2004) con-

cluded that raffonei should be changed into raf-

foneae. However, Article 31.1.2 simply states

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160 J. Speybroeck et al.

how species-group names should originally be

formed. The article does not provide grounds

for subsequent changes to the name. Further-

more, justiﬁed emendations can only be made

to ‘spellings that must be corrected’ (Article

32.5, see also 33.2.2), in which these cases are

not included. Even though Michels and Bauer

(2004) are linguistically correct (Arribas, 2017),

amending raffonei to raffoneae is unjustiﬁed

(Article 33.2.3), making the latter an unjustiﬁed

emendation and hence a junior objective syn-

onym of the former (see also Dubois, 2007).

We thus recommend to maintain the original

spelling P. raffonei. As a side note, we highlight

that the results of Senczuk et al. (2019) also cast

serious doubts on the validity of the species sta-

tus of P. raffonei, but we refrain here from ad-

vocating any change for now.

Investigating the species group comprising

Podarcis cretensis,P. levendis and P. pelo-

ponnesiacus, Spilani et al. (2019) found well-

supported differentiation within the latter

species. Based on 17 microsatellites, the east-

ern populations diverged 1.86 Mya. Most note-

worthy, the eastern and western lineages occur

in actual or near syntopy without any apparent

sign of admixture. The observed level of diver-

gence, together with evidence of reproductive

isolation, warrant a species-level split within P.

peloponnesiacus. However, Spilani et al. (2019)

refrain from making any taxonomic change. The

TC follows this position, as we await a forth-

coming morphological study, prior to proposing

a formal change.

From a long line of studies on the Iberian

Podarcis species, the acceptance of Podarcis li-

olepis and the redeﬁnition of Podarcis hispan-

icus sensu stricto were the most recent steps

that were adopted by SBC2010. At the time,

two additional species were sufﬁciently sub-

stantiated, genetically as well as morphologi-

cally. Awaiting formal naming, they were listed

as Podarcis hispanicus type 1 (north-western

Iberian Peninsula and central Iberian moun-

tain chains) and type 2 (parts of central and

south-western Iberian Peninsula). Geniez et al.

(2014) provided additional morphological anal-

yses, and formally named them as Podarcis

guadarramae (former type 1) and P. virescens

(former type 2). Morphological and molecular

differentiation within Podarcis guadarramae re-

sulted in the description of the subspecies P. g.

lusitanicus from northern Portugal and north-

western Spain. We accept P. guadarramae and

P. virescens as valid species, while the status of

lusitanicus will be the focus of a forthcoming

study (G. Dias and C. Pinho, pers. comm.).

Although sometimes credited to Gray, 1825

(e.g. Sindaco and Jeremˇ

cenko, 2008), the fam-

ily name Scincidae must be credited to Oppel

(1811), as adopted by Hedges (2014) and oth-

ers. Oppel (1811) created the name as “Familia.

Scincoides”. Article 11.7.3 of the Code (see

family Agamidae above) makes it clear that the

nomen is made available by its publication in

Oppel (1811), even if the sufﬁx must be cor-

rected.

A split of the Scincidae into seven families

would affect the systematics of the European

genera, with Ablepharus being placed into Eu-

gongylidae, Heremites into Mabuyidae and the

remaining genera Chalcides,Ophiomorus and

Eumeces remaining in Scincidae s.s. (Hedges

and Conn, 2012; Hedges, 2014). This was nei-

ther adopted in the large-scale squamate phy-

logeny of Zheng and Wiens (2016), nor in

the online Reptile Database (Uetz, Freed and

Hošek, 2019). As most skinks are readily recog-

nisable as belonging to this group, and the

monophyly of Scincidae sensu lato is well es-

tablished, we follow the argument of the ‘phe-

notypic diagnosability’ taxon naming criterion

of Vences et al. (2013a) for the time being, keep-

ing all species (even if there are many) within a

single family. Distinction of subfamilies, how-

ever, certainly makes sense as it provides more

manageable units for taxonomic revision and

other purposes.

For a long time, the skink genus Mabuya in-

cluded numerous species from South America,

Africa and Asia. A series of phylogenetic works

(e.g. Mausfeld et al., 2000, 2002; Carranza and

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Species list of European herpetofauna 161

Arnold, 2003; Mausfeld and Schmitz, 2003;

Whiting et al., 2006) conﬁrmed the monophyly

of the genus, but also uncovered deep evolu-

tionary divergence between its main radiations,

which also differ consistently in several mor-

phological and life-history traits. The initial di-

vergence within the group is estimated to date

back to the Eocene, between 40 and 50 million

years ago (Karin et al., 2016). Mausfeld et al.

(2002) were the ﬁrst to suggest a split of the

genus Mabuya to reﬂect the long evolutionary

divergence of its main lineages, a proposal that

has been universally adopted since. The Middle

Eastern Mabuya species (M. aurata,M. septem-

taeniata and M. vittata) form a well-supported

monophyletic group in all phylogenies. They

have been classiﬁed with the African species

in the genus Trachylepis for several years but,

even if their phylogenetic position within the

radiation of the former Mabuya is still poorly

resolved, they are not part of the radiation of

African species in the genus Trachylepis (Karin

et al., 2016). Their origin is quite ancient (esti-

mated around the late Oligocene, more than 30

million years ago) and their clade is thus now

afforded genus rank by all recent works on the

subject, a position that we follow here. The valid

genus-group nomen of the Middle-East clade is

Heremites Gray, 1845 (masculine, see Karin et

al., 2016). We thus recommend calling the Mid-

dle East species Heremites auratus,H. vittatus

and the extralimital H. septemtaeniatus.

The phylogeography of Ophiomorus punc-

tatissimus was examined by Poulakakis et al.

(2008) using three mitochondrial fragments.

On the basis of divergence rates of mitochon-

drial genes calibrated in other groups of squa-

mates, they uncovered a deep divergence be-

tween the Asian (Anatolian) and European

populations of the species, estimated to have

occurred around 10 Mya. This date ﬁts nicely

with the opening of the mid-Aegean trench,

estimated from geological information to have

started around 12 Mya and being completed

around 9 Mya. More recently, Kornilios et al.

(2018) used mtDNA and nuclear DNA data

(three nuclear genes) for single-locus and mul-

tilocus coalescent-based species delimitation.

The established COI p-distance is substantial

(12.7%). No allele sharing was found, and the

Asian and European lineages appeared recipro-

cally monophyletic for all three nuclear genes,

conﬁrming their long-term isolation. Even if

suboptimal preservation of specimens and/or

morphological conservatism in these fossorial

species did not allow identifying any morpholo-

gical differentiation between the two allopatric

Ophiomorus lineages, the authors advocated

recognition of the eastern lineage from Ana-

tolia as a new species, which they described

as O. kardesi. The status of several mitochon-

drial lineages recovered within O. punctatis-

simus awaits further study. Based on the diver-

gence time and level of genetic divergence be-

tween these two cryptic species, we accept the

species status of O. kardesi. In our area, the new

species occurs on the Greek island of Kastel-

lorizo (Kornilios et al., 2018).

Phylogenetic analysis based on mitochon-

drial and nuclear markers (Skourtanioti et al.,

2016) showed strong divergence, supporting

the independent history of a clade of snake-

eyed skinks (Ablepharus) from Kastellorizo and

southwestern Turkey. The samples of this clade

originate from an area where the only taxon

known to occur according to Schmidtler (1997)

is Ablepharus budaki anatolicus. As a conse-

quence, Skourtanioti et al. (2016) suggested

that the name anatolicus is available for this

new species-level lineage, which splits from a

more basal node than all other but one (A. pan-

nonicus from Iran) species of the genus, and

whose divergence was estimated to date back to

the mid-Miocene. As the genetic differentiation

and deep phylogenetic divergence of the new

species was conﬁrmed by independent analy-

ses of both mitochondrial and nuclear genes, we

agree that the genetic data strongly support the

recognition of an additional Ablepharus species

from SW Turkey (including Kastellorizo and

hence entering our area). However, we refrain

from formally accepting a new species here and

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162 J. Speybroeck et al.

prefer to wait until genetic data conﬁrm that ani-

mals from the type locality of anatolicus indeed

represent this new lineage.

Sindaco and Jeremˇ

cenko (2008), who pro-

vided the starting point of our taxonomy for

areas not covered by SBC2010 or Gasc et

al. (1997), follow Pasteur, Keymar and Perret

(1988) in treating the Gran Canaria skinks as

two species, Chalcides bistriatus in the north-

east of the island and C. sexlineatus in the south-

west. This classiﬁcation was based on a sharp

geographic transition in morphology and (es-

pecially) colour pattern between the skinks in-

habiting the mesic and xeric habitats on Gran

Canaria (see Brown and Thorpe, 1991a, b, al-

though these authors rejected the hypothesis of

speciation between these morphotypes). Studies

on the evolutionary history of these populations

using allozymes (Mayer and Tiedemann, 1991)

or a combination of mtDNA and microsatel-

lites (Suárez, Pestano and Brown, 2014) estab-

lished that this differentiation has a genetic ba-

sis, originating from allopatric diversiﬁcation

within Gran Canaria. This divergence is linked

to volcanic activity 1.5-3 Mya and is currently

maintained by strong selection on coloration,

in spite of considerable introgression and gene

ﬂow between the two morphs. While the deepest

divergence between the mtDNA clades within

Gran Canaria is dated to 1.5 Mya, the diver-

gence between the two morphs was dated to

260,000 years ago. This discrepancy could re-

ﬂect real differences between coalescence time

of mtDNA lineages versus divergence time of

populations, or analytical issues with one of the

methods. Either way, the available data suggest

a young divergence of C. s. sexlineatus and C.

s. bistriatus, together with a lack of reproduc-

tive isolation. We thus recommend to treat these

taxa as conspeciﬁc as C. s. sexlineatus and C. s.

bistriatus, as adopted by Mayer and Tiedemann

(1991), Carranza et al. (2008) and Salvador and

Brown (2015).

Mitochondrial DNA data of the Canarian

skinks establish with a very high support that

the Chalcides from Tenerife (C. v. viridanus)

and La Gomera and El Hierro (C. v. coeruleop-

unctatus) are not each other’s closest relative,

with coeruleopunctatus being more closely re-

lated to the taxa from Gran Canaria (sexlinea-

tus and bistriatus) than to viridanus (Brown and

Pestano, 1998; Carranza et al., 2008). These re-

lationships rule out maintaining coeruleopunc-

tatus as a subspecies of viridanus, as was tra-

ditionally the case based on their morpholo-

gical similarity. In addition, the amount of di-

vergence between C. sexlineatus,C. viridanus

and C. coeruleopunctatus is typical of interspe-

ciﬁc divergences in the genus Chalcides (Car-

ranza et al., 2008). We thus accept the conclu-

sion of Carranza et al. (2008) and treat the an-

imals from La Gomera and El Hierro as Chal-

cides coeruleopunctatus.

The publication dates of the works of Daudin

have been worked out by Harper (1940). The

volume 4 of “Histoire Naturelle des Reptiles”,

containing the description of Eumeces schnei-

derii, was published in August 1802. The spe-

ciﬁc epithet is often spelled schneideri,even

if both schneideri and schneiderii areinuse.

The original spelling is schneiderii (see Daudin,

1802), with schneideri as an incorrect subse-

quent spelling. The only reason to use this

spelling rather than the original spelling would

be if the incorrect subsequent spelling schnei-

deri was in prevailing usage (see Art. 33.3.1

of the Code). Prevailing usage is deﬁned by

the Code as “that usage of the name which is

adopted by at least a substantial majority of the

most recent authors concerned with the rele-

vant taxon”. We checked the number of publica-

tions using either spelling after 2000 using the

electronic version of the Zoological Record and

found that 17 works published after 2000 used

schneiderii while 28 used schneideri. We inter-

pret this as a lack of clear support for a prevail-

ing usage of the incorrect subsequent spelling

and thus adopt the original spelling E. schnei-

derii.

Considerable divergence within the genus

Anguis has been revealed by means of genetic

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Species list of European herpetofauna 163

and morphological analyses (Cabela and Gril-

litsch, 1989; Gvoždík et al., 2010). This led to

the recognition of A. graeca and A. colchica,re-

sulting in the acceptance of at least four Anguis

species in Europe by Gvoždík et al. (2010) and,

subsequently, SBC2010. Although no Italian

specimens were incorporated in previous anal-

yses, Anguis fragilis was hitherto presumed to

inhabit much of western, central and eastern Eu-

rope, including the Italian Peninsula (Gvoždík

et al., 2010). More recently, Gvoždík et al.

(2013) showed Italian Anguis to represent a

deeply differentiated mtDNA clade, which pre-

sumably diverged during or shortly after the

basal radiation within the genus. Genetic dis-

tances between the Italian clade and all recog-

nised species are larger than the interspeciﬁc

distances among A. fragilis,A. graeca and A.

colchica. Hence, Gvoždík et al. (2013) proposed

to recognise the populations from Italy and ad-

jacent extreme south-eastern France as Anguis

veronensis. Two specimens (one from NE Italy

and one from Slovenia) were heterozygous for

the most common A. fragilis and A. veronen-

sis PRLR haplotypes and carried A. fragilis

mtDNA, indicating hybridisation. While mor-

phological differentiation between A. veronen-

sis and A. fragilis is signiﬁcant, the ranges of all

studied characters overlap. In light of the earlier

accepted splits among the European Anguis by

SBC2010, the divergence during the basal ra-

diation of the genus, and the genetic and mor-

phological evidence presented by Gvoždík et

al. (2013), we recognise the Italian and south-

eastern French populations as A. veronensis.

Albert and Fernández (2009) partitioned the

Iberian Worm Lizard Blanus cinereus into two

species, attributing southwestern Iberian popu-

lations to the cryptic Blanus mariae and desig-

nating a lectotype for B. cinereus. Their lecto-

type designation was, however, found to be in-

valid, due to the lack of explicitly stated tax-

onomic purpose, while they did not establish

whether the original type series consists of one

holotype or several syntypes, nor examined the

status of other, older nomina in the synonymy of

Iberian Blanus (SBC2010). These issues were

tackled by Ceríaco and Bauer (2018), who ar-

gued that the type locality of Amphisbaena

cinerea Vandelli, 1797 falls within the range of

populations attributed to B. mariae by Albert

and Fernández (2009). To further strengthen

their interpretation, they made a valid neotype

designation for B. cinereus, selecting a neo-

type belonging to the south-western Iberian

species. This effectively places B. mariae in

the synonymy of B. cinereus, which becomes

the valid name for the south-western Iberian

Blanus species. In agreement with Sampaio et

al. (2015), who conﬁrmed that the two Iberian

Blanus clades show high levels of mitochon-

drial differentiation and no signs of nuclear

haplotype sharing, Ceríaco and Bauer (2018)

accepted the existence of two Iberian Blanus

species. However, they deemed no name to

be valid for the Central Iberian species. Yet,

two older names, Amphisbaena oxyura Wagler,

1824 and Amphisbaena rufa Hemprich, 1820,

are available and relate to Blanus from Spain.

The type specimen of A. oxyura is lost, while

its type locality (‘Spain’, according to Wagler,

1830) makes it impossible to link this name

to either of the two Spanish Blanus species.

The name A. rufa was based on a holotype that

is still present in the Zoological Museum in

Berlin (ZMB), but the catalogue entry for this

specimen shows that it originates from south-

ern Spain, where both Blanus clades occur. As

morphological identiﬁcation of the two Span-

ish Blanus species is currently impossible, ro-

bust attribution of A. rufa to either clade re-

quires molecular data. Yet, this is problematic

with the currently available methods (Ceríaco

and Bauer, 2018). Ceríaco and Bauer (2018)

thus treated these two names as nomina du-

bia and created the new nomen Blanus vandel-

lii. SBC2010 had already recognised the two

Iberian Blanus clades at species level. We fol-

low this arrangement, and accept that B. mariae

is a junior synonym of B. cinereus, with the

latter being the valid name for the southwest-

ern Blanus clade (Ceríaco and Bauer, 2018).

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164 J. Speybroeck et al.

However, instead of introducing a new name in

the form of B. vandellii, we promote the use

of the oldest available name, Blanus rufus,as

the valid name for the central clade until in-

formation on the identity of the A. rufa holo-

type can be obtained. Rather than discarding

the old name until proven to apply, we advocate

that when an available name exists, it should be

used until proven not to apply to the relevant

taxon. We thus list the two Iberian species as

B. cinereus (south-western species) and B. ru-

fus (north-eastern species).

Hedges et al. (2014) present molecular and

morphological data to clarify the phylogeny of

the Typhlopidae and split the large genus Ty-

phlops into a number of well-deﬁned new gene-

ra. As a consequence, the European blind snake

species is now referred to as Xerotyphlops ver-

micularis. Additionally, according to Hedges et

al. (2014), the non-native Flowerpot Snake, re-

cently introduced to Italy and Spain (Zamora-

Camacho, 2017; Paolino, Scotti and Grano,

2019), is now called Indotyphlops braminus.

The name of Hierophis viridiﬂavus has been

involved in nomenclatural debate over the last

few decades, but are now solved (ICZN Opin-

ions 1463 and 1686). In 1833, Charles-Lucien

Bonaparte (the nephew of Napoleon) described

Coluber viridi-ﬂavus carbonarius for the east-

ern Italian populations that are completely or

almost completely black. The subspecies H. v.

carbonarius was considered valid, until Schätti

and Vanni (1986) placed it in the synonymy of

the nominal form. A series of phylogeographic

studies based on mtDNA and nuclear DNA con-

ﬁrmed that viridiﬂavus and carbonarius are dis-

tinct evolutionary lineages which form recip-

rocally monophyletic clades in mtDNA (Nagy

et al., 2001; Rato et al., 2009; Mezzasalma et

al., 2015; Avella, Castiglia and Senczuk, 2017).

These two lineages differ in the morphology of

their W chromosomes (submetacentric in the E

clade and telocentric in the W clade – Mezza-

salma et al., 2015). As a consequence, they have

been treated as valid subspecies since Nagy et

al. (2001). In 2015, Mezzasalma et al. (2015)

proposed to treat H. viridiﬂavus and H. car-

bonarius as different species on the basis of

(i) different morphology of the W sex chro-

mosome, (ii) reciprocal monophyly in mtDNA

with a genetic divergence of 4% in both cyt

band ND4, and (iii) morphological differen-

tiation consistent with mtDNA differentiation.

However, Rato et al. (2009) found a lack of

general agreement between colour pattern and

mtDNA clade in Italy, and no differences in

the nuclear intron beta-ﬁbrinogen intron 7 be-

tween the two lineages. The amount of diver-

gence in mtDNA between H. v. carbonarius and

H. v. viridiﬂavus is much lower than between

H. gemonensis and H. viridiﬂavus, while the

amount of divergence in PRLR between them

resembles the divergence within H. gemonensis.

The sample size for the chromosomes is quite

low as well. The split of H. v. carbonarius from

H. v. viridiﬂavus did not reach broad support in

the TC, as several members felt the evidence

is still insufﬁcient, and they prefer to wait for

information on genetic variation near the con-

tact zones. We thus recommend to maintain Hi-

erophis viridiﬂavus carbonarius as a subspecies

for the time being.

Jablonski et al. (2019) applied phylogenetic

and morphological analyses to range-wide sam-

pled data, aiming to better understand the in-

traspeciﬁc relationships and biogeography of

Elaphe sauromates, and subsequently revised

its taxonomy. Known from the Balkans, Ana-

tolia, the Caucasus, the Ponto-Caspian steppes,

and the Levant, this species has been suspected

to be composed of two or more genetically

diverse populations (Lenk, Joger and Wink,

2001). Sequences from 63 specimens and mor-

phological data from 95 specimens were anal-

ysed. The authors found two distinct evolution-

ary lineages, one of which represents a new

species, Elaphe urartica. The new species is

distributed in eastern Turkey, Georgia, Armenia,

Azerbaijan, Nagorno-Karabakh, Iran, and Rus-

sia (Dagestan). The mtDNA genetic distances

between E. sauromates and E. urartica (6-8%

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Species list of European herpetofauna 165

in COI and ND4) were interpreted as indicat-

ing a split at the Miocene-Pliocene boundary

(5-8 Mya; see Kornilios et al., 2014), thus they

presumably separated not very long after their

common ancestors separated from E. quatuor-

lineata. Two out of four analysed nuclear genes

were also clearly differentiated, though closely

related, between E. urartica and E. sauromates,

while the remaining two showed signs of in-

complete lineage sorting. Both lineages are also

moderately morphologically differentiated and,

while none of the characters are exclusively di-

agnostic, their combination can be used for con-

ﬁdent lineage identiﬁcation. Conclusively, we

accept Elaphe urartica as a separate species.

The ladder snake Rhinechis scalaris is char-

acterised by a backward-oriented rostral scale

that is wedged between the internasal scales.

Apart from this striking feature, external mor-

phology of R. scalaris has long been acknowl-

edged to closely match that of Zamenis rat

snakes (Salvi et al., 2018). Multiple recent

phylogenetic assessments have revealed that

Rhinechis (with the sole member R. scalaris)

and Zamenis are closely related (Burbrink and

Lawson, 2007; Pyron et al., 2011; Pyron, Bur-

brink and Wiens, 2013; Zheng and Wiens,

2016). Some of these studies found Zamenis to

be paraphyletic in respect to Rhinechis, while

others placed Rhinechis as sister to all Zamenis

species, albeit with low support (reviewed by

Salvi et al., 2018). The latter authors argue that

this phylogenetic uncertainty reﬂects an evo-

lutionary scenario in which early cladogenetic

events took place at the same time or within a

short time frame, making it hard to disentangle

which (if any) lineage branched off ﬁrst. Us-

ing two mitochondrial and ﬁve nuclear genes,

Salvi et al. (2018) then inferred phylogenetic

relationships between Zamenis and Rhinechis.

They recovered a strongly supported clade con-

sisting of Rhinechis scalaris and Zamenis, while

the monophyly of Zamenis without scalaris re-

mained poorly supported. As a result, these au-

thors proposed to include scalaris in Zamenis.

In agreement with the ‘clade stability’ taxon

naming criterion of Vences et al. (2013a) and as

a named genus of uncontroversial monophyly is

obtained, we here follow this.

The California Kingsnake Lampropeltis

getula californiae has been introduced to Gran

Canaria, where it turned into a problematic in-

vasive species (Monzón-Arguëllo et al., 2015;

Fisher et al., 2019). Based on mtDNA data and

morphological variation, Pyron and Burbrink

(2009) elevated L. g. californiae and four other

subspecies to species status. While the basal di-

vision in their phylogeny has L. g. getula and L.

g. nigra in the one main branch and L. g. hol-

brooki,L. g. splendida and L. g. californiae in

the other, the division between L. g. splendida

and L. g. californiae is retrieved as the most re-

cent one. Pending the publication of a multilo-

cus analysis or a detailed study of the various

contact zones, we maintain L. g. californiae as

a subspecies of the Common Kingsnake Lam-

propeltis getula for the time being.

The groundwork of contemporary elucidation

of the Natrix natrix complex was initiated by

the mitochondrial phylogeography of Kindler

et al. (2013), leading to rejection of Natrix

megalocephala and the identiﬁcation of multi-

ple contact zones, three of which were further

investigated more recently (see below). Subse-

quently, a virtual lack of gene ﬂow between

N. natrix helvetica and N. natrix astreptophora

and the differentiation between both taxa es-

tablished on multiple sources of evidence, such

as microsatellites, mtDNA, osteology and exter-

nal morphology, led to suggesting species status

for Natrix astreptophora (Pokrant et al., 2016).

Within central Europe, Kindler et al. (2017) in-

vestigated two macro-scale transects across Na-

trix transition zones. The eastern contact zone

between two lineages identiﬁed by Kindler et

al. (2013) as running from Central Europe to

the southern Balkans displayed wide-reaching

introgression. In contrast, the western contact

zone between the traditional subspecies N. n.

natrix and N. n. helvetica in western Germany

was shown to be narrow (microsatellite cline

width of about 40 km), with a deﬁcit of admixed

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166 J. Speybroeck et al.

individuals generating a bimodal hybrid zone,

indicative of strongly restricted gene ﬂow. We

therefore accept N. astreptophora and N. hel-

vetica as valid species in addition to N. natrix.

Ferchaud et al. (2012) provided a well-

resolved mitochondrial phylogeny of the Vipera

ursinii complex, which indicates that the re-

nardi clade, the ursinii clade and the graeca

clade diverged approximately 3-4 Mya. The di-

vergence between these three clades is of the

same magnitude as between the two widely ac-

cepted species V. berus and V. seoanei.There-

nardi and ursinii clades do not seem to share

haplotypes, although they inhabit similar habi-

tats along the north-western Black Sea coast

where Vipera ursinii moldavica occurs. Given

the level of genetic divergence, lack of haplo-

type sharing, marked morphological divergence

and lack of well supported morphological in-

tergradation, it seems more adequate to treat V.

renardi as speciﬁcally distinct from V. ursinii.

Given its well-supported position at the base

of a renardi +ursinii clade and although we

veriﬁed a very low genetic distance between

ursinii and graeca in cyt b(3.2%), this arrange-

ment makes it necessary to treat V. graeca as a

distinct species as well. Additional morpholo-

gical and mtDNA and nuclear DNA data sup-

port the distinct position of the graeca clade

(Mizsei et al., 2017). Several other taxa from

the Caucasus and the Middle East have been

put forward as valid species, two of which oc-

cur in our area: the Caucasian lotievi and the re-

cently described shemakhensis (Tuniyev et al.,

2013) from north-eastern Azerbaijan. A recent

phylogenomic (RADseq data) analysis of mul-

tiple specimens of renardi and lotievi by Zi-

nenko et al. (2016) showed a lack of genetic dif-

ferentiation at the genome scale between spec-

imens collected in the lowlands (renardi) and

Caucasian uplands (lotievi). There is no genetic

information for shemakhensis, which is, how-

ever, very similar to lotievi morphologically. In

conclusion, we consider three European species

within the Vipera ursinii complex: V. graeca,V.

renardi and V. ursinii, and recommend treating

V. r. lotievi and V. r. shemakhensis as conspeciﬁc

with V. renardi.

Tuniyev and Ostrovskikh (2001) described

Vipera orlovi and V. magniﬁca, two viper

species from the Russian Caucasus that are re-

lated to Vipera kaznakovi.Vipera orlovi inhabits

both slopes of the northwesternmost Caucasus,

while V. magniﬁca is described from the Malaja

Laba river valley and restricted to Krasnodar

territory. As no molecular data was provided,

and it was later shown that both taxa share

mtDNA haplotypes with V. kaznakovi (Zinenko

et al., 2015), further enquiry was needed. Zi-

nenko et al. (2016) used genomic methods to

demonstrate the hybrid origin of these taxa, with

parapatric V. kaznakovi and V. renardi as likely

parental species. Low admixture and limited ge-

netic differentiation led these authors to treat V.

magniﬁca as a synonym of V. kaznakovi. The au-

thors build a case for not revoking the species

status of V. orlovi by advocating caution with re-

gards to its conservation status. While we do not

reject species of hybrid origin per se, the 2016

data does not suggest reproductive isolation or

strong genetic differentiation of either taxon in

relation to V. kaznakovi. Thus, we do not treat

V. magniﬁca or V. orlovi as valid species. As

they are more closely related to V. kaznakovi

than to V. renardi, we treat both as conspeciﬁc

with V. kaznakovi. In addition, we disagree with

Zinenko et al. (2016) that the Code does not al-

low the recognition of taxa with hybrid ancestry.

The Code rules nomenclature, not systematics,

and nothing in the Code precludes treating taxa

of admixed ancestry as valid. As both orlovi and

magniﬁca were described on the base of dis-

tinct phenotypic features, we tentatively main-

tain them as subspecies of V. kaznakovi (V. k.

orlovi and V. k. magniﬁca) until a detailed exam-

ination of morphological variation might show

them not to be morphologically distinct.

A new viper species, named Vipera walser,

was described from northern Piedmont, Italy

(Ghielmi et al., 2016) based on mtDNA, nuclear

DNA and morphological discriminant analy-

ses. Remarkably, while morphologically very

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Species list of European herpetofauna 167

similar to north Italian V. berus, mtDNA data

showed the new taxon to be nested with geo-

graphically distant taxa of the ursinii-kaznakovi

complex. For two protein-coding nuclear genes

(BACH2 and RAG1), V. wa l se r does not share

haplotypes with V. berus. However, only single

individuals of both species have so far been se-

quenced. Moreover, for RAG1, the new species

is similar to V. berus, while no sequences of the

ursinii complex exist for this gene. Therefore,

the existence of cytonuclear discordance can-

not be discarded. Until additional nuclear DNA

data conﬁrms the distinct nature of the walser

population, we consider acceptance of the new

species to be premature, and tentatively regard

it as a population of V. berus with distinct, per-

haps introgressed, mtDNA. Morphologically, V.

walser does not differ much from the V. berus

populations inhabiting the Italian Alps (the Ital-

ian clade of V. berus in Ursenbacher et al., 2006)

but it is easily distinguishable from the Cen-

tral and Northern European populations of V.

berus. Thus, the Italian Alps populations of V.

berus could be treated as a subspecies under the

name V. b. wa l se r . However, we refrain from

making formal recommendations here, pending

a detailed analysis of morphological variation

between populations of V. berus from the Italian

Alps (including the populations described as V.

walser) and the rest of the V. berus populations.

The populations of mostly black adders in-

habiting the forest-steppe zone of the south

and south-eastern parts of the East European

Plain have been described as Vipera nikol-

skii Vedmederya, Grubant and Rudaeva, 1986.

The distribution and morphological characters

of nikolskii compared to berus have been re-

viewed by Bakiev, Böhme and Joger (2005),

Milto and Zinenko (2005) and Zinenko, ¸Tur-

canu and Strugariu (2010, who showed that

populations of nikolskii from the western parts

of its range can be mostly composed of non-

melanistic specimens). According to Milto and

Zinenko (2005), several older names created by

Pallas for black adders from southern Russia

were based on populations resulting from in-

tergradation between berus and nikolskii and

cannot apply to the later taxon, leaving nikol-

skii as the valid nomen for this taxon. While

it was ﬁrst described at species rank, nikolskii

is now variously treated as a full species (e.g.

Bakiev, Böhme and Joger, 2005; Uetz, Freed

and Hošek, 2019) or as a subspecies of Vipera

berus (e.g. SBC2010; Zinenko, ¸Turcanu and

Strugariu, 2010; Mizsei et al., 2017; Crnobrnja

Isailovic et al., 2019 by implication). Depend-

ing on their geographical origin, specimens of

nikolskii either carry berus mtDNA haplotypes,

or haplotypes grouping (within the diversity of

the berus group) close to V. b. barani (Joger

et al., 1997; Kalyabina-Hauf et al., 2004). A

multilocus phylogeny places nikolskii within V.

berus (Mizsei et al., 2017). Detailed morpholo-

gical studies show extensive introgression in the

contact zones with V. b. berus (Zinenko, 2004;

Milto and Zinenko, 2005; Zinenko, ¸Turcanu and

Strugariu, 2010). We thus maintain the treat-

ment of SBC2010 and consider nikolskii as con-

speciﬁc with V. berus, supporting its recognition

as V. b. nikolskii to acknowledge its morpholo-

gical and ecological peculiarities.

The correct spelling of the speciﬁc epithet of

the Lataste’s viper has been a matter of long

lasting debates because Boscá (1878) used the

spelling “Vipera Latasti” in the text and “Vipera

latastei” in the caption for the accompanying

plate, with both spellings in current use in the

literature (see Alonso-Zarazaga, 2013 and Sal-

vador et al., 2014 for summaries). Following an

application to the International Commission on

Zoological Nomenclature (ICZN) by Salvador

et al. (2014), the ICZN (2017) conﬁrmed that

Vipera latastei Boscá, 1878 is the correct origi-

nal spelling of the speciﬁc name.

Frétey (2019) provided compelling argumen-

tation of why the capitalised epithet Lebetinus

in Coluber Lebetinus Linnaeus, 1758 is a noun

in apposition and must be treated as invari-

able. We agree with his interpretation, and ac-

cept that the Blunt-nosed Viper should be called

Macrovipera lebetinus.

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168 J. Speybroeck et al.

Based on morphology, Nilson and Andrén

(1988) elevated Macrovipera lebetinus schweiz-

eri to species rank. Stümpel (2012), however,

found that the mtDNA haplotypes found in

M. schweizeri are not limited to the Milos

archipelago, but are also present near Mersin on

the Turkish mainland. This author thus pointed

out that treating M. schweizeri as a valid species

might render M. lebetinus paraphyletic. Further-

more, they showed that in their mtDNA trees

schweizeri is neither basal to the rest of the

M. lebetinus haplotypes, nor highly divergent.

Yet, while Stümpel’s dissertation is available,

and a (brief) sentence in Stümpel and Joger

(2009) refers to these results, these ﬁndings

have not yet been properly published in a peer-

reviewed journal, and the relevant Macrovipera

sequences are not yet available in GenBank

either. As a consequence, many TC members

felt that a formal taxonomic decision on this

case would be premature, and, while it is likely

that schweizeri is best treated as a subspecies,

the decision can only be taken after the evi-

dence has been properly published, or the data

has been made publicly available. Conclusively,

we maintain Macrovipera schweizeri at species

level for now.

The occurrence of the genus Gloydius in-

side the area covered by this list is unclear.

The only European area where the occurrence

of the genus has been reported is north of the

Caspian Sea, between the Volga and Ural val-

leys. As explained by Conant (1982) and Dirk-

sen and Böhme (2005), all records supported

by specimens in museum collections are either

on the east side of the Ural River, hence out-

side our area (Induski Hills =Indersky Moun-

tains), or cannot be precisely localised (one

specimen from “Gur’jev”, now Atyrau, which

lies at the mouth of the Ural River on both sides

of the river). Attempts to locate Gloydius west

of the Ural River have failed (Conant, 1982;

Bakiev, Ratnikov and Zinenko, 2007; Sarayev

and Pestov, 2010; Simonov, pers. comm.). Thus,

no modern records of the presence of the genus

in Europe exist, and no record is supported by

a collected specimen. The possible occurrence

of Gloydius within Europe is based on several

ancient sources that report the genus between

the Volga and Ural rivers in the dry steppes or

deserts east of Astrakhan (see Conant, 1982 and

Bour, 1993 for details). They all seem to orig-

inate from just two original sources: Gmelin

(1789), who mentions the arid deserts of As-

trakhan for Coluber halys, and Pallas (1799),

who, in his book devoted to his second journey

to the southern areas of Russia, mentions (on p.

112) the species “Berus” and “Halys” were seen

in an area called “Saltan-Murat” and located

east of the Volga River, in an area of dry steppes

close to Astrakhan (more precisely north-east

of present-day Krasnyy Yar, see Conant, 1982

and Bour, 1993). All subsequent references are

based on these two sources. It is even unclear if

the mention of the deserts of Astrakhan in J.F.

Gmelin’s book is based on original information

he received from his uncle J.G. Gmelin, who

also travelled widely in Russia including the

Caspian Sea area, from Pallas himself, or from

any other source. J.F. Gmelin never visited these

areas himself. We may never know how reli-

able these old records are. However, the other

venomous species mentioned by Pallas (1799)

from the same area, Vipera berus, is currently

absent from the dry areas between the Ural and

Volga Rivers. This casts doubt on the reliability

of these observations. In summary, the presence

of the genus Gloydius west of the Ural River

entirely rests on unsubstantiated records that

cannot be veriﬁed. We thus believe the species

should be formally removed from the list of Eu-

ropean species. In addition, we note that the

systematics and nomenclature of the

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

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