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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).
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Amphibia-Reptilia 41 (2020): 139-189
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,
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
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,
Corresponding author;
©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 (, 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
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-
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 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.
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
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).
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 -
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
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.
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 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. (“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. 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
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 bischof(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-
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-
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-
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