ArticlePDF Available

Podarcis vaucheri (Sauria: Lacertidae) far away from home: A new invasive species in Greece


Abstract and Figures

In this study we aimed to clarify the identity of a wall lizard population that deviates phenotypically from the other Podarcis lizards that occur in the broader area (Athens, Greece). To this end we used molecular techniques. Most surprisingly, we identified the focal population as Podarcis vaucheri, a species far away from its natural range. Molecular results suggest an Iberian origin of this population. To the best of our knowledge, this is the first report of P. vaucheri outside its original range. The new population should be attributed to human-mediated introduction. The future interaction of this introduced species with native lizards, many of which are endemic to Greece, is of critical importance.
Content may be subject to copyright.
Amphibia-Reptilia (2018) DOI:10.1163/15685381-18000002
Podarcis vaucheri (Sauria: Lacertidae) far away from home:
a new invasive species in Greece
Loukia Spilani1,2, Ilias Strachinis3, Andreas Lampropoulos4,PavlosTsigas
5, Nikos Poulakakis1,2,
Panayiotis Pafilis4,
Abstract. In this study we aimed to clarify the identity of a wall lizard population that deviates phenotypically from the other
Podarcis lizards that occur in the broader area (Athens, Greece). To this end we used molecular techniques. Most surprisingly,
we identified the focal population as Podarcis vaucheri, a species far away from its natural range. Molecular results suggest
an Iberian origin of this population. To the best of our knowledge, this is the first report of P. vaucheri outside its original
range. The new population should be attributed to human-mediated introduction. The future interaction of this introduced
species with native lizards, many of which are endemic to Greece, is of critical importance.
Keywords: alien species, human introduction, mitochondrial DNA, wall lizards.
With 86 species, Greece hosts one of the richest
herpetofaunas in Europe. Thirteen of the species
are endemic, whereas for 13 more, Greece hosts
the only European populations (Pafilis, 2010).
Though reptilian and amphibian species have
been moved around Mediterranean Basin for
long time, this trend has been considerably ac-
celerated nowadays as a consequence of human
activities (on the other hand the more exten-
sive contemporary research efforts might also
account for the new records). As such, many
species have widened their traditional range,
colonizing new locations within the country
(Hill and Mayer, 2004; Troidl and Troidl, 2008;
Belasen, Li and Foufopoulos, 2012; Spaneli and
1 - Natural History Museum of Crete, School of Sciences
and Engineering, University of Crete, Knossos Avenue,
Irakleio 71409, Greece
2 - Department of Biology, School of Sciences and Engi-
neering, University of Crete, Vassilika Vouton, Irakleio
70013, Greece
3 - Section of Genetics, Development and Molecular Bi-
ology, Department of Biology, Aristotle University of
Thessaloniki, 541 24 Thessaloniki, Greece
4 - Section of Zoology and Marine Biology, Department
of Biology, National and Kapodistrian University of
Athens, 157 84 Panepistimioupolis, Ilisia, Athens,
5 - 32, N. Plastira str., Agioi Anargyroi, Athens, Greece
Corresponding author; e-mail:
Lymberakis, 2014; Itescu et al., 2016; Kornil-
ios and Thanou, 2016; Mizerakis and Strachi-
nis, 2017). Interestingly, the Athens metropoli-
tan area (capital city of Greece) is on the front-
line of this trend and new lizard populations
of either native or exotic origin have been re-
ported from there (Adamopoulou, 2015; Hed-
man et al., 2017; Karameta and Pafilis, 2017;
Strachinis and Pafilis, 2018).
Podarcis vaucheri has a wide range that
includes southern Spain, central and north-
ern Morocco, northern Algeria and northern
Tunisia, occurring in a variety of habitats such
as Mediterranean type vegetation, rocky areas,
pastureland, rural gardens and urban areas from
sea level up to 3100 m (Mateo et al., 2009).
To the best of our knowledge, this species has
never been reported outside its range. In a sur-
vey conducted at the western outskirts of Athens
(suburbs Agioi Anargyroi and Aigaleo), we en-
countered common wall lizards (Podarcis mu-
ralis) verifying previous reports (Karameta and
Pafilis, 2017), but at Agioi Anargyroi we found
numerous lizards that did not resembled P. m u-
ralis. These individuals had a general mor-
phological appearance closer to Podarcis tau-
ricus: dorsal coloration in males bright green
with many black spots, head, hind legs and tail
brown, whereas flanks had dark and brownish
©Koninklijke Brill NV, Leiden, 2018. DOI:10.1163/15685381-18000002
2Short Notes
spots and blotches in a brown-yellowish back-
ground. Females had less or no green coloration
dorsally, while both sexes had whitish bellies
(Valakos et al., 2008). Nonetheless, the lizards
lacked the typical serrated collar of P. tauri-
We captured 17 individuals from Agioi Anar-
gyroi that were deposited in the Herpetologi-
cal Collection of the Natural History Museum
of Crete (NHMC), University of Crete (voucher
numbers NHMC:,
604, and Our aim was to
identify the collected, unknown lizards. To
this end we first focused on pholidosis. How-
ever pholidotic characters did not indicate any
Greek Podarcis species. Therefore, we em-
ployed molecular techniques to unravel the
identity of the focal lizards.
Total genomic DNA was extracted from the
17 specimens with ambiguous assignment from
the wider region of Attica using a standard
ammonium acetate protocol (Bruford, Hanotte
and Burke, 1998). The mitochondrial gene
(mtDNA) encoding the cytochrome b (cyt b)
(430 bp) was amplified through PCR using
primers GLUDG and CB2 (Palumbi, 1996) and
conditions described in Psonis et al. (2017).
Single stranded sequencing of the PCR prod-
ucts was performed using the Big-Dye Termina-
tor v.3.1 Cycle Sequencing kit®on an ABI3730
automated sequencer following the manufac-
turer’s protocol and using the PCR primers. Se-
quences were edited using CodonCode Aligner
v.3.7.1 (CodonCode Corporation®) and the au-
thenticity and homology to the targeted locus
was evaluated with a BLAST search in the
NCBI genetic database (http://blast.ncbi.nlm. The above search revealed
that 10 of the amplified cyt b sequences had high
similarity with other available cyt b sequences
of Podarcis vaucheri in GenBank (E-value <
10167), while the remaining seven sequences
had high similarity with P. muralis (E-value <
5×10167). In both cases, the similarity dropped
dramatically when compared to other Podarcis
To confirm the above results, a phylogenetic
tree was built using a dataset with all repre-
sentatives of the autochthonous Balkan species
(Poulakakis et al., 2003, 2005a, 2005b; Pso-
nis et al., 2017) as well as sequences from all
main P. vaucheri lineages and its sister species
P. hispanicus (Carranza et al., 2004; Busack et
al., 2005; Pinho et al., 2006; Kaliontzopoulou
et al., 2011). Two sequences of Lacerta ag-
ilis were used as outgroup. Sequence align-
ment was performed using the ClustalW im-
plemented in MEGA v.6 (Tamura et al., 2013)
and the nucleotide substitution model selection
test was carried out using PartitionFinder (PF)
v.2.1 (Guindon et al., 2010; Lanfear et al., 2012,
2016). The dataset was partitioned as specified
by PF, with the following parameters: linked
branch length; MrBayes models; BIC model se-
lection; greedy search algorithm; each codon as
a data block.
Phylogenetic reconstruction was conducted
using Bayesian Inference (BI) and the analy-
sis was performed in MrBayes v.3.2.6 (Ron-
quist et al., 2012), with four runs and eight
chains for each run. Each chain run for 107gen-
erations sampling every 103generations. Sev-
eral MCMC convergence diagnostics were used
to check for convergence and stationarity fol-
lowing the manual’s instructions. The first 25%
trees were discarded as burn-in, as a measure
to sample from the stationary distribution and
avoid the possibility of including random, sub-
optimal trees. A majority rule consensus tree
was then produced from the posterior distribu-
tion of trees, and the posterior probabilities were
calculated as the percentage of samples recover-
ing any particular clade. Posterior probabilities
0.95 indicate statistically significant support
(Huelsenbeck and Ronquist, 2001).
In total, 423 base pairs (bp) of cyt bse-
quences were obtained from all examined spec-
imens. The alignment contained 156 variable
and 144 parsimony informative sites (160 and
151, respectively when the outgroup was also
included). Uncorrected pairwise genetic dis-
tances (p-distances) varied from 0 to 18.6%
Short Notes 3
Tab l e 1. Genetic p-distances (%) among the main clades/lineages for cyt b.
1. P. vaucheri (Greece)
2. P. muralis 15.8
3. P. cretensis 15.413.3
4. P. erhardii 15.412.812.4
5. P. gaigeae 15.815.214.614.8
6. P. hispanicus 12.413.415.314.914.3
7. P. levendis 12.713.48.912.115.016.4
8. P. melisellensis 16.211.714.611.
9. P. milensis 17.013.413.612.19.414.014.09.4
10. P. peloponnesiacus 13.813.56.711.214.716.16.714.313.6
11. P. tauricus 16.213.713.312.910.816.212.810.610.513.5
12. P. vaucheri (Morocco) 4.913.714.915.015.511.114.815.315.914.715.3
13. P. vaucheri (Spain) 1.315.915.715.815.412.513.316.016.714.116.24.7
when the outgroup was not included. The mean
distance between the 10 P. vaucheri specimens
from Agioi Anargyroi and the ones from Spain,
Algeria, Tunisia, Morocco varied from 1.3%
to 12.4% with the spanish lineage being the
closest. Furthermore, the distances among the
specimens in question and the Balkan Podar-
cis species varied between 12.4% and 17% (ta-
ble 1).
The best-fit partitioning scheme and the nu-
cleotide substitution model selected by PF was
K80 +γfor the 1st codon position, HKY +I
for the 2nd codon position and GTR +γfor
the third codon position. In the MrBayes anal-
ysis (arithmetic mean ln L =2749.53), the
MCMC convergence diagnostics did not pro-
vide any clues of non-convergence and indi-
cated stationarity. Considering the P. vaucheri
specimens from Attiki, they form a highly sup-
ported monophyletic group [posterior probabil-
ity (p.p.) =0.99], which, in turn, form another
highly monophyletic group (p.p. =0.98) when
coupled with the P. vaucheri and P. hispanicus
sequences retrieved from GenBank. Addition-
ally, the Greek P. vaucheri lineage seems to be
more closely related to the Spanish lineage as
they form a very well supported clade (p.p. =1)
(fig. 1).
Podarcis vaucheri is a highly diverse species
(Pinho, Ferrand and Harris, 2006; Lima et al.,
2009) that has been raised to specific level from
the Podarcis hispanicus species complex (Oliv-
ero et al., 2000; Busack, Lawson and Arjo,
2005; Arnold et al., 2007). Though it was con-
sidered to be a North African species, new re-
search provided evidence that P. vaucheri in-
vaded Africa from the Iberian Peninsula from
where it originates (Kaliontzopoulou et al.,
2011). To the best of our knowledge, until now
P. vaucheri has never been recorded outside its
original range. Thus, our finding was quite un-
expected, particularly due to the remoteness:
distance between the Athens population and
the easternmost native Spanish population in
Almeria (Rivera, Simón and Arribas, 2009; Fer-
nández Guiberteau and González de la Vega,
2012) is some 2315 km. Apparently, the estab-
lishment of the new population should be at-
tributed to human transportation. Certain Po-
darcis species (e.g. P. muralis,P. pityusensis,
P. siculus) expand their distribution thanks to
direct or indirect anthropogenic means of dis-
persal: railways, pet trade, cargo, merchant or
touristic vessels, timber trade, plant trade, build-
ing materials (Valdeón et al., 2010; Rivera et
al., 2011; Hodgkins, Davis and Foster, 2012;
Silva-Rocha et al., 2014). In at least one case, P.
vaucheri has been reported to form a new popu-
lation as a consequence of human-mediated in-
troduction (Renoult et al., 2010). The thriving
Athenian population (we counted over 60 indi-
viduals of different age classes) might have fol-
lowed a similar way.
4Short Notes
Figure 1. Bayesian Inference tree based on cyt b sequences. The posterior probabilities (>0.95) are given near the branches.
No values means low statistical support.
During the last years several exotic species
invaded the country (Adamopoulou and Lega-
kis, 2016). The proven negative consequences
that some of them (e.g. P. siculus,Lithobates
catesbeianus) may induce to native species
(Kiesecker and Blaustein, 1998; Downes and
Bauwens, 2002), underscore the need for high
alert. We do not know yet whether P. vaucheri
represents a threat for the seven endemic lacer-
tids or the other lizards of Greece. A continuous
monitoring protocol will provide valuable infor-
mation on the potential of P. vaucheri to estab-
lish new populations, occupy new habitats and
outcompete other lizards.
Adamopoulou, C. (2015): First record of Podarcis siculus
(Rafinesque-Schmaltz, 1810) from Greece. Herpetozoa
27: 187-188.
Adamopoulou, C., Legakis, A. (2016): First account on
the occurrence of selected invasive alien vertebrates in
Greece. BioInvasions Rec. 5: 189-196.
Arnold, E.N., Arribas, O., Carranza, S. (2007): Systematics
of the Palaearctic and Oriental lizard tribe Lacertini
(Squamata: Lacertidae: Lacertinae), with descriptions of
eight new genera. Zootaxa 1430: 1-86.
Belasen, A., Li, B., Foufopoulos, J. (2012): Chalcides ocel-
latus (ocellated skink). Greece: Cyclades, Naxos Island.
Herpetol. Rev. 43: 102.
Bruford, M.W., Hanotte, O., Burke, T. (1998): Multi and
single locus DNA fingerprinting. In: Molecular Genetic
Analysis of Populations: a Practical Approach, p. 225-
269. Hoelzel, A.R., Ed., IRL Press.
Short Notes 5
Busack, S.D., Lawson, R., Arjo, W.M. (2005): Mitochon-
drial DNA, allozymes, morphology and historical bio-
geography in the Podarcis vaucheri (Lacertidae) species
complex. Amphibia-Reptilia 26: 239-256.
Downes, S., Bauwens, D. (2002): An experimental demon-
stration of direct behavioural interference in two
Mediterranean lacertid lizard species. Anim. Behav. 63:
Fernández Guiberteau, F., González de la Vega, J.P. (2012):
Confirmación de la presencia de población extralimi-
tal de lagartija andaluza Podarcis vaucheri (Boulenger,
1905) en Almería (sureste de la Península Ibérica). Bull.
Soc. Cat. Herp. 20: 47-53.
Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M.,
Hordijk, W., Gascuel, O. (2010): New algorithms and
methods to estimate maximum-likelihood phylogenies:
assessing the performance of PhyML 3.0. Syst. Biol. 59:
Hedman, H., Kapsalas, G., Karameta, E., Psonis, N.,
Poulakakiks, N., Foufopoulos, J., Pafilis, P. (2017):
A new locality record and range extension for Podarcis
peloponnesiacus (Bibron and Bory, 1833) (Sauria: Lac-
ertidae) from Athens, Greece. Herpetozoa 29: 190-193.
Hill, J., Mayer, W. (2004): First record of the wall lizard Po-
darcis muralis (Laurenti, 1768), from the Ionian Island
of Corfu. Herpetozoa 17: 94-96.
Hodgkins, J., Davis, C., Foster, J. (2012): Successful rapid
response to an accidental introduction of non-native
lizards Podarcis siculus in Buckinghamshire, UK. Con-
serv. Evid. 9: 63-66.
Huelsenbeck, J.P., Ronquist, F. (2001): MRBAYES:
Bayesian inference of phylogenetic trees. Bioinformat-
ics 17: 754-755.
Itescu, Y., Slavenko, A., Schwarz, R., Meiri, S., Pafilis,
P. (2016): A new island record for Chalcides ocellatus
(Forskål, 1775) from Kythnos, Greece. Herpetozoa 29:
Kaliontzopoulou, A., Pinho, C., Harris, D.J., Carretero,
M.A. (2011): When cryptic diversity blurs the picture: a
cautionary tail from Iberian and North African Podarcis
wall lizards. Biol. J. Linn. Soc. 103: 779-800.
Karameta, E., Pafilis, P. (2017): The first record of Podarcis
muralis (LAURENT I, 1768) from Athens, Greece. Her-
petozoa 30: 87-88.
Kiesecker, J.M., Blaustein, A.R. (1998): Effects of intro-
duced bullfrogs and smallmouth bass on microhabitat
use, growth, and survival of native red-legged frogs
(Rana aurora). Cons. Biol. 12: 776-787.
Kornilios, P., Thanou, E. (2016): Two additions to the
herpetofauna of Kasos (Aegean Sea, Greece) and the
role of human-mediated dispersals. Herpetol. Rev. 47:
Lanfear, R., Calcott, B., Ho, S.Y.W., Guindon, S. (2012):
PartitionFinder: combined selection of partitioning
schemes and substitution models for phylogenetic anal-
yses. Mol. Biol. Evol. 29: 1695-1701.
Lima, A., Pinho, C., Larbes, S., Carretero, M.A., Brito,
J.C., Harris, D.J. (2009): Relationships of Podarcis wall
lizards from Algeria based on mtDNA data. Amphibia-
Reptilia 30: 483-492.
Mateo, J.A., Cheylan, M., Nouira, M.S., Joger, U., Sá-
Sousa, P., Pérez Mellado, V., Martinez Solano, I. (2009):
Podarcis vaucheri. The IUCN Red List of Threatened
Species 2009: e.T61556A86439309.
Mizerakis, V., Strachinis, I. (2017): New record of Tarentola
mauritanica (Squamata: Phyllodactylidae) from Lesvos
island, Greece. Herpetol. Notes 10: 157-159.
Oliverio, M., Bologna, M.A., Mariottini, P. (2000): Molec-
ular biogeography of the Mediterranean lizards Podar-
cis Wagler, 1830 and Teira Gray, 1838 (Reptilia, Lacer-
tidae). J. Biogeogr. 27: 1403-1420.
Pafilis, P. (2010): A brief history of Greek herpetology.
Bonn Zool. Bull. 57: 329-345.
Palumbi, S.R. (1996): Nucleic acids II: the polymerase
chain reaction. In: Molecular Systematics, p. 205-248.
Hillis, D.M., Moritz, C., Mable, B.K., Eds, Sinauer,
Pinho, C., Ferrand, N., Harris, D.J. (2006): Reexamination
of the Iberian and North African Podarcis (Squamata:
Lacertidae) phylogeny based on increased mitochondrial
DNA sequencing. Mol. Phyl. Evol. 38: 266-273.
Psonis, N., Antoniou, A., Kukushkin, O., Jablonski, D.,
Petrov, B., Crnobrnja-Isailovi´
c, J., Sotiropoulos, K.,
Gherghel, I., Lymberakis, P., Poulakakis, N. (2017):
Hidden diversity in the Podarcis tauricus (Sauria, Lac-
ertidae) species subgroup in the light of multilocus phy-
logeny and species delimitation. Molecular phylogenet-
ics and evolution 106: 6-17.
Renoult, J.P., Geniez, P., Beddek, M., Crochet, P.-A. (2010):
An isolated population of Podarcis vaucheri (Sauria:
Lacertidae) in south-eastern Spain: genetic data suggest
human-mediated range expansion. Amphibia-Reptilia
31: 287-293.
Rivera, X., Simón, J.G., Arribas, O. (2009): Troballa d’una
població extralimital de sargantana de Vaucher Podarcis
vaucheri (Boulenger, 1905) a Almeria (sud-est de la
península Ibèrica). Bull. Soc. Cat. Herp. 18: 37-38.
Rivera, X., Arribas, O., Carranza, S., Maluquer-Margalef, J.
(2011): An introduction of Podarcis sicula in Catalonia
(NE Iberian Peninsula) on imported olive trees. Bull.
Soc. Cat. Herp. 19: 79-85.
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L.,
Darling, A., Hohna, S., Larget, B., Liu, L., Suchard,
M.A., Huelsenbeck, J.P. (2012): MrBayes 3.2: effi-
cient Bayesian phylogenetic inference and model choice
across a large model space. Syst. Biol. 61: 539-542.
Silva-Rocha, I., Salvi, D., Harris, D.J., Freitas, S., Davis,
C., Foster, J., Deichsel, G., Adamopoulou, C., Carretero,
M.A. (2014): Molecular assessment of Podarcis sicula
populations in Britain, Greece and Turkey reinforces
a multiple-origin invasion pattern in this species. Acta
Herpetol. 9: 253-258.
Spaneli, V., Lymberakis, P. (2014): First record of Stel-
lagama stellio (Linnaeus, 1758) from Crete, Greece.
Herpetol. Notes 7: 367-369.
Strachinis, I., Pafilis, P. (2017): First record of Tarentola
mauritanica (Linnaeus, 1758) from Athens, Greece.
Herpetozoa, under review.
Tamura, K., Stecher, G., Peterson, D., Filipski, A., Kumar,
S. (2013): MEGA6: molecular evolutionary genetics
analysis version 6.0. Mol. Biol. Evol. 30: 2725-2729.
6Short Notes
Troidl, A., Troidl, S. (2008): Fotoexkursion zu den
Hardunen auf der Kykladeninsel Tinos (Griechenland).
Iguana Rundschreiben 20: 4-10.
Valakos, E.D., Pafilis, P., Sotiropoulos, K., Lymberakis, P.,
Maragou, P., Foufopoulos, J. (2008): The Amphibians
and Reptiles of Greece. Chimaira, Frankfurt am Main,
Valdeón, A., Perera, A., Costa, S., Sampaio, F., Carretero,
M.A. (2010): Evidencia de una introducción de Podarcis
sicula desde Italia a España asociada a una importación
de olivos (Olea europaea). Bol. Asoc.Herpetol. Esp. 21:
Submitted: January 11, 2018. Final revision received:
April 20, 2018. Accepted: May 9, 2018.
Associate Editor: Sylvain Dubey.
... The wall lizard, Podarcis vaucheri, has a wide native distribution range comprising southern Spain, central and northern Morocco, northern Algeria and northern Tunisia (Mateo et al. 2009). Up till now, records of this wall lizard species outside its native range include Athens (continental Greece) (Spilani et al. 2018) and Asilah (Morocco) (Busack et al. 2005), apart from Alborán. However, a population of P. vaucheri clustering with North African individuals was once found in southeastern Iberia, resulting from human-mediated introductions (Renoult et al. 2010). ...
... Apart from the populations discovered in Greece (Spilani et al. 2018) and Morocco (Busack et al. 2005), Alborán now represents the third locality where P. vaucheri is found outside its native geographic range. Surprisingly, both the Greek and Moroccan wall lizard specimens belong also to the Southern Spain clade, hence, like the individuals from this study. ...
Full-text available
Alborán is a small volcanic island located in the center of the Alborán Sea, W Mediterranean, 85km from the Iberian Peninsula (Europe) and 55.5km from the NW African coast line. Despite its small size (605x265m), four distinct exotic lizard species have already been reported there, one of them presumably extinct (Saurodactylus mauritanicus). Throughout history, the island has maintained an intermittent human presence, being now permanently occupied by a military detachment connected to the Iberian Peninsula mainly by means of cargo shipments. Hence, this communication between the island and the continent has fuelled human-mediated species introductions. Populations of the geckos Tarentola mauritanica and Hemidactylus turcicus and the most recently reported wall lizard Podarcis vaucheri still remain in the island. Here, we used molecular tools to assess species identity and putative invasion pathways. As expected, results confirm the initial species assignement and indicate southern Iberia as the source area. However, surprisingly, two clades of T. mauritanica were found, the European clade commonly introduced elsewhere, and the Iberian clade, reported here for the first time outside its native distribution. Moreover, the reported southern Iberian lineage of P. vaucheri in Alborán, represents the third case of introduction of this species. This amazing concentration of alien species in such a reduced territory highlights how prone small islands are to be sucessfully invaded, and that these should not be neglected in invasion biology. The effects of these alien reptile species over native taxa are yet unknown, although an impact over several endemic invertebrates cannot be ruled out. As such, regular monitoring should be implemented to prevent they act as hubs for future new introductions.
... The southern Balkans host 10 native Podarcis species, five of which are endemic to Greek islands and the Peloponnese: P. cretensis (Wettstein, 1952) on the island of Crete, P. gaigeae (Werner, 1930) on the Skyros island group, P. levendis Lymberakis et al., 2008, on Pori and Lagouvardos islets, P. milensis (Bedriaga, 1882) on the Milos island group and P. peloponnesiacus on the Peloponnese (Lymberakis & Poulakakis, 2010;Pafilis, 2010;Speybroeck et al., 2020;Uetz et al., 2020); additionally, two alien Podarcis species (P. vaucheri and P. siculus) have also been found in Greece (Adamopoulou, 2015;Spilani et al., 2018). The species found in the Balkans (excluding the alien ones) are divided into three groups (Psonis et al., 2021). ...
The Peloponnese wall lizard, Podarcis peloponnesiacus, is endemic to the Peloponnese. Although the phylogeny and species diversity of the Balkan species of Podarcis have been extensively studied, the intraspecific relationships of P. peloponnesiacus are not yet well defined. The aim of this study was to investigate the intraspecific diversity in this species and clarify its taxonomic status by analyzing eight gene fragments (two mitochondrial and six nuclear) and several morphological traits, typically used for systematic inferences within the genus Podarcis. Together with ecological niche modeling, we provided an integrative evaluation of the differentiation between lineages. The combination of several phylogenetic, species delimitation, and chronophylogenetic analyses revealed the existence of two highly supported and divergent clades with a distinct geographical and parapatric distribution and high niche overlap. The differentiation of the two clades dates back to the Pleistocene and is probably correlated with the paleogeography of the Peloponnese. These clades also differed in several phenotypic traits, which, however, exhibit extensive overlap and are not fully diagnostic. The combination of the above results allowed us to identify the two revealed clades as distinct species. We investigated the intraspecific phylogenetic relationships and clarified the taxonomic status of Podarcis peloponnesiacus (endemic species to the Peloponnese, Greece) by analyzing eight gene fragments (two mitochondrial and six nuclear) and several morphological traits, together with ecological niche modeling. We consistently uncovered two, highly supported, clades within P. peloponnesiacus with distinct geographic distribution, which have diverged in Pleistocene. The combination of our results allowed us to identify the two revealed clades as distinct species: P. peloponnesiacus and P. thais.
... Genetic tools have been instrumental in demonstrating the well-known ancient introductions of North African species of amphibians (e.g., frogs of the genus Discoglossus: Francesca, Roberta and Giuseppe, 2006; tree frogs of the genus Hyla: Recuero et al., 2007;Dufresnes and Alard, 2020) or reptiles (snakes of the genera Macroprotodon or Hemorrhois: Carranza et al., 2004;Carranza, Arnold and Pleguezuelos, 2006; lizards of the genus Teira: Silva-Rocha et al., 2016) in Europe. Genetic data have also helped to identify cryptic introduction of individuals of various species of reptiles from North Africa on islands or mainland across Europe: Podarcis vaucheri from Spain in Greece (Spilani et al., 2018) or from Northern Morocco in Spain (Renoult et al., 2010), Tarentola mauritanica on Corfu (Mačát et al., 2014) or Northern African individuals of the Mediterranean pond turtle Mauremys leprosa in southern France and Spain (Palacios et al., 2015). ...
We report the discovery of a population of the exotic North African Water Frog Pelophylax saharicus around the Etang de Berre, on the Mediterranean coast of France, about 25 km north-west of Marseille. The animals had been originally identified as P. perezi or P. kl. grafi by a combination of acoustic and morphological characters and their true identity was not revealed until three samples from one locality were included in a large-scale genomic work dedicated to the genus Pelophylax. Mitochondrial barcoding of the samples from other areas around Etang de Berre did not detect any native P. perezi or P. kl. grafi and confirmed that P. saharicus has spread to several localities, does reproduce and has been present since 2011 at least. We suggest that dedicated field work is needed as soon as possible to i) map the extant of P. saharicus’s distribution around the Etang de Berre, ii) establish if populations of the native P. perezi – kl. grafi system still persist around the Etang de Berre or not and iii) check if P. saharicus has spread to neighboring areas or not. Depending on the answers to these three questions, local conservationists will need to evaluate the feasibility and relevance of any action to control the spread of this new invasive species and attempt to eradicate it.
... This could be of high priority if we take into account the fact that two exotic species of Podarcis (P. vaucheri, and P. siculus) have invaded the southern Balkans (Adamopoulou, 2015;Spilani et al., 2018). The proven negative consequences that some of them (e.g., P. siculus) may induce to native species (Downes and Bauwens, 2002) underscore the need for high alert. ...
Wall lizards of the genus Podarcis (Sauria, Lacertidae) are the predominant reptile group in southern Europe, including 24 recognized species. Mitochondrial DNA data have shown that, with the exception of P. muralis, the Podarcis species distributed in the Balkan peninsula form a species group that is further sub-divided into two subgroups: the one of “P. tauricus” consisting of P. tauricus, P. milensis, P. gaigeae, and P. melisellensis, and the other of “P. erhardii” comprising P. erhardii, P. levendis, P. cretensis, and P. peloponnesiacus. In an attempt to explore the Balkan Podarcis phylogenomic relationships, assess the levels of genetic structure and to re-evaluate the number of extant species, we employed phylogenomic and admixture approaches on ddRADseq (double digested Restriction site Associated DNA sequencing) genomic data. With this efficient Next Generation Sequencing approach, we were able to obtain a large number of genomic loci randomly distributed throughout the genome and use them to resolve the previously obscure phylogenetic relationships among the different Podarcis species distributed in the Balkans. The obtained phylogenomic relationships support the monophyly of both aforementioned subgroups and revealed several divergent lineages within each subgroup, stressing the need for taxonomic re-evaluation of Podarcis’ species in Balkans. The phylogenomic trees and the species delimitation analyses confirmed all recently recognized species (P. levendis, P. cretensis, and P. ionicus) and showed the presence of at least two more species, one in P. erhardii and the other in P. peloponnesiacus.
... Two invasive species that were recently recorded in the Athens metropolitan area, the Italian wall lizard (Podarcis siculus) (Adamopoulou 2015) and the Andalusian wall lizard (Podarcis vaucheri) (Spilani et al. 2018) have seemingly not dispersed much beyond the locations of the original sightings. The same goes for other native species with different origin that were found inside the Athenian urban districts like the common wall gecko (Tarentola mauritanica) , the Peloponnese wall lizard (Podarcis peloponnesiacus) (Hedman et al. 2017) and the Greek Algyroides (Algyroides moreoticus) (Deimezis-Tsikoutas et al. 2020). ...
Full-text available
Although the insular distribution of the rich herpetofauna of Greece has been studied more extensively, the mainland one is still underexplored. The region of Attica in central Greece represents one of these “black spots” in the mainland. Thus, in 2019 we surveyed for the first time the herpetofauna of Attica in a systematic way. We collected 794 field records of 31 species (4 amphibians, 27 reptiles); Testudo graeca and Trachemys elegans were documented for the first time for Attica.
... In recent years, new sightings of reptiles and amphibians from Greece are frequently published, both from the mainland (e.g., Adamopoulou, 2015;Andriopoulos and Pafilis, 2016;Hedman et al., 2017;Karameta and Pafilis, 2017;Spilani et al., 2018;Strachinis and Pafilis, 2018;Deimezis Tsikoutas et al., 2019) as well as the islands (e.g., Belasen et al., 2012;Spaneli and Lymberakis, 2014;Itescu et al., 2016;Bogaerts et al., 2018;Christopoulos, 2018). This is an impressive fact, since the Aegean Sea is considered well studied (Pafilis, 2010; Sfenthourakis et al., 2018). ...
In this note we report for the first time the occurrence of the Roughtail Rock Agama (Stellagama stellio) from Kaprathos Island, Greece. Greece represents the westernmost range limit of the animal and hosts its only European populations. Although Karpathos is a relatively well-studied island in terms of its herpetofauna, this is the second remarkable herpetological finding within the past year, following the rediscovery of the Grass Snake more than 50 years after it was first described from the island. This report for S. stellio expands the range of the species and it is added to a long series of new records in the Greek herpetofauna that occurred during the last few years.
... Our finding represents the easternmost population of A. nigropunctatus, as the documented eastern limit of its natural range is located in the prefecture of Fthiotida (Andriopoulos and Pafilis, 2016), more than 180 km west of Athens. Such reports are of particular interest, as they add to our knowledge on animal mobility, in the constantly changing biogeography of the Mediterranean basin (Spilani et al., 2018). ...
A small but growing population of the Dalmatian Algyroides (Algyroides nigropunctatus) was found in a suburb of Athens (Greece), a long way from the species’ known range. This apparently introduced population increases the number of new lizard species documented in the Athens metropolitan area during the past few years.
... Thus, the populations of Aetoloakarnania could have originated from colonizers from either/both the Peloponnese and Ionian islands, most likely through human mediated transportation. However, introductions from other countries cannot be excluded as recent introductions of lizard species have been observed from the city of Athens, namely, Podarcis siculus (Rafinesque-Schmaltz, 1810) (Adamopoulou, 2015) and Podarcis vaucheri (Boulenger, 1905) (Spilani et al., 2018). What is for certain, is that the Moorish Gecko has overcome geographical barriers to actively disperse onto the Greek mainland which is no surprise given it is a known successful and adaptive colonizer. ...
Tarentola mauritanica, also known as Moorish Gecko, is a trans-Mediterranean gecko with a large distribution, ranging from Israel and North Africa, to southern Europe, as well as on many large Mediterranean islands. In Greece, the Moorish Gecko is distributed across several islands, and the only mainland occurrences reported are from the Peloponnese (in the west and north) and very recently, from Athens. Here we present three new records of the Moorish Gecko from localities in the Aetoloakarnania prefecture (Western Greece), and briefly discuss potential scenarios of introduction and expansion of this species on the Greek mainland.
... The negative association patterns found in the co-occurrence analysis suggest that the introduction of alien congeneric species could have a very negative effect on microinsular endemic lizards. Accidental translocation of several species of Podarcis has been documented in the Mediterranean (Silva-Rocha et al., 2014;Spilani et al., 2018). The results of this study indicate that in the event of the introduction of alien lizards to small islets, their rapid control and eradication is advisable. ...
Full-text available
Mediterranean islands have complex reptile assemblages, but little is known about the factors that determine their organization. In this study, the structure of assemblages of Squamata was evaluated based on their species richness and two measures of phylogenetic diversity (variability and clustering). I evaluated the composition of the assemblages comparing distinct biogeographic subregions within the Mediterranean: Adriatic, Aegean, Balearic, Corsica–Sardinia, Crete, Gulf of Gabés, Ionian Sea, Ligurian Sea, Malta, Sicily, and Tyrrhenian Sea. The effect of island environments and geographical isolation on the diversity metrics was assessed using generalized linear models. The analyses indicated that species richness was mostly influenced by island area and geographical isolation. Assemblages on smaller islands were poorer in species and phylogenetically dispersed, possibly as an effect of interspecific competition. The species composition of the assemblages was determined by similar environmental drivers within the biogeographic subregions, including island area, island elevation, geographical isolation, and aridity. In several subregions, significant patterns of phylogenetic attraction were found in species co-occurrences, caused by the limits imposed by the island size on large predatory species.
... Human translocations with North African origin are proposed for the Iberian populations of the chameleon Chamaeleo chamaeleon (Paulo et al., 2002) and the treefrog Hyla meridionalis (Recuero et al., 2007). Other dispersals, on the other hand, have their origins in the Iberian Peninsula, for example, the translocation of the lizard Podarcis vaucheri in Greece (Spilani et al., 2018). In the Western Mediterranean context, the humanmediated introductions of herpetofauna in the Balearic Islands stand out, where the successive arrivals of different human groups led to the extirpation of native species in most of the main islands (Alytes muletensis and Podarcis lilfordi), besides the introduction of new species from other Mediterranean regions (e.g. ...
Human movements in the regions surrounding the Mediterranean Sea have caused a great impact in the composition of terrestrial fauna due to the introductions of several allochthonous species, intentionally or not. Reptiles are one of the groups where this anthropic impact is most evident, owing to the extensive intra-Mediterranean dispersals of recent chronologies. Chalcides ocellatus is a widespread skink with a natural distribution that covers almost the entire Mediterranean Basin. Two hypotheses have been proposed to explain its origin: natural dispersions and human translocations. Previous molecular data suggest the occurrence of a recent dispersal phenomenon across the Mediterranean Sea. In this study we present the first record of this species in the Iberian Peninsula, in Serra del Molar (South-east Spain). We combined molecular analyses and archaeological records to study the origin of this population. The molecular results indicate that the population is phylogenetically closely related to specimens from north-eastern Egypt and southern Red Sea. We suggest that the species arrived at the Iberian Peninsula most likely through human-mediated dispersal by using the trade routes. Between the Iron to Middle Ages, even now, the region surrounding Serra del Molar has been the destination of human groups and commercial goods of Egyptian origins, in which Chalcides ocellatus could have arrived as stowaways. The regional geomorphological evolution would have restricted its expansion out of Serra del Molar. These findings provide new data about the impact of human movements on faunal introductions and present new information relating to mechanisms of long-distance translocations.
Full-text available
In this document we present the discovery of a new established population of the Moorish Gecko (Tarentola mauritanica) in Athens at the regions of Sepolia and Aigaleo. This is the first time the species is reported in mainland Greece, outside its known distribution in the W Peloponnese. As a strongly synanthropic species, the Moorish Gecko must have been introduced in Athens through human activities.
Full-text available
This is the first attempt to outline the occurrence of selected invasive alien vertebrates in Greece, since up to now, there are no " official " or scientific reports except from sporadic sightings and anecdotal stories. Records on the occurrence of: Lithobates catesbeianus, Trachemys scripta (T. s. elegans and T. s. scripta), Neovison vison, Myocastor coypus, Nyctereutes procyonoides and Ondatra zibethicus were requested through a pan-Hellenic survey. According to the results, the coypu (Myocastor coypus) appears to be the most widely distributed of all species, having conquered practically all wetlands of Western and Central Greece with populations exceeding, in most cases, 20 individuals each. On the contrary, there is merely one unconfirmed record of the raccoon dog (Nyctereutes procyonoides) while there were no sightings at all for the muskrat (Ondatra zibethicus). The American mink (Neovison vison) was recorded in the northwestern part of Greece. Regarding the alien herpetofauna, the bullfrog (Lithobates catesbeianus) seems to be still confined in Crete where it was originally introduced, while the red-eared slider (Trachemys scripta) appears mostly in Crete, Attiki prefecture, and a few more places, showing most probably an underestimated distribution.