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

May 2018
Amphibia-Reptilia 39(3):363 – 368

DOI:10.1163/15685381-18000002

Authors:

Loukia Spilani

Institute of Evolutionary Biology

Ilias Strachinis

Aristotle University of Thessaloniki

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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.

Bayesian Inference tree based on cyt b sequences. The posterior probabilities (>0.95) are given near the branches. No values means low statistical support.

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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,

Greece

5 - 32, N. Plastira str., Agioi Anargyroi, Athens, Greece

∗Corresponding author; e-mail: ppaﬁl@biol.uoa.gr

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-

cus.

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: 80.3.183.1-10, 80.3.53.603-

604, and 80.3.53.670-674). 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.

nih.gov/Blast.cgi). 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 <

10−167), while the remaining seven sequences

had high similarity with P. muralis (E-value <

5×10−167). In both cases, the similarity dropped

dramatically when compared to other Podarcis

sequences.

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.

123456789101112

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 16.211.714.611.89.015.013.3

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.

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The lacertid wall lizards of the genus Podarcis (Wagler, 1830) originate from Western Europe and are divided into 24-25 species (Speybroeck et al., 2020). Despite the abundance of phylogenetic studies referring to the genus, the relationships among certain species, as well as the taxonomic status of some genealogical lineages, remain unclear due to great genetic diversity. The common wall lizard Podarcis muralis has a relatively wide distribution and despite the existence of various molecular studies focused on this species, its Greek populations had never been thoroughly sampled until now. To fill in this geographical gap, we sampled the species’ Greek distribution and constructed phylogenies that uncovered the presence of at least five monophyletic lineages that correspond to different geographic regions. Furthermore, species delimitation analyses assigned all lineages to a single species diversifying during the early Pleistocene (c. 1.93 mya). The strong association of the genetic lineages with specific geographical regions coupled with the Pleistocene diversification of the group imply the presence of multiple refugia within Greece and, by extension, the Balkan peninsula, supporting a refugia-within-refugia scenario. Finally, in an effort to clarify the position of these new samples within the larger phylogeny of P. muralis , a larger phylogeny was constructed which indicated that the Greek populations cluster with the central European, Italian and Turkish populations of the species.

Tarentola mauritanica (Squamata: Phyllodactylidae) in Greece: an update on the species' distribution, including new records

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Jun 2023

The Moorish Gecko (Tarentola mauritanica) is a successful colonizer that has been introduced in many European countries. In Greece, the species' presence is documented for at least 150 years now and it has been only recently introduced on the islands of Corfu and Lesvos, and the prefectures of Aetoloakarnania and Attica (Athens city). In this document we unify the current knowledge on the Moorish Gecko's occurrence in Greece, with new, unpublished records, museum archives and citizen science data, in order to generate an updated image of the species' extant distribution. This merge of data fills several distributional gaps and also extends the Moorish Gecko's range in two new areas, namely Fokida prefecture and Kos Island. Concerns about possible threats to native herpetofauna are also discussed.

Evidence of cryptic diversity in Podarcis peloponnesiacus and re‐evaluation of its current taxonomy; insights from genetic, morphological, and ecological data

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Full-text available

Oct 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.

Discovery of a Pelophylax saharicus (Anura, Ranidae) population in Southern France: a new potentially invasive species of water frogs in Europe

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Jul 2021
AMPHIBIA-REPTILIA

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.

The wall lizards of the Balkan peninsula: Tackling questions at the interface of phylogenomics and population genomics

Article

Feb 2021

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.

Guide de biodiversité de la Medjerda

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Full-text available

Feb 2022

Ce livre, conçu par l’association Exploralis dans le cadre du projet Med’Ex (Medjerda Expédition), soutenu par le programme PPI-OSCAN et financé par la fondation MAVA et le Fonds Français pour l’Environnement Mondial, se concentre sur la conservation de l’écosystème complexe de la Medjerda en Tunisie. Depuis 2016, Med’Ex observe la dynamique écologique et la biodiversité de la Medjerda, offrant un équilibre entre rigueur scientifique et vulgarisation pour le grand public. Destiné aux amateurs, ce guide simplifié aide à identifier les espèces tout en les sensibilisant à la richesse de la région. Il évite le jargon technique, se basant sur des références fiables et la liste rouge de l’UICN pour des informations sur la conservation des espèces. Le livre présente les espèces communes, vulnérables, endémiques et menacées, en fournissant des descriptions et des photographies accessibles. Son objectif est d’encourager l’observation respectueuse de la nature plutôt que la collecte de spécimens. En célébrant la biodiversité locale, il appelle à une appréciation accrue face à la perte mondiale de biodiversité. Exploralis remercie tous les contributeurs, y compris les jeunes experts, les institutions et la société civile, pour leur engagement dans ce projet. Un remerciement particulier est adressé à M. Zekher Bouragaoui et M. Mehdi Boujemâa pour leur soutien inestimable

First record of Anatololacerta pelasgiana (Mertens, 1959) in mainland Greece: another new species in Athens

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Full-text available

Dec 2022

Urban habitats receive an increasing number of species due to anthropogenic activities, mainly transportations. Here, we report a new addition to the herpetofauna of Athens (Greece): a small population of the Pelasgian wall lizard (Anatololacerta pelasgiana) was found in a suburb of the Athenian metropolitan area. The species normally occurs in southwestern Anatolia and southeastern Aegean islands and this is the first record in the Greek mainland. Allochthonous species that successfully colonize cities raise new challenges to urban ecology

Rivera, X;,Simón, J.G. & ARRIBAS, O. (2009): Troballa d’una població extralimital de sargantana de Vaucher Podarcis vaucheri (Boulenger, 1905) a Almeria (sudest de la península Ibèrica) Butll. Soc. Cat. Herp., 18 (2009): 37-38. [ISSN: 0212-6648]

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Jan 2009

First record of Tarentola mauritanica (Linnaeus, 1758) from Athens, Greece

Article

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Aug 2018

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.

Podarcis vaucheri

Chapter

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Dec 2009

First record of Podarcis muralis (Laurenti, 1768), from athens, Greece

Article

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Aug 2017

New record of Tarentola mauritanica (Squamata: Phyllodactylidae) from Lesvos island, Greece

Article

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Apr 2017

First account on the occurrence of selected invasive alien vertebrates in Greece

Article

Full-text available

Nov 2016

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.

Two Additions to the Herpetofauna of Kasos (Aegean Sea, Greece) and the Role of Human-mediated Dispersals

Article

Dec 2016

Multilocus and single-locus DNA fingerprinting

Chapter

Mar 1998

Methods enabling the direct study of genetic variation in natural populations have improved considerably. The new edition explores these updated techniques in DNA analysis and provides a revised and refined laboratory guide to investigating variation in DNA molecules.

Single-locus and multilocus DNA fingerprinting

Article