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New records and an updated list of the herpetofauna of Kastellorizo and the adjacent islet Psomi (Dodecanese, SE Greece)

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Based on several recent herpetological surveys on the Kastellorizo archipelago in the Eastern Mediterranean, we present a review of the herpetofauna of the area. Active searching and visual encounter surveys on Kastellorizo confirmed the presence of two species of amphibians and thirteen species of reptiles, and in conjunction with the available literature data, brought the terrestrial herpetofauna of the island to a total of 17 species. In addition, during a one-day herpetological survey on Psomi islet, we encountered individuals of Anatololacerta sp. and Mediodactylus cf. danilewskii. Based on the analysis of the mitochondrial gene cytb, the recently discovered lizard population from the settlement of Megisti was identified as Anatololacerta pelasgiana (Mertens, 1959), while that of Psomi islet as Anatololacerta budaki (Eiselt and Schmidtler, 1986), a new addition to the herpetofauna of Greece. We reported on the extant herpetofauna of Kastellorizo and Psomi islet providing a species list that includes four new species records, namely A. pelasgiana, A. budaki, Platyceps najadum (Eichwald, 1831), and Telescopus fallax (Fleischmann, 1831). Our results expand our knowledge on the biogeography of the region and the insular distribution of the species. We also discuss the possible introduction ways of the non-native lizard A. pelasgiana, recent taxonomic revisions, and possible erroneous records.
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Introduction
Kastellorizo (also known as Megisti) is a small
Greek island belonging to the Dodecanese and the
Rhodes regional unit with an approximate area of 12
km2, located 125 km east of Rhodes Island and about
2 km south of the nearest Turkish coast (Anatolian
coastal town of Kaş). The archipelago composed of the
islands Kastellorizo, Ro, Strongili, and numerous islets
(Agrielia, Psomi, Psoradia, and others) represents the
easternmost point of Greece (Fig. 1). It is a limestone
island composed of karstic limestone with a highest
elevation of 273 m above sea level (Vigla) and steep,
rocky coastlines. The island lacks springs and is mostly
covered with phrygana vegetation, which surrounds the
few cultivated areas (Paysant, 2005; Fig. 2A). There
is only one settlement on the island, namely Megisti,
with a population of 492 residents as per the 2011
census (Hellenic Statistical Authority, 2011). Psomi is
a small islet in the archipelago of Kastellorizo with an
approximate area of 0.003 km2, located 1 km northeast
of Megisti and around 3 km of the Turkish coast (Fig.
3A). It is composed of karstic limestone and is covered
mostly with chasmophytic vegetation.
Most of the previous herpetological surveys on the
island (e.g., Kiortsis et al., 1974; Tzannetatou-Polymeni,
1986; Veith et al., 2001) focused mainly on the study of
Lyciasalamandra luschani basoglui (Baran and Atatür,
1980). Literature regarding the island’s herpetofauna
is restricted (Valakos and Papapanagiotou, 1985;
Paysant, 1999, 2005). Early surveys (Valakos and
Papapanagiotou, 1985) have led to the listing of nine
reptile and one amphibian species on Kastellorizo.
Later, Van Wingerde (1986) reported the presence
of Montivipera xanthina (Gray, 1849) on the island,
Herpetology Notes, volume 11: 1009-1019 (2018) (published online on 27 November 2018)
New records and an updated list of the herpetofauna
of Kastellorizo and the adjacent islet Psomi
(Dodecanese, SE Greece)
Konstantinos Kalaentzis1,*, Ilias Strachinis1, Philippos Katsiyiannis2, Peter Oefinger3, and Christos Kazilas1
1 School of Biology, Aristotle University of Thessaloniki
(AUTH), 54124 Thessaloniki, Greece.
2 Present address: Kato Platanovrysi, 25008 Achaia, Greece.
3 Present address: Philipp-Reis-Straße 9, 40215 Düsseldorf,
Germany.
* Corresponding author. E-mail: konstakal95@gmail.com
Abstract. Based on several recent herpetological surveys on the Kastellorizo archipelago in the Eastern Mediterranean, we
present a review of the herpetofauna of the area. Active searching and visual encounter surveys on Kastellorizo confirmed the
presence of two species of amphibians and thirteen species of reptiles, and in conjunction with the available literature data,
brought the terrestrial herpetofauna of the island to a total of 17 species. In addition, during a one-day herpetological survey
on Psomi islet, we encountered individuals of Anatololacerta sp. and Mediodactylus cf. danilewskii. Based on the analysis
of the mitochondrial gene cytb, the recently discovered lizard population from the settlement of Megisti was identified as
Anatololacerta pelasgiana (Mertens, 1959), while that of Psomi islet as Anatololacerta budaki (Eiselt and Schmidtler, 1986), a
new addition to the herpetofauna of Greece. We reported on the extant herpetofauna of Kastellorizo and Psomi islet providing
a species list that includes four new species records, namely A. pelasgiana, A. budaki, Platyceps najadum (Eichwald, 1831),
and Telescopus fallax (Fleischmann, 1831). Our results expand our knowledge on the biogeography of the region and the
insular distribution of the species. We also discuss the possible introduction ways of the non-native lizard A. pelasgiana, recent
taxonomic revisions, and possible erroneous records.
Keywords. Megisti, herpetological survey, species identification, Anatololacerta, cytb, Bayesian inference
Konstantinos Kalaentzis et al.
1010
however with some doubts. More recent herpetological
surveys (Paysant, 1999, 2005) revealed the occurrence
of four additional species on the island, namely
Bufotes viridis (Laurenti, 1768), Heremites auratus
(Linnaeus, 1758), Xerotyphlops vermicularis (Merrem,
1820), and Dolichophis jugularis (Linnaeus, 1758).
Regarding the herpetofauna of Psomi islet, individuals
of Anatololacerta sp. and Mediodactylus cf. danilewskii
have been recorded by the Natural History Museum of
Crete in 1996 (Petros Lymberakis, pers. comm.).
Here, we present an updated list of the herpetofauna
of Kastellorizo and Psomi, based on six herpetological
surveys conducted over a period of nine years (2009-
2017) and available literature data, including four new
species records, namely Anatololacerta pelasgiana
(Mertens, 1959), Anatololacerta budaki (Eiselt &
Schmidtler, 1986), Platyceps najadum (Eichwald,
1831), and Telescopus fallax (Fleischmann, 1831).
Materials and Methods
Systematic surveys were conducted during the past
years, namely in September 2009 (3 d), March 2011 (3
d), April 2015 (4 d), March 2017 (3 d), May 2017 (5 d),
and August 2017 (9 d). Psomi islet was surveyed on 3
May 2017 (1d).
Observations were made by visual encounter surveys
and active searching during day and night, in a variety of
habitats and climatic conditions, as well as by identifying
road-killed specimens. Individuals of all the observed
amphibian and reptile species were photographed using
digital SLR cameras and in some cases specimens were
collected. In specific, specimens of Anatololacerta sp.
and Mediodactylus cf. danilewskii from Psomi islet,
as well as one specimen of Anatololacerta sp. from
the settlement of Megisti, were collected. Voucher
specimens and photographic vouchers were deposited
at the Herpetological Collection of the Natural History
Museum of Crete, Greece (NHMC). All of the
specimens were collected under the Greek Presidential
Decree 67/81.
Muscle tissue of the Anatololacerta specimens
from Kastellorizo and Psomi islet was obtained.
Whole genome DNA was extracted following the
method described by Hillis et al. (1996), based on
cetyltrimethylammonium bromide (CTAB) reagent.
Polymerase Chain Reactions (PCR) were performed for
the amplification of a partial sequence (420bp) of the
mitochondrial protein-coding cytochrome b gene (cytb),
using the forward primer GLUDG-L (Kocher et al.,
1989) and the reverse primer CB2-H (Palumbi, 1996).
PCR consisted of an initial activation step of 10 min at
95°C, 40 cycles of steps a) 94°C for 1 min, b) 48,5°C
for 1 min and c) 72°C for 1 min, and then 10 min at
72°C. Subsequently, PCR products were purified using
the NucleoSpin Gel and PCR Clean-up kit (Macherey-
Nagel), and Sanger sequencing was performed by
Genewiz Inc. Chromatograms were checked using
Geneious v10.2.3 (Biomatters Ldt.). Cytb sequences
of the Anatololacerta species group and two outgroup
species, i.e. Hellenolacerta graeca (Bedriaga, 1886and
Parvilacerta parva (Boulenger, 1887) were retrieved
from the nucleotide database of NCBI (GenBank).
Τhesequences werealigned usingClustalW (7homson
et al., 1994) and phylogenetic trees were constructed
using Maximum Likelihood (ML) and BayesiaQ
Inference (BI) methods. The model selection was based
on the Bayesian Information Criterion (BIC; Schwarz,
1978) using jModelTest v2.1.10 (Posada, 2008; Darriba
et al., 2012). ML analysis was performed in MEGA
v7.0.26 (Kumar et al., 2016), and the confidence of each
node of the best ML tree was examined with a bootstrap
analysis of 1000 pseudo-replicates (Felsenstein, 1985).
BI analysis was performed in MrBayes v3.2.6 (Ronquist
et al., 2012) for 106 generations, and the current tree
was saved to file every 500 generations. The first
25% of the trees were discarded by default, to avoid
the possibility of including random sub-optimal trees.
The obtained sequences of Anatololacerta specimens
from Kastellorizo and Psomi islet were deposited in
Figure 1. (A) Map showing the location of Kastellorizo
Island, Greece. (B) A more focused view of the island group
comprising Kastellorizo, Ro, and Strongili islands. The
dotted line represents the national border between Greece and
Turkey.
the GenBank nucleotide database with the accession
numbers MH041496 and MH041497 respectively..
Results and Discussion
Kastellorizo
In total, two species of amphibians and thirteen species
of terrestrial reptiles were observed and photographed
by the authors on Kastellorizo Island during six field
expeditions between September 2009 and August 2017
(Table 1).
Lyciasalamandra luschani (Steindachner, 1891),
Luschan’s Salamander (Fig. 4A).—The presence
of this species on the island has already been reported
several times (e.g., Kiortsis et al., 1974; Valakos and
Papapanagiotou, 1985; Tzannetatou-Polymeni, 1986;
Paysant, 2005). Luschan’s Salamander is a very
common species that was found almost everywhere on
the island. Large numbers of this species were observed
in a variety of habitats during nocturnal surveys and
even during daytime it was found under stones on
humid substrate. Most observations were made at night.
New records and an updated list of the herpetofauna of Kastellorizo and Psomi 1011
Figure 2. Typical landscape covered with phryganic vegetation on the central plateau of Kastellorizo (A) and the three new
records from the island (B-D). (B) Pelasgian Rock Lizard (Anatololacerta pelasgiana) in the settlement of Megisti in May 2017.
(C) A dead specimen of Dahl’s Whip Snake (Platyceps najadum). (D) A dead specimen of European Cat Snake (Telescopus
fallax). Photos by Ilias Strachinis (A, B), and Philippos Katsiyiannis (C, D).
However, during a rainy day in March 2017 numerous
salamanders were found, at daytime, on the ground or
climbing on walls. In a single night in May 2017 (1717°C,
humidity 88%, and calm wind), over 300 individuals,
including many salamanders in amplexus position, were
recorded on a 200 m distance track on a dirt road inside
the settlement. Individuals of L. luschani were found
around the settlement of Megisti, where the higher
numbers were recorded, but also on the island’s plateau
and Acheres plain.
Bufotes viridis (Laurenti, 1768), European Green
Toad (Fig. 4B).—The European Green Toad was first
mentioned on the island of Kastellorizo by Paysant
(2005). During our expeditions, we observed few
adult individuals of B. viridis around the settlement
of Megisti, road-killed specimens on Acheres plain,
and several tadpoles inside an old metal barrel on the
island’s plateau. Since their reproduction cycle is
completed in water, toads of this species are dependent
on the presence of water bodies. The main sites of water
sources on the island consist of artificial reservoirs set
to collect rainwater, temporary flooded fields on the
island’s plateau, and shallow limestone depressions.
Apart from the scarcity of reproduction sites on the
Konstantinos Kalaentzis et al.
1012
Figure 3. (A) Psomi islet and the adjacent Turkish coast as seen from Megisti. (B) Mediterranean Thin-toed Gecko (Mediodactylus
cf. danilewskii) collected from Psomi. (C) The specimen of Budak’s Rock Lizard (Anatololacerta budaki) collected from Psomi.
Photos by Peter Oefinger (A), Konstantinos Kalaentzis (B), and ,lias Strachinis(C).
island, individuals of the species also face a threat while
using the artificial water tanks as egg laying sites, since
their vertical walls can prevent them from escaping
(Paysant, 2005).
Stellagama stellio (Linnaeus, 1758), Starred
Agama.—This species is reported from Kastellorizo
several times (e.g., Valakos and Papapanagiotou, 1985;
Paysant, 1999, 2005). It seems to be quite common
on the island, since we encountered numerous Starred
Agamas in almost every site we visited.
Hemidactylus turcicus (Linnaeus, 1758), Turkish
Gecko.—The Turkish Gecko is a very common
species in the Mediterranean region, especially around
human settlements and dwellings. It is known from the
settlement of Megisti (Valakos and Papapanagiotou,
1985). We observed the species in great numbers in
Megisti during nighttime surveys, but also in several
localities on the island’s plateau at daytime, during our
expeditions in September 2009, as well as in May and
August 2017.
Mediodactylus danilewskii (Strauch, 1887),
Mediterranean Thin-toed Gecko.—Valakos and
Papapanagiotou (1985) mentioned the presence of
the species, then known as Tenuidactylus kotschyi
(Steindachner, 1870) from Agios Ioannis and Acheres.
Kotsakiozi et al. (2018) based on a multilocus
phylogenetic analysis proposed the recognition of
five taxa as full species inside the M. kotschyi species
complex, namely M. kotschyi, M. orientalis (Štěpánek,
1937),, M. danilewskii (Strauch, 1887),, M. bartoni
(Štěpánek, 1934), and M. oertzeni (Boettger, 1888). In
the aforementioned study, Mediodactylus populations
from the island of Kastellorizo were classified as M.
danilewskii. We observed individuals of the species
on the island during our expeditions in March 2011,
as well as in May and August 2017. According to our
New records and an updated list of the herpetofauna of Kastellorizo and Psomi 1013
Table 1. List of amphibian and reptile species of Kastellorizo and Psomi islet (Dodecanese, Greece) encountered during the
herpetological surveys conducted by the authors (indicated with X). Numbers indicate the references that mention previous
records, including 1—Valakos and Papapanagiotou (1985), 2a—Paysant (1999), and 2b—Paysant (2005). Montivipera xanthina,
mentioned in Van Wingerde (1986) from Kastellorizo, is not included because we consider it unreliable (see Results and
Discussion).
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observations and experience, it seems that this species
is less common on Kastellorizo compared to other
Dodecanese Islands (e.g., Karpathos, Symi).
Anatololacerta pelasgiana (Mertens, 1959),
Pelasgian Rock Lizard.—PO spotted one individual
of Anatololacerta sp. at Agia Triada Monastery, near
Megisti, in March 2011, without knowing at that time if
this was a single individual or if there was an established
population on the island. Later, in March 2017, the same
author encountered three individuals of the species in
the settlement. In May 2017, we observed at least five
of these lizards in Megisti (Fig. 6A), photographed them
and collected one specimen (NHMC 80.3.62.63; Fig.
2B). The rarity of these lizards on Kastellorizo, the fact
that they have never been reported on the island before,
and their proximity to the island’s commercial port,
possibly indicate that their arrival is recent, probably
due to accidental introduction.
Our phylogenetic analysis agreed with this hypothesis,
since the specimen from Megisti was grouped in the
same subclade with three specimens of A. pelasgiana
(Mertens, 1959) from Rhodes; this result was supporteG
with significantly high posterior probabilities and
bootstrap values (Fig. 5). Consequently, we consider
the Kastellorizo population introduced, possibly from
Rhodes. It could have arrived on the island by human-
mediated transports, e.g. ferry boats, trade of plants,
building material etc. It is worth mentioning that
Rhodes is the primary supply and transportation hub
of Kastellorizo. In addition, a similar recent finding of
A. pelasgiana on another Dodecanese island (Kassos)
supports the scenario of anthropogenic transportation
(Kornilios and Thanou, 2016).
We follow the definition of A. pelasgiana proposed by
Bellati et al. (2015), who elevated A. oertzeni pelasgiana
to full species status. This is a new species record for
Kastellorizo, as it has never been recorded on the island
by previous herpetofaunal surveys (e.g., Valakos and
Papapanagiotou, 1985; Paysant, 2005).
Ablepharus anatolicus Schmidtler, 1997, Lycian
Snake-eyed Skink (Fig. 4C).—The species is already
mentioned on the island as Ablepharus kitaibelii (Bibron
and Bory St-Vincent, 1833), by previous investigators
(Valakos and Papapanagiotou, 1985; Paysant, 2005).
Poulakakis et al. (2005) showed that A. kitaibelii is not
a monophyletic taxon, and that the population of the
Kastellorizo island group is particularly distinct and
deserves species status. A more recent phylogenetic
analysis based on morphological features and genetic
markers (mtDNA, nuDNA) proposed the raise of
Ablepharus populations from Kastellorizo island
group and southwest Turkey to species level, namely
A. anatolicus (Skourtanioti et al., 2016). The large
ear opening is a common morphological trait the
populations of Ablepharus from the island group of
Kastellorizo and southern Anatolia share (Schmidtler,
1997; Poulakakis et al., 2005; Fig. 4C). We observed
this species in several localities, during our visits in
March 2011, April 2015, as well as in March, May, and
August 2017, mostly near abandoned dwellings around
the town of Megisti. The species seems to occur in high
densities in the area.
Heremites auratus (Linnaeus, 1758), Levant
Skink.—This species is known from the island by
Paysant (2005), as Euprepes auratus. In 2016, Karin et
al., using molecular data, resurrected genus Heremites
and grouped the Middle East species of Trachylepis
within Heremites. In May 2017, we encountered six
individuals in different localities throughout the island.
More specifically, four individuals were found around
the settlement and near the Acheres plain, while the
other two were spotted basking by low stone walls on
the island’s plateau.
Ophiomorus kardesi (Kornilios et al., 2018),
Anatolian Limbless Skink (Fig. 4D).— A very recent
genetic analysis (Kornilios et al., 2018) based on a
multilocus DNA dataset, comprising of mitochondrial
and nuclear genetic markers, described the Ophiomorus
populations of Kastellorizo and SW Turkey as a new
species, namely O. kardesi. The Anatolian Limbless
Skink is mentioned by Valakos and Papapanagiotou
(1985), who collected one specimen from the island’s
plateau. Several individuals of this species were found
under stones, at daytime, during our surveys in March
and May 2017.
Blanus strauchi (Bedriaga, 1884), Anatolian
Worm Lizard.—Populations of Kastellorizo and SW
Anatolia belong to the subspecies B. strauchi bedriagae
Boulenger, 1884, according to Alexander (1966) and
Sindaco et al. (2014). It has been mentioned from
previous expeditions (e.g., Valakos and Papapanagiotou,
1985; Paysant, 2005). We also encountered it in March
and May 2017. During our visit in May 2017 one of
the four Anatolian Worm Lizards observed, presented
piebald morphotype (Kazilas et al., 2018).
Xerotyphlops vermicularis (Merrem, 1820),
Eurasian Blind Snake (Fig. 4E).—The species is
Konstantinos Kalaentzis et al.
1014
already mentioned from Kastellorizo (Paysant, 1999,
2005). We encountered several individuals in March and
May 2017 in different localities around the settlement
and on the island’s plateau. Individuals of this species,
due to their fossorial nature, were mostly found under
stones, at daytime. However, we also spotted and
photographed two active individuals on the surface at
night near the settlement in May 2017.
Dolichophis jugularis (Linnaeus, 1758), Large
Whip Snake.—The species was first mentioned on the
island by Paysant (1999). Both alive individuals and
dead specimens were observed during our expeditions in
March 2011 and May 2017, mostly near the settlement
of Megisti.
Eirenis modestus (Martin, 1838), Ring-headed
Dwarf Snake (Fig. 4F).— Its presence on the
island is known from previous expeditions (Valakos
and Papapanagiotou, 1985; Paysant, 2005). It was
encountered in April 2015, as well as in March and May
2017, during daytime searches under stones. We found
specimens throughout the island including the vicinity
of the settlement, the Acheres plain, and on the island’s
central plateau. It seems to be a very common species
on Kastellorizo. Two of the individuals we encountered,
one juvenile in April 2015 and one adult in May 2017,
had unusual dark colourations resembling melanistic
morphotypes (Kalaentzis et al., 2018).
Hemorrhois nummifer (Reuss, 1834), Coin-marked
Snake.—The Coin-marked Snake is mentioned from
literature: Valakos and Papapanagiotou (1985) collected
two specimens from cultivated fields on the island’s
plateau and Paysant (2005) found one individual near
the monastery of Saint George. We did not find it in any
of our expeditions.
Platyceps najadum (Eichwald, 1831), Dahl’s
Whip Snake.—During a visit in September 2009, KK
observed and collected a specimen of P. najadum in
the settlement of Megisti (NHMC 80.3.132.13), a new
species record for the area. An additional dead, adult
specimen was found in April 2015 (Fig. 2C). However,
we failed to encounter the species in any of our further
expeditions.
Telescopus fallax (Fleischmann, 1831), European
Cat Snake.—A road killed specimen of T. fallax was
photographed near the settlement of Megisti, in August
2017 (Fig. 2D; NHMC 80.3.38.130). The identification
was based on the shape of head, body, and tail, pholidosis
characteristics of the head, the dark transverse bars on
the back of the body, and on the characteristic dark
collar, which extended to the middle of the head. This
is the first time this species is reported from the island.
However, since there are no other records of the species
on Kastellorizo, one cannot exclude an isolated event
of accidental translocation and further investigation is
required in order to confirm an established population
on the island.
Malpolon insignitus (Geoffroy Saint-Hilaire,
1827), Eastern Montpellier Snake.—There is a
single report of this species from Kastellorizo by
Valakos and Papapanagiotou (1985), who collected one
specimen from Acheres plain. However, no individual
of M. insignitus was encountered on the island during
any of our surveys. We suggest that supplementary
investigation is needed in order to confirm the presence
of the species on the island.
Montivipera xanthina (Gray, 1849), Ottoman
Viper.—Van Wingerde (1986) suggested the possible
occurrence of the Ottoman Viper on the island, based
on a dubious observation of a fast moving, stout bodied
snake with a black and off-white pattern. This doubtful
record was later mentioned by Dimitropoulos (1986)
and Dimitropoulos et al. (1988). The species was not
observed by the authors, nor by other herpetologists
(Valakos and Papapanagiotou, 1985; Paysant, 1999,
2005). For all the above and due to lack of any evidence
on the presence of the species on the island (specimen or
photographic vouchers), we propose that M. xanthina is
not included in the island’s herpetofauna list, until solid
proof of its occurrence is presented.
Psomi
During a one-day expedition on Psomi islet (northeast
of Megisti; Fig. 6A) in May 2017, two reptile species
were observed and voucher specimens were collected
(Table 1).
Mediodactylus cf. danilewskii (Strauch, 1887),
Mediterranean Thin-toed Gecko.—The presence of
this species on Psomi islet was already detected by a
group of scientists from the NHMC (Petros Lymberakis,
pers. comm.). We photographed and collected two
specimens on the islet (NHMC 80.3.85.1937-8; Fig.
3B) during our visit in May 2017. Population density on
Psomi islet seems to be higher than that on Kastellorizo.
The population of this species on Psomi islet should
possibly be attributed to M. danilewskii but further
research is needed in order to confirm this.
New records and an updated list of the herpetofauna of Kastellorizo and Psomi 1015
Konstantinos Kalaentzis et al.
1016
Figure 4. Two amphibian and four reptile species of the total 15 observed on the island of Kastellorizo, during six herpetological
surveys. (A) A female Luschan’s Salamander (Lyciasalamandra luschani basoglui). (B) A female European Green Toad (Bufotes
viridis). (C) An adult Lycian Snake-eyed Skink (Ablepharus anatolicus). The main morphological trait of this species, a large ear
opening, is denoted. (D) Anatolian Limbless Skink (Ophiomorus kardesi). (E) Eurasian Blind Snake (Xerotyphlops vermicularis).
(F) An adult Ring-headed Dwarf Snake with rows of dark blotches in the first half of the body (Eirenis modestus) found on the
island’s plateau. Photos by Christos Kazilas (A, F), Konstantinos Kalaentzis (B, C, D), and Ilias Strachinis (E).
Anatololacerta budaki (Eiselt & Schmidtler, 1986),
Budak’s Rock Lizard.—A population of Anatololacerta
sp. lizards was already known from an expedition of
NHMC (Petros Lymberakis, pers. comm.). During our
visit on the islet, we spotted numerous individuals and
managed to collect one specimen (NHMC 80.3.62.64;
Fig. 3C). We noticed that the colour of all the observed
individuals was brownish and unusually dark compared
to those from Kastellorizo and Rhodes.
Our phylogenetic analysis showed a clear distinction
between two main clades that was supported with
significantly high values in both trees (Fig. 5). The first
clade was comprised of A. pelasgiana specimens from
Turkey and Rhodes, while the specimen from Psomi
islet was grouped together in the second clade with
specimens from the adjacent Turkish mainland, which
belong to the former taxon Anatololacerta oertzeni
finikensis (Eiselt and Schmidtler, 1986).
Bellati et al. (2015) investigated the relationships
between the four species in the Anatololacerta species
complex; of these Anatololacerta budaki (Eiselt and
Schmidtler, 1986) includes the previous taxa A. oertzeni
budaki (Eiselt and Schmidtler, 1986) and A. oertzeni
finikensis that both occur in SW Anatolia. Although the
taxon of A. budaki is still a subject of discussion with
regard to its geographical distribution and the taxonomic
status of A. oertzeni ibrahimi (Eiselt and Schmidtler,
1986), we follow the definition of A. budaki proposed by
Bellati et al. (2015). Previous to this finding, the species
was unknown from Greece (Valakos et al., 2008). This
is the first record of the species on Greek territory.
Conclusions
The current study reveals the presence of two amphibian
and 13 reptile species on the island of Kastellorizo
and, in conjunction with the available literature data
from previous herpetofaunal expeditions, brings its
terrestrial herpetofauna to a total of 17 species (Table
1). Regarding the species mentioned in the literature
but not encountered in any of our surveys, namely H.
nummifer, M. insignitus, and M. xanthina, only the
latter is considered doubtful and thus is not included
in the list of the island’s herpetofauna (see Results and
Discussion). In addition, our list includes three new
species records for the island, of which the presence
of P. najadum and T. fallax is of no surprise, since the
New records and an updated list of the herpetofauna of Kastellorizo and Psomi 1017
Figure 5. Bayesian Inference tree based on a 383 bp fragment of cytb focusing on the identification of the two Anatololacerta
specimens collected from the Kastellorizo archipelago. The posterior probabilities (BI) and bootstrap support (ML) are given near
the branches.
surrounding area falls within the distribution range of
both species. On the other hand, the Anatololacerta
lizards from Kastellorizo seem to be introduced, since
our phylogenetic analysis shows a clear clustering of the
obtained sequence into a single clade with A. pelasgiana
specimens from Rhodes (Fig. 5).
On Psomi islet, we found two reptile species, including
Anatololacerta lizards that were identified as A. budaki
and represent the first record of this species on Greek
territory. Since no herpetofaunal surveys have been
conducted on the other islands (Ro, Strongili) or islets
of the archipelago, this should be the focus of future
surveys in the area.
The richness of amphibians and reptiles on the island
of Kastellorizo is high considering its very small size.
This can be explained mainly by the short distance of
the island from the neighboring Anatolia and its recent
separation from it. Furthermore, human activities may
account for the introduction of non-native species (e.g.
ferry boats, trade of building material, vehicles etc.),
though the short distance (only 2 km off the Turkish
coast) does not exclude the possibility of natural
transportations (“rafts”). Transmarine dispersal is not
unusual and has been recently demonstrated for Crete
(Kyriazi et al., 2013).
Acknowledgements. We thank Petros Lymberakis (NHMC)
for providing useful information, but also for his assistance and
advice. We are grateful to Alexandros Triantafyllidis (AUTH) for
allowing us to complete the present genetic analysis in his lab.
We also thank Kostas Chousalas for his company and suggestions
during our stay on the island.
References
Alexander, A.A. (1966): Taxonomy and variation of Blanus
strauchi (Amphisbaenia, Reptilia). Copeia 1966: 205–224.
Bellati, A., Carranza, S., Garcia-Porta, J., Fasola, M., Sindaco,
R. (2015): Cryptic diversity within the Anatololacerta species
complex (Squamata: Lacertidae) in the Anatolian Peninsula:
Evidence from a multi-locus approach. Molecular Phylogenetics
and Evolution 82: 219–233.
Darriba, D., Taboada, G.L., Doallo, R., Posada, D. (2012):
jModelTest 2: more models, new heuristics and parallel
computing. Nature Methods 9: 772–772.
Dimitropoulos, A. (1986): Some notes on the colour and
pattern of the Greek snake fauna in relation to geographic
distribution. Biologia Gallo-Hellenica 12: 463–471.
Dimitropoulos, A., Giannatos, G., Mertzanis, G. (1988): An
additional record of Ottoman Viper, Vipera xanthina (Gray,
1849) from mainland Greece. Herptile, London 13: 99–103.
Felsenstein, J. (1985): Confidence limits on phylogenies: an
approach using the bootstrap. Evolution 39: 783–791.
Hellenic Statistical Authority (2011): 2011 Population-Housing
Census. Available at: http://www.statistics.gr. Accessed on 15
March 2018.
Hillis, D.M., Moritz, C., Mable, B.K. (1996): Molecular
Systematics. Sunderland, USA, Sinauer Associates.
Kalaentzis, K., Kazilas, C., Strachinis, I. (2018): Two cases of
melanism in Ring-headed Dwarf Snake Eirenis modestus
(Martin, 1838) (Serpentes: Colubridae) from Kastellorizo,
Greece. Herpetology Notes 11: 175–178.
Karin, B.R., Metallinou, M., Weinell, J.L., Jackman, T.R.,
Bauer, A.M. (2016): Resolving the higher-order phylogenetic
relationships of the circumtropical Mabuya group (Squamata:
Scincidae): An out-of-Asia diversification. Molecular
Phylogenetics and Evolution 102: 220–232.
Kazilas, C., Kalaentzis, K., Strachinis, I. (2018): A case of
piebaldism in the Anatolian Worm Lizard, Blanus strauchi
(Bedriaga, 1884) (Squamata: Blanidae), from Kastellorizo,
Greece. Herpetology Notes 11: 527–529.
Kiortsis, V., Koutsaftikis, A., Matsakis, J. (1974): Les
Mertensielle de Grèce I. Nouvelle forme de Mertensiella
Konstantinos Kalaentzis et al.
1018
Figure 6. (A) Partial topographic map of the Kastellorizo
archipelago showing the location of the islets surrounding
Megisti. Sampling localities of Anatololacerta budaki,
represented by red circle (Psomi islet), and Anatololacerta
pelasgiana, represented by blue circle (Megisti), are shown.
Additional localities where individuals of A. pelasgiana were
also encountered during our expeditions are indicated with
smaller, transparent, blue circles. The location of the island’s
harbour and airport are labelled. (B) Map of SE Aegean and
SW Anatolia, displaying the sampling locations of every
specimen that was used in our phylogenetic analysis, including
those acquired from the nucleotide database of GenBank. A.
pelasgiana is represented by blue circles, while A. budaki by
red.
luschani (Steindachner) dans l’Archipel Hellénique (Note
préliminaire). Biologia Gallo-Hellenica 5: 355–358.
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo,
S., Villablanca, F.X., Wilson, A.C. (1989): Dynamics of
mitochondrial DNA evolution in animals: amplification and
sequencing with conserved primers. Proceedings of the National
Academy of Sciences 86: 6196–6200..
Kornilios, P., Kumlutaş, Y., Lymberakis, P., Ilgaz, Ç. (2018):
Cryptic diversity and molecular systematics of the Aegean
Ophiomorus skinks (Reptilia: Squamata), with the description
of a new species. Journal of Zoological Systematics and
Evolutionary Research 2018: 1–18.
Kornilios, P., Thanou, E. (2016): Two Additions to the Herpetofauna
of Kassos (Aegean Sea, Greece) and the Role of Human-
mediated Dispersals. Herpetological Review 47: 633–635.
Kotsakiozi, P., Jablonski, D., Ilgaz, C., Kumlutaş, Y., Avcı, A.,
Meiri, S., et al. (2018): Multilocus phylogeny and coalescent
species delimitation in Kotschy’s gecko, Mediodactylus kotschyi:
hidden diversity and cryptic species. Molecular Phylogenetics
and Evolution 125: 177–187.
Kumar, S., Stecher, G., Tamura, K. (2016): MEGA7: Molecular
Evolutionary Genetics Analysis version 7.0 for bigger datasets.
Molecular Biology and Evolution 33: 1870–1874.
Kyriazi, P., Kornilios, P., Nagy, Z.T., Poulakakis, N., Kumlutaş,
Y., Ilgaz, Ç., et al. (2013): Comparative phylogeography reveals
distinct colonization patterns of Cretan snakes. Journal of
Biogeography 40: 1143–1155.
Palumbi, S.R. (1996): Nucleic acids II: The Polymerase Chain
Reaction. In: Molecular Systematics, p. 205-248. Hills, D.
& Moritz, C., Mable, B.K., Eds., Sunderland, USA, Sinaeur
Associates.
Paysant, F. (1999): Nouvelles données sur l’herpétofaune de
Castellorizzo (sud-est de l’archipel Egéen, Grèce). Bulletin de
la Société herpétologique de France 91: 5–12.
Paysant, F. (2005): Herpetological notes on the island Kastellorizo
(South-east Aegean, Greece). Herpetozoa 18: 80–83.
Posada, D. (2008): jModelTest: phylogenetic model averaging.
Molecular Biology and Evolution 25: 1253–1256.
Poulakakis, N., Lymberakis, P., Tsigenopoulos, C.S., Magoulas,
A., Mylonas, M. (2005): Phylogenetic relationships and
evolutionary history of snake-eyed skink Ablepharus kitaibelii
(Sauria: Scincidae). Molecular Phylogenetics and Evolution 34:
245–256.
Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling,
A., Höhna, S., et al. (2012): MrBayes 3.2: efficient Bayesian
phylogenetic inference and model choice across a large model
space. Systematic Biology 61: 539–542.
Schmidtler, J.F. (1997): The Ablepharus kitaibelii-group in southern
Anatolia and adjacent territories. Herpetozoa 10: 35–63.
Schwarz, G. (1978): Estimating the dimension of a model. The
Annals of Statistics 6: 461–464.
Sindaco, R., Kornilios, P., Sacchi, R., Lymberakis, P. (2014):
Taxonomic reassessment of Blanus strauchi (Bedriaga, 1884)
(Squamata: Amphisbaenia: Blanidae), with the description of a
new species from south-east Anatolia (Turkey). Zootaxa 3795:
311–326.
Skourtanioti, E., Kapli, P., Ilgaz, Ç., Kumlutaş, Y., Avcı, A.,
Ahmadzadeh, F., et al. (2016): A reinvestigation of phylogeny
and divergence times of the Ablepharus kitaibelii species
complex (Sauria, Scincidae) based on mtDNA and nuDNA
genes. Molecular Phylogenetics and Evolution 103: 199–214.
Thomas, K.R., Thomas, R. (1978): Locomotor activity responses
to photoperiod in four West Indian fossorial Squamates of the
genera Amphisbaena and Typhlops (Reptilia, Lacertilia). Journal
of Herpetology 12: 35–41.
Thomson J.D., Higgins D.G., Gibson T.J. (1994): CLUSTAL
W: improving the sensitivity of progressive multiple sequence
alignment through sequence weighting, position-specific gap
penalties and weight matrix choice. Nucleic Acids Research 22:
4673–4680.
Tzannetatou-Polymeni, R. (1986): Observations comparatives
sur les populations de Mertensiella luschani (Steindachner),
des iles Kastellorizo, Karpathos, Kassos et Saria (Amphibia,
Salamandridae). Biologia Gallo-Hellenica 12: 445–450.
Valakos, E.D., Pafilis, P., Lymberakis, P., Maragou, P.,
Sotiropoulos, K., Foufopoulos, J. (2008): The amphibians and
reptiles of Greece. Frankfurt, Germany,Edition Chimaira.
Valakos, S., Papapanagiotou, D. (1985): The herpetofauna of
Kastellorizo (Megisti) Island (SE Aegean archipelago). British
Herpetological Bulletin 13: 9–12.
Veith, M., Baran, İ., Godmann, O., Kiefer, A., Öz, M., Tunç, M.R.
(2001): A revision of population designation and geographic
distribution of the Lycian salamander Mertensiella luschani
(Steindachner, 1891). Zoology in the Middle East 22: 67–82.
Van Wingerde, J. (1986): The distribution of Vipera xanthina (Gray,
1849) on the East Aegean Islands and in Thrace. Litteratura
Serpentium 6: 131–139.
New records and an updated list of the herpetofauna of Kastellorizo and Psomi 1019
Accepted by Hendrik Müller
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