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Mediodactylus kotschyi in the Peloponnese peninsula, Greece: Distribution and habitat

  • Leibniz-Institute of Freshwater Ecology and Inland Fisheries and Freie Universität Berlin

Abstract and Figures

The gecko Mediodactylus kotschyi is considered rare in mainland Greece, yet it is very abundant on the Aegean islands. It has been thought to be saxicolous throughout much of its range. In a recent survey on the Pelopon-nese peninsula, however, we encountered it mainly on trees, and with higher frequency than previously reported. We combined our observations of localities in which we detected this gecko, and places where we failed to detect it, with data about its occurrence from the literature and museum collections. We posited two hypotheses as possible causes for the apparent relative scarcity of M. kotschyi in the Peloponnese: that it is associated with low precipitation and that it has an aversion to limestone rock. We predicted that M. kotschyi would be more likely to be found in arid places and where limestone is not the dominant type of rock, since it has been reported that this substrate is less suitable for this species. Moreover, we predicted that geckos occurring in limestone regions would be found on trees rather than under rocks. Geckos were indeed found mainly in the more arid parts of the Peloponnese, but not exclusively so. We found no evidence of limestone avoidance. We suggest that, because M. kotschyi is better known as being mostly saxi-colous over most of its range, and exclusively so on the Greek islands, in the Peloponnese the search for this species has been restricted to a single habitat type, i.e., under rocks and not on trees. It may thus inhabit more localities in the Peloponnese and be more abundant there than has previously been thought.
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Acta Herpetologica 11(2): 179-187, 2016
ISSN 1827-9635 (print) © Firenze University Press
ISSN 1827-9643 (online)
DOI: 10.13128/Acta_Herpetol-18176
Mediodactylus kotschyi in the Peloponnese peninsula, Greece:
distribution and habitat
R S,*, I-A G, Y I, S J, K S, S
M, P P
1 Department of Zoology, Tel Aviv University, Tel Aviv 6997801, Israel. * Corresponding author. E-mail:
2 Department of Zoology and Marine Biology, School of Biology, University of Athens, Panepistimioupolis, Ilissia, Greece
3 Department of Human and Animal Physiology, School of Biology, University of Athens, Panepistimioupolis, Ilissia, Greece
Submitted on 2016, 14th April; revised on 2016, 03th July; accepted on 2016, 20th August
Editor: Marco Sannolo
Abstract. e gecko Mediodactylus kotschyi is considered rare in mainland Greece, yet it is very abundant on the
Aegean islands. It has been thought to be saxicolous throughout much of its range. In a recent survey on the Pelopon-
nese peninsula, however, we encountered it mainly on trees, and with higher frequency than previously reported. We
combined our observations of localities in which we detected this gecko, and places where we failed to detect it, with
data about its occurrence from the literature and museum collections. We posited two hypotheses as possible causes
for the apparent relative scarcity of M. kotschyi in the Peloponnese: that it is associated with low precipitation and that
it has an aversion to limestone rock. We predicted that M. kotschyi would be more likely to be found in arid places
and where limestone is not the dominant type of rock, since it has been reported that this substrate is less suitable for
this species. Moreover, we predicted that geckos occurring in limestone regions would be found on trees rather than
under rocks. Geckos were indeed found mainly in the more arid parts of the Peloponnese, but not exclusively so. We
found no evidence of limestone avoidance. We suggest that, because M. kotschyi is better known as being mostly saxi-
colous over most of its range, and exclusively so on the Greek islands, in the Peloponnese the search for this species
has been restricted to a single habitat type, i.e., under rocks and not on trees. It may thus inhabit more localities in the
Peloponnese and be more abundant there than has previously been thought.
Keywords. Arboreality, habitat preferences, Mediodactylus kotschyi, Peloponnese, rock type.
e genus Mediodactylus is predominantly Asian, with
only one of 13 species being found in Europe. e Mediter-
ranean thin-toed gecko, Mediodactylus kotschyi (Steindach-
ner, 1870) (Reptilia: Gekkonidae), has a discontinuous dis-
tribution incorporating southern Italy, through to the Bal-
kans and the Crimean peninsula, to Israel and Iran (Arnold
and Ovenden, 2002; Sindaco and Jeremcenko, 2008).
In Greece, M. kotschy is ubiquitous and highly
abundant on the Sporades, Cyclades, and south Aegean
islands as well as around Crete, where its distribution is
well documented (e.g., Wettstein, 1937; Beutler and Gru-
ber, 1977; Beutler, 1981; Chondropoulos, 1986), and its
diversity is high: 13 subspecies are currently recognized
from the Greek islands (Karandinos and Paraschi, 1992;
Kasapidis et al., 2005; Uetz and Hošek, 2016). However,
it is considered to be rare on the Greek mainland and
throughout the Balkans (Stojanov et al., 2011; Tomovic et
al., 2014), and is oen absent from Peloponnese species
checklists (e.g., Werner, 1929; Cyrén, 1935; Bischo and
Bischo, 1980; Henle, 1989; Pèrez-Mellado et al., 1999).
e Peloponnese peninsula nonetheless forms a major
part of M. kotschyi’s distribution on mainland Greece
180 Rachel Schwarz et alii
(Valakos et al., 2008). It has been recorded from sev-
eral locations (e.g., Monemvasia, the terra typica of M. k.
bibroni, Beutler and Gruber, 1977; Sparta and Kalamata,
Stepánek, 1937), and is thought to be widespread in the
western Peloponnese (Valakos et al. 2008). It is nonethe-
less considered rare, exhibiting a low population density
almost everywhere on the Greek mainland (Ajtić, 2014).
M. kotschyi is described as being mainly saxicolous
across much of its range, being found under rocks and
stone piles, on dry stone walls and even on the external
walls of houses and other buildings (Beutler, 1981 and
citations therein; Musters and In den Bosch, 1982; Arnold
and Ovenden, 2002; Ajtic 2014). In Greece it is described
as preferring dry areas with phrygana (=dwarf shrub
steppe) vegetation, although it also inhabits cultivated
areas (Beutler, 1981). Phrygana is common on the Aegean
islands, but is rare on mainland Greece, and this has been
claimed to be the main reason for its rarity on the main-
land (Beutler, 1981). In Israel, in contrast, M. kotschyi is
almost obligatorily arboreal, and its Hebrew name (“םיצע
תיממש” = “tree-gecko”) reects this (Werner, 1993; Bar
and Haimovitch, 2012; and our pers. obs.). In Israel,
southern Turkey and Iraq it can be found on some com-
mon Mediterranean trees such as oak, olive, g, almond
and carob, as well as on introduced species such as Euca-
lyptus (Weber, 1960; Beutler, 1981; and pers. obs.).
Another major factor thought to inuence the distri-
bution of M. kotschyi is that of substrate. Like all mem-
bers of its genus, this species lacks adhesive toe pads
(Gamble et al., 2012). Many pad-less saxicolous species
are associated with rough rock surfaces (Higham, 2015),
such as sandstone (Russell et al., 2007), perhaps because
they are able to attain a secure grip on these types of
rocks compared to smoother ones. Beutler (1981) sug-
gested that in the Cyclades, where M. kotschyi is very
abundant, the ground is comprised mainly of slate, gran-
ite, mica, marble and volcanic rocks. However, the main
rock type on the Greek mainland is limestone, which
according to Beutler is less suitable for M. kotschyi. Con-
sequently, he has claimed that the only mainland region
where M. kotschyi is abundant is in the Taygetos moun-
tain range south of Sparta, where the dominant rock
types are shale and sandstone (Beutler, 1981).
e range of precipitation for Mediterranean veg-
etation is ~250-800 mm, the lower limit of which cor-
responds to phrygana vegetation (Aschmann, 1973),
which covers the Peloponnese peninsula’s coastline from
south to east (Mavromatis, 1978). We hypothesized that,
because M. kotschyi is thought to be strongly associated
with sparse phrygana, it would be more common in the
eastern Peloponnese, where the climate is drier due to
the rain shadow cast by the central mountains (Kotini-
Zabaka, 1983; Bringsoe, 1985). We further hypothesized
that M. kotschyi would be rare in areas where the main
type of rock is limestone (Beutler, 1981), and that in wet
regions, and where limestone predominates, it would be
more likely to occur on trees than among rocks.
To test our predictions we constructed a presence and
absence distribution map of M. kotschyi, incorporating 68 loca-
tions from across the entire Peloponnese. Sixteen of these obser-
vations (eight presences and eight absences, Table 1a, b) are
derived from eldwork we conducted in June and October 2015,
including two locations from which the gecko had been previ-
ously reported (Maragou et al., 2015) as well as six new locali-
ties in both phrygana habitats and tree groves. e remaining 52
locations were sourced from publications, museum records and
localities surveyed independently by KS (Table 2).
We searched for M. kotschyi by turning over rocks and
visually scanning tree trunks during daylight hours (M. kotschyi
is mostly diurnal, active during all but the hottest hours of the
day, Beutler, 1981; Valakos et al., 2008; Baier et al., 2009; and
pers. obs.). In each locality two to four people searched for
geckos for at least 30 minutes. Weather conditions were favour-
able for M. kotschyi activity throughout. If no individuals were
observed in a locality we considered it to be absent, although
we acknowledge that false absences are a possibility.
We recorded the habitat and type of substrate on which
the individuals were found. We recorded GPS coordinates of
all locations surveyed, for both presence and absence of M.
kotschyi, and assembled them on a map using ArcGIS (ESRI,
2011). Most literature-based locations (Table 2) are provided
only as verbal descriptions (usually the name of a town). We
digitized the coordinates of these using Google Maps (2015).
To determine whether a connection exists between the dis-
tribution of M. kotschyi and aridity, we assigned mean annual
precipitation data (from Worldclim, Hijmans et al., 2005) to
sampled localities (presence and absence, Tables 1, 2). We also
recorded rock type for all such locations using geological maps
(Higgins and Higgins, 1996) in order to test for substrate pref-
erences. We performed statistical tests of rock type associations
only for the presence locations for which substrate data were
specied. We used χ2 tests for goodness of t to compare obser-
vations from wet and arid regions, Student’s t tests were applied
to test for a connection between substrate type and precipita-
tion, and to compare climatic conditions at sites with and with-
out geckos. We used Fisher’s exact test to search for a connection
between rock type and preferred substrate. All analyses were car-
ried out using R version 3.0.1 (R Development Core Team, 2013).
During our 2015 survey we encountered geckos
in eight locations but failed to encounter them in the
remaining eight (Table 1). Most records of M. kotschyi
Mediodactylus kotschyi in the Peloponnese peninsula
are from the central and eastern parts of the peninsula
(Fig. 1). e 800 mm isohyet divides the Peloponnese
into roughly equal areas (above 800 mm: 10,134.45 km2;
below 800 mm: 11,113.96 km2), and thus the null expec-
tation would be for 10:11 number of observations from
arid and wet regions. Nevertheless, most records of pres-
ence (77%, 46 out of 60, Table 1 a) are from where annu-
al mean precipitation is lower than 800 mm (χ2 goodness
of t test, χ2 = 14.21, n = 60, P = 0.0002). Fourteen loca-
tions where presence has been recorded are from regions
Table 1. Presence (a) and absence (b) of Mediodactylus kotschyi at sites surveyed during our eld work in June and October 2015 in the
Peloponnese. Data on substrate and rock type, annual precipitation (mm) and GPS coordinates were combined with the data presented in
Table 2 for map construction.
a. presence
Location Habitat searched Substrate Rock type
No. individuals
caught (rocks/
Latitude Longitude
North west of
Neapolis, Malea
Peninsula, Laconia
Sparse phrygana
with carob trees and
eucalypt logs
On carob and olive
trees, and eucalypt logs Limestone 564 (2/7) 36.5631N 23.0040E
Neapolis, Malea
Peninsula, Laconia Olive grove
On olive and carob
trees and on a
building’s wall
Alluvium 548 (2/4) 36.5339N 23.0421E
North west of Neapoli,
Malea Peninsula,
Dense phrygana Under a rock Limestone 585 (1/0) 36.5690N 22.9890E
Gera, near
Monemvasia, Malea
Peninsula, Laconia
Dense phrygana with
carob trees
Under rocks
and on carob trees Limestone 543 (3/2) 36.6867N 23.0368E
North west of
Monemvasia, Malea
Peninsula, Laconia
Eucalypt, almond and
carob grove
On eucalypt, almond
and carob trees Phyillites 572 (0/17) 36.7291N 22.9802E
North east of Geraki,
Phrygana with carob
trees bordering an
olive grove
On carob and olive
trees Alluvium 688 (0/5) 36.999N 22.722E
Kato Vervena, Arcadia Olive grove On olive trees Alluvium 600 (0/5) 37.4396N 22.7370E
Kalogria, south west of
Patras, Achaea
Eucalypt and pine
Under rock piles and
on eucalypt trees Alluvium 766 (4/2) 38.13N 21.37E
b. absence
Location Habitat searched Rock type Annual precipitation
(mm) Latitude Longitude
Lagokili, Mani Peninsula Olive grove (both on trees and
under rocks) Limestone- marble 707 36.6502N 22.4017E
South of Platsa, Mani Peninsula,
Messenia Stone piles Limestone- marble 752 36.800N 22.318E
Kardamyli, Mani Peninsula Eucalypt grove (trees only) Neogene sediments 719 36.891N 22.233E
West of Prosilio, Mani Peninsula Phrygana
(rocks only) Limestone 770 36.9134N 22.2240E
South east of Agii Anargiri,
(rocks only) Neogene sediments 676 37.0160N 22.6360E
South west of Kosmas, Laconia Stream bed (rocks only) Limestone- marble 849 37.0800N 22.7200E
East of Kalogria, Achaea Phrygana (rocks only) Limestone 793 38.1605N 21.3847E
Trochalia, Malea Peninsula,
Laconia Eucalypt grove (trees only) Alluvium 546 36.6535N 23.0241E
182 Rachel Schwarz et alii
Table 2. Locations, substrate and rock type, annual precipitation (mm), sources of data, and estimated GPS coordinates (longitude and lati-
tude in decimal degrees) used for map construction.
Location name Substrate Rock type
Latitude Longitude Source
Lachos, Mani Peninsula Unspecied Limestone-marble 688 36.48N 22.37E Valakos et al. 2008
Kokkala, Mani Peninsula Unspecied Limestone-marble 651 36.52N 22.47E Valakos et al. 2008
Kato Kastania, Malea Peninsula,
Phrygana with
many rocks Limestone 596 36.52N 23.11E Bringsoe 1985
Ano Kastania, Malea Peninsula,
Laconia Unspecied Phyllites 606 36.537N 23.102E Valakos et al. 2008
Lira, Malea Peninsula, Laconia Unspecied Phyllites 637 36.640N 22.964E Valakos et al. 2008
Loutsa, Mani peninsula Unspecied Limestone-marble 711 36.643N 22.474E Valakos et al. 2008
Monemvasia, Malea Peninsula,
Laconia Unspecied Limestone 546 36.69N 23.05E Beutler and Gruber
South of Agios Ioannis, Malea
Peninsula, Laconia Unspecied Limestone 557 36.726N 23.007E Valakos et al. 2008
5 km north of Monemvasia,
Malea Peninsula, Laconia Building Limestone 549 36.73N 23.02E Bringsoe 1985
Methoni, Messenia Unspecied Alluvium 742 36.82N 21.704E Valakos et al. 2008
Kastania, Kariofouni and Driopi
area, Laconia
Area with stone
walls Limestone-marble 762 36.84N 22.35E Bauer 2004; Valakos et
al. 2008
Between Saidona and Kastane,
Mani Peninsula Unspecied Limestone-marble 797 36.87N 22.29E Bringsoe 1985; Valakos
et al. 2008
Lakkos, Mani Peninsula On trees in
forest Neogene sediments 768 36.893N 22.259E Pers. obs. Kostas
Sagonas 2014
Exochori (Taygetos), Mani
Peninsula Unspecied Neogene sediments 768 36.90N 22.26E Werner 1937; Valakos et
al. 2008
Mandina near Kampos, Mani
Peninsula Unspecied Flysch 781 36.93N 22.20E Naturhistorisches
Museum Wien
Kalamata, Messenia Unspecied Alluvium 762 37.04N 22.11E Stepánek 1937; Valakos
et al. 2008
2 km south of Gargalianoi,
On the ground
in open eld Peridotite and serpentinite 813 37.049N 21.634E Pers. obs. Kostas
Sagonas 2014
Mystras, Laconia Unspecied Limestone- marble 776 37.06N 22.37E
Beutler and Gruber
1977; Stepánek 1937;
Valakos et al. 2008
10 km west of Sparta, Laconia Unspecied Limestone-marble 913 37.064N 22.305E Valakos et al. 2008
Pyrgos, Elis Near stone
terraces Neogene sediments 839 37.07N 21.69E Bringsoe 1985
Sparta, Laconia Unspecied Alluvium 712 37.071N 22.430E Valakos et al. 2008
5 km north east of Kosmas,
Eggs under a at
rock Limestone 715 37.12N 22.78E Bringsoe 1985
Rouzaki, Messenia On trees in
forest Neogene sediments 790 37.236N 21.662E Pers. obs. Kostas
Sagonas 2014
Agii Asomatoi, Arkadia Under stones in
maquis Flysch 671 37.332N 22.699E Pers. obs. Kostas
Sagonas 2014
Tegea, Arcadia Unspecied Alluvium 798 37.45N 22.41E Naturhistorisches
Museum Wien
Didima, Corinthia Unspecied Limestone 527 37.461N 23.171E Valakos et al. 2008
Tripoli, Arcadia Unspecied Limestone 807 37.507N 22.371E Valakos et al. 2008
1 km south west of Mainalo,
Under stones in
maquis Limestone 836 37.529N 22.299E Pers. obs. Kostas
Sagonas 2014
Methanon, Malea Peninsula,
Laconia Unspecied Volcanic rocks 452 37.58N 23.39E Naturhistorisches
Museum Wien
Mediodactylus kotschyi in the Peloponnese peninsula
with > 800 mm precipitation annually (Fig. 1), although
only seven of these are from areas with > 850 mm pre-
cipitation annually.
In seven of the eight locations (Table 1a) in which we
encountered the species in our 2015 eldwork we found
geckos on trees (in three of them exclusively on trees),
and in ve locations we encountered them under rocks
(in one exclusively under rocks). Absence from both
microhabitats was also recorded (Table 1b).
irty-two percent of gecko localities in the Pelopon-
nese are located in regions where limestone is the domi-
nant rock type (combined data from Tables 1a and 2). We
found no connection between rock type and arboreality
(Fisher’s exact test, two on trees and nine under rocks
in limestone habitats, seven on trees and 11 under rocks
in other rock types, P = 0.41), or between habitat type
(rocks or trees) and precipitation (trees: 649 ± 35 mm,
rocks: 692.1 ± 28 mm; t-test assuming unequal variances,
t = 2.1, n = 29, P = 0.34).
Twenty ve percent of our absence ndings were in
localities in which limestone is the dominant rock type
(although we did not try to identify the type of the rocks
under which we searched for geckos), and all of them
were for locations in which we searched for geckos under
Location name Substrate Rock type
Latitude Longitude Source
Tiryntha, Argolis Peninsula Unspecied Alluvium 583 37.59N 22.80E Beutler and Gruber
1977; Valakos et al. 2008
Argos, Argolis Peninsula Unspecied Alluvium 603 37.632N 22.732E Valakos et al. 2008
Palea Epidavros, Argolis
Peninsula Unspecied Alluvium 502 37.635N 23.153E Valakos et al. 2008
Epidavros, Argolis Peninsula Unspecied Limestone 560 37.65N 23.14E Beutler and Gruber
Archea Olympia, Elis Under stones in
phrygana Alluvium 781 37.651N 21.618E Pers. obs. Kostas
Sagonas 2014
Inachos, Corinthia Unspecied Alluvium 603 37.659N 22.750E Valakos et al. 2008
Nea Epidavros, Argolis Peninsula Unspecied Peridotite and serpentinite 555 37.675N 23.126E Valakos et al. 2008
Nea Epidavros, Corinthia Under stones in
phrygana Peridotite and serpentinite 538 37.675N 23.134E Pers. obs. Kostas
Sagonas 2014
Levidi, Arcadia Unspecied Limestone 873 37.68N 22.29E Beutler and Gruber
1977; Valakos et al. 2008
Kamenitsa, Laconia Unspecied Limestone 833 37.72N 22.19E Bringsoe 1985; Valakos
et al. 2008
Tropaia, Arcadia Unspecied Limestone 864 37.730N 21.954E Valakos et al. 2008
3 km north east of Soko,
Corinthia Stone terrace Limestone 639 37.81N 23.08E Bringsoe 1985
9 km east of Lampeia, Archaia
4 eggs under a
at rock Limestone 927 37.85N 21.91E Bringsoe 1985
Ano Tripotama, Achaea Unspecied Limestone 887 37.857N 21.912E Valakos et al. 2008
Archea Korinthos, Achaea Unspecied Neogene and Pleistocene
sediments 587 37.903N 22.882E Valakos et al. 2008
Korinthos, Achaea Unspecied Neogene and Pleistocene
sediments 566 37.936N 22.927E Valakos et al. 2008
Feneos, Corinthia On walls Alluvium 835 37.950N 22.325E Koppitz 2013
Kokkoni, Achaea Unspecied Alluvium 600 37.966N 22.779E Valakos et al. 2008
2.5 km east of Karia, Corinthia On a wall in
phrygana Alluvium 607 38.009N 22.442E Pers. obs. Kostas
Sagonas 2014
4 km south west of Kalavrita,
Achaea Unspecied Limestone 883 38.010N 22.079E Valakos et al. 2008
2 km south of Kalavrita, Achaea Unspecied Neogene sediments 865 38.018N 22.102E Valakos et al. 2008
Mega Spilaio, Achaea Unspecied Neogene sediments 848 38.08N 22.17E Stepánek 1937
Trapeza, Achaea On a wall Neogene sediments 728 38.172N 22.229E Pers. obs. Kostas
Sagonas 2014
184 Rachel Schwarz et alii
rocks (Table 1b). Precipitation was not signicantly great-
er in localities where geckos were not present (Table 1b)
than where they were encountered (Table 1a; presence:
690 ± 23 mm, absence: 727 ± 32 mm; t-test assuming
unequal variances, t = 2.13, n = 33, P = 0.38).
Our hypotheses were only partially supported.
Geckos were indeed more common in the more arid are-
as of the Peloponnese. However, we did not nd evidence
of limestone avoidance. e high frequency of occur-
rence of M. kotschyi on trees in the Peloponnese contrasts
with that on the Greek islands. During eldwork in the
Cyclades (once or twice from 2013 to 2015; e.g., Slavenko
et al., 2015) we observed M. kotschyi on a tree trunk only
twice (on Ano Koufonisi, in May 2013, and on Kimolos
Island, in May 2015, Fig. 2). All our other observations
(~ 800, from 40 islands) of this species were on and
under rocks, in stone piles, on dry stone walls and on
low building walls, under various objects of refuse and in
abandoned stone shelters (see also Arnold and Ovenden,
2002; Beutler, 1981; Musters and In den Bosch, 1982 and
citations therein). During our 2015 survey in the Pelo-
ponnese, most specimens (81%) were found on trees,
especially on almond, olive and eucalypts (Table 1a). In
only three locations were specimens found under rocks
(Table 1a), despite searching localities with apparently
suitable phrygana habitats.
Because our sole criterion for establishing absence
was that we did not nd the species following a search
under what we considered to be suitable conditions for
M. kotschyi, we are well aware that some absences may
Annual precipitation(mm)
Fig. 1. Presence and absence localities derived from the literature and our eld observations plotted on an annual mean temperature map of
the Peloponnese (adopted from Worldclim, Hijmans et al. 2005). Squares () designate specimens found among rocks; triangles (∆) desig-
nate specimens found on trees; circles (o) designate specimens for which substrate was not specied; dots inside the symbols (•) represent
published, museum and observational data (otherwise: our data); Blue: presence; Red: absence.
Mediodactylus kotschyi in the Peloponnese peninsula
very well be false-absences. is can only be supported
(or refuted), however, by future surveys. at said, we
have no reason to believe that reported absences are more
likely for either the tree or for the rock microhabitat, or
for dierent geographic locations, and thus false absences
are unlikely to alter our conclusions.
Werner (1993) described M. kotschyi as being a “par-
adoxical” species. He contended that, in Israel and Iraq,
it lives mainly on large tree trunks with exfoliating bark,
such as carob, eucalypts and oak (Werner, 1993), even
though it lacks the characteristic adhesive toe pads of
other arboreal geckos (Gamble et al., 2012). M. kotschyi
is nonetheless superbly camouaged against the back-
ground pattern of tree trunks (Werner, 1993; Baier et al.,
2009; Bar and Haimovitch, 2012; see also Fig. 2), making
it hard to dismiss the idea that it is well adapted to living
on trees as well as on rocks.
According to the most comprehensive phylogenies
available (Pyron and Burbrink, 2014), the closest rela-
tives of M. kotschyi are the arboreal Mediodactylus sag-
ittifer and the saxicolous Mediodactylus heteropholis
and Mediodactylus heterocercus (M. kotschyi is sister
to a clade containing all three). More distantly-related
allies (Pyron and Burbrink, 2014) include members of
the mostly saxicolous and terrestrial genera Tenuidacty-
lus and Cyrtopodion (note that Tenuidactylus caspius is
described as arboreal, saxicolous and terrestrial, Rogner
1997), and the mostly terrestrial Bunopus, Agamura and
Crossobamon. e ancestral state of M. kotschyi is thus
most likely terrestrial or saxicolous although an arboreal
ancestor cannot be ruled out. e fact that M. kotschyi is
saxicolous over most of its distribution may imply that
this species was originally saxicolous and later adapted
to inhabit trees too.
Current data on the occurrence of this species in the
Peloponnese (Tables 1 and 2) do not suggest a strong
preference of M. kotschyi for a specic type of substrate,
and we did not detect an aversion to limestone. e thin-
toed gecko does occur in places where the mean annual
precipitation is greater than 850 mm, although it is prob-
ably scarce in such regions. It is certainly not obligatorily
associated with phrygana, in contrast to what was previ-
ously suggested (Beutler, 1981). Our ndings, along with
our observations of this species on trees, lead us to sug-
gest that M. kotschyi is highly exible and adaptable in
its habitat preference, which may have contributed to its
successful establishment and broad range.
Our observations indicate that M. kotschyi is rela-
tively abundant on trees in the Peloponnese, whereas it is
extremely abundant and conspicuous on and under rocks
and on stone walls in the Cyclades. is might have led
to the general misconception that it is purely saxicolous.
Although it is certainly much more abundant on islands
(as many lizards are, Novosolov et al., 2013), we suggest
that M. kotschyi is more common in the Peloponnese
than has previously been considered, because it was for-
merly sought mostly on and under rocks.
This research was done under permit number
20305/824 from the Ministry of the Environment. We
wish to thank Oliver Tallowin, Anat Feldman and Maria
Novosolov for help with GIS construction. We wish to
thank three anonymous referees for comments on a pre-
vious version of this manuscript. is study is funded by
an ISF grant #1005/12 to SM.
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... Mediodactylus kotschyi is a small (mean snoutvent-length [SVL] 42.2 6 4.3 mm, mean mass 2.7 6 0.9 g, Itescu et al. 2017), padless, cathemeral, and insectivorous gecko, ranging from the southern Balkans, throughout Greece, including the Cyclades and North Aegean archipelagos (Kotsakiozi et al. 2018;Lymberakis et al. 2018). In the Peloponnese region (mainland Greece) M. kotschyi is almost exclusively arboreal (Schwarz et al. 2016;Fig. 1), while on the small, treeless Aegean islands it is exclusively saxicolous, inhabiting rocks, stone piles, and drystone walls (Beutler 1981;Valakos et al. 2008;Schwarz et al. 2016;Fig. ...
... In the Peloponnese region (mainland Greece) M. kotschyi is almost exclusively arboreal (Schwarz et al. 2016;Fig. 1), while on the small, treeless Aegean islands it is exclusively saxicolous, inhabiting rocks, stone piles, and drystone walls (Beutler 1981;Valakos et al. 2008;Schwarz et al. 2016;Fig. 1). ...
... These results demonstrate that individuals of this species are neither rigidly associated with the substrate they usually occupy in nature nor specifically adapted to it, and that they will use substrates that are not present in their natural environment. This implies that these geckos are opportunistic in their habitat use, which agrees with their widespread distribution and the high densities they reach within their range (Valakos et al. 2008;Schwarz et al. 2016;Itescu et al. 2018). This, potentially along with its preference for trees (albeit not statistically significant in our analyses, and thus requiring further examination), may explain why M. kotschyi was observed by Pafilis et al. (2020) on olive trees planted on the small (<1 km 2 ) islet of Monolia in the Lichadonissia Archipelago. ...
Populations of the same species occupying different microhabitats can either exhibit generalized traits across them or display intraspecific variability, adapting to each microhabitat in order to maximize performance. Intraspecific variability contributes to the generation of diversity, following selection and adaptation, and understanding such variability is important for comprehending how individuals choose their microhabitats. Compared to interspecific variability, however, intraspecific variability in functional morphology and its relationship with microhabitat preference and use have been relatively little studied. Here we examined whether populations of the gecko Mediodactylus kotschyi that differ in the substrates they occupy, display habitat-specific behaviours and differing morphologies associated with functional adaptation to their microhabitats. We collected 207 geckos from under or on rocks or on trees from seven populations in Greece. On large islands individuals occupy both substrates; whereas small islets are devoid of trees and the geckos are restricted to rocks, while on the mainland they are only found on trees.We determined gecko substrate preferences in the lab, together with their clinging abilities to the different substrates. We measured their limbs, digits and claws and assessed how these measurements relate to clinging ability.Geckos from all populations preferred the tree made available to them, but this preference was not statistically significant. Geckos from both large and small islands clung better to the tree than to the rock in the lab, while those from the mainland clung similarly to both substrates. Geckos collected from trees had longer manual digits and hind limbs. Geckos collected from large and small islands had taller (longer on the dorso-ventral axis; henceforth "deeper") claws. Longer digits and deeper but shorter claws were associated with a better ability to cling to rocks. Our findings suggest that while M. kotschyi is potentially preferentially arboreal, due to the great variation and plasticity it possesses, it can successfully also exploit the habitats available on the smallest, treeless islets in the Aegean Sea. Our study suggests that the dichotomous use of generalist vs. specialist in describing species' habitat use is oversimplified, and we suggest the use of a generalist-specialist gradient instead.
... Mediodactylus geckos are distributed across the Balkan Peninsula, including the Aegean Sea islands and mainland Greece (Ajtić, 2014;Schwarz et al., 2016;Roll et al., 2017;Kotsakiozi et al., 2018). The closely related Mediodactylus kotschyi (Steindachner, 1870) and Mediodactylus oertzeni (Boettger, 1888) are widespread on Aegean islands and islets (Kotsakiozi et al., 2018). ...
... Each island is inhabited by only one species of Mediodactylus. The geckos vary greatly between islands in morphology, behaviour, diet, abundance and life history (Valakos, 1989;Valakos & Polymeni, 1990;Mollov, 2011;Slavenko et al., 2015;Schwarz et al., 2016;Itescu et al., 2017Itescu et al., , 2018. This variation makes them excellent organisms with which to study the effects of island attributes on life-history traits. ...
Insular animals are thought to be under weak predation pressure and increased intraspecific competition compared with those on the mainland. Thus, insular populations are predicted to evolve 'slow' life histories characterized by fewer and smaller clutches of larger eggs, a pattern called the 'island syndrome'. To test this pattern, we collected data on egg volume, clutch size and laying frequency of 31 Aegean Island populations of the closely related geckos of the Mediodactylus kotschyi species complex. We tested how predation pressure, resource abundance, island area and isolation influenced reproductive traits. Isolation and predation were the main drivers of variation in life-history traits. Higher predator richness seemed to promote faster life histories, perhaps owing to predation on adults, whereas the presence of boas promoted slower life histories, perhaps owing to release from predation by rats on the eggs of geckos. Insular geckos followed only some of the predictions of the 'island syndrome'. Predation pressure seemed to be more complex than expected and drove life histories of species in two opposing directions. Our results highlight the importance of considering the identity of specific predators in ecological studies. ADDITIONAL KEYWORDS: Aegean Islands-clutch frequency-clutch size-egg volume-geckos-island biology-island syndrome-life-history traits.
... A bulgáriai populációk épületeken is előfordulnak, de a Balkán többi részén többnyire sziklás domboldalakon, kőkerítéseken látható. Az egykor ebbe a fajba sorolt Mediodactylus orientalis viszont főleg idős fák törzsén él (Werner 1993), de Görögországban Kotschy-gekkókat is figyeltek már meg fatörzseken (Schwarz et al. 2016). A nőstény meszes héjú, 8,0-9,5 x 6,8-7,3 milliméteres (M. ...
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Data on the occurrences of the introduced Kotschy's gecko, Mediodactylus kotschyi (Stein-dachner, 1870) in Hungary.) Invasive alien species are among the most prominent threats to biodiversity. In general, information on introduced species is still inadequate. Many potential invasive species are still in a lag phase, and thus elude attention. The number of established alien reptile populations in Hungary is small. One of them is the Mediterranean Kotschy's gecko, Mediodactylus kotschyi with synanthropic tendencies, which inhabits buildings in parts of its distribution range. In part from these populations, four colonies have been established in Hungary; three on isolated residential houses and one extending to several building blocks in the city centre of Budapest. Their winter survival probably depends on the heating of these buildings. The future of the colonies on residential area houses is uncertain but the colony living in the city may be able to persist into the future and even spread to new compounds. Their effect as competitors on the local herpetofauna will probably be insignificant but they may host parasites, which may spread to indigenous reptiles, especially to the commonly urban dweller Podarcis muralis.
... We observed this behavior in the Plakida Mediodactylus population only in response to the removal of cover under which they hided, which we think the geckos interpret as posing predation threat. The common antipredator mechanisms of Mediodactylus geckos are camouflage (especially where they are arboreal, Schwarz et al., 2016) and tail autotomy . Our observation thus potentially expands the known repertoire of antipredatory tactics in this species. ...
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Body size evolution on islands is widely studied and hotly debated. Gigantism and dwarfism are thought to evolve under strong natural selection, especially on small remote islands. We report a curious co-occurrence of both dwarf and giant lizards on the same small, remote island (Plakida): the largest Podarcis erhardii (Lacertidae) and smallest Mediodactylus kotschyi sensu lato; Gekkonidae — the two commonest insular reptiles in the Aegean Sea. The geckos of Plakida have a peculiar tail-waving behavior, documented here for the first time in this genus. We suspect that P. erhardii evolved large size to consume geckos and the geckos evolved a unique tail-waving behavior as a defensive mechanism.
... Thus we assume that this cryptic gecko was observed in our survey because we have previous experience locating it (e.g. Slavenko et al., 2015;Schwarz et al., 2016;Itescu et al., 2017). As long as surveys such as this are not performed by the same team members, differences in the detectability of the reptiles are likely to have an effect on the results (Mazerolle et al., 2007;Lardner et al., 2015). ...
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The south-facing slopes in canyons, oriented along an east-west axis north of the equator, are often hotter and drier than north-facing slopes, promoting differences in the biotic and abiotic characteristics of the opposing slopes. We studied how diversity and abundance patterns have changed in Oren stream (Carmel Mountains, Israel) during the last 25 years. We tested whether temperature and habitat preferences of reptiles affected observation frequencies, to assess potential effects of global warming on the reptiles. We compared the results of a 1993–1994 survey in Oren stream to a survey we conducted during 2017–2018, using similar methods, survey area and effort. Species composition and abundance in Oren stream did not significantly change between studies, but the proportion of observations differed significantly across slopes for four out of the six most abundant species. The number of observations increased monotonically with increasing temperatures on the south-facing slope, but decreased on the north-facing slope above a temperature of 22°C. The major biome species inhabit globally was unrelated to the number of observations across slopes or studies, but species inhabiting warmer ranges were more frequently observed in the current survey. Our results suggest that as global temperatures rise, reptile species which can tolerate higher temperatures, and those which can avoid the hottest temperatures of the day, may be able to cope better. These results however may also derive from better detection ability of some species over others between study teams.
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Terrestrial reptiles are threatened by numerous anthropogenic activities, including agriculture. Many agricultural methods and techniques affect the herpetofauna located in the oldest known tree crops in the Mediterranean Basin, olive trees. For the first time, we present a case of unintentional capture (and killing) of 12 snake-eyed skinks Ablepharus kitaibelii (Bibron & Bory de Saint-Vincent, 1833) on an insect control sticky trap in an olive grove in central Greece.
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billeder der er taget af den nyligt genfundne skivetung ede frø, Discoglossus nigriuenter. En art man regnede for uddød, da den ikke var fundet siden 1955. Arten er endemisk for Israel og blev genfundet i 2011. Det undrer mig dog en del, hvorfor de har valgt ikke at medtage de fem arter havskildpadder som er registreret fra Israels kyster, da disse for-I 2011 udkom "A Field Guide to Reptiles and mentligt også ville vaere relevante for øko-Amphibians of Israel" skrevet af Aviad Bar turister med hang til badning/dykning ved og Guy Haimovitch. Det er mig bekendt den kysterne. første engelsksprogede feltguide til Israels Bogen er gennemgående velgjort, med en spaendende herpetofauna. praesentation af hver art på to sider. Her bru-ges en side på beskrivelse af dyret, lidt infor-Israel er et af de lande i middelhavsområdet mation om artens levevis og dens udbredelse i der har flest hjemhørende arter af krybdyr Israel, samt listning af eventuelle underarter. og padder-kun overgået af Marokko, som Ligeledes vises et lille kort over artens udbre-har nogle få arter mere. Den israelske her-delse i landet. Anden side i praesentationen petofauna er interessant, fordi den består af viser typisk to billeder af den omhandlede art. såvel typiske europaeiske arter, men også af Beskrivelserne er mig en smule korte og det typiske mellemøstlige og nordafrikanske fau-kan godt aegre mig lidt, at forfatterne vaelger naelementer. Israel er med andre ord et sam-at bruge plads på artens udbredelse udenfor lingspunkt mellem disse tre regioner. I den Israel, da den plads der bruges til dette, nemt nordlige del af Israel er der kølige bjerge og kunne vaere brugt til at informere lidt mere frodige skovområder, i den sydlige del varm om selve artens levevis i Israel, og dermed og sandet ørken, alt sammen indenfor en gøre bogen mere brugbar i søgningen efter afstand, der er til at overkomme i bil. Altså den pågaeldende art. et meget varieret miljø, som nemt kan nås på en ferie. Alt dette indenfor en rimelig rejseaf-Ved praesentationen af f.eks. Eumeces schruei-stand af Danmark og derfor blot endnu mere deri vises to bitleder af to temmelig forskel-interessant for danske feltherpetologer med lige skinker, den ene en hun og den anden en rejselyst. At det derudover skulle vaere bil-han, men ved at sammenligne den medføl-ligt at komme der til og forholdsvist simpelt gende billedtekst, med lokalitet for de to dyr, at leje bil og køre rundt i Israel gør det ikke synes det oplagt at de begge repraesenterer mindre interessant. denne art, men to forskellige underarter. Det Den her omhandrede rertguide er derror Lff;fl"lXl"f:m}||åffi'''fl:ålJå11# kaerkommen og laenge ventet. Den er skre-. arter oS o; hr;;;;r ligeså _ eller om de to vet af to aktive israelske feltherpeloger, med underarters hanner og hunner adskiller sig grundig viden og indsigt i deres lands herpe-fra hinanden. Der gives ingen karakterer til tofauna' Bogen er rigt illustretet med farve-ui.t de to underarter desvaerre. BIot fotos afhøj kvalitet. Stort set alle disse er U"."u" udfra udbredelseskortet håbe på at taget af forfatterne selv, hvilket siger noget man kan underartsbestemme en eventuelt set om deres engagement i området' Interessant skink. Her kunne jeg godt have ønsket mig er det også i bogen at finde nogle afde første Nordisk Herpetologisk Forening
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summary: The distribution range of Kotschy's gecko (Cyrtopodion kotschyi) extends from southern Italy, across the Balkan Peninsula, to Asia Minor. To date, more than 25 subspecies have been described within this species. In this study, we analysed approximately 300 samples belonging to nine subspecies, from populations inhabiting the mainland portion of the species' distribution range. The specimens analysed herein belong to the following subspecies: Cyrtopodion kotschyi kotschyi, C. k. bibroni, C. k. rumelicus, C. k. skopjensis, C. k. danilewskii, C. k. lycaonicus, C. k. steindachneri, C. k. orientalis and C. k. syriacus. Morphological analyses included 23 morphometric, seven meristic and five quantitative traits. In analysing these data, several statistical procedures were used (discriminant analysis, cluster and correspondence analyses), in order to define or delineate populations at multivariate levels. Results from discriminant analysis of morphometric traits revealed highly complex differentiation of the analysed groups, and no clear pattern of geographic variability. Results from correspondent analysis of meristic and qualitative traits showed that different combinations of these traits and their states can define mainland geographical groups from the Balkan Peninsula in comparison to populations from the Near East. With this approach, we proved the validity of the so-called kotschyi and danilewskii subspecies' groups. Previously, researchers defined these two groups of subspecies on a descriptive, morphological level. In this study, we also discussed the validity of some of the previously described subspecies belonging to these two basic groups. The results of the study showed that there are no explicit diagnostic characters which could clearly define the subspecies within the kotschyi and danilewskii groups. This implies that the majority of the previously described subspecies are in fact synonyms within the two groups. We compared the results of our morphological analyses with previously published phylogeographical studies. Behavioural observations performed in the field (at study sites near Prizren, which lies at the edge of this species' distribution range in the Balkans) and in captivity (terraria), showed that the activity of C. kotschyi depends on temperature and daylight regimes, as well as prey activity. In the surroundings of Prizren, we studied
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The island syndrome describes the evolution of slow life history traits in insular environments. Animals are thought to evolve smaller clutches of larger offspring on islands in response to release from predation pressure and interspecific competition, and the resulting increases in population density and intraspecific competition. These forces become more pronounced with diminishing island size, and life histories are thus expected to become slowest on small, isolated islands. We measured clutch sizes in 12 insular populations of Mediodactylus kotschyi, a small gecko from the Cyclades Archipelago, a set of land-bridge islands in the Aegean Sea (Greece). We analyse variation in clutch size in relation to island area, island age, maternal body size, the presence of putative competitors and nesting seabirds (which increase resource abundance in the form of marine subsidies), and richness of predators. Clutch size of M. kotschyi decreases with increasing island area, in departure from classic island syndrome predictions, sug-gesting the evolution of faster life histories on smaller islands. There are no relationships between clutch size and island age, maternal size, the presence of competitors or predator richness. Instead, larger clutches on small islands could simply reflect the beneficial effect of marine subsidies derived from resident seabird colonies. Indeed, populations of M. kotschyi on islands with nesting seabirds have clutch sizes 30.9 % larger (1.82 vs. 1.39 eggs) than populations on islands without nesting seabirds. Thus, our data suggest that bottom-up effects of marine subsidies may supersede the expression of a simple island syndrome in the Aegean M. kotschyi.
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In this paper we present confirmed and potential distribution ranges of all native Serbian reptile species. The information provided herein presents the combination of the newly collected faunistic data and previously published records. The centres of reptilian diversity in Serbia were evaluated, in order to focus future conservation efforts on the regions of particular importance for the protection of this understudied group of animals. We found four main centres of species richness: three in Metohija and one in Šumadija, with 17–21 species per 50 × 50 km square. Analysis of the similarity of species composition in different regions of Serbia showed that South-eastern Serbia is the most distinct from other regions of Serbia, due to high number of Mediterranean species, with two being found exclusively in that region (E. quatuorlineata and P. najadum). Metohija is also very distinct, due to the highest number of species and significant number of specific (Mediterranean) faunal elements. The remaining geographic regions are grouped into three clusters, with Kosovo-Southern Serbia cluster being especially distinct in the terms of species richness and the presence of Mediterranean species. We also compared Serbian herpetofauna with those in other Balkan countries, considering species’ numbers and zoogeographic herpetofaunistic elements. Serbian herpetofauna is closest to the Romanian. Zoogeographic analysis showed that reptilian fauna of Serbia consists of eight chorotypes, with the Eastern-Mediterranean (nine species) and Southern-European (five species) as the most dominant ones.