Acta Herpetologica 11(2): 179-187, 2016 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-18176 Mediodactylus kotschyi in the Peloponnese peninsula, Greece: distribution and habitat Rachel Schwarz1,*, Ioanna-Aikaterini Gavriilidi2, Yuval Itescu1, Simon Jamison1, Kostas Sagonas3, Shai Meiri1, Panayiotis Pafilis2 1 Department of Zoology, Tel Aviv University, Tel Aviv 6997801, Israel. * Corresponding author. E-mail: rachelschwarz13@gmail.com 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. 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. Keywords. Arboreality, habitat preferences, Mediodactylus kotschyi, Peloponnese, rock type. INTRODUCTION The genus Mediodactylus is predominantly Asian, with only one of 13 species being found in Europe. The 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 often absent from Peloponnese species checklists (e.g., Werner, 1929; Cyrén, 1935; Bischoff and Bischoff, 1980; Henle, 1989; Pèrez-Mellado et al., 1999). The 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 (“עצים tree-gecko”) reflects this (Werner, 1993; Bar“ = ”שממית 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, fig, 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 influence 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). The 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. MATERIAL AND METHODS 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 fieldwork 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. The 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 specified. We used χ2 tests for goodness of fit 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). RESULTS 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 181Mediodactylus kotschyi in the Peloponnese peninsula are from the central and eastern parts of the peninsula (Fig. 1). The 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 fit 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 field 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 Annual precipitation (mm) No. individuals caught (rocks/ trees) 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, Laconia Dense phrygana Under a rock Limestone 585 (1/0) 36.5690N 22.9890E Gefira, 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, Laconia 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 forest 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, Laconia Phrygana (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 Annual prec. (mm) Latitude Longitude Source Lachos, Mani Peninsula Unspecified Limestone-marble 688 36.48N 22.37E Valakos et al. 2008 Kokkala, Mani Peninsula Unspecified Limestone-marble 651 36.52N 22.47E Valakos et al. 2008 Kato Kastania, Malea Peninsula, Laconia Phrygana with many rocks Limestone 596 36.52N 23.11E Bringsoe 1985 Ano Kastania, Malea Peninsula, Laconia Unspecified Phyllites 606 36.537N 23.102E Valakos et al. 2008 Lira, Malea Peninsula, Laconia Unspecified Phyllites 637 36.640N 22.964E Valakos et al. 2008 Loutsa, Mani peninsula Unspecified Limestone-marble 711 36.643N 22.474E Valakos et al. 2008 Monemvasia, Malea Peninsula, Laconia Unspecified Limestone 546 36.69N 23.05E Beutler and Gruber 1977 South of Agios Ioannis, Malea Peninsula, Laconia Unspecified 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 Unspecified 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 Unspecified 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 Unspecified Neogene sediments 768 36.90N 22.26E Werner 1937; Valakos et al. 2008 Mandina near Kampos, Mani Peninsula Unspecified Flysch 781 36.93N 22.20E Naturhistorisches Museum Wien Kalamata, Messenia Unspecified Alluvium 762 37.04N 22.11E Stepánek 1937; Valakos et al. 2008 2 km south of Gargalianoi, Messenia On the ground in open field Peridotite and serpentinite 813 37.049N 21.634E Pers. obs. Kostas Sagonas 2014 Mystras, Laconia Unspecified Limestone- marble 776 37.06N 22.37E Beutler and Gruber 1977; Stepánek 1937; Valakos et al. 2008 10 km west of Sparta, Laconia Unspecified 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 Unspecified Alluvium 712 37.071N 22.430E Valakos et al. 2008 5 km north east of Kosmas, Arcadia Eggs under a flat 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 Unspecified Alluvium 798 37.45N 22.41E Naturhistorisches Museum Wien Didima, Corinthia Unspecified Limestone 527 37.461N 23.171E Valakos et al. 2008 Tripoli, Arcadia Unspecified Limestone 807 37.507N 22.371E Valakos et al. 2008 1 km south west of Mainalo, Arkadia Under stones in maquis Limestone 836 37.529N 22.299E Pers. obs. Kostas Sagonas 2014 Methanon, Malea Peninsula, Laconia Unspecified Volcanic rocks 452 37.58N 23.39E Naturhistorisches Museum Wien 183Mediodactylus 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 fieldwork we found geckos on trees (in three of them exclusively on trees), and in five locations we encountered them under rocks (in one exclusively under rocks). Absence from both microhabitats was also recorded (Table 1b). Thirty-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 five percent of our absence findings 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 Annual prec. (mm) Latitude Longitude Source Tiryntha, Argolis Peninsula Unspecified Alluvium 583 37.59N 22.80E Beutler and Gruber 1977; Valakos et al. 2008 Argos, Argolis Peninsula Unspecified Alluvium 603 37.632N 22.732E Valakos et al. 2008 Palea Epidavros, Argolis Peninsula Unspecified Alluvium 502 37.635N 23.153E Valakos et al. 2008 Epidavros, Argolis Peninsula Unspecified Limestone 560 37.65N 23.14E Beutler and Gruber 1977 Archea Olympia, Elis Under stones in phrygana Alluvium 781 37.651N 21.618E Pers. obs. Kostas Sagonas 2014 Inachos, Corinthia Unspecified Alluvium 603 37.659N 22.750E Valakos et al. 2008 Nea Epidavros, Argolis Peninsula Unspecified 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 Unspecified Limestone 873 37.68N 22.29E Beutler and Gruber 1977; Valakos et al. 2008 Kamenitsa, Laconia Unspecified Limestone 833 37.72N 22.19E Bringsoe 1985; Valakos et al. 2008 Tropaia, Arcadia Unspecified Limestone 864 37.730N 21.954E Valakos et al. 2008 3 km north east of Sofiko, Corinthia Stone terrace Limestone 639 37.81N 23.08E Bringsoe 1985 9 km east of Lampeia, Archaia Olympia 4 eggs under a flat rock Limestone 927 37.85N 21.91E Bringsoe 1985 Ano Tripotama, Achaea Unspecified Limestone 887 37.857N 21.912E Valakos et al. 2008 Archea Korinthos, Achaea Unspecified Neogene and Pleistocene sediments 587 37.903N 22.882E Valakos et al. 2008 Korinthos, Achaea Unspecified 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 Unspecified 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 Unspecified Limestone 883 38.010N 22.079E Valakos et al. 2008 2 km south of Kalavrita, Achaea Unspecified Neogene sediments 865 38.018N 22.102E Valakos et al. 2008 Mega Spilaio, Achaea Unspecified 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 significantly 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). DISCUSSION Our hypotheses were only partially supported. Geckos were indeed more common in the more arid are- as of the Peloponnese. However, we did not find evidence of limestone avoidance. The high frequency of occur- rence of M. kotschyi on trees in the Peloponnese contrasts with that on the Greek islands. During fieldwork 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 find the species following a search under what we considered to be suitable conditions for M. kotschyi, we are well aware that some absences may 1 2 Annual precipitation (mm) Fig. 1. Presence and absence localities derived from the literature and our field 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 specified; dots inside the symbols (•) represent published, museum and observational data (otherwise: our data); Blue: presence; Red: absence. 185Mediodactylus kotschyi in the Peloponnese peninsula very well be false-absences. This can only be supported (or refuted), however, by future surveys. That said, we have no reason to believe that reported absences are more likely for either the tree or for the rock microhabitat, or for different 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 camouflaged 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. The ancestral state of M. kotschyi is thus most likely terrestrial or saxicolous although an arboreal ancestor cannot be ruled out. The 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 specific type of substrate, and we did not detect an aversion to limestone. The 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 findings, along with our observations of this species on trees, lead us to sug- gest that M. kotschyi is highly flexible 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. This 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. ACKNOWLEDGEMENTS 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. This study is funded by an ISF grant #1005/12 to SM. REFERENCES Ajtić, R. (2014): Morphological, biogeographical and eco- logical characteristics of Kotschy’s gecko (Cyrtodacty- lus kotschyi Steindachner, 1870 Gekkonidae) from the mainland portion of its distribution range. Fauna Bal- kana 3: 1-70. Arnold, E.N., Ovenden, D. 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