HerPeToZoA 27 (1/2): 39 - 63 39 Wien, 30. Juli 2014

The distribution of amphibians and on island () (Amphibia: reptilia)

Die Verbreitung der Amphibien und reptilien auf der insel Samos (Griechenland) (Amphibia: reptilia)

Jeroen SPeyBroeCK & D Aniel BoHle & e DoArDo rAZZeTTi & M AriA DiMAKi & M Arlene KATHArinA KirCHner & W ouTer BeuKeMA

KurZFASSunG Die Arbeit faßt 1580 Funddaten von reptilien und Amphibien auf der insel Samos (Griechenland) aus den letzten Jahren (1993-2012) zusammen. Alle Funde wurden auf einem 1 km x 1 km raster verortet, wobei 40 % der Zellen (261 von insgesamt 648) zumindest eine Beobachtung einer Art aufwiesen. Auf dieser Datengrundlage wurden Verbreitungsmuster und ökologische Beobachtungen mit Vorkommen auf anderen inseln der Ägäis ver - glichen. Das in der Arbeit erstmals publizierte Vorkommen der Münzennatter nummifer (r euSS , 1834) auf dieser insel hebt die Anzahl der bekannten Arten für Samos auf 28 (4 Amphibien- und 24 reptilienarten). ABSTrACT A collection of 1580 recent data records (1993-2012) of amphibians and reptiles from the Greek island of Samos was compiled. All records were mapped onto a grid of 1 km by 1 km, with 261 of the total 648 cells (40 %) of the island holding at least one observation. Subsequently, the distribution patterns and ecological observa - tions were compared with data from other islands in the Aegean Basin. The discovery of the Coin-marked (r euSS , 1834), published herein after, constitutes a new island record, setting the herpeto - fauna list to 28 confirmed (4 amphibians, 24 reptiles). KeyWorDS Amphibia, reptilia, amphibians, reptiles, distribution, mapping, large contemporary dataset, Samos, Greece, Aegean, island herpetofauna, new island record, Hemorrhois nummifer

inTroDuCTion

Samos is a Greek island in the eastern ber of perennial and intermittent streams, , separated from Asia Minor by coastal marshes as well as man-made water the 1.3 km wide Mycale Strait (Figs. 1A, reservoirs are of particular importance to the 1B). The island has a surface of 476 km², anuran species of Samos, as well as a num - roughly 44 km from west to east by 19 km ber of species dependent on water from north to south ( STAMATelAToS & habitats (namely, Grass Snake and terra - VAMVA -S TAMATelATou 2006). Built up by pins). The island’s climate is of mild Medi - largely mountainous and hilly terrain with a terranean type, with monthly average tem - maximal elevation of 1,434 m (Mt. Kerkis), peratures ranging from 10.3 (January) to pine forest dominates considerable sections 28.4 °C (July) and monthly rainfall ranging of the inland. Additionally, the island hosts from 0.5 (July) to 164 mm (December) ( Hel - relatively large flatlands, especially in the leniC nATionAl MeTeoroloGiCAl Ser ViCe southeast. These areas are characterized by 2013). its heterogeneous landscape, mild non-intensively managed olive orchards climate and close proximity to the Anatolian with rich herbaceous undergrowth bordered mainland has resulted in a distinct and high by dry-stone walls, providing refuge for biodiversity. wildlife. As sustainable amphibian popula - The herpetofauna of Samos is one of tions rely on water for reproduction, a num - the most diverse of all Greek islands, likely 40 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA to be only surpassed by that of lepis aurata for the first time. The first (Kerkyra) ( TóTH et al. 2002; rAZZeTTi et al. explicit reports of Montivipera xanthina 2006). (1888) set the basis were published in the same year within of herpetological studies on Samos, describ - three different publications: VAn WinGerDe ing specimens collected by eberhard Von (1986), nilSon & A nDrén (1986) and oerTZen in 1887, thus presenting more or TieDeMAnn & G rilliTSCH (1986). The less the first reference to a list of Samos presence of the species on Samos was, how - amphibian and reptile species (listed here by ever, (indirectly) noted much earlier. in their current names): Bufo bufo , Pelophylax 1857, the italian explorer and zoologist bedriagae , Chamaeleo chamaeleon , Lau - orazio AnTinori was bitten in his right dakia stellio , Ophisops elegans , Anatolola - hand and suffered partial paralysis of the certa anatolica , Typhlops vermicularis and arm and discomfort for nearly two years modestus . Subsequently, enrica (AMBroGi 1992; MAZZoTTi 2011). Natrix CAlABreSi (1923) published a list of speci - tessellata was listed as occurring on Samos mens collected on the island in 1893 by by CHonDroPouloS (1989), by authority of Forsyth MAJor , adding Mediodactylus an observation made by S. VAlAKoS in kotschyi , Hemidactylus turcicus and Able - 1984, but has been revoked by the latter (S. pharus kitaibelii . unfortunately, the speci - VAlAKoS in verbis ). BuTTle (1990) men - mens collected by F.S.C. MAJor , until today tioned Blanus strauchi for Samos by author - present in the Museo di Storia naturale “la ity of a personal communication of Specola” of Firenze, cannot be assigned un- DiMiTroPouloS , which was subsequently ambiguously to Samos, as the locality re- indicated as erroneous by the latter (A. ported on the labels is “Samos e isole vicine” DiMiTroPouloS in verbis ). Bol (1992) was (= Samos and nearby islands) (Anna maria the first to mention Eryx jaculus for Samos niSTri , in litt. 2013). CAlABreSi ’s first from a 1989 find, although an earlier obser - mention of the presence of Mediodactylus vation (1988) was reported two years later kotschyi (cited as Gymnodactylus oertzeni ) (ioAnniDeS et al. 1994). ioAnniDeS et al. was later also listed by (1994) also published the presence of Hyla (1930, 1933), and with indication of doubt arborea (see also ioAnniDeS & D iMAKi adopted by BeuTler & G ruBer (1977) and 1996, 1997) and Malpolon insignitus for the CHonDroPouloS (1986). However, since first time. The last addition to the herpeto - the exact origin of the data is unclear and fauna list of the island was Emys orbicu - more recent records of M. kotschyi are lack - laris , published by Meyer & F riTZ (1996). ing from the available literature, there leaving sea turtles aside, and exclud - seems to be no convincing evidence that this ing three unconfirmed species ( Mediodacty - species is present on Samos. in contrast to lus kotschyi , Blanus strauchi and Natrix tes - CAlABreSi ’s citation of M. kotschyi , subse - sellata ), prior to this paper, twenty-six quent records confirmed the presence on species have been found, confirmed, pub - Samos of Hemidactylus turcicus (Werner lished and repeatedly recorded on Samos, 1935) and Ablepharus kitaibelii (Werner which includes four species of amphibians 1930). A third major addition to the list of and twenty-two species of reptiles. As some herpetofauna species of Samos was made by of these species occur within Greece only a number of papers by Werner (1930, on Samos and other islands off the Turkish 1933, 1935, 1938), adding Bufo viridis, western , the herpetofauna of the island Mauremys rivulata , Testudo graeca , Pseu - has been identified as holding an ‘Asian’ dopus apodus , Dolichophis caspius , Zame - character, in contrast to islands west of the nis situla and Natrix natrix to the faunal Mid-Aegean Trench, with a more ‘- inventory. From the second half of the last pean’ fauna composition ( PAFiliS 2010). century onwards, the majority of the species The aim of this paper is to collect, dis - seemed to have been discovered and report - cuss and map recent distribution records, ed, and new discoveries became less numer - most of which have never been published ous. DAAn (1967) added Lacerta trilineata , before, in order to enhance knowledge Platyceps najadum and Telescopus fallax, about presence, abundance, seasonality and while BeuTler (1979) mentioned Trachy - distribution of the herpetofauna of Samos. Distribution of amphibians and reptiles on Samos island 41 0 0 0 2 0 6 4

1 MATeriAlS AnD MeTHoDS 4 0 3 0 % 1 1 1 1 1 2 1 1 l 0 7 0 5 7 5 2 4

a Data 0 6 8 t 4 6 6 5 9 5 1 2 6 4 5 o 1 1 1 1 1 4 2 t 1 Data was collected opportunistically 7 8 ? 1 5 5 from 1993 to 2012 (Table 1) by the authors as well as from numerous contributors (see 2 2 7 0 2 acknowledgements). Table 1 summarizes 3 1 4 1 1 7 ‘ the temporal spread of the collected data,

9 with spring (April-May) and a number of 1 2 7 8 1 1 4 1 5 3 1 1 2 1 7 ‘ 1 years with dedicated field trips taking up large portions of the total data. 7 0 5 9 3 2 1 2 5 8 1 7 5 3 1

‘ The degree of accuracy differs within 1 the data, ranging from exact GPS coordi - 5 0 9 3 2 0 1 3 2 4 6

0 nates, estimated sites of location in Google 3 1 ‘ 1 1 earth and other available detailed mapping 4 8 8 3 0 8 4 tools, to only the indication of the grid cell 3 8 4 0 2 1 2 4 2 ‘ 1

. (see below) of observation. Such data was h t n of sufficient quality for the present mapping 7 o 3 5 1 1 0 1 7 8 ‘ m purposes, while less precise observations d n were omitted. a 6

9 0 . 1 r t 0 8 9 repeated sightings of the same spe - ‘ a a e n y o cies at the same site were not lumped, in y M 5 9 0 b

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2 3 d ‘ d n e and observational preferences. Densities u t a r r l h 4 6 h 4 (number of observed individuals per obser - u 2 a 2 7 a 0 b 5 J J ‘ 1 1 a

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t (Figs. 5 - 8) and calculate the portion of e o c u p e u u o ? D n o S A J J M M A t occupied grid cells within the total grid. 42 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Fig. 1: A & B - Geographic location of the island of Samos (east Aegean, Greece) and basic topography and hydrology of the island; C - Density distribution of collected herpetological data, represented by different shades of brown. The legend shows the number of species within each grid cell. Abb. 1: A. & B - Die geographische lage von Samos, sowie eine vereinfachte Darstellung der Topographie und Hydrologie; C - Dichte der gesammelten herpetologischen Daten, verdeutlicht durch unterschiedliche Braun-Schattierungen. Die legende zeigt die Anzahl der Arten innerhalb jeder Zelle. Distribution of amphibians and reptiles on Samos island 43

Altitudinal information was not avail - tained from online catalogues. unfortunate - able for the records. This is why for each ly, some of these queries re mained unan - species, the occupied grid cells were com - swered. Concerning the use of museum bined with altitude descriptors, viz. average acronyms, standard symbolic codes for insti - altitude ± SD ( HiJMAnS et al. 2013) to dis - tutional resource collections as listed in play the species’ vertical distribution. Mean SABAJ PéreZ (2013) were adopted: AMnH - altitude ± SD of the entire island, based on American Museum of natural History, new the averages of all its 648 grid cells is york; GnHM - Goulandris natural History 304.25 ± 263.59 m, and of the territory cov - Museum, Athinai; MCZ - Museo Civico ered by occupied grid cells, based on the di Zoologia, roma; MHnG - Muséum averages of the 261 occupied grid cells is d’Histoire naturelle, Genève; MnHn - Mu - 116 ± 150.37 m. séum national d’Histoire naturelle, ; Bibliographic data and references to MSnF - Museum of natural Sciences, museum specimens were included in each Firenze; nHMAS - natural History Museum species account. To reflect true data accu - of the Aegean, Mitilini; nHMC - natural mulation, it was attempted to only list papers History Museum of , iraklió; nHMW - providing new distributional data, excluding naturhistorisches Muse um, Wien; nrM - secondary sources like reviews such as Swedish Museum of natural History, CHonDroPouloS (1986, 1989). Specimen Stockholm; yPM - yale university Peabody information was requested from numerous Museum, new Haven. museum collections, while some were ob -

reSulTS AnD DiSCuSSion

A total number of 1,580 records (spe- one or more observations. important (in cies observation at a given site and date), most cases mountainous) parts of the center belonging to 28 species, were collected and central south of the island are readily (Table 2). Merging multiple observations of visible as gaps in the mapping effort, with the same species within the same grid cell, available data partially concentrated around resulted in 1,019 species-cell observations commonly used tourist accommodation remaining. facilities, serving as the working base of Species frequently represented by most of the field trips, which contributed to numerous records per grid cell showed this data compilation. higher values in the percentage of observa - The frequency distribution of specific tions in comparison to the percentage of grid occupancy rates (number of grid cells occupied grid cells (e.g., Pelophylax bedria - by number of herpetological species) is given gae and especially Mauremys rivulata ), or in Fig. 3. The highest number of species vice versa, occupied less grid cells than the within a single cell was 20. general overall trend would predict, judging The altitudinal distribution of each their number of observations. This is likely species is illustrated in Fig. 4 and will be to be due to both habitat specificity as well discussed in the species accounts below. As as relatively high ease of observation; other more observations and surveying activities types of biased sampling (seasonal and were done in lower, coastal and more acces - behavioral effects) may add to this imbal - sible areas, sampling is unbalanced with re- ance. Figure 2 illustrates the relation be - gard to medium and higher elevations. tween the tabulated percentages. The graph in the following section, the species indicates that the more frequently observed are discussed in detail. Available references species tended to be observed more often in to bibliography and museum specimens are the same grid cell than seemingly rarer given. species. Mean altitudes of the sites of observa - As shown in Fig. 1C, coverage is in - tion are represented by the mean altitudes of complete at the chosen grid resolution: 261 the corresponding grids ± 1 Standard Devia- grid cells of a total of 648 cells (40%) hold tion given in m a.s.l.. 44 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Table 2: Amphibians and reptiles of Samos island (Greece). overview of available records per species, including number of records, proportion of records in total dataset ( n = 1,580), number of occupied grid cells (1 km x 1 km), and proportion of occupied grid cells in total number of grid cells ( n = 648). Tab. 2: Amphibien und reptilien von Samos (Griechenland). Überblick der vorhandenen nachweise. Angegeben werden für jede Art: Anzahl der nachweise, Prozentsatz der nachweise bezogen auf den Gesamt- datensatz ( n = 1580), Anzahl der belegten rasterfelder (1 km x 1 km), Prozentsatz der belegten rasterfelder bezo - gen auf die Gesamtanzahl der rasterfelder ( n = 648).

Species number of Proportion relative number of Proportion relative - records / to all records (%) / grid cells / to all grid cells (%) / - Anzahl der Anteil an allen Anzahl der Anteil an allen - nachweise Beobachtungen (%) rasterfelder rasterfeldern (%) Bufo bufo 34 2.2 26 4.0 Bufo viridis 22 1.4 17 2.6 Hyla arborea 13 0.8 10 1.5 Pelophylax bedriagae 117 7.4 57 8.8 Testudo graeca 26 1.6 24 3.7 Mauremys rivulata 68 4.3 23 3.5 Emys orbicularis 7 0.4 4 0.6 Trachemys scripta 3 0.2 1 0.2 Laudakia stellio 182 11.5 93 14.4 Chamaeleo chamaeleon 162 10.3 86 13.3 Hemidactylus turcicus 48 3.0 31 4.8 Anatololacerta anatolica 45 2.8 30 4.6 Lacerta trilineata 100 6.3 60 9.3 Ophisops elegans 154 9.7 98 15.1 Ablepharus kitaibelii 47 3.0 40 6.2 aurata 23 1.5 20 3.1 Pseudopus apodus 66 4.2 46 7.1 Typhlops vermicularis 44 2.8 35 5.4 Eryx jaculus 24 1.5 23 3.5 Malpolon insignitus 18 1.1 17 2.6 Natrix natrix 30 1.9 18 2.8 Dolichophis caspius 106 6.7 67 10.3 Eirenis modestus 82 5.2 65 10.0 Hemorrhois nummifer 9 0.6 7 1.1 Platyceps najadum 54 3.4 47 7.3 Telescopus fallax 32 2.0 28 4.3 Zamenis situla 13 0.8 11 1.7 Montivipera xanthina 51 3.2 35 5.4 Total Σ = 1580 x = 3.56 Σ = 1019 x = 5.61

AMPHiBiAnS od), in combination with lack of interest for nocturnal searches during rain, and tadpoles as helpful means of detection. Being a spe- Bufo bufo cies of typically temperate humid climates, (linnAeuS , 1758) (Fig. 5) this toad is absent from most Greek islands (VAlAKoS et al. 2008). The relatively high even though it was one of the first mean altitude of the sites of observation species recorded ( BoeTTGer 1888), subse - (160 ± 27 m) coincides with a more inland quent sightings remained fairly scarce. distribution and could also be related to the only 34 records were collected, correspon - species’ occurrence in more temperate habi - ding to 2.2 % of the data. These fall within tats such as woodland. As inland records 26 grid cells (4.0 % of entire grid). The are relatively scarce, this species’ distribu - present records are scattered across different tion is likely insufficiently sampled. no parts of Samos. Possibly, a certain bias is obvious seasonal pattern seems to emerge caused by most of the data originating from from the data; the percentage of observa - months of suboptimal conditions for this tions per month does not strongly differ species to be active (after reproductive peri - from that of the entire dataset. Distribution of amphibians and reptiles on Samos island 45

Fig. 2: relation between percentages of observations (% obs) and occupied grid cells (% grid) for 28 herpetological species on the island of Samos. Abb. 2: Die Beziehung zwischen prozentualer Anzahl der Beobachtungen (% obs) und prozentualer Anzahl der besetzten rasterfelder (% grid) für 28 herpetologische Arten auf der insel Samos.

Fig. 3: Frequency distribution of specific grid occupancy rates (number of grid cells by number of herpetological species). Abb. 3: Häufigkeitsverteilung der Belegungsdichten der rasterzellen mit unterschiedlichen Artenzahlen (Anzahl von rasterzellen bezogen auf die Anzahl der herpetologischen Arten). 46 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Fig. 4: Altitudinal distribution of the amphibian and reptile species records ( n = 1019 grid cells) on the island of Samos. Triangles indicate individual data points, squares are species means. Abb. 4: Höhenverteilung der Amphibien- und reptiliennachweise ( n = 1019 rasterfelder) auf der insel Samos. Dreiecke kennzeichnen einzelne Fundpunkte, Quadrate zeigen den Durchschnittswert pro Spezies an.

Museum specimens: nHMAS 27-28, apply here as well. Sightings are relatively 4103, nHMW 27904, 27905, 30980:1-65. uncommon and seem to display some bias to Bibliographic data: BoeTTGer (1888), the more intensively sampled south-eastern Werner (1938), ioAnniDeS et al. (1994), parts of Samos, coinciding with a mean alti - ioAnniDeS & D iMAKi (1996), ClArK (2000), tude of observation (92 ± 25 m) which is CATTAneo (2003). lower than in B. bufo . Although the rather small number of observations for this Bufo viridis species may be misleading, its detectability (lAurenTi , 1768) (Fig. 5) may be higher in spring – while only 14 % of all available data was collected in April, only 22 records, or 1.4 % of the data, 32 % of the records of this species were col - relate to this species. These fall within 17 lected during that same month. The species grid cells (2.6 % of entire grid). While this may be recorded more efficiently earlier in rather xerophilic species is adapted to more the year. temporal, shallow and even brackish waters remarks: STöCK et al. (2006) pro - as breeding grounds than the previous posed to split Bufo viridis into a number of species, and consequentially occurs on many different species, of which Bufo variabilis of the bigger Greek islands ( VAlAKoS et al. (PAllAS , 1769) would be the taxon inhabit - 2008), other above raised hypotheses might ing Samos. However, as at present this Distribution of amphibians and reptiles on Samos island 47 arrangement is not generally accepted, the belonging to it. These fall within 57 grid authors follow SPeyBroeCK et al. (2010), cells (8.8 % of the entire grid), indicating a awaiting further support for the species rank rather high number of observations per grid of B. variabilis and maintaining the use of cell. in contrast to the other amphibians of the name Bufo . Samos, this species displayed a fairly regu - Museum specimens: MCZ A-19372, lar presence and abundance. Most records nHMA S 4106, nHMW 30818, 30991. are from sites at relatively low altitude (76 ± Bibliographic data: Werner (1933), 10 m, maximum of 610 m). ioAnniDeS et al. (1994), ioAnniDeS & D i- remarks: The validity of this species MAKi (1996), CATTAneo (2003). is under debate. HoTZ et al. (2013) suggest - ed Pelophylax bedriagae possibly should be Hyla arborea treated as a synonym of Pelophylax ridibun - BeDriAGA , 1890 (Fig. 5) dus while other authors like BÜlBÜl et al. (2011) confirmed the validity of the species only the larger, more humid Greek based on analyses of mtDnA sequences. islands are inhabited by this species ( VAlA- Museum specimens: AMnH 37088, et al. 2008). recorded for the first time nHMW 30817:1-4, 30990:1-7. as late as the first half of the 1990s ( ioAn- Bibliographic data: BoeTTGer (1888), niDeS et al. 1994; ioAnniDeS & D iMAKi 1996 CAlABreSi (1923), W erner (1938), B ol 1997), Hyla arborea seems localized and (1992), i oAnniDeS et al. (1994), ioAnniDeS rare on Samos (13 records (0.8 %) relate to & D iMAKi (1996), C lArK (2000), C ATTAneo the species), in contrast to its widespread (2003). and abundant nature in many other parts of the range. As these 13 observations fall within 10 different grid squares (1.5 % of rePTileS total grid), a possibly relatively large yet localized range of H. arborea seems to be Testudo graeca the case, at relatively low altitude: 53 ± 15 linnAeuS , 1758 (Fig. 5) m, maximum of 161 m. records appear to be clearly restricted to the south-eastern From the authors’ experience in other parts of the island. While this area does parts of Greece and in similar habi - indeed provide suitable wetland areas as tat and comparable seasonal conditions, habitat, it cannot be ruled out that other numbers of this species seem relatively low parts of Samos have been surveyed incom - on Samos: 26 sightings (1.6 % of collected pletely for this species. data) were made. These fall within 24 grid remarks: STöCK et al. (2008, 2011 cells, corresponding to 3.7 % of the entire and 2012) assigned the tree frogs of the area grid. A possibly biased image appears, as in which Samos is located (including sam - sightings seem absent from the northwest of ples from the Anatolian mainland as well as Samos. The altitudinal range of this species island) to Hyla orientalis BeDriAGA , is rather wide (188 ± 43 m). 1890. As taxonomical discussion is clearly Museum specimens: nHMA S 8102. not in the scope of the present paper, the Bibliographic data: Werner (1935), Samos populations were conservatively as - onDriAS (1968), i oAnniDeS et al. (1994), signed to H. arborea . MÜller (1995), i oAnniDeS & D iMAKi Bibliographic data: ioAnniDeS et al. (1996), B roGGi (1997), C lArK (2000), C AT- (1994), ioAnniDeS & D iMAKi (1996, 1997). TAneo (2003). Pelophylax bedriagae Mauremys rivulata (CAMerAno , 1882) (Fig. 5) (VAlenCienneS , 1833) (Fig. 5) While generally restricted to the water This species seems largely restricted sources of the island, this species was easily to the south-eastern wetland areas of the spotted, and thus commonly encountered, island. interesting in this regard, is a single with 117 records (7.4 % of collected data) record from the north-western coast near 48 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Karlovasi. A relatively high tolerance for Laudakia stellio elevated levels of salinity and eutrophica - (linnAeuS , 1758) (Fig. 6) tion make this species fairly abundant with - in suitable habitats. While local populations This most commonly recorded lizard, appear to be safe, the number of records accounting for 11.5 % of all collected data remains limited (68 records, 4.3 % of col - (182 records) and inhabiting the ample lected data). Furthermore, these fall within exposed, rocky habitats on the island, was 23 grid cells (3.5 % of the entire grid), thus indeed one of the most conspicuous, wide - indicating large numbers of observations spread and abundant species. observations per grid cell. From the collected data, this is fell within 93 grid cells (14.4 % of entire a lowland species (27 ± 6 m). grid), which indicates a rather high number Museum specimens: nHMC 80.3.15. of observations per grid cell. Altitudinal 101-104, nHMW 33342:1, nrM 6707. distribution was concentrated at 128 ±13 m, Bibliographic data: Werner (1935, with the maximum altitude recorded as high 1938), B ol (1992), i oAnniDeS et al. (1994), as at 999 m. ioAnniDeS & D iMAKi (1996), C lArK (2000), remarks: in the collection of the CATTAneo (2003). Muséum national d’Histoire naturelle (Paris) there is a stuffed specimen (MnHn Emys orbicularis 7494) of Stellio vulgaris collected by (linnAeuS , 1758) (Fig. 5) Admiral Jules DuMonT D’u rVille (1790- 1842) (ivan ineiCH in litteris , 2013). it was This species is extremely rare on probably collected during the hydrographi - Samos. Seven records were available (0.4 cal survey (1819-1820) of the Aegean Sea % of collected data), including the first by the ship Chevrette ( GoePP & C orDier sighting on the island ( Meyer & F riTZ 1877), thus this specimen represents the 1996), which is the only previously pub - oldest museum sample from Samos, appar - lished record. The other records refer to two ently predating the oldest available pub - additional coastal sites (4 ± 1 m). The ob- lished record ( BoeTTGer 1888). servations fall within 4 grid cells, corre - BAiG et al. (2012) proposed the assign- sponding to 0.6 % of the entire grid. Since ment of this species to the genus Stell - numbers of this species are reported by our agama . As evidence of the alleged para - observers as well as ClArK (2000) to be ex- phyly of the genus Laudakia sensu lato is tremely low, the conservation of the species limited ( MACey et al. 2000), the authors on Samos deserves attention. occurrence of refrain from the use of the new name. the species in the adjacent part of Turkey is Museum specimens: GnHM 5092, also restriced ( eiSelT & S PiTZenBerGer 7081, MCZ r-37019, MnHn 7494, MSnF 1967). 727, 13691-92, nHMAS 23-24, 4107, Museum specimens: GnHM 4104, 70011, nHMW 7297, 27584, 30820:1-2, nHMAS 4104. 30989:4, 33040:2, 35403:12-14, yPM Bibliographic data: Meyer & F riTZ Herr 005835. (1996), C lArK (2000). Bibliographic data: BoeTTGer (1888), CAlABreSi (1923), W erner (1933), o nDri- Trachemys scripta AS (1968), B ol (1992), i oAnniDeS et al. (SCHoePFF , 1792) (Fig. 5) (1994), ioAnniDeS & D iMAKi (1996), C lArK (2000), C ATTAneo (2003). Three records of this alien species, subspecies elegans (WieD , 1838) , at a sin - Chamaeleo chamaeleon gle site are available to the authors. As this (linnAeuS , 1758) (Fig. 6) site is also inhabited by Mauremys rivulata and close to one of the very few sites of As its current status on the islands of Emys orbicularis , eradication of this poten - and Crete needs further investigation, tially harmful species (e.g., CADi & J oly Samos currently holds the only stable and 2004; P olo -C AViA et al. 2011) may be con - confirmed population of this species in sidered. Greece ( DiMiTroPouloS 1987; V AlAKoS et Distribution of amphibians and reptiles on Samos island 49

Fig. 5: island of Samos, distribution maps – amphibians, tortoises and terrapins. Abb. 5: insel Samos, artspezifische Verbreitungskarten – Amphibien, land- und Sumpfschildkröten. 50 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA al. 2008). Chameleons are cryptic and easi - island, this species’ distribution seems to ly overlooked, thus often recorded as single indicate sampling bias, related to its noctur - individuals only. yet, as part of the present nal activity pattern and its presence in data originated from specific research effort human environments (among other places). (DiMAKi 2008; D iMAKi et al. 2010), the its actual presence might in reality take up a species takes up 10.3 % of the data (162 much more significant portion of the coastal observations). These fall within 86 grid lowlands, including less developed areas. cells, corresponding to 13.3 % of the entire Museum specimens: nHMAS 8103, grid. While also recorded in many places in nHMW 30995:4-6, MSnF 703. the interior of the island, most sightings Bibliographic data: CAlABreSi (1923), were made in or near wetlands fairly close Werner (1935), i oAnniDeS et al. (1994), to the coast, in concordance with the pres - ioAnniDeS & D iMAKi (1996), C ATTAneo ence of water, soft substrate (for nesting) (2003). and warmer habitats. its altitudinal range is accordingly (84 m ± 10 m). While only 10 Anatololacerta anatolica % of the entire dataset stemmed from (Werner , 1902) (Fig. 6) August, 47 % of the observations of this species were made in that month. Although While from the authors’ experience, a certain bias originated from the nature of seemingly less abundant than the related species-specific data collection, this seems Anatololacerta oertzeni (W erner , 1904) on to indicate that this species is one of the few other Greek islands like or , 45 more readily encountered during warmer the records (2.8 %) display a fairly wide - months. This is related to mating and breed - spread distribution across the island, in a ing activities making adults (especially wide range of rocky, often vegetated habi - males but also ground-dwelling, nesting tats. The observations fall within 30 grid females in autumn) more conspicuous, cells, corresponding to 4.6 % of the entire while in September, hatchlings add to the grid. Samos holds the only Greek popula - population density (e.g., BonS & B onS tion of this species ( VAlAKoS et al. 2008). 1960; D íAZ -P AniAGuA et al. 2002; DiMAKi Vertical distribution seems mainly, but not 2008). exclusively, related to medium and lower Museum specimens: GnHM 1-3, 19- altitudes (164 ± 27 m). 23, 78-83, 183-200, 268, 358-367, 369-411, remarks: BoeTTGer (1888) recorded 538-539, 541, 4097, 7083, 7094, 8082-8083, Lacerta danfordi (GÜnTHer , 1876) among 8085, 8087, 8091-8092, 8121, MHnG the specimens he studied from Samos. 2679.95-96, nHMAS 14-17, 7103, 8001, CAlABreSi (1923) omitted to take the for - 8003-8009, 9001, 71012, 80010-80012, mer paper into account, and assigned the 14 80014-80016, 81011, 81014-81015, nHMC specimens collected by J. C. F. MAJor to 80.3.91.5, 80.3.91.16-18, nHMW 7398, Lacerta muralis lAur . var. erhardi BeDr ., a 30987. taxon at the time known from Seriphos Bibliographic data: BoeTTGer (1888), island (e.g., SCHreiBer 1912). Werner (1930), W eTTSTein (1953), o nDri- Museum specimens: nHMW 30981: AS (1968), i oAnniDeS et al. (1994), ioAnni- 1-6, MSnF 745 e 18021-32. DeS & D iMAKi (1996), DiMAKi et al. (2000), Bibliographic data: BoeTTGer (1888), DiMAKi (2001, 2008), DiMAKi et al. (2010). CAlABreSi (1923), B ol (1992), i oAnniDeS et al. (1994), ioAnniDeS & D iMAKi (1996), Hemidactylus turcicus CATTAneo (2003). (linnAeuS , 1758) (Fig. 6) Lacerta trilineata Forty-eight observations (3.0 % of the BeDriAGA , 1886 (Fig. 6) total) were available in the collected data. These fall within 31 grid cells (4.8 % of the With 100 records (6.3 %) and 60 occu - entire grid). pied grid cells (9.3 %), this species is fairly Seemingly restricted to coastal loca - frequently sighted and widespread. While tions (41 ± 8 m) in limited parts of the 36 % of all data was collected in May and Distribution of amphibians and reptiles on Samos island 51

Fig. 6: island of Samos, distribution maps – lizards except Pseudopus apodus . Abb. 6: insel Samos, artspezifische Verbreitungskarten – echsen außer Pseudopus apodus . 52 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

June, as much as 54 % of the observations tribution pattern similar to that of the most of this species were made in these months, widespread taxa Ophisops elegans and indicating a spring activity peak. Alti - Laudakia stellio . This rather indicates sam - tudinal range is fairly wide (122 ± 18 m, pling bias than actual presence and is most from 2 to 968 m). likely due to its inconspicuous and rather Museum specimens: nHMAS 18, 7104, secretive nature. The observations fall with - nHMC 80.3.60.286, nHMW 30816:1-4, in 40 grid cells, corresponding to 6.2 % of 30988:1-3, nrM 6703. the entire grid (altitudinal range: 123 ± 24 Bibliographic data: DAAn (1967), B ol m). remarkably, 40 % of the records were (1992), i oAnniDeS et al. (1994), ioAnniD eS collected in April, whereas only 14 % of the & D iMAKi (1996), ClArK (2000), C ATTAneo entire dataset originated from that month. (2003). Museum specimens: MSnF 864, 12763-77, nHMC 80.3.82.54, nHMW Ophisops elegans 30993:1-2. MénéTrieS , 1832 (Fig. 6) Bibliographic data: CAlABreSi (1923), Werner (1930, 1935, 1938), i oAnniDeS et only Laudakia stellio and (the specif - al. (1994), ioAnniDeS & D iMAKi (1996), ically intensively sampled) Chamaeleo ClArK (2000), C ATTAneo (2 003). chamaeleon were sighted more often than this species (154 site records, 9.7 % of col - Trachylepis aurata lected data). The observations fall within 98 (linnAeuS , 1758) (Fig. 6) grid cells, corresponding to 15.1 % of the entire grid. As in L. stellio , observational While records of this species are scat - bias of sighting the same species more than tered, it is recorded infrequently (23 rec- once in limited surveyed areas seems to be ords, 1.5 %; 20 grid cells, 3.1 %; altitudinal the case. Laudakia stellio and O. elegans range: 160 ± 27 m) and is unclear to which often co-occur in dry and hot environments, extent this is due to its secretive habits, actu - the latter being the most common small lac - al rareness or other factors. While 26 % of ertid, and clearly more ground-dwelling all data was collected in May, 35 % of than Anatololacerta anatolica. The map for observations of this species were made in this species shows a broad occurrence that month, possibly suggesting a spring throughout the island, albeit with significant activity peak. gaps, most likely due to lack of sampling. Museum specimens: GnHM 7096, records are most numerous from lower lo- nHMAS 80017. cations (152 ± 15 m), but range as wide as Bibliographic data: BeuTler (1979), from 0 to 1,047 m a.s.l., the latter represent - Bol (1992), i oAnniDeS et al. (1994), ioAn- ing the highest record throughout the niDeS & D iMAKi (1996), C ATTAneo (2003). dataset. Museum specimens: MCZ r-37052, Pseudopus apodus 38505, nHMAS 19, nHMC 80.3.70.120, (PAllAS , 1775) (Fig. 7) 80.3.70.350, 80.3.70.351, 80.3.70.352, 80.3.70.421, nHMW 11921:1-2, 30819:1-5, Commonly recorded (66 records, 4.2 30967:4, 30985:4-5, yPM Herr 005801. %; 46 grid cells, 7.1 %; altitudinal range: Bibliographic data: BoeTTGer (1888), 107 ± 16 m), the map of this species high - Werner (1902, 1933), C yrén (1941), B ol lights most likely sampling intensity rather (1992), i oAnniDeS et al. (1994), ioAnniDeS than actual presence. Per sonal experience & D iMAKi (1996), C lArK (2000), C ATTAneo suggests abundance values similar as in (2003). places outside Samos. While 26 % of all data was collected in May, as much as 48 % Ablepharus kitaibelii BiBron & B ory of observations of this species were made in De SAinT -V inCenT , 1833 (Fig. 6) that month, indicating a spring activity peak. Museum specimens: GnHM 5091, This species, while observed less fre - nHMAS 29, 5004, nHMW 30962:1-2, quently (47 records, 3.0 %), showed a dis - 30998. Distribution of amphibians and reptiles on Samos island 53

Bibliographic data: Werner (1933), snake, the rather limited data availability Bol (1992), i oAnniDeS et al. (1994), ioAn- was not in contrast to what is known for niDeS & D iMAKi (1996), C ATTAneo (2003). other Greek islands (e.g., TóTH et al. 2002; rAZZeTTi et al. 2006). The data did not Typhlops vermicularis allow clear identification of patterns. Most MerreM , 1820 (Fig. 7) of the individuals have been observed on the inner side of coastal lagoons or at the border While bias may exist due to the fact of cultivated areas, as loose soils are usual - that finding this species requires turning ly preferred by this species. stones and other cover objects, this is a com - Museum specimens: GnHM 6, monly recorded snake species (44 records, nHMAS 7106, 7109, 50013. 2.8 % of data; 35 grid cells, 5.4 % of entire Bibliographic data: Bol (1992), i oAn- grid; altitudinal range: 119 ± 18 m), occur - niDeS et al. (1994), ioAnniDeS & D iMAKi ring throughout the island. While its fosso - (1996), ClArK (2000), C ATTAneo (2003). rial habits and apparent seasonal behavior (including aestivation or deeper under - Malpolon insignitus ground activity) make this species hard to (GeoFFroy SAinT -H ilAire , 1827) encounter during summer or autumn, spring (Fig. 7) sightings are rather common. As much as 89 % of records relate to observations in eighteen records (1.1 %) were avail - April or May, while these months together able to this study, which fall in 17 different hold only 40 % of the entire dataset. grid cells (2.6 % of entire grid), suggesting remarks: local folklore seemed to widespread occurrence, albeit concentrated incorrectly attribute a venomous character to in the eastern parts of the island. The few this species, as Franz Werner (1932), fifty records mainly originated from low-lying years before the first scientific record about areas (altitudinal range: 125 ± 24 m). As the the presence of a viperid snake on Samos species of this genus are known as oppor - wrote: “Das angebliche Vorkommen von tunistic and often abundant , being Giftschlan gen auf Samos bezieht sich nicht able to thrive in anthropogenic environ - auf Vipern, sondern auf den allgemein sehr ge- ments (e.g., PleGueZueloS 2003), its appar - fürchteten Typhlops vermicularis ” [The al - ent absence from much of the island could leged occurrence of venomous snakes on Sa- be related to competition with sympatric mos does not relate to vipers, but to the in gen- Dolichophis caspius (G Melin , 1789) , as eral greatly feared Typhlops vermicularis ]. both species are large diurnal, mostly terres - Museum specimens: GnHM 7992, trial snakes ( ArnolD & o VenDen 2002). nHMW 15419:4 (collected 1901), 30972:1- Since island ecological networks are simpli - 7, 30997:3-7. fied, with species broadening their niche, Bibliographic data: BoeTTGer (1888), competition can be particularly strong (e.g., Werner (1902, 1930), B ureSCH & Z onKoV liSTer 1976; C ASe & B olGer 1991; B o- (1934), W erner (1938), i oAnniDeS et al. BACK 2003). Published sightings are limit - (1994), ioAnniDeS & D iMAKi (1996), ed: ioAnniDeS et al. (1994) found a dead CATTA neo (2003). specimen near Karlovasi and an additional specimen near Mytilini, while CATTAneo Eryx jaculus (2003) found two specimens near Maratho- (linnAeuS , 1758) (Fig. 7) kampos. From the latter locality, one ani - mal was also collected in 1979 by H. With first records as recent as 1988 MAliCKy and donated to the nHMW and 1989 ( Bol 1992; i oAnniDeS et al. (27588). The 18 presently evaluated data 1994), this was one of the more infrequent - add strongly to the available records. note - ly recorded species on Samos (24 records, worthy is that 14 of these relate to 1.5 % of data; altitudinal range: 148 ± 25 found as traffic victims. m). interestingly, nearly all records fell in Museum specimens: GnHM 36-38, different grid cells, 23 of which are occu - 10971, nHMAS 4109, 5009, 7009, 50010, pied (3.5 %). Being a secretive burrowing nHMW 27588. 54 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Fig. 7: island of Samos, distribution maps – snakes part 1 and Pseudopus apodus . Abb. 7: insel Samos, artspezifische Verbreitungskarten – Schlangen Teil 1 und Pseudopus apodus . Distribution of amphibians and reptiles on Samos island 55

Bibliographic data: ioAnniDeS et al. 50011, 50018, 50020, 70014, 71011, nHMC (1994), ioAnniDeS & D iMAKi (1996), CATTA - 80.3.117.25, nrM 6702. neo (2003). Bibliographic data: Werner (1933), Natrix natrix BureSCH & Z onKoV (1934), C lArK (1968, (linnAeuS , 1758) (Fig. 7) 2000), S CHÄTTi (1988), B ol (1992), i oAnni- DeS et al. (1994), ioAnniDeS & D iMAKi As amphibians often build up a signif - (1996), CATTAneo (2003). icant portion of its diet, this snake is usually (but not always) recorded near water. Eirenis modestus Natrix natrix is among the most readily (MArTin , 1838) (Fig. 7) recorded species in other parts of Greece and Turkey (e.g., VAlAKoS et al. 2004), yet This was, from the data analyzed, the on other islands the species appears to be second most recorded and widespread snake rare and localized (e.g., - BAier et species (82 records, 5.2 %; 65 grid cells, al. 2009). records for Samos seem rather 10.0 %), mirroring its mainland abundance infrequent from the data processed (30 (e.g. FrAnZen et al. 2008). The species records, 1.9 %; 18 grid cells, 2.8 %). seems to display a rather wide altitudinal observation sites are mainly located in the range (153 ± 18 m). southeast of the island, which might be remarks: From the “Samos region”, related to a relatively higher sampling effort WeTTSTein (1937) described Contia modes - in that area, as well as a locally higher avail - ta werneri from Alazonisi on Fourni island. ability of suitable habitat. As to be expect - This taxon was later considered invalid by ed from multiple records near low-lying BArAn (1976). wetlands and other water sources, the col - Museum specimens: GnHM 8, 4942, lected data for this species is mainly restrict - 6942, 7993, MCZ r-37005-37006, ed to low elevations (55 ± 13 m). nHMAS 21, 25, 5102, 7002, 7101-7102, Museum specimens: GnHM 7082, 7108, 8002, 50015, 50019, 70010, 70013, 7084, nHMA S 5003, 7005, nHMC 80013, nHMW 30971:1-4, 31000:1-4, 80.3.34.7. nrM 6705, 6706, 6708, yPM Herr Bibliographic data: Werner (1933), 005751. ClArK (1968), B ol (1992), i oAnniDeS et al. Bibliographic data: BoeTTGer (1888), (1994), ioAnniDeS & D iMAKi (1996), C lArK CAlABreSi (1923), W erner (1902, 1933), (2000), C ATTAneo (2003). ClArK (1968), B ol (1992), i oAnniDeS et al. (1994), ioAnniDeS & D iMAKi (1996), Dolichophis caspius CATTA neo (2003). (GMelin , 1789) (Fig. 7) Hemorrhois nummifer This is the most often recorded snake (reuSS , 1834) (Figs. 7, 8) species in the dataset analyzed (106 records, 6.7 %; 67 grid cells, 10.3 %), in concor - The present data includes what is most dance with the authors’ experience from likely the first record of this species on the other parts of southeast europe, including island, made available by Andreas Meyer Greece, Bul garia, Montenegro and romania and dating back to 1996. Also for the east (SPey BroeCK , unpublished data). it occurs of lesbos ( HoFSTrA 2008), through out the island. This species is fre - Samothraki and Chios ( STrACHiniS & l yM - quently recorded as a traffic victim, with 62 BerAKiS 2013 ), this species represents a records relating to such findings. Altitud - recent addition to the known fauna. inal data points towards low elevations (117 on some islands, this species could be ± 13 m). difficult to detect, as according to SCHÄTTi Museum specimens: GnHM 7, 33-34, & S iGG (1989), from Cypriot data, H. num - 39, 4943, 5002, 6941, 6984-6985, 7001, mifer has reclusive habits, even though it 7095, 7994, 8084, 70014, MCZ r-37003, can be locally abundant. nHMAS 22, 5001, 5006, 5008, 5101, 6001, observers suggested that this species 7007, 7107, 8101, 41010-41011, 41013, might be easier to record in autumn than in 56 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Fig. 8: Hemorrhois nummifer (reuSS , 1834), from Samos, found and photographed by first author (JS), autumn 2009. Abb. 8: Hemorrhois nummifer (reuSS , 1834) von Samos, gefunden und fotografiert vom erstautor (JS), Herbst 2009.

Fig. 9: island of Samos, distribution maps – snakes part 2. Abb. 9: insel Samos, artspezifische Verbreitungskarten – Schlangen Teil 2. Distribution of amphibians and reptiles on Samos island 57

spring, whereas herpetofauna searches often (KirCHner 2009). Altitudinal distribution focus on the spring months April and May. seems relatively wide (195 ± 31 m, from 8 in this regard, it deserves mentioning that to 695 m a.s.l). five (55 %) of the available records (9 Museum specimens: GnHM 40, records, 0.6 %; 7 grid cells, 1.1 %) stem 4941, 9091, nHMAS 5002, 41012, 50016- from autumn observations. This could be 50017, 81012, nHMC 80.3.38.36, nHMW related to the dispersion of juveniles. As 30969. eight out of nine records originate from the Bibliographic data: DAAn (1967), central southern part of the island, a 2011 ioAnniDeS et al. (1994), ioAnniDeS & D i - western observation from near Agia Kiriaki MAKi (1996). stands out. The limited available data show an average altitude which is, together with Zamenis situla that of Zamenis situla (l innAeuS , 1758) , the (linnAeuS , 1758) (Fig. 9) highest among all Samos herpetofauna species (237 ± 63 m). This species, known as secretive, is generally observed in relatively low abun - Platyceps najadum dances ( VAlAKoS et al. 2008) . Thus, the (eiCHWAlD , 1831) (Fig. 9) number of 13 records (0.8 %; 11 grid cells, 1.7 %) was not particularly low in relation This species is ffrequently encoun - to those of other snake species. Available tered (54 records, 3.4 %; 47 grid cells, 7.3 records occur across most of the island. As %), displaying a wide distribution in warm, such, Z. situla is possibly not particularly rocky habitats across the island. While 40 rare on Samos. While 14 % of all processed % of all data analyzed was collected in April records originated from April, 31 % of the and May, 56 % of observations of this Z. situla records fell in that same month, species were made in those months, sug - suggesting some seasonality in the chance gesting a spring activity peak. The limited to observe this species. This species was, available data point to a relatively low (to together with Hemorrhois nummifer , found medium) altitudinal distribution (103 ± 15 at the highest mean elevation (237 ± 69 m). m). Museum specimens: GnHM 6983, Museum specimens: GnHM 9118, nHMAS 20, 7001, nHMC 80.3.30.39. nHMAS 26, 5007, 7105, 50014, nHMW Bibliographic data: Werner (1933), 30968, 30999:3. BureSCH & Z onKoV (1934), i oAnniDeS et Bibliographic data: DAAn (1967), B ol al. (1994), ioAnniDeS & D iMAKi (1996), (1992), i oAnniDeS et al. (1994), ioAnniDeS CATTAneo (2003). & D iMAKi (1996), C lArK (2000), C ATTAneo (2003 ). Montivipera xanthina (GrAy , 1849) (Fig. 9) Telescopus fallax (FleiSCHMAnn , 1831) (Fig. 9) Directed searching by one of the authors (MD) may have added some bias to Since this species is nocturnal and the dataset processed (51 records, 3.2 %; 35 secretive, the number of 32 records (2.0 %; grid cells, 5.4 %), as, from the rest of the 28 grid cells, 4.3 %) is in fact relatively data, this species appeared infrequently high, in relation to species which are usual - sighted. This might, in part, explain its ly en countered much more frequently else - rather recent first reference ( nilSon & where, like N. natrix and M. insignitus . AnDrén 1986; T ieDeMAnn & G rilliTSCH Available records occur across most of the 1986; VAn WinGerDe 1986; yet, ClArK island. As such, this species is possibly not 1989 refers to a 1984 sighting of the species particularly rare on Samos. The fact that it near Marathokampos). Montivipera xanthi - is often recorded as traffic victim, with 21 na is frequently recorded as traffic victim, records relating to such findings, is also with 34 records relating to such findings. exemplified by its recent first time discov - Despite geographically biased sampling, the ery for Chios by one of the authors species seems to be more frequently 58 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA encountered in the southern half of the island, easily observed by one of the authors (er) mainly at relatively low elevation (111 ± 14 during a short survey. As noted before, the m). apparent rarity of some species on Samos Museum specimens: GnHM 4096, could be explained by interspecific compe - 6986, 8081, 9942, nHMAS 7003-7004, tition on this island, however, these hypo - 7006, 7008, 8104, nHMW 27587, 30963. theses remain to be verified. Bibliographic data: VAn WinGerDe From the three species listed above, (1986), n ilSon & A nDrén (1986), T ieDe- M. kotschyi deserves some additional atten - MAnn & G rilliTSCH (1986), C lArK (1989), tion. As described in the introduction, ioAnniDeS et al. (1994), ioAnniDeS & D iMA- uncertainty with the initial citation of this Ki (1996), C lArK (2000), C ATTAneo (2003). species from Samos ( CAlABreSi 1923), as well as the lack of any subsequent published The high herpetofauna diversity ob - records, makes its presence in need of con - served on Samos is largely the result of the firmation, if not doubtful. However, three complex paleogeography and geological specimens are listed in the catalog of the history of the island, and the Aegean region natural History Museum of Crete (nHMC in general, especially during the glaciations 80.3.85.1193, 80.3.85.1236, 80.3.85.1237). (PAniTSA et al. 2010). The sea level changes The animals were collected in 2007 at Psili during late Pleistocene and Holocene are Ammos in pitfall traps used by Dimitris particularly relevant as they exposed a large KAlTSAS in his PhD research on the beetle part of the continental shelf, thus connecting fauna of Juniperus phoenicea ecosystems in Samos and adjacent islands, including the Aegean region. Both D. KAlTSAS and , Fourni, , lipsi, and Petros lyMBerAKiS (who provided the , to the Anatolian coast ( AKSu et authors with photos of the severely desic - al. 1995; lAMBeCK 1995; F ouFoPouloS & cated specimens) see no logical reason to iVeS 1999). Since Samos is separated from assume any error in this record. Thus, this the Turkish coast by the shallow Mycale might represent the first record in over 80 Strait (with a maximum depth of less than years of the presence of this species on 20 m) it was probably still connected to the Samos, if not ever. establishing the absence mainland 9,000 years B.P. when the sea level of a species in a given area is difficult, if not was about 35 m lower than now ( AKSu et al. logically impossible, as a lack of observa - 1995). The herpetofauna of western Turkey tions for a species (even with large series of and of the entire Anatolian Peninsula is par - field data) does not necessarily imply its ticularly rich because of its geographical absence ( rAZZeTTi & S inDACo 2006). in location functioning as a bridge or barrier any case, M. kotschyi is at best rare and for species dispersal between Asia and localized on the island, although suitable europe ( SinDACo et al. 2000). Moreover, habitat seems amply available. yet, local - western Anatolia acted as a refugium during ized and poorly recorded populations of M. the Ages and later allowed the re-colo - kotschyi also exist on the large and variably- nization of northern areas during the inter - structured island of Crete (Petros lyMBe- glacial periods ( HeWiTT 1996). According - rAKiS , in litt .). Conclusively, the authors ly, these connections promoted the crossing consider the presence of Mediodactylus of species from the Anatolian mainland onto kotschyi on Samos in continued need of Samos, giving the island its ‘Asian charac - confirmation. ter’ ( PAFiliS 2010). The obvious absence of The species cited by BoeTTGer (1888) several species to be expected, including include those that were most common in the Blanus strauchi (B eDriAGA , 1884) , Medio - authors’ dataset. While this data is certain - dactylus kotschyi (S TeinDACHner , 1870) ly imperfect for scientific evaluation of and Natrix tessellata (l AurenTi , 1768) on abundance, several species appear to be Samos, is not self-evident and asks for ex- scarcer and less widespread on Samos than planation. it may be the result of secondary in many other parts of their range. regard - stochastic events of extinction, as these ing conservation issues it should be stressed species occur abundantly on the Anatolian that frequent and widespread fires affected mainland ( FrAnZen et al. 2008) and were large parts of the island during the last Distribution of amphibians and reptiles on Samos island 59 decades. in 1983, 2,000 ha of forest (ca. 4.2 Mauremys rivulata (as Mauremys caspica ), % of the island’s surface) were burned, the Anatololacerta anato lica (as Lacerta dan - 2000 fire burned about one third of the fordi ), Lacerta triline ata , Ophisops elegans , island ( eDenFelD et al. 2000; KAlABoKiDiS Ablepharus kitai belii , Laudakia stellio (as et al. 2008; Dimaki 2008), and an addition - Stellio stellio ), Chamaeleo chamaeleon , al major fire hit the island in 2010. ob - Pseudopus apodus (as Ophisaurus apodus ), viously, these frequent and repeated fires Dolichophis caspi us (as Coluber caspius ), are of particu lar, and potentially damaging Platyceps najadum (as Coluber najadum ), effect to less mobile elements of the fauna Hemorrhois nummifer (as Coluber num - including various amphibians and reptiles mifer ), Eirenis modestus (as Eirenis modes - (Fig. 10) ( SAnToS & P oQueT 2010 ). Tourist ta ), Zamenis situla (as Elaphe situla ), industry development seems to represent Telescopus fallax (as Te lescopus falax ), another threat. As tourism is an important Macrovipera xanthina (as Vipera xanthina ), economic factor in Greece, it seems not in - Eryx jaculus , Bufo viridis and Hyla arborea . frequently considered prevalent over nature Hence, since a large portion of the Samos preservation. This is consistently evident in herpetofauna is listed in one or both Habitat coastal areas. Several coastal and lowland Directive annexes, a clear legal obligation habitats (e.g., wetlands such as marshes and exists to balance human activities against lagoons) are particularly vulnerable. While protection and conservation of the diverse their flat topography makes them suitable herpetofauna of Samos. in particular, this sites for construction of e.g., houses and refers to two listed species which appear touristic facilities, their unique natural re - very rare on the island – Emys orbicularis sources often are unrecognized and neglect - and Hemorrhois nummifer . eradication of ed. This is in line with the conclusions of exogenous spe cies like Trachemys scripta BroGGi (1994), who highlighted the impor - should be considered. tance of threatened Greek island biotopes such as mouths of creeks, cisterns and pits Perspectives and mountain creeks. As these and other threats impact the This paper is not considered an end - herpetofauna of Samos, it is important to point in data acquisition, therefore, potential stress that many of the island’s amphibian contributors to the herpetofauna of Samos and reptile species are protected under the are kindly invited to get in touch with european Habitat Directive (92/43/Cee). Daniel BoHle at < [email protected] >. The list of protected species was updated on The authors of this paper are also open to november 20, 2006, after the accession of disseminate the collected data (at grid cell romania and Bulgaria to the european level resolution) to any Greek or interna - union (Council Directive 2006/ 105/eC). tional scientific initiative upon simple Four Samos reptile species are listed in the request. Annex ii “ and plant species of com - in summary, the currently known her - munity interest whose conservation requires petofauna of Samos comprises 28 species, the designation of special areas of conserva - including one that has not been published tion”: T. graeca , E. orbicu laris , M. rivulata before ( Hemorrhois nummifer ). Three addi - (as Mauremys caspica ) and Z. situla (as tional species ( Blanus strauchi , Mediodac - Elaphe situla ). in addition, 20 of the 28 tylus kotschyi and Natrix tessellata ), listed amphibian and reptile species of Samos are elsewhere for the island, require confirma - listed in Annex iV “animal and plant species tion of their presence, until which time they of community interest in need of strict pro - are disregarded as contemporary parts of the tection”: Testudo graeca , Emys orbicularis , herpetofauna of Samos.

ACKnoWleDGMenTS

in order of decreasing number of contributed Stefan Dummermuth, Maarten Gilbert, Buttle, data records, we are grateful for their contributions to Arjan van der lugt, Markus Grimm, Thomas reich, 60 J. S PeyBroeCK & D. B oHle & e. r AZZeTTi & M. D iMAKi & M. K. KirCHner & W. B euKeMA

Fig. 10: Testudo graeca linnAeuS , 1758, from Samos with obvious burn marks, photographed by third author (er). Abb. 10: Testudo graeca linnAeuS , 1758 von Samos mit offensichtlichen Brandverletzungen, fotografiert vom Drittautor (er).

Andreas Meyer, Miloslav Homolka, Peter oefinger, Havlicek). David Bird, Sergé Bogaerts, Francesca Cat- Bobby Bok, nikolai Babiniuk, rainer Kastl, Jon taneo and Mario Schweiger provided literature. Webster, Melina Baurecht, ronald laan, Michael Glaß, Curators of several natural History museums Thomas Athineou, Kristian Munkholm, Ben Verboom, provided data about reptile and amphibian specimens in Hero Moorlag, Aaron lutziger, Dieter Fritsch, Peter the collections, for which we are particularly grateful to Molz, Georg Wiesinger, Michael Zeder, Christian Massimo Capula (MCZ), Andreas Schmitz (MHnG), Koerner, Johan F. Storm and a limited number of ivan ineich (MnHn), Annamaria nistri (MSnF), Giu - unnamed data providers, as well as to those who joined liano Doria (MSnG), Stefano Scali (MSnM), Franco the authors during their field work activities (to DB: Andreone (MrSn), Petros lymberakis (nHMC) and Marita Wenzel; to er: Francesca Baccalini, Massimo Heinz Grillitsch, Silke Schweiger and richard Gemel Delfino, Anna rita Di Cerbo, roberto Sacchi; to JS and (nHMW). DB: Anniek Aerden, Tekla Boersma, Bobby Bok, Gijs Special thanks go to Andreas Meyer for allowing Damen, Gerd Dossche, Peter engelen, Jos lycops, liam the authors to publish his discovery of Hemorrhois num - russell, ilias Strachinis, lea Sylva, Jan Van Der Voort, mifer in this paper, as well as to Christoph riegler for leonard Zammit; to MKK: the Grimm family, nicolá allowing reference to his discovery of the same species lutzmann, Michael Barej, Karin Kirchner and Michael on Chios.

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DATe oF SuBMiSSion: April 22, 2013 Corresponding editor: Heinz Grillitsch

AuTHorS: Jeroen SPeyBroeCK (Corresponding author < [email protected]; jeroenspeybroeck @hotmail.com >) – research institute for nature and Forest, Kliniekstraat 25, 1070 Brussel, Belgium & Daniel BoHle – Mommsenstrasse 20, 10629 Berlin, & edoardo rAZZeTTi – Museo di Storia naturale, university of Pavia, Piazza Botta 9-10, 27100 Pavia, & Maria DiMAKi – Goulandris natural History Museum, 100, othonos St. 145 62 Kifissia, Greece & Marlene Katharina KirCHner – university of natural resources and life Sciences, Division of livestock Sciences, Gregor-Mendel-Straße 33, 1180 Wien, & Wouter BeuKeMA – Cátedra rui nabeiro, universidade de évora (CiBio), Casa Cordovil, rua Dr. Joaquim Henrique da Fonseca, 7000- 890 évora, Portugal.