Acta Herpetologica 17(2): 187-195, 2022 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.36253/a_h-11547 One site, three species, three stories: syntopy of geckoes Euleptes europaea (Gené, 1839), Hemidactylus turcicus (Linnaeus, 1758), Tarentola mauritanica (Linnaeus, 1758) in a coastal area of southern Tuscany (central Italy) Giacomo Radi1,2, Marco A.L. Zuffi1,* 1 Museum Natural History, University of Pisa, via Roma 79, I-56011 Calci (Pisa) 2 loc. Biagioni 7, 58020 Scarlino (Grosseto) *Corresponding author. Email: marco.zuffi@unipi.it Submitted on: 2021, 30th July; revised on: 2022, 10th June; accepted on: 2022, 30th June Editor: Fabio M. Guarino Abstract. Ecological aspects of syntopic geckoes were rarely addressed in the Mediterranean basin. We reported basic information on habitat use, and activity patterns of three species found in syntopy in Cala Violina site (divided in three subareas), a highly touristic beach located in southern Tuscany, central Italy, during 2009-2010. The most abun- dant species at first capture is Hemidactylus turcicus (94 individuals), while Tarentola mauritanica and Euleptes euro- paea are less represented (28 animals in both cases). Total captures and recaptures were 175. Sex ratio did not differ from 1:1 in all the species, nor sexes of adults did differ in size. Ambient temperatures did not differ in T. mauritanica and E. europaea, while were different in H. turcicus. Despite the humidity of capture sites did not vary among species, we recorded the highest number of E. europaea at 95% and H. turcicus at 62% humidity. Wind influenced negatively T. mauritanica and H. turcicus presence, not on E. europaea. Higher observation rate took place between 21:00 and 22:00. After 23:00, only Euleptes was active. Height from the ground was different only in H. turcicus. General Linear Models showed that interaction substrate-height at capture was important for Euleptes, not for the other two species. Along the area, E. europaea was more concentrated in the northern patch, while T. mauritanica and H. turcicus dis- tributed more homogeneously. We suggest limitation of human presence for conservation purposes. Keywords. Syntopy, geckoes, Tarentola, Hemidactylus, Euleptes, central Italy. INTRODUCTION The study of the ecology of the Italian reptile species is particularly advanced for some groups, especially for tortoises (e.g., Chelazzi and Carlà, 1986; Rugiero and Lui- selli, 2006), pond turtles (e.g., Rovero and Chelazzi, 1996; Lebboroni and Chelazzi, 1998; Zuffi et al., 2004, 2007), as well as for snakes (e.g., Luiselli et al., 1996; Zuffi, 2008; Zuffi et al. 2009; Scali et al., 2011) and lizards (e.g., Perez- Mellado and Corti, 1993; Sacchi et al., 2007; Salvidio and Oneto, 2008; Bombi et al., 2009; Zuffi et al., 2011, 2012). Most information about Italian lacertilia sensu lato has been provided in Atlases and Distributive Maps (Vanni and Nistri 2006; Corti et al., 2011), despite quite anecdotal and descriptive. On the contrary, complete and scrutinized data concern phylogeographic and taxonomic features (Harris et al., 1998; Oliverio et al., 1998; Gamble et al., 2008), and, partially, ecological-behavioural fea- tures (Vervust et al., 2007; Biaggini et al., 2009; Marsili et al., 2009; Sacchi et al., 2015; Scali et al., 2016). However, research considering comparative aspects in different rep- tile species are relatively limited (i.e., Capula, Luiselli and 188 Giacomo Radi, Marco A.L. Zuffi Rugiero, 1993; Capula and Luiselli, 1994; Carvalho Jr et al., 2008; Gordon et al., 2010; Maura et al., 2011; Simbula et al., 2019) and further studies are strongly needed. There are four gecko species in Italy (Corti et al., 2011): Tarentola mauritanica and Hemidactylus turcicus, distributed in most of the Mediterranean coastal environ- ments, and Euleptes europaea and Mediodactylus kotschyi more localised, in western Mediterranean Italy and in south-eastern Italy (Apulia), respectively. Although the distribution of Tarentola, Hemidactylus and Euleptes is to some extent overlapped in north-western Italy, namely in coastal Tuscany (Vanni and Nistri, 2006), the only site where the three species actually occur in sympatry and in syntopy is in Southern Tuscany, in the municipality of Scarlino, province of Grosseto (Radi, 2013). On average, available data on Euleptes europaea refer to a few sites only in Liguria (Tinetto, Tino, Genoa), Sardinia (Sassari, Gallura) and Tuscany (Castiglione della Pescaia and the Tuscan Archipelago) and regard morphology and popu- lation dynamics (Salvidio and Delaugerre, 2003; Salvid- io and Oneto, 2008; Salvidio et al., 2011, for a review). Ecological data on Hemidactylus turcicus and Tarentola mauritanica of Italy are quite descriptive (Capula and Luiselli, 1994; Luiselli and Capizzi, 1999; Aprea et al., 2011; Zuffi et al., 2011) with the exception of a few stud- ies on sympatric geckoes in Italy and Croatia (Lisicic et al., 2012; Simbula et al., 2019). Our research, as far as we are aware, is likely the first one aimed at comparing three Gekkota species in sympatry and syntopy, and analysing and comparing biometrical features, population structure and ecology patterns in a quite unique zoogeographic context. MATERIAL AND METHODS Study area Study area is in Cala Violina, municipality of Scar- lino (province of Grosseto), which is a highly frequent- ed touristic place during summer. This area extends for 1000 × 300 m and falls within the “Monte d’Alma” 108 SIR (Sito di Interesse Regionale, Regional Interest Site; IT51A0008), and pSIC (Sito di Interesse Comuni- tario, EU Interest Site; Natura 2000 IT51A0008) and the A.N.P.I.L. “Costiere di Scarlino” (Area Naturale Protet- ta di Interesse Locale, Protected Natural Area of Local Interest) (42.856850°N, 10.774386°E) (Fig. 1). We have focused the field activity on the maximum area exten- sion, which is about 670 m long sector; the area is char- acterized by a central sandy part. Proceeding towards the far ends of the promontories, that are quite high and boulder-like, the sandy part gradually changes into coarse soiled sandstone cliffs. Going down from the shore to the Mediterranean scrub it is possible to find dissolved bedrock and sandstone slopes, transitional environments in which geckoes live. Climate is Mediterranean, with average rainfall of 600-800 mm during winter, and aver- age temperatures of 14 °C (Selvi and Stefanini, 2005). Specifically, in Follonica, the closest meteorological sta- tion to the study area, average rainfall and temperatures are 655.2 mm and 15.7 °C respectively (Barazzuoli et al., 1993). Sampling and measurements Sampling was carried out with censuses during two annual sessions, in 2009, from 23rd July to 22nd Novem- ber and in 2010, from 07th April to 30th August. We did 20 sampling days in 2009 (18 out of 20 during the night) and 22 in 2010 (20 out of 22 during the night) for 42 sampling days. Each sampling lasted five hours on aver- age for a total of 210 hours of field night sampling. Sam- pling occurred from 20:00 to 02:00 solar hour to avoid touristic disturbance and to match species’ activity. The area is naturally divided in three sectors T1, North, ca 220 m long T2, central, ca 70 m long, and T3 south, ca 350 m long, by two forest tracks (the first 24 m wide, the second 4 m wide) loading to the beach from the forest, for a total of 670 m transect length (Fig. 2). We have con- sidered the three sectors as a unique survey area. We captured geckoes by hand, or with a noose on a long stick, and placed in cotton sacks before data recording. At each capture, we registered solar time, substrate type (sandstone, loose ground, boulders, sand, vegetation), ambient temperature, humidity (thermo- Fig. 1. Satellite picture (source: Google Earth) of Follonica Gulf delimited by Piombino promontory (LI) to the North, and by Punta Ala promontory (GR) to the South. Cala Violina is pointed by a white pin. 189Three geckoes syntopy on coastal Tuscany hygrometer HANNA, HI9565, precision 0.1 °C, 0.1% humidity), wind (with the empirical Beaufort scale), animal position (height from the ground, distance from transect starting point). Morphological data were snout to vent length, tail length, head length, width and height, eye diameter, distance between eye and nostril, internarial distance, interorbital distance. We also determined size and sex class, as described in full by Atzori et al. (2007). We therefore considered males and females, juveniles (medium size, unsexable) and new-borns (very small size, unsexable). Geckoes were marked with acrylic water pens for short term recog- nition and with a cut of coded sub-digital scales (see Atzori et al., 2007) for a long-term recognition. We did not apply sub-digital marking to the new-borns of all the three species due to their markedly small size, and to the adults of the European leaf-toad gecko, due to the extremely reduced size of lamellae and very thin fingers. We assessed female reproductive status by manual pal- pation and, in some cases, using a direct light placed on the female vent, to detect eggs for transparency. In the whole period, we captured 175 geckoes: 110 Hemi- dactylus, 34 Euleptes and 31 Tarentola. We excluded recaptures and visual encounters from this research. We therefore analyzed 159 unique gecko records, 100 of which were Hemidactylus (59 in 2009, 41 in 2010), 31 were Euleptes (21 in 2009, 10 in 2010), and 28 Tarentola (14 in 2009, 14 in 2010). We defined three age classes: males, females, and juveniles. We considered juveniles and new-borns together as juvenile category. We tested sex-ratio differences within and among species using a log-linear model (with binomial distribution). We tested differences in size and biometry (all variables were nor- mally distributed, Kolmogorov-Smirnov test, P > 0.05) with a General Linear Model (multivariate GLM, with species and sex as fixed factors and their interaction). We applied this analysis only to the adults. Given all environmental variables, we used a mul- tivariate GLM to test if species do differ in some way between years and among them. In addition, to describe ecological relationships among gecko species, we applied a Principal Component Analysis (Varimax Procedure, eigenvalue ≥ 1, rotated matrix), to all environmental variables and two biometrical features (SVL, body mass), extracting the most correlated variables within each main component. Therefore, we were able to describe the com- ponents driving the average ecology of the three species of geckos. To analyse the spatial distribution of captured and observed geckoes, we normalized the three subsec- tors, creating a unique transect. We also considered the width of the two tracks used to reach the beach: 26 and 4 metres, respectively. Temperature, humidity, wind presence and hourly distribution of captures were not normally distributed, and we therefore considered them in GLM and PCA analyses. We carried out univariate and multivariate anal- yses with SPSS 20.0 release. RESULTS We sampled 31 Euleptes (17 males, eight females and six juveniles), 100 Hemidactylus (35 males, 36 females and 29 juveniles, and 28 Tarentola (nine males, seven females and 12 juveniles). Sex ratio of adults (juveniles were excluded) did not differ from 1:1 in each of the three species (Wald test = 0.785, df = 1, P = 0.376). The three species are markedly different for all con- sidered biometric features (Table 1). Euleptes is the small- est, Hemidactylus is intermediate and Tarentola the larg- est (all with P values < 0.0001, but inter-nasal P = 0.015). They do not show sexual difference (P values from 0.093 for head width to 0.663 for SVL) nor sex × species inter- action (P values from 0.130 for head width to 0.695 for inter-orbital). All statistics are reported in Supplementary Table 1. Ecological variables recorded at each gecko capture (Table 2) showed differences in most cases: multivari- ate GLM showed a marked difference between 2009 and 2010 for wind (F2,131 = 5.807, P = 0.017) and humidity (F2,131 = 36.593, P < 0.0001). Species differed in site posi- tion (F2,131 = 4.011, P = 0.020), height from the ground (F2,131 = 11.670, P < 0.0001) and hour of capture (F2,131 = Fig. 2. Satellite image (Google Earth) of Cala Violina. White lines show three transects (T1, T2, T3), A1 and A2 indicate access to the beach. 190 Giacomo Radi, Marco A.L. Zuffi 8.587, P < 0.0001). Year × species interaction was signifi- cant only for humidity (F2,131 = 3.781, P < 0.025). PCA (sampling adequacy = 0.551; sphericity Bartlett test = 424.286, P < 0.0001) extracted four main compo- nents, explaining about 68% of total variance (Table 3). The rotated matrix showed the first component describ- ing species and body size, the second describing period, site position, ground type and height from the ground, the third describing wind and humidity, the fourth describing temperature and ground type (Table 4). The distribution of the three species of geckos accord- ing to four components are shown in figures 3-5. Figure 3 shows that smaller geckoes, as Euleptes and smaller Hemidactylus and Tarentola, tend to be distributed to the northern part of the study area. Figure 4 shows that Euleptes is associated to low or no wind but with high humidity, while the other two species are more associ- ated to a relative absence of humidity. Figure 5 shows that Table 1. Average in mm and grams ± 1 SD of selected variables for each gecko species. Variable Taxon Average SD Head length Euleptes 10.230 0.312 Hemidactylus 13.996 0.173 Tarentola 16.419 0.367 Head width Euleptes 6.809 0.191 Hemidactylus 9.251 0.106 Tarentola 11.767 0.224 Head height Euleptes 3.471 0.126 Hemidactylus 5.561 0.070 Tarentola 6.965 0.149 Eye diameter Euleptes 2.036 0.059 Hemidactylus 2.701 0.033 Tarentola 3.012 0.069 Nostril eye Euleptes 2.795 0.087 Hemidactylus 3.695 0.048 Tarentola 5.010 0.102 Inter-nasal Euleptes 1.735 0.052 Hemidactylus 1.863 0.029 Tarentola 1.964 0.061 Inter-orbital Euleptes 4.567 0.132 Hemidactylus 4.702 0.073 Tarentola 6.630 0.155 bmass Euleptes 1.292 0.235 Hemidactylus 3.152 0.130 Tarentola 5.208 0.276 SVL Euleptes 38.939 1.086 Hemidactylus 50.534 0.601 Tarentola 55.377 1.276 Table 2. Variability of ecological variables recorded at gecko cap- ture. Wind in m/sec, umidity in %, site position and Hmslm in m, Hour as in hours (solar time). Variable Species Average ± SD Sample Wind Euleptes Hemidactylus Tarentola 0.28 ± 0.45 0.22 ± 0.45 0.16 ± 0.37 29 77 25 Umidity Euleptes Hemidactylus Tarentola 77.38 ± 17.70 69.66 ± 18.17 72.20 ± 20.60 29 77 25 Site position Euleptes Hemidactylus Tarentola 157.21 ± 45.14 119.65 ± 69.67 119.24 ± 60.10 29 77 25 Hmslm Euleptes Hemidactylus Tarentola 2.74 ± 1.15 1.61 ± 1.04 1.72 ± 1.05 29 77 25 Hour Euleptes Hemidactylus Tarentola 17:55 ± 08:13 21:22 ± 00:32 21:14 ± 00:17 29 77 25 Table 3. First four principal components explaining about 68% of total variance. Component Eigenvalues Weights of rotated factors total % variance % cumulated total % variance % cumulated 1 2.653 26.533 26.533 2.405 24.048 24.048 2 1.687 16.873 43.407 1.705 17.053 41.101 3 1.359 13.592 56.999 1.377 13.767 54.868 4 1.105 11.052 68.051 1.318 13.183 68.051 Table 4. Rotated component matrix and main contribution of each variable (in bold) to each component. Variable Component 1 2 3 4 species 0.679 0.157 0.074 -0.270 month -0.202 0.622 0.206 -0.008 wind -0.179 0.365 -0.771 -0.087 T° 0.124 -0.041 -0.032 0.853 umidity -0.225 0.264 0.817 -0.187 siteposition -0.228 -0.717 0.208 -0.099 ground -0.026 0.518 -0.073 0.586 hmslm -,484 -0.548 0.077 0.078 bmass 0.898 -0.022 -0.049 0.199 SVL 0.844 -0.076 -0.089 0.276 191Three geckoes syntopy on coastal Tuscany Euleptes and Hemidactylus are active at lower tempera- tures and on similar substrate ground than Tarentola. Multiple histograms show the occurrence of captures of the three species along the normalized transect (670 m long). Red bars indicate and delimitate the accesses to Cala Violina (Fig. 6). All the species showed a different distribution, Taren- tola and Hemidactylus were quite homogeneous, while Euleptes was more concentrated in the first patch of the area (transect 1). Spatial distribution was not normal (Kolmogorov Smirnov Z = 2.149, P < 0.001), and the observed differences in spatial distribution were signifi- cant (P = 0.007). DISCUSSION The gecko community in this area shows a sex-ratio not differing from the expected 1:1, for all the species considered. Average body size of the leaf-toad gecko and of the Turkish gecko falls within the range of the spe- cies in Italy (Corti et al., 2011), while the Moorish gecko results smaller in size with respect to available data (Tus- cany: Atzori et al., 2007). In our studied sample, we did not find any significant sexual difference within each species. This matter may occur in some other popula- tions of Euleptes (Delaugerre, 1985), likely due to the biotic capacity of the site (Salvidio et al., 2011). Hemi- dactylus males generally present much larger heads than the females, while size is similar for the two sexes (Corti et al., 2011). In Tarentola, sexual dimorphism is sig- nificantly marked for some features: larger eye diameter and bigger head in males (more voluminous head than equal sized females) and, same SVL, lighter body mass in females (Zuffi et al., 2011). In our study area, the differ- ence between males and females may be underestimated because the number of the two sexes of juvenile age, or as young adults, probably does not present yet the strongly different sexual dimorphism when considering adults. Interestingly, as pointed out by Simbula et al. (2019) where diurnal vs nocturnal Tarentola were considered, sexual dimorphism was not significant, a similar mat- ter as occurred in our population. Furthermore, noctur- Fig. 3. Geckos distribution along the Component 1-Component 2 relationship. Fig. 4. Geckos distribution along the Component 1-Component 3 relationship. Fig. 5. Geckos distribution along the Component 1-Component 4 relationship. Fig. 6. Capture frequency distribution of target species in the study area. Red lines represent the borders of forest tracks to the beach. 192 Giacomo Radi, Marco A.L. Zuffi nal individuals attained a smaller body size than diurnal ones (Simbula et al., 2019), suggesting an analogous pat- tern also in our sample. Body size did differ for almost all parameters among the species. We have recorded, on average, a different association of species as regards site position, height from the ground and hour of observation. Between the two years humid- ity and wind were different, with a significant interaction among species and year for humidity only. To date, ecology studies on these three species of geckoes are relatively scarce, covering trophic ecology (Capula and Luiselli, 1994; Luiselli and Capizzi, 1999; Hòdar et al., 2006), or underlining competition for spa- tial niches in sympatric populations of Hemidactylus and Tarentola in Croatia (Lisicic et al., 2012), and on lizard and geckoes’ community in central Italy (Simbula et al., 2019). Overall, our research is among the very few field works on species assemblages and, actually, it is the first work on comparative ecology of these three species of geckoes in condition of syntopy. Tarentola seems to prevail in the northern portion of the area, while Hemidactylus is more common in the northern and southern portion, as supported by GLM and Chi square analyses. According to our data, wind and humidity had more effects on Tarentola and on Hemidactylus respectively (Table 2), the species are dif- ferently distributed in the area (e.g., site position) and placed at different heights from the ground, Euleptes being relatively higher than the other two species, with a different hour distribution of observation. However, in accordance with data and analyses, the three species seem to occupy and frequent the area not in a markedly differ- ent way. PCA results showed a different distribution of age classes (as size component) relatively to the site posi- tion especially for Hemidactylus and Tarentola (Fig. 3) and a relatively importance of wind, humidity, tempera- ture and ground type for geckoes observation (Figures 4-5). The whole scenario is in accordance with previ- ous ecological observations (see for instance Delaugerre, 1984; Simbula et al., 2019). The similar occurrence of species along the transect despite the slight, significant, differences among them in many ecological parameters, resembles the pattern found and underlined by Simbula et al. (2019). Specifically, the “The observed overlap in spatial resource use was higher than expected by chance, thus showing a shared resource use instead of a partitioning pattern” (Simbula et al., 2019). For what we know so far, Cala Violina is the only syntopy area for Euleptes europaea, Hemidactylus turcicus and Tarentola mauritanica, and it arises a pivotal impor- tance for the conservation of the three species. On aver- age, we must stress that human presence in this area, as a marked touristic presence for most of the geckoes active season, is on one side undoubtedly a risk for the three species. On the other side, according to recorded data, it is not possible yet to assert if studied populations of the three species have suffered numerical losses because of direct anthropic actions (killings, capture, and removal) or indirect (alteration or damage to habitat, increment of tourism), because there are not yet studies underlining the risk factors. Locally, arsons, inappropriate woodcutting and touristic impacts are the strongest risk factors in the study area. The touristic activity on the seaside probably plays a negative role on daily activity of Tarentola mauri- tanica. In fact, the study area is characterised by intensive daily seaside activity in the spring-summer seasons (from June to September). Cala Violina is featured in S.I.R. (Site of Regional Interest) 108 “Monte d’Alma” (IT51A0008), in homonym pS.I.C. (cod. nature 2000 IT51A0008), and in A.N.P.I.L. (Area Naturale Protetta di Interesse Locale, “Protected Natural Area of Local Interest”) “Costiere di Scarlino”, but the actual presence of Euleptes europaea justifies the opportunity to propose the realization of a Biotope. The “Rete Ecologica Regionale” (Ecological Regional Network), together with the “Piano Territori- ale di Coordinamento” (PTC, “Territorial Coordinational Plan) applicable in the Grosseto Province, represent two essential instruments enforcing the importance of envi- ronmental connectivity and natural resources protection. Biotopes, in fact, are an innovative aspect in the field of planning management, to define further restrictions con- gruent to the latest acts of territory planning. In a conser- vation and management planning for this area, priority should be given to intervention finalized to protect slopes and cliffs in which these geckoes live and spent most of their life history traits. Protective intervention for slopes and cliffs is desirable to avoid further human disturbance, and particularly to avoid damages to natural fissures and crevices (abandon of garbage and other objects in fissures, destruction of portions of loose soil by sun beds, chairs and other), likely used for egg deposition, and surely for daily hideaway. These protective measures could be made up by wood barriers adequately distant from sides (0.5 – 1 m), which do not consist of a landscape obstacle, and with explicative posters making visitors aware of the site pecu- liarities, fauna richness and of all the protection meas- ures. Besides, it would be appropriate to evaluate through a dedicated study how much touristic impact influences these geckoes and other herpetofauna activity in the area, and to limit, if necessary, the daily access to Cala Violina with a maximum number of people per day (a maximum of 700/day since the last year; P. Biagini pers. comm). Measure of human disturbance on a lizard community has been recently carried out in Spain on a population of 193Three geckoes syntopy on coastal Tuscany wall lizard (Podarcis muralis), in a strongly touristic area (Amo, Lopez and Martin, 2006), where the authors point- ed out that tourism had harmful effects on physical condi- tion and relations host-parasite in this reptile. Therefore, it would be desirable to verify the actual situation in Cala Violina, because of this and other studies (e.g., Attum et al., 2006; French et al., 2008). ACKNOWLEDGEMENTS We thanks all friends and colleagues, which sup- ported and helped in the field: Giovanni Bencini, Marco Porciani, Nicola Destefano, Giuliano Franchi, Matteo Bencini, Lorenzo Saccucci, Emanuele Biggi, Flavio Lo Scalzo, Pietro Giovacchini, Fausto Corsi, Marco Dragon- etti, Sara Costa, Marco Balzarini, Elisa Riservato, Roberto Sacchi, Roberto Sindaco, Anna Rita Di Cerbo. Thanks to Comune di Scarlino and Complesso Agricolo Forestale Regionale “Bandite di Scarlino” (Dr. Patrizio Biagini) for Permissions to enter the protected area and to capture and handle the protected species. SUPPLEMENTARY MATERIAL Supplementary material associated with this article can be found at manuscript number 11547 REFERENCES Amo, L., Lopez, P., Martin, J. (2006): Nature-based tour- ism as a form of predation risk affects body condition and health state of Podarcis muralis lizards. Biol. Con- serv. 131: 402-409. Attum, O., Eason, P., Cobbs, G., Baha El Din, S.M. (2006): Response of a desert lizard community to habitat degradation: Do ideas about habitat special- ists/generalists hold? Biol. Conserv. 133: 52-62. Atzori, A., Berti, F., Cencetti, T., Fornasiero, S., Tambur- ini, M., Zuffi, M.A.L. (2007): Advances in methodolo- gies of sexing and marking less dimorphic gekkonid lizards: the study case of the Moorish gecko, Tarentola mauritanica. Amphibia-Reptilia 28: 449-454. Barazzuoli, P., Guasparri, G., Salleolini, M. (1993): Il cli- ma. In: La storia naturale della Toscana meridionale, pp: 141-171. Giusti, F., Ed, Monte dei Paschi di Siena, Pizzi Editore. Biaggini, M., Berti, R., Corti, C. (2009): Different pres- sures? Analysis of escape behaviour and ectoparasite load in Podarcis sicula (Lacertidae) populations in different agricultural habitats. Amphibia-Reptilia 30: 453-461. Bombi, P., Salvi, D., Luiselli, L., Bologna, M.A. (2009): Modelling correlated of microhabitat use of two sym- patric lizards: a model selection approach. Animal Biol. 59: 109-126. Capula, M., Luiselli, L., Rugiero, L. (1993): Comparative ecology in sympatric Podarcis muralis and P. sicula (Reptilia: Lacertidae) from the historical centre of Rome: What about competition and niche segregation in an urban habitat? Boll. Zool. 60: 287-291. Capula, M., Luiselli, L. (1994): Trophic niche overlap in sympatric Tarentola mauritanica and Hemidactylus turcicus: a preliminary study. Herpetol. J. 4: 24-25. Carvalho Jr., E.A.R., Lima, A.P., Magnusson, W.E., Alber- naz, L.K.M. (2008): Long-term effect of forest frag- mentation on the Amazonian gekkonid lizards, Coelo- dactylus amazonicus and Gonatodes humeralis. Austral Ecol. 33: 723-729. Chelazzi, G., Carlà, M. (1986): Mechanism allowing home range stability in Testudo hermanni Gmelin (Reptilia Testudinidae): field study and simulation. Monitore Zool. Ital. (N. ser.). 20: 349-370. Corti, C., Capula, M., Luiselli, L., Razzetti, E., Sindaco, R. (2011): Reptilia, Fauna d’Italia vol. XLV. Calderini, Bologna, XII + 869 pp. Delauguerre, M. (1984): Sur l’ecologie termique des geckos Phyllodactylus europeus, Hemidactylys turcicus et Tarentola mauritanica; rythmes d’activité, témpera- tures et activité, répartition altitudinale. - Travaus sci- entifiques du parc naturel régional de Corse 3: 96-121. Delaugerre, M. (1985): La variation géographique chez Phyllodactylus europaeus Gené (Reptilia, Sauria, Gek- konidae) - étude de la population de l’îlot Sperduto Grande (Sud de la Corse, Réserve naturelle des Iles Lavezzi). Bull. mensuel Soc. Linnéenne Lyon 54: 262- 269. French, S.S., Fokidis, H.B., Moore, M.C. (2008): Varia- tion in stress and innate immunity in the tree lizard (Urosaurus ornatus) across an urban-rural gradient. J. Comp. Physiol. B 178: 997-1005. Gamble, T., Bauer, A.M., Greenbaum, E., Jackman, T.R. (2008): Out of the blue: a novel, trans-Atlantic clade of geckos (Gekkota, Squamata). Zool. Scripta 37: 355- 366. Gordon, C.E., Dickman, C.R., Thompson, M.B. (2010): Partitioning of temporal activity among desert lizards in relation to prey availability and temperature. Aus- tral Ecol. 35: 41-52. Harris, D.J., Arnold, E.N., Thomas, R.H. (1998): Relation- ships of lacertid lizards (Reptilia: Lacertidae) estimat- 194 Giacomo Radi, Marco A.L. Zuffi ed from mitochondrial DNA sequences and morphol- ogy. Proc. R. Soc. – Biol. Sci. 265: 1939-1948. Lebboroni, M., Chelazzi, G. (1998): Habitat use, repro- duction, and conservation of Emys orbicularis in a pond system in Central Italy. In: Ponds and Pond landscapes of Europe. Procedings, International Con- ference of the pond life project, Boothby, J., Ed, Maas- tricht, The Netherlands 30th August 2nd September 1998. Lisicic, D., Drakulic, S., Herrel, A., Dikic, D., Benkovic, V., Tadic, Z. (2012): Effect of competition on habitat utilization in two temperate climate gecko species. Ecol. Res. 27: 551-560. Luiselli, L., Capizzi, D. (1999): Ecological distribution of the geckoes Tarentola mauritanica and Hemidactylus turcicus in the urban area of Rome in relation to age of building and conditions of the walls. J. Herpetol. 33: 316-319. Luiselli, L., Capula, M., Shine, R. (1996): Reproductive output, costs of reproduction, and ecology of the smooth snake, Coronella austriaca, in the eastern Ital- ian Alps. Oecologia 106: 100-110. Marsili, L., Casini, S., Mori, G., Ancora, S., Bianchi, N., D’Agostino, A., Ferraro, M., Fossi, M. C. (2009): The Italian wall lizard (Podarcis sicula) as a bioindicator of oil field activity. Sci. Total Environ. 407: 3597-3604. Maura, M., Vignoli, L., Bologna, M., Rugiero, L., Luiselli, L. (2011): Population density of syntopic, differently sized lizards in three fragmented woodlands from Mediterranean Central Italy. Community Ecol. 12: 249-258. Perez-Mellado, V., Corti, C. (1993): Dietary adaptation and herbivory in Lacertid lizards of the genus Podar- cis from Western Mediterranean islands (Reptilia: Sauria). Bonn. zool. Beitr. 44: 193-220. Radi, G. (2013): Ecologia comparata in tre generi sintopi- ci di Gekkota [Euleptes europaea (Gené, 1839), Hemi- dactylus turcicus (Linnaeus, 1758), Tarentola mau- ritanica (Linnaeus, 1758)] in una zona costiera della Toscana meridionale (Italia centrale). Unpubl. Master thesis, University of Pisa. Rovero, F., Chelazzi, G. (1996): Nesting migrations in a population of the European pond turtle Emys orbicu- laris (L.) (Chelonia Emydidae) from central Italy. Ethol. Ecol. Evol. 8: 297-304. Rugiero, L., Luiselli, L. (2006): Ecological modelling of a habitat use and the annual activity patterns in an urban populations of the tortoise, Testudo hermanni. Ital. J. Zool. 73: 219-225. Sacchi, R., Mangiacotti, M., Scali, S., Sannolo, M., Pel- litteri-Rosa, D., Bellati, A., Zuffi, M.A.L., Galeotti, P., Fasola, M. (2015): Context-dependent expression of sexual dimorphism in common wall lizards (Podarcis muralis). A test with island populations. Biol. J. Linn. Soc. 114: 552-565. Sacchi, R., Scali, S., Pupin, F., Gentilli, A., Galeotti, P., Fasola, M. (2007): Microgeographic variation of col- our morph frequency and biometry of common wall lizards. J. Zool. Lond. 273: 389-396. Salvidio, S., Delaugerre, M. (2003): Population dynamics of the European leaf-toed gecko (Euleptes europaea) in NW Italy: implications for conservation. Herpetol. J., 13: 81-88. Salvidio, S., Lanza, B., Delaugerre, M. (2011): Euleptes europaea (Gené, 1839). In: Reptilia, Fauna d’Italia vol. XLV, pp.: 258-270. Corti, C., Capula, M., Luiselli, L., Razzetti, E., Sindaco, R., Eds, Calderini, Bologna. Salvidio, S., Oneto, F. (2008): Density regulation in the Mediterranean leaf-toed gecko Euleptes europea. Ecol. Res. 23: 1051-1055. Scali, S., Mangiacotti, M., Sacchi, R., Gentilli, A. (2011): A tribute to Hubert Saint Girons: niche separation between Vipera aspis and V. berus on the basis of dis- tribution models. Amphibia-Reptilia 32: 223-233. Scali, S., Sacchi, R., Mangiacotti, M., Pupin, F., Gentilli, A., Zucchi, C., Sannolo, M., Pavesi, M., Zuffi, M.A.L. (2016): Does a polymorphic species have a “polymor- phic” diet? A case study from a lacertid lizard. Biol. J. Linn. Soc. 117: 492-502. Selvi, F., Stefanini, P. (2005): Biotopi naturali ed aree pro- tette nella Provincia di Grosseto: componenti floris- tiche e ambienti vegetazionali. Provincia di Grosseto, Unità Operativa Aree Protette e Biodiversità, Quad- erni delle Aree Protette, vol. 1. Simbula, G., Luiselli, L., Vignoli, L. (2019): Lizards and the city: A community study of Lacertidae and Gek- konidae from an archaeological park in Rome. Zool. Anz. 283: 20-26. Vanni, S., Nistri, A. (2006): Atlante degli Anfibi e dei Ret- tili della Toscana. Regione Toscana, Università degli Studi di Firenze, Museo di Storia Naturale, Sezione Zoologica “La Specola”, Firenze. Vervust, B., Grbac, I., Van Damme, R. (2007): Differences in morphology, performance and behaviour between recently diverged populations of Podarcis sicula mirror differences in predation pressure. Oikos 116: 1343-1352. Zuffi, M.A.L. (2008): Colour pattern variation in popula- tions of the European Whip snake, Hierophis viridifla- vus: does geography explain everything? Amphibia- Reptilia 29: 229-233. Zuffi, M.A.L., Casu, V., Marino, S. (2012): The Italian wall lizard, Podarcis siculus, along the Tuscanian coast of central Italy: biometrical features, and phenotypic pat- terns. Herpetol. J. 22: 207-212. 195Three geckoes syntopy on coastal Tuscany Zuffi, M.A.L., Celani, A., Foschi, E., Tripepi, S. (2007): Reproductive strategies and body shape in the Euro- pean pond turtle (Emys orbicularis) from contrasting habitats in Italy. J. Zool. Lond. 271: 218-224. Zuffi, M.A.L., Di Benedetto, M.F., Foschi, E. (2004): The reproductive strategies in neighbouring populations of the European pond turtle, Emys orbicularis, in central Italy. Ital. J. Zool. 71 (suppl. 3): 101-104. Zuffi, M.A.L., Gentilli, A., Cecchinelli, E., Pupin, F., Bon- net, X., Filippi, E., Luiselli, L.M., Barbanera, F., Dini, F., Fasola, M. (2009): Geographic variation of body size and reproductive patterns in Continental versus Mediterranean asp vipers, Vipera aspis. Biol. J. Linn. Soc. 96: 383-391. Zuffi, M.A.L., Sacchi, R., Pupin, F., Cencetti, T. (2011): Sexual size and shape dimorphism in the Moorish gecko (Tarentola mauritanica, Gekkota, Phyllodactyli- dae). North-West. J. Zool. 7: 189-197. Acta Herpetologica Vol. 17, n. 2 - December 2022 Firenze University Press Cryptic diversity in pygmy chameleons (Chamaeleonidae: Rhampholeon) of the Eastern Arc Mountains of Tanzania, with description of six new species Michele Menegon1,2,*, John V. Lyakurwa3,4, Simon P. Loader5, Krystal A. Tolley6,7 Preliminary genetic characterisation of Southern Smooth Snake Coronella girondica (Serpentes, Colubridae) populations in Italy, with some considerations on their alpine distribution Matteo R. Di Nicola1, Raffaella Melfi2, Francesco P. Faraone3,*, Daniel L. N. Iversen4, Gabriele Giacalone5, Giovanni Paolino1, Mario Lo Valvo6 Species diversity and distribution of amphibians and reptiles in Sardinia, Italy Claudia Corti1,2,*, Marta Biaggini1, Valeria Nulchis2, Roberto Cogoni2, Ilaria Maria Cossu2, Salvatore Frau4, Manuela Mulargia2, Enrico Lunghi2, Lara Bassu2. The Italian wall lizard, Podarcis siculus campestris, unexpected presence on Gorgona Island (Tuscan Archipelago) Marco A.L. Zuffi1,*, Alan J. Coladonato2, Gianluca Lombardo3, Antonio Torroni3, Matilde Boschetti1, Stefano Scali4, Marco Mangiacotti2, Roberto Sacchi2 Molecular analysis of recently introduced populations of the Italian wall lizard (Podarcis siculus) Oleksandra Oskyrko1,2,*, Lekshmi B. Sreelatha1,12,13, Iolanda Silva-Rocha1, Tibor Sos3,4, Sabina E. Vlad5,6,7, Dan Cogălniceanu5,6, Florina Stănescu6,7,8, Tavakkul M. Iskenderov9, Igor V. Doronin10, Duje Lisičić11, Miguel A. Carretero1,12,13 Sunny-side up: ontogenetic variation in egg mass temperatures of the wood frog Rana sylvatica Ryan Calsbeek*, Ava Calsbeek, Isabel Calsbeek Ecological niche differentiation in the Anatolian rock lizards (Genus: Anatololacerta) (Reptilia: Lacertidae) of the Anatolian Peninsula and Aegean Islands Mehmet Kürşat Şahin1,*, Kamil Candan2,3, Danae Karakasi4, Petros Lymberakis4, Nikos Poulakakis4,5,6, Yusuf Kumlutaş2,3, Elif Yıldırım2,3, Çetin Ilgaz2,3 Occupancy and probability of detection of the introduced population of Eleutherodactylus coqui in Turrialba, Costa Rica Jimmy Barrantes-Madrigal1,*, Manuel Spínola Parallada1, Gilbert Alvarado 2, Víctor J. Acosta- Chaves3,4. One site, three species, three stories: syntopy of geckoes Euleptes europaea (Gené, 1839), Hemidactylus turcicus (Linnaeus, 1758), Tarentola mauritanica (Linnaeus, 1758) in a coastal area of southern Tuscany (central Italy) Giacomo Radi1,2, Marco A.L. Zuffi1,* Comparative cytogenetics on Zamenis lineatus and Elaphe quatuorlineata (Serpentes: Colubridae) Marcello Mezzasalma1,* , Elvira Brunelli1, Gaetano Odierna2, Fabio M. Guarino2