Acta Herpetologica 12(1): 49-54, 2017 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-18354 Feeding ecology of two sympatric geckos in an urban area of Northeastern Brazil José Guilherme G. Sousa1, Adonias A. Martins Teixeira2,*, Diêgo Alves Teles2, João Antônio Araújo-Filho2, Robson Waldemar Ávila3 ¹ Programa de Pós-Graduação em Ecologia e Recursos Naturais, Departamento de Ciências Biológicas, Universidade Federal do Ceará, Campus Universitário do Pici, CEP 60021970, Fortaleza, Ceará, Brazil ² Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento de Sistemática e Ecologia – DSE, Centro de Ciências Exatas e da Natureza – CCEN, Universidade Federal da Paraíba – UFPB, Cidade Universitária, Campus I, CEP 58059-900, João Pes- soa, PB, Brazil. *Corresponding author. E-mail: adoniasteixeira01@gmail.com ³ Laboratório de Herpetologia, Departamento de Ciências Biológicas, Universidade Regional do Cariri, Rua Cel. Antônio Luiz, 1161, Campus do Pimenta, 63105-000, Crato, Brazil; and Departamento de Ciências Biológicas, Universidade Federal do Ceará, Campus Universitário do Pici, CEP 60021970, Fortaleza, Ceará, Brazil Submitted on 2016, 31st May; revised on 2016, 11th November; accepted on 2016, 16th December Editor: Sandra Hochsheid Abstract. The diets of two sympatric gecko species, Hemidactylus mabouia and Phyllopezus pollicaris, were studied from an urban area of the Crato municipality, Northeastern Brazil. While the house gecko H. mabouia is an intro- duced species widely distributed in North, Central, and South America, the Brazilian gecko P. pollicaris is a native spe- cies distributed along the great diagonal of open formations of South America. The diets of both species were mainly composed by arthropods, Diptera was the most important item for both species, corroborating others studies with liz- ards in urban areas. Male and female adults of both H. mabouia and P. pollicaris use similar microhabitats which can explain the high sexual and interspecific trophic niche overlap. In these populations from an urban area of the Crato municipality, the alien H. mabouia seems to have not negatively affected the trophic niche of the native P. pollicaris. Keywords. Trophic ecology, diet composition, Gekkota, Hemidactylus mabouia, invasive lizard, Phyllopezus polli- caris, body size. INTRODUCTION Gekkota is a species-rich clade of lizards with a great potential to invade new habitats, frequently introduced in anthropogenically disturbed areas and/or urban habitats (Hanley et al., 1998; Carranza and Arnold, 2006; Gam- ble et al., 2008). Its success in colonizing and establish- ing populations in new habitats is due in part to its great plasticity, which led to a wide distribution of geckos, especially Hemidactylus, in the old and new world (Rocha et al., 1994; Hanley et al., 1998; Vences et al., 2004; Gam- ble et al., 2008; Meshaka, 2011). The tropical house gecko Hemidactylus mabouia is native to Africa and has successfully colonized South, Central, and North America (Rocha et al., 2011). In Bra- zil, H. mabouia is frequently found in human-altered are- as, but also in pristine habitats in the Amazon, Atlantic Forest, Cerrado, and Caatinga (Vanzolini, 1978; Vanzolini et al., 1980; Araújo, 1991; Rocha et al., 2000; Rocha et al., 2002; Rocha et al., 2011; Albuquerque et al., 2013). Several studies have demonstrated the role of Hemi- dactylus spp. in niche displacement of resident geckos by means of direct exploitative competition, indirect competition, aggression, and predation (Meshaka, 1995; 50 J.G.G. Sousa et alii Meshaka and Moody, 1996; Meshaka, 2000; Meshaka et al., 2005; Hoskin, 2011; Hughes et al., 2015). Buildings in Brazil are colonized by many native gecko species, but the co-occurrence with H. mabouia caused exclusion by competition in some species, such as Thecadactylus rapi- cauda (Vitt and Zani, 1997). However, Phyllopezus polli- caris, a medium-sized gecko inhabiting boulder slabs in the dry formations of South America, is also frequently found in human habitations syntopic with H. mabouia (Vanzolini et al., 1980; Vitt, 1995; Sousa et al., 2010; Recoder et al., 2012). Plasticity of feeding habits, associated with ecological and biological conditions, has been considered the main cause of H. mabouia´s invasion success (Zamprogno and Teixeira, 1998; Bonfiglio et al., 2006). Thus, diet stud- ies can provide valuable information on the importance of prey types on the mechanism by which Hemidactylus geckos interact with other lizard species (Belver and Avi- la, 2001). Herein, we present data on the diets of H. mabouia and P. pollicaris and test if the invasive species has affect- ed the native lizard, considering the trophic niche, in order to answer the following questions: i) is there sexual dimorphism between H. mabouia and P. pollicaris and between species? ii) how do the diets of these two geckos differ regarding composition? iii) what are the aver- age niche breadths and the niche overlaps between these species? iv) is niche overlap between the studied species caused by chance? MATERIALS AND METHODS Lizard specimens were collected from March to Novem- ber 2011 in human habitations in the city of Crato (7°14’S and 39°24’W, datum WGS84), Ceará state, Northeastern Brazil. The regional climate is predominantly tropical, hot, and sub-humid (average annual temperatures vary from 24 to 26 °C). The rainy period occurs between January and May, with an average annu- al rainfall of 1,091 mm (IPECE, 2013). The lizards were collected manually, humanely killed with lethal doses of 2% lidocaine, fixed in 10% formaldehyde, and preserved in 70% ethyl alcohol. The specimens were sacrificed for another study on parasitism which has been already pub- lished (Sousa et al., 2014). Snout-vent length (SVL), jaw width (JW), mouth length (ML), and head length (HL) of each lizard were measured using a digital caliper (± 0.01 mm; Table 1). Voucher specimens were deposited in the Coleção Herpetológi- ca da Universidade Regional do Cariri - URCA. To test for morphological variation due to the sex of each species, between adults of H. mabouia and P. pollicaris in body shape (using all morphometric variables), and between each individual variable, we performed a multivariate discriminant function analysis with residuals of morphological variables of each sampled lizard species. Residuals were obtained from a simple linear regression following Sousa and Ávila (2015). Dis- criminant function analysis was executed using Statistica ver- sion 10.0 (Statsoft, 2011). Lizards were dissected in the laboratory and their stomach contents were analyzed under a stereomicroscope. Prey items were identified and classified to the order level; length and width were measured with a digital caliper (precision 0.01mm). The percentage of lizards with empty stomachs was calculated and only lizards with identifiable items were used. The volume of each prey item was estimated by the ellipsoid formula: V = 4 3 π L 2 ⎛ ⎝ ⎜ ⎞ ⎠ ⎟ W 2 ⎛ ⎝ ⎜ ⎞ ⎠ ⎟ 2 , where L = length and W = width. The Importance Value Index (I) was estimated by the following formula (Powell et al., 1990): I = F%+N%+V% 3 , where F% is the relative frequency, N% is the numerical per- centage, and V% is the volumetric percentage of each prey item. Trophic niche breadth (both numerical and volumetrically) was calculated by the inverse of the Simpson’s diversity index (Simpson, 1949): B=1/ Pi 2 i=1 n ∑ , where p is the numeric or volumetric proportion of prey cat- egory i and n is the number of categories (Pianka, 1973). The value of niche breadth varies from 1 to n, where the lowest values represent a more specialized diet and the highest values indicate a generalist diet. Differences in number and volume of prey items between males and females were tested by non- parametric Mann-Whitney U test using Statistica version 10.0 (Statsoft-Inc, 2011). Trophic niche overlap between sexes and species was calcu- lated using the Pianka’s overlap index (Pianka, 1973): ∅ik = PijPik i=1 n ∑ (Pij2)(Pik2) i=1 n ∑ , where Pij and Pik are the rate consumption of prey category i, with j and k representing the compared species and sexes. Pian- ka’s overlap index varies from zero (no overlap) to one (com- plete overlap). Finally, we compared the observed niche over- lap values for H. mabouia and P. pollicaris against a null model (1,000 interactions) generated by the randomization algorithm 3 (RA3) (Lawlor, 1980). The use of RA3 seems adequate because it retains the niche breadth of each species, but randomizes which particular resource states are used, allowing the species to potentially use other resource states (Winemiller and Pianka, 1990). Niche overlaps and null models were calculated using EcosimR - R code for null model analysis (Gotelli and Ellison, 2013). Differences between proportions of each prey category among the two gecko species were tested by a non-parametric 51Feeding ecology of two sympatric geckos multivariate analysis of similarity (ANOSIM), using the Bray- Curtis similarity coefficient and 9,999 permutations in the soft- ware PAST 3.0 (Hammer et al., 2001). A similarity percentage analysis (SIMPER) was performed to determine which preys were responsible for dissimilarity in diets between the two gecko species. Data matrix for ANOSIM and SIMPER were standardized (as a percentage) to minimize the discrepancy between samples. RESULTS We collected 58 H. mabouia, 10 adult males (mean ± standard deviation) (SVL = 56.35 ± 9.76 mm), 21 adult females (SVL = 53.69 ± 6.21 mm), and 27 juveniles (SVL = 30.92 ± 5.36 mm). Of the species P. pollicaris, we col- lected 100 specimen, 38 adult males (SVL = 65.48 ± 9.06 mm), 54 adult females (SVL = 64.58 ± 12.62 mm), and 8 juveniles (SVL =34.54 ± 4.32 mm). Of these, 18 (31.03%) H. mabouia (2 males, 7 females, and 9 juveniles) and 45 (45%) P. pollicaris had empty stomachs (12 males, 27 females, and 6 juveniles). There were no significant sexual differences in mor- phometric variables of both species (Table 1). On the other hand, adults of P. pollicaris were significantly larger than H. mabouia in body shape, with SVL and JW of P. pollicaris greater than those of H. mabouia (Table 1). The diet of both species consisted of 10 arthropod prey items: Diptera, Hymenoptera, and Coleoptera were the most important items for H. mabouia and Diptera, Coleoptera, and Hemiptera for P. pollicaris (Table 2). Hemidactylus mabouia had numerical and volumetrical- ly niche breadth values of 1.904 and 1.979, respectively, which was smaller than the niche breadth of P. pollicaris: 4.05 and 3.036 (Table 2). The average number of prey items per stomach (only those individuals whose stomachs were not empty) was similar for both species: 3.86 ± 8.22 for H. mabouia and 3.47 ± 4.52 for P. pollicaris, with no significant difference (U = 1,017.5; P = 0.387). However, the average number of prey categories per stomach was 1.54 ± 0.61 and 1.24 ± 0.51 for H. mabouia and P. pollicaris, respectively, and there was a significant difference (U = 769.5; P = 0.014). Males and females of P. pollicaris ingested more prey items (4.26 ± 6.02 and 3.18 ± 3.07, respectively) than those of H. mabouia (2.2 ± 2.69 and 1.68 ± 1.21), but there were no statistical differences in prey items ingest- ed by females of P. pollicaris and H. mabouia (U= 230.5; P= 0.836) neither between males of both species (U = 32; P= 0.081). Also, there were no significant differences in number (U = 117; P = 0.385) or volume (U = 111; P = 0.3) of the prey items between H. mabouia males and females, between P. pollicaris males and females (Num- ber: U = 624.50; P= 0.323; Volume U = 605; P = 0.25), or in the number between adult individuals of both spe- cies (U = 995; P = 0.21). On the other hand, there was a significant difference in volume between adults of P. pol- licaris (211.858 ± 435.062) and H. mabouia and (36.749 ± 141.807) (U = 697; P < 0.001). Niche overlap between the sexes was 0.9159 in H. mabouia and 0.9526 in P. pollicaris; and between the two gecko species, niche overlap was 0.9094. The high niche overlap between adult individuals of both studied spe- cies could be due to chance (average overlap simulated = 0.2740; P = 0.99). There was no significant difference Table 1. Morphometric data (mean ± standard deviation) and results of discriminant function analysis. Hm= Hemidactylus mabouia; Pp= Phyllopezus pollicaris. M= Males; F= Females; Body Shape= pooled variables; SVL= Snout-Vent Lenght; JW= Jaw width; ML= Mouth lenght; HL= Head lenght; λ = Wilks' Lambda. Hm Pp Hm Pp M F M F Adults Adults SVL 53.4 ± 12.9 53.6 ± 6.8 65.9 ± 11.8 65.8 ± 11.5 55.1 ± 8.3 65.8 ± 11.6 JW 9.6 ± 2.2 9.6 ± 1.0 11.7 ± 2.1 11.8 ± 2.0 9.6 ± 1.3 11.7 ± 2.0 ML 10.2 ± 2.2 10.5 ± 1.2 12.0 ± 2.3 12.0 ± 2.0 10.4 ± 1.4 12.0 ± 2.1 HL 14.2 ± 2.5 14.6 ± 1.4 16.7 ± 3.0 16.8 ± 2.6 14.5 ± 1.7 16.7 ± 2.7 λ (M vs F) P λ (M vs F) P λ (Hm vs Pp) P Body shape 0.951 0.884 0.983 0.860 0.753 <0.0000* SVL 0.997 0.802 0.983 0.989 0.908 0.000* JW 0.965 0.385 0.994 0.361 0.835 0.001* ML 1.000 0.970 0.989 0.498 0.773 0.100 HL 0.996 0.775 0.986 0.636 0.756 0.563 52 J.G.G. Sousa et alii between the proportions in prey use (ANOSIM, R = 0.7037; P = 0.2035). The results of the SIMPER analysis also showed a small dissimilarity between diet composi- tion (32.69%), with Diptera, Hemiptera, and Coleoptera as prey groups that contribute most to dissimilarities between the two geckos, explaining 32.2, 26.7 and 11.1% of the variation. DISCUSSION The diets of the sympatric H. mabouia and P. polli- caris in urban habitats of Northeastern Brazil were very similar, with both species consuming 10 prey categories. Diptera, Coleoptera, Hemiptera, and Hymenoptera were the most frequently ingested preys; Diptera were the most important prey for both species. However, Hymenoptera and Coleoptera were the second most important prey for H. mabouia and P. pollicaris, respectively. These insects are commonly found in urban habitats, mainly close to streetlights (Robinson, 2005). Food composition of urban populations of lizards may differ from that of pristine habitats (Hódar and Ple- guezuelos, 1999). Moreover, the opportunistic behavior of these two gecko species, plus changes in diversity and abundance of prey, may contribute to differences between study sites (Zamprogno and Teixeira, 1998). Vitt (1995), studying the diet of H. mabouia and P. pollicaris at Caat- inga habitats from Exu, found that insect larvae were the most important prey for both species. In the coastal plain of the Espírito Santo state, Zamprogno and Teix- eira (1998) found Araneae, Homoptera, and Isopoda as the most important prey for H. mabouia, while Aptery- gota, Araneae, and Orthoptera were the main items for P. pollicaris at Cerrado habitats in the state of Tocantins (Recoder et al., 2012). Albuquerque et al. (2013) found cannibalistic habits for H. mabouia, where H. mabouia was the most important item, followed by Formicidae and Hemiptera; for P. pollicaris, the most important items were Coleoptera, Araneae, and Homoptera in perian- tropic environments in the state Mato Grosso do Sul. Our results corroborate the study of Bonfiglio et al. (2006), in an urban area in Rio Grande do Sul, who found that Dip- tera were the most important item in the diets of the two species studied here. That convergence in diet composi- tion may be due to the fact that Diptera is often present at urban environments, mainly close to streetlights as mentioned above. In this case, perhaps geckos could act as a control of Diptera in cities. Energetic balance in a given population of lizards can be estimated by the proportion of empty stomachs, where nocturnal lizards have a higher proportion of empty stomachs (24.1%) than diurnal lizards (10.5%); the mean percentage of empty stomachs for nocturnal geckos is 21.2% (Huey and Pianka 1981). In the present study, H. mabouia (31.01%) and P. pollicaris (42.6%) showed high- er proportions of empty stomachs than stated by Huey and Pianka (1981). This may be explained by the time Table 2. Diet composition and numerical and volumetric niche breadth of Hemidactylus mabouia and Phyllopezus pollicaris from an urban area of Crato city, Ceará state, Brazil. n= total of analyzed individuals. F = frequency of occurrence, N = number of preys, V= volume (mm³) and I = relative importance index. Category H. mabouia (n = 58) P. pollicaris (n = 100) F F % N N% V V % I F F % N N% V V % I Araneae 4 7.02 4 2.80 28.83 2.12 3.98 8 10.96 8 3.98 364.37 1.55 5.50 Blatodae - - - - - - - 1 1.37 1 0.50 51.92 0.22 0.70 Coleoptera 8 14.04 13 9.09 160.26 11.78 11.64 13 17.81 51 25.37 2605.41 11.10 18.09 Diptera 24 42.11 102 71.33 936.65 68.86 60.76 26 35.62 79 39.30 11319.23 48.15 41.05 H. mabouia shed skin 1 1.75 3 2.10 10.45 0.77 1.54 - - - - - - - Vertebrate eggshell 1 1.75 1 0.70 8.83 0.65 1.03 1 1.37 3 1.49 131.19 0.56 1.14 Hemiptera - - - - - - - 8 10.96 27 13.43 6544.06 27.89 17.43 Hymenoptera 10 17.54 10 6.99 173.73 12.77 12.44 6 8.22 15 7.46 1804.91 7.69 7.79 Isoptera 1 1.75 1 0.70 3.00 0.22 0.89 - - - - - - - Insect larvae 5 8.77 6 4.20 18.57 1.37 4.78 4 5.48 5 2.49 265.74 1.13 3.03 Lepidoptera 1 1.75 1 0.70 12.42 0.91 1.12 2 2.74 8 3.98 293.33 1.25 2.66 Ortoptera 2 3.51 2 1.40 7.49 0.55 1.82 4 5.48 4 1.99 83.53 0.36 2.61 Total 57 100 143 100 1360.24 100 100 73 100 201 100 23463.69 100 100 Niche breadth 1.90 1.98 4.05 3.04 Empty stomachs 18 45 53Feeding ecology of two sympatric geckos when the lizards were captured, i.e., in the first hours of the night. The lack of sexual differences in the diets of the spe- cies we studied is probably a result of similar use of space and time, which led to the use of the same prey types for both sexes and higher dietary overlap (see Rocha and Anjos, 2007). Dietary overlap was higher even when we paired both species (0.91), and the null model showed that high niche overlap between them is due to chance, indicat- ing that the two species may eventually compete for prey. However, P. pollicaris presented higher prey volume and niche breadth than H. mabouia (probably due to its larger body) and despite the expected, H. mabouia was apparent- ly not significantly affected by the trophic niche of P. pol- licaris. Dissimilarity between the diets of these two geckos was about 30% in the present study, but this pattern can vary according to prey availability and/or greater time of colonization by H. mabouia. These factors can make this invasive species compete more intensely, damaging the spatial and/or trophic niche of native species (see Rodder et al., 2008; Rocha et al., 2011). Additionally, further stud- ies are necessary to understand the role of competition for food and/or space in the coexistence of native and invader lizard species in disturbed and natural areas in Brazil. ACKNOWLEDGMENTS We thank the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES for providing a fellowship to JGGS, AAMT, and JAAF, the Conselho Nacional de Desenvolvimento Científico (CNPq) for a research grant to RWA (process # 303622/2015-6) and a fellowship to DAT. We also thank Sandra Hochscheid and two anonymous reviewers for their valuable sugges- tions. The specimens were collected under the license of the Instituto Chico Mendes de Conservação da Biodiver- sidade-ICMBio 29613-1. REFERENCES Albuquerque, N.R.D., Costa-Urquiza, A.D.S., Soares, M.P., Alves, L.S., Urquiza, M.V.S. (2013): Diet of two sit-and-wait lizards, Phyllopezus pollicaris (Spix, 1825) (Phyllodactylidae) and Hemidactylus mabouia (Moreau de Jonnès, 1818) (Gekkonidae) in a perian- thropic area of Mato Grosso do Sul, western Brazil. Biota Neotrop. 13: 376-381. Araújo, A.F. (1991): Structure of a white sand-dune liz- ard community of coastal Brazil. Rev. Brasil. Biol. 51: 857-865. Belver, L.C., Avila L.J. (2001): Ritmo de actividad diaria y estacional de Cnemidophorus longicaudus (Squamata: Teiidae: Teiinae) en el Norte de La Rioja, Argentina. Bol. Soc. Biol. Concepción Chile 72: 31-36. Bonfiglio, F., Balestrin, R.L., Cappellari, L.H. (2006): Diet of Hemidactylus mabouia (Sauria, Gekkonidae) in urban area of southern Brazil. Biociências (On-line) 14: 107-111. Carranza, S., Arnold, E. (2006): Systematics, biogeogra- phy, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Mol. Phylogenet. Evol. 38: 531-545. Gamble, T., Bauer, A.M., Greenbaum, E., Jackman, T.R. (2008): Evidence for Gondwanan vicariance in an ancient clade of gecko lizards. J. Biogeogr. 35: 88-104. Gotelli, N.J., Ellison, A.M. (2013): EcoSimR. Version 1.00. Computer software. http://www.uvm.edu/~ngotelli/ EcoSim/EcoSim.html. (accessed on 12-04-2012). Hammer, Ø., Harper, D.A.T., Ryan, P.D. (2001): PAST: Paleontological Statistics Software Package for educa- tion and data analysis. Palaeontol. Electron. 4: 1-9. Hanley, K.A., Petren, K., Case, T.J. (1998): An experimen- tal investigation of the competitive displacement of a native gecko by an invading gecko: no role for para- sites. Oecologia 115: 196-205. Hódar, J., Pleguezuelos, J. (1999): Diet of the Moorish gecko Tarentola mauritanica in an arid zone of south- eastern Spain. Herpetol. J. 9: 29-32. Hoskin, C.J. (2011): The invasion and potential impact of the Asian House Gecko (Hemidactylus frenatus) in Australia. Austral Ecol. 36: 240-251. Huey, R.B., Pianka, E.R. (1981): Ecological consequences of foraging mode. Ecology 62: 991-999. Hughes, D.F., Meshaka We, J.R., Van Buurt, G. (2015): The superior colonizing gecko Hemidactylus mabouia on Curaçao: Conservation implications for the native gecko Phyllodactylus martini. J. Herpetol. 49: 60-63. IPECE. (2013): Crato. Anuário Estatístico do Ceará. Per- fil Básico dos Municípios. http://www.ipece.ce.gov.br. (accessed on 15-03-2013). Meshaka We, J.R. (1995): Reproductive cycle and coloniza- tion ability of the Mediterranean gecko (Hemidactylus turcicus) in south-central Florida. Fla. Sci. 58: 10-15. Meshaka We, J.R., Moody, B.A. (1996): The old world tropical housegecko (Hemidactylus mabouia) on the Dry Tortugas. Fla. Sci. 59: 115-117. Meshaka We, J.R. (2000): Colonization dynamics of two exotic geckos (Hemidactylus garnotii and H. mabouia) in Everglades National Park. J. Herpetol. 34: 163-168. Meshaka We, J.R., Smith, H.T., Severson, R., Severson, M.A. (2005): Spatial picture of a gecko assemblage in flux. Fla. Sci. 68: 53-55. 54 J.G.G. Sousa et alii Meshaka We, J.R. (2011): A runaway train in the making: the exotic amphibians, reptiles, turtles, and crocodil- ians of Florida. Monograph 1. Herpetol. Conserv. Biol. 6: 1-101. Pianka, E.R. (1973): The structure of lizard communities. Annu. Rev. Ecol. Syst. 4: 53-74. Powell, R., Parmelee, J.S., Rice, M.A., Smith, D.D. (1990): Ecological observations of Hemidactylus brooki hai- tianus Meerwath (Sauria: Gekkonidae) from Hispan- iola. Caribbean J. Sci. 26: 67-70. Recoder, R., Junior, M.T., Camacho, A., Rodrigues, M.T. (2012): Natural history of the tropical gecko Phyl- lopezus pollicaris (Squamata, Phyllodactylidae) from a sandstone outcrop in Central Brazil. Herpetol. Notes 5: 49-58. Robinson, W.H. (2005): Urban insects and arachnids: a handbook of urban entomology. Cambridge Univer- sity Press. New York. Rocha, C.F.D., Anjos, L.A. (2007): Feeding ecology of a nocturnal invasive alien lizard species, Hemidactylus mabouia Moreau de Jonnès, 1818 (Gekkonidae), liv- ing in an outcrop rocky area in southeastern Brazil. Braz. J. Biol. 67: 485-491. Rocha, C.F.D., Dutra, G., Vrcibradic, D., Menezes, V. (2002): The terrestrial reptile fauna of the Abrolhos Archipelago: species list and ecological aspects. Braz. J. Biol. 62: 285-291. Rocha, C.F.D., Vrcibradic, D., Araújo, A., Esteves, F., Lacerda, L. (2000): Ecofisiologia de répteis de rest- ingas brasileiras. In: Ecologia de Restingas e Lagoas Costeiras, pp. 117-149. Esteves, F.V., Lacerda, L.D., Eds, NUPEN-UFRJ, Macaé. Rocha, C.F.D., Anjos, L.A., Bergallo, H.G. (2011): Con- quering Brazil: the invasion by the exotic gekkonid lizard Hemidactylus mabouia (Squamata) in Brazilian natural environments. Zoologia 28: 747-754. Rocha, C.F.D., Nascimento, L.B., Bernardes, A.T., Cotta, G.A. (1994): Introdução a  ecologia de lagartos bra- sileiros. Herpetologia no Brasil 1. Pontifícia Universi- dade Católica de Minas Gerais/PUC-MG, Belo Hori- zonte. Rödder, D., Solé, M., Böhme, W. (2008): Predicting the potential distributions of two alien invasive House- geckos (Gekkonidae: Hemidactylus frenatus, Hemidac- tylus mabouia. North-West J. Zool. 4: 236-246. Simpson, E.H. (1949): Measurement of diversity. Nature 163: 688. Sousa, J.G.G., Ribeiro, S.C., Roberto, I.J., Teles, D.A., Almeida, W.O. (2010): Ocorrência de pentastomídeos (Metameria: Ecdysozoa) no lagarto Phyllopezus polli- caris (Spix, 1825). Cad. Cult. Cienc. 2: 64-71. Sousa, J.G.G., Brito, S.V., Ávila, R.W., Teles, D.A., Arau- jo-Filho, J.A., Teixeira, A.A.M., Anjos, L.A., Almeida, W.O. (2014). Helminths and Pentastomida of two synanthropic gecko lizards, Hemidactylus mabouia and Phyllopezus pollicaris, in an urban area in North- eastern Brazil. Braz. J. Biol. 74: 943-948. Sousa, J.G.G., Ávila, R.W. (2015). Body size, reproduc- tion and feeding ecology of Pleurodema diplolister (Amphibia: Anura: Leiuperidae) from Caatinga, Per- nambuco state, Northeastern Brazil.  Acta Herpe- tol. 10: 129-134. Statsoft, Inc. (2011): Statistica for Windows [Computer program manual]. Versão 10.0. Tulsa: StatSoft. Vanzolini, P.E. (1978): On South American Hemidactylus (Sauria, Gekkonidae). Arq. Zool. Estado São Paulo 17: 01-84. Vanzolini, P.E., Ramos-Costa, A.M.M., Vitt, L.J. (1980): Réptéis das Caatingas. Academia Brasileira Ciéncias, Rio de Janeiro. Vences, M., Wanke, S., Vieites, D.R., Branch, W.R., Glaw, F., Meyer, A. (2004): Natural colonization or introduc- tion? Phylogeographical relationships and morpho- logical differentiation of house geckos (Hemidactylus) from Madagascar. Biol. J. Linn. Soc. 83: 115-130. Vitt, L.J. (1995): The ecology of tropical lizards in the caatinga of northeast Brazil. Oklahoma Museum of Natural History, University of Oklahoma. Vitt, L.J., Zani, P.A. (1997): Ecology of the nocturnal liz- ard Thecadactylus rapicauda (Sauria: Gekkonidae) in the Amazon region. Herpetologica 53: 165-179. Zamprogno, C., Teixeira, R. (1998): Hábitos alimentares da lagartixa de parede Hemidactylus mabouia (Rep- tilia, Gekkonidae) da planície litorânea do norte do Espírito Santo, Brasil. Braz. J. Biol. 58: 143-150. Acta Herpetologica Vol. 12, n. 1 - June 2017 Firenze University Press Morphological variation and sexual dimorphism in Liolaemus wiegmannii (Duméril & Bibron, 1837) (Squamata: Liolaemidae) from Uruguay Joaquín Villamil1,*, Arley Camargo2, Raúl Maneyro1 Sex does not affect tail autotomy in lacertid lizards Panayiotis Pafilis1,*, Kostas Sagonas2, Grigoris Kapsalas1, Johannes Foufopoulos3, Efstratios D. Valakos4 Fire salamander (Salamandra salamandra) males’ activity during breeding season: effects of microhabitat features and body size Raoul Manenti*, Andrea Conti, Roberta Pennati Call variation and vocalizations of the stealthy litter frog Ischnocnema abdita (Anura: Brachycephalidae) Pedro Carvalho Rocha1,2,*, João Victor A. Lacerda2,3, Rafael Félix de Magalhães2,3, Clarissa Canedo4, Bruno V. S. Pimenta5, Rodrigo Carrara Heitor6, Paulo Christiano de Anchieta Garcia2,3 Feeding ecology of two sympatric geckos in an urban area of Northeastern Brazil José Guilherme G. Sousa1, Adonias A. Martins Teixeira2,*, Diêgo Alves Teles2, João Antônio Araújo-Filho2 and Robson Waldemar Ávila3 A pattern-based tool for long-term, large-sample capture-mark-recapture studies of fire salamanders Salamandra species (Amphibia: Urodela: Salamandridae) Jeroen Speybroeck1,*, Koen Steenhoudt2,$ Skeletal variation within the darwinii group of Liolaemus (Iguania: Liolaemidae): new characters, identification of polymorphisms and new synapomorphies for subclades Linda Díaz-Fernández*, Andrés S. Quinteros, Fernando Lobo Marking techniques in the Marbled Newt (Triturus marmoratus): PIT-Tag and tracking device implant protocols Hugo Le Chevalier1, Olivier Calvez2, Albert Martínez-Silvestre3, Damien Picard4, Sandra Guérin4,5, Francis Isselin-Nondedeu6, Alexandre Ribéron1, Audrey Trochet1,2,* Evidence for directional testes asymmetry in Hyla gongshanensis jindongensis Qing Gui Wu1, Wen Bo Liao2,* The advertisement call of Pristimantis subsigillatus (Anura, Craugastoridae) Florina Stănescu1,4, Rafael Márquez2, Paul Székely3,4,*, Dan Cogălniceanu1,4,5 Nematodes Infecting Anotosaura vanzolinia (Squamata: Gymnophthalmidae) from Caatinga, northeastern Brazil Bruno Halluan S. Oliveira¹, Adonias A. Martins Teixeira¹,*, Romilda Narciza M. Queiroz¹, João A. Araujo-Filho¹, Diêgo A. Teles¹, Samuel V. Brito2, Daniel O. Mesquita¹ Where is my place? Quick chorus structure assembly in the European tree frog Michal Berec A possible mutualistic interaction between vertebrates: frogs use water buffaloes as a foraging place Piotr Zduniak1,*, Kiraz Erciyas-Yavuz2, Piotr Tryjanowski3 Good vibrations: A novel method for sexing turtles Donald T. McKnight1,2,*, Hunter J. Howell3, Ethan C. Hollender1, and Day B. Ligon1 Errata to Mecke, S., Hartmann, L., Mader, F., Kieckbusch, M., Kaiser, H. (2016): Redescription of Cyrtodactylus fumosus (Müller, 1895) (Reptilia: Squamata: Gekkonidae), with a revised identification key to the bent-toed geckos of Sulawesi. Acta Herpetologica 11(2):