Natural History of the Tropical Gecko Phyllopezus Pollicaris (Squamata, Phyllodactylidae) from a Sandstone Outcrop in Central Brazil

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Herpetology Notes, volume 5: 49-58 (2012) (published online on 18 March 2012)

Natural history of the tropical gecko Phyllopezus pollicaris (Squamata, Phyllodactylidae) from a sandstone outcrop in Central Brazil.

Renato Recoder1*, Mauro Teixeira Junior1, Agustín Camacho1 and Miguel Trefaut Rodrigues1

Abstract. Natural history aspects of the Neotropical gecko Phyllopezus pollicaris were studied at Estação Ecológica Serra Geral do Tocantins, in the Cerrado region of Central Brazil. Despite initial prospection at different types of habitats, all individuals were collected at sandstone outcrops within savannahs. Most individuals were observed at night, but several specimens were found active during daytime. Body temperatures were significantly higher in day-active individuals. We did not detect sexual dimorphism in size, shape, weight, or body condition. All adult males were reproductively mature, in contrast to just two adult females (11%), one of which contained two oviductal eggs. Dietary data indicates that P. pollicaris feeds upon a variety of arthropods. Dietary overlap between sexes and age classes was moderate to high. The rate of caudal autotomy varied between age classes but not between sexes. Our data, the first for a population ofP. pollicaris from a savannah habitat, are in overall agreement with observations made in populations from Caatinga and Dry Forest, except for microhabitat use and reproductive cycle.

Keywords. Cerrado, lizard, local variation, niche breadth, thermal ecology, sexual dimorphism, tail autotomy.

Introduction information about aspects of the natural history (habitat Phyllopezus pollicaris (Spix, 1825) is a large-sized, use, morphology, diet, temperatures, reproductive nocturnal and insectivorous gecko native to central condition and caudal autotomy) of a population of South America (Rodrigues, 1986; Vanzolini, Costa P. pollicaris from Central Brazil. We discuss local and Vitt, 1980; Vitt, 1995). Currently, P. pollicaris is variation and compare our results with available data. considered to represent a complex of cryptic species (Gamble et al., 2012) which is widespread along the Materials and methods South American great diagonal of open formations

(Vanzolini, 1968; 1988) ranging from the Chaco, Study area through the Brazilian Cerrado and Caatinga eastwards Field work was carried out in the Estação Ecológica Serra Geral to the coastal Atlantic Forest of north-eastern Brazil do Tocantins (EESGT), a large conservation unit in the Cerrado (Vanzolini, 1968; Rodrigues, 1986; Cei, 1993). The region of Central Brazil. The expedition was conducted with the presence of P. pollicaris has been frequently associated main objective of making an inventory of the herpetofauna of with rock outcrops (Rodrigues, 1986; Cei, 1993; Vitt, the EESGT (results in Recoder et al., 2011; Valdujo et al., 2011). The area is characterized by typical Cerrado vegetation compo- 1995; Werneck, Colli and Vitt, 2009). sed of extensive grasslands and savannahs, interspersed by palm Phyllopezus pollicaris is locally common but, despite marshes and gallery forests. The EESGT region has three main its widespread distribution, ecological knowledge levels of altimetric surfaces. The first is composed of a sandstone on the species is scarce. The available information plateau with elevations ranging between 800 and 900 m, which is based on a limited number of studied populations occupies the eastern portion contiguous with Planalto dos Gerais. (Rodrigues, 1986; Vitt, 1986; 1995; Dias and Lira- Isolated sandstone relicts are also present at the central portion of da-Silva, 1998; Werneck, Colli and Vitt, 2009) or on the Ecological Station. The second level composes an extensive sandy depression which resulted from erosion of the plateau, with anecdotal data (Vanzolini 1974; Vanzolini, Costa and elevations ranging between 400 and 550 m in the central, western Vitt, 1980; Cei, 1993). Thus, this work aims to provide and southern portions. The final level is an intermediary plateau crossing the station oriented NNW-SSE direction, with elevations ranging between 600 and 700 m.

1 Laboratório de Herpetologia, Departamento de Zoologia, Field data collection Instituto de Biociências, Rua do Matão, no 101, CEP 05508- Field work was conducted from the end of January through the 090, Universidade de São Paulo, São Paulo, Brazil; beginning of February 2008, at the end of the rainy season (Figure *Corresponding author; e-mail: [email protected] 50 Renato Recoder et al.

Table 1. Morphological measurements, weight and body condition estimates of adult males, adult females and juveniles of Phyllopezus pollicaris from the EESGT. Data presents the average, followed by standard deviation, and minimum and maximum values between parentheses.

Adult males Adult females Juveniles N=9 N=18 N=6 64.4 + 8.9 68.4+ 7.4 40.5+3.1 SVL (mm) (54.3–77.9) (56.3–78.4) (34.1–44.3) 28.6+3.9 30.1+3.5 18.1+1.7 TRL (mm) (23.9–34.7) (23.8–33.8) (15.7–20.1) 12.7+ 1.6 13.4+ 1.4 8.6+ 0.5 HW (mm) (10.6–15.2) (11.1–15.9) (7.7–8.9) 15.9+ 1.8 16.8+ 1.7 11.0 + 0.4 HL (mm) (13.8–18.5) (14.0–19.2) (10.3–11.5) 10.5+ 1.3 11.2+ 1.0 7.3+ 0.3 ML (mm) (9.0–12.5) (9.3–12.6) (6.8–7.7) 12.1+ 1.5 12.5+ 1.2 7.7+ 0.6 FL (mm) (10.2–14.2) (10.6–13.8) (6.8–8.2) 11.7+ 1.5 12.2+ 1.1 7.1+ 0.6 TL (mm) (10.0–14.0) (10.5–13.4) (6.2–7.6) 2.0 + 0.4 2.2+ 0.3 1.2+ 0.1 FTW (mm) (1.7–2.6) (1.7–2.6) (1.1–1.3) 6.7 + 2.8 8.2 + 2.8 1.6 + 0.6 Weight (g) (3.7–11.0) (4.1–11.5) (0.9–2.3) 3.35 + 1.49 3.81 +1.44 1.39+ 0.55 Body Condition (W/SVL) (1.78–5.95) (1.84–6.43) (0.70–1.90)

1). Two main sites were sampled: (1) the southern portion of the EESGT (general coordinates 11o 11’ S; 46o 50’ W), within the municipality of Almas, state of Tocantins, and (2) the eastern portion (general coordinates 10o 40’ S; 46o 09’ W), within the municipality of Formosa do Rio Preto, state of Bahia. Each locality was sampled during nine consecutive days by means of a set of nine lines of pitfall traps (40 buckets each) bordered by drift fences. The main types of available habitats (i.e. grasslands, typical open-savannahs, rocky savannahs, palm marshes and gallery forest) were sampled in each locality. Diurnal and nocturnal active search was performed haphazardly in the vicinities of pitfall traps. Efforts of active search were not standardized, as the primary intention was to complement the results of pitfall trap sampling by means of searches for arboreal, rock-dwelling and large sized species. o Body temperatures (Tb; nearest 0,1 C) were taken with a quick- reading thermometer immediately upon capture. Subsequently,

microhabitat temperature (Tss; temperature of the surface where

the individual was first observed) and air temperature (Ta; temperature five centimetres above substrate where the individual was Figure 1. Climate diagram of the municipality of Mateiros, first observed) were measured. Additionally, the hour of activity state of Tocantins, Brazil. Squares represent maximum and was recorded. Several available substrate temperatures were re- minimum temperatures for each month, bars represent mean corded at random within the sampling area with an infrared por- rain accumulated. Black arrow indicated period in which field table thermometer: temperatures of rock surface (Ts), temperatures work was carried out. in crevices less than five centimetres deep (T<5), and temperatures Natural history of the tropical gecko Phyllopezus pollicaris 51

Figure 2. Typical open savannah with sandstone outcrops where individuals were observed, at eastern portion of the Estação Ecológica Serra Geral do Tocantins (top); individual active by morning on rock surfaces (bottom). 52 Renato Recoder et al.

Figure 3. The results of a principal component analysis on size-adjusted morphometric variables of adult male (black triangles) and females (gray circles) Phyllopezus pollicaris from the EESGT. The contribution of each component to total variance is presented between parentheses.

Table 2. Scores of the Principal Component Analysis performed on size-adjusted morphometric variables of adult Phyllopezus pollicaris. The coefficients with significant correlation with the component are in bold. Individual contribution to total variation and cumulative contribution of each component are shown in the bottom.

Characters PC1 PC2 TRL 0.17425 -0.42491 HW 0.49568 0.59285 HL 0.29050 0.72333 ML -0.02148 0.50903 FL 0.78709 -0.18713 TL 0.83678 -0.43406 FTW 0.71711 0.12758 % of Variance 31.36 22.20 % Cumulative 31.36 53.56 Natural history of the tropical gecko Phyllopezus pollicaris 53

in crevices deeper than five centimetres (T>5). Available substrate tivariate analysis of variance (ANOVA and MANOVA). Shape temperatures were taken at night (22:00 pm) and morning (11:00 variation was summarized with a Principal Component Analysis am) periods. Variation in air temperature was recorded hourly (PCA) performed using the residuals of the linear regression of with a HOBO data logger for eight consecutive days. the morphometric data on the SVL. The effect of environmental temperatures over body temperatures was investigated with mul- Laboratory data collection tiple linear regression analysis. Differences in tail-loss frequen- Specimens were euthanized by a lethal injection of anaesthetic, cies between sexes and age groups were tested using chi-square weighed to the nearest 0.1 g using Pesola© string balances, fixed test (χ2). We calculate estimations of complete tails for individuals using a 10% formalin solution and preserved in 70% ethanol. with tail broken, with the coefficients of the linear regression of Eight morphometric measures were taken with a digital calliper to intact tails with SVL. The relative regeneration of tail and the the nearest 0.1 mm: snout-vent length (SVL), head length (HL), frequency of tail-break in each quart of tail length (from proximal head width (HW), mandible length (ML), trunk length between to distal portion of complete tails) were calculated based on the members (TRL), femur length (FL), tibia length (TL) and fourth estimations of complete tails. Statistical analyses were performed toe width (FTW). Individual body condition was estimated by di- with SPSS v.15.0 and in R (R Development Core Team, 2010), viding the total body mass by SVL. Intact and regenerated porti- with a 0.05 significance level. ons of tails were measured to estimate rates of tail autotomy. Lizards were dissected to remove stomach contents for dietary Results analysis. Prey items were identified to the lowest taxonomic level.

We calculated an index of importance (Ix; modified from Howard Sample distribution et al., 1999) to represent the relative importance of each prey item We obtained 33 individuals of Phyllopezus pollicarus in relation to the entire diet: in the EESGT. All lizards were hand-collected during active search. Most individuals (31 specimens; 93.9%) were obtained within the eastern portion of the station. Two individuals (6.1%) were obtained at the central portion (“Morro do Fumo” hill). No observations of where nx and fx are number and frequency of the prey category x, and N and F the equivalents for all prey items. We estimated the P. pollicarus were made in the southern portion of the niche breadth for numerical data with the inverse of Simpson’s station. All individuals were collected on sandstone diversity index (Simpson, 1949): outcrops in typical savannah habitat (Fig. 2). Morphological variation and sexual dimorphism The largest female measured 78.4 mm SVL and weighted 11.5 g, the largest male 77.9 mm SVL and where p is the proportion of utilization of prey item i, and n is weighted 11.0 g (see Table 1 for descriptive statistics). the number of categories. B varies from 1 (utilization of a single The smallest individual measured 34.1 mm and prey category) to n (even utilization of all prey categories). We weighted 0.9 g. Adult females and males did not differ calculate dietary overlap between sexes and age groups with the in size (ANOVA; F1,25 = 1.538, P = 0.226), weight Pianka’s niche overlap index (Pianka, 1974): (F1,25 = 1.685, P = 0.206) or body condition (F1,25 = 1.794, P = 0.192). The sexes did not differ in any other

morphometric character (MANOVA; F7,19 = 1.779, P = 0.150). Overall shape variation was summarized by PCA with size-adjusted data, which did not reveal any

where pik is the proportion of utilization of prey item i by species conspicuous morphological differentiation between k and pij is the proportion of utilization of prey item i by species sexes (Fig. 3, Table 2). j. Dietary overlap varies from 0 (no overlap) to 1 (complete overlap). Reproductive conditions Dissected lizards were sexed through examination of gonads. Females were considered reproductive if vitellogenic follicles, The sample consisted of six immature juveniles and corpora lutea and/or oviductal eggs were present. Males were twenty seven adult individuals. The smallest juvenile considered reproductive by presence of convoluted epididymides could not be accurately sexed. Of the eighteen adult and enlarged testes. Voucher specimens are deposited in the her- females analysed, two were reproductive (11.0%), petological collection of the Museu de Zoologia da Universidade one presenting a single vitellogenic follicle, the other de São Paulo, São Paulo, Brazil. containing two enlarged oviductal eggs. Corpora lutea was not observed. Statistical analysis Intraspecific sex variation in size, weight, body condition and Of the thirteen males, eight presented convoluted overall morphology was investigated with univariate and mul- epididymides and enlarged testes (mean ratio of testicle 54 Renato Recoder et al.

Table 3. Selected temperatures at microhabitat (body Tb, substrate Tss and air Ta) and available environmental temperatures (rock surfaces Ts, crevices less than five centimetres deep <5T , and crevices deeper than five centimetres >5T ). Data show averages followed by standard deviation, and minimum and maximum values in parentheses.

Body Microhabitat temperatures Environmental temperatures temperatures

Tb Tss Ta Ts T<5 T>5 24.4 + 1.0 22.9 + 1.0 21.9 + 1.0 22.7 + 0.7 23.2 + 0.7 23.5 + 0.6 Night (23.0–26.6) (21.0–25.2) (20.0–24.4) (21.1–23.7) (21.8–25.4) (22.6–25.1) 28.5 + 2.2 26.5 + 1.8 28.1 + 2.5 27.5 + 2.4 26.3 + 2.4 23.7 + 1.9 Morning (25.0–31.2) (25.0–31.2) (24.9–30.8) (24.2–32.0) (23.0–32.0) (22.0–29.0)

Table 4. Number (N), percentage (%N), frequency (F), frequency in percentage (%F) and importance index (Ix) of prey categories in the diet of Phyllopezus pollicaris in the EESGT. In the bottom is presented the dietary niche breadth.

Prey Type N %N F %F Ix INSECTA Orthoptera 8 8.3 7 29.2 0.19 Blattodea 5 6.0 4 16.7 0.11 Coleoptera 3 3.6 2 8.3 0.06 Coleoptera larvae 2 2.4 1 4.2 0.03 Hemiptera Heteroptera 3 3.6 3 12.5 0.08 Homoptera 3 3.6 2 8.3 0.06 Hymenoptera Formicidae 14 16.7 5 20.8 0.19 Formicidae alate 1 1.2 1 4.2 0.03 Isoptera 17 20.2 4 16.7 0.18 Apterygota 11 13.1 8 33.3 0.23 Diptera 1 1.2 1 4.2 0.03 Mantodea 1 1.2 1 4.2 0.03 Lepidoptera larvae 2 2.4 2 8.3 0.05 ARACHNIDA Araneae 9 10.7 8 33.3 0.22 Scorpionida 1 1.2 1 4.2 0.03 MIRIAPODA Diplopoda 1 1.2 1 4.2 0.03 OTHERS Lizard skin 2 2.4 2 8.3 0.05 Egg shell pieces 1 1.2 1 4.2 0.03 Total 85 24 Niche Breadth 8.80 Natural history of the tropical gecko Phyllopezus pollicaris 55

Table 5. Niche breadth and niche overlap between adult females, males and juveniles.

Niche Overlap Adult males Adult females Juveniles Adult males - 0.55 0.71 Adult females - - 0.57 Niche Breadth 7.05 7.05 5.63

Figure 4. Daily variation in environmental temperatures, recorded hourly for eight consecutive days with a Hobo data logger. The average values are in black, the maximum and minimum are in gray. Bars represent standard deviations.

diameter/SVL = 0.079 + 0.006 mm) and were considered were captured at night (between 21:10 and 23:35 pm) to be reproductive (69.2%). One sub adult had slightly from rock surfaces and large crevices. + o convoluted epididymides and moderately enlarged Body temperatures (Tb) averaged 24.4 1.0 C (N = testes (ratio of testicle diameter/SVL = 0.055). Four 22, range = 23.0–26.6oC) in nocturnal individuals, and juvenile males did not have developed testicles (ratio 28.9 + 1.9oC (N = 10, range = 25.0–32.2oC) in diurnal of testicle diameter/SVL < 0.015) thus were considered individuals. Substrate temperatures (Ts) averaged 22.9 immature. + 1.0oC (range = 21.0–25.2oC) by night, and 26.3 + 0.5oC (range = 25.6–27.0oC) by day, air temperatures + o o Activity and temperatures (Ta) averaged 21.9 1.0 C (range = 20.0–24.4 C) by Six individuals were captured during the morning night, and 27.5 + 1.9oC (range = 24.9–30.8oC) by day. (between 11:00 and 11:30 am), of which four were Body temperatures were significantly higher in diurnal directly exposed to sunlight (Fig. 2). Four individuals individuals when compared to nocturnal specimens were captured between 17:00 and 17:30 pm hidden (ANOVA, F1,30 = 73.21, P < 0.01). under rocks and crevices. The remaining 23 individuals 56 Renato Recoder et al.

The multiple regression model indicated that estimated complete tail. Tail-break occurred more body temperature at night is weakly associated with frequently in the first quart of the tail (59.3%, N=16); environmental temperatures (r2 = 0.292; F = 2.290; P less frequently in the second (37.0%, N=10) and = 0.147). Body temperatures of lizards active between on a single occasion in the third quarter (3.7%). 11.00-11.30 am is significantly associated with air temperature (r2 = 0.864; F = 9.515; P = 0.05). Discussion Daily variation in air temperatures has been In agreement with several other South American summarized in Figure 4. Temperatures of selected Phyllodactylids (sensu Gamble et al., 2008; 2011) which microhabitats (T ) did not differ from random substrate ss occur in similar habitats, the savannah population of P. temperatures at night (ANOVA; F = 2.138, P = 1,44 pollicaris studied here occurred in close association 0.150) and morning (F = 2.116, P = 0.156). However, 1,28 with rocks (e.g. Vanzolini, 1982; Marquet et al., temperatures of selected microhabitats by active lizards 1990; Cei, 1993; Colli et al., 2003; Koch, Venegas at morning are significantly lower than temperatures of and Böhme, 2006; Cacciali, Ávila and Bauer, 2007; adjacent rock surfaces exposed to sun (F = 8.360, P 1,28 Blanco, Villavicencio and Acosta, 2009; Werneck, < 0.01). Table 3 summarizes the selected and available Colli and Vitt, 2009). Nevertheless, P. pollicaris can substrate temperatures. be found within a wide range of microhabitats (e.g. tree trunks, fallen logs, cactuses, palm leaves, rock Diet crevices, bushes, bark, fences) in the Caatinga and We analysed 33 stomach contents and identified 85 Atlantic Forest domains (Rodrigues, 1986; 2003; pers. items in 18 prey categories. Nine stomachs were empty obs.). Such environmental plasticity has also been (27.3%). The prey items with the highest importance observed in geckos of the genus Gymnodactylus. While index were Apterygota (0.23), Araneae (0.22), G. amarali seems to be a microhabitat specialist within Orthoptera (0.19), Formicidae (0.19) and Isoptera the Cerrado (Colli et al., 2003; Vitt et al., 2007), G. (0.18) respectively (Table 4). Niche breadth based on geckoides utilizes a wide range of microhabitats in the numerical data was 8.80. Caatinga (Vitt, 1995; Rodrigues, 2003). Microhabitat Overlap in dietary niches between adult males and choice by lizards may be constrained in habitats with juveniles was high (0.712), but moderate between high temperature variation to sites that offer sufficient adult females and males (0.556), or adult females and opportunities for thermoregulation (Marquet et al., juveniles (0.579) (Table 5). Niche breadth was similar 1990; Kearney, 2002; Hitchcock and McBreyer, 2006; for adults of both sexes (7.05) but lower for juveniles Aguilar and Cruz, 2010). Thus, rocks may constitute (5.63). We found no evidence for intra-population a thermally suitable microhabitat for nocturnal lizards niche partitioning between sexes or between adults and in the Cerrado, by providing refuges with a gradient of juveniles in diet. temperatures (Kearney and Predavec, 2000; Kearney, 2002; Vitt et al., 2007). Although we took temperatures Tail autotomy and regeneration of several day and night-active individuals of P. Six specimens had intact tails (18.2%), five individuals pollicaris, by lack of evidence and lack of an adequate had broken tails (15.1%) and the remaining twenty two experimental design, we can not address whether the lizards (66.7%) possessed regenerated tails. Excluding studied specimens were thermoconformers in nocturnal individuals with broken tails (in which case tail loss activity and if individuals thermoregulate during the day cannot be readily attributed to natural causes), we found as our empirical observation suggests, and as do other a frequency of regenerated tails of 78.6% in the sample. New World nocturnal geckos (e.g. Vitt 1995; Vitt and Tail loss did not differ between sexes (χ2 = 0.330; P = Zani, 1997; Colli et al., 2003). 0.565), but the frequency was significantly higher in Our results show that P. pollicaris is an opportunistic adults when compared to juveniles (χ2 = 10.889; P < predator that feeds upon a variety of active arthropods. 0.001). Upon comparison of our data with other three A linear regression of intact tails with SVL as populations for which dietary data are available, we independent variable shows that size is a strong found that a similar variety of prey items were ingested. factor in explaining tail length (r2= 0.981; F = 4,1 In all cases, crickets, ants, termites, spiders, insect 202.87; P < 0.001). The relative regeneration varied larvae and beetles represent the most abundant prey from 47.5% to 100% (mean = 82.6 + 0.14%) of the items (Rodrigues, 1986; Vitt, 1995; Dias and Lira-da- Natural history of the tropical gecko Phyllopezus pollicaris 57

Silva, 1998; Werneck, Colli and Vitt, 2009). Also, P. of regenerated tails more likely reflects escape success pollicaris has been commonly placed among the species from predation attempts than intra-specific aggression with widest dietary niche in lizard communities they (Vitt, 1983; Medel, 1988; Vitt and Zani, 1997; Cooper occur, with estimated niche breadths varying between Jr., Pérez-Mellado and Vitt, 2004). 4.90 and 9.09 (Vitt, 1995; Dias and Lira-da-Silva, 1998; Acknowledgements. We thank Cristiano Nogueira, Paula Werneck, Colli and Vitt, 2009). Valdujo, Pedro Nunes, Tami Mott and José Mario Ghellere for field The results presented herein regarding female assistance and logistical support, and Rodrigo Dios and Silvio Nihei reproductive state are in agreement with those of whom kindly helped with identification of prey items. We thank Rodrigues (1986), who found no signs of reproductive Wouter Beukema for his critical review and valuable suggestions activity in a sample of ten adult P. pollicaris obtained that improved the manuscript. We also thank the “Instituto Chico during February in the Caatinga. In contrast, Vitt (1986, Mendes de Conservação da Biodiversidade” (ICMBIO) for allowing the faunal survey in EESGT and providing collection 1995) studied a lizard community in the Caatinga for over permit (12187-1). The Fundação Boticário de Conservação da a year, and found that four sympatric geckos, including Natureza, “Conselho Nacional de Desenvolvimento Científico P. pollicaris, show nearly continuous reproduction. e Tecnológico” (CNPq) and “Fundação de Apoio à Pesquisa do Seasonal reproduction can however be expected for P. Estado de São Paulo” (FAPESP) provided financial support. pollicaris populations within the Cerrado, as this pattern has been observed in most lizards from the domain, References due to its highly marked seasonality in precipitation and/or food resources (Colli, 1991; Van Sluys, 1993; Aguilar, R., Cruz, F.B. (2010): Refuge use in a patagonian noc- Wiederhecker, Pinto and Colli, 2002; Colli et al., 2003; turnal lizard, Homonota darwini: the role of temperature. J. Herpetol. 44: 236-241. Mesquita and Colli, 2003). Arnold, E.N. (1984): Evolutionary aspects of tail shedding in li- In consensus with our results, Vitt (1986) did not recover zards and their relatives. J. Nat. Hist. 18: 127-169. sexual dimorphism in a populations of P. pollicaris Bateman, P.W., Fleming, P.A. (2009): To cut a long tail short: a from the Caatinga. Lack of the same phenomenon was review of lizard caudal autotomy studies carried out over the found in two other nocturnal Geckonidae, while sexual last 20 years. J. Zool. 277: 1-14. dimorphism was present in two diurnal species (Vitt, Blanco, G., Villavicencio, H.J., Acosta, J.C. (2009): Field body 1986). The latter author stated that sexually dimorphic temperature, diet, and reproduction of Homonota andicola (Gekkonidae) in Catamarca, Argentina. Herp. Rev. 40: 156- characters may differ in visually-oriented compared to 158. auditory-oriented tropical geckos, what may explain the Cacciali, P., Ávila, I., Bauer, F. (2007): A new species of Homo- lack of sexual dimorphism in the nocturnal species (Vitt, nota (Squamata, Gekkonidae) from Paraguay, with a key to the 1986). genus. Phyllomedusa 6: 137-146. The high frequencies of tail-break were similar to Cei, J.M. (1993): Reptiles del noroeste, nordeste y este de la Ar- those reported for other tropical geckos, which varies gentina. Museo Regionale di Scienze Naturali, Torino. Mono- between 32.3% in Hemidactylus palaichthus (Zani, grafie14 : 1-949. Colli, G.R. (1991): Reproductive ecology of Ameiva ameiva 1996), 65.6% in Thecadactylus rapicauda (Vitt and (Sauria: Teiidae) in the cerrado of central Brazil. Copeia 1991: Zani, 1997), to 72.5% in Gymnodactylus amarali (Vitt 1002-1012. et al., 2007). High incidence of tail-break in P. pollicaris Colli, G.R., Mesquita, D.O., Rodrigues, P.V.V., Kitayama, K. may be a phylogenetically conserved propensity for (2003): Ecology of the Gecko Gymnodactylus geckoides ama- caudal autotomy in geckos (Arnold, 1984; Van Sluys, rali in a Neotropical Savanna. J. Herpetol. 37: 694-706. Vrcibradic and Rocha, 2002; Bateman and Fleming, Cooper Jr., W.E., Pérez-Mellado, V., Vitt, L.J. (2004): Ease and 2009). We also found a high relative tail regeneration effectiveness of costly autotomy vary with predation intensity among lizard populations. J. Zool. 262: 243-255. in the population, reinforcing the importance of tail Dias, E.J.R., Lira–da–Silva, R. (1998): Utilização dos recursos autotomy as a defensive strategy for this species (Vitt, alimentares por quatro espécies de lagartos (Phyllopezus polli- Congdon and Dickson, 1977). Although males P. caris, Tropidurus hispidus, Mabuya machrorhyncha e Vanzos- pollicaris occasionally engage in aggressive encounters saura rubricauda) da Caatinga UHE-Xingó. Braz. J. Ecol. (Vitt, 1983), no differences in tail-break frequencies was 1998: 97-101. observed between sexes. Nevertheless, frequencies are Gamble, T., Bauer, A.M., Greenbaum, E., Jackman, T.R. (2008): significantly higher in adults than juveniles. This pattern Out of the blue: a novel, trans-Atlantic clade of geckos (Gekko- ta, Squamata). Zool. Scripta 37: 355-366. has also been observed in other tropical geckos (e.g. Vitt, Gamble, T., Bauer, A.M., Colli, G.R. Greenbaum, E. Jackman, 1983; Vitt and Zani, 1997). Thus, variation in frequency T.R. Vitt, L.J., Simons, A.M. (2011): Coming to America: mul- 58 Renato Recoder et al.

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Accepted by Wouter Beukema