Phyllomedusa 16(2):255–267, 2017 © 2017 Universidade de São Paulo - ESALQ ISSN 1519-1397 (print) / ISSN 2316-9079 (online) doi: http://dx.doi.org/10.11606/issn.2316-9079.v16i2p255-267 Nesting in the lizard Phyllopezus pollicaris (Squamata: Phyllodactylidae) and a phylogenetic perspective on communal nesting in the family

Fabricius M. C. B. Domingos,1,2 Ísis C. Arantes,3 Renan J. Bosque,3 and Marcella G. Santos3 1 Programa de Pós-Graduação em Ecologia e Conservação, Universidade do Estado de Mato Grosso, 78690-000 Nova Xavantina, MT, Brazil. E-mail: [email protected]. 2 3 Department of Biology, University of Mississippi, Oxford, Mississippi 38677, USA.

Abstract Nesting in the lizard Phyllopezus pollicaris (Squamata: Phyllodactylidae) and a phylogenetic perspective on communal nesting in the family. Communal nesting occurs in many reptile species. The hypotheses that explain the evolution of such behaviours are still controversial, but will be better understood as more communal nesting records are a communal nesting record for squamate species in a Cerrado core area. We provide nest photographs and detailed neonate measurements and weight. Nests were found during the dry season, in contrast with the reproduction pattern previously described for the species in the Cerrado periphery. We also conducted an extensive literature review seeking all available information on communal nesting information in Phyllodactylidae, and present this information in the context of a phylogenetic tree of phyllodactylid genera. We suggest that studying the correlation between communal nesting evolution and reproductive available in the literature.

Keywords: Brazil, Cerrado, lizards, nest, phylogeny, reproduction.

Resumo Ninhadas de Phyllopezus pollicaris de ninhadas comunais para a família. serão melhor compreendidas à medida que mais registros de ninhadas comunais forem descritos na literatura. Neste estudo, registramos duas ninhadas comunais de , incluindo a

Received 19 January 2017 Accepted 27 July 2017 Distributed December 2017

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encontradas durante a estação seca, em contraste com o padrão reprodutivo previamente descrito respeito de ninhadas comunais na família Phyllodactylidae e apresentamos os resultados no contexto entre a evolução de ninhadas comunais e o investimento reprodutivo se tornará um campo de estudo

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Introduction (Doody 2009), but appears to be particularly prevalent in geckos (Graves and Duvall 1995, Communal nesting, the phenomenon by Doody 2009). Clutch size is a phylo- which different females of one or more species genetically conserved attribute in geckos (Vitt deposit eggs at a shared nest (Noble and Mason 1986, Sinervo 1994), and most species produce 1933), occurs in many squamate species (Graves only one or two eggs at a time (Mesquita and Duvall 1995). The evolutionary and ecolo- 2015, 2016a), which is a relatively small number gical processes resulting in this behavior are compared to most other lizard families (Mesquita hypothesized to be related to either environmental 2016b). The repeated evolution of constraints or selective advantage (Doody communal nesting in geckos might be a 2009). Early attempts to explain squamate behavioral adaptation in response to this clutch communal nesting suggested that this pheno- size constraint, as it potentially increases female menon was simply a factor of nesting site scarcity (Rand 1967, Vitt 1986). More recent (Blouin-Demers 2004, Radder and Shine studies, however, advocate that communal 2007). nesting evolved through natural selection, since A few Neotropical geckos are known to nest there are potential energetic and reproductive communally, such as advantages for females that use communal nests. (Boulenger, 1888) in the Caatinga (Oliveira These advantages could be related to less time 2015), Vanzolini, 1978 spent searching and building nests (Graves and in the Caatinga (Bezerra 2011), Duvall 1995), and to increased offspring Rodrigues, 1986 in the Caatinga (Lima performance and/or survivorship (Radder and 2011), and (Guichenot, 1855) in the Amazon (Vitt collected data and laboratory experiments 1997, Oda 2004), among others. To the best of support the latter hypothesis, and found that our knowledge, the only known gecko that produces communal nests in the Cerrado, the the involved females (Blouin-Demers Brazilian savanna, is 2004, Radder and Shine 2007). Nonetheless, the (Spix, 1825) (Righi 2004). Despite the fact relative prevalence of these alternative processes that the Cerrado is a biodiversity hotspot (Myers has not been investigated in a comparative ma- 2000) and houses a high proportion of croevolutionary approach (Doody 2009). South American lizard diversity (Colli Communal nesting has evolved independently 2002, Nogueira 2011), natural history several times during the history of squamates information on the reproductive ecology of

Phyllomedusa - 16(2), December 2017 256 Phyllopezus pollicaris

Cerrado lizards are still scarce (e.g., Colli 1991, Table 1. Descriptive statistics of body measurements Wiederhecker 2002, Colli 2003). (mm) and mass (g) of Phyllopezus pollicaris The lizard is distri- neonates (N = 22) from two communal nests in the Cerrado biome. buted across the dry-diagonal biomes of South America (Caatinga, Cerrado, and Chaco) Neonates measurements Mean ± SD (Werneck and Colli 2006). Although there are no recent taxonomic changes related to the status Mass 0.88 ± 0.06 of it is certainly a species complex Snout vent length 32.53 ± 1.12 composed of many cryptic lineages (Gamble 2012, Werneck 2012). In terms of Tail length 34.19 ± 1.53 habitat use, is mostly found on rock Body width 6.65 ± 0.65 outcrops (Werneck 2009), either granite (Vitt 1986) or sandstone (Recoder 2012), Body height 4.57 ± 0.48 where they forage and lay their eggs (Righi Head width 6.90 ± 0.40 2004, Recoder 2012). Individuals of this species are surface active during the night and Head height 3.82 ± 0.30 usually found within rock crevices during the Head length 9.80 ± 0.58 day (Recoder 2012). According to literature records, is only known to produce Anterior limb length 9.91 ± 0.70 two eggs (Vitt 1986, Ávila and Cunha-Avellar Posterior limb length 13.62 ± 0.71 2005, Recoder 2012, Righi 2012). that will nest communally, and found nests with a maximum number of six Materials and Methods eggs (probably produced by three females) in the Caatinga biome. Ávila and Cunha-Avellar We searched for nests (2005) recorded a similar instance (single nest for three consecutive days (18–20 May 2013) at with six eggs) in seasonally-dry forests in Parque Nacional Chapada dos Veadeiros (Alto western Brazil. Finally, Righi (2004) found Paraíso de Goiás, Goiás state, Brazil), a large protected area at the core of the Cerrado biome. peripheral area, southeastern Brazil. The only area (14°9'47.26'' S, 47°37'9.46'' W), which available information on neonates of this species consists of typical cerrado (savanna) vegetation is average snout vent length and weight, which growing amidst sandstone rock outcrops (Figure were based on a limited number of individuals 1A, B). Sandstone rocks are found in many (Vitt 1986), and snout vent length and tail length different sizes, and it is common for lizards to of two individuals (Gomides and Garcia 2014). hide and nest under rocks laying on top of larger Herein, we report the largest communal nest rocks (Recoder 2012). Hence, we spent ~8 found for hours per day searching for eggs in rock crevices, squamate communal nesting in a Cerrado core and also lifting smaller rocks, since nests could area, and visual records of such nests in the potentially be found under such rocks. When biome. We also provide detailed neonate found, all eggs from a nest were collected, taken measurements and weight (Table 1, Appendix I). to the laboratory to hatch, and kept at room Finally, we present this information with a temperature (~21°C) in separate closed plastic communal nesting phylogenetic perspective for containers (lids had holes to allow for gas the family Phyllodactylidae. exchange). Since there was no soft substrate in

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the rocks where eggs were collected (e.g., litter found on 19 May 2013 under stacked rocks. All or sand), they were kept in exfoliated vermiculite. eggs were hard shelled and oviposition occurred Neonates were killed immediately after directly on the rocks. From Nest 1, 80% of the hatching using a 2% lidocaine hydrochloride eggs hatched (N = 20), and from Nest 2 50% (N injection. Body measurements of the neonates were taken to the nearest 0.01 mm using a digital ranged from 3 to 112 days (mean ± SD: 30 ± caliper (Mitutoyo®), and weighing to the nearest 32.5 days; Appendix I). Neonates measurement 0.01 g was performed using a digital scale (A&D and mass descriptive statistics can be found in Company®). To allow for future genetic Table 1. investigations, we also extracted the liver from In our literature review, out of the 135 all lizards after measurements were performed. currently described phyllodactylid species, we Livers were preserved in 99% ethanol, and found 21 species for which communal nesting evidence is available (Table 2), distributed and tissue samples were deposited at Coleção among seven out of the 10 phyllodactylid genera Herpetológica da Universidade de Brasília (Figure 2, Appendix II). (CHUNB) (Appendix I). To place our results in a broader evolutionary Discussion perspective within the Phyllodactylidae, we conducted a literature review of published Females of may take information on communal nesting for this family. up to three months to produce eggs when We sought published papers and notes on nests reproduction season starts (i.e., time between of all synonyms of the 135 currently described species of Phyllodactylidae (Uetz 2016). Since are found) (Righi 2012), and it took the absence of communal nesting behavior is between three and 112 days for our captured virtually impossible to determine, it becomes eggs to hatch in captivity. Hence, it is not impractical to reconstruct the evolution of this possible, at this stage, to evaluate whether at trait in a phylogenetic context using only the least some ovipositions in communal relatively few cases where communal nesting is nests were produced by the same females. We actually present. Hence, following the approach believe that future genetic investigations using of Doody (2009), we plotted the available the collected tissue samples, and possibly other information in a phylogenetic tree of all currently nests, might clarify this issue. recognized phyllodactylid genera, in order to The uneven number of eggs in Nest 1 is visualize the macroevolutionary patterns of this somewhat surprising, because behavior. We used the most recent Squamate is believed to always produce two phylogenetic estimate (Tonini 2016), and eggs (Vitt 1986, Righi pruned the tree so it would only depict the might be explained by egg predation, or simply relationships among phyllodactylid genera, using the package (Paradis 2004) in R separate location, since no empty shells were version 3.3.1 (R Core Team 2016). found at Nest 1. Although very unlikely, the fact that we found the same incubation time for three Results eggs in two distinct cases (Appendix I), and the arrangement of two rows of three eggs each in We found two Nest 2 (Figure 1E) might suggest that communal nests, one containing 25 eggs (Figure could produce up to three eggs in our 1C, D), hereafter Nest 1, and another containing study site. We visually inspected all Chapada six eggs (Figure 1E), Nest 2. Both nests were dos Veadeiros specimens deposited

Phyllomedusa - 16(2), December 2017 258 Phyllopezus pollicaris

A B

C D

E

Figure 1. Phyllopezus pollicaris communal nests. (A–B) The ‘cerrado rupestre’ habitat at Parque Nacional da Chapada dos Veadeiros. Nest 1 with 25 eggs, (C) 22 at the bottom rock and (D) three stuck at the top rock. (E) Nest 2 with six eggs disposed in two rows of three eggs each. Photographs of nests were taken after the removal of the top rocks during the searches.

at CHUNB (N = 22) but, unfortunately, there possibly suitable nesting sites and found only were no pregnant females that could shed light two nests. Since nesting sites appear to be on this question. This hypothesis might be tested abundant in the area, and considering that one when more communal nests are nest contained such a large number of eggs (25), found, and specimens are collected during the reproductive season at Chapada dos Veadeiros. true for living in the Cerrado. The Vitt (1986) suggested that communal nesting alternative explanation, that communal nesting by geckos in arid areas might simply be a factor might increase of the limited amount of nesting sites. However, because of egg insulation, predation abatement, we spent ~ 24 working hours checking many increased hatchling quality, or other mechanisms,

Phyllomedusa - 16(2), December 2017 259 et al. Table 2. Table Thecadactylus rapicauda Tarentola mauritanica Tarentola crombiei Tarentola americana Ptyodactylus oudrii Ptyodactylus hasselquistii Ptyodactylus guttatus Phyllopezus pollicaris Phyllopezus periosus Phyllodactylus wirshingi Phyllodactylus transversalis Phyllodactylus reissii Pozo-Andrade, 2013 Phyllodactylus leoni Phyllodactylus kofordi Phyllodactylus interandinus Phyllodactylus galapagensis Homonota uruguayensis Homonota borellii Gymnodactylus geckoides Gymnodactylus darwinii Gymnodactylus amarali Species Phyllodactylidae species known to lay eggs communally, and reference list from where communal nesting information was obtained. was information nesting communal where from list reference and communally, eggs lay to known species Phyllodactylidae

(Peracca, 1897)

Diaz & Hedges, 2008 Lataste, 1880

Torres-Carvajal, Carvajal-Campos Barnes, Nicholls and

(Gray, 1831) Peters, 1862 Heyden, 1827 Rodrigues, 1986

(Linnaeus, 1758) (Spix, 1825) Dixon & Huey, 1970

Barbour, 1925

(Vaz-Ferreira & Sierra de Soriano, 1961) Kerster & Smith, 1955 (Gray, 1845)

(Houttuyn, 1782) (Donndorff, 1798)

Spix, 1825

Huey, 1975 Dixon & Huey, 1970 Peters, 1869 (Vitt 1986, Righi (Werner 1965, Werner 1972, 1986, Roobol (Schwartz 1968, Novo 1985, Schwartz and Henderson 1991) (Bons 1959) (Petzold 1982, González de la Vega 1988) (Diaz and Hedges 2008, Alfonso et al. (Vitt pers. comm. in Doody (Vitt pers. comm. in Doody (Godoy and Pincheira-Donoso 2009) References for communal nesting (Cassimiro and Rodrigues 2010) 2004, Ávila and Cunha-Avellar 2005, Gomides (García-Márquez (Torres-Carvajal (López-Victoria (Graves and Duvall 1995) Garcia 2014, this study) (Dixon and Huey 1970) (Dixon and Huey 1970) (Dixon and Huey 1970) (Lima (Disi (Vitt (Gudynas 1986) (Kushlan 1981) et al. et al. et al. 1997) et al. 2001) et al. 2011) et al. 2013) 2013) et al. et al. 1996) et al. 2009) 2009) 2012) et al. 2002)

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biome, where reproduction is concentrated in a few months. On the other hand, we found nests at the start of the dry season, outside the estimated reproduction time for in another Cerrado location (Diamantina, Minas Gerais state) at the southeastern border of the biome (Righi 2012). It is beyond the scope of this manuscript to infer reproduction seasonality aspects of , but such large difference might be related to climatic differences between both localities, or to the cryptic lineages within the species (Werneck 2012). Coincidently, reproduction of other phyllodactylid species, , is also concentrated during the dry season in Cerrado core areas (Colli 2003). The same is not true for other lizard species such as Figure 2. Phylogenetic relationships among currently (Linnaeus, 1758) (Colli 1991) and recognized genera of Phyllodactylidae (Wied-Neuwied, 1820) (number of species know to nest communally/ (Wiederhecker 2002) which reproduce number of species in the genus). Tree adapted from Tonini et al. (2016). Genera known to during the wet season at the core of Cerrado. lay eggs communally are in black, and genera Hence, there is also the possibility that for which no evidence of communal nesting is differences between the peripheral and core available are in gray. reproduction patterns of phyllodactylid species (Furley 1999, Souza 2015). Communal nests may provide thermal should be further studied to test this suggestion insulation for eggs, which could in turn lead to (Doody 2009). Interestingly, repro- advantage for females (Blouin-Demers duces throughout the year in the Caatinga (Vitt 2004). This is apparently true in temperate 1986), but only between August and December regions, but could be an important selective in the Cerrado (Righi 2012), during the pressure in the cold nights of the Cerrado dry end of the dry season and through part of the season. If this hypothesis is true, we predict wet season (Eiten 1972). The same pattern is that: (1) within populations, neonates from nests true for two other phyllodactylid sister species: with fewer eggs would be smaller than neonates Spix, 1825, which from nests with more eggs; and (2) between reproduces continuously in the Caatinga, and Barbour, 1925 which reproduces in colder regions as opposed to nests with fewer during the Cerrado dry season (Colli eggs in warmer regions. We could not detect a 2003). This pattern is probably related to the from the two nests sampled in this study (results lizards to constantly invest in reproduction (Colli 1991, Mesquita 2016a), and also the other hand, we registered the largest suggests that large Phyllodactylidae communal communal nest to date, as opposed to nests will be mostly found in the Cerrado smaller nests found both in the Caatinga (Vitt

Phyllomedusa - 16(2), December 2017 261 et al.

1986, Gomides and Garcia 2014), which presents The relatively high proportion of species know warmer climate, and in the Cerrado during the to nest communally in Phyllodactylidae (Figure 2) wet season (Righi 2012), which is usually shows that such behavior is particularly prevalent warmer than the dry season. Average snout vent in the family. Apart from the species-rich genus length of neonates in this study all genera have between 1 and 3 (32.53 ± 1.12 mm) was similar to the species known to lay eggs communally (Figure 2, measurements obtained by Vitt (1986) in a Appendix II). The phylogenetic distribution of Caatinga area (31.0 ± 0.55 mm). However, a communal nesting in phyllodactylid genera sample of two laboratory hatched strongly suggests that this is an ancient behavior, also from the Caatinga were considerably smaller (24.82 ± 0.04 mm) (Gomides and Garcia 2014). particularly surprising, given that communal nesting behavior is known for members of most communal nesting on neonate size would require currently recognized Gekkota families (Doody a comprehensive database of hatchlings size and 2009). Communal nesting has not been adult females among populations of recorded in or (and closely related species), and our complete species, which could suggest that such behavior set of measurements will be a valuable source of was lost at least twice during the evolution of the information for future comparative studies. This family (Figure 2). However, there is not much hypothesis should also be explored in an ecological information on the few species in these integrative approach, by testing the effects of genera (Marquet 1990, Razzetti 2011), temperature on embryo development in the and future detailed studies may unearth this laboratory, and by assessing nests microclimate behavior in these genera. Communal nesting is a recurrent feature of Alternatively, communal nesting might have gecko evolution (Graves and Duvall 1995, little or no effect on neonate size in Doody 2009) and, notably, species of the Phyllodactylidae, since the rigid eggshell acts family Phyllodactylidae display some of the as an important constraint in this trait (Pike highest levels of investment per progeny of all lizard families (Mesquita 2016b). allocating resources for multiple clutches per Therefore, t reproductive season is a common strategy to nesting might be related to the high reproductive increase reproductive success (Meiri investment in relation to the low female longevity 2012, 2015). Genetic analysis of the samples found in species (Mesquita collected in this study will provide a rare 2016b). The same relationship might explain opportunity to detect if females are producing why this behavior is present in so many multiple clutches and using the same nesting phyllodactylid species (Figure 2). Although, at site. Nevertheless, considering the constraints this stage, this should be interpreted only as a in the reproductive biology of phyllodactylids, rudimentary association, studying the correlation between communal nesting evolution and individuals. reproductive investment should become a fertile Some of the adaptive hypotheses that might explain this phenomenon in the species include becomes available in the literature. reduction in eggs predation, reduction in Acknowledgments and increasing survival of embryos and hatchlings (Graves and Duvall 1995, Doody Animals were collected under Instituto Chico 2009). Mendes de Conservação da Biodiversidade

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Phyllomedusa - 16(2), December 2017 265 et al. measured (CHUNB 72555, incubation = 7 days), and is not included in this table; two other neonates (CHUNB 72588 and 72589) coul 72589) and 72588 (CHUNB neonates other two table; this in included not is and days), 7 = incubation 72555, (CHUNB measured CHUNB-72547 Museum voucher CHUNB-72548 CHUNB-72549 CHUNB-72551 CHUNB-72552 CHUNB-72553 CHUNB-72554 CHUNB-72561 CHUNB-72562 CHUNB-72563 CHUNB-72580 CHUNB-72581 CHUNB-72582 CHUNB-72583 CHUNB-72585 CHUNB-72586 CHUNB-72588 CHUNB-72589 CHUNB-72590 CHUNB-72591 CHUNB-72599 CHUNB-72601 Appendix I. Appendix day recorded (hatching occurred between July 10 July between occurred (hatching recorded day Body measurements (mm), mass (g), and incubation time since field collection (days) of (days) collection field since time incubation and (g), mass (mm), measurements Body Mass 0.90 0.90 0.90 0.90 0.80 0.90 0.90 0.94 0.96 0.96 0.92 0.91 0.85 0.95 0.91 0.80 0.75 0.80 0.83 0.84 0.84 0.79 Snout vent length 32.61 31.35 31.78 33.84 32.86 33.48 32.30 34.17 30.08 32.14 33.46 33.17 32.24 32.21 33.40 31.43 34.35 33.98 32.25 30.95 32.12 31.49 length 33.96 33.80 34.03 36.18 35.65 34.10 32.80 33.91 35.65 35.99 35.82 32.44 33.82 34.82 35.88 31.45 34.44 35.59 33.41 34.89 33.23 30.36 Tail th width and 31 and Body 6.91 7.42 7.16 7.37 7.00 7.14 7.69 7.04 6.70 7.35 6.70 6.11 6.20 6.81 5.72 5.62 5.71 6.04 7.00 6.96 5.62 6.08 st 2013) and were not included in the average incubation time calculation (30.0 calculation time incubation average the in included not were and 2013) height Body 5.02 5.31 3.87 4.35 4.15 4.86 4.18 4.86 5.10 5.15 4.51 4.17 4.53 4.03 4.03 4.10 4.75 3.83 4.83 4.51 5.06 5.35 width Head 7.28 6.83 7.03 6.75 6.83 6.77 6.99 6.76 6.77 6.66 7.55 7.08 6.10 7.38 6.65 6.51 7.10 7.29 7.60 6.60 6.07 7.09 height Head 3.69 3.43 4.16 3.43 3.56 3.95 3.56 3.78 3.51 3.78 3.47 3.95 4.17 4.32 3.70 3.88 4.54 3.97 3.92 3.94 3.44 3.86 length 10.24 10.23 10.14 10.65 10.27 10.48 10.61 Head 9.89 9.54 9.68 9.66 9.98 9.83 9.87 9.53 9.55 9.55 7.89 9.83 9.46 9.31 9.45 Phyllopezus pollicaris Phyllopezus limb length Anterior 9.61 9.41 9.24 9.80 9.13 9.91 9.41 9.30 9.26 9.35 10.94 10.17 8.81 9.04 10.60 10.27 10.93 10.21 10.99 11.02 10.38 10.26 limb length Posterior 14.16 13.93 12.57 13.53 13.89 14.32 12.32 14.38 12.42 14.15 14.31 14.16 13.82 12.60 13.57 13.55 14.42 14.06 12.91 14.27 12.65 13.60 neonates. One neonate was not was neonate One neonates. Communal nest (# eggs in nest) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 1 (25) Nest 2 (6) Nest 2 (6) Nest 2 (6) d not have their hatching their have not d ± 32.5 days). 32.5 Incubation collection 52 to 72 52 to 72 since 112 15 15 15 29 29 31 31 32 36 74 74 99 3 3 3 4 4 7 7

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Appendix II. Phylogenetic tree depicting all currently recognized species in the family Phyllodactylidae, extracted from Tonini et al. (2016). Species known to lay eggs communally are in black, and species for which no evidence of communal nesting is available are in gray.

Tarentola americana Tarentola parvicarinata Tarentola boettgeri Tarentola pastoria Tarentola delalandii Tarentola chazaliae Tarentola bischoffi Tarentola gomerensis Tarentola nicolauensis Tarentola rudis Tarentola protogigas Tarentola maioensis Tarentola gigas Tarentola fogoensis Tarentola substituta Tarentola raziana Tarentola caboverdiana Tarentola darwini Tarentola bocagei Tarentola boavistensis Tarentola ephippiata Tarentola annularis Tarentola crombiei Tarentola neglecta Tarentola mindiae Tarentola fascicularis Tarentola deserti Tarentola boehmei Tarentola mauritanica Tarentola angustimentalis Tarentola albertschwartzi Phyllopezus pollicaris Phyllopezus periosus Phyllopezus maranjonensis Phyllopezus lutzae Gymnodactylus darwinii Gymnodactylus vanzolinii Gymnodactylus guttulatus Gymnodactylus geckoides Gymnodactylus amarali Phyllodactylus wirshingi Phyllodactylus transversalis Phyllodactylus paralepis Phyllodactylus ventralis Phyllodactylus papenfussi Phyllodactylus rutteni Phyllodactylus microphyllus Phyllodactylus palmeus Phyllodactylus lepidopygus Phyllodactylus tuberculosus Phyllodactylus muralis Phyllodactylus johnwrighti Phyllodactylus tinklei Phyllodactylus interandinus Phyllodactylus santacruzensis Phyllodactylus insularis Phyllodactylus sentosus Phyllodactylus inaequalis Phyllodactylus pulcher Phyllodactylus heterurus Phyllodactylus partidus Phyllodactylus gerrhopygus Phyllodactylus paucituberculatus Phyllodactylus delsolari Phyllodactylus kofordi Phyllodactylus pumilus Phyllodactylus leoni Phyllodactylus darwini Phyllodactylus dixoni Phyllodactylus clinatus Phyllodactylus delcampoi Phyllodactylus homolepidurus Phyllodactylus duellmani Phyllodactylus davisi Phyllodactylus xanti Phyllodactylus unctus Phyllodactylus nocticolus Phyllodactylus bugastrolepis Phyllodactylus martini Phyllodactylus lanei Phyllodactylus bordai Phyllodactylus reissii Phyllodactylus leei Phyllodactylus baurii Phyllodactylus galapagensis Phyllodactylus barringtonensis Phyllodactylus julieni Phyllodactylus apricus Phyllodactylus thompsoni Phyllodactylus angustidigitus Phyllodactylus gilberti Phyllodactylus angelensis Homonota underwoodi Homonota fasciata Homonota williamsii Homonota uruguayensis Homonota taragui Homonota rupicola Homonota borellii Homonota darwinii Homonota whitii Homonota andicola Garthia penai Garthia gaudichaudii Thecadactylus solimoensis Thecadactylus rapicauda Thecadactylus oskrobapreinorum Ptyodactylus ragazzii Ptyodactylus homolepis Ptyodactylus oudrii Ptyodactylus dhofarensis Ptyodactylus orlovi Ptyodactylus puiseuxi Ptyodactylus hasselquistii Ptyodactylus guttatus Ptyodactylus ananjevae Haemodracon trachyrhinus Haemodracon riebeckii Asaccus montanus Asaccus tangestanensis Asaccus nasrullahi Asaccus iranicus Asaccus barani Asaccus zagrosicus Asaccus kermanshahensis Asaccus kurdistanensis Asaccus griseonotus Asaccus elisae Asaccus saffinae Asaccus granularis Asaccus platyrhynchus Asaccus gallagheri Asaccus caudivolvulus Asaccus andersoni

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