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 Phyllomedusa - 16(2), December 2017 255 et al. 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 Palavras-chave: 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 Phyllomedusa - 16(2), December 2017 257 et al. 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).