title: Sharing refuges on arid islands: ecological and social influence on aggregation behaviour of wall geckos.
abstract: BACKGROUND
  The extent of social behaviour among reptiles is underappreciated. Two types of aggregations are recognized in lizards: ecological and social, i.e., related to the attraction to a site or to animals of the same species, respectively. As most lizards are territorial, aggregations increase the probability of aggressive interactions among individuals, a density-dependent behaviour.
  
  
  METHODS
  After some spurious observations of aggregation behaviour in the endemic Cabo Verde nocturnal geckoTarentola substituta,we conducted a field-based study in order to thoroughly characterize it. We sampled 48 transects and 40 10 × 10 m quadrats on São Vicente Island to describe the incidence, size and composition of aggregations and to study the effect of gecko and refuge density, plus refuge quality, on refuge sharing. We hypothesize that when density of animals and scarcity of high-quality refuges is higher, lizards have increased probability of aggregating. We also predict a consistent pattern of size and composition of groups (male-female pairs, only one adult male per group) throughout the year if there is a selected behaviour to avoid agonistic interactions, and low thermal advantage to aggregating individuals.
  
  
  RESULTS
  We present one of the first evidences of aggregation for Phyllodactylidae geckos. We found thatT. substitutaforms aggregations around 30-40% of the time, and that refuges are almost always shared by a female-male pair, sometimes with a juvenile, probably a mechanism to avoid aggressive interactions. We also observed that refuge sharing is dependent on refuge quality, as medium-large (thermally more stable and positively selected) rocks are shared much more frequently than small ones, but independent of adult sizes. Refuge sharing is also directly related to the density of geckos and inversely related to the density of high-quality refuges. We found no relation between body temperatures of geckos and refuge sharing when controlling the effect of rock/air temperature, suggesting that huddling does not improve thermoregulation.
  
  
  CONCLUSIONS
  Our results suggest that in this harsh environment (rocks reach 46 °C) aggregation incidence is mainly driven by an ecological factor (scarcity of high-quality refuges) and its intersexual composition by social factors (avoidance of agonistic interactions by males, and possible increased reproductive success of the pair). This study sheds some light on the little explored gecko aggregation behaviour and other studies should follow.
authors.affiliation: CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Universidade do Porto, Vairão, Portugal; IBE, Institute of Evolutionary Biology (CSIC-Universitat Pompeu Fabra), Barcelona, Spain.
authors.forename: Raquel
authors.initials: R
authors.lastname: Vasconcelos
authors.affiliation: Departamento de Bioquímica, Genética e Inmunología, Facultad de Biología, Universidad de Vigo , Vigo , Spain.
authors.forename: Sara
authors.initials: S
authors.lastname: Rocha
authors.affiliation: CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Universidade do Porto , Vairão , Portugal.
authors.forename: Xavier
authors.initials: X
authors.lastname: Santos
keywords: Cabo verde; Density-dependent; Male–female pairs; Refuge sharing; Reptile; Tarentola substituta; Thermobiology