Thesis (Ph.D.) -- Flinders University, School of Biological Sciences. / Typescript (bound). Includes bibliographical references. Also available online.

Research on squamate evolution of Australia has predominantly focused on the eastern portion of the continent, whereas little is known about the record from Western Australia. Deposits in Devils Lair provide a glimpse of late Pleistocene Egernia group skink representation from the Cape Leeuwin-Naturaliste region of southwestern Western Australia. Previous studies of fossils from the cave indicate a peak in the number of lizard remains during the last glacial maximum, a time when mammal and charcoal remains suggest an open, dry environment. Previous studies of Devils Lair have not attempted to identify lizard remains beyond 'lizard'. Here, we identify Egernia group lizard remains as Liopholis spp., Egernia spp., Lissolepis spp. and Tiliqua rugosa, which range in age from ca 48000 to 13000 yr B.P.

When species compete for resources, in a stable homogeneous environment, there are two possible outcomes. The first is that one species will out-compete the other and exclude it from the environment. This is known as the competitive exclusion principle. The second is that both species will manage to coexist. Coexistence can only occur if the species’ niches are differentiated such that interspecific competition is minimised, or eliminated. This outcome is known as resource partitioning. Two closely related Australian skink species of the Egernia genus, Egernia multiscutata and Egernia whitii, are abundant and sympatric on Wedge Island in South Australia’s Spencer Gulf. The species are morphologically very similar and appear to have very similar life histories and habitat requirements. Ostensibly, they would compete for limiting resources in this environment. This thesis is the first investigation into resource partitioning in this previously unstudied model organism. I report the results of multi-faceted investigations into the coexistence of the skinks, E. multiscutata and E. whitii on Wedge Island and the evidence for, and mechanisms of, any facultative resource partitioning between them. Study methods involved a transect survey of most of Wedge Island to determine the species’ distributions and any evidence for resource partitioning; a morphological comparison to investigate any potential competitive advantages of either species; a habitat choice experiment to establish retreat-site preferences in the absence of interspecific interference; and, a series of staged dyadic encounter experiments to investigate interspecific competitive interactions. Resource partitioning was evidenced by differential distributions of the species among substrates containing the elements required for permanent refuge shelters. This partitioning was not mediated by avoidance of particular substrates but by the presence of the opponent species, combined with attraction to suitable substrates. Asymmetries in some morphological characters were found to confer a potential competitive advantage to E. multiscutata in agonistic encounters with E. whitii. Both species were found to have the same refuge site preferences when interference competition was experimentally removed. This result was not concordant with observed resource partitioning in the field and suggests that the habitat choices of both species are modified by the presence of the opponent species. Analyses of staged dyadic encounter experiments showed that E. multiscutata was more likely to gain greater access to a contested habitat resource and more likely to exclude E. whitii from the resource than vice-versa. Nevertheless, the outcome of competitive interactions was not completely deterministic and there was some tolerance of co-habitation. E. multiscutata’s competitive advantage was attributable largely to its greater mass and head dimensions relative to snout to vent length. However, differential behavioural responses to the threat of larger opponent size also played an important part in resource partitioning between the species.

ABSTRACT Social relationships, habitat utilisation and life history characteristics provide a framework which enables the survival of populations in fluctuating ecological conditions. An understanding of behavioural ecology is critical to the implementation of Natural Resource Management strategies if they are to succeed in their conservation efforts during the emergence of climate change. Egernia whitii from Wedge Island in the Spencer Gulf of South Australia were used as a model system to investigate the interaction of life history traits, scat piling behaviour and chemosensory communication in social lizards. Juveniles typically took ¡Ý 3 years to reach sexual maturity and the results of skeletochronological studies suggested longevity of ¡Ý 13 years. Combined with a mean litter size of 2.2, a pregnancy rate estimated at 75% of eligible females during short-term studies, and highly stable groups, this information suggests several life history features. Prolonged juvenile development and adult longevity may be prerequisite to the development of parental care. Parental care may, in turn, be the determining factor that facilitates the formation of small family groups. In E. whitii parental care takes the form of foetal and neonatal provisioning and tolerance of juveniles by small family or social groups within established resource areas. Presumably, resident juveniles also benefit from adult territorialism. Research on birds suggests that low adult mortality predisposes cooperative breeding or social grouping in birds, and life history traits and ecological factors appear to act together to facilitate cooperative systems. E. whitii practice scat piling both individually and in small groups. Social benefits arising from signalling could confer both cooperative and competitive benefits. Permanent territorial markers have the potential to benefit conspecifics, congenerics and other species. The high incidence of a skink species (E. whitii) refuging with a gecko species (N. milii) on Wedge Island provides an example of interspecific cooperation. The diurnal refuge of the nocturnal gecko is a useful transient shelter for the diurnal skink. Scat piling may release a species ¡®signature¡¯ for each group that allows mutual recognition. Scat piling also facilitates intraspecific scent marking by individual members, which has the potential to indicate relatedness, or social or sexual status within the group. The discovery of cloacal scent marking activity is new to the Egernia genus. E. Whitii differentiate between their own scats, and conspecific and congeneric scats. They scent mark at the site of conspecific scats, and males and females differ in their response to scent cues over time. Scat piling has the potential to make information concerning the social environment available to dispersing transient and potential immigrant conspecifics, enabling settlement choices to be made. This thesis explores some of the behavioural strategies employed by E. whitii to reduce risks to individuals within groups and between groups. Scents eliciting a range of behavioural responses relevant to the formation of adaptive social groupings, reproductive activity, and juvenile protection until maturity and dispersal are likely to be present in this species. Tests confirming chemosensory cues that differentiate sex, kin and age would be an interesting addition to current knowledge. The interaction of delayed maturity, parental care, sociality, chemosensory communication and scat piling highlights the sophistication of this species¡¯ behaviour. An alternative method for permanently marking lizards was developed. Persistence, reliability and individual discrimination were demonstrated using photographic identification and the method was shown to be reliable for broad-scale application by researchers. Naturally occurring toe loss in the field provided a context against which to examine this alternative identification method and revealed the need to further investigate the consequences of routine toe clipping, as this practice appears to diminish survivorship.