Experimental analysis of retreat‐site selection by thick‐tailed geckos Nephrurus milii

Abstract Although animals use habitats non‐randomly in the wild, complex correlations among environmental features mean that proximate influences on habitat selection can be identified only by experimental manipulation of potential cues. Thick‐tailed geckos Nephrurus milii are large lizards that are widely distributed through southern Australia. These nocturnally active animals typically spend daylight hours under surface rocks. We presented captive geckos with alternative retreat‐sites (rock crevices) differing in attributes potentially relevant to habitat selection. The lizards showed strong preference for shelter‐sites that enhanced thermoregulation (warm rather than cool) and that reduced the animal's vulnerability to predators (narrow crevices with small openings and not containing the scent of a predatory snake). Horizontal rather than sloping crevices were also preferred. Overall crevice size and thickness of the overlying rock did not influence retreat‐site selection in the laboratory, but could be important in the field because of their influence on thermal regimes under rocks. The present study supports the idea that nocturnal reptiles base their selection of diurnal shelters on multiple aspects related to the fitness consequences of occupancy of alternative available retreat‐sites.     

Forest‐fire regimes affect thermoregulatory opportunities for terrestrial ectotherms

Fire‐induced changes in canopy openness may affect sunlight penetration to the forest floor, and thus the operative temperatures available to terrestrial ectotherms. We examined thermal regimes for two types of ectotherms: diurnally active species that utilize sun‐exposed patches to regulate their body temperatures, and nocturnally active species that depend upon solar radiation striking the rocks under which they shelter. We measured canopy openness, shrub height, radiation transmission and operative environmental temperatures in the open and inside reptile retreat‐sites, at 24 study sites in eucalypt forests in two regions (Gosford and Yengo) in south‐eastern Australia. All sites were last burnt in 2000–2001, but had experienced different fire frequencies (1–4 fires over the previous 37 years). In Gosford, higher fire frequencies reduced canopy openness and radiation transmission at ground and shrub level, and thus reduced environmental temperatures and the thermal quality of reptile habitats. Our modelling based on thermal preferenda of an endangered snake species (the broad‐headed snake Hoplocephalus bungaroides) suggests that increased fire frequency at Gosford halved the amount of time an animal could spend within its preferred (set‐point) range, regardless of whether it thermoregulated beneath rocks or basked out in the open. At Yengo, however, fire frequency did not affect the thermal quality of reptile habitats. Thus, the effects of fire frequency on forest structure and the thermal environment at ground level differed between adjacent areas, and relatively small changes in canopy openness translated into major effects on thermoregulatory opportunities for reptiles. Although fire is a useful management tool for creating open habitats, we need to understand more about the effects of fire frequency on vegetation structure and thermal environment before we can use fire to manage habitats for reptiles.     

Does rock disturbance by superb lyrebirds (Menura novaehollandiae) influence habitat selection by juvenile snakes?

Abstract: Vertebrates that destroy or disturb habitats used by other animals may influence habitat selection by sympatric taxa. In south‐east Australian forests, superb lyrebirds (Menura novaehollandiae) displace soil, leaf litter and rocks during their daily foraging activities. We investigated whether superb lyrebirds disturb small sandstone rocks that endangered broad‐headed snakes Hoplocephalus bungaroides and common small‐eyed snakes Cryptophis nigrescens use as diurnal thermoregulatory sites. To estimate the frequency of lyrebird rock disturbance, and to assess whether lyrebirds also attack small snakes, we placed 900 plasticine snake replicas under stones on rock and soil substrates along transects on three sandstone plateaux. Because juvenile snakes must select retreat sites that simultaneously allow them to thermoregulate and minimize predation risk, we quantified the thermal environments underneath stones on rock and soil substrates. During the 6‐week experiment, animals disturbed rocks on soil substrates twice as often (16.9%) as rocks lying on rock substrates (8.2%). Disturbed rocks were significantly smaller and lighter than undisturbed rocks on both substrates. Lyrebirds were the major agents of disturbance, and attacked 40% of snake models under disturbed rocks. Rocks on soil substrates conferred the greatest thermal benefits to snakes, but both species of snake avoided these microhabitats in the field. Instead, juvenile snakes selected rocks on rock substrates, and sheltered under stones that were too heavy for superb lyrebirds to disturb. By disturbing rocks over millennia, superb lyrebirds not only have shaped the physical landscape, but also may have exerted strong selection on habitat selection by sympatric snakes.     

Habitat selection in a rocky landscape: experimentally decoupling the influence of retreat site attributes from that of landscape features

Organisms selecting retreat sites may evaluate not only the quality of the specific shelter, but also the proximity of that site to resources in the surrounding area. Distinguishing between habitat selection at these two spatial scales is complicated by co-variation among microhabitat factors (i.e., the attributes of individual retreat sites often correlate with their proximity to landscape features). Disentangling this co-variation may facilitate the restoration or conservation of threatened systems. To experimentally examine the role of landscape attributes in determining retreat-site quality for saxicolous ectotherms, we deployed 198 identical artificial rocks in open (sun-exposed) sites on sandstone outcrops in southeastern Australia, and recorded faunal usage of those retreat sites over the next 29 months. Several landscape-scale attributes were associated with occupancy of experimental rocks, but different features were important for different species. For example, endangered broad-headed snakes (Hoplocephalus bungaroides) preferred retreat sites close to cliff edges, flat rock spiders (Hemicloea major) preferred small outcrops, and velvet geckos (Oedura lesueurii) preferred rocks close to the cliff edge with higher-than-average sun exposure. Standardized retreat sites can provide robust experimental data on the effects of landscape-scale attributes on retreat site selection, revealing interspecific divergences among sympatric taxa that use similar habitats.     

Factors affecting retreat‐site selection by coppertail skinks (Ctenotus taeniolatus) from sandstone outcrops in eastern Australia

Abstract Coppertail skinks (Ctenotus taeniolatus) are medium‐sized diurnal lizards that are widespread in eastern Australia. Field surveys on sandstone outcrops in the Sydney region showed that these animals use available rocks non‐randomly: coppertails were found under large, sun‐exposed rocks on deep soil. Choice trials in the laboratory documented selection of retreat sites that were warm, with substrate preferences shifting in a diel cycle. Lizards selected retreats with rock substrates during the day but with sandy substrates at night. Rocky retreats attain higher temperatures during daylight hours, but burrows in sand beneath rocks may provide greater protection against ingress by nocturnal predators. During fieldwork we rarely found lizards under rocks with either ants or centipedes, suggesting that coppertails may avoid these predatory invertebrates. Tongue‐flick trials showed that the lizards could discriminate among common ant species based on chemosensory cues, but apparently could not detect centipede chemical cues. In experimental trials, the lizards did not avoid retreat sites scented by either ants or centipedes. Our data thus suggest that retreat‐site selection in coppertails is driven by abiotic cues (rock size, sun exposure and substrate type) that may confer fitness benefits in terms of thermoregulation and predator avoidance, with biotic cues playing a less important role.     

Mapping microhabitat thermal patterns in artificial breakwaters: Alteration of intertidal biodiversity by higher rock temperature

Urbanization is altering community structure and functioning in marine ecosystems, but knowledge about the mechanisms driving loss of species diversity is still limited. Here, we examine rock thermal patterns in artificial breakwaters and test whether they have higher and spatially less variable rock temperature than natural adjacent habitats, which corresponds with lower biodiversity patterns. We estimated rock temperatures at mid‐high intertidal using infrared thermography during mid‐day in summer, in both artificial (Rip‐raps) and natural (boulder fields) habitats. We also conducted diurnal thermal surveys (every 4 hr) in four seasons at one study site. Concurrent sampling of air and seawater temperature, wind velocity, and topographic structure of habitats were considered to explore their influence on rock temperature. Rock temperature was in average 3.7°C higher in the artificial breakwater in two of the three study sites, while air temperature was about 1.5–4°C higher at this habitat at summer. Thermal patterns were more homogeneous across the artificial habitat. Lower species abundance and richness in the artificial breakwaters were associated with higher rock temperature. Mechanism underlying enhanced substrate temperature in the artificial structures seems related to their lower small‐scale spatial heterogeneity. Our study thus highlighted that higher rock temperature in artificial breakwaters can contribute to loss of biodiversity and that integrated artificial structures may alter coastal urban microclimates, a matter that should be considered in the spatial planning of urban coastal ecosystems.     

Differential Avoidance of Snake Odours by a Lizard: Evidence for Prioritized Avoidance Based on Risk

Most studies of predator avoidance behaviours have focussed on single‐predator systems, despite the fact that prey often are confronted with predator rich environments. In the presence of more than one predator, prey may have to choose between avoiding one predator over another. How prey cope with exposure to several enemies simultaneously remains largely untested. In this study I set out to investigate if skinks showed preferential avoidance of snake odours based on the relative predation risk posed by different snake species. This relative predation risk was estimated using information on density, diet specificity and foraging habit of each snake species. I tested retreat‐site selection in two‐choice tests, where lizards chose between different combinations of control and snake treated retreat‐sites as well as two retreat‐sites treated with different snake species odours. Lizards preferred control–treated retreat‐sites to those treated with snake odours and showed a differential avoidance response to refuges treated with odours from different snake species. There was strong evidence to suggest that lizards preferentially avoided refuges with the odours of the snake that posed the greatest predation risk, the white‐lipped snake (Drysdalia coronoides). Naïve juvenile lizards were also tested and their response was similar to the adults demonstrating that the behaviour is innate and not the result of higher encounter rates of more common snake odours. To my knowledge this is one of the first studies to demonstrate that prey can prioritize avoidance to a single most dangerous predator in the face of several predators and conflicting avoidance responses.     

Effect of disturbance on assemblages: an example using porifera

Extensive sponge assemblages are found in a number of habitats at Lough Hyne Marine Nature Reserve. These habitats are unusual in experiencing a range of environmental conditions, even though they are only separated by small geographic distances (1-500 m), reducing the possibility of confounding effects between study sites (e.g., silica concentrations and temperature). Sponge assemblages were examined on ephemeral (rocks), stable (cliffs), and artificial (slate panels) hard substrata from high- and low-energy environments that were used to represent two measures of disturbance (flow rate and habitat stability). Sponge assemblages varied considerably between habitat types such that only 26% (25 species) of species reported were common to both rock and cliff habitats. Seven species (of a total of 96 species) were found in the least-developed assemblages (slate panels) and were common to all habitats. Sponge assemblages on rocks and panels varied little between high- and low-energy environments, whereas assemblages inhabiting cliffs varied considerably. Assemblage composition was visualized using Bray-Curtis similarity analysis and Multi-Dimensional Scaling, which enabled differences and similarities between sponge assemblages to be visualized. Cliffs from high- and low-energy sites had different assemblage compositions compared to large rocks, small rocks, and panels, all of which had similar assemblages irrespective of environmental conditions. Differences in assemblages were partially attributed to sponge morphology (shape), as certain morphologies (e.g., arborescent species) were excluded from 2-D rock habitats. Other mechanisms were also considered responsible for the sponge assemblages associated with different habitats.     

Nesting in a thermally challenging environment: nest‐site selection in a rock‐dwelling gecko,Oedura lesueurii (Reptilia: Gekkonidae)

In egg‐laying species, maternal oviposition choice can influence egg survival and offspring phenotypes. According to the maternal‐preference offspring‐performance hypothesis, females should choose oviposition sites that are optimal for offspring fitness. However, in thermally challenging environments, maternal oviposition behaviour may be constrained by the limited availability of suitable oviposition sites. We investigated nest‐site selection in a nocturnal lizard [velvet gecko Oedura lesueurii (Duméril and Bibron)] that inhabits a thermally challenging environment in south‐eastern Australia. The viability of these gecko populations is critical for the persistence of an endangered snake species (Hoplocephalus bungaroides Wagler) that feeds heavily on velvet geckos. Female geckos chose nest sites nonrandomly, with 87% of nests (N = 30) being laid in deep crevices. By contrast, only 13% of clutches were laid under rocks, which were the most readily available potential nest sites. Nest success in crevices was high (100%), but no eggs hatched from nests under rocks. Temperatures in nest crevices remained relatively low and constant throughout the incubation period (mean = 22.7 °C, range 21.0–24.5 °C), whereas thermal regimes under rocks showed large diurnal fluctuations. Geckos selected crevices that were deeper, had less canopy cover, and were warmer than most available crevices; in 85% of cases, such crevices were used simultaneously by more than one female. The thermally distinctive attributes of nest sites, and their frequent communal use, suggest that nest sites are a scarce resource for female velvet geckos, and that the shading of rock outcrops through vegetation encroachment may influence nest success in this species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 250–259.     

Plant species coexistence in cliff habitats

An ecological analysis of the ledge and crevice vegetation of cliffs in north-east Ireland was carried out using a sampling programme based on multivariate land classification. The composition of the vegetation was shown to be determined primarily by distance from the sea, elevation and rock type. The chemical composition and physical structure of the cliffs, together with environmental gradients associated with aspect, elevation and nearness to the sea provide a wide range of habitats in which plants coexist. Certain types of cliff vegetation contained associations of species with a contrasting ecology, such as arctic–montane, maritime, woodland, calcicole and calcifuge plants. The positions of these species in ordination space were found to be unique but overlapping. A model is described to account for these observations. It proposes that species dominance in cliff habitats is prevented by factors such as physical disturbance and that plants are able to coexist in cliff habitats which meet their common environmental tolerances and preferences. The model predicts that cliff habitats with the greatest diversity of species are likely to be subject to less extremes of environmental variability. These habitats include inland sites with nutrient enrichment by sea spray, north-facing sites, seepage sites, base-rich sites, low elevation sites and sites with a heterogeneous physical structure.     

A cost‐effective method of assessing thermal habitat quality for endotherms

The conservation of many endothermic species depends critically on the availability of suitable retreat sites, yet we know little about the variation in thermal quality of such microhabitats. Studies of thermal habitat suitability for birds and mammals must account for the effect of endothermic heat production on their microclimates. For example, endotherms may significantly raise the air temperature in their retreat sites and this effect must be considered when assessing retreat site quality. We devised an inexpensive means by which to construct pseudo‐endothermic ‘environmental temperature’ models with the use of disposable heat pads. We applied this technique to investigate thermal aspects of nest box design, illustrating the potential positive and negative effects of nest box insulation depending on the environmental context. We suggest that, from a thermal perspective, the avoidance of heat stress is an important and underappreciated issue in the retreat site selection of endotherms.     

Benefits of parental care: Do juvenile lizards obtain better‐quality habitat by remaining with their parents?

Abstract Evolutionary theory suggests that parental care is favoured by natural selection when the benefits to offspring fitness outweigh the costs of parental expenditure. The nature of such benefits may differ among species, however, especially in species reflecting independent evolutionary origins of parental care. Black rock skinks (Egernia saxatilis, Scincidae) of south‐eastern Australia are viviparous rock‐dwelling lizards with prolonged parent–offspring association; adult pairs vigorously defend their home range – and, when present, their offspring – against conspecifics. We addressed the hypothesis that, by remaining within their parents' (vigorously defended) home range, juveniles thereby obtain access to better‐quality habitats. Measurements of biologically significant variables (crevice size, sun exposure, vegetation cover) revealed little difference between shelter‐rocks used by solitary (‘orphan’) juveniles and those within family groups containing adults. Indeed, the only consistent differences involved larger (and therefore, less predator‐proof) crevices for juveniles within family groups than for solitary conspecifics. Our data thus falsify the hypothesis that parental care evolves because of benefits associated with habitat quality; instead, it appears that parental protection of juveniles against infanticidal conspecifics may be the most plausible benefit to parent–offspring association in this system.     

Habitat use affects morphological diversification in dragon lizards

Habitat use may lead to variation in diversity among evolutionary lineages because habitats differ in the variety of ways they allow for species to make a living. Here, we show that structural habitats contribute to differential diversification of limb and body form in dragon lizards (Agamidae). Based on phylogenetic analysis and ancestral state reconstructions for 90 species, we find that multiple lineages have independently adopted each of four habitat use types: rock‐dwelling, terrestriality, semi‐arboreality and arboreality. Given these reconstructions, we fit models of evolution to species’ morphological trait values and find that rock‐dwelling and arboreality limit diversification relative to terrestriality and semi‐arboreality. Models preferred by Akaike information criterion infer slower rates of size and shape evolution in lineages inferred to occupy rocks and trees, and model‐averaged rate estimates are slowest for these habitat types. These results suggest that ground‐dwelling facilitates ecomorphological differentiation and that use of trees or rocks impedes diversification.     

Fish assemblages associated with urban structures and natural reefs in Sydney,Australia

Abstract Fish ecology in urban estuaries is poorly understood. As coastal landscapes are transformed, recognizing the impact that urban structures, such as marinas, seawalls and wharfs, have on local fish populations is becoming increasingly important. The extent to which fish are able to maintain natural ecological assemblages can be measured, to a certain extent, by how closely they mimic natural habitats. In Sydney Harbour, assemblages of fish associated with artificial structures were compared with those associated with natural rocky reefs. Sampling was carried out in five locations, each with a marina, swimming enclosure and natural reef. In each location, different habitats supported different assemblages, but differences between habitats were not consistent among locations. Subsequent sampling compared artificial and natural sites in three different areas in each of three different estuaries. Results indicated that differences in fish assemblages between artificial and natural sites were greater than differences between sites within each habitat, but there were no patterns among different positions in an estuary or from estuary to estuary. This study provides initial evidence that, although artificial habitats generally support the same species as found on natural reefs, assemblages usually differed between natural and artificial habitats. In addition, without knowing if these habitats do, in fact, sustain viable populations of fish, it would be premature to label artificial structures as effective habitat for fish.     

The importance of water‐retaining features for biodiversity on artificial intertidal coastal defence structures

Aim

Artificial coastal defence structures are proliferating in response to rising and stormier seas. These structures provide habitat for many species but generally support lower biodiversity than natural habitats. This is primarily due to the absence of environmental heterogeneity and water‐retaining features on artificial structures. We compared the epibiotic communities associated with artificial coastal defence structures and natural habitats to ask the following questions: (1) is species richness on emergent substrata greater in natural than artificial habitats and is the magnitude of this difference greater at mid than upper tidal levels; (2) is species richness greater in rock pools than emergent substrata and is the magnitude of this difference greater in artificial than natural habitats; and (3) in artificial habitats, is species richness in rock pools greater at mid than upper tidal levels?

Location

British Isles.

Methods

Standard non‐destructive random sampling compared the effect of habitat type and tidal height on epibiota on natural rocky shores and artificial coastal defence structures.

Results

Natural emergent substrata supported greater species richness than artificial substrata. Species richness was greater at mid than upper tidal levels, particularly in artificial habitats. Rock pools supported greater species richness than emergent substrata, and this difference was more pronounced in artificial than natural habitats. Rock pools in artificial habitats supported greater species richness at mid than upper tidal levels.

Main conclusions

Artificial structures support lower biodiversity than natural habitats. This is primarily due to the lack of habitat heterogeneity in artificial habitats. Artificial structures can be modified to provide rock pools that promote biodiversity. The effect of rock pool creation will be more pronounced at mid than upper tidal levels. The challenge now is to establish at what tidal height the effect of pools becomes negligible and to determine the rock pool dimensions for optimum habitat enhancement.

     

Variation in subsurface thermal characteristics of microrefuges used by range core and peripheral populations of the American pika (Ochotona princeps)

Microrefuges provide microclimates decoupled from inhospitable regional climate regimes that enable range‐peripheral populations to persist and are important to cold‐adapted species in an era of accelerated climate change. However, identifying and describing the thermal characteristics of microrefuge habitats is challenging, particularly for mobile organisms in cryptic, patchy habitats. We examined variation in subsurface thermal conditions of microrefuge habitats among different rock substrate types used by the American pika (Ochotona princeps ), a climate‐sensitive, rock‐dwelling Lagomorph. We compared subsurface temperatures in talus and lava substrates in pika survey sites in two US national park units; one park study area on the range periphery and the other in the range core. We deployed paired sensors to examine within‐site temperature variation. We hypothesized that subsurface temperatures within occupied sites and structurally complex substrates would be cooler in summer and warmer in winter than unoccupied and less complex sites. Although within‐site variability was high, with correlations between paired sensors as low as 47%, we found compelling evidence that pikas occupy microrefuge habitats where subsurface conditions provide more thermal stability than in unoccupied microhabitats. The percentage of days in which microhabitat temperatures were between ?2.5 and 25.5°C was significantly higher in occupied sites. Interestingly, thermal conditions were substantially more stable (p  < .05) in the lava substrate type identified to be preferentially used by pikas (pahoehoe vs. a'a) in a previous study. Our study and others suggest that thermal stability appears to be the defining characteristic of subsurface microrefuges used by American pikas and is a likely explanation for enigmatic population persistence at the range periphery. Our study exemplifies an integrated approach for studying complex microhabitat conditions, paired with site use surveys and contextualized with information about gene flow provided by complementary studies.     

How should a clever baboon choose and move rocks?

Chacma baboons (Papio ursinus) intentionally overturn rocks to feed on the invertebrates beneath. However, baboons do not move all the rocks they encounter, with this presumably reflecting cost–benefit (or effort–reward) trade‐offs in their foraging behavior. We ask, how do “clever baboons” choose rock sizes and shapes and move these rocks? Using optimal foraging theory, we predicted that baboons would prefer to move medium‐sized rocks, a trade‐off between moving larger rocks that might require more effort to move, and smaller rocks that likely do not provide enough prey (the reward) to make the effort worthwhile. We also expected baboons to prefer rounded rocks as these will require less energy to move by rolling (rather than being flipped as for flat rocks) and that the effort of rock movement might be offset by moving rocks along the shortest axis. We show that baboons have clear preferences for specific rock sizes (medium‐sized) and shapes (angular and flat when these were medium‐sized), and the way in which rocks are moved (along the shortest axis). Prey occurred infrequently under rocks. The low predictability of prey beneath rocks suggests that such prey, when encountered, is of considerable value to baboons for them to expend the search effort, and also explains the extensive nature of rock movement by baboons in the landscape. Our study provides a novel application of the optimal foraging theory concept and has important implications for understanding and predicting how animals choose to move rocks.     

Multiple habitat use by declining migratory birds necessitates joined‐up conservation

Many species depend on multiple habitats at different points in space and time. Their effective conservation requires an understanding of how and when each habitat is used, coupled with adequate protection. Migratory shorebirds use intertidal and supratidal wetlands, both of which are affected by coastal landscape change. Yet the extent to which shorebirds use artificial supratidal habitats, particularly at highly developed stopover sites, remains poorly understood leading to potential deficiencies in habitat management. We surveyed shorebirds on their southward migration in southern Jiangsu, a critical stopover region in the East Asian Australasian Flyway (EAAF), to measure their use of artificial supratidal habitats and assess linkages between intertidal and supratidal habitat use. To inform management, we examined how biophysical features influenced occupancy of supratidal habitats, and whether these habitats were used for roosting or foraging. We found that shorebirds at four of five sites were limited to artificial supratidal habitats at high tide for ~11–25 days per month because natural intertidal flats were completely covered by seawater. Within the supratidal landscape, at least 37 shorebird species aggregated on artificial wetlands, and shorebirds were more abundant on larger ponds with less water cover, less vegetation, at least one unvegetated bund, and fewer built structures nearby. Artificial supratidal habitats were rarely used for foraging and rarely occupied when intertidal flats were available, underscoring the complementarity between supratidal roosting habitat and intertidal foraging habitat. Joined‐up artificial supratidal management and natural intertidal habitat conservation are clearly required at our study site given the simultaneous dependence by over 35,000 migrating shorebirds on both habitats. Guided by observed patterns of habitat use, there is a clear opportunity to improve habitat condition by working with local land custodians to consider shorebird habitat requirements when managing supratidal ponds. This approach is likely applicable to shorebird sites throughout the EAAF.     

Habitat selection by grassland snakes at northern range limits: Implications for conservation

Animal populations at northern range limits may use habitat differently from those at range cores, requiring distinct conservation plans. Snakes are ectotherms that often have very specific requirements, but few studies have focused on the effect of northern latitudes on habitat selection by grassland snakes. We studied movement and habitat selection of 2 sympatric snake species at their northern range limits on the North American Great Plains: the eastern yellow-bellied racer (Coluber constrictor flaviventris, hereafter racer), a Threatened species in Canada, and the bullsnake (Pituophis catenifer sayi), which is listed as Data Deficient. Both of these species are potentially vulnerable to extinction in Canada because of habitat loss. Snakes from our study populations traveled up to 10-times farther from winter dens and occupied home ranges 3–104 times larger than populations further south. Both snake species moved from winter dens in the slopes of a major river valley to habitat in adjacent lowlands, including riparian zones (racers) and hilly areas with native grass species (bullsnakes). Multivariate modeling revealed that proximity to retreat sites was a significant predictor of snake site use for both species. Considering the need for winter dens and summering areas, our data suggest that snakes in northern latitudes should ideally have much larger protected areas compared to snakes near the core of their range. An alternative strategy is to conserve corridors linking wintering dens and summer habitats. Retreat sites such as burrows and shrubs are critical components of local habitat and should be included in conservation plans. © 2011 The Wildlife Society.     

The social elite: Habitat heterogeneity,complexity and quality in granite inselbergs influence patterns of aggregation in Egernia striolata (Lygosominae: Scincidae)

Habitat heterogeneity, structural complexity and habitat quality are key features of the environment that drive species' distribution and patterns of biological organization. Traditionally, pattern‐based studies have focused on faunal responses to biological systems. However, the influence of non‐biological environments such as insular rock outcrops on patterns of vertebrate distribution is conceivably as important, but has received less attention. Granite inselbergs are a naturally heterogeneous and spatially‐limited habitat. As such, they provide an opportunity for investigating whether environmental attributes influence social behaviour in animals that use these kinds of habitat, particularly lizards that are well adapted to saxicoline environments. We applied ecological theory to investigate the influence of habitat heterogeneity, structural complexity and habitat quality on patterns of home‐site occupancy in the crevice skink Egernia striolata (Lygosominea: Scincidae) from insular granite outcrops located within fragmented agricultural landscapes. We compared home‐site occupancy among solitary juveniles, solitary adults and lizard aggregations. We found significant differences in home‐site occupancy between aggregations and solitary lizard outcrop attributes measured at multiple spatial scales. The probability of a home‐site being occupied by an aggregation increased where large rock masses were present, on northern aspects near the core of the outcrop and in structurally variegated landscapes. Significantly more aggregations occupied home‐sites surrounded by high boulder cover and crevice microhabitat. We provide evidence that geophysical attributes of granite inselbergs and landscape context can influence patterns of lizard aggregation. Thus, we clearly document the environmental correlations of variability in sociality among subpopulations of Egernia striolata.