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Correlates of Active Body Temperatures and Microhabitat Occupation in Nine Species of Central Australian Agamid Lizards

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Correlates of Active Body Temperatures and Microhabitat Occupation in Nine Species of Central Australian Agamid Lizards Austral Ecology (2001) 26, 660–669 Correlates of active body temperatures and microhabitat occupation in nine species of central Australian agamid lizards JANE MELVILLE* AND JAMES A. SCHULTE II Department of Biology, Washington University, Campus Box 1137, St Louis, Missouri 63130, USA (Email: [email protected]) Abstract Body temperatures of active lizards and their correspondence with microhabitat occupation were studied for nine species of agamid lizards in the central Australian arid zone. Thermoregulatory behaviour was also documented using several measures, such as the use of shade and perch height. The effects of thermal environ- ment on lizard habitat occupation were hypothesized to be significant, because desert regions experience daily and seasonal extremes of temperature that are well in excess of a lizard’s preferred temperature range. All species, except Ctenophorus isolepis and Diporiphora winneckei, were found to have body temperatures that corresponded closely to ground and surface temperatures. Thermoregulatory behaviour was also found to be important through- out a lizard’s daily activity; all study species, other than Ctenophorus isolepis, were found to increase their perch height in the middle of the day. Ctenophorus isolepis was shown to be a strictly terrestrial species that uses the shade of spinifex in its thermoregulatory behaviour. Species exhibited a non-random selection of microhabitats and a preference for a particular set of thermal and structural factors. In this study, it was shown that structural factors were particularly important in microhabitat occupation. Thermal factors accounted for a smaller proportion of variance in microhabitat occupation, but still played a considerable role in the microhabitat use in central Australian agamids. Key words: activity patterns, Agamidae, arid-zone lizards, ecology, microhabitat occupation, thermoregulatory behaviour. INTRODUCTION lizards are constrained to occupy specific thermal microclimates when they are active: to feed, defend Lizard ecologists have placed much importance on the territories, acquire mates and avoid predators. These study of microhabitat occupation, usually by investi- constraints are especially important for desert lizards gating resource partitioning and interspecific compe- that occupy an environment of extreme temperatures, tition (Toft 1985). Despite the importance of so that failure to select an appropriate thermal micro- temperature in lizard ecology, comparatively few climate for even a few minutes can result in danger- studies have considered the influence of thermoregu- ously high body temperatures or even death. lation on habitat use in desert regions (Huey & Pianka Consequently, the availability of thermally appropri- 1977; Avery 1978; Adolph 1990). Physiological and ate microhabitats is potentially a major limiting factor biochemical processes are strongly influenced by body affecting lizards in desert regions. Thermal constraints temperature (Huey et al. 1991). Reproduction, rates of vary in time and space because the physical environ- growth, feeding and mobility are whole-body functions ment is temporally and spatially variable with respect dramatically affected by temperature (Huey 1982). to a variety of factors, such as the thermal inertia of Consequently, temperature is the most important substrates, air temperature and vegetation structure physical factor in the ecology of lizards (Heatwole (Hertz et al. 1993). These climatic factors interact with 1976) and a large proportion of a lizard’s day is spent the surface heat transfer properties of a lizard to pro- responding to the thermal environment. duce a distribution of available temperatures in time In desert regions, with daily and seasonal extremes and space within which a lizard can survive (Stevenson of weather, thermal biology may restrict the use of 1985). Thus, a lizard will occupy a restricted range of microhabitats by lizards. Thermal microhabitat con- microhabitats within its overall habitat. straints arise because the body temperature of an There have been very few detailed studies examin- ectotherm is a complex function of its biophysical ing the habitat occupation of agamid lizards, especially environment (Gates 1980). Thus, thermoregulating those in desert regions and our study provides a quantification of structural and thermal microhabitat *Corresponding author. occupation in a number of these species. Agamids Accepted for publication May 2001. occupy a wide variety of habitats in Australia, such as BODY TEMPERATURE AND ECOLOGY OF DESERT LIZARDS 661 tropical rainforests (Hypsilurus boydii), cool temperate Field work woodlands (Tympanocryptis diemensis), stony deserts (Tympanocryptis lineata), desert dunes (Ctenophorus Structural and thermal microhabitat characteristics isolepis) and dry salt lakes (Ctenophorus maculosus). were recorded at four field sites (Fig. 1). Field sites were However, they reach their highest diversity in the semi- chosen to provide a wide range of habitat types, includ- arid and arid regions of Australia. The alpha diversity ing flat spinifex desert (sites 2, 3 and 4), sand ridges of lizards in arid Australia is high relative to deserts on (sites 3 and 4), rocky spinifex desert (site 1), desert other continents (Pianka 1986). This has been ranges (sites 1 and 3), gibber plains (site 4), acacia attributed to the diversity of microhabitats and the woodlands (sites 1, 2, 3 and 4) and riverine eucalypts diversity of food resources (Pianka 1986), which (sites 1, 2 and 3). The data were recorded by driving vary dramatically with habitat type (Morton & James or walking throughout each field site from approxi- 1988). For this reason, and the fact that temperature mately 07.00 until 19.00 hours. This method ensured plays an important role in desert environments, we coverage of the entire range of structural and thermal tested the hypothesis that active body temperature is microhabitats available. This was done over a 6-week correlated with microhabitat occupation in nine species period from October to December 1999, with approxi- of desert agamids. We examined this by recording mately 10 days spent at each location. Data were only active field body temperatures and quantifying the collected on sunny, warm days when lizards could be thermal and structural microhabitat occupation of fully active. the study species. Active lizards were caught with the use of a noose or by hand. Body temperature was recorded by using a METHODS Study species Agamids are diurnal heliotherms that are primarily sit-and-wait predators (Huey & Pianka 1981). They comprise approximately 20% of the lizard fauna in desert regions of Australia (Pianka 1986). We studied nine agamid species at four sites in central Australia (Fig. 1). Eleven agamid species were caught in the present study but the only species included in the analy- sis were those for which we caught greater than 15 indi- viduals. The two species not included were Ctenophorus clayi (one individual was caught at field site 3) and Pogona mitchelli (two individuals were caught at field Fig. 1. Map of the four field sites. All field work was con- site 1). The nine agamid species included in the pre- ducted in National Parks or on leasehold land. Field site 1 sent study are made up of five genera: Ctenophorus, included Larapinta and Namatjira Drives, West MacDonnell Diporiphora, Lophognathus, Pogona and Tympanocryptis. Ranges National Park and Finke Gorge National Park; field site 2 included the Old South Road, Roe Creek and Stuart All species, except C. pictus, were caught at more that Highway (north of Alice Springs); field site 3 included one field site (Table 1). The Tympanocryptis lineata Watarrka National Park; field site 4 included the four-wheel- caught in the present study all belonged to the sub- drive track between Mount Dare Station and Andado species centralis. Station. Ta bl e 1 . Presence of agamid species at each field site and general description of habitats in which the species occurs Species Field site no. Habitat category General habitat description Ctenophorus caudicinctus 1, 3 Saxicolous Desert ranges; rocky outcrops Ctenophorus isolepis 2, 4 Ground-dwelling Sandy spinifex desert Ctenophorus nuchalis 1, 2, 3, 4 Ground-dwelling All open habitats with non-rocky substrate Ctenophorus pictus 4 Ground-dwelling Sandy substrates with open shrub cover Diporiphora winneckei 3, 4 Semi-arboreal Sandy spinifex desert Lophognathus gilberti 1, 2 Semi-arboreal Mulga woodlands Lophognathus longirostris 1, 2, 3 Semi-arboreal Mulga and riverine woodlands Pogona vitticeps 1, 2, 4 Semi-arboreal Open habitats with heterogeneous vegetation Tympanocryptis lineata 1, 4 Saxicolous Rocky spinifex desert 662 J. MELVILLE AND J. A. SCHULTE II thermistor and lizards were sexed before being released. number of components that describes the underlying Ten structural and three thermal characteristics of the dimensions in the data. The principal components were microhabitat for each lizard caught were recorded. The extracted from a correlation matrix of the raw data. The structural variables were: 1, visual estimate of percent number of principal components (henceforth PCs) canopy cover in a 3-m2 radius surrounding lizard; 2, utilized in the analysis was determined by using the visual estimate of percentage shrub cover in a 3-m2 scree test of eigenvalues plotted against factors, maxi- radius surrounding lizard; 3, visual estimate of percen- mizing the adequacy of extraction. Principal compon- tage ground and grass cover in
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