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The Effects of Prey, Habitat Heterogeneity and Fire on the Spatial

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The Effects of Prey, Habitat Heterogeneity and Fire on the Spatial bs_bs_banner Austral Ecology (2014) 39, 181–189 The effects of prey, habitat heterogeneity and fire on the spatial ecology of peninsular Diamond Pythons (Morelia spilota spilota: Pythonidae) DAMIAN R. MICHAEL,* ROSS B. CUNNINGHAM, CHRISTOPHER MACGREGOR, DARREN BROWN AND DAVID B. LINDENMAYER Fenner School of Environment and Society, The Australian National University, Canberra, ACT 0200, Australia (Email: [email protected]) Abstract Understanding an organism’s home range is an important component of effective wildlife management. However, home ranges can vary spatially and temporally within and between populations. Landscape ecology theory can provide a framework for understanding spatio-temporal variability in animal traits. We used radio- telemetry in a population of diamond python Morelia spilota spilota Lacépède (Pythonidae) from a biologically rich and structurally heterogeneous reserve in eastern Australia to explore the relationship between home range size, optimal foraging theory and vegetation mosaic theory.Twelve adult snakes were tracked between September 2004 and February 2008. Male home ranges were significantly larger (P < 0.05) and more variable (41 Ϯ 30 ha) than female home ranges (23 Ϯ 5 ha), and males moved further between observations (123 m c.f. 65 m). Core activity centres varied significantly among habitat (P < 0.05) with larger home ranges observed in heathland, a vegetation community which supported comparatively low mammal diversity. No other variables examined including number of fixes, body length, prey abundance, vegetation heterogeneity or fire history explained home range variability. In this system, relatively high mammalian prey diversity and rapid post-fire vegetation succession may limit prey availability and fire effects as being significant determinants of home range variability in M. spilota. Key words: home range, optimal foraging theory, prey availability, pyro-diversity, vegetation heterogeneity. INTRODUCTION reptiles have the ability to reduce metabolic costs when there is a shortage of food, and incredibly high densi- Effective wildlife management is underpinned by ties of some species such as the red-sided garter snake knowledge of species resource requirements, including Thamnophis sirtalis (Gregory & Stewart 1975) may be an understanding of the factors that influence move- accounted for by this reptilian trait. ment patterns and space utilization (Caughley & Many early studies of snakes reported strong differ- Sinclair 1994). Developments in radio-telemetry over ences in home range size between sexes (Shine 1987; the past three decades have enabled researchers to Macartney et al. 1988; Slip & Shine 1988c; Weatherhead study space use in a wide range of organisms (White & Hoysak 1989), non-significant differences in home & Garrott 1990). Understanding differential spatial range size between sexes (e.g. Natrix natrix Madsen 1984) responses in vertebrates can result from examin- and contrasting home range size sex differences between ing density-dependent predator-prey relationships different populations (e.g. Morelia spilota imbricata (Tilman & Kareiva 1997). Many studies on large Pearson et al. 2005). These conflicting accounts obscure mammals have found inverse relationships with home general spatial patterns from being identified based solely range size and prey density. For example, home ranges on gender (Macartney et al. 1988). Clearly, there are of the Eurasian lynx Lynx lynx increased in response to other factors that influence space utilization in snakes, low densities of roe deer Capreolus capreolus (Herfindal including migratory behaviour (Gregory & Stewart 1975; et al. 2005). Litvaitis et al. (1986) found home ranges Larsen 1987; Neumann & Mebert 2011), agonistic inter- of the bobcat Felis rufus increased in response to actions (Whitaker & Shine 2003), landscape hetero- declining numbers of the snowshoe hare Lepus geneity (Hoss et al. 2010; Kapfer et al. 2010), and prey americanus. These patterns are often attributed to the distribution and abundance (Schwaner 1991; Madsen & high metabolic rate and energy expenditure costs Shine 1996; Whitaker & Shine 2003; Heard et al. 2004). that are characteristic of large carnivorous mammals Prey diversity can fluctuate over various spatial and (Carbone et al. 2011). In contrast, unlike mammals, temporal scales. For example, the relationship between fluctuating small mammal populations and climatic *Corresponding author. conditions is well documented (Dickman et al. 1999; Accepted for publication April 2013. Letnic & Dickman 2006; Greenville et al. 2012). © 2013 The Authors doi:10.1111/aec.12056 Austral Ecology © 2013 Ecological Society of Australia 182 D. R. MICHAEL ET AL. Resources and conditions that fluctuate unpredictably context of landscape ecology theories, as well as com- may cause individuals to utilize more space depending paring sex effects with other studies. on seasonal conditions (Brown & Shine 2002; Roe 1. Do home ranges differ between males and et al. 2004; Schofield et al. 2010). Whitaker and Shine females? We predicted that male home ranges will (2003) found home ranges in male eastern brown be larger than females because reproductive males snakes Pseudonaja textilis increased when populations tend to monopolize areas with access to many of the house mouse Mus musculus declined with females (Slip & Shine 1988c). drought conditions. Resource productivity also can be 2. Is there a relationship between prey abundance marginal near the edge of a species geographical range and home range size (sensu optimal foraging resulting in large home ranges (Arvisais et al. 2002; theory)? Degregorio et al. 2011). Optimal foraging theory 3. Is there a relationship between habitat attributes (sensu MacArthur & Pianka 1966) predicts that space (e.g. vegetation association and vegetation hetero- use will be related to the distribution and abundance geneity) and home range size (sensu vegetation of resources over large spatial-scales. Because animal mosaic theory)? movements often come at a cost of increased predation 4. Is there a relationship between fire attributes and risk and energy expenditure, one hypothesis is that home range size (sensu pyro-diversity theory)? animals will occupy the smallest home range that A central prediction of this study was that pythons which includes adequate critical resources, such as prey and occupy home ranges in burnt, highly heterogeneous veg- shelter sites, thereby minimizing movement associ- etation communities or in areas with high prey diversity ated costs (Hoss et al. 2010). Consequently, if these will have smaller home ranges than pythons which occupy resources are more accessible or abundant in particu- home ranges in unburnt, homogeneous vegetation com- lar environments then home range size may be directly munities or in areas with low prey diversity. related to prey abundance (Hoss et al. 2010). Variability in the abundance of prey for snakes such as terrestrial mammals can be a function of land use METHODS history such as fire regimes (Andersen et al. 2005), as well as intrinsic landscape features which are known to Study animal influence vegetation patterns such as soil and moisture gradients (Feldhamer 2010). These concepts give rise The diamond python Morelia spilota spilota is a colour morph to vegetation mosaic theory (sensu Wiens 1995) and of the widely distributed Australian carpet python Morelia pyro-diversity theory (sensu Martin & Sapsis 1992), spilota complex (Taylor 2005) and can attain a length of which predict that high variability in patterns of veg- 3.5 m (Wilson & Swan 2010). The species is semi-arboreal etation or fire can promote high levels of biotic diver- and can withstand long periods of reduced metabolic activity sity, which may in turn influence predator – prey space during the winter months (Slip & Shine 1988a). Morelia use dynamics. The relative effects of these ecological spilota is a non-venomous, ambush predator preying on a theories are often difficult to unravel as the effects of wide range of small mammals and lizards as a juvenile and fire on vegetation (and associated fauna) in heteroge- transitioning to prey such as medium-sized mammals and neous landscapes can produce complex and confound- nestling birds as an adult (Slip & Shine 1988b). The species is generally solitary but individuals aggregate during the ing interactions (Lindenmayer et al. 2008a; Williams breeding season and in some areas, spend the winter in et al. 2012). hibernacula such as deep rock crevices (Slip & Shine 1988a). The python Morelia spilota is an ideal species in Females lay a single clutch of up to 54 eggs in a nest of leaf which to study spatial ecology for several reasons. litter beneath dense vegetation, in tree cavities or hollow logs (i) Home ranges have previously been reported in (Slip & Shine 1988c). Incubation lasts for 60–80 days and M. spilota (e.g. Slip & Shine 1988a; Shine & Fitzgerald hatching occurs between late February and March. Along the 1996; Pearson et al. 2005; Corey & Doody 2010), east coast of Australia, M. spilota can attain high densities in enabling direct comparison with other studies. (ii) suitable habitat (Slip & Shine 1988a; Shine & Fitzgerald Home ranges are variable both within and between 1996; Fearn et al. 2001) despite evidence of a broader in- populations (Pearson et al. 2005) providing a founda- land decline (Shine 1994; Heard et al. 2004; Michael & Lindenmayer 2008; Corey & Doody 2010; Michael 2011). tion for testing the validity of ecological theory in
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