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INTRODUCTION
Habitat use is the way an animal uses physical and biological resources (Krausman 1999). Habitat selection by a species can be considered as a multi-level hierarchical process (Johnson 1980). Selection of specific habitats by a species can be considered on multiple levels - the landscape where it is distributed and hence evolved as a species over millions of years, the site that it selects as a home range as an individual, and the microhabitats that are used for food, survival and reproduction, again at the individual level. Ungulates of different size and metabolic rates have specific requirements in terms of nutrition and to fulfil these requirements occupy various spatial and temporal niches (Jarman 1974, Newing 2001). Ungulates also adopt specific anti-predatory strategies to protect themselves and their young ones from predators (Broom & Ruxton 2005, Caro et al. 2004, Leuthold 1977).
At the landscape level, the four-horned antelope (Tetracerus quadricornis Blainville 1816) uses tropical dry deciduous forest habitats (Baskaran et al. 2011, Berwick 1974, Krishna et al. 2008, 2009; Sharma 2006). Recent studies also indicate that its abundance and distribution is considerably affected by habitat changes because of local and landscape-level factors (Krishna et al. 2009). However, even in forest patches typified as tropical dry deciduous, the distribution of four-horned antelope is often not uniform. For instance, in the current study the four-horned antelope was not encountered in most areas within the lower plateau of Panna Tiger Reserve (PTR), despite being a tropical dry deciduous forest.
Several environmental, anthropogenic and ecological variables were believed to govern the spatial distribution of the four-horned antelope within a tropical dry deciduous forest. While some literature suggested that the species is found in undulating terrain (Brander 1923, Prater 1980), others suggested that its distribution is limited by water (Brander 1923, Prater 1980) and within well-wooded forests (Prater 1980, Rahmani 2001). Krishna et al. (2008) and Baskaran et al. (2011) highlighted its preference for open habitats, whereas Sharma et al. (2009) underscored the importance of undisturbed habitat and year-round fruit and flower availability. Although predators can also have strong indirect effects on habitat use by ungulates (Valeix et al. 2009), we assumed the predation pressure to be mostly uniform across the...