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402 E. K. Madsen and F. Broekhuis The combined map shows that site use is highest east and


FIG. 3 Combined probability of site use values for the six species (the cheetah, elephant, hyaena, leopard, lion and wild dog; Fig. 2), outside the protected areas of the Maasai Mara (Fig. 1).


may not be the case as they are able to persist in human- dominated areas outside protected areas. Rivers influenced the presence of lions, leopards, wild


dogs and elephants and this could be related to the dense vegetation found beside rivers. Leopards, wild dogs and lions have been shown to select for similar habitats, includ- ing rivers, as they provide a cool environment during the day, denning opportunities for females with offspring, and increased hunting opportunities (Spong, 2002). De Knegt et al. (2011) theorized that, on a fine scale, elephants do not always need to be close to a water source as long as there is one within a day’s walking distance. In the Mara, water sources are fairly common and accessible and there- fore water availability is unlikely to be a limiting factor. It is therefore likely that the selection for rivers is a reflection of the habitat type occurring beside them. The individual species distribution maps are supported


by both sightings and data collected by GPS collars, in par- ticular those that have been fitted on cheetahs (Klaassen & Broekhuis, 2018) and wild dogs (Masenga et al., 2016). Lions have a much wider distribution than leopards, despite these species selecting for similar habitat (Balme et al., 2017b) and this probably reflects the higher densities at which lions occur in the Mara (Elliot & Gopalaswamy, 2017). Cheetahs and lions occur in similar areas and, although lions are a di- rect threat to cheetahs, these species can co-exist through fine-scale avoidance behaviour (Broekhuis et al., 2013). The inverse distribution of wild dogs and lions is similar to findings from Botswana (Cozzi, 2012) and South Africa (Darnell et al., 2014). Such avoidance behaviour affects the patterns of co-occurrence (MacKenzie et al., 2004). This illustrates the importance of taking a multi-species, rather than a single species, approach to management as the ex- pansion of protected areas could increase lion presence and numbers, which could have a negative impact on the population of wild dogs in the Mara. Potential species in- teractions therefore need to be taken into account when considering management practices.


north of the protected areas. Compared to other areas with a similar distance to protected areas, the area north of Naboisho and east of Olare-Motorogi conservancies (also known as the Pardamat Plains) had a low level of site use by all six species. This is probably because the open habitat provides little cover for wildlife. The area south of Naboisho and Ol Kinyei conservancies appears to have the highest site use for all species. This could prove problematic as the Sekanani road, which passes through this area, was, at the time of writing, being tarmacked. This could have negative consequences for the dispersal and movement of these species, and others, unless adequately mitigated. The covariates used in this studywere on a relatively broad


scale and it is possible that there are others affecting wildlife distribution on a finer scale. For example, elephants have been shown to have seasonal habitat preferences and fine scale preferences with respect to plant composition (Galanti et al., 2006; Shannon et al., 2006). Gradient and elevation, which were not taken into account in this study, influence leopard site use (Balme et al., 2017b). There could also be other factors that affect the respondents’ probability of seeing a species: not all pastoralistswill spend equal amounts of time with their livestock. One potential limitation of our study is that it was difficult to ensure that the interviewees reported only on their site and did not include sightings from other areas. However, the false positive model should have ac- counted for the potential for a small number of interviewees reporting on a wider area than intended. Interviewmethodology provides an opportunity to rapid-


ly assess the distribution of wildlife in unprotected areas and has the potential to be developed into a long-term mon- itoring programme. The use of the false positive model increases the robustness of the results derived from un- trained individuals (Petracca et al., 2018). In general, our findings corroborate published literature and expert opinion. We recommend the areas to the east, between the main pro- tected areas and the outer conservancies, which have the highest level of use, as a conservation priority, which could include the expansion of current conservancies and the cre- ation of new conservancies and wildlife corridors. This is particularly important in light of the increasing development in this area, including the erection of fences (Løvschal et al., 2017) and the tarmacking of a main road that passes through this region. However, increasing the amount of protected land can have a negative conservation outcome if local peo- ple become resentful, especially if they are displaced or access to resources is restricted. For successful conservation, land protection schemes need to take into account the needs of both wildlife and people (Roe, 2008).


Acknowledgements A component of this study was carried out in fulfilment of the Wild Animal Biology MSc (EKM) at the Royal Veterinary College and the Zoological Society of London. We thank


Oryx, 2020, 54(3), 395–404 © 2018 Fauna & Flora International doi:10.1017/S0030605318000297


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