Multi-species site use 399
hyaenas, one did not converge so only the top model was used for prediction of site use. For all species the detection probability coefficients improved the predictive ability of the model compared to the null model. The proportion of open habitat had the best predictive ability for the probabil- ity of detecting cheetahs, lions and wild dogs. For lions the probability of detection increased with proportion of open habitat, and for wild dogs and cheetahs the probability of detection decreased, but minimally (Table 2). For elephants and leopards both the respondent’s occupation and the proportion of open habitat influenced the detection prob- ability. The proportion of open habitat increased the probability of detecting elephants, but was less important for leopards, with confidence intervals crossing zero. For hyaenas only the occupation of the respondent was in the final model. All species except hyaenas had a habitat parameter in the
final occupancy models (Table 2). In the univariate habitat covariate selection, the probabilities of site use by lions and wild dogs were best predicted by the proportion of semi- closed habitat, with both having a positive relationship, as predicted (Table 1). Site use by cheetahs, elephants and leo- pards, however, was best predicted by the proportion of open habitat. Probability of site use by cheetahs increased with proportion of open habitat, which was expected (Table 1), whereas the probability of site use by both ele- phants and leopards decreased with the proportion of open habitat. All six species were affected by human pres- ence (Table 2). For cheetahs, lions, leopards and hyaenas the human disturbance covariate with the best fit was the mean human distance in the univariate analysis, with all showing a preference for sites further from human presence, which was expected for all species except leopards (Table 1). However, for both lions and leopards this effect was min- imal. The probability of site use by both elephants and wild dogs decreased with an increased proportion of fences, which had the best predictive value of the human distur- bance covariates. All six species contained the mean dis- tance to protected area in their top models, with all except hyaenas decreasing in probability of site use with distance from protected area, although confidence intervals span zero for cheetahs, lions and wild dogs (Table 2). The confi- dence intervals for hyaenas and wild dogs also indicate that the coefficients may be zero but showed relatively even pro- portions in both directions. Elephants, leopards, lions and wild dogs all had the distance to nearest river in their top models, with the probability of site use decreasing with in- creased distance (Table 2), as predicted for these species (Table 1). The predicted values of the probability of site use for each
of the species were mapped to show their possible distribu- tions (Fig. 2). Both elephants (mean probability of site use of 0.553) and hyaenas (0.910) had a wide distribution in the un- protected areas whereas leopards had the most restricted
distribution (0.130). The distribution of wild dogs (0.176) appears to be in two distinct areas, in the south-east and in the north, whereas lions (0.547) and cheetahs (0.598) are present around the boundaries of the protected areas. These species-specific maps were summed to generate a combined map (Fig. 3), which highlights an important wild- life area to the east of the Maasai Mara National Reserve.
Discussion
The main aim of this study was to identify areas outside pro- tected areas with the highest levels of use by cheetahs, ele- phants, spotted hyaenas, leopards, lions and wild dogs, and thus to determine key wildlife areas for future land pro- tection schemes.We also aimed to identify the main covari- ates that influenced site use by each species, to aid in management decisions for these species. Site use varied greatly between species, possibly as a result of their differ- ences in behavioural patterns and resource requirements. Distance from humans was the most informative of the human disturbance covariates for all species, indicating they avoid any human presence irrespective of density. Additionally, the two human disturbance density covariates only accounted for human disturbance within a site whereas the mean human distance also takes into account the sur- rounding sites. This could indicate that these species take into account human disturbance on a wider scale and not just in their vicinity. The avoidance of humans has been shown for multiple large carnivore species (e.g. Schuette et al., 2013, Klaassen & Broekhuis, 2018) and is possibly a re- sult of negative interactions with people (Loveridge et al., 2017). Leopards may be the exception, as they persist in many human-dominated landscapes (Athreya et al., 2013), but this was not reflected in our study, possibly because the high levels of grazing and agriculture reduce the amount of suitable habitat for them in unprotected areas or because they are difficult to detect. For elephants and wild dogs, the proportion of fences negatively influenced presence, al- though the effect was marginal. Previous studies have shown that fences, even though elephants have the ability to break through them, can have a strong negative impact on elephant movement (Thouless & Sakwa, 1995; Loarie et al., 2009). The increase in fencing around the Mara (Løvschal et al., 2017) could therefore prove problematic for elephant movement. In the Mara wild dogs are rarely re- ported within protected areas, but they are found in human- dominated areas. Our findings indicate that the presence of wild dogs is more likely to be influenced by fences than by human presence, possibly because fences restrict their wide- ranging behaviour. Although hyaenas strongly avoided human presence, they still had the widest distribution out- side the protected areas. Other studies have shown that, rather than avoiding areas of high human and livestock
Oryx, 2020, 54(3), 395–404 © 2018 Fauna & Flora International doi:10.1017/S0030605318000297
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