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524 M. A. Vinks et al.


TABLE 3 Densities of primary leopard prey in north-central Kafue National Park, and the proportion of the Kafue leopard diet comprised of each species, from Vinks et al. (2020) and Creel et al. (2018).


Preferred prey species Puku Kobus vardonii


Impala Aepyceros melampus Warthog Phacochoerus africanus


Mean individual density (95% CI), per km2


11.54 (10.27–12.82) 6.25 (5.99–6.51) 2.51 (2.36–2.67)


appear to favour leopards in small areas that maintain rela- tively higher densities of their favoured prey (Fig. 2). Nonetheless, average leopard density across the entire Greater Kafue Ecosystem would still be expected to reflect low preferred prey abundance as a whole (Stander et al., 1997). A direct examination of these issues near the Park border and within the Game Management Areas would be of value. Lions can limit the distribution and abundance of leo-


pards (Stander et al., 1997; Balme et al., 2013), although re- cent findings suggest that interference competition with lions may be weaker than previously thought, particularly in secured protected areas with intact prey populations (Rosenblatt et al., 2016; Balme et al., 2017; Miller et al., 2018). The effects of intraguild competitionmight be stronger in prey-depleted ecosystems, particularly when large-bodied prey densities are disproportionally reduced (Balme et al., 2017; Creel et al., 2018). The diets of leopards and lions over- lap extensively within our study area (Creel et al., 2018) be- cause the larger-bodied prey preferred by lions have become rare (Vinks et al., 2020). This niche compression reduces the scope for dietary niche partitioning, and thus could strengthen the limiting effect of competition on subordinate competitors (Creel et al., 2018). However, we did not find evidence for this effect in Kafue. The distribution of leopard detections across space paralleled the core utilization distri- bution (60%isopleth of a kernel utilization distribution) for two resident lion prides in our study area (Fig. 3), suggesting that leopards did not avoid areas heavily used by lions throughout the dry season. However, lion density in Kafue is low (3.43 individuals per 100 km2, 95%CI 2.79–4.23) and pride size is small, with an average of 2.89 adult females per pride (Vinks et al., 2021). Accordingly, the ratio of leopards to lions per 100 km2 in this part of Kafue is 2.3:1 (HMMDM) and 0.97:1 (MMDM), compared to 0.82:1 (HMMDM) and 0.47:1 (MMDM) in a part of Zambia’s South Luangwa National Park with favourable ecological conditions, where leopard and lion densities are both high (Rosenblatt et al., 2014, 2016). Such a substantial reduction in lion density could partially offset the anticipated effects of low prey densities on leopards in this part of the Park and could help explain a lack of evidence for intraguild competition. Survival rates in our core study area were similar to esti-


mates from another Zambian leopard population with better protection (Rosenblatt et al., 2016) and thus do not


Mean herd density (95% CI), per km2


1.31 (1.20–1.42) 0.74 (0.72–0.77) 0.66 (0.63–0.70)


Proportion of leopard diet


0.34 0.20 0.26


appear to be reduced by low prey density. Prior research with lions in this system also found that survival for resident individuals was high, but that lion density and cub recruit- mentwere both low (Vinks et al., 2021). Thus measurements of adult survival may not be a good tool to evaluate the effects of prey depletion on large carnivore populations. Our results have several important limitations. We were


unable to monitor leopard recruitment effectively, and re- cruitment could be affected by low prey density (Balme et al., 2013).Our survival estimates were not partitioned across age classes (because we could not reliably age leopards using camera-trap photographs), which could have obscured vari- ation in survival among age–sex classes.We note that survival rates for leopards outside our core study area, where human pressures are stronger (Watson et al., 2014;Overton et al., 2017), could be lower given lower overall leopard survival in human dominated landscapes (Swanepoel et al., 2015a,b). Wire-snare poaching could also be a source of mortality for leopards (Becker et al., 2013; Rosenblatt et al., 2014; Schuette et al., 2018), although we did not detect any snared leopards on our study site during the course of the study and snared leopards are rarely detected in Kafue, in contrast to snared wild dogs, spotted hyenas, and lions (Zambian Carnivore Programme, unpubl. data). Limited observations of snared leopards could be due in part to individuals succumbing to snares without ever breaking loose (Loveridge et al., 2020), as well as their cryptic nature. Another limitation of our study was the inability to esti-


mate abundance in multiple years, because of insufficient recapture rates. As a consequence, we were unable to evalu- ate potential annual trends in abundance, limiting our infer- ences about density to a single year. With that caveat, we did not observe substantial fluctuations in the number of leo- pards detected throughout the course of the study, and known individuals were consistently detected across mul- tiple years (Table 1). Sample sizes were small, which may lead to unreliable abundance estimates (White et al., 1982), although precision was comparable to other published stud- ies (Henschel et al., 2011; Stein et al., 2011; Maputla et al., 2013; Swanepoel et al., 2015a,b). Nonetheless, given that estimates of population size for leopards in miombo wood- land are limited (Balme et al., 2007; Rosenblatt et al., 2016), especially where prey densities are disproportionately re- duced, our density estimates are of high value from a con- servation and management perspective.


Oryx, 2022, 56(4), 518–527 © The Author(s), 2021. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321000223


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