Leopard density 523


TABLE 1 Leopard Panthera pardus camera-trap detections during 2013–2019.We considered leopards new in this study if they had not been previously documented on the study site in 2013–2019 (through either opportunistic observations or camera traps).


Year 2013


2014 2015 2016 2017 2018 2019


Total detections 21


15 13 19 11 10 12


Individuals detected 10


10 8


12 7 9 7


of prey depletion on leopards could be partially offset by a decrease in the limiting effect of competition, if any exists (Rosenblattetal., 2016; Balme et al., 2017; Miller et al., 2018).


This study provided baseline estimates of sex-specific


annual survival rates and population density for leopards in north-central Kafue, and is one of few studies to provide such results for leopards in miombo woodland, which con- stitutes a large portion of the species’ range across Zambia and Tanzania (Balme et al., 2007; Rosenblatt et al., 2016). Contrary to our predictions, density estimates were compar- able to those in ecosystems with relatively greater protec- tion, where ecological conditions (notably preferred prey abundance) remain favourable for leopards (Ray, 2011; Du Preez et al., 2014; Swanepoel et al., 2015a,b; Rosenblatt et al., 2016, 2019). Although leopard density in our study site was not precariously low, this site is located in the heart of northern Kafue, with relatively high densities of preferred leopard prey, along a major perennial river, and mostly removed from negative anthropogenic edge effects typical outside the core of the Park. Densities of preferred leopard prey are substantially reduced in other parts of the Park and adjacent Game Management Areas (Creel et al., 2018; Schuette et al., 2018; Vinks et al., 2020), and there is evidence of wire snare poaching bycatch in addition to targeted poaching in areas of the Greater Kafue Ecosystem with


TABLE 2 The best-supported Cormack–Jolly–Seber models of leo- pard survival, as determined by Akaike’s information criteria cor- rected for small sample size (AICc).


Model1 w(.) P(.)


w(.) P(*sex) w(*sex) P(.) w(.) P(.) π(.)


K2 ΔAICc3 2 0


3 1.834 3 1.991 3 2.214


w(.) P(*study year) 8 2.716 w(*sex) P(*sex) w(.) P(*sex) π(.)


4 3.547 4 4.127


Weight Deviance 0.307


0.123 0.114 0.102 0.079 0.052 0.039


145.735 145.355 145.513 145.735 133.889 144.775 145.355


1w, apparent survival rate; P, detection probability; π, mixture parameter. 2K, number of parameters. 3ΔAICc, difference in AICc to the best performing model.


Female 7


4 4 9 2 3 5


Male 3


4 4 3 4 4 0


Unknown sex 0


2 0 0 1 2 2


New in this study 10


6 2 3 1 5 2


less protection (Overton et al., 2017). Thus, extrapolating our density estimate across other portions of the Park or Game Management Areas, where ecological conditions are less favourable and human pressures are known to be stron- ger (Watson et al., 2014; Overton et al., 2017; Schuette et al., 2018), would not be valid (Trouwborst et al., 2019). We did not fit spatial capture–recapture models, which


are widely used in conjunction with leopard camera-trap data (Du Preez et al., 2014; Goldberg et al., 2015; Balme et al., 2019; Devens et al., 2019).When the number of rede- tections is low, the data do not allow good estimates of spa- tial capture–recapture model parameters, yielding a biased and imprecise density estimate (Efford et al., 2004;Du Preez et al., 2014; Paterson et al., 2019). Instead, we used a randomized grid with a spacing tailored to leopard move- ments to produce a representative estimate of density for the sampled area. As with other large carnivores, leopard density is gener-


ally expected to be positively correlated with preferred prey biomass at a system-wide scale (Stander et al., 1997;Marker & Dickman, 2005). This logic leads to an expectation that Kafue leopard density should be low (Henschel et al., 2011; Balme et al., 2013; Rosenblatt et al., 2016), but it was not. The relatively higher localized densities of preferred leopard prey (Table 3) remaining within our 228 km2 camera-trap area, compared to the rest of the Park, could explain the relatively high observed leopard density. This explanation is sup- ported by the observation that we obtained more leopard detections at sites with higher localized densities of pre- ferred prey (Fig. 2). These results also suggest that leopards may maintain higher densities where preferred prey re- mains abundant at small spatial scales within larger areas with lower prey density overall, such as Kafue. Leopards may kill smaller prey in other parts of the ecosystem, in- cluding the common duiker Sylvicapra grimmia, which re- main relatively abundant throughout Kafue (Schuette et al., 2018; Vinks et al., 2020). However, dependency on such small prey is also expected to reduce population density (Henschel et al., 2011). Although the Greater Kafue Ecosystem as a whole is affected by prey depletion, pockets of higher relative prey abundance could affect large carni- vore dynamics across the Greater Kafue Ecosystem, and


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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