Leopard density in southern Mozambique 409
each flank; Karanth, 1995), allowing us to survey a larger area and thus increase the number of captured individuals, with limited resources. Although we believe that this was the best approach in our case, the trade-offs between surveying a lar- ger area and obtaining higher identification rates should be considered on a case-by-case basis. Although males commonly occupy territories overlap-
ping with those of 2–4 females (Bailey, 1993), more males (n = 5) than females (n = 2) were captured during our study (the sex of two individuals could not be determined). Maputla et al. (2013) also recorded a male-bias in capture rates and cited several potential reasons for this, including heterogeneity in behaviour between sexes in the vicinity of the trap, and in tendencies to use specific trap locations, such as roads (Krebs, 1999). The model with the highest support was that in which sex
did not influence the detection (g0) or scale (σ) parameters. However, rather than indicating the absence of widely de- scribed sex-dependent heterogeneity in behaviour and ran- ging patterns (Bailey, 1993; Kittle et al., 2017), we believe that the relatively small sample size did not provide enough data to facilitate the inference and modelling of sex-specific dif- ferences in detectability and ranging patterns.
Implications for conservation policy in Mozambique
Trophy hunting has the potential to benefit the conservation of large carnivores (Lindsey et al., 2007; Loveridge et al., 2007), and it is estimated that each leopard hunted in Mozambique could contribute c. USD 24,000 to the local and national economy (Jorge et al., 2013). However, if hunting is poorly managed or adds to other sources of anthropogenic mortality, it can reduce numbers to such an extent that a population is no longer viable in the long term. Demonstrating that hunting practices, including quo- tas, are biologically sustainable is therefore essential for tro- phy hunting to be an effective tool in the management and conservation of large African carnivores (Swanepoel et al., 2014; Braczkowski et al., 2015). Our results lead us to question the reliability of the es-
timates employed to set quotas for hunting leopards in Mozambique. The study by Martin & de Meulenaer (1988), quoted as the primary justification for a recent in- crease of the trophy export quota in Mozambique (from 60 to 120 individuals per annum; CITES, 2007), states that up to 80% of the country supports leopard densities of 3–10 individuals/100 km2. It also suggests that only 3%of the country’s total land area should have leopard population densities lower than that found in our study. However, our estimate of 2.60 leopards/100 km2 in Xonghile Game Reserve, one of the better protected areas in the country, sug- gests that it is unlikely thatmany areas in Mozambique sup- port the densities cited in the application for a revision of the hunting quota. Although some landscapes will have higher
primary productivity levels than the Reserve, it is likely that high levels of anthropogenic disturbances in large parts of the country (Hatton et al., 2001) would more than counteract this. Thus, although we appreciate that trophy hunting has not taken place in the Reserve for nearly 10 years (LA & KTE, unpubl. data) and we acknowledge the limitations of our study in terms of the number of indivi- duals encountered relative to the overall range and total size of the population, we believe it is unlikely that leopard densities as high as those cited in the application for a quota increase are commonin areas where hunting currently takes place. Wetherefore recommend further assessments of leopard population status and densities across different habitats and land-use types across the country, in both hunting and protected areas. This would be an important step to- wards the development of a sustainable and empirical quota allocation system, similar to that currently being developed for South Africa, which includes hunting re- gulations based on leopards’ age, adaptive management strategies, and dynamic, evidence-based quota systems (Department of Environmental Affairs, 2017). Camera trap- ping surveys are a rapid method for obtaining robust esti- mates of leopard numbers at a moderate cost (Balme et al., 2009) and, if followed by effectivemanagement inter- ventions, could play an important role in the species’ recov- ery and conservation in many post-conflict landscapes across the country. The conservation challengeswe have identified are not ex-
clusive toMozambique, with Tanzania andNamibia also em- ploying the density estimates of Martin & de Meulenaer (1988) for justifying modifications, approved by CITES (CITES, 2002, 2004), of quotas for hunting leopards. Our re- sults reinforce the need for caution when setting hunting quotas for leopards, and the importance of reliable popula- tion estimates across the species’ range.We recommend simi- lar research be carried out in other regions where such estimates are used to set harvest quotas, to support a shift to- wards evidence-based guidance of management and policy.
Acknowledgements We thank the Director ofNationalConservation Areas Mozambique for granting KE and LA research permits (005-2011/003-2012), the shareholders of Xonghile Game Reserve for allowing access, the Centre for Wildlife Management, University of Pretoria, and Wilderness Trust for financial support, and Valeria Boron and Charlotte E. Searle for their help with the analyses.
Author contributions Data collection: LA and KE; analyses: PS; writing: all authors.
Conflicts of interest None.
Ethical standards All research carried out complied with the Oryx Code of Conduct.
Oryx, 2020, 54(3), 405–411 © 2018 Fauna & Flora International doi:10.1017/S0030605318000121
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