Assessing cheetah reintroduction success 511


TABLE 3 Mean demographic parameters for cheetahs in Liwonde National Park and various South African (SA) reserves in the Cheetah Metapopulation Project (Bissett & Bernard, 2011), Pilanesberg National Park (Power et al., 2019) and Mun-Ya-Wana Conservancy (Gigliotti et al., 2020b). Sample sizes are the number of times each event was recorded (i.e. number of litters or number of females). For interbirth interval, the first sample size is for the number of intervals, and the second is the number of females.


Variable


Age at first reproduction (months)


Interbirth interval (months)


Litter size (number of cubs at emergence)


Litter size (number of cubs at independence)1


Age of independence of cubs (months)


Liwonde National Park


27.7 (n = 3)


SA Reserves (with lions; n = 4)


26.9 (n = 8)


17.7 (n = 1;1) 18.6 (n = 7;3) 4.0 (n = 4)


2.0 (n = 3) 15.6 (n = 2)


% cub survival to independence 60 (n = 3)


3.8 (n = 26) 2.9 (n = 17) 18.8 (n = 8)


SA Reserves (no lions; n = 3)


28.2 (n = 5) 17.0 (n = 5;4)


4.6 (n = 11) 4.7 (n = 7) 14.9 (n = 4)


Pilanesberg National Park, SA


Mun-Ya-Wana Conservancy, SA


28.1 (n = 14)


17.0–18.0 (n = 1;3) 19.4 (n = 19;1) 4.0 (n = 3)


3.3 (n = 61)


2.0 (n = 3) 16.2 (n = 2) 50 (n = 3)


16.7 (n = 14) 42 (n = 61)


1Some litters were still dependent on their mothers at the end of these studies, therefore the litter numbers at independence may be fewer than at emergence.


African fenced reserves (84%; Marnewick et al., 2009). Even with the loss of CM3 and CM4 (57%), survival in Liwonde was higher than in Matusadona National Park, Zimbabwe (36%; Purchase & Vhurumuku, 2005). In addition, cub survival in Liwonde (60%) was higher than that recorded in the South African source population (42–50%; Table 3). Notably, we recorded no mortalities caused by lions,


although lions were reintroduced to Liwonde only 107–354 days after the cheetahs. Lions are a major cause of mortality for cheetahs (Buk et al., 2018;Gigliotti et al., 2020b), and it has been recommended that cheetahs intended for release into areas with resident lions have prior knowledge of lions (Hayward et al., 2007b). The fact that 57%(n = 4) of the chee- tahs were sourced fromreserveswith resident lions may have contributed to them successfully adapting their behaviour in the presence of lions. Ongoing analyses of habitat use before and after lion reintroduction should provide further insights on this particular aspect.


Reintroduction assessment and perspectives


Reintroduction success can be viewed on three scales: an in- dividual’s settlement, the establishment of a population and overall population persistence (Armstrong&Seddon, 2007). If post-release movements and mortality are low, individ- uals will settle sooner, allowing the population to establish from a small number of founders (e.g. Hayward et al., 2007a; Briers-Louw et al., 2019). The reintroduction of chee- tahs into Liwonde was considered successful in establishing a breeding population, with five individuals developing home ranges and all females breeding. The fact that founder populations in most large carnivore reintroductions are small (Breitenmoser et al., 2001) emphasizes the importance of analysing post-release monitoring data, which will facili- tate the comparison across multiple case studies and varying factors over time.


Successful establishment is not a guarantee for popu-


lation persistence. Populations monitored by the Cheetah Metapopulation Project that did not persist in the long termwere extirpated, on average, 8.4 years after reintroduction (Buk et al., 2018). Park management must therefore consider long-term populationmonitoring to establish persistence, and address factors that may cause extirpation of this small popu- lation,which is inherently susceptible to stochastic events such as disease outbreaks (Davidson-Phillips et al., 2019). Also, the death of CF3 in an old wire snare set for bushmeat hunting demonstrates that although Liwonde’s law enforcement ran- gers removed .27,000 snares in the 2 years prior to this reintroduction, snaring remains a problem. Given the geographical isolation of Liwonde’s cheetahs,


and the small founder population, inbreeding depression is a significant risk (Frankham, 2010; Naude et al., 2020). Supplementation of Liwonde’s current population with un- related individuals is strongly recommended, to mimic nat- ural immigration (Ferreira & Hofmeyr, 2014) and ensure the long-term genetic diversification of the population (Gustafson et al., 2017). This could include individuals from the more recent reintroduction of unrelated cheetahs to Majete Wildlife Reserve, Malawi, through the develop- ment of a managed metapopulation node in Malawi.


Conclusion


As the conservation value of protected areas in Malawi has been declining over recent decades because of lack of fund- ing, high levels of poaching and anthropogenic encroach- ment, certain protected areas in Malawi are now shifting towards fenced systems. This approach aims to curtail off- take of natural resources and edge effect pressures caused by the activities of surrounding communities, thereby opening opportunities for species reintroductions (Packer et al., 2013). We recommend that future reintroduction


Oryx, 2022, 56(4), 505–513 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321000788


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