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500 E. H. Walker et al.


release, and one male (#NAAJU1616 ofCM4) was killed by a leopard Panthera pardus before the coalition had achieved independence. Considering only cheetahs that underwent final release, independence was achieved by 96% of released individuals. Each individual or group achieving independ- ence was supplemental fed on average 17 times (median 7 feedings, range 1–112 feedings) and made its first known kill after 18 days (median 7 days, range 3–113 days). FC1 struggled post-release, probably because of her dependent cubs, and as a result was fed more frequently (112 feedings) to ensure that the cubs were properly nourished. Most indi- viduals or groups (25 cheetahs) achieved independence within 2 weeks after release and only required 1–3 supple- mental feedings before making their first kills.


Survival The eight individuals that failed their training–re- leasetrialswerenot included in thesurvivalanalyses, reducing the total number of individuals used for analysis to 28 (11 M, 17 F).Of these, 68%(6 M, 13 F)were recorded dead at the time of analysis,and thefateof 32%(5 M, 4 F) was unknown because of collar failure or depleted batteries. The annual Kaplan– Meier survival estimate for all released individuals was 0.61 (95%CI 0.45–0.84; Fig. 2, Table 4). Social grouping did not significantly affect survival, but males in coalitions tended to live longer compared to females in coalitions and single fe- males. The six deaths recorded on farmland were attributed to both anthropogenic (34%) and natural causes (64%). Of the four cheetahs that died of natural causes, two were from hunting injuries and two were killed by leopards. For the 13 recorded deaths in a reserve, 69% were attributed to natural causes and 31% to unknown causes. Of the nine cheetahs that died of natural causes, two died of old age, two of hunting injuries, one was killed by baboons Papio ursinus and four by competing predators (two leopard, one lion Panthera leo,one spotted hyaena Crocuta crocuta). No differences in survival were found for the variables age when orphaned, time spent in captivity, and training–release (Fig. 2, Table 4). No deaths occurred as a result of starvation, suggesting that cheetahs, with strategic support provided by post-release monitoring, can acquire the skills required to support themselves despite their history.Overall survival estimates at 3, 6, 9 and 12 months remained stable, without any increasedmortality in the first 3 months post-release (Fig. 3). Survival in the first and second years did not differ significantly (χ2 = 0,df = 1,P = 0.8). The increase in survivorship after 15 months is a result of five in- dividuals (18%) surviving beyond 2 years-post release.


Reproductive success Successful reproduction is the ulti- mate measure of the success of release efforts. Although our data is insufficient for statistical analyses, the following observations weremade: SF5,SF6,SF10 and SF11 mated with and were impregnated by wild males. SF5 raised two female cubs to independence, SF10 gave birth to a litter of cubs


before her collar failed. During the necropsy on SF6, five un- born cubs were found. SF11 gave birth to two litters, lost the first but raised the second to 11 months of age before being killed by a leopard. CM4 were seen on several occasions with female cheetahs, and were observed mating with a wild female.


Discussion


Following our release protocol, the success rate of animals achieving independencewas high. Furthermore, annual post- release survival ratewas similar to other cheetahs translocated into a free-ranging reserve (NamibRand) in Namibia (an- nual Kaplan–Meier survival estimate 0.57, 95%CI 0.35–0.76; Weise et al., 2015) and mortality was comparable to cheetahs in a fenced reserve in South Africa (Bissett&Bernard, 2011). Independence was achieved quickly for most individuals (,2 weeks) and we had evidence of successful reproduction events for some released individuals. Neither age when orphaned nor time in captivity affected


the survival probability of released cheetahs. Therefore, our protocol for pre-release management to achieve release candidate status for individuals orphaned younger than 6 months through bonding them to candidates orphaned at an older age seems effective. These findings are contrary to those of previous studies in which time in captivity had significant negative influence over survival, as they show success in releasing captive-raised individuals (Jule et al., 2008;Weise et al., 2015; Boast et al., 2018). Although the training–releases did not significantly im-


prove survival, one advantage they may offer is the oppor- tunity to screen candidates that meet requirements but that would benefit from additional assessment to confirm their ability to reach independence (Gusset et al., 2006; Houser et al., 2011). However, we suggest caution when employing such a technique to provide pre-release hunting training to large carnivores. As natural predator–prey interactions are difficult to recreate, the behaviour and strategy developed within a training environment may not be adequate for a final release.With intensive post-release monitoring accord- ing to our protocol, an animal can be reliably supported with supplemental feeding/water and other interventions (see Methods) as necessary until it has reached independence. Thus,we suggest that intensive post-releasemonitoringwith- in the release site provides the animal with the safest and most stable conditions for acquiring the necessary hunting and survival skills relevant to that release site. Past studies of carnivore releases have demonstrated the


challenges associated with homing behaviour (Linnell et al., 1997; Yiu et al., 2015), which has also been observed in captive-raised and released animals of other species (Henshaw & Stephenson, 1974; Ridgway & Robison, 1985). Although we could not test for differences between soft and hard release, we recommend implementing soft releases


Oryx, 2022, 56(4), 495–504 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321000235


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