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Assessing cheetah reintroduction success 507


twice per week, locating cheetahs through VHF track- ing (receiver: Communication Specialists, Orange, USA; H-Type antenna: Telonics, Mesa, USA). We supplemented these observations with opportunistic sightings. We also in- spected dens within the first 2 weeks of denning to record location, litter size and cub survival. To minimize distur- bance, dens were checked by a single person while the female was hunting, and we did not handle the cubs (Laurenson & Caro, 1994). To replace GPS collars before battery depletion, animals were immobilized by chemical capture and then re-fitted with VHF collars (Sirtrack, Hawkes Bay, New Zealand) modified with a long-range geolocation trans- mitter (LoRa; Smart Parks, Rotterdam, The Netherlands). Re-fitted collars (359 g) facilitated continued checks of physical condition and dens.


Data analysis


We analysed the first year (range 297–365 days) of post- release movements for five cheetahs. Two cheetahs (CM3 and CM4) were omitted from the analysis because of in- sufficient data from the VHF collars. To ensure consistent sampling across individuals, we selected the GPS fix closest to 12.00 for analysis. Although two mixed-sex groupings were released from the boma, the females separated from their groupings 2 and 19 days post-release, respectively; we therefore analysed each GPS-collared animal as a singleton adult or female with dependent cubs. We conducted all analyses in R 3.5.1 (R Core Team, 2018). We also compared demographic parameters to those documented within source populations of the Cheetah Metapopulation Project.


Post-release exploration We investigated initial post- release exploration by calculating daily displacement for each individual as the Euclidean distance between two suc- cessive locations in a 24-hour cycle using the adehabitatLT package in R (Calenge, 2006). We developed a linear mixed model using the lme4 package in R (Bates et al., 2015), to as- sess drivers of post-release exploration. Variables included in the model were sex, age at translocation, duration of the pre-release holding period in Liwonde, and days since release. We included individual identity as a random intercept.


Settlement For the first year post-release, we calculated progressive home ranges for each of the five cheetahs, to determine the duration of exploratory behaviour and time of settlement (Weise et al., 2015a). We defined progressive home ranges as the minimum convex polygon around GPS fixes over a moving window shifting continuously by 1 day. Because of collar failure, we used a moving window of 11 days for the calculation of aminimum convex polygon


based on aminimum of five GPS fixes. We used 100%min- imum convex polygons as they include potential exploratory forays of interest. We also calculated the net squared dis- placement between each location and the first location col- lected post-release to determine site fidelity and settlement (Börger & Fryxell, 2012).We visually assessed the progressive minimum convex polygon and net squared displacement curves to identify when patterns of space use stabilized, indi- cative of individuals having settled, similar to natal dispersers (Fattebert et al., 2015;Weise et al., 2015a,b).


Results


Post-release exploration Males moved longer daily distances (3.6 ± SD 3.3 km, n = 2) than females (1.2 ± SD 1.2 km, n = 3; βmale = 2.5, 95% CI = 1.99–3.14,P,0.001; Fig. 2). Older individuals moved


shorter daily distances than younger ones (βage =−0.008, 95%CI=−0.015–0.002,P= 0.01), and cheetahsmoved short- er distances as more time elapsed post-release (βtime =−0.005, 95%CI =−0.006–0.004,P,0.001). Duration of pre-release holding periods did not significantly affect daily movements post-release (Table 2).


Settlement Reintroduced GPS-collared cheetahs (n = 5) developed home ranges and settled on average 103 ± SD 68 days post- release (range 29–174 days). The three females developed home ranges within 3 months post-release (55 ± SD 23 days; 29–73 days; Figs 3 & 4). Before the estimated start of the denning period, both CF1 and CF2 demonstrated a period of early settlement prior to the final settlement phase between 23–44 and 34–55 days post-release, respect- ively. Males settled 174 days post-release (Figs 3 & 4), after initial extensive exploratory movements (Fig. 2).


Survival and reproduction


Four of the five GPS-collared cheetahs survived their first year post-release. CF3 was killed in a wire snare 307 days post-release, and CM4 and CM3 were recorded missing 85 and 152 days post-release, respectively. All three females gave birth to their first litter after home range development 131 ± SD 7 days post-release (range 123–138 days). Four birth- ing events were recorded over the 2-year monitoring period. Dens were located within 2.2 ± SD 0.5 km (n = 4) of the boma. Denning lasted 55 ± SD 1 days (53–56 days; n = 4 lit- ters). Mean litter size was 4 ± SD 1.4 cubs (range 3–6;n = 4), which was consistent with mean litter sizes recorded in source populations (4.3–3.3 cubs; Table 3). All litters reached the age of emergence from the den, but cub mortality was


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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