The introduction of African cheetahs to India was planned without considering their spatial ecology

KNP might not result in a long-term reduction of livestock losses or human-cheetah conflicts. If India further follows their plans of establishing a metapopulation in their country by introducing cheetahs to several parks in India (Jhala et al., 2021), we argue that the socio-spatial organization of cheetahs needs to be considered. Our predictive approach has the potential of tackling pro-actively farmer-cheetah conflicts, enhancing our knowledge of cheetahs establishing territories in new areas and to assess the success of potential future trans-continental introductions.

In this letter, we focus on the socio-spatial organization of cheetahs to predict the outcome of the experimental cheetah translocation. The socio-spatial organization is an important aspect of cheetah behavior which has not been considered for the translocation so far. Here, we derive six predictions from empirical data on how we expect the introduced cheetahs to distribute themselves in the KNP. The socio-spatial organization of cheetahs is characterized by (1) adult cheetah males exhibiting two distinct spatial tactics, being either territory holders or "floaters" and by (2) territories being distributed in the landscape with distances of 20 to 23 km between territory centers (Caro, 1994;Melzheimer et al., 2018). The large areas between the territories are not defended by any males but used by females and floaters (Caro, 1994;Melzheimer et al., 2018). In Namibia, territory holders occupy $380 km 2 , while floaters roam in overlapping home ranges of $1600 km 2 encompassing two to four territories and females use home ranges of $650 km 2 between the male territories (Melzheimer et al., 2018(Melzheimer et al., , 2020. In another ecosystem, the Serengeti NP in East Africa, the territories are only $50 km 2 in size (Caro, 1994;Caro & Collins, 1987). This difference might be based on differences in prey availability, humancarnivore conflict, predator competition or habitat composition of the two ecosystems or in the use of different collar types for recording cheetah locations. Remarkably, the centers of the territories in both ecosystems are separated by 20-23 km with large undefended areas between the territories (Caro, 1994;Caro & Collins, 1987;Melzheimer et al., 2020).
We therefore expect that the spatial tactic of cheetah males, the distribution of male territories in the landscape, and the distance between the territories is also observed in the KNP in India. The eight cheetahs from Namibia brought to the KNP comprised a solitary male, two brothers, and five unrelated females. Thus, we predict that the solitary male and the two brothers will establish a territory each which will be separated by 20-23 km (Prediction 1). We further predict that irrespective of the territory size, these three males will occupy the entire KNP which is $17 km Â 44 km in size (Prediction 2), thus not leaving space for additional territories for males introduced from South Africa.
The process of establishing territories in a new area is currently unknown, but translocated cheetahs in Namibia have exhibited extensive exploration movements of several thousand km 2 during the first 6 months (Weise et al., 2015). We therefore predict that the eight cheetahs will conduct extensive excursions outside the KNP during their exploration phase (Prediction 3), potentially coming into conflict with livestock farmers. We further predict that it will take the males many months to adjust the distance between themselves and settle, and the females to settle between the males (Prediction 4).
Territories are valuable because their core areas function as "communication hubs" where territorial males, floaters, and females exchange olfactory information on their presence and reproductive status (Caro, 1994;Melzheimer et al., 2020). Thus we predict that additional males brought in or born in KNP will settle at a distance of $20-23 km away from the first two established territories, coming into conflict with livestock farmers (Prediction 5). The distribution of additional territories would give the five females and additional females new opportunities to establish their home ranges between the territories. Thus we predict that females will also move out of KNP and coming also into conflict with livestock farmers (Prediction 6).
In the communication hubs, the cheetah density is much higher than between the hubs because cheetahs move in and out to exchange olfactory information (Melzheimer et al., 2020). This has consequences on livestock predation. In Namibia, cattle calf losses are highest in the communication hubs (Melzheimer et al., 2020). Shifting breeding herds away from the hubs reduces calf losses by 86% (Melzheimer et al., 2020). When territorial cheetah males are removed, floaters occupy the vacant territories quickly (Melzheimer et al., 2020). This suggests that translocating cheetahs back into the KNP might not result in a long-term reduction of livestock losses or human-cheetah conflicts.
If India further follows their plans of establishing a metapopulation in their country by introducing cheetahs to several parks in India (Jhala et al., 2021), we argue that the socio-spatial organization of cheetahs needs to be considered. Our predictive approach has the potential of tackling pro-actively farmer-cheetah conflicts, enhancing our knowledge of cheetahs establishing territories in new areas and to assess the success of potential future transcontinental introductions.

ACKNOWLEDGMENTS
We thank the Messerli Foundation Switzerland for their long-term funding support of our Cheetah Research Project in Namibia, the Namibian Ministry of Environment, Forestry and Tourism for permission to conduct the study, the farmers in Namibia for cooperation and all team members for data collection and analyses. Open Access funding enabled and organized by Projekt DEAL. The publication of this article was funded by the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation)project number 491292795.