Corridors of Tolerance 849 Following dispersal out of protected areas, these individ-
ual lions resided on unprotected community lands amongst high densities of humans and livestock. The lions Ndelie, Selenkay and their offspring were responsible for a min- imum of 146 depredation events, totalling losses of at least 216 head of livestock. We documented these individuals being hunted by Maasai warriors a minimum of 53 times. Although a total of 267 hunts were recorded within the study area during 2010–2018, ,4% resulted in a lion be- ing killed. More than 95% of these hunts were halted, pri- marily by non-governmental conservation organizations and theKenya Wildlife Service(Hazzah et al., 2014). Inside the study area, before conservation interventions (2001–2002), there were approximately 20–30 lions killed each year on the Group Ranches (Hazzah et al., 2014). Once conservation initiatives facilitated tolerance towards lions, the number reduced to 0–2 lions per year (Hazzah et al., 2014). Outside the study area, a minimum of 38 (20%) of known dispersers were killed (poisoned, snared or speared) whilst dispersing. Osapuku has spent his entire life outside protected
areas. We postulate that because of living in a landscape where humans, not lions, are the apex predator, Osapuku and the other lions learned how to move and subsist near people, allowing him to traverse a densely populated area before arriving at an area of refuge within the Kapiti region (Mogensen et al., 2011; Valeix et al., 2012; Ordiz et al., 2013). Although Maasai pastoralists of the Amboseli region
had decimated the lion population by the early years of the 21st century (Chardonnet, 2002), current tolerance of lions by the human communities, presumably because of conser- vation initiatives (Hazzah et al., 2014), has seemingly allowed these lions to survive to adulthood, breed and successfully disperse (Packer et al., 1991; Björklund, 2003; Trinkel et al., 2008; Dolrenry et al., 2016). As shown in previous studies within the ecosystem (Okello, 2009; Hazzah et al., 2014; Dolrenry et al., 2016), with high levels of local participation in conservation and a greater sense of ownership of their environment, tolerance for lions increases. To our knowledge, this is the first time that links have
been observed between the lion populations of Tsavo West, Amboseli and Nairobi National Parks. We believe the in- creased and sustained human tolerance over several lion gen- erations, in addition to continued availability of habitat and prey, have contributed to the survival of dispersers into and out of the study population, which, as shown by other studies (Andrewartha, 1954; den Boer, 1968, 1981, 1990; Hansson, 1991; Fahrig & Merriam, 1994;Sweanor et al., 2000), contri- butes to increased viability of the lion metapopulation. In addition to habitat preservation, promoting connectivity by increasing the tolerance of human communities for, and acceptance of, large carnivores and concurrently allowing the carnivores to learn how to coexist in human- and live- stock-dominated landscapes, facilitates a more connected metapopulation (Carpenter et al., 2000; Crooks & Sanjayan,
2006;Groom&Harris, 2008;Maclennan et al., 2009;Hazzah et al., 2013, 2014; Dolrenry et al., 2014; Blackburn et al., 2016). The future of African lions lies in the hands of the human communities (Adams&McShane, 1996;Western&Wright, 2013) and the stories of the successful dispersal of these lions provide an example of how human tolerance can engender connectivity. In conclusion, we suggest a broadening of the definition of corridors, particularly for the large carnivore species that are a challenge to human–wildlife coexistence. Additionally, we urge conservationists to establish compre- hensive databases that promote consistent data structure for shared and verifiable research. Such databases should capture the necessary individual information and facilitate engagement in broad-scale collaborations that create op- portunities for the exchange of knowledge and best prac- tices, particularly in identifying dispersing animals so as to improve the understanding of connections between sites (Dolrenry et al., 2014). Nevertheless, without forbearing communities exemplifying the necessary tolerance to allow lions to move through their space, the functional meta- population model of lions in East Africa could be lost.
Acknowledgements We thank the Kenya Wildlife Service and the Government of Kenya for permissions and support, the group ranch committees for allowing access to their lands, the Maasai communities for being welcoming and supportive, the Lion Guardians for their dedi- cation to lions and this research, M. Mbithi for sharing his observations and knowledge, P. Briggs, L. Nkiinti, L. Mamaai and E. Kesoi for their dedication, S. Chandra and N. R. de Souza for editing and map- making, respectively, and two anonymous reviewers and the Editor for their suggestions. This study was funded by National Science Foundation Fellowship Grant No. 2008058924, Woodtiger Fund, Panthera, Denver Zoo, Philadelphia Zoo, Flora Family Foundation, S. Gold, National Geographic, and Rufford Small Grants.
Author contributions Study conception and design: all authors; analysis: SD, LH; data collection, analysis and writing: SD.
Conflicts of interest None.
Ethical standards This research abided by the Oryx guidelines on ethical standards, and was conducted under Permit No. MOEST 13/ C689, Animal Use Protocols R191 University of California, Berkeley, and L400 University of Wisconsin, Madison, USA.
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Oryx, 2020, 54(6), 847–850 © 2020 Fauna & Flora International doi:10.1017/S0030605319000656
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