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Savannah and forest elephants in Uganda 189


FIG. 1 The Democratic Republic of the Congo–Uganda border, and the Sebitoli area in Kibale National Park (Uganda), with locations of the camera traps.


et al., 2002;Mondol et al., 2015; Kim &Wasser, 2019); along the border of the DRC and Uganda; the northern Central African Republic; the Gourma region in Mali; and along the Pendjari–Arli complex of West Africa on the Benin– Burkina Faso border (Mondol et al., 2015; Kim & Wasser, 2019). To date, no study combining genetics and morphology


has been able to shed light on the phenotype of hybrid ele- phants. Yet the study of phenotypes (the set of observable characteristics of all or part of an organism) is central to many disciplines in biology (Houle et al., 2010). In addition, we have no information about whether hybrids could be phenotypically intermediate between their parents, closer to one parent or outside the range of variation of both par- ents’ species. As hybrids are fertile (Mondol et al., 2015), we can assume they have a wide range of phenotypes depending on their hybridization rate and whether their mates are hy- brids or pure forest or savannah elephants (Roca et al., 2005; Mondol et al., 2015). Our study aims to confirmthat the two species of African elephant can be discriminated using a set of six proposed morphological criteria, and that in a hybrid- ization zone both species can be found as well as individuals differing from both the savannah and the forest phenotype, which could emerge from the reproduction of the two species. This study took place in the hybridization zone at the


DRC–Uganda border in the forested Sebitoli area in the north of Kibale National Park, Uganda. This zone is located 60 km north of the nearest area of savannah in Queen Elizabeth National Park and is linked to it by a corridor that facilitates the movement of elephants between the two Parks (Fig. 1). The presence of the two species and the hybrids has already been documented in the south of Kibale National Park. However, neither the proportions of each species nor the history or dynamics of the populations are


known (Mondol et al., 2015). Local farmers and villagers in the Sebitoli area have also reported the presence of two types of elephants distinguishable by their size, colour and behaviour (Capelot, 2013; SK, pers. comm., 2012; JB, pers. comm., 2019). As direct observations and access to biolog- ical samples are challenging because of the dense under- growth, swamps and mountains of the Sebitoli terrain, all of which add to the danger of closely approaching elephants, which are likely to charge if threatened (Omeja et al., 2016), we used camera traps and developed an assignment key for the elephant species to document phenotypes in this area.


Study area


The 795 km2 Kibale National Park in south-west Uganda is a mosaic of mature forest, successional forest (formerly used for agriculture), grassland, woodland, lakes and wetlands (Chapman & Lambert, 2000). The movement of elephants between Kibale National Park (forested area) and DRC can only take place in the south via the connection between Queen Elizabeth National Park (savannah area) and Virunga National Park (mixed forest–savannah area) be- cause of the high altitude of the Rwenzori Mountains be- tween the two countries. Migrations between the Virunga and Queen Elizabeth National Parks have been observed since the 1960s, mainly from the DRC to Uganda (Keigwin et al., 2016). In the extreme north of Kibale National Park, the Sebitoli research area, monitored by the Sebitoli Chimpanzee Project, covers c. 25 km2 of forest logged in the 1970s. Today, this area comprises regenerating forest (70%), with only 14% being old-growth forest, surrounded by farm- lands and tea and eucalyptus plantations (Bortolamiol et al., 2014).


Oryx, 2023, 57(2), 188–195 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321001605


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