Divergent seed dispersal outcomes: Interactions between seed, disperser, and forest traits.

Animals disperse seeds in various ways that affect seed deposition sites and seed survival, ultimately shaping plant species distribution, community composition, and ecosystem structure. Some animal species can disperse seeds through multiple pathways (e.g., defecation, regurgitation, epizoochory), each likely producing distinct seed dispersal outcomes. We studied how seed traits (size and toughness) interact with disperser species to influence seed dispersal pathway and how this ultimately shapes the proportion of seeds deposited in various habitat types. We focused on three frugivorous species of duikers (African forest antelopes) in the Dja Faunal Reserve, a tropical rainforest in southern Cameroon. Duikers can both defecate and regurgitate seeds, the latter predominantly occurring during rumination at their bedding sites (or "nests"). We located duiker nests and dungs along 18 linear 1-km-transects to assess: (1) how seed traits affect the likelihood of dispersal via defecation versus regurgitation, (2) if defecated versus regurgitated seeds are deposited at different rates in different forest types (assessed by indigenous Baka), microhabitats, and forest structural attributes (measured by drone lidar), and (3) if these differ between three duiker species that vary in size and diel activity patterns. We found that duikers predominantly defecated small seeds (<3 mm length) and regurgitated larger and tougher seeds (>10 mm length), the latter including 25 different plant species. The three duiker species varied in their nesting habits, with nocturnal bay duikers (Cephalophus dorsalis) nesting in dense understory vegetation at proportions 3-4 times higher than Peter's and yellow-backed duikers (Cephalophus callipygus and Cephalophus silvicultor). As a result, bay duikers deposited larger regurgitated seeds at a higher rate in habitats with denser understory where lianas and palms predominate and near fallen trees. This directed regurgitation seed deposition likely plays an important and unique role in forest succession and structure. This study highlights the importance of ungulate seed dispersal by regurgitation, a vastly understudied process that could impact many ecosystems given the prevalence of ruminating ungulates worldwide.