To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.
Early successional tree species typically attract numerous generalist frugivores, many of which serve as seed dispersers. This study evaluated whether several such dispersers facilitated the invasion of an exotic, early successional tree species, Maesopsis eminii (Rhamnaceae), in submontane forests of the East Usambara Mountains, Tanzania. The primary seed dispersers and their dispersal effectiveness were determined through focal watches at eight trees. Only 3 of 13 frugivores that consumed Maesopsis fruits functioned as seed dispersers (silvery-cheeked hornbill Ceratogymna brevis, Fischer's turaco Tauraco fischeri and blue monkey Cercopithecus mitis). On average, hornbills dispersed > 26 times more seeds than monkeys, and > 3 times as many seeds as turacos per visit and were thus considered the most important disperser. Seed germination was not affected by gut passage through hornbills. Germination in four microhabitats was experimentally evaluated to determine whether microhabitat influences germination success. Large gaps and forest edges had significantly higher germination rates compared with understorey locations; the germination rate in small gaps was not significantly different from that in the understorey. Expansive Maesopsis invasion in the East Usambaras was likely enhanced in both rapidity and scale by the presence of an extremely effective dispersal agent, the silvery-cheeked hornbill.
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