Nesting distributions of Galápagos boobies (Aves: Sulidae): an apparent case of amensalism.

Blue-footed boobies (Sula nebouxii) in the Galápagos Islands nest at coastal sites such as cliff edges if Nazca boobies (S. granti) are absent. However, if sympatric with nesting Nazca boobies, they nest nearby, but farther inland, in areas with little topographical relief. Nazca boobies nest at the coastal sites whether blue-footed boobies are present or not. The segregated nesting pattern of these two species offers a model system to investigate factors influencing community structure. We tested a non-interactive hypothesis, in which different fundamental niches generate the non-overlapping distributions, and an interactive hypothesis, in which the two fundamental niches overlap and an interaction between the two species causes the segregation. Data on three factors considered as likely parameters differentiating fundamental niches (nest microclimate, nature of the nesting substrate, and ease of taking flight from nest sites) failed to support the non-interactive hypothesis. These results suggest that the two species have indistinguishable fundamental niches with respect to these parameters, but different realized niches. Researchers studying resource partitioning by ecologically similar species often only consider competition (a "-/-" interaction) to explain situations like this, ignoring the more parsimonious amensal (0/-) possibility. Nesting segregation in this situation is apparently caused by attacks of non-breeding adult Nazca boobies on blue-footed booby nestlings, injuring nestlings and ultimately preventing them from fledging. The interaction does not result in any discernible costs or benefits (i.e., effects on fecundity or survival) for the adult Nazca boobies, so it is best described as an amensal interaction. This interaction provides a sufficient explanation of the observed nesting segregation, and precludes present competition for nesting space.