About 13 percent of bird species, including most seabirds, nest in colonies. Colonial nesting evolves in response to a combination of two environmental conditions: (1) a shortage of nesting sites that are safe from predators and (2) abundant or unpredictable food that is distant from safe nest sites. First and foremost, individual birds are safer in colonies that are inaccessible to predators, as on small rocky islands. In addition, colonial birds detect predators more quickly than do small groups or pairs and can drive the predators from the vicinity of the nesting area. Because nests at the edges of breeding colonies are more vulnerable to predators than those in the centers, the preference for advantageous central sites promotes dense centralized packing of nests.
The yellow-rumped cacique, which nests in colonies in Amazonian Peru, demonstrates how colonial birds prevent predation. These tropical blackbirds defend their closed, pouchlike nests against predators in three ways. First, by nesting on islands and near wasp nests, caciques are safe from arboreal mammals such primates. Second, caciques mob predators (work together as a group to attack predators). The effectiveness of mobbing increases with group size, which increases with colony size. Third, caciques hide their nests from predators by mixing active nests with abandoned nests. Overall, nests in cluster on islands and near wasp nests suffer the least predation.
Coordinated social interactions tend to be weak when a colony is first forming, but true colonies provide extra benefits. Synchronized nesting, for example, produces a sudden abundance of eggs and chicks that exceeds the daily needs of local predators. Additionally, colonial neighbors can improve their foraging by watching others. This behavior is especially valuable when the off-site food supplies are restricted or variable in location, as are swarms of aerial insects harvested by swallows. The colonies of American cliff swallows, for example, serve as information centers from which unsuccessful individual birds follow successful neighbors to good feeding sites. Cliff swallows that are unable to find food return to their colony, locate a neighbor that has been successful, and then follow that neighbor to its food source. All birds in the colony are equally likely to follow or to be followed and thus contribute to the sharing of information that helps to ensure their reproductive success. As a result of their enhanced foraging efficiency, parent swallows in large colonies return with food for their nestlings more often and bring more food each trip than do parents in small colonies.
To support large congregations of birds, suitable colony sites must be near rich, clumped food supplies. Colonies of pinyon jays and red crossbills settle near seed-rich conifer forests, and wattled starlings nest in large colonies near locust outbreaks. The huge colonies of guanay cormorants and other seabirds that nest on the coast of Peru depend on the productive cold waters of the Humboldt Current. The combination of abundant food in the Humboldt Current and the vastness of oceanic habitat can support enormous populations of seabirds, which concentrate at the few available nesting locations. The populations crash when their food supplies decline during El Nino years.
Among the costs, colonial nesting leads to increased competition for nest sites and mates, the stealing of nest materials, and increased physical interference among other effects. In spite of food abundance, large colonies sometimes exhaust their local food supplies and abandon their nests. Large groups also attract predators, especially raptors, and facilitate the spread of parasites and diseases. The globular mud nests in large colonies of the American cliff swallow, for example, are more likely to be infested by fleas or other bloodsucking parasites than are nests in small colonies. Experiments in which some burrows were fumigated to kill the parasites showed that these parasites lowered survivorship by as much as 50 percent in large colonies but not significantly in small ones. The swallows inspect and then select parasite-free nests in large colonies, they tend to build new nests rather than use old, infested ones. On balance, the advantages of colonial nesting clearly outweigh the disadvantages, given the many times at which colonial nesting has evolved independently among different groups of birds. Still lacking, however, is a general framework for testing different hypothesis for the evolution of coloniality.
5.According to paragraph 3, cliff swallows closely observe other members of their colony in order to