Why come back home? Breeding-site fidelity varies with group size and parasite load in a colonial bird.

Fidelity to a past breeding site is widespread among animals and may confer both costs and benefits. Colonial species occur at specific sites that can accommodate multiple breeders, and the choice of whether to return to last year's site or disperse elsewhere can affect colony site use, the colony size distribution and individual fitness. For the colonial cliff swallow, Petrochelidon pyrrhonota, which occupies colonies of widely different sizes, we used a 30-year field study in western Nebraska to investigate how the extent of infestation by ectoparasites and colony size affected breeders' colony site fidelity between years. We compared philopatry at colonies where parasitic swallow bugs, Oeciacus vicarius, had been removed by fumigation with that at nonfumigated sites exposed to natural levels of ectoparasites. About 25% of birds at nonfumigated colonies returned to their previous year's site, whereas about 69% of birds at fumigated colonies did so. Site fidelity was greatest at nonfumigated sites that changed the least in size between years. Birds were less likely to return to a nonfumigated site as the colony there became increasingly larger. Individuals philopatric to both nonfumigated and fumigated sites resided in colonies more similar in size between years than did dispersing birds. Most cliff swallows settled within 6 km of their previous year's site, indicating that many nonphilopatric birds still may have had some familiarity with the local landscape surrounding the site to which they moved. Removal of ectoparasites at a site allows large colonies to persist there perennially, probably contributing to higher philopatry because such large colonies are rare and would have been difficult to find had the residents dispersed. Cliff swallows are likely to be sensitive to both colony size and general familiarity with a given site or landscape region, and probably integrate these with other cues to select breeding colonies.

Fluctuating survival selection explains variation in avian group size.

Most animal groups vary extensively in size. Because individuals in certain sizes of groups often have higher apparent fitness than those in other groups, why wide group size variation persists in most populations remains unexplained. We used a 30-y mark-recapture study of colonially breeding cliff swallows (Petrochelidon pyrrhonota) to show that the survival advantages of different colony sizes fluctuated among years. Colony size was under both stabilizing and directional selection in different years, and reversals in the sign of directional selection regularly occurred. Directional selection was predicted in part by drought conditions: birds in larger colonies tended to be favored in cooler and wetter years, and birds in smaller colonies in hotter and drier years. Oscillating selection on colony size likely reflected annual differences in food availability and the consequent importance of information transfer, and/or the level of ectoparasitism, with the net benefit of sociality varying under these different conditions. Averaged across years, there was no net directional change in selection on colony size. The wide range in cliff swallow group size is probably maintained by fluctuating survival selection and represents the first case, to our knowledge, in which fitness advantages of different group sizes regularly oscillate over time in a natural vertebrate population.

Costs and benefits of late nesting in cliff swallows.

Many organisms of temperate latitudes exhibit declines in reproductive success as the breeding season advances. Experiments can delay the onset of reproduction for early breeders to investigate the consequences of late nesting, but it is rarely possible to observe a distinct second round of nesting in species that normally nest only once. The colonial cliff swallow (Petrochelidon pyrrhonota) is a migratory songbird that has a relatively short breeding season in the western Great Plains, USA, with birds rarely nesting late in the summer. Previous work suggested that ectoparasitism is a primary reason why reproductive success in this species declines over the summer. At colony sites where nests were fumigated to remove ectoparasitic swallow bugs (Oeciacus vicarius), cliff swallows frequently undertook a distinct round of late nesting after previously fledging young that year. Mark-recapture revealed that late-nesting pairs at these colonies produced fewer offspring that survived to the next breeding season, and that survival of late-nesting adults was lower during the next year, relative to pairs nesting earlier in the season. These reproductive costs applied in the absence of ectoparasites and likely reflect other environmental costs of late nesting such as seasonal declines in food availability or a delayed start of fall migration. Despite the costs, the estimated fitness for perennial early-and-late nesters in the absence of ectoparasites was equivalent to that of birds that nested only early in the season. The collective disadvantages of late nesting likely constrain most cliff swallows to raising a single brood in the middle latitudes of North America.

Variation in age composition among colony sizes in Cliff Swallows.

Variation in group size is characteristic of most social species. The extent to which individuals sort among group sizes based on age may yield insight into why groups vary in size and the age-specific costs and benefits of different social environments. We investigated the age composition of Cliff Swallow (Petrochelidon pyrrhonota) colonies of different sizes over 18 years at a long-term study site in western Nebraska, USA. Using years elapsed since banding as a relative measure of age for over 194,000 birds, we found that the proportion of age-class-1 swallows (birds banded as nestlings or juveniles or adults in the year of banding) of both sexes increased in larger colonies and at colony sites becoming active later in the summer. Age composition was unrelated to how often a particular colony site was used. The effect of colony size most likely reflected the fact that older birds return to the same colony site in successive years even when the colony size there decreases, and that yearlings and immigrants benefit more from larger colonies than do older, more experienced individuals. The date effect probably resulted in part from later spring arrival by younger and/or immigrant swallows. At fumigated sites where ectoparasitic swallow bugs (Oeciacus vicarius) had been removed, age composition did not vary with either colony size or colony initiation date. The patterns reported here appear to be driven partially by the presence of ectoparasites and suggest that the hematophagous bugs influence variation in Cliff Swallow group composition. Our results are consistent with the hypothesis that variation in colony size reflects, in part, age-based sorting of individuals among groups.

Recapture heterogeneity in cliff swallows: increased exposure to mist nets leads to net avoidance.

Ecologists often use mark-recapture to estimate demographic variables such as abundance, growth rate, or survival for samples of wild animal populations. A common assumption underlying mark-recapture is that all animals have an equal probability of detection, and failure to meet or correct for this assumption-as when certain members of the population are either easier or more difficult to capture than other animals-can lead to biased and inaccurate demographic estimates. We built within-year and among-years Cormack-Jolly-Seber recaptures-only models to identify causes of capture heterogeneity for a population of colonially nesting cliff swallows (Petrochelidon pyrrhonota) caught using mist-netting as a part of a 20-year mark-recapture study in southwestern Nebraska, U.S.A. Daily detection of cliff swallows caught in stationary mist nets at their colony sites declined as the birds got older and as the frequency of netting at a site within a season increased. Experienced birds' avoidance of the net could be countered by sudden disturbances that startled them into a net, such as when we dropped a net over the side of a bridge or flushed nesting cliff swallows into a stationary net positioned at a colony entrance. Our results support the widely held, but seldom tested, belief that birds learn to avoid stationary mist nets over time, but also show that modifications of traditional field methods can reduce this source of recapture heterogeneity.

Heritable choice of colony size in cliff swallows: does experience trump genetics in older birds?

The variation in breeding-colony size seen in populations of most colonial birds may reflect heritable choices made by individuals who are phenotypically specialized for particular social environments. Although a few studies have reported evidence for genetically based choice of group sizes in birds, we know relatively little about the extent to which animals potentially rely on experience versus innate preferences in deciding with how many conspecifics to settle at different times of their lives. We conducted a cross-fostering experiment in 1997-1998 on cliff swallows (Petrochelidon pyrrhonota) in southwestern Nebraska, USA, in which some individuals were reared in colonies different in size from those in which they were born. Breeding-colony sizes chosen by this cohort of birds were monitored by mark-recapture throughout their lives. A multistate mark-recapture analysis revealed that birds in their first breeding year chose colony sizes similar to those of their birth, regardless of their rearing environment, confirming a previous analysis. Beyond the first breeding year, however, cliff swallows' colony choice was less dependent on where they were born. Birds born in small colonies and reared in large colonies showed evidence of a delayed rearing effect, with these birds overwhelmingly choosing large colonies in later years. Heritabilities suggested strong genetic effects on first-year colony choice but not in later years. Cliff swallows' genetically based colony-size preferences their first year could be a way to ensure matching of their phenotype to an appropriate social environment as yearlings. In later years, familiarity with particular colony sites and available information on site quality may override innate group-size preferences when birds choose colonies.