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Connor E. Armstad

Publications and source records attributed to Connor E. Armstad.

3 recordsLinked to original sources

Within-group relatedness and patterns of reproductive sharing and cooperation in the tropical chestnut-crested yuhina

In cooperatively breeding animals, genetic relatedness among group members often determines the extent of reproductive sharing, cooperation and competition within a group. Studies of species for which cooperative behaviour is not entirely based on kinship are key for understanding the benefits favouring the evolution and maintenance of cooperative breeding among nonrelatives. In the cooperatively breeding chestnut-crested yuhina, Yuhina everetti , a songbird endemic to Borneo, we tested whether unrelated helpers are more likely to gain parentage than are related helpers consistent with the hypothesis that inbreeding risk constrains reproduction by related helpers. We also examined whether related or unrelated helpers provision broods more because of differences in their potential indirect or direct fitness benefits of helping. Kin structure of breeding groups (breeding pair and up to eight helpers of both sexes, median = 2 helpers, 96% of 57 pairs had helpers) based on genetic analysis was mixed; 48% of 76 breeder/helper dyads were first-order (26%) or second-order (22%) relatives of one or both members of the breeding pair, and 52% were nonrelatives. Only unrelated male and female helpers gained parentage, and helpers did not differ in their provisioning rate according to their relatedness to the broods. We documented quasi-parasitism or co-breeding by female helpers in 14% of 29 broods and extrapair paternity by male helpers in 21% of 47 broods. This rate of extrapair paternity is relatively high among the few tropical species examined but fit with predictions for mixed-kin groups where inbreeding is avoided. These findings support the emerging pattern for cooperative breeding in birds with mixed-kin groups, wherein unrelated helpers are more likely to gain parentage than are related helpers and helping effort is not necessarily predicted by kinship.

Animal Behaviour

Direct fitness benefits and kinship of social foraging groups in an Old World tropical babbler

Molecular studies have revealed that social groups composed mainly of nonrelatives may be widespread in group-living vertebrates, but the benefits favoring such sociality are not well understood. In the Old World, birds often form conspecific foraging groups that are maintained year-round and offspring usually disperse to other social groups. We tested the hypothesis that nonbreeding group members are largely unrelated and gain direct fitness benefits through breeding opportunities (males) and brood parasitism (females) in the tropical gray-throated babbler, Stachyris nigriceps , in Malaysian Borneo. Babblers foraged in social groups containing one or more breeding pairs (median = 8 group members of equal sex ratio), but group members rarely assisted with breeding (9% of 67 breeding pairs had a third helper; exhibiting facultative cooperative breeding). Although 20% of 266 group member dyads were first-order relatives of one or both members of the breeding pairs, 80% were unrelated. Male group members gained direct fitness benefits through extrapair and extra-group paternity (25% of 73 offspring), which was independent of their relatedness to the breeding pair and increased with decreasing group size. In contrast, females did not gain direct fitness benefits through brood parasitism. The low levels of relatedness and helping in social groups suggest that most group members do not gain indirect fitness benefits by helping to raise unrelated offspring. These findings highlight the importance of examining benefits of sociality for unrelated individuals that largely do not help and broaden the direct fitness benefits of group foraging beyond assumed survival benefits.

Behavioral Ecology

Enclosed nests may provide greater thermal than nest predation benefits compared with open nests across latitudes

Nest structure is thought to provide benefits that have fitness consequences for several taxa. Traditionally, reduced nest predation has been considered the primary benefit underlying evolution of nest structure, whereas thermal benefits have been considered a secondary or even non-existent factor. Yet, the relative roles of these factors on nest structures remain largely unexplored. Enclosed nests have a constructed or natural roof connected to sides that allow a restricted opening or tube entrance that provides cover in all directions except the entrance, whereas open nests are cups or platforms that are open above. We show that construction of enclosed nests is more common among songbirds (Passeriformes) in tropical and southern hemisphere regions than in north temperate regions. This geographic pattern may reflect selection from predation risk, under long-standing assumptions that nest predation rates are higher in southern regions and that enclosed nests reduce predation risk compared with open cup nests. We therefore compared nest predation rates between enclosed vs. open nests in 114 songbird species that do not nest in tree holes among five communities of coexisting birds, and for 205 non-hole-nesting species from the literature, across northern temperate, tropical, and southern hemisphere regions. Among coexisting species, enclosed nests had lower nest predation rates than open nests in two south temperate sites, but not in either of two tropical sites or a north temperate site. Nest predation did not differ between nest types at any latitude based on literature data. Among 319 species from both our field studies and the literature, enclosed nests did not show consistent benefits of reduced predation and, in fact, predation was not consistently higher in the tropics, contrary to long-standing perspectives. Thermal benefits of enclosed nests were indicated based on three indirect results. First, species that built enclosed nests were smaller than species using open nests both among coexisting species and among species from the literature. Smaller species lose heat fastest and thereby may gain important thermal benefits from reduced convective cooling. Second, eggs were warmed by parents for less time in species with enclosed nests, as can be expected if egg cooling rates are slower. Finally, species using enclosed nests exhibited enhanced growth of mass and wings compared with species using open nests, suggesting reduced thermoregulatory costs allowed increased energy for growth. Enclosed nests may therefore provide more consistent thermal than nest predation benefits, counter to long-standing perspectives.

Functional Ecology