The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

The use of conspecific reproductive success for breeding patch selection in terrestrial migratory species

Classical models of breeding habitat selection rarely deal with the question of information gathering for patch quality assessment. In this paper, we present two models comparing the fitness outcomes of behavioural strategies based on conspecific reproductive success as a cue to assess local environmental quality before selecting a new breeding habitat. The models deal with two phases of the life-cycle of a territorial migratory species: recruitment to a breeding population (model 1) and breeding site fidelity of subsequent breeding attempts (model 2). The first model shows that prospecting breeding patches before recruiting is the best strategy if the environment is predictable and contains a low proportion of good patches, even if it implies losing a breeding opportunity. The second model shows that dispersing after a breeding attempt according to the patch's breeding success rather than the individual's own success is the bests own success is the best strategy if the environment is patchy. These results underline the importance of studying the spatio-temporal variations of factors affecting reproductive success when considering the importance of habitat selection strategies based on conspecifics. Moreover, they allow the understanding of individual behaviour patterns observed in natural populations and their potential consequences at the metapopulation level.     

Ectoparasites cause increased bilateral asymmetry of naturally selected traits in a colonial bird

Abstract Parasitism has been shown to correlate with levels of bilateral symmetry in some organisms, with more asymmetric individuals often having more parasites. However, few studies have shown experimentally that parasitism directly causes increased asymmetry. By fumigating some cliff swallow (Petrochelidon pyrrhonota) colonies and leaving others untreated, we investigated experimentally whether ectoparasitism by the cimicid swallow bug led to higher levels of asymmetry in length of wings, outer tail feathers, and tarsus among juvenile and adult birds. Juveniles from fumigated colonies measured soon after fledging had significantly less asymmetry in wing and outer tail length than juveniles from nonfumigated colonies; asymmetry in tarsus length was unaffected by parasitism. Adults that had undergone one or more post‐juvenal molts on the wintering grounds showed no differences in asymmetry between those reared in fumigated vs. nonfumigated colonies. These results show that ectoparasitism directly leads to increased feather asymmetry in cliff swallows, probably through parasite‐induced nutritional stress. Because wing and tail asymmetry impair flight performance and reduce foraging efficiency, the increased asymmetry caused by parasites represents a fitness cost to cliff swallows. This is among the few experimental studies to show an effect of parasites on asymmetry of naturally selected characters.     

Kinship affects investment by helpers in a cooperatively breeding bird

Helping behaviour in cooperative breeding systems has been attributed to kin selection, but the relative roles of direct and indirect fitness benefits in the evolution of such systems remain a matter of debate. In theory, helpers could maximize the indirect fitness benefits of cooperation by investing more in broods with whom they are more closely related, but there is little evidence for such fine-scale adjustment in helper effort among cooperative vertebrates. In this study, we used the unusual cooperative breeding system of the long-tailed tit Aegithalos caudatus to test the hypothesis that the provisioning effort of helpers was positively correlated with their kinship to broods. We first use pedigrees and microsatellite genotypes to characterize the relatedness between helpers and breeders from a 14 year field study. We used both pedigree and genetic approaches because long-tailed tits have access to pedigree information acquired through social relationships, but any fitness consequences will be determined by genetic relatedness. We then show using both pedigrees and genetic relatedness estimates that alloparental investment by helpers increases as their relatedness to the recipients of their care increases. We conclude that kin selection has played a critical role in moulding the investment decisions of helpers in this cooperatively breeding species.     

Experimental test on public information use in the colonial Lesser Kestrel

The formation of colonies and the evolution of coloniality have been suggested to be a by-product of the use of public information to select breeding habitats. In this study we performed an experimental test to investigate whether the Lesser Kestrel (Falco naumanni) uses breeding success of conspecifics as a source of public information to select a breeding colony. We considered four potential cues based on reproductive performance of conspecifics in each colony: mean and variance of breeding success measured in both all pairs (TRS, total reproductive success), and only successful pairs (PRS, partial reproductive success). Both mean PRS and mean TRS could be predictive cues of the future reproductive output at a given site, because they varied among colonies and were autocorrelated from one year to the next. To dissociate any relationship between habitat quality and reproductive success we manipulated the mean brood sizes by transferring chicks among ten colonies. In five colonies, all the broods with at least two nestlings were reduced in one chick and in other five colonies all broods were enlarged by one chick. In the following year, we found that the number of adult immigrant kestrels settled at a colony was only explained by mean PRS in the previous year. We also examined the settlement of yearlings, which lack of information on breeding success in previous years. Unlike adult immigrant kestrels, yearlings tended to settle more frequently in those colonies with low mean PRS in the previous year, probably because of the lower pressure for occupation in those colonies. A multiple regression analysis showed that the growth of colonies was positively dependent on the mean PRS in the experimental year, and negatively on the number of predated nests. This study provides a solid support to the hypothesis that colonial species can use mean breeding success of successful pairs as a source of public information to make settlement decisions.     

Bateman's principle is reversed in a cooperatively breeding bird

Bateman''s principle is not only used to explain sex differences in mating behaviour, but also to determine which sex has the greater opportunity for sexual selection. It predicts that the relationship between the number of mates and the number of offspring produced should be stronger for males than for females. Yet, it is unclear whether Bateman''s principle holds in cooperatively breeding systems where the strength of selection on traits used in intrasexual competition is high in both sexes. We tested Bateman''s principle in the cooperatively breeding superb starling (Lamprotornis superbus), finding that only females showed a significant, positive Bateman gradient. We also found that the opportunity for selection was on average higher in females, but that its strength and direction oscillated through time. These data are consistent with the hypothesis that sexual selection underlies the female trait elaboration observed in superb starlings and other cooperative breeders. Even though the Bateman gradient was steeper for females than for males, the year-to-year oscillation in the strength and direction of the opportunity for selection likely explains why cooperative breeders do not exhibit sexual role reversal. Thus, Bateman''s principle may not hold in cooperative breeders where both sexes appear to be under mutually strong sexual selection.     

睾酮对鸟类繁殖影响研究进展

类固醇激素睾酮是影响鸟类繁殖最重要的性激素之一,与鸟类的繁殖行为的各个方面息息相关。睾酮通过影响鸟类的羽色、鸣声等来影响鸟类的配偶选择,同时睾酮可以调节配偶选择和繁殖投入之间的平衡。睾酮水平影响出雏数、出飞数、孵化率成功率等繁殖成效。睾酮还对个体的免疫活性和个体的存活率等产生影响。目前关于睾酮对鸟类繁殖影响的研究大多是通过外源性植入睾酮的方式来改变个体睾酮的浓度,其研究结果也常出现相互矛盾之处,对于自然状态下影响睾酮水平变化的因素尚缺乏了解,睾酮对雌雄鸟在繁殖过程中的影响也不尽相同,有必要继续深入研究。     

Public information and conspecific nest parasitism in goldeneyes: targeting safe nests by parasites

Conspecific nest parasitism (CNP) is a widespread alternativereproductive tactic in birds. Several hypotheses have been putforward to explain the evolution and occurrence of CNP, butno generally applicable hypothesis exists. Recent experimentalresults from the common goldeneye (Bucephala clangula), a cavity-nestingduck, have revealed that parasitic females preferentially layeggs in safe nest-sites, implying that nest predation risk isan important ecological determinant of CNP. The present studyfocuses on the mechanisms by which parasites identify safe nest-sites.Predation risk of a given nest-site was predictable betweensuccessive breeding seasons. At the end of the nesting season,females prospected active nest-sites more frequently than nest-sitesthat did not have a nest in the current season. Nest-sites thathad been prospected more frequently by females in year t hada higher probability to be parasitized in year t + 1. The resultssuggest that the use of public information, derived throughnest-site prospecting, enabled parasites to target safe nests.These findings provide a new and potentially generally applicableperspective to understand the evolution and occurrence of CNP.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

Colony size selection determines adult survival and dispersal preferences: allee effects in a colonial bird

Avian coloniality traditionally has been investigated by examining how breeding success varies with colony size, but other crucial fitness components rarely have been examined. This may lead to wrong conclusions because unmeasured parameters may change the final fitness balance. We used multistate capture-recapture models to investigate adult survival and dispersal in relation to colony size within a long-term monitored population of lesser kestrels (Falco naumanni). Nest predation probability decreases with colony size, and adult survival is predicted to show the same trend because adults are exposed to the same suite of predators. As expected, survival probability was higher in large colonies (0.72+/-0.015; mean+/-SE) than in medium or small colonies (0.65+/-0.02). Additionally, dispersal probabilities were higher going from small to large colonies (0.20+/-0.01) than from large to small (0.08+/-0.01), as predicted by theory of habitat selection shaped by fitness maximization. These asymmetries are likely to generate size-specific colony population dynamics, so they should be taken into account in studies of colonial birds and other metapopulation-like systems. Allee effects, that is, positive density dependence, appear to be the cause of the evolution of dispersal behavior and may explain the maintenance of coloniality in this species.     

Post-breeding information gathering and breeding territory shifts in northern wheatears

1. Prospecting non-breeding individuals have been shown to collect information on breeding sites a year ahead of breeding, but whether experienced breeders prospect future breeding sites is less well-known. Using data on post-breeding movements and between-year site shifts from a long-term population study of the migratory northern wheatear (Oenanthe oenanthe L.), we investigated (1) whether breeding territory selection of experienced breeders was a two-step process, made partly in the post-breeding period and partly at the time of territory establishment in the subsequent year; (2) predictions of which factors and cues correlate to site shifts at these two selection periods; and (3) consequences of territory shifts. 2. Many wheatears stayed close to their breeding site during the post-breeding period, but about 20% shifted to new potential breeding sites. Males that shifted to a new post-breeding location were also more likely to shift territory between years. 3. Factors linked to site shifts differed in the two investigated episodes of site selection. Post-breeding site shifts were linked to an environmental predictor of individual fitness, which is also related to foraging conditions (i.e. wheatears moved from tall to short field layers). Post-breeding site shifts, however, were not more frequent among young and failed breeders which may benefit most from prospecting for alternative breeding sites. Instead, in line with predictions of improving breeding conditions between-year site shifts were more frequent among young (only males) and failed breeders, whereas the link to territory field layer height disappeared. 4. Young males occupied more attractive sites and were more likely to breed successfully in year t + 1, but this improvement could also be expected from random choice. 5. Our results suggest that even though site shifts are determined partly during the post-breeding period, post-breeding movements of experienced breeders may be costly, especially when the breeding site is also a good foraging and moulting site in the post-breeding season. Under such circumstances an individual's choice of breeding site may often be determined at arrival in the subsequent year.     

Dispersal of sibling coalitions promotes helping among immigrants in a cooperatively breeding bird

Kin selection is a major force in social evolution, but dispersal is often assumed to reduce its impact by diluting kinship. In most cooperatively breeding vertebrates, in which more than two individuals care for young, juveniles delay dispersal and become helpers in family groups. In long-tailed tits (Aegithalos caudatus), however, offspring disperse to breed and helpers are failed breeders that preferentially aid kin. Helping also occurs among immigrants, but their origins are unknown and cooperation in these cases is poorly understood. Here, we combine long-term demographic and genetic data from our study population to investigate immigration and helping in this species. We first used a novel application of parentage analysis to discriminate between immigrants and unknown philopatric recruits. We then cross-checked sibship reconstruction with pairwise relatedness estimates to show that immigrants disperse in sibling coalitions and helping among them is kin biased. These results indicate that dispersal need not preclude sociality, and dispersal of kin coalitions may help maintain kin-selected cooperation in the absence of delayed dispersal.