Kim Jaatinen

Stress responsiveness, age and body condition interactively affect flight initiation distance in breeding female eiders

Predation may drive differential selection among personality types, but the mechanism linking personality with predation risk is poorly understood. One such mechanism may be provided by stress hormones (corticosterone in birds), which are linked to boldness towards predators. However, because of feedbacks between boldness and future fitness expectations, the relationship between boldness and stress physiology may be modulated by individual quality. We investigated flight initiation distances (FIDs) of incubating eider, Somateria mollissima, females in relation to handling-induced corticosterone concentrations and individual quality (female breeding experience, body condition). We investigated whether FIDs were repeatable, and whether stress responsiveness and individual quality attributes, either independently or interactively, affected FIDs. We also analysed whether incubation duration, a period of peak predation pressure on females, would depend on female boldness towards predators (FID), controlling for individual quality. FIDs were repeatable within and between seasons, and females with higher handling-induced corticosterone concentrations generally had longer FIDs. However, this relationship was modulated by interactions between stress responsiveness and individual quality. High stress responsiveness was associated with longer FIDs in younger females, while the opposite was found for the oldest females. Furthermore, the FIDs of females in good body condition increased less strongly with increasing stress-induced circulating corticosterone. Shy females (long FIDs) and those in poor body condition had shorter incubation periods. Boldness is thus linked to stress responsiveness, helping to explain why stress responses may be correlated with survival. However, physiological stress effects on boldness cannot be understood in isolation from effects of individual quality.

Causes and consequences of fine-scale breeding dispersal in a female-philopatric species

The potentially confounded effects of factors affecting breeding dispersal have rarely been simultaneously examined. The consequences of breeding dispersal are even less studied, presenting a paradox: breeding dispersal seldom seems to improve breeding success, despite its presumed adaptiveness. We studied the causes and consequences of breeding dispersal in female-philopatric eiders (Somateria mollissima) in relation to the spatiotemporal predictability of nest success. Previous nest fate, breeding experience, and breeding density simultaneously affected breeding dispersal. Dispersal distances were longer among inexperienced breeders and after failed breeding. Individual dispersal distances decreased with increasing nest-site-specific breeding density, whereas island-specific nesting success peaked at intermediate densities. The fate of neighbouring nests (‘public information’) did not influence dispersal. Breeding dispersal was unrelated to subsequent hatching success, controlling for individual quality (body condition, breeding experience, previous nest fate), while it delayed hatch date, which is likely to impair reproductive success. This delay may result from the loss of acquired information of local breeding conditions, prolonging nest prospecting and establishment, also helping explain why breeding dispersal did not increase at high breeding densities, despite a potential reduction in nesting success. In long-lived species, however, dispersal-induced reductions in reproductive output in one season could be offset by improved parental survival prospects. Careful nest prospecting may be profitable, because overall nest success had a strong island-specific component but showed weak temporal variation, and successive individual nest fates were predictable between years. Once a safe nest site is found, females may breed at the same place successfully for many years.

Adult predation risk drives shifts in parental care strategies: a long‐term study

1. Grouping provides antipredatory benefits, and therefore aggregation tendencies increase under heightened predation risk. In socially breeding groups, however, conflicts over reproductive shares or safety tend to disintegrate groups. Group formation thereby involves a balance between the antipredatory benefits of aggregation and the destabilizing effect of reproductive conflict. 2. We study the grouping behaviour of a facultatively social precocial sea duck with uniparental female care, the eider (Somateria mollissima Linnaeus). Females tend their young solitarily or in groups of 2-5 females. Here, we focus on the effect predation on adults has on group-formation decisions of brood-caring females. 3. By modifying an existing bidding game over care, we model the effects of predation risk on the width of the window of selfishness, which delimits the reproductive sharing allowing cooperation within brood-rearing coalitions, and generate predictions about the relative frequencies of solitary versus cooperative parental care modes. Furthermore, we model the dilution effect as a function of female group size and predation risk. 4. The window of selfishness widens with increasing predation risk, and the dilution of predation risk increases with both female group size and increasing predation risk, yielding the following predictions: (i) cooperative brood care becomes more prevalent and, conversely, solitary brood care less prevalent under heightened predation risk and (ii) group sizes increase concomitantly. 5. We tested these predictions using 13 years of data on female grouping decisions and annual predation rates, while controlling for the potentially confounding effect of female body condition. 6. Our data supported both predictions, where heightened predation risk of nesting females, but not changes in their condition, increased the relative frequency of cooperative brood care. Increased female nesting mortality also resulted in larger groups of cooperative females. 7. The predation risk of incubating females has long-term implications for later parental care decisions. We discuss the potential consequences of predation-induced shifts in group size on per capita fitness and population-wide productivity.

Experience attracts: the role of age in the formation of cooperative brood-rearing coalitions in eiders

Lacking experience represents a constraint on solitary breeding, which may be overcome by joining groups of more experienced breeders. Also, biological market theory predicts preference for partnerships with conspecifics of the highest value. We examined the formation of cooperative brood-rearing coalitions in facultatively social eider females, Somateria mollissima. Our aim was to elucidate the hitherto neglected role of age on female group size and condition differences within groups, and to explore whether older females represent attractive coalition partners. Based on previous work, we hypothesized that older breeders are found in smaller groups, and that the attraction of older partners would override the predicted negative effect of differences in body condition on female group size. We found that older females occurred in smaller groups and that the negative relationship between condition differences and group size became less steep with increasing age of the oldest group member. We also found that the maximum body condition found in a brood-rearing coalition increased with the age of the oldest female, despite a sharp decline in the probability of the oldest individual being the one in best condition. These results demonstrate the importance of age in the formation of cooperative brood-rearing coalitions and the increase in female quality with age. The experience of older females is a valuable commodity, which attracts younger prospective coalition partners in good condition regardless of body condition differences. Our results illustrate the general principle that grouping decisions cannot be understood by viewing partner choice criteria in isolation.

Clutch Desertion in Barrow's Goldeneyes (Bucephala islandica) — Effects of Non-Natal Eggs, the Environment and Host Female Characteristics

Addition of eggs to nests of conspecifics is a common avian alternative breeding strategy, called conspecific brood parasitism. The consequences of this breeding strategy on recipient breeding success have seldom been quantified, while taking into account environmental factors and host female characteristics. We study the occurrence of nest parasitism and, using an information theoretic approach, the most important factors responsible for nest desertion in female Barrows goldeneyes (Bucephala islandica). Nest parasitism is common in the study populationpopulation, and 58% of the nests contained non-natal eggs, representing 20% of all eggs. A prime factor explaining nest desertion was the number of non-natal eggs. There were also significant effects of year and own clutch size. By contrast, ambient temperature and female laying date did not influence nest desertion. These results provide one of the first demonstrations that non-natal eggs can have substantial negative effects also in precocial species.

Safety in numbers: the dilution effect and other drivers of group life in the face of danger

Animals can congregate in groups for many reasons, from reproductive assurance to improved foraging or predation efficiency, to avoiding themselves becoming the target of predation by other animals. It is the last category that is the focus of this review: group living as protection from predation. The drivers of group life in the face of danger are at the same time diverse and interlinked, with much potential for confusion between concepts. Here we review these concepts, using the dilution effect as a starting point. We construct a mathematical model that allows us to examine various features of the dilution effect and their connection to ecology. We also show the importance of including a time scale when modelling the dilution effect and how this translates into more realistic estimation of the fitness consequences of a diluted predation risk. The central role of the dilution effect in creating safety in numbers is underlined by showing how it may affect life-history evolution and result in the emergence of gregarious life-history strategies, even among sessile organisms limited in their abilities to exhibit behavioural responses to predation. Finally, we review the other central processes underpinning group protection from predation: the satiation effect, selfish herding, the confusion effect and group vigilance.

Do female ornaments indicate quality in eider ducks

The fitness consequences of female ornamentation remain little studied and the results are often contradictory. Female ornamentation may be an artefact of a genetic correlation with male ornamentation, but this possibility can be disregarded if the ornament only occurs in females. Female-specific white wing bars in eiders (Somateria mollissima) have been suggested to indicate individual quality, and we studied size variation in this trait in relation to key fitness components and quality attributes. We found that clutch size, body condition, female age, hatching date and success were unrelated to female ornament size; ornament size was explained by its size in the previous year. In contrast, good body condition was associated with hatching success. These results suggest that the breadth of the white wing bars does not indicate individual quality in our study population.

Kin association during brood care in a facultatively social bird: active discrimination or by‐product of partner choice and demography?

Intra‐group relatedness does not necessarily imply kin selection, a leading explanation for social evolution. An overlooked mechanism for generating population genetic structure is variation in longevity and fecundity, referred to as individual quality, affecting kin structure and the potential for cooperation. Individual quality also affects choosiness in partner choice, a key process explaining cooperation through direct fitness benefits. Reproductive skew theory predicts that relatedness decreases with increasing group size, but this relationship could also arise because of quality‐dependent demography and partner choice, without active kin association. We addressed whether brood‐rearing eider (Somateria mollissima) females preferentially associated with kin using a 6‐year data set with individuals genotyped at 19 microsatellite loci and tested whether relatedness decreased with increasing female group size. We also determined the relationship between local relatedness and indices of female age and body condition. We further examined whether the level of female intracoalition relatedness differed from background relatedness in any year. As predicted, median female intra‐group relatedness decreased with increasing female group size. However, the proportion of related individuals increased with advancing female age, and older females prefer smaller brood‐rearing coalitions, potentially producing a negative relationship between group size and relatedness. There were considerable annual fluctuations in the level of relatedness between coalition‐forming females, and in 1 year this level exceeded that expected by random association. Thus, both passive and active mechanisms govern kin associations in brood‐rearing eiders. Eiders apparently can discriminate between kin, but the benefits of doing so may vary over time.

Differential responses to related hosts by nesting and non-nesting parasites in a brood-parasitic duck.

Host–parasite relatedness may facilitate the evolution of conspecific brood parasitism, but empirical support for this contention remains inconclusive. One reason for this disparity may relate to the diversity of parasitic tactics, a key distinguishing feature being whether the parasite has a nest of her own. Previous work suggests that parasites without nests of their own may be of inferior phenotypic quality, but because of difficulties in identifying these parasitic individuals, little is known about their host selection criteria. We used high‐resolution molecular maternity tests to assign parasitic offspring to known parasites with and without their own nests in a population of Barrow’s goldeneyes (Bucephala islandica). We determined whether parasite nesting status, host–parasite relatedness and distance between host and parasite nests affected the probability of parasitizing a host and the number of eggs laid per host. We also investigated whether nesting parasites, conventionally nesting females and non‐nesting parasites differed regarding their age, structural size, body condition, nesting phenology or total brood size. The probability of engaging in parasitism increased with host–parasite relatedness and spatial proximity to host nests for nesting and non‐nesting females alike. However, nesting parasites increased the number of eggs donated with relatedness to the host, while non‐nesting parasites did not do so. Non‐nesting parasites laid fewer eggs in total, but did not differ by any of the other quality measures from conventional nesters or nesting parasites. Our study provides the first demonstration that nesting and non‐nesting parasites from the same population may use different host selection criteria.

Brood size matching: a novel perspective on predator dilution.

A primary benefit of grouping is diluting the individual risk of attack by predators. However, the fact that groups are formed not always by solitary adults but also by subgroups (e.g., families) has been overlooked. The subgroup-specific benefit of predator dilution depends on its relative contribution to total group size. Therefore, the willingness of a subgroup to merge with others should increase the less it contributes to total group size, but the conflicting preferences of partners may result in the preferential merger of similar-sized subgroups. Here, we evaluate how the proportional contribution of subgroups to diluting risk affects group formation. We generate predictions using a bidding game over parental care and test them using data on common eiders (Somateria mollissima), in which females with variable-sized broods may form brood-rearing coalitions. The predictions (1) that size-matched subgroups should have a higher propensity to merge, (2) that predation should increase group formation propensity, and (3) that increased bargaining power, as proxied by female body condition, should increase the time needed to establish partnerships were all supported. Partners do negotiate over their relative contributions to predator dilution, accepting or rejecting partnerships on the basis of this criterion. Our results show that consideration of the size of subgroups before merger is critical in understanding the process of group formation under the threat of predation.