Do common eiders nest in kin groups? Microgeographic genetic structure in a philopatric sea duck

We investigated local genetic associations among female Pacific common eiders (Somateria mollissima v‐nigrum) nesting in a stochastic Arctic environment within two groups of barrier islands (Simpson Lagoon and Mikkelsen Bay) in the Beaufort Sea, Alaska. Nonrandom genetic associations were observed among nesting females using regional spatial autocorrelation analyses for distance classes up to 1000 m in Simpson Lagoon. Nearest‐neighbour analyses identified clusters of genetically related females with positive lr values observed for 0–13% and 0–7% of the comparisons in Simpson Lagoon and Mikkelsen Bay, respectively, across years. These results indicate that a proportion of females are nesting in close proximity to more genetically related individuals, albeit at low frequency. Such kin groupings may form through active association between relatives or through natal philopatry and breeding site fidelity. Eiders nest in close association with driftwood, which is redistributed annually by seasonal storms. Yet, genetic associations were still observed. Microgeographic structure may thus be more attributable to kin association than natal philopatry and site fidelity. However, habitat availability may also influence the level of structure observed. Regional structure was present only within Simpson Lagoon and this island group includes at least three islands with sufficient driftwood for colonies, whereas only one island at Mikkelsen Bay has these features. A long‐term demographic study is needed to understand more fully the mechanisms that lead to fine‐scale genetic structure observed in common eiders breeding in the Beaufort Sea.     

Philopatry and within-colony movements in Columbian ground squirrels

Philopatry and dispersal result in selection of habitat locations that may differ in resources and social environment and thus should influence fitness components like survival and reproduction. We examined short-distance movements of young and adult females from natal or previous nesting sites within a colony of Columbian ground squirrels (Urocitellus columbianus) in the Rocky Mountains of Alberta, Canada, over a 17-year period. Females of all ages were strongly philopatric, yet a few (10-15%) exhibited movements that took them to new home ranges. We tested three hypotheses to explain the pattern of female natal and breeding movements: (1) that movements of philopatric females promote proximity to close kin; (2) that range shifts favour close kin via bequeathal of territory and (3) that dispersers move to lower density areas where competition for resources is lower. Tests of these three hypotheses revealed that: (1) philopatry and movements of young and older philopatric females led to proximity to mothers and local presence of close kin; (2) breeding dispersal did not result in bequeathal of home range to daughters, but movements of philopatric females suggested that they shared space with close kin and (3) adult females moved to new ranges with lower local densities, though dispersing females also left ranges where local density was significantly lower than for philopatric females. Natal and breeding movements among years produced two opportunities for territorial females: close spatial proximity to close kin via short philopatric movements, and habitats with fewer competitors via longer dispersal movements.     

Fine-scale study finds limited evidence of kin clustering in a gray seal colony

Microsatellites were used to investigate fine‐scale spatial and temporal genetic structure of a gray seal breeding colony, using samples collected throughout the colony (1997) and more extensive local sampling (2000–2002). Previous behavior of breeding seals on North Rona, including philopatry and high breeding site fidelity, suggested female kin may cluster together. However, low F_ST scores indicated no genetic differentiation between the major breeding aggregations in different regions of the colony. Nevertheless, the pairwise relatedness of mothers within regions was significantly higher than the relatedness of mothers between regions. Evidence of kin clustering occurred within one region in the colony. Within this region, mothers' pairwise relatedness decreased significantly with increasing distance between pupping sites. Pairwise relatedness of neighboring females within this region was also higher than expected. Conversely, in the other regions mothers that were considered likely to have social interactions, based on their spatial and temporal proximity, were not more related to each other than random. Therefore, sustained philopatry appears sufficient to produce differences in relatedness at a within‐colony scale (>500 m), but not at finer scales. This suggests that long‐term associations of mothers on North Rona detected previously are unlikely to be formed solely by close kin.     

Experimental evaluation of nest shelter effects on weight loss in incubating common eiders Somateria mollissima

Evaluating consequences of habitat selection is an important step in understanding life history strategies and behavioural decisions of animals. Kilpi and Lindström (1997) found that incubating common eiders Somateria mollissima on exposed, treeless islands lost weight faster than females nesting on wooded islands and proposed that this difference was due to adverse incubation conditions at exposed nests. Therefore, we tested whether common eiders gained an advantage when nesting in sheltered habitats by placing artificial shelters over randomly-selected females after the onset of incubation within an eider colony in arctic Canada. We predicted that sheltered females would be heavier on completion of incubation than control hens lacking shelters. Females nesting in artificial shelters experienced a more moderate thermal environment at both cold and warm temperature extremes. Eiders nesting in shelters were heavier than control females during mid incubation, consistent with habitat-specific rates of weight loss reported by Kilpi and Lindström (1997) . Natural overhead cover was available at potential nests but few eiders used those sites. We suspect that microclimatic advantages offered by sheltered sites may be offset by costs of increased female vulnerability to predators. Further work is needed to test this hypothesis, and to determine mechanisms responsible for lower weight loss in eiders attending well concealed nests.     

Female distribution, genetic relatedness, and fostering behaviour in harbour seals, Phoca vitulina

Although harbour seals may not recognize their relatives, relatives could be chosen preferentially for fostering (i.e. kin selection) if harbour seals display natal philopatry coupled with breeding site fidelity, and thus kin are clustered within the colony. We used behavioural and genetic data to investigate population structure within the Sable Island breeding colony and to test whether harbour seals tend to foster related pups. Adult females on Sable Island showed a high level of breeding-colony site fidelity but low levels of within-colony site fidelity both within and between years. Similarly, although lactating females showed a clumped distribution, group composition was highly variable, suggesting that this study colony was not composed of groups of related animals. DNA fingerprint data supported the hypothesis that female distribution within the colony was not correlated with genetic relatedness. Furthermore, the mean DNA band sharing among foster dyads did not differ significantly from that for unrelated animals. These results indicate that among harbour seals, related pups are not usually chosen preferentially for fostering and hence, kin selection is not likely to be influencing the occurrence of this behaviour. Copyright 1999 The Association for the Study of Animal Behaviour.     

Genetic evidence for kin-based female social structure in common eiders (Somateria mollissima)

Kin-based social groups are commonly studied among cooperativelybreeding species but have been less studied in "nontraditional"group breeding systems. We investigated the presence of kin-basedsociality among females in the common eider (Somateria mollisima),a colonial nesting sea duck that exhibits high levels of natalphilopatry in females. Previous studies of female socialityin common eiders have been restricted to observations duringbrood rearing. However, aggregations of female common eidersare also observed during other periods of the life cycle suchas colony arrival and nesting. Here we apply a novel, empiricalframework using molecular markers and field sampling to geneticallycharacterize female social groups at several stages of the commoneider life cycle. When compared with mean estimates of interindividualrelatedness for the entire colony, significantly higher levelsof relatedness were found between females within groups arrivingto the colony in flight, between females and nearest neighborsat the time of nest site selection, and between groups of femalesdeparting the colony with ducklings. Both full-sibling and half-siblingequivalent relationships were also found within these groups.Therefore, throughout each of several stages including in-flightcolony arrival, nesting, and brood rearing, we provide the firstgenetically confirmed evidence of female kin-based social groupsin common eiders and anseriformes in general.     

Complex social organization reflects genetic structure and relatedness in the cooperatively breeding bell miner, Manorina melanophrys

The cooperatively breeding bell miner, Manorina melanophrys, differs from most other cooperative breeding species in the complexity of its social system, where discrete social organization occurs on at least three levels. Microsatellite markers were used to investigate the degree of genetic structure underlying the social organization of M. melanophrys by comparing colonies, coteries and nest contingents. The genetic data confirmed behavioural observations of M. melanophrys living in male kin-based groups between which females disperse short distances to breed. Estimates of FST revealed restricted gene flow between eight colonies located within 30 km that was significantly associated with geographical distance when the two most distant colonies were included. Within a high density colony significant differences were found between coteries; analysis of the degree of relatedness between coterie members showed that this is due to related individuals associating preferentially with each other. Similarly, the contingent of individuals attending a nest were generally close relatives of the young they were aiding, supporting models invoking kin selection as the selective agency mediating helping.     

Kin grouping is insufficient to explain the inclusive fitness gains of conspecific brood parasitism in the common eider

Conspecific brood parasitism allows females to exploit other females' nests and enhance their reproductive output. Here, we test a recent theoretical model of how host females gain inclusive fitness from brood parasitism. High levels of relatedness between host and parasitizer can be maintained either by: (a) kin recognizing and parasitizing each other as a form of cooperative breeding or (b) natal philopatry and nest site fidelity facilitating the formation of kin groups, thereby increasing the probability of parasitism between relatives nesting in close proximity. To address these two hypotheses we genotyped feathers and hatch membranes of common eiders (Somateria mollissima) from western Hudson Bay, Canada, using a noninvasive sampling methodology. We found that most instances of brood parasitism do result in inclusive fitness gains. Furthermore, females with failed nests moved an average of 492 m from their previous year's nest site, while successful females only moved an average of 13 m. Therefore, we observed host–parasite relatedness can occur at levels higher than would be expected by chance even in the absence of kin grouping, suggesting that closely related females nesting near one another is not essential to maintain high host–parasitizer relatedness. In addition, kin grouping is only a transient phenomenon that cannot occur every year due to the propensity for females of failed nests to nest farther away from their nest site in subsequent years than females with successful nests, which provides support for kin recognition as a more likely mechanism to maintain high host–parasitizer relatedness over time.     

Age-related reproductive success in solitarily and communally nesting female dormice (Glis glis)

The factors underlying the occurrence of communal nesting in a natural population of fat dormouse ( Glis glis ) breeding in nest-boxes have been analysed on the assumption that such behaviour is adaptive. Since co-nesting females were previously shown to be close kin (prevalently mother-daughter pairs), they were expected to obtain kin-selected benefits not enjoyed by solitary breeders. The only advantage so far detected accrues to the younger partner, which gives birth, on average, several days earlier than it would as a solitary breeder and thus allows its progeny to attain a heavier weight by the time of hibernation. On the other hand, pup predation in the nest, presumably caused by the older female towards the younger's litter, was occasionally recorded. On the whole, it is perhaps more plausible that communal nesting results from delayed dispersal of yearling females when a high population density limits breeding space, as proposed for other small rodents.     

Foraging locations of female New Zealand sea lions (

Figure of Eight Island is located in the southern end of the Auckland Islands and hosts the smallest breeding colony of New Zealand (NZ) sea lions (Phocarctos hookeri). Between 1995/96 and 2005/06, pup production in this colony decreased by 57% (from 144 to 62 pups). In contrast, there was a 30% decrease in pup production in the largest colony in the north-east of the Auckland Islands over the same period. NZ sea lions in the Auckland Islands area are subject to by-catch deaths and resource competition from subantarctic trawl fisheries. The present study investigated where four lactating females from Figure of Eight Island foraged during the austral summer of 2007/08 and compared their foraging areas with female NZ sea lions from the northern Auckland Islands breeding locations (Enderby and Dundas islands) and with fisheries activities. Females foraged south of Adams Island (the southernmost Auckland Island), predominantly at the edge of the Auckland Islands shelf, but those from Figure of Eight Island made shorter foraging trips within more concentrated areas than females from Enderby or Dundas islands. The 59 female NZ sea lions satellite-tracked to date from Figure of Eight, Enderby and Dundas islands foraged over the entire area of the Auckland Islands shelf and many (including three of the four females from Figure of Eight Island) had extensive overlap with subantarctic trawl fisheries. Further research is needed to determine whether the foraging behaviour of females from Figure of Eight Island is linked to their greater decline in pup production.     

Matrilineal kin clusters and their effect on reproductive success in the field vole Microtus agrestis

Kin dusters of philopatric females are thought to form the basicsocial organization of breeding females in various vole species.The spatial association between young breeding females and theirmothers was studied in five 2500–m² enclosures in naturalhabitats during mid- and late breeding season in summer 1992.The mothers (founder females) were introduced into the enclosuresin die beginning of the study in June, whereas the young breedingfemales were born within the enclosures. Matrilineal kinshipwas determined by using radionudides. In July-August, 58% ofyoung breeding females were philopatric, but 82% of the mother-daughterassociations broke up until August-September. Of the young femalesthat started to breed in August-September, only 12% were philopatricThus the kin clusters found were short Hved and few in numberduring the late breeding season. The reduction in the incidenceof the kin clusters coincided with an increase of populationdensity. The mean reproductive success of females breeding inkin clusters was better than that of those breeding separately.In addition, among the nonphilopatric females there was a positive,significant correlation between reproductive success and thedistance to the nearest unrelated founder female. Such correlationwas not found among the philopatric females.     

Adoption of chicks and the level of relatedness in common gull, Larus canus, colonies: DNA fingerprinting analyses

In common gull colonies on islands of the Vistula River, Poland, adoption of chicks is common. In 1997, we observed 81 chicks from 35 nests. Of these, 19 (23.4%) left their natal broods and were adopted by other pairs. Another 11 (31.4%) were driven from the foreign territory by the owners. Foreign chicks were adopted by 15 pairs (42.9%). Eleven pairs (31.4%) drove foreign chicks from the territory. To test if the frequent adoptions in these colonies could be explained by kin selection or the occurrence of kin groups, we calculated band-sharing coefficients and genetic relatedness (r) between interacting birds (neighbours and non-neighbours). Adults that adopted were most often neighbours of the biological parents of adopted chicks, whereas spatially segregated birds, nesting further away, usually drove off the chicks. Band-sharing coefficients between males, but not females, were higher with decreasing internest distances. The band-sharing coefficients for adopted chicks and foster parents were significantly higher than for adopted chicks and randomly selected, spatially segregated pairs from the same and another colony. Band-sharing coefficients of adopted chicks and adopting neighbours (males: r=0.20; females: r=0.16) also tended to be higher than those of rejected chicks and rejecting neighbours (both sexes: r=0.08). Our results suggest that kin groups of neighbours do occur in common gull colonies. Such social structure might lead to indirect inclusive fitness benefits of adopting pairs. Differences in genetic similarity between chicks and adopting or rejecting neighbours show that at least in common gulls we should consider kin altruism as a factor in adoptions. Copyright 2000 The Association for the Study of Animal Behaviour.     

Pre-reproductive alliance formation in female wild house mice (<Emphasis Type="Italic">Mus domesticus</Emphasis>): the effects of familiarity and age disparity

Female house mice (Mus domesticus) are known to perform communal breeding more often with kin than with non-kin partners. When mice are grouped in semi-natural enclosures, related females develop pre-reproductive alliances more frequently than unrelated ones. However, little is known about the behavioural mechanisms and the factors promoting kin preferential cooperative associations in female mice. Here we evaluate the relative importance of familiarity and age disparity on the pre-reproductive development of agonistic behaviour and spatial associations within groups of three related females, freely interacting in semi-natural indoor enclosures. We found that familiarity clearly promoted female alliance formation by reducing aggression and enhancing spatial cohabitation, while genetic relatedness per se did not alleviate the effects of unfamiliarity on female grouping. Older sisters clearly dominated younger ones, even if they had lower body weights, or if they had to confront two allied littermate sisters. Hence, our findings corroborate previous findings on familiarity as a proxy for kin-preferential alliance formation in female mice. Moreover, we observed a strong impact of age stratification on female aggression-mediated dominance development. We suggest that this age effect could be expected from a queuing-for-reproduction view of younger house mouse females.Communicated by I. Schlupp     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.     

Complex pattern of genetic structuring in the Atlantic salmon (Salmo salar L.) of the River Foyle system in northwest Ireland: disentangling the evolutionary signal from population stochasticity

Little is known about the microevolutionary processes shaping within river population genetic structure of aquatic organisms characterized by high levels of homing and spawning site fidelity. Using a microsatellite panel, we observed complex and highly significant levels of intrariver population genetic substructure and Isolation-by-Distance, in the Atlantic salmon stock of a large river system. Two evolutionary models have been considered explaining mechanisms promoting genetic substructuring in Atlantic salmon, the member-vagrant and metapopulation models. We show that both models can be simultaneously used to explain patterns and levels of population structuring within the Foyle system. We show that anthropogenic factors have had a large influence on contemporary population structure observed. In an analytical development, we found that the frequently used estimator of genetic differentiation, F(ST), routinely underestimated genetic differentiation by a factor three to four compared to the equivalent statistic Jost's D(est) (Jost 2008). These statistics also showed a near-perfect correlation. Despite ongoing discussions regarding the usefulness of "adjusted"F(ST) statistics, we argue that these could be useful to identify and quantify qualitative differences between populations, which are important from management and conservation perspectives as an indicator of existence of biologically significant variation among tributary populations or a warning of critical environmental damage.     

Colony kin structure and host‐parasite relatedness in the barnacle goose

Conspecific brood parasitism (CBP), females laying eggs in the nest of other ‘host’ females of the same species, is a common alternative reproductive tactic among birds. For hosts there are likely costs of incubating and rearing foreign offspring, but costs may be low in species with precocial chicks such as waterfowl, among which CBP is common. Waterfowl show strong female natal philopatry, and spatial relatedness among females may influence the evolution of CBP. Here we investigate fine‐scale kin structure in a Baltic colony of barnacle geese, Branta leucopsis, estimating female spatial relatedness using protein fingerprints of egg albumen, and testing the performance of this estimator in known mother‐daughter pairs. Relatedness was significantly higher between neighbour females (nesting ≤ 40 metres from each other) than between females nesting farther apart, but there was no further distance trend in relatedness. This pattern may be explained by earlier observations of females nesting close to their mother or brood sisters, even when far from the birth nest. Hosts and parasites were on average not more closely related than neighbour females. In 25 of 35 sampled parasitized nests, parasitic eggs were laid after the host female finished laying, too late to develop and hatch. Timely parasites, laying eggs in the host’s laying sequence, had similar relatedness to hosts as that between neighbours. Females laying late parasitic eggs tended to be less related to the host, but not significantly so. Our results suggest that CBP in barnacle geese might represent different tactical life‐history responses.     

Mating patterns, relatedness and the basis of natal philopatry in the brown long-eared bat, Plecotus auritus

The brown long-eared bat, Plecotus auritus, is unusual among temperate zone bats in that summer maternity colonies are composed of adult males and females, with both sexes displaying natal philopatry and long-term association with a colony. Here, we describe the use of microsatellite analysis to investigate colony relatedness and mating patterns, with the aim of identifying the evolutionary determinants of social organization in P. auritus. Mean colony relatedness was found to be low (R=0.033 +/- 0.002), with pairwise estimates of R within colonies ranging from -0.4 to 0.9. The proportion of young fathered by males in their own colony was investigated using a Bayesian approach, incorporating parameters detailing the number of untyped individuals. This analysis revealed that most offspring were fathered by males originating from a different colony to their own. In addition, we determined that the number of paternal half-sibs among cohorts of young was low, inferring little or no skew in male reproductive success. The results of this study suggest that kin selection cannot account for colony stability and natal philopatry in P. auritus, which may instead be explained by advantages accrued through the use of familiar and successful roost sites, and through long-term associations with conspecifics. Moreover, because the underlying causes of male natal dispersal in mammals, such as risk of inbreeding or competition for mates, appear to be avoided via extra-colony copulation and low male reproductive skew, both P. auritus males and females are able to benefit from long-term association with the natal colony.     

Agonistic display and spacing in the black-headed gull, Larus ridibundus

Black-headed gulls nesting on islands in a marsh habitat had a higher breeding success than those nesting on the periphery of the colony. Island birds maintained close inter-nest distances whereas peripheral nests were spaced further apart as predation of eggs increased. These changes in spacing corresponded with changes in the response distances of agonistic displays. There was no change during incubation for island birds but peripheral birds displayed to intruders at much greater distances than during pair formation. Island birds were more likely to attack and display more frequently to intruders than peripheral birds. However, in one season the peripheral birds were more aggressive during pair formation because of an increase in intrusions by unpaired females.     

Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

Communal Nesting in Asocial Abert's Squirrels: the Role of Social Thermoregulation and Breeding Strategy

The social thermoregulation hypothesis states that endothermic species will communally nest to reduce energy expenditures on thermoregulation. The hypothesis predicts that the frequency of communal nesting should increase with decreasing ambient temperature. The potential costs of communal nesting (e.g., increased predation risk, resource competition, cuckoldry, parasite/disease transmission, or infanticide) could decrease the frequency of communal nesting especially for asocial breeding females with dependent offspring. We examined the effects of ambient temperature and seasonal reproductive activities on the probability of communal nesting in Abert's squirrels (Sciurus aberti) in the Pinaleño Mountains, Arizona. Most squirrels nested consistently with the same partner in mixed‐sex pairs. The proportion of individuals engaging in communal nesting increased with decreasing ambient temperature as predicted by the social thermoregulation hypothesis. The onset of the breeding season greatly reduced the proportion of individuals communally nesting. The negative relationship between ambient temperature and communal nesting supports the use of communal nesting in Abert's squirrels as a mechanism to reduce thermoregulatory costs during cold conditions. The abrupt drop in the frequency of communal nesting during the breeding season is likely due to female abandonment of this behavior. By avoiding communally nesting during the breeding season, females may prevent males from mating with them outside of mating chases, reduce resource competition, and protect offspring from infanticide, diseases, and parasites. Males may gain additional fitness benefits from nesting with females because familiarity with females increases dominance rank in mating activities.