Animals forming social groups that include breeders and nonbreeders present evolutionary paradoxes; why do breeders tolerate nonbreeders? and why do nonbreeders tolerate their situation? Both paradoxes are often explained with kin selection. Kin selection is, however, assumed to play little or no role in social group formation of marine organisms with dispersive larval phases. Yet, in some marine organisms, recent evidence suggests small-scale patterns of relatedness, meaning that this assumption must always be tested. Here, we investigated the genetic relatedness of social groups of the emerald coral goby, Paragobiodon xanthosoma. We genotyped 73 individuals from 16 groups in Kimbe Bay, Papua New Guinea, at 20 microsatellite loci and estimated pairwise relatedness among all individuals. We found that estimated pairwise relatedness among individuals within groups was significantly higher than the pairwise relatedness among individuals from the same reef, and pairwise relatedness among individuals from the same reef was significantly higher than the pairwise relatedness among individuals from different reefs. This spatial signature suggests that there may be very limited dispersal in this species. The slightly positive relatedness within groups creates the potential for weak kin selection, which may help to resolve the paradox of why breeders tolerate subordinates in P. xanthosoma. The other paradox, why nonbreeders tolerate their situation, is better explained by alternative hypotheses such as territory inheritance, and ecological and social constraints. We show that even in marine animals with dispersive larval phases, kin selection needs to be considered to explain the evolution of complex social groups.

A central issue in evolutionary ecology is how patterns of dispersal influence patterns of relatedness in populations. In terrestrial organisms, limited dispersal of offspring leads to groups of related individuals. By contrast, for most marine organisms, larval dispersal in open waters is thought to minimize kin associations within populations. However, recent molecular evidence and theoretical approaches have shown that limited dispersal, sibling cohesion and/or differential reproductive success can lead to kin association and elevated relatedness. Here, we tested the hypothesis that limited dispersal explains small-scale patterns of relatedness in the pajama cardinalfish Sphaeramia nematoptera. We used 19 microsatellite markers to assess parentage of 233 juveniles and pairwise relatedness among 527 individuals from 41 groups in Kimbe Bay, Papua New Guinea. Our findings support three predictions of the limited dispersal hypothesis: (i) elevated relatedness within groups, compared with among groups and elevated relatedness within reefs compared with among reefs; (ii) a weak negative correlation of relatedness with distance; (iii) more juveniles than would be expected by chance in the same group and the same reef as their parents. We provide the first example for natal philopatry at the group level causing small-scale patterns of genetic relatedness in a marine fish.

Plasticity, the capacity of individuals to respond to changing environments by modifying traits, may be critically important for population persistence by allowing for adaptive responses on shorter timescales than genetic change. Here, we use the clown anemonefish Amphiprion percula, whose access to resources is constrained by their anemones, to test the role of plasticity in generating variation in reproductive success among groups. We surveyed a wild clownfish population and found positive correlations between anemone area, fish size, reproduction and parental care. We used structural equation modeling to test the hypothesis that these correlations emanate from variation in anemone area and found support for a pathway linking anemone area to female investment, female investment to male investment and male investment to embryo survival. Next, we experimentally tested whether plasticity in response to resource availability can result in variation in parental traits using a feeding manipulation and found substantial plasticity in reproduction and parental care in response to changes in the availability of food resources. The results of this study reveal the role of plasticity in response to local resource availability in generating variation among individuals in reproductive strategies, linking studies of behavior and demography in this model species, and ultimately contributing to our ability to predict how populations might cope with environmental changes.

. In social groups, high reproductive skew is predicted to arise when the reproductive output of a group is limited, and dominant individuals can suppress subordinate reproductive efforts. Reproductive suppression is often assumed to occur via overt aggression or the threat of eviction. It is unclear, however, whether the threat of eviction alone is sufficient to induce reproductive restraint by subordinates. Here, we test two assumptions of the restraint model of reproductive skew by investigating whether resource limitation generates reproductive competition and whether the threat of eviction leads to reproductive restraint in the clown anemonefish
. Amphiprion percula
. First, we use a feeding experiment to test whether reproduction is resource limited, which would create an incentive for the dominant pair to suppress subordinate reproduction. We show that the number of eggs laid increased in the population over the study period, but the per cent increase in fed groups was more than twice that in unfed groups (205% and 78%, respectively). Second, we use an eviction experiment to test whether the dominant pair evicts mature subordinates, which would create an incentive for the subordinates to forgo reproduction. We show that mature subordinates are seven times more likely to be evicted than immature subordinates of the same size. In summary, we provide experimental support for the assumptions of the restraint model by showing that resource limitation creates reproductive competition and a credible threat of eviction helps explain why subordinates forego reproduction. Transactional models of reproductive skew may apply well to this and other simple systems.
.