Predation cues amplify the effects of parasites on the personality of a keystone grazer.

Parasites can alter species interactions either by modifying infected host behaviour or by influencing behavioural responses in uninfected individuals. Salt marsh ecosystems are characterized by a predator-prey interaction between the keystone grazer, Littoraria irrorata, and its main predator, Callinectes sapidus, both integral players in mediating the productivity of these habitats. Littoraria also acts as the first intermediate host for at least four species of digenetic trematode. Parasite infection has been shown to decrease grazing and climbing in populations of Littoraria, although effects on infected host response to predators have not been investigated. Moreover, how infection might increase or decrease among-individual variation in behaviour (i.e. animal personality) is still unknown. Here we ask how trematode infection affects the expression of boldness in the anti-predator responses of L. irrorata in both the absence and presence of a predator cue. We find that individual boldness varies substantially, and repeatability tends to increase as the number of stressors increases, with infected individuals exposed to a predator cue showing the strongest expression of behavioural types. Parasitism amplifies this effect, although the parasite itself does not appear to directly induce behavioural changes: infected snails show no evidence of decreased climbing or differences in refuge use as compared to their uninfected counterparts. Infection might therefore drive the expression of condition-dependent personality differences evident only under high-risk conditions. Group infection status strongly influenced behavioural reaction norms: uninfected individuals grouped with an infected snail were more responsive to predation risk, exhibiting increased climbing behaviour and spending less time in the water. Here parasites are influencing personality indirectly by inducing avoidance behaviours in healthy individuals, although only in high-risk environments. The potential for exposure to parasites and predators fluctuates greatly across marsh ecosystems. Given the ecological importance of this predator-prey relationship, trematode infection can act as an important, although indirect, determinant of overall salt marsh community structure, health and function.

The Maternal Legacy: Female Identity Predicts Offspring Sex Ratio in the Loggerhead Sea Turtle.

In organisms with temperature-dependent sex determination, the incubation environment plays a key role in determining offspring sex ratios. Given that global temperatures have warmed approximately 0.6 °C in the last century, it is necessary to consider how organisms will adjust to climate change. To better understand the degree to which mothers influence the sex ratios of their offspring, we use 24 years of nesting data for individual female loggerhead sea turtles (Caretta caretta) observed on Bald Head Island, North Carolina. We find that maternal identity is the best predictor of nest sex ratio in univariate and multivariate predictive models. We find significant variability in estimated nest sex ratios among mothers, but a high degree of consistency within mothers, despite substantial spatial and temporal thermal variation. Our results suggest that individual differences in nesting preferences are the main driver behind divergences in nest sex ratios. As such, a female's ability to plastically adjust her nest sex ratios in response to environmental conditions is constrained, potentially limiting how individuals behaviorally mitigate the effects of environmental change. Given that many loggerhead populations already show female-biased offspring sex ratios, understanding maternal behavioral responses is critical for predicting the future of long-lived species vulnerable to extinction.

Kinship and the evolution of social behaviours in the sea.

Until recently, little attention has been paid to the existence of kin structure in the sea, despite the fact that many marine organisms are sessile or sedentary. This lack of attention to kin structure, and its impacts on social evolution, historically stems from the pervasive assumption that the dispersal of gametes and larvae is almost always sufficient to prevent any persistent associations of closely related offspring or adults. However, growing evidence, both theoretical and empirical, casts doubt on the generality of this assumption, not only in species with limited dispersal, but also in species with long dispersive phases. Moreover, many marine organisms either internally brood their progeny or package them in nurseries, both of which provide ample opportunities for kinship to influence the nature and outcomes of social interactions among family members. As the evidence for kin structure within marine populations mounts, it follows that kin selection may play a far greater role in the evolution of both behaviours and life histories of marine organisms than is presently appreciated.

Genetic relatedness influences plant biomass accumulation in eelgrass (Zostera marina).

In multispecies assemblages, phylogenetic relatedness often predicts total community biomass. In assemblages dominated by a single species, increasing the number of genotypes increases total production, but the role of genetic relatedness is unknown. We used data from three published experiments and a field survey of eelgrass (Zostera marina), a habitat-forming marine angiosperm, to examine the strength and direction of the relationship between genetic relatedness and plant biomass. The genetic relatedness of an assemblage strongly predicted its biomass, more so than the number of genotypes. However, contrary to the pattern observed in multispecies assemblages, maximum biomass occurred in assemblages of more closely related individuals. The mechanisms underlying this pattern remain unclear; however, our data support a role for both trait differentiation and cooperation among kin. Many habitat-forming species interact intensely with conspecifics of varying relatedness; thus, genetic relatedness could influence the functioning of ecosystems dominated by such species.

Exclusive male care despite extreme female promiscuity and low paternity in a marine snail.

Males exhibit striking variation in the degree to which they invest in offspring, from merely provisioning females with sperm, to providing exclusive post-zygotic care. Paternity assurance is often invoked to explain this variation: the greater a male's confidence of paternity, the more he should be willing to provide care. Here, we report a striking exception to expectations based on paternity assurance: despite high levels of female promiscuity, males of a marine snail provide exclusive, and costly, care of offspring. Remarkably, genetic paternity analyses reveal cuckoldry in all broods, with fewer than 25% of offspring being sired by the caring male, although caring males sired proportionally more offspring in a given clutch than any other fathers did individually. This system presents the most extreme example of the coexistence of high levels of female promiscuity, low paternity, and costly male care, and emphasises the still unresolved roles of natural and sexual selection in the evolution of male parental care.

Reproductive biology, family conflict, and size of offspring in marine invertebrates.

All organisms face two fundamental trade-offs in the allocation of energetic resources: one between many small versus a few large offspring, and the second between present and future reproduction. Nowhere are these trade-offs more apparent than in the vast range of variation in the sizes of eggs and offspring exhibited among species of marine invertebrates. It has become increasingly clear that, in many taxa of marine organisms, there is also substantial intraspecific variation in the size of eggs and hatchings. This variation has largely been attributed to adaptive maternal effects. In theory, however, the inevitable conflicts of interest that arise in families of sexually reproducing organisms over the optimal distribution of parental resources among siblings could also account for much of this variation in egg and offspring size. Here, we explore the potential impacts of family conflict on offspring traits by comparing the life histories of two exemplar species of marine organisms, the polychaete Boccardia proboscidea and the gastropod Solenosteira macrospira, emphasizing how differences in modes of fertilization and parental care might influence the phenotype and, consequently, the fitness of offspring.

Inter-seasonal maintenance of individual nest site preferences in hawksbill sea turtles.

Within a single population of hawksbill sea turtles (Eretmochelys imbricata), we found a behavioral polymorphism for maternal nest site choice with respect to beach microhabitat characteristics. Some females preferred to nest in littoral forest and in places with overstory vegetation cover, and others preferred to nest in more open, deforested areas. Nest site choice was consistent within and between nesting seasons two years apart. This was not a result of females simply returning to the same location along the shoreline; beach sections used by individual turtles varied between seasons. Nest site choice was not influenced by changes in beach environment (e.g., beach width and foliage cover) or by changes in females' reproductive output (e.g., clutch size), suggesting that fidelity to particular microhabitats is a major determinant of the observed nesting patterns. Because hawksbills exhibit temperature-dependent sex determination, if the behavioral polymorphism in nest site choice has a genetic basis, as is plausible, then this would have implications for sex ratio evolution and offspring survival. By taking an individual-based approach to the study of maternal behavior we reveal previously overlooked individual variation and hope to provide some impetus for more detailed studies of nest site choice.