Cooperative interactions among females can lead to even more extraordinary sex ratios.

Hamilton's local mate competition theory provided an explanation for extraordinary female-biased sex ratios in a range of organisms. When mating takes place locally, in structured populations, a female-biased sex ratio is favored to reduce competition between related males, and to provide more mates for males. However, there are a number of wasp species in which the sex ratios appear to more female biased than predicted by Hamilton's theory. It has been hypothesized that the additional female bias in these wasp species results from cooperative interactions between females. We investigated theoretically the extent to which cooperation between related females can interact with local mate competition to favor even more female-biased sex ratios. We found that (i) cooperation between females can lead to sex ratios that are more female biased than predicted by local competition theory alone, and (ii) sex ratios can be more female biased when the cooperation occurs from offspring to mothers before dispersal, rather than cooperation between siblings after dispersal. Our models formally confirm the verbal predictions made in previous experimental studies, which could be applied to a range of organisms. Specifically, cooperation can help explain sex ratio biases in Sclerodermus and Melittobia wasps, although quantitative comparisons between predictions and data suggest that some additional factors may be operating.

A solution to a sex ratio puzzle in Melittobia wasps.

The puzzling sex ratio behavior of Melittobia wasps has long posed one of the greatest questions in the field of sex allocation. Laboratory experiments have found that, in contrast to the predictions of theory and the behavior of numerous other organisms, Melittobia females do not produce fewer female-biased offspring sex ratios when more females lay eggs on a patch. We solve this puzzle by showing that, in nature, females of Melittobia australica have a sophisticated sex ratio behavior, in which their strategy also depends on whether they have dispersed from the patch where they emerged. When females have not dispersed, they lay eggs with close relatives, which keeps local mate competition high even with multiple females, and therefore, they are selected to produce consistently female-biased sex ratios. Laboratory experiments mimic these conditions. In contrast, when females disperse, they interact with nonrelatives, and thus adjust their sex ratio depending on the number of females laying eggs. Consequently, females appear to use dispersal status as an indirect cue of relatedness and whether they should adjust their sex ratio in response to the number of females laying eggs on the patch.

Reproductive interference hampers species coexistence despite conspecific sperm precedence.

Negative interspecific mating interactions, known as reproductive interference, can hamper species coexistence in a local patch and promote niche partitioning or geographical segregation of closely related species. Conspecific sperm precedence (CSP), which occurs when females that have mated with both conspecific and heterospecific males preferentially use conspecific sperm for fertilization, might contribute to species coexistence by mitigating the costs of interspecific mating and hybridization. We discussed whether two species exhibiting CSP can coexist in a local environment in the presence of reproductive interference. First, using a behaviorally explicit mathematical model, we demonstrated that two species characterized by negative mating interactions are unlikely to coexist because the costs of reproductive interference, such as loss of mating opportunity with conspecific partners, are inevitably incurred when individuals of both species are present. Second, we experimentally examined differences in mating activity and preference in two Harmonia ladybird species known to exhibit CSP. These behavioral differences may lead to local extinction of H. yedoensis because of reproductive interference by H. axyridis. This prediction is consistent with field observations that H. axyridis uses various food sources and habitats whereas H. yedoensis is confined to a less preferred prey item and a pine tree habitat. Finally, by a comparative approach, we observed that niche partitioning or parapatric distribution, but not sympatric coexistence in the same habitat, is maintained between species with CSP belonging to a wide range of taxa, including vertebrates and invertebrates living in aquatic or terrestrial environments. Taken together, it is possible that reproductive interference may destabilize local coexistence even in closely related species that exhibit CSP.

Temporal changes in spatial variation: partitioning the extinction and colonisation components of beta diversity.

The last two decades have witnessed unprecedented changes in beta diversity, the spatial variation in species composition, from local to global scales. However, analytical challenges have hampered empirical ecologists from quantifying the extinction and colonisation processes behind these changing beta diversity patterns. Here, we develop a novel numerical method to additively partition the temporal changes in beta diversity into components that reflect local extinctions and colonisations. By applying this method to empirical datasets, we revealed spatiotemporal community dynamics that were otherwise undetectable. In mature forests, we found that local extinctions resulted in tree communities becoming more spatially heterogeneous, while colonisations simultaneously caused them to homogenise. In coral communities, we detected non-random community disassembly and reassembly following an environmental perturbation, with a temporally varying balance between extinctions and colonisations. Partitioning the dynamic processes that underlie beta diversity can provide more mechanistic insights into the spatiotemporal organisation of biodiversity.

Intraspecific Adaptation Load: A Mechanism for Species Coexistence.

Evolutionary ecological theory suggests that selection arising from interactions with conspecifics, such as sexual and kin selection, may result in evolution of intraspecific conflicts and evolutionary 'tragedy of the commons'. Here, we propose that such an evolution of conspecific conflicts may affect population dynamics in a way that enhances species coexistence. Empirical evidence and theoretical models suggest that more abundant species is more susceptible to invasion of 'selfish' individuals that increase their own reproductive success at the expense of population growth (intraspecific adaptation load). The density-dependent intraspecific adaptation load gives rise to a self-regulation mechanism at the population level, and stabilizes species coexistence at the community level by negative frequency-dependence.

The evolution of stage-specific virulence: Differential selection of parasites in juveniles.

The impact of infectious disease is often very different in juveniles and adults, but theory has focused on the drivers of stage-dependent defense in hosts rather than the potential for stage-dependent virulence evolution in parasites. Stage structure has the potential to be important to the evolution of pathogens because it exposes parasites to heterogeneous environments in terms of both host characteristics and transmission pathways. We develop a stage-structured (juvenile-adult) epidemiological model and examine the evolutionary outcomes of stage-specific virulence under the classic assumption of a transmission-virulence trade-off. We show that selection on virulence against adults remains consistent with the classic theory. However, the evolution of juvenile virulence is sensitive to both demography and transmission pathway with higher virulence against juveniles being favored either when the transmission pathway is assortative (juveniles preferentially interact together) and the juvenile stage is long, or in contrast when the transmission pathway is disassortative and the juvenile stage is short. These results highlight the potentially profound effects of host stage structure on determining parasite virulence in nature. This new perspective may have broad implications for both understanding and managing disease severity.

Host manipulation by parasites as a cryptic driver of energy flow through food webs.

Manipulative parasites alter predator-prey interactions, and thus may facilitate, shift or create energy flow pathways through food webs (referred to hereafter as manipulation-mediated energy flow, MMEF). The ecological significance of MMEF would be determined not only by the strength of host manipulation, but also ecological and epidemiological factors, including host biomass, parasite incidence, and trophic position of the host-parasite association in their food webs. While previous theory has predicted that strong manipulation will destabilize host-parasite dynamics, a recently proposed theoretical framework claims that a switching strategy (sequential manipulation from predation suppression to enhancement) should allow parasites to induce strong predation enhancement and thus large MMEF. We formally outline the current and future directions to better understand the causes and consequences of MMEF across biological hierarchies.

Understanding the role of eco-evolutionary feedbacks in host-parasite coevolution.

It is widely recognised that eco-evolutionary feedbacks can have important implications for evolution. However, many models of host-parasite coevolution omit eco-evolutionary feedbacks for the sake of simplicity, typically by assuming the population sizes of both species are constant. It is often difficult to determine whether the results of these models are qualitatively robust if eco-evolutionary feedbacks are included. Here, by allowing interspecific encounter probabilities to depend on population densities without otherwise varying the structure of the models, we provide a simple method that can test whether eco-evolutionary feedbacks per se affect evolutionary outcomes. Applying this approach to explicit genetic and quantitative trait models from the literature, our framework shows that qualitative changes to the outcome can be directly attributable to eco-evolutionary feedbacks. For example, shifting the dynamics between stable monomorphism or polymorphism and cycling, as well as changing the nature of the cycles. Our approach, which can be readily applied to many different models of host-parasite coevolution, offers a straightforward method for testing whether eco-evolutionary feedbacks qualitatively change coevolutionary outcomes.

Host-Manipulation by Trophically Transmitted Parasites: The Switcher-Paradigm.

Host-manipulation by trophically transmitted parasites is thought to always predispose the intermediate hosts to enhanced predation by definitive hosts ('enhancement'). However, theory predicts that enhancement can disrupt stable, bottom-heavy predator-prey ratios, leading to fluctuation-driven extinction of intermediate hosts and parasites. How then can enhancement persist in nature despite this apparent instability? We address this paradox and conceptualize the 'switcher-paradigm', a novel framework incorporating sequential phases of reduced predation ('suppression') followed by enhancement. Theoretical models within the framework that consider 'switching' from suppression to enhancement indicate that switching likely increases parasite persistence and, in some circumstances, cancels out the effects of strong enhancement, leading to bottom-heavy predator-prey ratios. The switcher-paradigm confronts interdisciplinary research challenges, linking ecological processes across scales from within-host to community-wide dynamics.

Parasite infection drives the evolution of state-dependent dispersal of the host.

Dispersal plays a fundamental role in shaping the ecological processes such as host-parasite interactions, and the understanding of host dispersal tendency leads to that of parasites. Here, we present the result of our study on how the evolutionarily stable dispersal of a host would depend on parasite infection, considering kin competition among neighbours. We show that the evolving dispersal rate might be higher for susceptible than for infected individuals (S-biased dispersal) or vice versa (I-biased dispersal). S-biased dispersal is favoured by strong virulence affecting competitive ability, by high rate of parasite release during dispersal, and by low virulence for infected emigrants (i.e. low virulence affecting dispersal ability), whereas I-biased dispersal is favoured in the opposite situation. We also discuss population structure or between-deme genetic differentiation of the host measured with Wright's FST. In I-biased dispersal, between-deme genetic differentiation decreases with the infection rate, while in S-biased dispersal, genetic differentiation increases with infection rate.