Samuel Lymbery

Female mate choice is a major mechanism of sexual selection, influencing the evolution of behaviour, morphology and life-history. Despite being so important, much about female mate choice remains unclear. Crucially, we have a limited understanding of how mating decisions are affected by the social environment within which these decisions inevitably occur. Here, we review how the social environment influences female mate choice and discuss the wider evolutionary consequences of these effects. We examine important social environment influences on female choosiness (i.e. how much females invest in assessing potential mates), the direction of female preferences (i.e. which phenotypes females prefer) and preference strength (i.e. how strident females are in their preferences). We also consider potential coevolution between mate choice and the social environment arising because both include genes. Finally, we highlight some outstanding questions that help us more fully understand social environment impacts on female mating decisions and emphasize the need for tests of current theoretical predictions.

Conservation, like life, is inherently unfair. Some species, through accidents of phylogeny or ecology, are provided with all the public attention and resources needed to develop and implement comprehensive conservation or recovery plans. Other species are at best ignored or at worst deliberately sacrificed at the altar of charisma. Parasite species are at a three-fold disadvantage in this conservation lottery. First, most of them are invertebrates...

In social animals, success can depend on the outcome of group battles. Theoretical models of warfare predict that group fighting ability is proportional to two key factors: the strength of each soldier in the group and group size. The relative importance of these factors is predicted to vary across environments [F. W. Lanchester, Aircraft in Warfare, the Dawn of the Fourth Arm (1916)]. Here, we provide an empirical validation of the theoretical prediction that open environments should favor superior numbers, whereas complex environments should favor stronger soldiers [R. N. Franks, L. W. Partridge, Anim. Behav. 45 , 197–199 (1993)]. We first demonstrate this pattern using simulated battles between relatively strong and weak soldiers in a computer-driven algorithm. We then validate this result in real animals using an ant model system: In battles in which the number of strong native meat ant Iridomyrmex purpureus workers is constant while the number of weak non-native invasive Argentine ant Linepithema humile workers increases across treatments, fatalities of I. purpureus are lower in complex than in simple arenas. Our results provide controlled experimental evidence that investing in stronger soldiers is more effective in complex environments. This is a significant advance in the empirical study of nonhuman warfare and is important for understanding the competitive balance among native and non-native invasive ant species.

Conditional strategies occur when the relative fitness pay-off from expressing a given phenotype is contingent upon environmental circumstances. This conditional strategy model underlies cases of alternative reproductive tactics, in which individuals of one sex employ different means to obtain reproduction. How kin structure affects the expression of alternative reproductive tactics remains unexplored. We address this using the mite Rhizoglyphus echinopus, in which large males develop into aggressive 'fighters' and small males develop into non-aggressive 'scramblers.' Because only fighters kill their rivals, they should incur a greater indirect fitness cost when competing with their relatives, and thus fighter expression could be reduced in the presence of relatives. We raised mites in full-sibling or mixed-sibship groups and found that fighters were more common at higher body weights in full-sibling groups, not less common as we predicted (small individuals were almost exclusively scramblers in both treatments). This result could be explained if relatedness and cue variability are interpreted signals of population density, since fighters are more common at low densities in this species. Alternatively, our results may indicate that males compete more intensely with relatives in this species. We provide the first evidence of kin-mediated plasticity in the expression of alternative reproductive tactics.

Reduced streamflow because of climate change presents a major threat to aquatic biodiversity in arid, semi-arid and Mediterranean climatic regions. Freshwater mussels are particularly sensitive to this threat, because of their sedentary nature and limited mobility as juveniles or adults. The freshwater mussel Westralunio carteri, which is endemic to south-western Australia, has undergone a 49% reduction in range in the last 50 years, and a drying climate presents substantial extinction risk, as highlighted by two recent cases of mass mortality. Experimental studies found that mussels respond to water emersion by first tracking receding water levels, then burrowing. The amount of horizontal movement by mussels was not affected by size, but smaller mussels initiated burrowing sooner and were also more likely to be predated if they remained on the surface. Burrowing and shading both significantly reduced mortality rate and increased survival time when mussels were exposed to drying; when shaded or allowed to burrow, mussels could survive at least 62 days out of water. Predicted future reductions in streamflow are likely to increase the mortality rate in W. carteri, but it may be possible to partially avert the adverse effects of drying rivers by increasing riparian shading.

Males of many species harm females as a byproduct of intrasexual competition, but this harm can be reduced if males are less competitive in the presence of familiar relatives. We determined the cue males use to identify competitors in this context. We assessed genetic variance in a putative kin recognition trait (cuticular hydrocarbons) in male seed beetles Callosobruchus maculatus and found that five hydrocarbons had significant components of additive genetic variance and could serve as relatedness cues. Next, we tested whether hydrocarbons were the mechanism males use to distinguish the social identities of competitors when strategically adjusting their competitiveness/harmfulness. Pairs of female and male C. maculatus were mated in the presence of hydrocarbons extracted from males that differed in their relatedness and familiarity to the focal male. Females were more productive after mating in the presence of extracts from the focal male's nonrelatives, if those extracts were also unfamiliar to the focal male. Relatedness had no effect on productivity when extracts were familiar to the focal male. These results may be reconciled with those of previous studies that manipulated the relatedness and familiarity of competing males if the difference between the effect of harmfulness on productivity following a single mating and the effect on lifetime reproductive fitness after multiple matings is accounted for. This study provides a novel demonstration of the mechanism of social recognition in the moderation of sexual conflict.

The outcome of sexual conflict can depend on the social environment, as males respond to changes in the inclusive fitness payoffs of harmfulness and harm females less when they compete with familiar relatives. Theoretical models also predict that if limited male dispersal predictably enhances local relatedness while maintaining global competition, kin selection can produce evolutionary divergences in male harmfulness among populations. Experimental tests of these predictions, however, are rare. We assessed rates of dispersal in female and male seed beetles Callosobruchus maculatus, a model species for studies of sexual conflict, in an experimental setting. Females dispersed significantly more often than males, but dispersing males travelled just as far as dispersing females. Next, we used experimental evolution to test whether limiting dispersal allowed the action of kin selection to affect divergence in male harmfulness and female resistance. Populations of C. maculatus were evolved for 20 and 25 generations under one of three dispersal regimens: completely free dispersal, limited dispersal and no dispersal. There was no divergence among treatments in female reproductive tract scarring, ejaculate size, mating behaviour, fitness of experimental females mated to stock males or fitness of stock females mated to experimental males. We suggest that this is likely due to insufficient strength of kin selection rather than a lack of genetic variation or time for selection. Limited dispersal alone is therefore not sufficient for kin selection to reduce male harmfulness in this species, consistent with general predictions that limited dispersal will only allow kin selection if local relatedness is independent of the intensity of competition among kin.

Although freshwater ecosystems are among the most diverse and endangered in the world, little attention has been paid to either the importance of parasitic disease as a threatening process for freshwater organisms, or the co-extinction risk of freshwater parasites. In this review, we use theoretical and empirical studies of host/parasite interactions to examine these issues, particularly with respect to the threat posed by climate change to fish and parasite communities in intermittent rivers. Intermittent rivers are those that cease to flow at any point in time or space, with isolated pools providing ecological refuges for freshwater biota between streamflow events. Intermittent rivers are the dominant river type in arid, semi-arid and Mediterranean regions; areas of the world that have experienced dramatic decreases in streamflow as a result of climate change. Reduced streamflow decreases the number, size and connectivity of refuge pools in intermittent rivers, with important consequences for free-living aquatic organisms, particularly fishes, and their parasitic fauna. As a result of more frequent and sustained periods of no flow, parasite diversity within refuge pools is expected to decrease, with a concomitant increase in the prevalence and intensity of those parasite species which do survive, particularly host generalists. Decreased connectivity between refuge pool communities should increase the spatial modularity of host/parasite interactions, leading to a greater structuring of host and parasite communities along the river. This increases the probability of species loss (for both hosts and their parasites), as local extinctions cannot be reversed by colonisation from other localities.

Males of many species adjust their reproductive investment to the number of rivals present simultaneously. However, few studies have investigated whether males sum previous encounters with rivals, and the total level of competition has never been explicitly separated from social familiarity. Social familiarity can be an important component of kin recognition and has been suggested as a cue that males use to avoid harming females when competing with relatives. Previous work has succeeded in independently manipulating social familiarity and relatedness among rivals, but experimental manipulations of familiarity are confounded with manipulations of the total number of rivals that males encounter. Using the seed beetle Callosobruchus maculatus, we manipulated three factors: familiarity among rival males, the maximum number of rivals encountered simultaneously and the total number of rivals encountered over a 48 h period. Males produced smaller ejaculates when exposed to more rivals in total, regardless of the maximum number of rivals they encountered simultaneously. Males did not respond to familiarity. Our results demonstrate that males of this species can sum the number of rivals encountered over separate days, and therefore the confounding of familiarity with the total level of competition in previous studies should not be ignored.

Recent interest has focused on the role of reactive oxygen species (ROS) as universal constraints in life-history evolution. Empirical studies have examined the oxidative costs of reproduction for females, with little work conducted on males. The male germline is thought to be particularly susceptible to oxidative damage because the testes, and the sperm themselves, can be prolific producers of ROS. We tested the hypothesis that protection of the male germline from oxidative damage represents a cost of reproduction for males. We fed male crickets, Teleoglyilus oceanicus, with one of two experimental diets in which we manipulated the availability of dietary antioxidants, and we induced variation in their expenditure on courtship effort by manipulating access to females. We measured the total antioxidant capacity, levels of ROS production and the amount of oxidative damage to proteins in both testis and thoracic muscle tissues. Dietary antioxidants contributed to positive oxidative balance in both tissue types. Although the testes had greater antioxidant defences than muscle tissue, they also produced considerably higher levels of ROS and sustained higher levels of oxidative damage. Courtship effort had no impact on any measure of oxidative balance. Our data confirm that the male germline is especially susceptible to oxidative stress and that dietary antioxidants can alleviate this oxidative cost of reproduction.