To harass or to respect: the economy of male persistence despite female refusal in a damselfly with scramble mate competition

In sexual conflict, males are often thought to gain fitness benefits from harassing females over mating. Yet when harassment itself incurs costs to males and if alternative, receptive females are available in a local population, theory predicts that when confronted with a female refusal, a male’s choice of persisting or retreating is determined in part by the likelihood of achieving a mating. We tested that prediction in the damselfly Enallagma hageni, whose males compete by intense scramble competition, resulting in widespread mating harassment toward females, which have a high level of control over mating. Using captive individuals of E. hageni in outdoor insectaries, we quantified male persistence in mating after refusals by pre- and post-oviposition focal females whose egg content we quantified after observations. We documented a novel, context-dependent head-turning refusal signal of sexual non-receptivity, most often displayed in tandem pairs by post-oviposition females that typically carried few mature eggs for males to fertilize. Male persistence was less likely to result in mating with post-oviposition females compared with pre-oviposition females carrying a clutch of mature eggs. Accordingly, males were less likely to persist following refusal signals given by post-oviposition females, supporting the theoretical prediction. Compared with a refusal signal known as wing spread, head-turning was significantly more effective in deterring harassing males. Our results suggest that despite on-going sexual conflict over mating, cooperation benefits both sexes when females use the honest signal of non-receptivity because they carry few mature eggs that males could fertilize.

Increased opposite sex association is linked with fitness benefits, otherwise sociality is subject to stabilising selection in a wild passerine

Animal sociality, an individuals propensity to association with others, has consequences for fitness, and particularly mate choice. For example, directly, by increasing the pool of prospective partners, and indirectly through increased survival. Individuals benefit from both over the short-term as these benefits are associated with mating status and subsequent fecundity, but whether animal sociality also translates into fitness is unknown. Here, we quantified social associations and their link with annual and lifetime fitness, measured as the number of recruits and in de-lifed fitness. We measured this in birds visiting a feeding station over two non-breeding periods, using social network analysis and a multi-generational genetic pedigree. We find high individual repeatability in sociality. We found that individuals with an average sociality had the highest fitness, and that birds with more opposite-sex associates had higher fitness, but this did not translate to improved lifetime fitness. For lifetime fitness, we found evidence for stabilizing selection on between sex sociality measures, suggesting that such benefits are only short-lived in a wild population.

Evolution of Social Learning with Payoff and Content Bias

There has been much theoretical work aimed at understanding the evolution of social learning; and in most of it, individual and social learning are treated as distinct processes. A number of authors have argued that this approach is faulty because the same psychological mechanisms underpin social and individual learning. In previous work, we analyzed a simple model in which both individual and social learning are the result of a single learning process. Here, we extend this approach by showing how payoff and content biases evolve. We show that payoff bias leads to higher average fitness when environments are noisy and change rapidly. Content bias always evolves when the expected fitness benefits of alternative traits differ.

When does cultural evolution become cumulative culture? A case study of humpback whale song

Culture presents a second inheritance system by which innovations can be transmitted between generations and among individuals. Some vocal behaviours present compelling examples of cultural evolution. Where modifications accumulate over time, such a process can become cumulative cultural evolution. The existence of cumulative cultural evolution in non-human animals is controversial. When physical products of such a process do not exist, modifications may not be clearly visible over time. Here, we investigate whether the constantly evolving songs of humpback whales ( Megaptera novaeangliae ) are indicative of cumulative cultural evolution. Using nine years of song data recorded from the New Caledonian humpback whale population, we quantified song evolution and complexity, and formally evaluated this process in light of criteria for cumulative cultural evolution. Song accumulates changes shown by an increase in complexity, but this process is punctuated by rapid loss of song material. While such changes tentatively satisfy the core criteria for cumulative cultural evolution, this claim hinges on the assumption that novel songs are preferred by females. While parsimonious, until such time as studies can link fitness benefits (reproductive success) to individual singers, any claims that humpback whale song evolution represents a form of cumulative cultural evolution may remain open to interpretation. This article is part of a discussion meeting issue ‘The emergence of collective knowledge and cumulative culture in animals, humans and machines’.

Fluid Nest Membership Drives Variable Relatedness in Groups of a Facultatively Social Bee

Kin selection theory has dominated our understanding of the evolution of group living. However, many animal groups form among non-relatives, which gain no indirect fitness benefits from cooperating with nestmates. In this study, we characterized the relatedness and inter-nest migration behavior of the facultatively social carpenter bee, Xylocopa sonorina. Nesting constraints due to costly nest construction in this species give rise to intense intraspecific competition over access to existing nests. We used mark-recapture techniques to characterize patterns of dispersal and nest relocation within a nesting aggregation of spatially clustered nests. Two-thirds of bees relocated at least once during the reproductive season, likely to seek reproductive opportunities in another nest. This fluid nest membership creates opportunities for association among non-relatives. To assess the effects of this dynamic nesting behavior on group relatedness, we used microsatellite analysis to estimate relative relatedness within and between nests in the aggregation. We found that relatedness was variable across sampling years, but that in many cases nestmates were no more related to one another than they were to non-nestmate bees in the population. Together, these results suggest that group composition in X. sonorina may result from strategies to maximize direct fitness. This study supports the hypothesis that factors beyond kinship, such as ecological constraints, are likely to drive group formation in this species.

Vegetative and Fecundity Fitness Benefit Found in a Glyphosate-Resistant Eleusine indica Population Caused by 5-Enolpyruvylshikimate-3-Phosphate Synthase Overexpression

Fitness is an important trait in weed species that have developed herbicide resistance, including resistance to the popular herbicide glyphosate. Fitness cost is commonly found in weeds with glyphosate resistance, which is caused by target-site mutations. In this study, the vegetative and fecundity fitness traits in a glyphosate-resistant (GR) Eleusine indica population caused by 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) overexpression were investigated under glyphosate-free conditions. The results showed that the resistance index of the population resistant (R) to glyphosate compared with that of the population susceptible (WT) to it was approximately 4.0. Furthermore, EPSPS expression level in the R plants was 20.1–82.7 times higher than that in the WT plants. The dry weight of the R population was significantly higher than that of the WT population at the later growth stage after planting; a similar trend was observed for leaf area. In addition, seed production in the R population was 1.4 times higher than that in the WT population. The R and WT populations showed similar maximum germination rates and T50 values. UPLC-MS/MS was performed for the metabolic extracts prepared from the leaves of R and WT populations to address changes in the metabolome. A total of 121 differential metabolites were identified between R and WT individuals. The levels of 6-hydroxy-1H-indole-3-acetamide and indole acetaldehyde, which are associated with auxin synthesis, were significantly higher in plants of the R population than in those of the WT population. However, some secondary metabolite levels were slightly lower in the R population than in the WT population. To conclude, in this study, vegetative and fecundity fitness benefits were found in the GR E. indica population. The results of metabolome analysis indicate that the increase in 6-hydroxy-1H-indole-3-acetamide and indole acetaldehyde levels may be the result of fitness benefit. Further studies should be conducted to confirm the functions of these metabolites.

Environmental quality mediates the ecological dominance of cooperatively breeding birds

Species as diverse as humans and ants are among the most abundant organisms on Earth, partly because of their ability to form cooperative societies1-3. Yet, animals form groups for many reasons4,5, and how these differences affect their ‘social conquests’2 remains unknown. Here we use a theoretical model to demonstrate that the different fitness benefits that animals receive by forming groups4,6 depend on the quality of their environment, which in turn impacts their ecological dominance and resilience to global change. Our model predicts species that group because of environmental hardships will have wider ecological niches, larger geographic ranges, and higher abundances than non-social species, whereas those that group because of intraspecific resource competition will not. As predicted, an analysis of >1500 avian species finds that cooperative breeders occurring in harsh and fluctuating environments have larger ranges and higher abundances than non-cooperative breeders, whereas cooperative breeders occurring in benign and stable environments do not. These results are consistent with our model predictions showing that species cooperating in harsh or fluctuating environments will be less vulnerable to climate change than non-social species and those cooperating against intra-specific competitors in benign or stable environments. Ultimately, by combining macroecological and sociobiological perspectives, our study helps understand and predict the past, present, and future state of social species, including our own.

Sex-Specific Routes to Independent Breeding in a Polygynous Cooperative Breeder

How can individuals obtain a breeding position and what are the benefits associated with philopatry compared to dispersal? These questions are particularly intriguing in polygamous cooperative breeders, where dispersal strategies reflect major life history decisions, and routes to independent breeding may utterly differ between the sexes. We scrutinized sex-dependent life-history routes by investigating dispersal patterns, growth rates and mortality in a wild colony of the cooperatively breeding cichlid Neolamprologus savoryi. Our data reveal that female helpers typically obtain dominant breeding positions immediately after reaching sexual maturity, which is associated with strongly reduced growth. In contrast, males obtain breeder status only at twice the age of females. After reaching sexual maturity, males follow one of two strategies: (i) they may retain their subordinate status within the harem of a dominant male, which may provide protection against predators but involves costs by helping in territory maintenance, defence and brood care; or (ii) they may disperse and adopt a solitary status, which diminishes survival chances and apparently reflects a best-of-a-bad-job strategy, as there are no obvious compensating future fitness benefits associated with this pathway. Our study illustrates that sex-dependent life history strategies strongly relate to specific social structures and mating patterns, with important implications for growth rates, the age at which breeding status is obtained, and survival.