Emily Burdfield-Steel

Experimental reduction of intromittent organ length reduces male reproductive success in a bug

It is now clear in many species that male and female genital evolution has been shaped by sexual selection. However, it has historically been difficult to confirm correlations between morphology and fitness, as genital traits are complex and manipulation tends to impair function significantly. In this study, we investigate the functional morphology of the elongate male intromittent organ (or processus) of the seed bug Lygaeus simulans, in two ways. We first use micro-computed tomography (micro-CT) and flash-freezing to reconstruct in high resolution the interaction between the male intromittent organ and the female internal reproductive anatomy during mating. We successfully trace the path of the male processus inside the female reproductive tract. We then confirm that male processus length influences sperm transfer by experimental ablation and show that males with shortened processi have significantly reduced post-copulatory reproductive success. Importantly, male insemination function is not affected by this manipulation per se. We thus present rare, direct experimental evidence that an internal genital trait functions to increase reproductive success and show that, with appropriate staining, micro-CT is an excellent tool for investigating the functional morphology of insect genitalia during copulation.

The extent and costs of reproductive interference among four species of true bug

Reproductive interference arises when individuals of one species engage in reproductive activities with individuals of another, leading to fitness costs in one or both species. Reproductive interference (RI) therefore has two components. First, there must be mis-directed mating interactions. Second, there must be costs associated with these mis-directed interactions. Here we consider RI between four species of true bug in the family Lygaeidae, focusing in particular on the fitness consequences to Lygaeus equestris. The species we consider vary in their relationships with each other, including species in the same or different genus, and with or without natural overlap in their geographic ranges. First we show that inter-specific mating interactions, although not a certain outcome, are common enough to perhaps influence mating behaviour in these species (arising in up to 10xa0% of inter-specific pairings). Second, we show that reproductive interference can seriously reduce female fitness in L. equestris. Importantly, different species impose different costs of RI on L. equestris, with interactions with male Spilostethus pandurus inflicting fitness costs of similar magnitude to the costs of mating with con-specifics. On the other hand, mating interactions with male Oncopeltus fasciatus appear to have no effect on female fitness. In a follow-up experiment, when we allowed competition amongst just females of S. pandurus and L. equestris, the fitness of the latter was not reduced, arguing more strongly for the role of reproductive interference. However, in our final experiments under mass mating conditions with extended ecological interactions (including scope for competition for resources and cannibalism), the costs of RI were less apparent. Our data therefore suggest that the costs of RI will be context-specific and may act in concert with, or be swamped by, other ecological effects. We suggest that comparative studies of this sort that both mimic naturally occurring reproductive interference events, and also artificially generate new ones, will be necessary if we are to better understand the ecological and evolutionary significance of reproductive interference.

How to fight multiple enemies: target-specific chemical defences in an aposematic moth

Animals have evolved different defensive strategies to survive predation, among which chemical defences are particularly widespread and diverse. Here we investigate the function of chemical defence diversity, hypothesizing that such diversity has evolved as a response to multiple enemies. The aposematic wood tiger moth (Arctia plantaginis) displays conspicuous hindwing coloration and secretes distinct defensive fluids from its thoracic glands and abdomen. We presented the two defensive fluids from laboratory-reared moths to two biologically relevant predators, birds and ants, and measured their reaction in controlled bioassays (no information on colour was provided). We found that defensive fluids are target-specific: thoracic fluids, and particularly 2-sec-butyl-3-methoxypyrazine, which they contain, deterred birds, but caused no aversive response in ants. By contrast, abdominal fluids were particularly deterrent to ants, while birds did not find them repellent. Our study, to our knowledge, is the first to show evidence of a single species producing separate chemical defences targeted to different predator types, highlighting the importance of taking into account complex predator communities in studies on the evolution of prey defence diversity.

Variation in social and sexual behaviour in four species of aposematic seed bugs (Hemiptera: Lygaeidae): The role of toxic and non-toxic food

Understanding variation in social behaviour both within and among species continues to be a challenge. Evolutionary or ecological theory typically predicts the optimal behaviour for an animal under a given set of circumstances, yet the real world presents much greater variation in behaviour than predicted. This variation is apparent in many social and sexual interactions, including mate choice, and has led to a renewed focus on individual variation in behaviour. Here we explore within and among species variation in social behaviour in four species of aposematic seed bug (Lygaeidae: Hemiptera). These species are Müllerian mimics, with characteristic warning colouration advertising their chemical toxicity. We examine the role of diet in generating variation in two key behaviours: social aggregation of nymphs and mate choice. We test how behaviour varies with exposure to either milkweed (a source of defensive compounds) or sunflower (that provides no defence). We show that although the four species vary in their food preferences, and diet influences their life-history (as highlighted by body size), social aggregation and mate choice is relatively unaffected by diet. We discuss our findings in terms of the evolution of aposematism, the importance of automimicry, and the role of diet in generating behavioural variation.

Sexual stereotypes: The case of sexual cannibalism

There is a long-standing debate within the field of sexual selection regarding the potential projection of stereotypical sex roles onto animals by researchers. It has been argued that this anthropomorphic view may be hampering research in this area, for example by prioritizing the study of male sexual adaptations over female ones. We investigated how males and females are described in the sexual cannibalism literature. Sexual cannibalism is a specific form of sexual conflict and is highly gendered, with females generally cannibalizing males. We found that females were more likely to be described using active words and males with reactive words. This is contrary to recent results from a survey of the sexual conflict literature. While this reversed gender bias may arise from the nature of sexual cannibalism, our results nevertheless indicate an alternative form of sexual stereotyping. A number of the words used to describe cannibalistic females were highly loaded and suggestive of a negative stereotype of sexually aggressive females. To make progress we suggest first that animal behaviour researchers recognize both the costs and benefits of looking for general patterns as part of the scientific method. Although necessary, the search for general patterns may validate existing stereotypes or provide the basis for new ones. Additionally, we suggest that the field of sexual behaviour research is neither wholly bad nor good in terms of language use but that we should work towards a consensus of how and when we use particular terms to describe sexual behaviour.

Do the benefits of polyandry scale with outbreeding

Lay Summary Mating can be both costly and dangerous. Despite this, females of many species typically mate more than once and with different males. We found that female seed bugs only benefit from mating with more than 1 male if one of her mates was a different species. Females needed only 1 mating with her own species to maximize her fitness. Thus, multiple mating may be adaptive in areas where the 2 species coexist.

Mate-guarding in a promiscuous insect: species discrimination influences context-dependent behaviour

Mating strategy is often informed by social context. However, information on social environment may be sensitive to interference by nearby heterospecifics, a process known as reproductive interference (RI). When heterospecific individuals are present in the environment, failures in species discrimination can lead to sub-optimal mating behaviours, such as misplaced courtship, misplaced rivalry behaviours, or heterospecific copulation attempts. All aspects of mating behaviour that are influenced by social context may be prone to RI, including copulatory behaviours associated with mate-guarding in the presence of possible competitors. Here we investigate the effect of three heterospecifics on the mate-guarding behaviour of male Lygaeus equestris seed bugs. We find that, despite previously reported heterospecific mating harassment amongst these species of lygaeid bug, male L. equestris are able to effectively distinguish rival conspecific males from heterospecifics. Thus, heterospecific mating attempts in this group may reflect selection on males to mate opportunistically, rather than a failure of species discrimination.

The evolutionary ecology of the Lygaeidae

The Lygaeidae (sensu lato) are a highly successful family of true bugs found worldwide, yet many aspects of their ecology and evolution remain obscure or unknown. While a few species have attracted considerable attention as model species for the study of insect physiology, it is only relatively recently that biologists have begun to explore aspects of their behavior, life history evolution, and patterns of intra- and interspecific ecological interactions across more species. As a result though, a range of new phenotypes and opportunities for addressing current questions in evolutionary ecology has been uncovered. For example, researchers have revealed hitherto unexpectedly rich patterns of bacterial symbiosis, begun to explore the evolutionary function of the familys complex genitalia, and also found evidence of parthenogenesis. Here we review our current understanding of the biology and ecology of the group as a whole, focusing on several of the best-studied characteristics of the group, including aposematism (i.e., the evolution of warning coloration), chemical communication, sexual selection (especially, postcopulatory sexual selection), sexual conflict, and patterns of host-endosymbiont coevolution. Importantly, many of these aspects of lygaeid biology are likely to interact, offering new avenues for research, for instance into how the evolution of aposematism influences sexual selection. With the growing availability of genomic tools for previously “non-model” organisms, combined with the relative ease of keeping many of the polyphagous species in the laboratory, we argue that these bugs offer many opportunities for behavioral and evolutionary ecologists.

Reproductive interference in insects: Reproductive interference in insects

1. Reproductive interference occurs when members of different species engage in reproductive interactions, leading to a fitness cost to one or both actors.

De novo Synthesis of Chemical Defenses in an Aposematic Moth

Abstract Many animals protect themselves from predation with chemicals, both self-made or sequestered from their diet. The potential drivers of the diversity of these chemicals have been long studied, but our knowledge of these chemicals and their acquisition mode is heavily based on specialist herbivores that sequester their defenses. The wood tiger moth (Arctia plantaginis, Linnaeus, 1758) is a well-studied aposematic species, but the nature of its chemical defenses has not been fully described . Here, we report the presence of two methoxypyrazines, 2-sec-butyl-3-methoxypyrazine and 2-isobutyl-3-methoxypyrazine, in the moths’ defensive secretions. By raising larvae on an artificial diet, we confirm, for the first time, that their defensive compounds are produced de novo rather than sequestered from their diet. Pyrazines are known for their defensive function in invertebrates due to their distinctive odor, inducing aversion and facilitating predator learning. While their synthesis has been suspected, it has never previously been experimentally confirmed. Our results highlight the importance of considering de novo synthesis, in addition to sequestration, when studying the defensive capabilities of insects and other invertebrates.