Comparative evidence supports a role for reproductive allocation in the evolution of female ornament diversity

1. Sexually selected ornaments are highly variable, even among closely related species, and the ultimate causes of variation in ornament evolution are unclear, including in rare cases of female ornament expression. One hypothesis is that differences across species in female reproductive allocation may help to explain patterns of female ornament expression among insects with nuptial gifts. 2. Dance flies (Diptera: Empididae: Empidinae) vary considerably among species in the presence and extravagance of female ornaments, which probably evolved through female contests for mates. In most dance flies, adult females appear to acquire all their dietary protein from nuptial gifts provided by males during mating. The importance of nuptial feeding on egg development is not yet known. 3. To test the prediction that the presence of female ornaments reflects differences in the degree to which females rely on nuptial feeding for egg development, egg development was examined in wild females of two species, one ornamented and the other unornamented. An ageing technique based on cuticular bands was validated, which permitted a regression of egg size on adult age. 4. We found that egg development depended on mating status in the ornamented species alone, meaning the eggs of unmated females of the ornamented species did not develop. This contrast across species is consistent with expectations that females of different species vary in their dependence on nuptial gifts for egg development. 5. These findings provide preliminary support for the hypothesis that differences in reproductive allocation mediate the intensity of female contests for nuptial gifts

Sexual selection on multiple female ornaments in dance flies

Sex-specific ornaments typically occur in males, but they can also develop in females. While there are several models concerning the evolution of male-specific ornaments, it is not clear how, or under what circumstances, those models apply to female-specific ornament evolution. Here, we present a manipulative field experiment that explores the theoretical ‘trait space’ of multiple female-specific ornaments to study how these unusual traits evolved. We measured the attractiveness of two female-specific ornaments (pinnate leg scales and inflatable abdominal sacs) in the dance fly Rhamphomyia longicauda in a wild mating swarm. We found significant directional preferences for larger ornaments of both types; however, variation in one of the ornaments (abdominal sacs) was almost three times more effective at improving attractiveness. The abdominal ornament was consistently effective in increasing attractiveness to males regardless of leg ornament expression, while leg ornament size was only effective if abdominal ornaments were very small. These results are consistent with predictions from a sexual conflict model of ornament expression in supporting the probable role of deception in the evolution of female-specific ornaments among dance flies. Sexual conflict can be an important force in generating elaborate sex-specific ornaments in females as well as males

Oecanthus nigricornis (Orthoptera: Gryllidae) as the first known host of Stylogaster neglecta (Diptera: Conopidae)

The conopid fly Stylogaster neglecta Williston (Diptera: Conopidae) is a parasitoid with no known host. We report this species as the first recorded dipteran parasitoid of Oecanthus nigricornis Walker (Orthoptera: Gryllidae) (black-horned tree crickets). We reared field-collected O. nigricornis juveniles over several months in 2017 and found that larval S. neglecta emerged from them during late July into August. We estimated the incubation period for S. neglecta larvae to be around 30 days based on the length of time it took for them to emerge from the host and pupate (subsequently all hosts died). We documented several cases of multiple parasitism. In 2018, we dissected O. nigricornis sampled from four sites across southern Ontario, Canada and upstate New York, United States of America and found that the percentage of juvenile O. nigricornis parasitised ranged 2–39%. Further sampling will be necessary to determine whether this variation represents consistent population differences or between-year variation in parasitism

Sperm competition in yellow dung flies: No consistent effect of sperm size

The male competition for fertilization that results from female multiple mating promotes the evolution of increased sperm numbers and can impact sperm morphology, with theory predicting that longer sperm can at times be advantageous during sperm competition. If so, males with longer sperm should sire more offspring than competitors with shorter sperm. Few studies have directly tested this prediction, and findings are inconsistent. Here we assessed whether longer sperm provide a competitive advantage in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae). Initially, we let brothers with different temperature-mediated mean sperm lengths compete – thus minimizing confounding effects of genetic background – and found no clear advantage of longer sperm. We then used flies from lines subjected to bidirectional selection on phenoloxidase activity that had shown correlated evolutionary responses in sperm and female spermathecal duct lengths. This experiment also yielded no main effect of sperm size on siring success. Instead, there was a trend for a shorter-sperm advantage, but only when competing in females with longer spermathecal ducts. Our data corroborated many previously reported findings (last-male precedence, effects of copula duration and body size), suggesting our failure to find sperm size effects is not inherently due to our experimental protocols. We conclude that longer sperm are not competitively superior in yellow dung flies under most circumstances, and that, consistent with previous work, in this species competitive fertilization success is primarily determined by the relative numbers of sperm competing

Competition for access to mates predicts female-specific ornamentation and male investment in relative testis size

Sexually selected ornaments are highly variable and the factors that drive variation in ornament expression are not always clear. Rare instances of female‐specific ornament evolution (such as in some dance fly species) are particularly puzzling. While some evidence suggests that such rare instances represent straightforward reversals of sexual selection intensity, the distinct nature of trade‐offs between ornaments and offspring pose special constraints in females. To examine whether competition for access to mates generally favours heightened ornament expression, we built a phylogeny and conducted a comparative analysis of Empidinae dance fly taxa that display female‐specific ornaments. We show that species with more female‐biased operational sex ratios in lek‐like mating swarms have greater female ornamentation, and in taxa with more ornate females, male relative testis investment is increased. These findings support the hypothesis that ornament diversity in dance flies depends on female receptivity to mates, which is associated with contests for nutritious nuptial gifts provided by males. Moreover, our results suggest that increases in female receptivity lead to higher levels of sperm competition among males. The incidence of both heightened pre‐mating sexual selection on females and post‐mating selection on males contradicts assertions that sex‐roles are straightforwardly reversed in dance flies

The impact of uncertainty on cooperation intent in a conservation conflict

1. Stakeholder cooperation can be vital in managing conservation conflicts. Laboratory experiments show cooperation is less likely in the presence of uncertainty. Much less is known about how stakeholders in real‐life conservation conflicts respond to different types of uncertainty. 2. We tested the effects of different sources of uncertainty on cooperative behaviour using a framed field experiment and interviews. The experiment compared a baseline scenario of perfect certainty with scenarios including either: (a) scientific uncertainty about the effectiveness of a conflict‐reduction intervention; (b) administrative uncertainty about intervention funding; or (c) political uncertainty about the extent of community support. We applied these scenarios to a conservation conflict in the Outer Hebrides, Scotland, involving the management of geese to simultaneously meet both conservation and farming objectives. We asked 149 crofters (small‐scale farmers) if they would commit to cooperate with others by helping fund a goose management plan given the three sources of uncertainty. 3. On average, intention to cooperate was highest (99%) in scenarios without uncertainty, and lowest under administrative uncertainty (77%). Scientific uncertainty and political uncertainty both had less of an effect, with over 95% of crofters predicted to be willing to cooperate in these scenarios. Crofters who indicated concern for other crofters suffering the impact of geese were more likely to cooperate. The longer an individual had been a crofter, the less likely they were to cooperate. 4. Synthesis and applications. Crofters’ intention to cooperate is high but lessened by uncertainty, especially over the commitment from other stakeholders such as government, to cooperate on goose management. Existing cooperation on goose management may be at risk if uncertainty is not reduced outright or commitments between parties are not strengthened. This has wide applicability, supporting the need for researchers and government advisers to: (a) determine how uncertainty will impact intention of stakeholders to cooperate; and (b) take steps (such as uncertainty reduction, communication or acceptance) to reduce the negative impact of uncertainty on cooperation

Growth rate mediates hidden developmental plasticity of female yellow dung fly reproductive morphology in response to environmental stressors

Understanding how environmental variation influences even cryptic traits is important to clarify the roles of selection and developmental constraints in past evolutionary divergence and to predict future adaptation under environmental change. Female yellow dung flies (Scathophaga stercoraria) typically have three sperm storage compartments (3S), but occasionally four (4S). More spermathecae are thought to be a female adaptation facilitating sperm sorting after mating, but the phenotype is very rare in nature. We manipulated the flies' developmental environment by food restriction, pesticides, and hot temperatures to investigate the nature and extent of developmental plasticity of this trait, and whether spermatheca expression correlates with measures of performance and developmental stability, as would be expected if 4S expression is a developmental aberration. The spermathecal polymorphism of yellow dung fly females is heritable, but also highly developmentally plastic, varying strongly with rearing conditions. 4S expression is tightly linked to growth rate, and weakly positively correlated with fluctuating asymmetry of wings and legs, suggesting that the production of a fourth spermatheca could be a nonadaptive developmental aberration. However, spermathecal plasticity is opposite in the closely related and ecologically similar Scathophaga suilla, demonstrating that overexpression of spermathecae under developmental stress is not universal. At the same time, we found overall mortality costs as well as benefits of 4S pheno- and genotypes (also affecting male siblings), suggesting that a life history trade-off may potentially moderate 4S expression. We conclude that the release of cryptic genetic variation in spermatheca number in the face of strong environmental variation may expose hidden traits (here reproductive morphology) to natural selection (here under climate warming or food augmentation). Once exposed, hidden traits can potentially undergo rapid genetic assimilation, even in cases when trait changes are first triggered by random errors that destabilize developmental processes

Senescence of the cellular immune response in Drosophila melanogaster

Immune system effectiveness generally declines as animals age, compromising disease resistance. In Drosophila, expression of a variety of immune-related genes elevates during ageing; however how this is linked to increasing pathogen susceptibility in older flies has remained unclear. We investigated whether changes in the Drosophila cellular immune response might contribute to immunosenescence. Experiments studied fly cohorts of different ages and compared the numbers and activity of the circulating haemocytes involved in pathogen defence. In female wildtype Samarkand and Oregon R flies the haemocyte population fell by 31.8% and 10.2% respectively during the first four weeks of adulthood. Interestingly we detected no such decline in male flies. The impact of ageing on the phagocytic activity of haemocytes was investigated by injecting flies with fluorescently labelled microbes or latex beads and assessing the ability of haemocytes to engulf them. For all immune challenges the proportion of actively phagocytosing haemocytes decreased as flies aged. Whilst 24.3% ± 1.15% of haemocytes in one-week-old flies phagocytosed Escherichia coli bacteria or Beauveria bassiana fungal spores, this decreased to 16.7% ± 0.99% in four-week-old flies. This clear senescence of the Drosophila cellular immune response may underpin increased disease susceptibility in older flies

The Effect of Diet Quality and Wing Morph on Male and Female Reproductive Investment in a Nuptial Feeding Ground Cricket

A common approach in the study of life-history trade-off evolution is to manipulate the nutrient content of diets during the life of an individual in order observe how the acquisition of resources influences the relationship between reproduction, lifespan and other life-history parameters such as dispersal. Here, we manipulate the quality of diet that replicate laboratory populations received as a thorough test of how diet quality influences the life-history trade-offs associated with reproductive investment in a nuptial feeding Australian ground cricket (Pteronemobius sp.). In this species, both males and females make significant contributions to the production of offspring, as males provide a nuptial gift by allowing females to chew on a modified tibial spur during copulation and feed directing on their haemolymph. Individuals also have two distinct wing morphs, a short-winged flightless morph and a long-winged morph that has the ability to disperse. By manipulating the quality of diet over seven generations, we found that the reproductive investment of males and females were affected differently by the diet quality treatment and wing morph of the individual. We discuss the broader implications of these findings including the differences in how males and females balance current and future reproductive effort in nuptial feeding insects, the changing nature of sexual selection when diets vary, and how the life-history trade-offs associated with the ability to disperse are expected to differ among populations

Mate choice for genetic quality when environments vary: suggestions for empirical progress

Mate choice for good-genes remains one of the most controversial evolutionary processes ever proposed. This is partly because strong directional choice should theoretically deplete the genetic variation that explains the evolution of this type of female mating preferences (the so-called lek paradox). Moreover, good-genes benefits are generally assumed to be too small to outweigh opposing direct selection on females. Here, we review recent progress in the study of mate choice for genetic quality, focussing particularly on the potential for genotype by environment interactions (GEIs) to rescue additive genetic variation for quality, and thereby resolve the lek paradox. We raise five questions that we think will stimulate empirical progress in this field, and suggest directions for research in each area: 1) How is condition-dependence affected by environmental variation? 2) How important are GEIs for maintaining additive genetic variance in condition? 3) How much do GEIs reduce the signalling value of male condition? 4) How does GEI affect the multivariate version of the lek paradox? 5) Have mating biases for high-condition males evolved because of indirect benefits