Contrasting sexual selection on males and females in a role-reversed swarming dance-fly, Rhamphomyia longicauda Loew (Diptera: Empididae)

Sex-specific ornamentation is widely known among male animals, but even among sex-role reversed species, ornamented females are rare. Although several hypotheses for this pattern exist, too few systems featuring female ornaments have been studied in detail to adequately test them. Empidine dance flies are exceptional in that many species show female ornamentation of wings, abdomens, or legs. Here we compare sexual selection in males and females of the long-tailed dance fly, Rhamphomyia longicauda Loew (Diptera: Empididae), a sex-role reversed fly in which swarming females aggregate in competition for the nuptial gifts provided by males during mating. Females in this species possess several secondary sex characters, including eversible abdominal sacs, enlarged wings, and decorated tibiae that may all function in mate attraction during swarming. Males preferentially approach large females in the swarm, but the strength and shape of selection on females and the degree to which selection is sex-specific are unknown. We estimated linear and nonlinear sexual selection on structures expressed in both male and female flies, and found contrasting patterns of sexual selection on wing length and tibia length in males and females. In females, long wings and short tibiae were associated with mating success, whereas selection on males was significantly different: males with short wings and long tibiae were most likely to mate (although tibia length was a marginally non-significant predictor of male mating success). We found no evidence for assortative or disassortative mating. Although the largest females occupied positions within the swarm closest to the entry point for choosy males, in contrast to selection for mating success these females tended to have larger tibiae than rivals. We discuss our findings in the context of the mating biology of R. longicauda compared to other empidine dance flies, and its relevance to the evolution of sexual dimorphism in general

Multiple differences in calling songs and other traits between solitary and gregarious Mormon crickets from allopatric mtDNA clades

The Linnaean Society, The British Ecological Society, The Russell Trust (St. Andrews), The Orthopterists' Society and an NSERC (Canada) grant to DTG. Gordon S. Brown provided assistance with digital imaging and morphological measurements.Background: In acoustic species, traits such as male calling song are likely to diverge quickly between allopatric populations due to sexual selection, and divergence in parameters such as carrier frequency, chirp structure, and other important song characters can influence sexual isolation. Here we make use of two forms of Mormon crickets to examine differences in a broad suite of traits that have the potential to influence speciation via sexual isolation. Mormon crickets in "gregarious" populations aggregate into dense migratory bands, and females are the sexually competitive Fsex ( sex- role reversal). There is also a non-outbreak "solitary" form. These two forms are largely but not perfectly correlated with a significant mtDNA subdivision within the species that is thought to have arisen in allopatry. Combined information about multiple, independently evolving traits, such as morphology and structural and behavioural differences in calling song, provides greater resolution of the overall differences between these allopatric populations, and allows us to assess their stage of divergence. We test two predictions, first that the forms differ in song and second that gregarious males are more reluctant to sing than solitary males due to sex role reversal. We also tested for a difference in the relationship between the size of the forewing resonator, the mirror, and carrier frequency, as most models of sound production in crickets indicate that mirror size should predict carrier frequency. Results: Multivariate analyses showed that solitary and gregarious individuals from different populations representing the two mtDNA clades had almost non-overlapping distributions based on multiple song and morphological measurements. Carrier frequency differed between the two, and gregarious males were more reluctant to sing overall. Mirror size predicted carrier frequency; however, the relationship between mirror size and surface area varied between solitary and gregarious forms, suggesting that factors above and beyond mirror size contribute to carrier frequency. Conclusion: The two clades of Mormon crickets differ in a broad suite of independent traits that probably justify subspecies status ( the two can successfully mate so may not be reproductively isolated). However, our results emphasize the importance of carefully distinguishing the ultimate causation of differences between traits used to delineate species or subspecies boundaries.Publisher PDFPeer reviewe

Sexual Selection for Male Mobility in a Giant Insect with Female‐Biased Size Dimorphism

Female-biased size dimorphism in which females are larger than males is prevalent in many animals, but the factors causing this pattern of dimorphism are still poorly understood. The agility hypothesis suggests that female-biased size dimorphism arises because smaller males are favoured in scramble competition for mates. Using radio telemetry, we assessed the agility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e. longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating) and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large and/or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males

Radio-telemetric evidence of migration in the gregarious but not the solitary morph of the Mormon cricket (Anabrus simplex : Orthoptera: Tettigoniidae

Abstract The Mormon cricket, Anabrus simplex, is one of just a few species of katydids (or bushcrickets, Orthoptera: Tettigoniidae) that, like migratory locusts, appear to have solitary and migratory morphs. Using radio telemetry we studied movements of individuals of two morphs of this flightless species. Individuals within each migratory band had similar rates of movements along similar directional headings whereas solitary individuals moved little and showed little evidence of directionality in movement. Our results also add to other recent radio-telemetry studies showing that flightless insects of 1-2 g in mass can be tracked successfully using these methods

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

Multiple differences in calling songs and other traits between solitary and gregarious Mormon crickets from allopatric mtDNA clades

BACKGROUND: In acoustic species, traits such as male calling song are likely to diverge quickly between allopatric populations due to sexual selection, and divergence in parameters such as carrier frequency, chirp structure, and other important song characters can influence sexual isolation. Here we make use of two forms of Mormon crickets to examine differences in a broad suite of traits that have the potential to influence speciation via sexual isolation. Mormon crickets in "gregarious" populations aggregate into dense migratory bands, and females are the sexually competitive sex (sex-role reversal). There is also a non-outbreak "solitary" form. These two forms are largely but not perfectly correlated with a significant mtDNA subdivision within the species that is thought to have arisen in allopatry. Combined information about multiple, independently evolving traits, such as morphology and structural and behavioural differences in calling song, provides greater resolution of the overall differences between these allopatric populations, and allows us to assess their stage of divergence. We test two predictions, first that the forms differ in song and second that gregarious males are more reluctant to sing than solitary males due to sex role reversal. We also tested for a difference in the relationship between the size of the forewing resonator, the mirror, and carrier frequency, as most models of sound production in crickets indicate that mirror size should predict carrier frequency. RESULTS: Multivariate analyses showed that solitary and gregarious individuals from different populations representing the two mtDNA clades had almost non-overlapping distributions based on multiple song and morphological measurements. Carrier frequency differed between the two, and gregarious males were more reluctant to sing overall. Mirror size predicted carrier frequency; however, the relationship between mirror size and surface area varied between solitary and gregarious forms, suggesting that factors above and beyond mirror size contribute to carrier frequency. CONCLUSION: The two clades of Mormon crickets differ in a broad suite of independent traits that probably justify subspecies status (the two can successfully mate so may not be reproductively isolated). However, our results emphasize the importance of carefully distinguishing the ultimate causation of differences between traits used to delineate species or subspecies boundaries

The role of functional constraints in nonrandom mating patterns for a dance fly with female ornaments

Most hypotheses to explain non‐random mating patterns invoke mate choice, particularly in species that display elaborate ornaments. However, conflicting selection pressures on traits can result in functional constraints that can also cause non‐random mating patterns. We tested for functional load‐lifting constraints during aerial copulation in Rhamphomyia longicauda, a species of dance fly that displays multiple extravagant female‐specific ornaments that are unusual among sexual traits because they are under stabilizing selection. R. longicauda males provide females with a nuptial gift before engaging in aerial mating, and the male bears the entire weight of the female and nuptial gift for the duration of copulation. In theory, a male's ability to carry females and nuptial gifts could constrain pairing opportunities for the heaviest females, as reported for non‐ornamented dance flies. In concert with directional preferences for large females with mature eggs, such a load‐lifting constraint could produce the stabilizing selection on female size previously observed in this species. We therefore tested whether wild‐caught male R. longicauda collected during copulation were experiencing load‐lift limitations by comparing the mass carried by males during copulation with the male's wing loading traits. We also performed permutation tests to determine whether the loads carried by males during copulation were lighter than expected. We found that heavier males are more often found mating with heavier females suggesting that while R. longicauda males do not experience a load‐lift constraint, there is a strong relationship of assortative mating by mass. We suggest that active male mate choice for intermediately adorned females is more likely to be causing the non‐random mating patterns observed in R. longicauda

Listening to male song induces female field crickets to differentially allocate reproductive resources

Differential investment in offspring by mothers is predicted when there is substantial variation in sire quality. Whether females invest more resources in the offspring of high-quality mates (differential allocation, DA) or in the offspring of low-quality mates (reproductive compensation, RC) is not consistent in the literature and both effects can be predicted by theoretical models. In the field cricket, Gryllus pennsylvanicus Burmeister, 1838 (Orthoptera: Gryllidae: Gryllinae), females are attracted more to calling songs of high-quality males than to those of low-quality males. We tested whether females invest reproductive resources differentially based on perceived mate quality. We manipulated female perception of male quality by allowing virgin females to approach a speaker broadcasting either high- or low-quality calling song (high or low calling effort respectively), and then mated them with a randomly chosen male that had been rendered incapable of calling. In the week following mating, females exposed to high-quality calling song gained less weight, laid more embryos, and laid larger embryos than females exposed to low-quality calling song, although only the first of these effects was statistically significant. These results support the DA hypothesis and suggest that females invest their reproductive output based on a trait (calling effort) that is an honest indicator of male quality

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