Static allometry suggests female ornaments in the long-tailed dance fly (Rhamphomyia longicauda) are not deceptive

Despite their prevalence in nature, the evolution of sex-specific female ornaments is still not well understood. Although in some cases (often carotenoid-based ornaments) they appear to honestly signal quality, such as fecundity, it has been suggested that some female ornaments have evolved to deceptively to obtain matings. We address these two hypotheses in the long-tailed dance fly ( Rhamphomyia longicauda ), where females possess two sex-specific ornaments: pinnate scales on the hind femur and tibia and abdominal sacs that are inflated in female-biased display swarms. Although several studies have suggested that female ornaments in this species are deceptive, evidence is mixed and requires further investigation. Here, we use static allometry (with body size as a proxy for condition) of both ornamental and non-ornamental traits in females (and homologous non-ornamental traits in males) in order to determine whether they are honest or deceptive signals of quality. Most male traits scaled isometrically with body size, however male leg hairs showed positive static allometry, probably because they are involved in nuptial-prey capture or in grasping mates. Ornamental traits in females (abdomen area and tibia scale length) showed significant positive allometry and had steep slopes relative to non-ornamental traits. As larger females invest more in ornamentation relative to smaller females, this suggests that these traits are likely honest, condition-dependent signals of quality. We note that honesty and deception are not mutually exclusive hypotheses. Individuals may vary in their signalling strategy, resulting in, for example, deception from some low condition individuals but honesty overall. Although our finding of positive allometry makes it unlikely this occurs in long tailed dance flies, simultaneous honesty and deception should be considered in future studies of female ornamentation.

Sex-specific ornament evolution is a consistent feature of climatic adaptation across space and time in dragonflies

Adaptation to different climates fuels the origins and maintenance of biodiversity. Detailing how organisms optimize fitness for their local climates is therefore an essential goal in biology. Although we increasingly understand how survival-related traits evolve as organisms adapt to climatic conditions, it is unclear whether organisms also optimize traits that coordinate mating between the sexes. Here, we show that dragonflies consistently adapt to warmer climates across space and time by evolving less male melanin ornamentation—a mating-related trait that also absorbs solar radiation and heats individuals above ambient temperatures. Continent-wide macroevolutionary analyses reveal that species inhabiting warmer climates evolve less male ornamentation. Community-science observations across 10 species indicate that populations adapt to warmer parts of species’ ranges through microevolution of smaller male ornaments. Observations from 2005 to 2019 detail that contemporary selective pressures oppose male ornaments in warmer years; and our climate-warming projections predict further decreases by 2070. Conversely, our analyses show that female ornamentation responds idiosyncratically to temperature across space and time, indicating the sexes evolve in different ways to meet the demands of the local climate. Overall, these macro- and microevolutionary findings demonstrate that organisms predictably optimize their mating-related traits for the climate just as they do their survival-related traits.

Synthesis and Outlook

In this final chapter I want to briefly recap what I presented in the previous chapters and provide a few ideas on what might be done in the future to move the field forward. All three factors I discussed as relevant in male mate choice—male investment in reproduction, sex ratios, and variability in partner quality—are still emerging fields in sexual selection research and need more theoretical and empirical work. I suggest that variability in female quality is more important and more complex than currently known. The same is true for sex ratios. On the other hand, I suggest that sheer investment in gametes may be a little less important than currently assumed. Most importantly we need to explore the interactions of these three pathways to male mate choice. Female competition and also female ornamentation are still somewhat enigmatic and both topics are likely to grow in importance for our understanding of sexual selection. I think considering male and female choice together, as well as female and male competition will ultimately provide a more complete picture of Darwinian sexual selection.

Male-like female morphs in hummingbirds: the evolution of a widespread sex-limited plumage polymorphism

Differences in the way males and females look or behave are common in animals. However, discrete variation within sexes (sex-limited polymorphism) also occurs in several vertebrate and invertebrate lineages. In birds, female-limited polymorphism (FLP) in which some females resemble males in coloration is most prominent in hummingbirds, a group known for its morphological and behavioural sexual dimorphism. Yet, it remains unclear whether this intrasexual colour variation in hummingbirds arises through direct selection on females, or indirectly as a non-adaptive byproduct resulting from selection on males. Here, we analysed specimens from more than 300 hummingbird species to determine the extent, evolutionary history and function of FLP. We found that FLP evolved independently in every major clade and occurs in nearly 25% of hummingbird species. Using phylogenetically informed analyses, we rejected non-adaptive hypotheses that FLP is the result of indirect selection or pleiotropy across species. Instead, FLP is associated with ecology, migratory status, and marginally with social dominance, suggesting a socioecological benefit to females. Ultimately, we show that FLP is not only widespread in hummingbirds and likely adaptive, but may also be useful for understanding the evolution of female ornamentation in systems under strong sexual selection.

Morphology of genitalia and non-genitalic contact structures in Trouessartia feather mites (Astigmata: Analgoidea: Trouessartiidae): is there evidence of correlated evolution between the sexes?

Positive correlations between the shapes of male and female sexual structures can be interpreted as cooperative or as combative. In the feather mite genus Trouessartia Canestrini, 1899, the spermaducts of females range from entirely internal to extending externally for varying lengths, whereas male primary genitalia range from gracile to massive. Males also possess a pair of adanal suckers used to hold onto the dorsal surface of the female during copulation. In the area of this attachment, females exhibit ornamentation and have strongly developed dorsal setae (setae h1), which we hypothesized serve to weaken the male’s hold during copulation. In male and female Trouessartia from 51 bird species, we compared female external spermaduct length and male genitalic “massiveness” and explored whether patterns of female dorsal ornamentation and (or) h1 seta size correlate with male adanal sucker size. Our results indicate that females with longer external spermaducts are associated with males with relatively massive genitalia. However, we found no significant relationship between male adanal sucker size and female ornamentation or h1 seta size. Further information regarding how the genitalia interact during sperm transfer is necessary to interpret correlations in genitalia size and strong intersexual differences in dorsal ornamentation and seta size in Trouessartia.

Does reproductive mode affect sexually-selected coloration? Evaluating UV–blue spots in parthenogenetic and bisexual lizards of the genus Darevskia

Sexual selection often leads to evolution of conspicuous signals, raising the chances of attracting not only potential mates, but also predators. In lacertid lizards, ultraviolet (UV)–blue spots on flanks and shoulders represent such a trait. Some level of correlation between male and female ornamentation is also known to exist. Therefore, the phenotype of females may change in the absence of sexual selection. We tested this hypothesis on a complex of parthenogenetic and bisexual lizards of the genus Darevskia. We evaluated area, counts, and chromatic properties (UV opponency, saturation) of UV–blue spots and compared the values between the clones and their bisexual progenitor species. We found a fair heterogeneity between the parthenogenetic species, but no general tendency toward higher crypsis or conspicuousness. Values of the parthenogens were not significantly different from the values of sexual females. A possible explanation is that the changes in selective forces associated with parthenogenetic reproduction are too small to affect the resulting pattern of selective pressures on the studied traits, or that the phenotypes of the parthenogens result from the unique combination of parental genomes and are conserved by clonal reproduction.

Competition for access to mates predicts female-specific ornamentation and polyandry

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 ornate 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, polyandry is increased. These findings support the hypothesis that ornament diversity in dance flies depends on female receptivity, 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 polyandry and 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.

No evidence for an adaptive role of early molt into breeding plumage in a female fairy wren

The evolution of ornaments as sexually selected signals is well understood in males, but female ornamentation remains understudied. Fairy wrens offer an excellent model system, given their complex social structure and mating systems, and the diversity of female ornamentation. We investigated whether early molt into ornamental breeding plumage plays an adaptive role in females of the monogamous purple-crowned fairy wren Malurus coronatus, the only fairy wren known to have female seasonal plumage. Using 6 years of monitoring, we found that the timing of female molt was similar to males, but there was no evidence for assortative mating. Like males (previous study), older and dominant individuals acquired their breeding plumage earlier; however, in contrast to males, early molt did not seem to be costly since unfavorable environmental conditions or previous reproductive effort did not delay molt. Early female molt was not associated with any indicator of reproductive quality nor did it attract additional offspring care by their partners. We also found no association between early molt and the likelihood of acquiring a dominant (breeding) position or with the presence or proximity to same-sex rivals. Our study results, which are similar to previous findings in conspecific males, suggest that directional selection for early molt might be relaxed in this species, in contrast to other genetically polygamous fairy wrens in which early molt predicts extrapair mating success in males. However, the finding that molt timing is status dependent raises the possibility that other attributes of the ornament may fulfill an adaptive function in females.