Male clasping ability, female polymorphism and sexual conflict: fine-scale elytral morphology as a sexually antagonistic adaptation in female diving beetles

During sexual conflict, males and females are expected to evolve traits and behaviours with a sexually antagonistic function. Recently, sexually antagonistic coevolution was proposed to occur between male and female diving beetles (Dytiscidae). Male diving beetles possess numerous suction cups on their forelegs whereas females commonly have rough structures on their elytra. These rough structures have been suggested to obstruct adhesion from male suction cups during mating attempts. However, some diving beetle species are dimorphic, where one female morph has a rough elytra and the other has a smooth elytra. Here, we used biomechanics to study the adhesive performance of male suction cups on the female morphs in two diving beetle species: Dytiscus lapponicus and Graphoderus zonatus . We compared adhesion on the rough and the smooth female morphs to infer the function of the rough elytral modifications. We found that the adhesive force on the rough structures was much lower than on other surfaces. These findings support the suggestion of sexual conflict in diving beetles and a sexually antagonistic function of the rough female structures. In addition, males differed in their adhesive capacity on different female surfaces, indicating a male trade-off between adhering to smooth and rough female morphs.

Frequency-dependent variation in mimetic fidelity in an intraspecific mimicry system

Contemporary theory predicts that the degree of mimetic similarity of mimics towards their model should increase as the mimic/model ratio increases. Thus, when the mimic/model ratio is high, then the mimic has to resemble the model very closely to still gain protection from the signal receiver. To date, empirical evidence of this effect is limited to a single example where mimicry occurs between species. Here, for the first time, we test whether mimetic fidelity varies with mimic/model ratios in an intraspecific mimicry system, in which signal receivers are the same species as the mimics and models. To this end, we studied a polymorphic damselfly with a single male phenotype and two female morphs, in which one morph resembles the male phenotype while the other does not. Phenotypic similarity of males to both female morphs was quantified using morphometric data for multiple populations with varying mimic/model ratios repeated over a 3 year period. Our results demonstrate that male-like females were overall closer in size to males than the other female morph. Furthermore, the extent of morphological similarity between male-like females and males, measured as Mahalanobis distances, was frequency-dependent in the direction predicted. Hence, this study provides direct quantitative support for the prediction that the mimetic similarity of mimics to their models increases as the mimic/model ratio increases. We suggest that the phenomenon may be widespread in a range of mimicry systems.

Multi-annual variation in female morph frequencies of the polymorphic damselfly, Nehalennia irene, at continental and regional scales

Female-limited polymorphism occurs in different animal taxa but is particularly abundant among species of damselflies (Insecta: Odonata), most likely as a consequence of selection to avoid excessive male harassment. Recent work on the damselfly Nehalennia irene indicated that within year spatial variation in female morph frequencies was limited in nearby populations (i.e. intra-regional scale), but large at a continental scale. As anticipated, some of the observed variation in morph frequency was correlated with variation in the estimated degree of male harassment towards female morphs, measured by male density and operational sex ratio. Here, we extended earlier work by quantifying variation in morph frequency over two to three years, allowing us to elucidate how morph frequencies vary temporally at both intra-regional and continental scales (data for 8 populations over three years and for 33 populations over two years, respectively). Annual variation in morph frequencies was relatively high at the intra-regional scale, but was never large enough to obscure the underlying spatial pattern at the continental scale. At both geographic scales, male density and operational sex ratio were highly variable between years. The estimated degree of male harassment correlated with variation in morph frequency within some regions, but not all. Together, the observed natural variation in female morph frequencies may be partly explained by variation in male harassment, but it appears that a complete understanding will require considering the role of other environmental factors.

Variation in flight morphology in a female polymorphic damselfly: intraspecific, intrasexual, and seasonal differences

In aerial animals, flight morphology needs to be designed to allow daily behavioural activities. Within species differences in behaviour can therefore be expected to relate to differences in flight morphology, not only between males and females but also between same-sex members when they use different behavioural strategies. In female polymorphic damselflies, one female morph is considered a male mimic that resembles the male’s body colour and behaviour (andromorph), whereas the other is dissimilar (gynomorph). Here, we questioned whether males, andromorphs, and gynomorphs of the damselfly Enallagma cyathigerum (Charpentier, 1840) differ in flight morphology, with andromorphs being more similar to males than gynomorphs. In addition, we evaluated whether differences in flight morphology are consistent or whether some morphs are more plastic in response to seasonal environmental fluctuations. Most morphometrics showed similar seasonal plasticity for males and both female morphs, which could only partly be explained from allometry. Consistent with high manoeuvrability in flight, males had broader wings and lower wing loading than females. Variation between female morphs was less pronounced, with no consistent differences in length, aspect ratio, total surface, and wing loading. However, we detected morph-specific differences in shape and width, with andromorphs having broader wings than gynomorphs similarly to males.

Female-limited polymorphism in a widespread damselfly: morph frequencies, male density, and phenotypic similarity of andromorphs to males

In several animal species, one male type coexists with two to several female types, a polymorphism often explained in the context of sexual selection. Where it occurs, one female morph typically resembles the conspecific male phenotype, but the degree of resemblance varies across species. Here, we question whether the degree of phenotypic similarity between male-like females and males varies within species. Phenotypic resemblance is hypothesized to depend on the potential for frequency- and density-dependent selection on male and (or) female phenotypes. We studied six populations of the damselfly Nehalennia irene (Hagen, 1861) that differed widely in estimates of morph frequency and male density. Male-like females resemble males more than another female type resembles males, across populations, when comparisons are based on abdominal patterns. Abdomen phenotype does matter in male–female interactions of damselflies. Furthermore, male-like females were more similar to males at low and high density sites compared with sites with intermediate densities, contrary to the hypothesis that the potential for male harassment influences the degree of phenotypic similarity. Additionally, male-like females of most populations converged on the abdominal phenotype of males of one population rather than on that of syntopic males; a problem that has not received any attention.

Seasonal and diurnal variation in the proportions of female morphs of the damselfly Enallagma cyathigerum

In many damselfly species a female-limited colour polymorphism is encountered which is assumed to be the result of sexual conflict. Typically, one morph resembles the male's body colouration (andromorph), while the other is dissimilar (heteromorph). Little is known about the extent of temporal variation in female morph proportions at the water where mating occurs. Knowledge about such variation should help to identify the factors that affect female morph proportion and the scales at which these factors operate. The objective of this study is to assess the occurrence of diurnal and seasonal variation in female morph proportions at the water for the damselfly Enallagma cyathigerum. Diurnal variation was evaluated at six nearby populations, while seasonal variation was examined at one of these populations. Furthermore, we considered temporal variation in female morph proportion in relation to proxies of male harassment (i.e., male density and operational sex ratio). Our findings indicate that female morph proportion varies throughout a day but is uniform on a seasonal scale. Variation in female morph proportions could not be explained by concomitant variation in male density or operational sex ratio. We suggest future study of male mate choice may consider temporal variation in female morph proportions at the water.