Sexual competition among females: What causes courtship-role reversal?

In most animals, males are the competitive sex whereas females are typically non-competitive and choosy of mates. In a variety of taxa, certain species (or populations within species) show a reversal in these typical courtship roles. Recent research with these organisms supports a central tenet of sexual selection theory: that it is the relative investment of the sexes in offspring that controls the number of males and females available for mating, and thus is the main determinant of the degree of sexual competition in each sex.     

Differential mating success of virgin female katydidsRequena verticalis (Orthoptera: Tettigoniidae)

Male katydids (Orthoptera: Tettigoniidae) invest in offspring through nutrients provided in a large spermatophore. Previous research with Requena verticalishad shown that almost all of the investment of males mating with recently mated (4 days previously) females is in eggs fertilized by the female's previous mate. Thus males are predicted to discriminate against such females as mates. In experiments placing males with both a virgin and a female mated 4–5 days previously, virgin females obtained almost all matings. Although male discrimination of mates was noted in the experiments, there was no evidence that such discrimination was against nonvirgins in both this experiment and one in which a single virgin or mated female was placed with a male. Instead, the results suggest that the differential mating was a result of interfemale competition. The mating advantage held by virgin females over nonvirgins appeared to be lost once the latter had oviposited. Finally, there was no evidence from both single- and paired-female experiments that males preferred larger females as mates.     

FEMALE CHOICE FOR AN INDICATOR OF MALE SIZE IN THE SONG OF THE BLACK-HORNED TREE CRICKET,OECANTHUS NIGRICORNIS (ORTHOPTERA: GRYLLIDAE: OECANTHINAE)

The calling song of male crickets, including Oecanthus nigricornis (Walker), attracts females for mating and provides a model system of sexual communication. We give the first conclusive identification of a feature of cricket song that is both attractive to females and indicates a phenotypic feature (body size) that determines male mating success and female reproductive benefits. We do this by first testing for correlations between song characteristics and aspects of male phenotype that are hypothesized to indicate male quality. We show that song is a reliable indicator of male size and male age, and that large male size is associated with increased female fecundity. We then use playbacks of synthetic songs that mimic natural variation in song parameters to study song preferences and we compare preferences under different presentation regimes to determine whether choices are based on relative song quality or some fixed criterion. Females show a preference for the lower frequency songs produced by large males, but only during simultaneous playbacks. Thus female choice is based on the relative quality of calls that can be sampled simultaneously. These results provide strong support for the hypothesis that females use variation in calling song to assess male mate quality.     

Mate feeding, offspring investment, and sexual differences in katydids (Orthoptera: Tettigoniidae)

Food stress in the katydid Requena veriicalis (Orthoptera: Tettigoniidae)decreases the relative availability of males able to supplynutritious spermatophores to females and increases the valueof the male courtship meal (i.e., relative male parental investment).These changes cause female sexual competition in katydid populations.Here we examine the effect of food stress on male and femaleinvestment in single offspring and test the prediction thatmale-derived nutrients in eggs increase relative to nutrientsfrom the female's reserves. We varied the diet of female R.veriicalis and determined the fate of nutrients from male andfemale sources using I4C and 3H radiolabeled amino acids. Low-dietfemales retained more nutrients from male and female sourcesin somatic tissues and invested less in reproduction both becausethey produced fewer eggs and because they invested less peroffspring (egg) than females maintained on a high-quality diet.Moreover, opposite to our prediction, relative male investmentin individual eggs decreased in foodstressed females; femalesretained more nutrients in somatic tissues from the male sourcethan the female source. Food-stressed females may retain nutrientreserves, particularly those from the male, as an adaptive trategyfor immediate survival needs and future reproduction. Such afemale strategy is unlikely to compromise male reproductivesuccess; first-male sperm precedence means that males matingwith virgin females are likely to father most eggs laid, evenin future reproductive bouts. The decrease in male investmentin eggs of low-diet females does not conflict with the contentionthat relative parental investment controls male intrasexualcompetition because, in mate-feeding species, male investmentinfluencing this competition includes more than investment incurrent offspring; females should compete for males if courtshipgifts aid survival and later reproduction.     

Darwinian balancing selection: Predation counters sexual selection in a wild insect

The potential viability costs of sexually selected traits are central to hypotheses about the evolution of exaggerated traits. Estimates of these costs in nature can come from selection analyses using multiple components of fitness during the same time frame. For a population of tree crickets (Oecanthus nigricornis: Gryllidae), we analyzed viability and sexual selection on male traits by comparing Oecanthus prey of a solitary wasp to those that survived, and comparing mating individuals to solitary males. We measured forewing width (sexually size dimorphic and used for singing), head width, pronotum length, and size of hind jumping legs as potential targets of selection. Supporting the hypothesis that sexually selected traits have viability costs, we found that significant directional sexual selection for wider heads was opposed by significant viability selection for narrower heads. Nonlinear selection revealed that individuals with wide heads and small legs were most attractive, but individuals with narrow heads, large legs, and intermediate pronotum length were most likely to survive. Successful mating may put males at greater risk of predation, especially if copulation per se is risky. Such balancing selection in tree crickets may have constrained the evolution of sexual dimorphism in head size—a condition seen in other gryllids and orthopterans.     

Egg Load Decreases Mobility and Increases Predation Risk in Female Black-Horned Tree Crickets (Oecanthus nigricornis)

Female-biased predation is an uncommon phenomenon in nature since males of many species take on riskier behaviours to gain more mates. Several species of sphecid wasps have been observed taking more female than male prey, and it is not fully understood why. The solitary sphecid Isodontia mexicana catches more adult female tree cricket (Oecanthus nigricornis) prey. Previous work has shown that, although female tree crickets are larger and thus likely to be more valuable as prey than males, body size alone cannot fully explain why wasps take more females. We tested the hypothesis that wasps catch adult female tree crickets more often because bearing eggs impedes a female’s ability to escape predation. We compared female survivors to prey of I. mexicana, and found that females carrying more eggs were significantly more likely to be caught by wasps, regardless of their body size and jumping leg mass. We also conducted laboratory experiments where females’ jumping responses to a simulated attack were measured and compared to her egg load and morphology. We found a significant negative relationship between egg load and jumping ability, and a positive relationship between body size and jumping ability. These findings support the hypothesis that ovarian eggs are a physical handicap that contributes to female-biased predation in this system. Predation on the most fecund females may have ecological-evolutionary consequences such as collapse of prey populations or selection for alternate life history strategies and behaviours.