One tool, many uses: precopulatory sexual selection on genital morphology in Aquarius remigis

While congruent evidence indicates that sexual selection is the most likely selective force explaining the rapid divergence of male genital morphology in insects, the mechanisms involved in this process remain unclear. In particular, little attention has been paid to precopulatory sexual selection. We examine sexual selection for mating success on male genital components in six populations of Aquarius remigis, a water strider characterized by unique genital morphology. Multivariate selection analysis confirms previous findings that precopulatory sexual selection favours longer external genitalia, and provides new evidence that this selection acts independently on external genital components. In contrast, the size of the major internal genital sclerite is not correlated with mating success. Thus, precopulatory sexual selection acts strongly on the size of the external genitalia, but not on the intromittent organ itself. These results highlight the multiple functions of genital organs and the importance of both precopulatory and post-copulatory sexual selection in shaping the remarkable diversity of male genitalia in insects.     

Wing dimorphism and the migratory syndrome: Correlated traits for migratory tendency in wing dimorphic insects

The hypothesis that the morphological, physiological, and behavioral traits comprising the migratory syndrome in insects are genetically correlated through pleiotropic effects of genes controlling the titre of a common hormonal determinant is explored. Evidence that juvenile hormone (JH) influences the component traits of the migratory syndrome is presented, and thus JH is assumed to be the underlying, common determinant. However, readers are cautioned that this does not imply that JH is solely responsible for these traits, nor is this necessary for the arguments presented. For wing dimorphic taxa, the “correlated traits hypothesis” predicts covariance within wing morphs between JH titre and the proportion winged. Four simple genetic models for wing-morph determination are considered: single-locus with short-winged (SW) dominant; single-locus with long-winged (LW) dominant; polygenic, fixed threshold, shifting distribution; and polygenic, shifting threshold, fixed distribution. In each case, wing morphology is assumed to be a threshold trait with the liability being JH titre at some critical stage of development. All models predict covariation between %LW and the mean JH titre of at least one of the wing morphs, but the form and direction of the relationship depends critically on the genetic model used. The results suggest that we should expect the traits associated with the migratory syndrome, and hence the trade-offs associated with the evolution of wing dimorphism, to be correlated with proportion winged and, in this sense, to be frequency-dependent.     

THE EVOLUTION OF ALTERNATE MORPHOLOGIES: FITNESS AND WING MORPHOLOGY IN MALE SAND CRICKETS

Many organisms show distinct morphological types. We argue that the evolution of these alternate morphologies depends upon both fitness differences between morphs within each sex and the genetic correlation between sexes. In this paper, we examine the evolution of alternate morphologies using wing dimorphism in insects as a model system. Many insect species are wing dimorphic, one morph having wings and being capable of flight, the other lacking functional wings. While there is a well established trade-off in females between macroptery and reproduction, there are few data on the possible costs in males. We examine trade-offs between macroptery and life-history traits in male sand crickets, Gryllus firmus, and estimate the genetic correlation of wing dimorphism between the sexes. Macropterous males develop faster than micropterous males and are either larger or the same size depending upon rearing conditions. There is no difference in absolute or relative testis size at eclosion or 7 d thereafter. Finally, there is no difference between macropterous and micropterous males in relative success at siring offspring. Thus, with respect to the above traits, there are no costs associated with being winged in male G. firmus. It is possible that there may be a trade-off between calling rate and macroptery. A comparison of the relative frequency of macroptery between males and female across different orders of insects supports this hypothesis. The genetic correlation of wing dimorphism between the sexes is high (r₈ = 0.86), and hence the frequency of macroptery in males may be strongly influenced by selection acting on females.     

Macroevolutionary patterns of bumblebee body size: detecting the interplay between natural and sexual selection

Bumblebees and other eusocial bees offer a unique opportunity to analyze the evolution of body size differences between sexes. The workers, being sterile females, are not subject to selection for reproductive function and thus provide a natural control for parsing the effects of selection on reproductive function (i.e., sexual and fecundity selection) from other natural selection. Using a phylogenetic comparative approach, we explored the allometric relationships among queens, males, and workers in 70 species of bumblebees (Bombus sp.). We found hyperallometry in thorax width for males relative to workers, indicating greater evolutionary divergence of body size in males than in sterile females. This is consistent with the hypothesis that selection for reproductive function, most probably sexual selection, has caused divergence in male size among species. The slope for males on workers was significantly steeper than that for queens on workers and the latter did not depart from isometry, providing further evidence of greater evolutionary divergence in male size than female size, and no evidence that reproductive selection has accelerated divergence of females. We did not detect significant hyperallometry when male size was regressed directly on queen size and our results thus add the genus Bombus to the increasing list of clades that have female-larger sexual size dimorphism and do not conform to Rensch's rule when analyzed according to standard methodology. Nevertheless, by using worker size as a common control, we were able to demonstrate that bumblee species do show the evolutionary pattern underlying Rensch's rule, that being correlated evolution of body size in males and females, but with greater evolutionary divergence in males.     

THE EVOLUTION OF THRESHOLD TRAITS: A QUANTITATIVE GENETIC ANALYSIS OF THE PHYSIOLOGICAL AND LIFE-HISTORY CORRELATES OF WING DIMORPHISM IN THE SAND CRICKET

Many traits are phenotypically discrete but polygenically determined. Such traits can be understood using the threshold model of quantitative genetics that posits a continuously distributed underlying trait, called the liability, and a threshold of response, individuals above the threshold displaying one morph and individuals below the threshold displaying the alternate morph. For many threshold traits the liability probably consists of a hormone or a suite of hormones. Previous experiments have implicated juvenile hormone esterase (JHE), a degratory enzyme of juvenile hormone, as a physiological determinant of wing dimorphism in the crickets Gryllus rubens and G. firmus. The present study uses a half-sib experiment to measure the heritability of JHE in the last nymphal stadium of G. firmus and its genetic correlation with fecundity, a trait that is itself genetically correlated with wing morph. The phenotypic and genetic parameters are consistent with the hypothesis that JHE is a significant component of the liability. Comparison of sire and dam estimates suggest that nonadditive effects may be important. Two models have been proposed to account for the fitness differences between morphs: the dichotomy model, which assumes that each morph can be characterized by a particular suite of traits, and the continuous model, which assumes that the associated fitness traits are correlated with the liability rather than the morphs themselves. The latter model predicts that the fitness differences will not be constant but change with the morph frequencies. Variation in fecundity and flight muscle histolysis are shown to be more consistent with the continuous model. Data from the present experiment on JHE are inconclusive, but results from a previous selection experiment also suggest that variation in JHE is consistent only with the continuous model.     

Allometry for sexual size dimorphism: testing two hypotheses for Rensch's rule in the water strider Aquarius remigis

Within any given clade, male size and female size typically covary, but male size often varies more than female size. This generates a pattern of allometry for sexual size dimorphism (SSD) known as Rensch's rule. I use allometry for SSD among populations of the water strider Aquarius remigis (Hemiptera, Gerridae) to test the hypothesis that Rensch's rule evolves in response to sexual selection on male secondary sexual traits and an alternative hypothesis that it is caused by greater phenotypic plasticity of body size in males. Comparisons of three populations reared under two temperature regimes are combined with an analysis of allometry for genital and somatic components of body size among 25 field populations. Contrary to the sexual-selection hypothesis, genital length, the target of sexual selection, shows the lowest allometric slope of all the assayed traits. Instead, the results support a novel interpretation of the differential-plasticity hypothesis: that the traits most closely associated with reproductive fitness (abdomen length in females and genital length in males) are "adaptively canalized." While this hypothesis is unlikely to explain Rensch's rule among species or higher clades, it may explain widespread patterns of intraspecific variation in SSD recently documented for many insect species.