SIGNALING EFFICACY DRIVES THE EVOLUTION OF LARGER SEXUAL ORNAMENTS BY SEXUAL SELECTION

Why are there so few small secondary sexual characters? Theoretical models predict that sexual selection should lead to reduction as often as exaggeration, and yet we mainly associate secondary sexual ornaments with exaggerated features such as the peacock's tail. We review the literature on mate choice experiments for evidence of reduced sexual traits. This shows that reduced ornamentation is effectively impossible in certain types of ornamental traits (behavioral, pheromonal, or color‐based traits, and morphological ornaments for which the natural selection optimum is no trait), but that there are many examples of morphological traits that would permit reduction. Yet small sexual traits are very rarely seen. We analyze a simple mathematical model of Fisher's runaway process (the null model for sexual selection). Our analysis shows that the imbalance cannot be wholly explained by larger ornaments being less costly than smaller ornaments, nor by preferences for larger ornaments being less costly than preferences for smaller ornaments. Instead, we suggest that asymmetry in signaling efficacy limits runaway to trait exaggeration.     

Female collared flycatchers learn to prefer males with an artificial novel ornament

We experimentally investigated whether learning from previousexperiences can lead to the establishment of a new mate preferencein a wild population of birds. During year one (2001), 63 femalecollared flycatchers (Ficedula albicollis) bred together withmales that we had provided with a novel trait, a red stripeon their white forehead patch (a sexually selected trait). Somecolor patterns of birds are largely determined by a few genes,and this experiment was designed to mimic the occurrence ofmutations in such genes. In the subsequent year (2002), we foundthat females with previous experience with red-striped maleswere more likely to pair with red-striped males (76%) than withcontrol males. By contrast, naïve females (i.e., with noprevious experience with red-striped males) were not more likelyto pair with red-striped males (44%) than with control males.Females paired with red-striped males produced more offspringthan females paired with control males, suggesting that maleswith the novel trait had become favored by selection. Thus,female collared flycatchers appear to quickly learn to associatea novel trait with a suitable mate that, in turn, leads to assortativemating between local mates (i.e., males with the new trait andfemales with previous experience of the new trait). Our resultsprovide support for the notion that learning may play an importantrole when the co-evolution of preferences and preferred traitstakes different routes in different populations of the samebird species.     

Carotenoids‐based reddish pelvic spines in nonreproducing female and male sticklebacks (Gasterosteus aculeatus) – Signalling social dominance?

Conspicuous ornaments are often considered a result of evolution by sexual selection. According to the social selection hypothesis, such conspicuous traits may also evolve as badges of status associated with increased boldness or aggression toward conspecifics in conflicts about ecological resources. This study tested predictions from the social selection hypothesis to explain evolution of conspicuous red color of the pelvic spines of the three‐spine stickleback (Gasterosteus aculeatus). Wild nonreproducing sticklebacks were presented to pairs of dummies which differed at their pelvic spines, having either (i) normal‐sized gray or red pelvic spines or (ii) normal‐sized gray or large red pelvic spines. The experimental tank was illuminated by white or green light, since green light impedes the sticklebacks’ ability to detect red color. The dummies moved slowly around in circles at each end of the experimental tank. We quantified the parameters (i) which of the two dummies was visited first, (ii) time taken before the first visit to a dummy, (iii) distribution of the focal sticklebacks in the two zones close to each of the two dummies and in the neutral zone of the tank, (iv) close to which of the two dummies did the focal fish eat its first food‐piece, and (v) time spent until the first piece of food was eaten. This was carried out for 22 females and 29 males sticklebacks. The results suggested no effect of the color or size of the dummies’ pelvic spines, on none of the five behavioral parameters. Moreover, neither the color of the pelvic spines of the focal sticklebacks themselves (as opposed to redness of the dummies’ spines) nor their body length was associated with behavior toward the dummies. Thus, this study did not support predictions from the social selection hypothesis to explain evolution of red pelvic spines in sticklebacks.     

Vicariance and convergence in Magellanic and New Zealand long‐looped brachiopod clades (Pan‐Brachiopoda: Terebratelloidea)

Phylogenetic reconstructions using parsimony, maximum likelihood (ML), and Bayesian relaxed molecular clock analyses of ～2850 nucleotides of nuclear‐encoded small and large subunit ribosomal DNAs (SSU and LSU rDNAs) from 14 long‐looped (terebratelloid) ingroup and four short‐looped (terebratulidine) outgroup brachiopod taxa, together with ML analyses of ～663 nucleotides of mitochondrial cytochrome oxidase subunit 1 (cox1) from 12 terebratelloid taxa, show that deep divergence separates taxa endemic to waters in the vicinity of New Zealand from those with a Magellanic distribution around South America and Antarctica. This deep divergence also separates Magellanic Terebratella dorsata from New Zealand Terebratella sanguinea, showing that they are not congeneric. Instead, they belong to separate ‘Magellanic’ (MAG) and ‘New Zealand’ (NZ) clades that first diverged about 82 Mya (95% highest posterior density, 48–120 Mya), correlating with separation between the NZ and Antarctic tectonic plates. Sequence analyses also reveal (1) that the Antarctic endemic taxa Magellania fragilis and Magellania joubini are not congeneric with Magellania venosa, suggesting that their previous placement in Aerothyris should be restored, and (2) that divergence between Antarctic and NZ species of the terebratulide Liothyrella occurred much later than plate separation, perhaps because of continuing gene flow caused by long‐lived larvae. The topology of the rDNA and cox1 gene trees implies that radial ornament of the shell (‘ribbing’) has been gained (and/or lost) independently within the MAG and NZ clades. Radial ribs are widespread in articulate brachiopods throughout the Phanerozoic, but no comparisons of brachiopod rib morphology and morphogenesis have been published. Our comparisons of transverse shell mid‐sections in the scanning electron microscope reveal no obvious evidence of differences in morphology between independently gained ribs. We also consider several ways in which ribs may affect fitness, including effects on hydrodynamics. Only scanty and inconclusive evidence is available, but we suggest that effects (if any) are likely to be of small magnitude; adaptive value of brachiopod shell ribs remains to be demonstrated. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 631–645.     

Effects of ornamentation and phylogeny on the evolution of wing shape in stalk‐eyed flies (Diopsidae)

Exaggerated male ornaments are predicted to be costly to their bearers, but these negative effects may be offset by the correlated evolution of compensatory traits. However, when locomotor systems, such as wings in flying species, evolve to decrease such costs, it remains unclear whether functional changes across related species are achieved via the same morphological route or via alternate changes that have similar function. We conducted a comparative analysis of wing shape in relation to eye‐stalk elongation across 24 species of stalk‐eyed flies, using geometric morphometrics to determine how species with increased eye span, a sexually selected trait, have modified wing morphology as a compensatory mechanism. Using traditional and phylogenetically informed multivariate analyses of shape in combination with phenotypic trajectory analysis, we found a strong phylogenetic signal in wing shape. However, dimorphic species possessed shifted wing veins with the result of lengthening and narrowing wings compared to monomorphic species. Dimorphic species also had changes that seem unrelated to wing size, but instead may govern wing flexion. Nevertheless, the lack of a uniform, compensatory pattern suggests that stalk‐eyed flies used alternative modifications in wing structure to increase wing area and aspect ratio, thus taking divergent morphological routes to compensate for exaggerated eye stalks.     

Testosterone levels and gular pouch coloration in courting magnificent frigatebird (Fregata magnificens): variation with age-class, visited status and blood parasite infection

Male magnificent frigatebirds (Fregata magnificens) possess a seasonally expressed skin ornament, namely the red and inflatable gular pouch, and are, therefore, a convenient model for the study of some theories related to the evolution of possible testosterone-dependent sexual skin coloration. Here we report the findings of a study performed over four consecutive mating seasons in the Mexican national park Isla Isabel. We investigated differences in testosterone level and gular pouch coloration in courting males in relation to the categories: age-class, visited status and blood parasite infection. Gular pouch color saturation increased with age-class. Investigated frigatebirds were infected only with Haemoproteus iwa (Haemosporida, Haemoproteidae), with an overall prevalence infection of 15.5%. Prevalence of the infection increased with birds' age-class. Testosterone levels were significantly higher in infected males, who also had lighter colored gular pouches. In non-infected males, those visited by a female had higher testosterone levels than non-visited males. Gular pouch lightness and redness were negatively correlated but only redness in non-visited non-infected males was positively correlated with testosterone levels. Gular pouch saturation in visited and infected males was positively correlated with body mass, which also increased with age-class. Mated males had lower testosterone levels and lighter, less red and saturated gular pouch coloration than courting males. In summary, we found that coloration of the male skin ornament could reflect age-class (saturation), parasite infection (lightness) and mated status (all), together with indications of condition and testosterone dependency of ornament expression.