Armaments and ornaments: an evolutionary explanation of traits of dual Utility

Secondary sexual characters in many species function both in male-male competition and as cues for female choice. Based on a literature compilation of existing knowledge of traits with this dual function, we propose that they commonly arise through intersexual selection processes and serve as honest signals to other males regarding fighting ability or dominance. Faking these traits, here called armaments, (i.e. weapons and status badges) is difficult, as they are constantly put to trial in male-male contests. Females that subsequently utilize them as indicators of male phenotypic quality when selecting a partner will benefit by acquiring males of higher quality to father their offspring. Thus, evolution of armaments through male-male competition is seen as a usually initiating process, whereas female choice later may assume a role as an additional selective factor. The reverse, that males use information from traits evolved through female choice, is, however, also possible. The traditional view of independently evolved and temporarily unordered intra- and intersexual selection processes fails to explain dual trait functions. Moreover, our model may more satisfyingly than traditional ones explain how trait honesty and trait genetic variance are maintained: theoretical and empirical evidence suggests that such honesty and variation are more easily maintained under male-male competition than under female choice.  

A theory of mate choice based on heterozygosity

In theories of mate choice that rely on genetic benefits, thenature of the"good genes" involved has received little attention.A review of genetic studies of mate choice in a variety of speciesand situations suggests that individual heterozygosity is moreimportant than previously realized. Females are predicted tovalue heterozygosity in their offspring and under some conditionsin their males. The expression of vigor, condition-sensitiveornaments, and symmetry in males may be a direct reflectionnot of "good genes" but of individual heterozygosity at keyloci or at many loci. Like sexuality itself, mate choice basedon heterozygosity and genic diversity may be an adaptation thatfavors the production of diverse and superior competitors. Femalechoice is made meaningful by sexuality, and the adaptive valueof choice probably depends on some of the same factors thatmaintain sexuality  

On the measurement and classification of colour in studies of animal colour patterns

In studies of animal colouration it is no longer necessary to rely on subjective assessments of colour and conspicuousness, nor on methods which rely upon human vision. This is important because animals vary greatly in colour vision and colour is context-dependent. New methods make it practical to measure the colour spectrum of pattern elements (patches) of animals and their visual backgrounds for the conditions under which patch spectra reach the conspecific's, predator's or prey's eyes. These methods can be used in both terrestrial and aquatic habitats. A patch's colour is dependent not only upon its reflectance spectrum, but also upon the ambient light spectrum, the transmission properties of air or water, and the veiling light spectrum. These factors change with time of day, weather, season and microhabitat, so colours must be measured under the conditions prevalent when colour patterns are normally used. Methods of measuring, classifying and comparing colours are presented, as well as techniques for assessing the conspicuousness of colour patterns as a whole. Some implications of the effect of environmental light and vision are also discussed.  

Structurally based plumage coloration is an honest signal of quality in male blue grosbeaks

We investigated the signaling function of blue plumage in maleblue grosbeaks (Guiraca caerulea) to determine if structurallybased coloration may act as a reliable signal of quality toconspecifics. Blue plumage results from the microstructureof feather barbules rather than from pigment granules, andthus it is possible that structurally based plumage ornamentsmay function differently from sexually selected ornamental coloration that is pigment based. The plumage of male blue grosbeaksreflects maximally in the blue-ultraviolet range, so most variationin plumage coloration among males is invisible to human observers.In previous research, we showed that increased area of blueplumage on the body is associated with a shift in the wavelengthof maximum feather reflectance toward the ultraviolet and withhigh intensity of light reflected at that maximum, and thatextreme expression of the male ornament is condition dependent.These observations suggest that blue plumage may be an honestadvertisement of male quality. We tested this hypothesis ina wild population of blue grosbeaks. We quantified male qualityin three broad categories. (1) Physical condition was assessed from subcutaneous fat deposits, ectoparasite load, and bodysize. (2) Territory quality was assessed from territory area,prey abundance, and predation risk. (3) Paternal investmentwas assessed from male feeding rate. We found that the bluestmales have the largest body size, maintain the largest territorieswith the greatest prey abundance, and feed nestlings in thefirst nest of the season at the highest rates. We conclude that structurally based plumage coloration functions as an honest,intraspecific signal of quality.  

Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration

In birds, carotenoid-based plumage coloration is more dependent on physical condition and foraging abilities and less constrained developmentally than is melanin-based coloration. Thus, female mate choice for honest signals should result in more intense sexual selection on carotenoid- than on melanin-based plumage coloration. Using variation in sexual dimorphism as an indirect measure of the intensity of sexual selection, we tested the prediction mat variation in sexual dimorphism is driven more by change in carotenoid-based coloration between males and females dian by change in melanin-based coloration. Examination of historical changes in carotenoid- versus melanin-based pigmentation in 126 extant species of Cardueline finches supported this prediction. We found that carotenoid-derived coloration changed more frequendy among congeners dian melanin-based coloration. In both sexes, increase in carotenoid-based coloration score, but not in melanin-based coloration score, was strongly associated with increase in sexual dichromatism. In addition, sexual dimorphism in carotenoid-based coloration contributed more to overall dichromatism than dimorphism in melanin-based plumage. Our results supported die hypothesis that melanin-based and carotenoid-based coloration have fundamentally different signal content and suggest that combining melanin-based and carotenoid-based coloration in comparative analyses is not appropriate.  

Antler size in red deer: heritability and selection but no evolution

We present estimates of the selection on and the heritability of a male secondary sexual weapon in a wild population: antler size in red deer. Male red deer with large antlers had increased lifetime breeding success, both before and after correcting for body size, generating a standardized selection gradient of 0.44 (+/- 0.18 SE). Despite substantial age- and environment-related variation, antler size was also heritable (heritability of antler mass = 0.33 +/- 0.12). However the observed selection did not generate an evolutionary response in antler size over the study period of nearly 30 years, and there was no evidence of a positive genetic correlation between antler size and fitness nor of a positive association between breeding values for antler size and fitness. Our results are consistent with the hypothesis that a heritable trait under directional selection will not evolve if associations between the measured trait and fitness are determined by environmental covariances: In red deer males, for example, both antler size and success in the fights for mates may be heavily dependent on an individual's nutritional state.  

Structural and melanin coloration indicate parental effort and reproductive success in male eastern bluebirds

Male eastern bluebirds (Sialia sialis) have two types of ornamentalplumage coloration: a brilliant blue-ultraviolet head, back,and wings, and a patch of chestnut breast feathers. The blue-UVcoloration is produced from feather microstructure, whereasthe chestnut coloration is produced by a combination of pheaomelaninand eumelanin pigments deposited in feathers. We tested thehypothesis that plumage coloration reflects male quality ineastern bluebirds, a socially monogamous, sexually dichromaticbird. We investigated whether male ornamentation correlateswith mate quality and parental effort. We quantified three aspectsof male ornament coloration: (1) size of the patch of chestnutbreast feathers, (2) reflectance properties of the chestnutplumage coloration, and (3) reflectance properties of the blue-ultravioletplumage coloration. We found that males with larger breast patchesand brighter plumage provisioned nestlings more often, fledgedheavier offspring, and paired with females that nested earlier.Males with plumage coloration that exhibit more ultraviolethues fledged more offspring. These results suggest that plumagecoloration is a reliable indicator of male mate quality andreproductive success. Both melanin-based and structural-basedplumages appear to be honest signals of male quality and parentalcare that can be assessed by competitors or by potential mates.  

Why do females mate multiply? A review of the genetic benefits

The aim of this review is to consider the potential benefits that females may gain from mating more than once in a single reproductive cycle. The relationship between non-genetic and genetic benefits is briefly explored. We suggest that multiple mating for purely non-genetic benefits is unlikely as it invariably leads to the possibility of genetic benefits as well. We begin by briefly reviewing the main models for genetic benefits to mate choice, and the supporting evidence that choice can increase offspring performance and the sexual attractiveness of sons. We then explain how multiple mating can elevate offspring fitness by increasing the number of potential sires that compete, when this occurs in conjunction with mechanisms of paternity biasing that function in copula or post-copulation. We begin by identifying cases where females use pre-copulatory cues to identify mates prior to remating. In the simplest case, females remate because they identify a superior mate and 'trade up' genetically. The main evidence for this process comes from extra-pair copulation in birds. Second, we note other cases where pre-copulatory cues may be less reliable and females mate with several males to promote post-copulatory mechanisms that bias paternity. Although a distinction is drawn between sperm competition and cryptic female choice, we point out that the genetic benefits to polyandry in terms of producing more viable or sexually attractive offspring do not depend on the exact mechanism that leads to biased paternity. Post-copulatory mechanisms of paternity biasing may: (1) reduce genetic incompatibility between male and female genetic contributions to offspring; (2) increase offspring viability if there is a positive correlation between traits favoured post-copulation and those that improve performance under natural selection; (3) increase the ability of sons to gain paternity when they mate with polyandrous females. A third possibility is that genetic diversity among offspring is directly favoured. This can be due to bet-hedging (due to mate assessment errors or temporal fluctuations in the environment), beneficial interactions between less related siblings or the opportunity to preferentially fertilise eggs with sperm of a specific genotype drawn from a range of stored sperm depending on prevailing environmental conditions. We use case studies from the social insects to provide some concrete examples of the role of genetic diversity among progeny in elevating fitness. We conclude that post-copulatory mechanisms provide a more reliable way of selecting a genetically compatible mate than pre-copulatory mate choice. Some of the best evidence for cryptic female choice by sperm selection is due to selection of more compatible sperm. Two future areas of research seem likely to be profitable. First, more experimental evidence is needed demonstrating that multiple mating increases offspring fitness via genetic gains. Second, the role of multiple mating in promoting assortative fertilization and increasing reproductive isolation between populations may help us to understand sympatric speciation.  

Mating systems, sperm competition, and the evolution of sexual dimorphism in birds

Comparative analyses suggest that a variety of factors influence the evolution of sexual dimorphism in birds. We analyzed the relative importance of social mating system and sperm competition to sexual differences in plumage and body size (mass and tail and wing length) of more than 1,000 species of birds from throughout the world. In these analyses we controlled for phylogeny and a variety of ecological and life-history variables. We used testis size (corrected for total body mass) as an index of sperm competition in each species, because testis size is correlated with levels of extrapair paternity and is available for a large number of species. In contrast to recent studies, we found strong and consistent effects of social mating system on most forms of dimorphism. Social mating system strongly influenced dimorphism in plumage, body mass, and wing length and had some effect on dimorphism in tail length. Sexual dimorphism was relatively greater in species with polygynous or lekking than monogamous mating systems. This was true when we used both species and phylogenetically independent contrasts for analysis. Relative testis size was also related positively to dimorphism in tail and wing length, but in most analyses it was a poorer predictor of plumage dimorphism than social mating system. There was no association between relative testis size and mass dimorphism. Geographic region and life history were also associated with the four types of dimorphism, although their influence varied between the different types of dimorphism. Although there is much interest in the effects of sperm competition on sexual dimorphism, we suggest that traditional explanations based on social mating systems are better predictors of dimorphism in birds.  

Sexual ornamentation and immunocompetence in the barn swallow

The handicap hypothesis of honest signaling suggests that secondarysexual characters reliably reflect phenotypic or genotypic qualityof signalers. This hypothesis is based on the assumptions thatsignals are costly to produce and/or maintain and the cost ofa given level of signaling is higher for low quality than forhigh quality signalers. We tested these assumptions in a fieldexperiment in which the size of a secondary sexual character[tail length in male barn swallows (Hirundo rustica)] was experimentallymanipulated. Males were randomly assigned to tail elongation,tail shortening, or two control treatments (tail manipulation,or just capture, ringing, and handling). Male barn swallowswere challenged with an injection of sheep red blood cells,and blood was sampled on the day of first capture and after3 to 4 weeks for determination of concentrations of gamma-globulins.Tail-elongated males did not increase levels of gamma-globulinswhile males of the other three groups demonstrated increases.Analyses of variation in gamma-globulins within treatment groupsrevealed a positive correlation between gamma-globulins andoriginal tail length among males with elongated tails. Theseresults suggest that tail length imposes an immu-nocompetencecost on males, and that males with naturally long tails aredifferentially better able to cope with this cost.