The evolution of sexual dimorphism in parasitic cuckoos: sexual selection or coevolution?

Sexual dimorphism is ubiquitous in animals and can result from selection pressure on one or both sexes. Sexual selection has become the predominant explanation for the evolution of sexual dimorphism, with strong selection on size-related mating success in males being the most common situation. The cuckoos (family Cuculidae) provide an exceptional case in which both sexes of many species are freed from the burden of parental care but where coevolution between parasitic cuckoos and their hosts also results in intense selection. Here, we show that size and plumage differences between the sexes in parasitic cuckoos are more likely the result of coevolution than sexual selection. While both sexes changed in size as brood parasitism evolved, we find no evidence for selection on males to become larger. Rather, our analysis indicates stronger selection on parasitic females to become smaller, resulting in a shift from dimorphism with larger females in cuckoos with parental care to dimorphism with larger males in parasitic species. In addition, the evolution of brood parasitism was associated with more cryptic plumage in both sexes, but especially in females, a result that contrasts with the strong plumage dimorphism seen in some other parasitic birds. Examination of the three independent origins of brood parasitism suggests that different parasitic cuckoo lineages followed divergent evolutionary pathways to successful brood parasitism. These results argue for the powerful role of parasite-host coevolution in shaping cuckoo life histories in general and sexual dimorphism in particular.     

Can the theory of evolution be falsified?

In this paper we discuss the epistemological positions of evolution theories. A sharp distinction is made between the theory that species evolved from common ancestors along specified lines of descent (here called the theory of common descent), and the theories intended as causal explanations of evolution (e.g. Lamarck's and Darwin's theory). The theory of common descent permits a large number of predictions of new results that would be improbable without evolution. For instance, (a) phylogenetic trees have been validated now; (b) the observed order in fossils of new species discovered since Darwin's time could be predicted from the theory of common descent; (c) owing to the theory of common descent, the degrees of similarity and difference in newly discovered properties of more or less related species could be predicted. Such observations can be regarded as attempts to falsify the theory of common descent. We conclude that the theory of common descent is an easily-falsifiable & often-tested & still-not-falsified theory, which is the strongest predicate a theory in an empirical science can obtain. Theories intended as causal explanations of evolution can be falsified essentially, and Lamarck's theory has been falsified actually. Several elements of Darwin's theory have been modified or falsified: new versions of a theory of evolution by natural selection are now the leading scientific theories on evolution. We have argued that the theory of common descent and Darwinism are ordinary, falsifiable scientific theories.     

Maternal inheritance and rapid evolution of sexual size dimorphism: passive effects or active strategies?

Adaptive evolution is often strongly influenced by maternal inheritance that transfers the parental strategies across generations. The consequences of maternal effects for the offspring generation depend on the between-generation similarity in environments and on the evolved sensitivity of the offspring's ontogeny to maternal effects. When these factors differ between sons and daughters, maternal effects can influence the evolution of sexual dimorphism. The establishment of house finch populations across western Montana during the last 30 years was accompanied by rapid evolutionary change in sexual size dimorphism. Here I show that traits that changed the most across generations were most influenced by maternal effects in males but not females. Maternal effects differentially affected sons' and daughters' survival; greater maternal effects were commonly associated with higher survival of sons, especially when maternal and offspring environments were similar. Stronger maternal effects extended preselection phenotypic variance in morphological traits of males, thereby producing some locally adaptive phenotypes and lessening juvenile mortality. Thus, the observed sex-specific maternal effects and their contribution to the evolution of sexual size dimorphism are likely a passive consequence of the distinct sensitivity of sons and daughters to maternal adaptations to breeding in ecologically distinct parts of the house finch's expanding range.     

Can sympatric speciation by disruptive sexual selection explain rapid evolution of cichlid diversity in Lake Victoria?

Rapid speciation can occur on ecological time scales and interfere with ecological processes, resulting in species distribution patterns that are difficult to reconcile with ecological theory. The haplochromine cichlids in East African lakes are an extreme example of rapid speciation. We analyse the causes of their high speciation rates. Various studies have identified disruptive sexual selection acting on colour polymorphisms that might cause sympatric speciation. Using data on geographical distribution, colouration and relatedness from 41 species endemic to Lake Victoria, we test predictions from this hypothesis. Plotting numbers of pairs of closely related species against the amount of distributional overlap between the species reveals a bimodal distribution with modes on allopatric and sympatric. The proportion of sister species pairs that are heteromorphic for the traits under disruptive selection is higher in sympatry than in allopatry. These data support the hypothesis that disruptive sexual selection on colour polymorphisms has caused sympatric speciation and help to explain the rapid radiation of haplochromine species flocks.     

Is sexual dimorphism affected by the combined action of prenatal stress and sex ratio?

During embryonic development, offspring are exposed to hormones of both maternal and sibling origin. Maternal stress increases offspring exposure to corticosterone, and, in polytocous animals, the sex ratio or intrauterine position can influence the levels of androgens and estrogens experienced by the offspring. Such hormone exposure has the potential to influence many important morphological and behavioural aspects of offspring, in particular sexually dimorphic traits. Although well known in rodents, the impact of prenatal hormone exposure in other vertebrates is poorly documented. We experimentally investigated the relationship between maternal stress, population density, sex ratio (a surrogate for the degree of exposure to steroids produced by siblings), and sexual dimorphism in a viviparous lizard, Lacerta vivipara. Our results show that prenatal sex ratios have consequences for sexually dimorphic morphology (ventral scale count) in both sexes, but with no effect of maternal stress or any interaction between the two. Embryonic steroid exposure can potentially be an important factor in generating individual variation in natural populations of viviparous animals.     

Time and energy constraints and the evolution of sexual size dimorphism — to eat or to mate?

Summary We present an empirical test of the Ghiselin—Reiss small-male hypothesis for the evolution of sexual size dimorphism (SSD). In mating systems dominated by scramble competition, where male reproductive success is a function of encounter rate with females, small males may be favoured when food is limiting because they require lower absolute amounts of food. Given a trade-off between time and energy devoted to foraging and to mate acquisition, small males should be able to devote more time to the latter. If at the same time larger females are favoured, this mechanism will contribute to the evolution of SSD and may be the major determinant of the female-biased SSDs that characterize most animal taxa. We tested this hypothesis using the water strider,Aquarius remigis (Heteroptera: Gerridae), a scramble competitor which mates many times over a prolonged mating season and which shows female-biased SSD. Laboratory experiments demonstrated that foraging success and giving up times (GUTs) are lower for males than for females during the reproductive season and that male water striders flexibly alter their time budgets under conditions of energy limitation. Controlled feeding experiments showed that male and female longevity, female fecundity and male mating success are positively related to food availability. As predicted, male body size is negatively correlated with several indices of male fitness (longevity, number of mating attempts and mating success), while female body size is positively correlated with longevity. These results are consistent with the hypothesis that scramble competition for mates favours small males in this species and provides empirical support for the Ghiselin—Reiss small-male hypothesis.     

The evolution of male-female dimorphism: Older than sex?

This contribution considers the evolution of a dimorphism with respect to cell fusion characteristics in a population of primitive cells. These cells reproduce exclusively asexually. The evolution towards asymmetric fusion behaviour of cells is driven by selection promoting horizontal transfer of an endosymbiontic replicator. It is concluded that evolution of asymmetric cell fusion in this scenario is more likely than evolution of sexual differentiation in a sexually reproducing population. Pre-existing dimorphism with respect to cell fusion may thus have been the basis for the establishment of sexual differentiation at the level of gamete fusion, and this in turn is fundamental to the evolution of two different sexes, male and female.     

Establishing sexual dimorphism: conservation amidst diversity?

The molecular mechanisms that control sexual dimorphism are very different in distantly related animals. Did sex determination arise several times with different regulatory mechanisms, or is it an ancient process with little surviving evidence of ancestral genes? The recent identification of related sexual regulators in different phyla indicates that some aspects of sexual regulation might be ancient. Studies of sex-determining mechanisms are beginning to reveal how sexual dimorphism arises and evolves.     

Does sexual dimorphism in human faces signal health?

Evolutionary psychologists suggest that a preference for sexually dimorphic traits in human faces is an adaptation for mate choice, because such traits reflect health during development. For male faces, this claim rests on the immunocompetence-handicap hypothesis, which states that the increased testosterone levels needed to develop large masculine traits stress the immune system. We examined whether masculine traits in adolescent male faces are associated with health during development, and also whether feminine traits in adolescent female faces signal health. Feminine traits are attractive, but it is less clear whether they should signal health. Rated masculinity in adolescent male faces correlated modestly with actual health, and was perceived as healthy, but not as attractive. Rated femininity in adolescent female faces did not correlate with actual health, although it was perceived as healthy and attractive. These results support the immunocompetence-handicap hypothesis for male faces in that masculine traits signalled health during adolescence. However, they suggest that any health-related evolutionary benefits obtained from preferences for attractive facial traits may be weak.     

Does sexual dimorphism vary by population? Laryngeal and ear anatomy in cricket frogs

Acoustic communication in many anuran species can show the effects of both natural and sexual selection. This is reflected in the sexually dimorphic an atomy of the lary nx and ear structures, as well as the allometric relationship of these morphological traits to head or body size. In this study, we examined laryngeal and ear structures of cricket frogs Acris crepitans not only as sexually dimorphic characteristics, but also as they differ across populations in environmentally different habitats. We used 2-way ANOVA to determine whether the volumetric or linear measurements of these structures differed by sex and population. Females have significantly larger body, head, and ear sizes, but significantly smaller larynges than males. Furthermore, females as well as males show larger body and head sizes, ears, and larynges in a dryer open habitat. An ANCOVA analysis shows that males, but not females, differ in laryngeal size across populations beyond the allometric changes attributable to head size alone indicating that males have a greater degree of laryngeal population variation. In contrast, our covariate analysis found that in both sexes many of the ear differences are non-sigrdficant once head size is accounted for, suggesting that most of the population-level ear variation is due to allometric effects of body size. We conclude that although both sexes show size differences in the larynx related to selection for larger body size in dry, open habitats, selection on males for larger larynx size related to the production of lower frequency calls in those habitats does not result in correlated changes in the female larynx. The results suggest that in anurans, selection for changes in body and head size affects both sexes equally, male calls and the vocal structures responsible for them can further diversify without concordant changes in females.     

Can fertility signals lead to quality signals? Insights from the evolution of primate sexual swellings

The sexual swellings of female primates have generated a great deal of interest in evolutionary biology. Two hypotheses recently proposed to elucidate their functional significance argue that maximal swelling size advertises either female fertility within a cycle or female quality across cycles. Published evidence favours the first hypothesis, and further indicates that larger swellings advertise higher fertility between cycles. If so, a male preference for large swellings might evolve, driving females to use swellings as quality indicators, as proposed by the second hypothesis. In this paper, we explore this possibility using a combination of empirical field data and mathematical modelling. We first test and find support for three key predictions of the female-quality hypothesis in wild chacma baboons (Papio ursinus): (i) inter-individual differences in swelling size are maintained across consecutive cycles, (ii) females in better condition have larger swellings and higher reproductive success, and (iii) males preferentially choose females with large swellings. We then develop an individual-based simulation model that indicates that females producing larger swellings can achieve higher mating success even when female–female competition is low and within-female variance in the trait is high. Taken together, our findings show that once sexual swellings have evolved as fertility signals, they might, in certain socio-sexual systems, be further selected to act as quality signals. These results, by reconciling two hypotheses, help to clarify the processes underlying sexual swelling evolution. More generally, our findings suggest that mate choice for direct benefits (fertility) can lead to indirect benefits (good genes).     

Did surface temperatures constrain microbial evolution?

The proposition that glaciation may not have occurred before the Cenozoic--albeit not yet a consensus position--nevertheless raises for reconsideration the surface temperature history of the earth. Glacial episodes, from the Huronian (2.3 billion years ago; BYA) through the late Paleozoic (320 to 250 million years ago; MYA) have been critical constraints on estimation of the upper bounds of temperature (Crowley 1983, Kasting and Toon 1989). Once removed, few if any constraints on the upper temperature limit other than life remain. Walker (1982) recognized that life provides an upper limit to temperature in the Precambrian. We propose a more radical concept: the upper temperature limit for viable growth of a given microbial group corresponds to the actual surface temperature at the time of the group's first appearance. In particular, we propose here that two major evolutionary developments--the emergence of cyanobacteria and aerobic eukaryotes--can be used to determine surface temperature in the Precambrian, and that only subsequent cooling mediated by higher plants and then angiosperms permitted what may possibly be the earth's first glaciation in the late Cenozoic.     

Can influenza epidemics be prevented by voluntary vaccination?

Previous modeling studies have identified the vaccination coverage level necessary for preventing influenza epidemics, but have not shown whether this critical coverage can be reached. Here we use computational modeling to determine, for the first time, whether the critical coverage for influenza can be achieved by voluntary vaccination. We construct a novel individual-level model of human cognition and behavior; individuals are characterized by two biological attributes (memory and adaptability) that they use when making vaccination decisions. We couple this model with a population-level model of influenza that includes vaccination dynamics. The coupled models allow individual-level decisions to influence influenza epidemiology and, conversely, influenza epidemiology to influence individual-level decisions. By including the effects of adaptive decision-making within an epidemic model, we can reproduce two essential characteristics of influenza epidemiology: annual variation in epidemic severity and sporadic occurrence of severe epidemics. We suggest that individual-level adaptive decision-making may be an important (previously overlooked) causal factor in driving influenza epidemiology. We find that severe epidemics cannot be prevented unless vaccination programs offer incentives. Frequency of severe epidemics could be reduced if programs provide, as an incentive to be vaccinated, several years of free vaccines to individuals who pay for one year of vaccination. Magnitude of epidemic amelioration will be determined by the number of years of free vaccination, an individuals' adaptability in decision-making, and their memory. This type of incentive program could control epidemics if individuals are very adaptable and have long-term memories. However, incentive-based programs that provide free vaccination for families could increase the frequency of severe epidemics. We conclude that incentive-based vaccination programs are necessary to control influenza, but some may be detrimental. Surprisingly, we find that individuals' memories and flexibility in adaptive decision-making can be extremely important factors in determining the success of influenza vaccination programs. Finally, we discuss the implication of our results for controlling pandemics.