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The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera.

Eyespots are found in a variety of animals, in particular lepidopterans. The role of eyespots as antipredator mechanisms has been discussed since the 19th Century, with two main hypotheses invoked to explain their occurrence. The first is that large, centrally located eyespots intimidate predators by resembling the eyes of the predators' own enemies; the second, though not necessarily conflicting, hypothesis is that small, peripherally located eyespots function as markers to deflect the attacks of predators to non-vital regions of the body. A third possibility is also proposed; that eyespots intimidate predators merely because they are novel or rarely encountered salient features. These hypotheses are reviewed, with special reference given to avian predators, since these are likely to be the principal visually hunting predators of the lepidopterans considered. Also highlighted is the necessity to consider the potential influence of sexual selection on lepidopteran wing patterns, and the genetics and development of eyespot formation.

Animals↗

Assortative mating and fertility in two Drosophila subobscura strains with different mitochondrial DNA haplotypes.

The mating pattern and female fertility on the two main mitochondrial DNA haplotypes (I and II) of Drosophila subobscura were studied, in an attempt to find possible differences between them in relation to sexual selection or isolation that could explain the populational dynamics and the co-existence of these two strains in nature. The mating pattern indicated an assortative mating in population cages, where couples of the same haplotype, mainly those of haplotype I, mated more often. However, the significations detected in laboratory conditions disappeared in wild populations, where random mating was the rule. The female fertility also showed differences in the laboratory compared to the wild, since couples with haplotype I males were more efficient in the laboratory populations. These results, together with others that we previously obtained, either point to selection acting directly on the mtDNA or to the presence of some kind of cytonuclear co-adaptation in these two haplotypes, although this must be modulated by other factors that change with the seasons and time. The end result could well be a balance of opposite forces acting on both haplotypes.

Animals↗

Condition-dependent signalling of genetic variation in stalk-eyed flies.

Handicap models of sexual selection predict that male sexual ornaments have strong condition-dependent expression and this allows females to evaluate male genetic quality. A number of previous experiments have demonstrated heightened condition-dependence of sexual ornaments in response to environmental stress. Here we show that genetic variation underlies the response to environmental stress (variable food quality) of a sexual ornament (male eye span) in the stalk-eyed fly Cyrtodiopsis dalmanni. Some male genotypes develop large eye span under all conditions, whereas other genotypes progressively reduce eye span as conditions deteriorate. Several non-sexual traits (female eye span, male and female wing length) also show genetic variation in condition-dependent expression, but their genetic response is entirely explained by scaling with body size. In contrast, the male sexual ornament still reveals genetic variation in the response to environmental stress after accounting for differences in body size. These results strongly support the hypothesis that female mate choice yields genetic benefits for offspring.

Animals↗

Polyandrous females avoid costs of inbreeding.

Why do females typically mate with more than one male? Female mating patterns have broad implications for sexual selection, speciation and conflicts of interest between the sexes, and yet they are poorly understood. Matings inevitably have costs, and for females, the benefits of taking more than one mate are rarely obvious. One possible explanation is that females gain benefits because they can avoid using sperm from genetically incompatible males, or invest less in the offspring of such males. It has been shown that mating with more than one male can increase offspring viability, but we present the first clear demonstration that this occurs because females with several mates avoid the negative effects of genetic incompatibility. We show that in crickets, the eggs of females that mate only with siblings have decreased hatching success. However, if females mate with both a sibling and a non-sibling they avoid altogether the low egg viability associated with sibling matings. If similar effects occur in other species, inbreeding avoidance may be important in understanding the prevalence of multiple mating.

Animals↗

Pigmentation and mate choice in Drosophila.

Many species of the fruitfly Drosophila are either sexually dimorphic for abdominal pigmentation (the posterior segments in males are black and those of females have thin dark stripes) or sexually monomorphic for this pigmentation (both sexes show striping). Kopp et al. report a correlation in two Drosophila clades between the expression of the bric-à-brac (bab) gene, which represses male-specific pigmentation in D. melanogaster females, and the presence of sexually dimorphic pigmentation. They suggest that sexual selection acted to produce sexual dichromatism in Drosophila by altering the regulation of bab, on the grounds that D. melanogaster males show a strong mate preference for females with lightly pigmented abdomens, and that this discrimination helps to maintain sexual dichromatism by preventing males from wasting time by courting other (darkly pigmented) males. Here we show that the mate discrimination observed by Kopp et al. may in fact have resulted from the nature of the strains and comparisons they used in their study and so could be irrelevant to mate choice in nature.

Abdomen↗

New perspectives on mate choice and the MHC.

A long series of studies on mice have shown that mate choice decisions can be made on the basis of individual genotype at the major histocompatibility complex (MHC), which accords well with the importance of immunocompetence in some theories of sexual selection. Recent work on other vertebrate species, including humans, indicates that MHC-based mate choice is not restricted to the genus Mus. However, its importance may vary among species as a result of differences in social and mating system structure, and perhaps genome structure. There appears to be a general preference expressed for MHC-dissimilar mates, and such MHC-disassortative mating may be involved in maintaining MHC and/or genome-wide diversity in natural populations. The strength and direction of MHC-based mating preference can vary, and may be modulated by factors such as genetic background, sex, and early life experience.

Animals↗

No evidence for parallel sympatric speciation in cichlid species of the genus Pseudotropheus from north-western Lake Malawi.

To test the hypothesis of parallel speciation by sexual selection, we examined length variation at six microsatellite loci of samples from four sites of four to six putative species belonging to two subgenera of rocky shore mbuna cichlids from Lake Malawi. Almost all fixation indices were significantly different from zero, suggesting that there is presently little or no gene flow among allopatric populations or sympatric species. Analysis of variance indicated that genetic distances among allopatric populations of putative conspecifics were significantly lower than among sympatric populations of heterospecifics. The topology of trees based on distance matrices was also largely consistent with the hypothesis that the putative species are monophyletic and have thus not evolved in parallel in their present locations. If parallel speciation does occur in Malawi cichlids, it may be on a larger spatial scale than investigated in our study.

Alleles↗

Parental investment, mate choice, and mate quality.

Current theory in sexual selection is extended to predict within-sex variability with regard to selectivity towards mates in different mating systems. Generally, the sex that invests more in the care of each offspring should be more selective of mates than the sex investing less. Within each sex, individuals of low male quality should be less selective than individuals of high quality, but there should be less variation in selectivity among individuals of the sex investing more. When only one sex contributes parental care, however, individuals of that sex should be uniformly selective, while the other sex is expected to mate indiscriminately. Using feral pigeons (Columba livia), these hypotheses are tested for the case in which both sexes contribute substantial parental care, but in which females contribute more than males. As predicted, females were found to be more selective of mates than males were. On certain criteria, males of lower quality were less selective of mates than males of higher quality.

Animals↗

Female preference for swords in Xiphophorus helleri reflects a bias for large apparent size.

Swordtail fish (Poeciliidae: genus Xiphophorus) are a paradigmatic case of sexual selection by sensory exploitation. Female preference for males with a conspicuous "sword" ornament is ancestral, suggesting that male morphology has evolved in response to a preexisting bias. The perceptual mechanisms underlying female mate choice have not been identified, complicating efforts to understand the selection pressures acting on ornament design. We consider two alternative models of receiver behavior, each consistent with previous results. Females could respond either to specific characteristics of the sword or to more general cues, such as the apparent size of potential mates. We showed female swordtails a series of computer-altered video sequences depicting a courting male. Footage of an intact male was preferred strongly to otherwise identical sequences in which portions of the sword had been deleted selectively, but a disembodied courting sword was less attractive than an intact male. There was no difference between responses to an isolated sword and to a swordless male of comparable length, or between an isolated sword and a homogenous background. Female preference for a sworded male was abolished by enlarging the image of a swordless male to compensate for the reduction in length caused by removing the ornament. This pattern of results is consistent with mate choice being mediated by a general preference for large males rather than by specific characters. Similar processes may account for the evolution of exaggerated traits in other systems.

Animals↗

A naked ape would have fewer parasites.

Unusually among the mammals, humans lack an outer layer of protective fur or hair. We propose the hypothesis that humans evolved hairlessness to reduce parasite loads, especially ectoparasites that may carry disease. We suggest that hairlessness is maintained by these naturally selected benefits and by sexual selection operating on both sexes. Hairlessness is made possible in humans owing to their unique abilities to regulate their environment via fire, shelter and clothing. Clothes and shelters allow a more flexible response to the external environment than a permanent layer of fur and can be changed or cleaned if infested with parasites. Naked molerats, another hairless and non-aquatic mammal species, also inhabit environments in which ectoparasite transmission is expected to be high, but in which temperatures are closely regulated. Our hypothesis explains features of human hairlessness-such as the marked sex difference in body hair, and its retention in the pubic regions-that are not explained by other theories.

Adaptation, Physiological↗

Good genes and the maternal effects of polyandry on offspring reproductive success in the bulb mite.

Genetic benefits are potentially the most robust explanation of the controversial issue of evolutionary maintenance of polyandry, but the unambiguous demonstration of such benefits has been hindered by the possibility of their confusion with maternal effects. Previous research has shown that polyandrous bulb mite females produce daughters with higher fecundity than monandrous females. Here, we investigate whether this effect arises because polyandrous females invest more in their offspring, or because their offspring inherit 'good genes' from their fathers. Females were mated with either one or four (different) males. However, by sterilizing three of the four males with ionizing radiation, we eliminated any chance of sexual selection (in the polyandrous treatment) so that any differences in the female mating regimes must have been owing to maternal effects. Polyandry had no significant effect on daughter fecundity, thus indicating that any previously documented effects must have been genetic. This was further supported by a significant association between fathers' offensive sperm-competitive ability and the fecundity of their daughters. The association with fathers' sperm defensive ability was not significant, and neither was the association between fathers' sperm competitiveness and sons' reproductive success. However, sons of polyandrous females had lower reproductive success than sons of monandrous females. This shows that the maternal effects of polyandry should be taken into account whenever its costs and benefits are being considered.

Analysis of Variance↗

Natural variation in male-induced 'cost-of-mating' and allele-specific association with male reproductive genes in Drosophila melanogaster.

One of the most sharply defined sexual conflicts arises when the act of mating is accompanied by an inflated risk of death. Several reports have documented an increased death rate of female Drosophila as a result of recurrent mating. Transgenic and mutation experiments have further identified components of seminal fluid that are at least in part responsible for this toxicity. Variation among males in their tendency for matings to be toxic to their partners has also been documented, but here for the first time we identify polymorphism within particular genes conferring differential post-mating female mortality. Such polymorphism is important, as it raises the challenge of whether sexual conflict models can provide means for maintenance of polymorphism. Using a set of second chromosome extraction lines, we scored differences in post-mating female fecundity and longevity subsequent to mating, and identified significant among-line differences. Seventy polymorphisms in ten male reproductive genes were scored and permutation tests were used to identify significant associations between genotype and phenotype. One polymorphism upstream of PEBII and an amino acid substitution in CG17331 were both associated with male-induced female mortality. The same allele of CG17331 that is toxic to females also induces greater refractoriness to remating in the females, providing an example of an allele-specific sexual conflict. Postcopulatory sexual selection could lead to sexual conflict by favouring males that prevent their mates from mating, even when there is a viability cost to those females.

Alleles↗

Species-specific genitalic copulatory courtship in sepsid flies (Diptera, Sepsidae, Microsepsis) and theories of genitalic evolution.

Males of Microsepsis eberhardi and M. armillata use their genitalic surstyli to rhythmically squeeze the female's abdomen with stereotyped movements during copulation. Squeezing movements did not begin until intromission had occurred and, contrary to predictions of the conflict-of-interest hypothesis for genitalic evolution, did not overcome morphological or behavioral female resistance. Contrary to predictions of the lock-and-key hypothesis, female morphology was uniform in the two species and could not mechanically exclude the genitalia of either species of male. The complex pattern of squeezing movements differed between the two species as predicted by the sexual selection hypothesis for genitalic evolution. Also, evolutionarily derived muscles and pseudoarticulations in the male's genitalic surstyli facilitated one type of movement, whose patterns were especially distinct. The data support the hypothesis that the male surstyli evolved to function as courtship devices.

Adaptation, Physiological↗

Multiple paternity in a natural population of a salamander with long-term sperm storage.

Sperm competition appears to be an important aspect of any mating system in which individual female organisms mate with multiple males and store sperm. Post-copulatory sexual selection may be particularly important in species that store sperm throughout long breeding seasons, because the lengthy storage period may permit extensive interactions among rival sperm. Few studies have addressed the potential for sperm competition in species exhibiting prolonged sperm storage. We used microsatellite markers to examine offspring paternity in field-collected clutches of the Ocoee salamander (Desmognathus ocoee), a species in which female organisms store sperm for up to 9 months prior to fertilization. We found that 96% of clutches were sired by multiple males, but that the majority of females used sperm from only two or three males to fertilize their eggs. The high rate of multiple mating by females suggests that sperm competition is an important aspect of this mating system. Comparison of our data with those of other parentage studies in salamanders and newts reveals that multiple mating may be common in urodele amphibians. Nevertheless, the number of males siring offspring per clutch in D. ocoee did not differ appreciably from that in other species of urodeles with shorter storage periods.

Animals↗

Microsatellite DNA analysis shows that greater sage grouse leks are not kin groups.

The spectacular social courtship displays of lekking birds are thought to evolve via sexual selection, but this view does not easily explain the participation of many males that apparently fail to mate. One of several proposed solutions to this 'lek skew paradox' is that kin selection favours low-ranking males joining leks to increase the fitness of closely related breeders. We investigated the potential for kin selection to operate in leks of the greater sage grouse, Centrocercus urophasianus, by estimating relatedness between lekking males using microsatellite DNA markers. We also calibrated these estimates using data from known families. Mean relatedness within leks was statistically indistinguishable from zero. We also found no evidence for local clustering of kin during lek display, although males tended to range closer to kin when off the lek. These results make kin selection an unlikely solution to the lek skew paradox in sage grouse. Together with other recent studies, they also raise the question of why kin selection apparently promotes social courtship in some lekking species, but not in others.

Animals↗

Divergence in replicated phylogenies: the evolution of partial post-mating prezygotic isolation in bean weevils.

By tradition, speciation research has been focused on processes leading to either premating or post-zygotic reproductive isolation. The processes which generate isolation after mating but before zygote formation are less well understood. Here, we study divergence in characters which contribute to post-mating prezygotic isolation, such as egg production and remating rate. We propose that 'replicated' laboratory phylogenies with known histories can be used to yield insights into the processes of divergence. We performed a series of cross-matings between populations within two strains of the bean weevil Callosobruchus maculatus. Each strain has a unique and independent origin and both have been kept in the same set of laboratories during the last few decades. Our results show that divergence has occurred between laboratory populations within strains with regards to the effects that mating has on female reproductive behaviour, showing that the evolution of partial post-mating prezygotic isolation can be rapid. More importantly, the pattern of divergence across populations was distinct in the two strains, suggesting that coevolutionary trajectories are not determined by environmental factors but are to some extent arbitrary. We discuss the limitations of the novel empirical strategy employed here, and conclude that our results lend support to the hypothesis that post-mating sexual selection is capable of rapidly generating post-mating prezygotic isolation.

Animals↗

Inheritance of song and stridulatory peg number divergence between Chorthippus brunneus and C. jacobsi, two naturally hybridizing grasshopper species (Orthoptera: Acrididae).

Knowledge of the genetic basis of divergence in mating signal characters that contribute to reproductive isolation is critical to understanding speciation. Here, we describe a semi-automated system for characterizing grasshopper acoustic signals. We used this system to study the genetic basis of divergence in three male calling song components [echeme (EL), syllable (SL) and phrase (PL) lengths] between Chorthippus brunneus and C. jacobsi, two species of grasshoppers that hybridize in northern Spain. We also studied the number of pegs in the stridulatory file. For all characters, additive effects accounted for most of the genetic differentiation between species. However, the three song components also showed small but significant epistatic effects. No sex linkage was detected. Wright-Castle-Lande estimates of the minimum numbers of genetic factors underlying song and peg number divergence were low: peg number (n(e)=5.87+/-5.84), SL (n(e)=2.37+/-4.79) and PL (n(e)=0.87+/-0.86). On the other hand, EL appeared to be controlled by many genes. These results suggest that divergence in SL and PL might be driven by sexual selection whereas EL might not be under selection. This is consistent with experimental results on female song preference in related species. However, the fact that few factors appear to underlie the differences in peg number is surprising. Peg number is not closely related to song characteristics. It often varies between closely related grasshopper species and it has been assumed to be a neutral character. The biometrical approaches used here tend to underestimate the number of factors influencing a trait but provide valuable background for subsequent quantitative trait loci analyses.

Animal Communication↗