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Sperm numbers vary between inter- and intra-population matings of the grasshopper Chorthippus parallelus.

Comparing the reproductive output of intra- and inter-population matings is the most common way to assess whether post-mating reproductive isolation is caused by genetic incompatibilities. Such genetic incompatibility can however, only assume that the quantity of the post-mating signals involved does not differ between intra- and inter-population matings. This assumption may not be true because sexual selection predicts reduced mating effort towards low-quality mates and in many circumstances, allopatric partners are low-quality mates. Post-mating efforts may, therefore, be reduced in inter- compared to intra-population matings. Here, I test this crucial assumption by studying variation in one post-mating trait, sperm number, in crosses of two parapatric grasshopper populations. In both populations, males transferred fewer sperm to allopatric than sympatric females. If such plasticity with respect to population is common in other post-mating traits, differences between inter- and intra-population crosses may be more frequently caused by differences in sperm number rather than gamete incompatibility. Additionally, I found that sperm numbers declined less rapidly in the female storage organ of allopatric than sympatric females but its rate differed markedly between populations. This is discussed with respect to female adaptations to male traits.

Animals↗

Bateman's principle and immunity.

The immunocompetence handicap hypothesis (ICHH) of Folstad and Karter has inspired a large number of studies that have tried to understand the causal basis of parasite-mediated sexual selection. Even though this hypothesis is based on the double function of testosterone, a hormone restricted to vertebrates, studies of invertebrates have tended to provide central support for specific predictions of the ICHH. I propose an alternative hypothesis that explains many of the findings without relying on testosterone or other biochemical feedback loops. This alternative is based on Bateman's principle, that males gain fitness by increasing their mating success whilst females increase fitness through longevity because their reproductive effort is much higher. Consequently, I predict that females should invest more in immunity than males. The extent of this dimorphism is determined by the mating system and the genetic correlation between males and females in immune traits. In support of my arguments, I mainly use studies on insects that share innate immunity with vertebrates and have the advantage that they are easier to study.

Animals↗

Rare male mating advantage for inversion karyotype in Drosophila ananassae.

These experiments were conducted to study frequency-dependent sexual selection on the alpha inversion in the left arm of the second chromosome (2L) in Drosophila ananassae. Two different strains, ST/ST-standard gene arrangement and AL/AL-alpha inversion in 2L were used. Female-choice experiments were carried out in five different male ratios. Two different methods of observation have been employed, one in which copulating pairs are aspirated out and the other, in which pairs are not removed. Results were analyzed by chi(2) tests, and log odds plus regression analyses. Regression analyses revealed the presence of one-sided rare male mating advantages for AL/AL males. Preferential mating is present within the strain having AL/AL karyotype. The minority male advantage and preferential mating found in the AL/AL strain are different components of mating success. ANOVA fails to show any significant difference between the outcomes of rare-male experiments using these two different methods.

Analysis of Variance↗

Differential male genetic success determines gene flow in an experimentally manipulated mouse population.

Sexual selection arises when genetically different males show heritable differences in reproductive success. Mouse mating behaviour involves both male competition and female choice. In this paper we show that introduced Y-linked DNA markers spread more extensively through a natural population than do genes inherited matrilineally. Differences in mating success between the sexes and among individual males may alter the pattern and rate of gene flow in natural populations. Another interesting possibility is that the success of the introduced Y chromosome may be attributable to so-called 'selfish' traits, such as sex-linked meiotic drive or intra-uterine competition. However, this study provides little unequivocal evidence to support this view. Differential success of introduced versus resident males may have implications for the reintroduction of endangered mammals into residual wild populations.

Animals↗

A novel song parameter correlates with extra-pair paternity and reflects male longevity.

Although elaborate bird song provides one of the prime examples of a trait that evolved under sexual selection, it is still unclear whether females judge the quality of males by attributes of their song and whether these song features honestly signal a male's genetic quality. We measured the ability of male dusky warblers Phylloscopus fuscatus to maintain a high sound amplitude during singing, which probably reflects an individual's physiological limitations. This new measure of singing performance was correlated with male longevity and with extra-pair paternity, indicating that females who copulated with better singers obtained 'good genes' for their offspring. Our findings are consistent with the idea that females assess male quality by subtle differences in their performance during the production of notes, rather than by the quantity or versatility of song. In addition, observations on territorial conflicts indicate that attractive males invest less in competition over territories because they can reproduce via extra-pair paternity.

Animals↗

A generalized female bias for long tails in a short-tailed widowbird.

Tail elongation in the polygynous widowbirds (Euplectes spp.) has evoked both adaptive and non-adaptive explanations. Female choice has been shown in the three longest tailed species (20-50 cm), whereas an agonistic function was proposed for a medium-tailed (10 cm) widowbird. To test the generality and directionality of sexual selection on tail length in widowbirds, we experimentally investigated selection in the relatively short-tailed (7 cm) red-shouldered widowbirds (E. axillaris). Prior to territory establishment, males were assigned to four tail-treatment groups; control, short, long and supernormal (similar to a sympatric long-tailed congener). No effects on male competition were detected as the groups were equally successful in acquiring territories of similar size and quality. However, mating success among the 92 territorial males was strongly skewed in favour of supernormal-tailed males (62% of active nests; 5.2 +/- 1.3 nests per territory). Long-tailed males also acquired more nests (1.9 +/- 0.7) than control (0.7 +/- 0.5) and short-tailed (0.5 +/- 0.3) males, while the latter two groups did not differ significantly. These results support a general, open-ended female preference for long tails in widowbirds and may represent a receiver bias that arose early in their divergence from the short-tailed weaverbirds (Ploceinae).

Animals↗

Premating, not postmating, barriers drive genetic dynamics in experimental hybrid populations of the endangered Sonoran topminnow.

The timing and pattern of reproductive barrier formation in allopatric populations has received much less attention than the accumulation of reproductive barriers in sympatry. The theory of allopatric speciation suggests that reproductive barriers evolve simply as by-products of overall genetic divergence. However, observations of enhanced premating barriers in allopatric populations suggest that sexual selection driven by intraspecific competition for mates may enhance species-specific signals and accelerate the speciation process. In a previous series of laboratory trials, we examined the strength of premating and postmating barriers in an allopatric species pair of the endangered Sonoran topminnow, Poeciliopsis occidentalis and P. sonoriensis. Behavioral observations provided evidence of asymmetrical assortative mating, while reduced brood sizes and male-biased F(1) sex ratios suggest postmating incompatibilities. Here we examine the combined effects of premating and postmating barriers on the genetic makeup of mixed populations, using cytonuclear genotype frequencies of first- and second-generation offspring. Observed genotype frequencies strongly reflect the directional assortative mating observed in behavioral trials, illustrating how isolating barriers that act earlier in the reproductive cycle will have a greater effect on total reproductive isolation and may be more important to speciation than subsequent postmating reproductive barriers.

Animals↗

Copulating with multiple mates enhances female fecundity but not egg-to-adult survival in the bruchid beetle Callosobruchus maculatus.

Postcopulatory sexual selection theory has come a long way since the evolutionary implications of sperm competition were first spelled out by Parker (1970). However, one of the most enduring questions remains: why do females copulate with multiple males? Here we show that females copulating with multiple males lay more eggs than those copulating repeatedly with the same male. We also show egg-to-adult survival to be more variable when females copulate multiply with different males and less variable when they copulate multiply with the same male. This supports the notion that egg-to-adult survival may depend on the genetic compatibility of males and females. However, pre-adult survival was highest when females copulated repeatedly with the same male rather than with different males. Thus, it would appear that polyandry in this species does not function to reduce the risk of embryo failure resulting from fertilization by genetically incompatible sperm.

Animals↗

Evolution of sexual isolation in laboratory populations: fitness differences between mating types and the associated hybrid incompatibilities.

Drosophila melanogaster is known to have two races in the incipient stages of speciation that exhibit strong asymmetric premating isolation: Zimbabwe (Z) and cosmopolitan (M). In a study examining the phenotypic and genotypic evolution after secondary contact, we found that despite strong sexual selection favoring the Z-type behavior, it is the M-type behavior that comes to dominate hybrid populations. This article examines the fitness costs associated with the Z-type behavior. We have discovered that these costs are great enough to explain the failure of the Z-type behavior to prosper. Here we report that Z-type females produce approximately half the number of offspring that M-type females produce. Furthermore, crosses between populations have revealed that Z-type females mated to M-type males have approximately 20% fewer offspring than the reciprocal crosses because of an inability of M-type sperm to successfully fertilize Z-type eggs. Hybrid crosses also exhibit much-reduced numbers of viable offspring in addition to reduced hybrid male fertility. These fitness effects suggest that multiple mechanisms of postmating isolation have evolved concurrently with the divergence in behavior.

Animals↗

Perception of female reproductive state from vocal cues in a mammal species.

While acoustic signalling by males is known to affect male-male competition, mate attraction and the timing of ovulation, the extent to which sexual selection has shaped the evolution of female acoustic signals is poorly understood. Among mammals, experimental evidence indicates that females attract mating partners by using olfactory and visual signals to advertise their reproductive state. Whether or not males ascertain female reproductive state from vocal signals has, however, never been systematically tested. In this study, we use playbacks of recorded vocalizations to demonstrate that male Barbary macaques, Macaca sylvanus, can discriminate between female copulation calls given at different stages of the oestrous cycle, responding more strongly to those given around the time when conception is most likely to occur. Acoustic analysis suggests that the mean dominant frequency of call units and a number of temporal parameters could provide males with the information necessary to discern the proximity of ovulation in this way Our results provide the first experimental evidence that the calls of female mammals may contain information on reproductive state, which males can perceive and use in such a way as to increase their reproductive success.

Animals↗

Chemical signal mediated premating reproductive isolation in a marine polychaete, Neanthes acuminata (arenaceodentata).

Neanthes acuminata Ehlers (1868) is a monogamous coastal polychaete with male parental care and a high level of sexual selection. We measured the level of prezygotic isolation among allopatric populations of N. acuminata; from the East and West Coast of the USA, a population from Hawaii, and a laboratory culture originating from Los Angeles, CA. All populations were found to preferably mate with members of their own population. Individuals from populations from Atlantic vs. Pacific Ocean failed to pair and to mate, either during the 10 min or 48 hr experiments. Instead, individuals showed high levels of aggressive behavior. Experiments measuring the levels of interpopulation aggression, established that individuals can recognize and discriminate among different populations of N. acuminata on the basis of olfactory cues. Aggressive behavior was induced by exposure of animals to seawater "conditioned" by individuals from the other populations, thus demonstrating the role of olfaction in the detection of "home" populations. The aggressive display was stronger upon exposure to seawater conditioned with "unrelated" populations and especially between Pacific and Atlantic populations.

Aggression↗

A female melanin ornament signals offspring fluctuating asymmetry in the barn owl.

Sexual selection theory predicts that males advertise quality by displaying extravagant ornaments. By contrast, whether phenotypic variation in females has a signalling function remains an open question. Here, to our knowledge, we provide the first evidence that a female plumage trait can signal fluctuating asymmetry in the offspring. We experimentally demonstrate in wild barn owls (Tyto alba) that the extent to which females display black spots on their plumage does not only signal offspring parasite resistance as shown in a previous study but also developmental homeostasis in the offspring. A greater number of spotted females produced offspring that had more symmetrical feathers during the period of growth. Males, that pair non-randomly with respect to female plumage spottiness therefore appear to gain substantial benefits by mating with heavily spotted females. Genetic variation in plumage spottiness is nevertheless maintained as the covariation between offspring body mass and mother plumage spottiness varies annually depending on environmental conditions.

Animals↗

Recognition of kin in bank voles (Clethrionomys glareolus).

The ability of animals to recognise their kin plays an important role in sexual selection. This study addressed the question of whether there is sibling recognition between young and adult bank vole females and males, and if so, how kin are recognised. The behavioural data from these experiments show that bank vole females can recognise their sibling brothers on the basis of their odours. In two-choice preference tests, prepubertal females were more attracted to the scent of socially known sibling and foster brothers than to the scent of unfamiliar males or of sibling brothers cross-fostered at birth and reared in a different nest. The females' odour preferences shifted with age. Mature females preferred the novel odour of unrelated males to that of sibling brothers regardless of whether the latter were reared with or apart from the tested females. These results indicate that two mechanisms of kin recognition co-exist in bank voles. Prepubertal females used a familiarity-based mechanism (recognition by social association) and adult used phenotype matching (genetic mechanism). The implications of these findings for bank vole biology are discussed.

Animals↗

Colourful male guppies produce faster and more viable sperm.

In guppies (Poecilia reticulata) precopulatory sexual selection (via female choice) and post-copulatory selection (via sperm competition) both favour males with relatively high levels of carotenoid (orange) pigmentation, suggesting that colourful males produce more competitive ejaculates. Here we test whether there is a positive association between male orange pigmentation and sperm quality. Our analysis of sperm quality focused on sperm swimming speeds (using CASA: computer-assisted sperm analysis to estimate three parameters of sperm velocity in vitro), sperm viability (proportion of live sperm per stripped ejaculate) and sperm lengths. We found that males with relatively large areas of orange pigmentation had significantly faster and more viable sperm than their less ornamented counterparts, suggesting a possible link between dietary carotenoid intake and sperm quality. By contrast, we found no relationship between sperm length (head length and total sperm length) and male phenotype. These findings, in conjunction with previous work showing that highly ornamented male guppies sire higher quality offspring, suggest that female preference for colourful males and sperm competition work in concert to favour intrinsically higher quality males.

Animal Communication↗

Tactic-specific success in sperm competition.

Sperm competition is a major force in sexual selection, but its implications for mating-system and life-history evolution are only beginning to be understood. The well-known sneak-guard model predicts that sneaks will win in sperm competition. We now provide empirical confirmation of this prediction. Bluegill sunfish (Lepomis macrochirus) have both sneak (cuckolder) and guard (parental) males. Guards make nests, court females and provide solitary parental care for the embryos. Sneaks include small cuckolders, which are termed 'sneakers', that dart in and out of nests in order to ejaculate between the spawning pair and larger cuckolders, which are termed 'satellites', that mimic females in order to ejaculate between the spawning pair. Using field behavioural data, genetic data and new mathematical models for paternity analyses, we show, for the first time to the authors' knowledge, that sneaks fertilize more eggs than guards during sperm competition. In addition, we show that sneakers are superior to satellites in sperm competition and, thus, that even among sneaks there are tactic-specific differences in competitive success.

Animals↗

Reversible frequency-dependent switches in male mate choice.

Current sexual-selection theories predict that mating should occur preferentially with the highest-quality partner, and assume that for distinguishing among potential mates the choosy sex applies an internal representation of the characteristics of the desired mate, i.e. a template. Binary choice experiments were performed to test male mate choice between two different female colour morphs in the damselfly Ischnura elegans. Choice experiments were conducted before and after an habituation period, during which males were exposed to only one female colour morph. Given the choice between the two female morphs, males did exhibit a choice for the most recently experienced female morph. This is the first evidence for a reversible switch in mate choice in a frequency-dependent way. In contrast with previous studies on mate choice, template formation in male I. elegans seems not to be based on quality. Switching mate choice in a frequency-dependent manner, choosing the most common morph, probably allows males to minimize their search efforts and to maximize fitness.

Animals↗

Nonlinear auditory mechanism enhances female sounds for male mosquitoes.

Sound plays an important role in the life history of mosquitoes. Male mosquitoes detect females by the sound generated by their wingbeat. Because female wings are weak acoustic radiators, males have been driven by sexual selection to evolve sensitive acoustic sensors. Mosquito antennae are very sensitive acoustic receivers, featuring up to 16,000 sensory cells, a number comparable with that contained in the human cochlea. The antennal sound receiver exhibits frequency selectivity, input amplification, and self-generated oscillations, features that parallel the functional sophistication of the cochlear amplifier. Although arguably the male antenna is well suited to receiving weak female sounds, the role of active mechanisms in mosquito hearing is far from understood. Previous mechanical studies on mosquito hearing largely focused on the steady-state antennal response to harmonic sounds, mostly evaluating the data through conventional Fourier transforms. Here, we report on the time-resolved mechanical behavior of the male antenna in response to female sounds. Crucially, stimuli were designed to reflect the temporal acoustic profile of a female flying by. With these stimuli, several previously unreported nonlinear features were unveiled, involving amplification, compression, and hysteresis. The time-resolved analysis reveals that, through the active participation of the sensory neurons, the antenna mechanically responds to enlarge its own range of detection. This behavior augments the capacity of the antennal receiver to detect female sounds, enhancing the male's chance to successfully pursue a passing female.

Acoustics↗

Sperm competitive ability in Drosophila melanogaster associated with variation in male reproductive proteins.

Multiple mating by females establishes the opportunity for postcopulatory sexual selection favoring males whose sperm is preferentially employed in fertilizations. Here we use natural variation in a wild population of Drosophila melanogaster to investigate the genetic basis of sperm competitive ability. Approximately 101 chromosome 2 substitution lines were scored for components of sperm competitive ability (P1', P2', fecundity, remating rate, and refractoriness), genotyped at 70 polymorphic markers in 10 male reproductive genes, and measured for transcript abundance of those genes. Permutation tests were applied to quantify the statistical significance of associations between genotype and phenotype. Nine significant associations were identified between polymorphisms in the male reproductive genes and sperm competitive ability and 13 were identified between genotype and transcript abundance, but no significant associations were found between transcript abundance and sperm competitive ability. Pleiotropy was evident in two genes: a polymorphism in Acp33A associated with both P1' and P2' and a polymorphism in CG17331 associated with both elevated P2' and reduced refractoriness. The latter case is consistent with antagonistic pleiotropy and may serve as a mechanism maintaining genetic variation.

Animals↗