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Honesty, perception and population divergence in sexually selected traits.

We investigate the evolution of female preference for one and two male ornaments, to address two issues in sexual selection: (i) what factors affect the evolution of female preferences; and (ii) how do preferences diverge between isolated populations, leading to speciation? We assume that the male traits are costly indicators of male condition ('handicaps'), and that females benefit directly from a high-condition mate. We find that optimal female preference for a single male trait equals (benefit of condition) x (detectability of male trait) x (honesty of male trait)/(costliness of preference). With two male traits to choose from, females should prefer the one with greatest honesty x detectability, and ignore the second. These results highlight the role of perception in the evolution of both male ornaments and female preferences, and provide a theoretical illustration of 'sensory drive'. They confirm that a less honest male trait can displace a more honest trait if its detectability is sufficiently high. Environmental differences can drive evolutionary divergence between populations in both the male trait and female preference. Even small differences between habitats in detectability of male traits can trigger dramatic change in the female preference. Finally, populations may drift apart in arbitrary directions if choice of different male traits yields equivalent benefits to females.

Animals↗

Genetic association between male attractiveness and female differential allocation.

Differential allocation of reproductive effort towards offspring of attractive mates is a form of post-copulatory mate choice. Although differential allocation has been demonstrated in many taxa, its evolutionary implications have received little attention. Theory predicts that mate choice will lead to a positive genetic correlation between female preference and male attractiveness. This prediction has been upheld for pre-copulatory mate choice, but whether such a relationship between male attractiveness and female differential allocation exists has never been tested. Here, we show that both female pre-copulatory mate choice and post-copulatory differential allocation are genetically associated with male attractiveness in house crickets, Acheta domesticus. Daughters of attractive males mated sooner and laid more eggs when paired with larger males. These forms of mate choice are strongest in large females, suggesting that costs decrease with increasing female size. The genetic association between attractiveness and differential allocation suggests potential for differential allocation to become exaggerated by coevolutionary runaway processes in an analogous manner to pre-copulatory choice. Sexual selection is thus likely to be stronger than predicted by pre-copulatory choice alone.

Animals↗

Sexual conflict and the evolution of female mate choice and male social dominance.

Conflicts between the sexes over control of reproduction are thought to lead to a cost of sexual selection through the evolution of male traits that manipulate female reproductive physiology and behaviour, and female traits that resist this manipulation. Although studies have begun to document negative fitness effects of sexual conflict, studies showing the expected association between sexual conflict and the specific behavioural mechanisms of sexual selection are lacking. Here we experimentally manipulated the opportunity for sexual conflict in the cockroach. Nauphoeta cinerea and showed that, for this species, odour cues in the social environment influence the behavioural strategies and fitness of males and females during sexual selection. Females provided with the opportunity for discriminating between males but not necessarily mating with preferred males produced fewer male offspring than females mated at random. The number of female offspring produced was not affected, nor was the viability of the offspring. Experimental modification of the composition of the males' pheromone showed that the fecundity effects were caused by exposure to the pheromone component that makes males attractive to females but also makes males less likely to be dominant. Female mate choice therefore carries a demographic cost but functions to avoid male manipulation and aggression. Male-male competition appears to function to circumvent mate choice rather than directly manipulating females, as the mate choice can be cryptic. The dynamic struggle between the sexes for control of mating opportunities and outcomes in N. cinerea therefore reveals a unique role for sexual conflict in the evolution of the behavioural components of sexual selection.

Animals↗

Sexual selection and speciation in mammals, butterflies and spiders.

Recently refined evolutionary theories propose that sexual selection and reproductive conflict could be drivers of speciation. Male and female reproductive optima invariably differ because the potential reproductive rate of males almost always exceeds that of females: females are selected to maximize mate 'quality', while males can increase fitness through mate 'quantity'. A dynamic, sexually selected conflict therefore exists in which 'competitive' males are selected to override the preference tactics evolved by 'choosy' females. The wide variation across taxa in mating systems therefore generates variance in the outcome of intrasexual conflict and the strength of sexual selection: monandry constrains reproductive heterozygosity and allows female choice to select and maintain particular (preferred) genes; polyandry promotes reproductive heterozygosity and will more likely override female choice. Two different theories predict how sexual selection might influence speciation. Traditional ideas indicate that increased sexual selection (and hence conflict) generates a greater diversity of male reproductive strategies to be counteracted by female mate preferences, thus providing elevated potentials for speciation as more evolutionary avenues of male-female interaction are created. A less intuitively obvious theory proposes that increased sexual selection and conflict constrains speciation by reducing the opportunities for female mate choice under polyandry. We use a comparative approach to test these theories by investigating whether two general measures of sexual selection and the potential for sexual conflict have influenced speciation. Sexual size dimorphism (across 480 mammalian genera, 105 butterfly genera and 148 spider genera) and degree of polyandry (measured as relative testes size in mammals (72 genera) and mating frequency in female butterflies (54 genera)) showed no associations with the variance in speciosity. Our results therefore show that speciation occurs independently of sexual selection.

Animals↗

Worker interests and male production in Polistes gallicus, a Mediterranean social wasp.

The resolution of social conflict in colonies may accord with the interests of the most numerous party. In social insect colonies with single once-mated queens, workers are more closely related to the workers' sons than they are to the queens' sons. Therefore, they should prefer workers to produce males, against the queen's interests. Workers are capable of producing males as they arise from unfertilized eggs. We found Polistes gallicus to have colonies of single, once-mated queens, as determined by microsatellite genotyping of the workers, so worker interests predict worker male production. In colonies lacking queens, workers produced the males, but not in colonies with original queens. Thus worker interests were expressed only when the queen was gone. The high fraction of missing queens and early end to the colony cycle relative to climate so early in the season is surprising and may indicate a forceful elimination of the queen.

Animals↗

Effects of central vasotocin and mesotocin manipulations on social behavior in male and female zebra finches.

Male and female zebra finches (Taeniopygia guttata; total n = 40) were fitted with chronic guide cannulae directed at the lateral ventricle and were tested for aggression, affiliation, and partner preference following infusions of mesotocin (MT), vasotocin (VT), their antagonists, and vehicle control. Aggressive behavior was tested in a mate competition paradigm and tests of intersexual affiliation and partner preference were conducted following 1 day of cohabitation with an opposite-sex individual. These tests also provided data on male courtship singing. The results demonstrate a modest dose-dependent facilitation of aggression by VT, but not MT, in both male and female finches. However, only males were sensitive to infusions of a vasopressin antagonist, suggesting that endogenous VT is more important for behavioral modulation in males. Peptide effects were specific to aggression, as no treatments influenced intersexual affiliation, partner preference, or male courtship singing. Thus, in contrast to rodents, partner preference is not readily induced by VT or MT in this species. However, the potential necessity of endogenous VT and MT for natural pair-bond formation remains to be tested.

Aggression↗

Change of mate and mating competition between males of Schistosoma intercalatum and S. mansoni.

Previous studies on mating behaviour of Schistosoma intercalatum and S. mansoni, two human schistosomes which may overlap in parts of Africa, have shown that in mixed infections in mice there are no physiological barriers preventing heterospecific pairings. However, when choice is possible, matings occur preferentially between partners of the same species. In this paper, further experimental studies on mating behaviour of the two species were conducted. Sequential infections showed that heterospecific pairs of worms change partners to homospecific pairs when given the opportunity. The change of mate is a progressive process requiring up to, at least 8 weeks, and this phenomenon is due to the male worm seeking the appropriate female. S. mansoni males are better at pairing with females than S. intercalatum males, and they will change partner to pair with homologous females in preference to heterologous females whenever given the opportunity. Moreover, in the absence of S. mansoni female worms, unpaired S. mansoni male worms pull away female S. intercalatum from male S. intercalatum. It appears from this study that S. mansoni males are much more competitive than S. intercalatum males at pairing with females, and this is a disadvantage for S. intercalatum in situations of sympatry.

Animals↗

Male ornament size as a reliable cue to enhanced offspring viability in the barn swallow.

Many extravagant secondary sexual characters are assumed to have evolved as a result of female choice, either because they attract females or because they reliably reflect the quality of males. Females mating with the most ornamented individuals with a superior genotype are expected to benefit by producing more viable offspring. A viability advantage associated with mate choice can be demonstrated only if (i) parent ornament size reliably reflects parent viability and (ii) offspring viability is directly related to the expression of the ornament of the parent. Barn swallows (Hirundo rustica) are monogamous passerine birds, which are sexually size dimorphic in tail length. Previous experiments and observations have shown that females prefer males with the largest tail ornaments and that male survivors have larger tail ornaments than nonsurvivors. Here I demonstrate that offspring longevity is positively related to ornament size of the male parent and that the longevity of sons is a trait with a statistically significant resemblance to that of their fathers. The viability effects could be entirely due to differences in quality of parental care. However, relative paternal provisioning of offspring was negatively related to the tail length of males, while total provisioning rate by both pair members, and thus offspring body size, body mass, and body condition, was unrelated to male tail length. Therefore, females may, through their mate choice, gain an indirect fitness advantage in terms of enhanced offspring viability.

Animals↗

Genetic and behavioral components of the cryptic species boundary between Laupala cerasina and L. kohalensis (Orthoptera: Gryllidae).

Cryptic species are often hypothesized on the basis of differences in courtship signals. These signal differences suggest that mate recognition systems, which include both courtship signals and responses to those signals, have diverged between genetically isolated populations. Cryptic species are therefore thought to represent distinct genetic units, the boundaries of which are maintained by premating incompatibilities, specifically by receiver preferences for conspecific signals. Laupala cerasina and L. kohalensis are sympatric species of swordtail crickets endemic to the big island of Hawaii, that are distinguishable by differences in male courtship song. We first tested whether groupings hypothesized by acoustic similarity reflect genetic groupings, using AFLP data to estimate genetic relationships. Second, we tested whether genetic boundaries are maintained by female preferences for conspecific song characteristics. Phonotaxis trials were used to determine the extent of female preferences for conspecific male song. Results generally support both hypotheses, but suggest the presence of porous species boundaries.

Acoustic Stimulation↗

Geographic variation of genetic and behavioral traits in northern and southern tüngara frogs.

We use a combination of microsatellite marker analysis and mate-choice behavior experiments to assess patterns of reproductive isolation of the túngara frog Physalaemus pustulosus along a 550-km transect of 25 populations in Costa Rica and Panama. Earlier studies using allozymes and mitochondrial DNA defined two genetic groups of túngara frogs, one ranging from Mexico to northern Costa Rica (northern group), the second ranging from Panama to northern South America (southern group). Our more fine-scale survey also shows that the northern and southern túngara frogs are genetically different and geographically separated by a gap in the distribution in central Pacific Costa Rica. Genetic differences among populations are highly correlated with geographic distances. Temporal call parameters differed among populations as well as between genetic groups. Differences in calls were explained better by geographic distance than by genetic distance. Phonotaxis experiments showed that females preferred calls of males from their own populations over calls of males from other populations in about two-thirds to three-fourths of the contrasts tested. In mating experiments, females and males from the same group and females from the north with males from the south produced nests and tadpoles. In contrast, females from the south did not produce nests or tadpoles with males from the north. Thus, northern and southern túngara frogs have diverged both genetically and bioacoustically. There is evidence for some prezygotic isolation due to differences in mate recognition and fertilization success, but such isolation is hardly complete. Our results support the general observation that significant differences in sexual signals are often not correlated with strong genetic differentiation.

Animals↗

Fitness effects of female mate choice: preferred males are detrimental for Drosophila melanogaster females.

The evolution of female mate choice, broadly defined to include any female behaviour or morphology which biases matings towards certain male phenotypes, is traditionally thought to result from direct or indirect benefits which females acquire when mating with preferred males. In contrast, new models have shown that female mate choice can be generated by sexual conflict, where preferred males may cause a fitness depression in females. Several studies have shown that female Drosophila melanogaster bias matings towards large males. Here, we use male size as a proxy for male attractiveness and test how female fitness is affected by reproducing with large or small males, under two different male densities. Females housed with large males had reduced lifespan and aged at an accelerated rate compared with females housed with small males, and increased male density depressed female fitness further. These fitness differences were due to effects on several different fitness components. Female fitness covaried negatively with male courtship rate, which suggests a cost of courtship. Mating rate increased with male size, whereas female fitness peaked at an intermediate mating rate. Our results suggest that female mate choice in D. melanogaster is, at least in part, a by-product of sexual conflict over the mating rate.

Animals↗

Evolutionary history of the hybridogenetic hybrid frog Rana esculenta as deduced from mtDNA analyses.

mtDNA of the hybridogenetic hybrid frog Rana esculenta from Switzerland, Austria, and Poland was compared to mtDNA of the parental species R. ridibunda and R. lessonae using electrophoretic analysis of restriction enzyme fragments. Two mtDNA phenotypes, with 3.4% sequence divergence, are present in R. lessonae: type C is found in Poland, and type D is found in Switzerland. Rana ridibunda from Poland has either of two mtDNA phenotypes: type A is the typical ridibunda mtDNA, and type B is a lessonae mitochondrial genome, introgressed into R. ridibunda, that differs from type C mtDNA of R. lessonae by only 0.3%. Each of the three lessonae genomes differs from A, the typical ridibunda mtDNA, by approximately 8%. All four types of mtDNA (A and B of R. ridibunda, C and D of R. lessonae) are found in R. esculenta. Of 62 R. esculenta from Poland, 58 had type C, three had type A, and one had type B mtDNA. All nine R. esculenta from Switzerland had type D mtDNA. All three R. esculenta from Austria, from a population in which males of R. esculenta are rare, had ridibunda mtDNA, two having type B and one having type A. Both field observations and studies of mating preference indicate that the primary hybridizations that produce R. esculenta are between R. ridibunda females and R. lessonae males; thereafter, R. esculenta lineages are usually maintained by matings of R. esculenta females with R. lessonae males. The presence of ridibunda mtDNA in the three R. esculenta sampled from Austria, its occasional presence in R. esculenta populations in Poland, and its absence from R. esculenta in Switzerland support both the direction of the original hybridization and the rarity of formation of new R. esculenta lineages. The preponderance of R. esculenta individuals with lessonae mtDNA in our samples from central Europe suggests that most lineages have gone through at least one mating between an R. lessonae female and an R. esculenta male. This reveals a greater reproductive role for R. esculenta males than their partial sterility and infrequent matings would suggest.

Animals↗

Reinforcement and sex linkage.

We present a general model for the effect of sex linkage on the evolution of reinforcement of mating preferences on an island. We find that the level of reinforcement can vary up to 80% depending on the mode of inheritance of the female preference and male trait. When reinforcement is driven mainly by selection in the male trait and intrinsic hybrid incompatibilities are weak, sex-linked preferences and autosomal male traits are the most conducive to reinforcement, whereas autosomal preferences and X-linked traits are the least. Surprisingly, the effect of mode of inheritance on reinforcement is poorly predicted by its effect on the genetic correlation between the male trait and female preference. Sex-linkage of genetic incompatibility loci increases reinforcement, though this is not due solely to the occurrence of Haldane's rule. We find that reinforcement can lead to complete reproductive isolation in some cases but not others and that the mode of inheritance can determine which outcome occurs.

Animals↗

Cultural transmission of vocal traditions in cowbirds (Molothrus ater) influences courtship patterns and mate preferences.

In this study, the authors tested the cultural transmission of vocal traditions in cowbirds (Molothrus ater). Young cowbirds from a South Dakota (SD) population were housed over winter with adults of the SD population or with adults from an Indiana (IN) population. Song differences between the original SD and IN adult models were acquired by South Dakota culture (SDC) and Indiana culture (INC) males, respectively, and were transmitted to a 2nd cultural generation of birds. During playback tests of SDC and INC songs, SD females gave more copulatory responses to SDC songs. Finally, males with SD-like songs courted SDC females preferentially in breeding season tests, whereas males with IN-like songs courted INC females preferentially. These results indicate that the transmission of vocal traditions plays a fundamental role in the courtship patterns and mating decisions of cowbirds.

Animals↗

Sex and context: hormones and primate sexual motivation.

Gonadal hormones regulate the ability to copulate in most mammalian species, but not in primates because copulatory ability has been emancipated from hormonal control. Instead, gonadal hormones primarily influence sexual motivation. This separation of mating ability from hormonally modulated mating interest allows social experience and context to powerfully influence the expression of sexual behavior in nonhuman primates, both developmentally and in adulthood. For example, male rhesus monkeys mount males and females equally as juveniles, but mount females almost exclusively as adults. Having ejaculated with a female better predicted this transition to female mounting partners than did increased pubertal testosterone (T). It is proposed that increased pubertal T stimulates male sexual motivation, increasing the male's probability of sexual experience with females, ultimately producing a sexual preference for females. Eliminating T in adulthood reduces male sexual motivation in both humans and rhesus monkeys, but does not eliminate the capacity to engage in sex. In male rhesus monkeys the effects of reduced androgens on sexual behavior vary with social status and sexual experience. Human sexual behavior also varies with hormonal state, social context, and cultural conventions. Ovarian hormones influence female sexual desire, but the specific sexual behaviors engaged in are affected by perceived pregnancy risk, suggesting that cognition plays an important role in human sexual behavior. How the physical capacity to mate became emancipated from hormonal regulation in primates is not understood. This emancipation, however, increases the importance of motivational systems and results in primate sexual behavior being strongly influenced by social context.

Animals↗

Ectoparasites of the whitespotted rabbitfish, Siganus sutor (Valenciennes, 1835) off the Kenyan Coast: distribution within the host population and site selection on the gills.

Different populations of the whitespotted rabbitfish, Siganus sutor, were examined for ectoparasites: adults from the Mombasa area (sampled in December 1990) and different age classes (adult, subadult and juveniles) from Gazi Bay (sampled in December 1992 and August 1993). The most common gill parasites were: the monogeneans Pseudohaliotrema sp., Tetrancistrum sigani and Microcotyle mouwoi, the copepods Hatschekia sp., Pseudolepeophtheirus sp. and juvenile Caligidae, and prazina larvae of the isopod Gnathia sp. Adult siganids had a higher parasite load than subadults. Juvenile rabbitfish did not harbour any gill parasites. Temporal differences in the parasite load of subadult rabbitfish were observed for M. mouwoi (highest in the December samples) and for juvenile Caligidae (highest in August). The microhabitat of the 5 most common gill parasites was species specific. Most parasite species showed distinct site preferences with respect to both gill arches and gill sectors, within the gill arches. Niche breadth of the different gill parasite species was independent of the abundance of any of the other species present. However, niche breadths of M. mouwoi, Tetrancistrum sp. and Hatschekia sp. increased with their own abundance. This suggests that interspecific competition for space is low and that intraspecific factors could play an important role in the microhabitat choice of these gill parasites. The hypothesis that niche restriction leads to higher intraspecific contact and an enhancement of chances to mate was tested on 2 monogenean species, Pseudohaliotrema sp. and Tetrancistrum sigani. Their highly aggregated distribution over the gill filaments, leading to increased intraspecific contact, is consistent with the hypothesis.

Animals↗

Simultaneous hermaphrodites reproducing in pairs self-fertilize some of their eggs: an experimental test of predictions of mixed-mating and Hermaphrodite's Dilemma theory.

Theory predicts (1) that mixed-mating systems (i.e. reproduction through both selfing and outcrossing) should usually not evolve and (2) that reproducing simultaneous hermaphrodites should be in a conflict over the preferred sexual role (The Hermaphrodite's Dilemma). In an in vitro system with the endoparasitic cestode Schistocephalus solidus, a simultaneous hermaphrodite, we tested predictions of both the mixed-mating and the Hermaphrodite's Dilemma theory. Using microsatellite markers, we measured the proportion of selfed offspring and the total reproductive output of each worm within pairs varying in mean weight and weight difference. Worms produced more outbred offspring not only with increasing total weight of the pair, but also with decreasing weight difference between the two paired worms. These results suggest: (1) that this parasite species reproduces by mixed-mating, which may be maintained by stochastic density fluctuations in the definitive host and hence unpredictability of self reproduction and (2) reproductive conflict may prevent worm pairs from achieving an optimal intermediate selfing rate.

Animals↗

Female preference for multiple partners: sperm competition in the hide beetle, Dermestes maculatus (DeGeer).

Explanations for the evolution of polyandry emphasize either direct benefits or indirect genetic benefits. The former class of explanation predicts that females may seek multiple copulations but not necessarily multiple partners, while the latter predicts that females will seek multiple copulations with different partners. We investigated these predictions experimentally in the hide beetle, in which females mate more than once during their reproductive cycle. Female hide beetles remated more readily with novel males than with their previous partners, suggesting they mate multiply in order to obtain indirect, genetic benefits. In double-mating experiments with sterile males, the proportion of eggs fertilized by the second male was highly variable; this could not be explained by variation in male body size, duration of copulation or mating order. Male hide beetles form postinsemination associations by riding on the backs of females. This behaviour was more prevalent in the presence of rival males and less prevalent in the presence of additional females, suggesting that the behaviour serves to increase paternity, perhaps by obtaining additional copulations. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗