How general is a female mating preference for clustered males in lekking species? a meta-analysis

A principal hypothesis for the evolution of leks (rare and intensely competitive territorial aggregations) is that leks result from females preferring to mate with clustered males. This hypothesis predicts more female visits and higher mating success per male on larger leks. Evidence for and against this hypothesis has been presented by different studies, primarily of individual populations, but its generality has not yet been formally investigated. We took a meta-analytical approach towards formally examining the generality of such a female bias in lekking species. Using available published data and using female visits as an index of female mating bias, we estimated the shape of the relationship between lek size and total female visits to a lek, female visits per lekking male and, where available, per capita male mating success. Individual analyses showed that female visits generally increased with lek size across the majority of taxa surveyed; the meta-analysis indicated that this relationship with lek size was disproportionately positive. The findings from analysing per capita female visits were mixed, with an increase with lek size detected in half of the species, which were, however, widely distributed taxonomically. Taken together, these findings suggest that a female bias for clustered males may be a general process across lekking species. Nevertheless, the substantial variation seen in these relationships implies that other processes are also important. Analyses of per capita copulation success suggested that, more generally, increased per capita mating benefits may be an important selective factor in lek maintenance.

No evidence for a genetic association between female mating preference and male secondary sexual trait in a Lake Victoria cichlid fish

Sexual selection by female mating preference for male nuptial coloration has been suggested as a driving force in the rapid speciation of Lake Victoria cichlid fish. This process could have been facilitated or accelerated by genetic associations between female preference loci and male coloration loci. Preferences, as well as coloration, are heritable traits and are probably determined by more than one gene. However, little is known about potential genetic associations between these traits. In turbid water, we found a population that is variable in male nuptial coloration from blue to yellow to red. Males at the extreme ends of the phenotype distribution resemble a reproductively isolated species pair in clear water that has diverged into one species with blue-grey mates and one species with bright red males. Females of the turbid water population vary in mating preference coinciding with the male phenotype distribution. For the current study, these females were mated to blue males. We measured the coloration of the sires and male offspring. Parents-offspring regression showed that the sires did not affect male offspring coloration, which confirms earlier findings that the blue species breeds true. In contrast, male offspring coloration was determined by the identity of the dams, which suggests that there is heritable variation in male color genes between females. However, we found that mating preferences of the dams were not correlated with male offspring coloration. Thus, there is no evidence for strong genetic linkage between mating preference and the preferred trait in this population [Current Zoology 56 (1): 57-64 2010].

Mapping Individual Variation in Male Mating Preference Space: Multiple Choice in a Color Polymorphic Cichlid Fish

Sexual selection theory largely rests on the assumption that populations contain individual variation in mating preferences and that individuals are consistent in their preferences. However, there are few empirical studies of within-population variation and even fewer have examined individual male mating preferences. Here, we studied a color polymorphic population of the Lake Victoria cichlid fish Neochromis omnicaeruleus, a species in which color morphs are associated with different sex-determining factors. Wild-caught males were tested in three-way choice trials with multiple combinations of different females belonging to the three color morphs. Compositional log-ratio techniques were applied to analyze individual male mating preferences. Large individual variation in consistency, strength, and direction of male mating preferences for female color morphs was found and hierarchical clustering of the compositional data revealed the presence of four distinct preference groups corresponding to the three color morphs in addition to a no-preference class. Consistency of individual male mating preferences was higher in males with strongest preferences. We discuss the implications of these findings for our understanding of the mechanisms underlying polymorphism in mating preferences.

A test of genetic association among male nuptial coloration, female mating preference, and male aggression bias within a polymorphic population of cichlid fish

Both inter- and intrasexual selection have been implicated in the origin and maintenance of species-rich taxa with diverse sexual traits. Simultaneous disruptive selection by female mate choice and male-male competition can, in theory, lead to speciation without geographical isolation if both act on the same male trait. Female mate choice can generate discontinuities in gene flow, while male-male competition can generate negative frequency-dependent selection stabilizing the male trait polymorphism. Speciation may be facilitated when mating preference and/or aggression bias are physically linked to the trait they operate on. We tested for genetic associations among female mating preference, male aggression bias and male coloration in the Lake Victoria cichlid Pundamilia. We crossed females from a phenotypically variable population with males from both extreme ends of the phenotype distribution in the same population (blue or red). Male offspring of a red sire were significantly redder than males of a blue sire, indicating that intra-population variation in male coloration is heritable. We tested mating preferences of female offspring and aggression biases of male offspring using binary choice tests. There was no evidence for associations at the family level between female mating preferences and coloration of sires, but dam identity had a significant effect on female mate preference. Sons of the red sire directed significantly more aggression to red than blue males, whereas sons of the blue sire did not show any bias. There was a positive correlation among individuals between male aggression bias and body coloration, possibly due to pleiotropy or physical linkage, which could facilitate the maintenance of color polymorphism.

Inheritance of female mating preference in a sympatric sibling species pair of Lake Victoria cichlids: implications for speciation

Female mate choice has often been proposed to play an important role in cases of rapid speciation, in particular in the explosively evolved haplochromine cichlid species flocks of the Great Lakes of East Africa. Little, if anything, is known in cichlid radiations about the heritability of female mating preferences. Entirely sympatric distribution, large ecological overlap and conspicuous differences in male nuptial coloration, and female preferences for these, make the sister species Pundamilia pundamilia and P. nyererei from Lake Victoria an ideally suited species pair to test assumptions on the genetics of mating preferences made in models of sympatric speciation. Female mate choice is necessary and sufficient to maintain reproductive isolation between these species, and it is perhaps not unlikely therefore, that female mate choice has been important during speciation. A prerequisite for this, which had remained untested in African cichlid fish, is that variation in female mating preferences is heritable. We investigated mating preferences of females of these sister species and their hybrids to test this assumption of most sympatric speciation models, and to further test the assumption of some models of sympatric speciation by sexual selection that female preference is a single-gene trait. We find that the differences in female mating preferences between the sister species are heritable, possibly with quite high heritabilities, and that few but probably more than one genetic loci contribute to this behavioural speciation trait with no apparent dominance. We discuss these results in the light of speciation models and the debate about the explosive radiation of cichlid fishes in Lake Victoria.

Female mating preference for bold males in the guppy, Poecilia reticulata.

Although females prefer to mate with brightly colored males in numerous species, the benefits accruing to such females are virtually unknown. According to one hypothesis of sexual selection theory, if the expression of costly preferred traits in males (such as conspicuous colors) is proportional to the male's overall quality or reveals his quality, a well-developed trait should indicate good condition and/or viability for example. A female choosing such a male would therefore stand to gain direct or indirect fitness benefits, or both. Among potential phenotypic indicators of an individual's quality are the amount and brightness of its carotenoid-based colors and its boldness, as measured by its willingness to risk approaching predators without being killed. Here, we show experimentally that in the Trinidadian guppy (Poecilia reticulata) the visual conspicuousness of the color pattern of males correlates positively with boldness toward, and with escape distance from, a cichlid fish predator. Bold individuals are thus more informed about nearby predators and more likely to survive encounters with them. Mate-choice experiments showed that females prefer colorful males as mates, but prefer bolder males irrespective of their coloration when given the opportunity to observe their behavior toward a potential fish predator. By preferentially mating with colorful males, female guppies are thus choosing on average, relatively bold, and perhaps more viable, individuals. In doing so, and to the extent that viability is heritable, they potentially gain indirect fitness benefits by producing more viable offspring than otherwise.

Preference polymorphism for coloration but no speciation in a population of Lake Victoria cichlids

Female mating preference based on male nuptial coloration has been suggested to be an important source of diversifying selection in the radiation of Lake Victoria cichlid fish. Initial variation in female preference is a prerequisite for diversifying selection; however, it is rarely studied in natural populations. In clear water areas of Lake Victoria, the sibling species Pundamilia pundamilia with blue males and Pundamilia nyererei with red males coexist, intermediate phenotypes are rare, and most females have species-assortative mating preferences. Here, we study a population of Pundamilia that inhabits turbid water where male coloration is variable from reddish to blue with most males intermediate. We investigated male phenotype distribution and female mating preferences. Male phenotype was unimodally distributed with a mode on intermediate color in 1 year and more blue-shifted in 2 other years. In mate choice experiments with females of the turbid water population and males from a clearer water population, we found females with a significant and consistent preference for P. pundamilia (blue) males, females with such preferences for P. nyererei (red) males, and many females without a preference. Hence, female mating preferences in this population could cause disruptive selection on male coloration that is probably constrained by the low signal transduction of the turbid water environment. We suggest that if environmental signal transduction was improved and the preference/color polymorphism was stabilized by negative frequency-dependent selection, divergent sexual selection might separate the 2 morphs into reproductively isolated species resembling the clear water species P. pundamilia and P. nyererei.

The shape of female mating preferences

The “shape” of a female mating preference is the relationship between a male trait and the probability of acceptance as a mating partner. The shape of preferences is important in many models of sexual selection, mate recognition, communication, and speciation, yet it has rarely been measured precisely. Here I examine preference shape for male calling song in a bushcricket (katydid). Preferences change dramatically between races of a species, from strongly directional to broadly stabilizing (but with a net directional effect). Preference shape generally matches the distribution of the male trait. This is compatible with a coevolutionary model of signal-preference evolution, although it does not rule out an alternative model, sensory exploitation. Preference shapes are shown to be genetic in origin.

Male mutation bias and possible long-term effects of human activities.

The ability of a population to adapt to changing environments depends critically on the amount and kind of genetic variability it possesses. Mutations are an important source of new genetic variability and may lead to new adaptations, especially if the population size is large. Mutation rates are extremely variable between and within species, and males usually have higher mutation rates as a result of elevated rates of male germ cell division. This male bias affects the overall mutation rate. We examined the factors that influence male mutation bias, and focused on the effects of classical life-history parameters, such as the average age at reproduction and elevated rates of sperm production in response to sexual selection and sperm competition. We argue that human-induced changes in age at reproduction or in sexual selection will affect male mutation biases and hence overall mutation rates. Depending on the effective population size, these changes are likely to influence the long-term persistence of a population.

Feasibility of Using a Caribbean Screwworm for SIT Campaigns in Brazil

The screwworm, Cochliomyia hominivorax (Coquerel), remains one of the most damaging parasites of livestock in South America, causing millions of dollars in annual losses to producers. Recently, South American countries demonstrated interest in controlling this pest using the Sterile Insect Technique, and a pilot-project was conducted near the Brazil-Uruguay border in 2009. Since molecular studies have suggested the existence of C. hominivorax regional groups, crossing tests were conducted to evaluate mating competitiveness, mating preference and reproductive compatibility between a C. hominivorax strain from the Caribbean (Jamaica-06) and one from Brazil. Mating rates between Jamaican males and Brazilian females ranged between 82 and 100%, and each male inseminated from 3.3 to 3.95 females. Sterile males, regardless of the strain, competed equally against the fertile males for Brazilian females. Jamaican sterile males and Brazilian fertile males mated randomly with fertile or sterile females. No evidence of genetic incompatibility or hybrid dysgenesis was found in the hybridization crosses. Mating barriers should not compromise the use of Jamaican sterile males for Sterile Insect Technique campaigns in Brazil.

Incipient speciation by sexual isolation in Drosophila: Concurrent evolution at multiple loci

Drosophila melanogaster from Zimbabwe and nearby regions shows strong but asymmetric sexual isolation from its cosmopolitan counterparts. By creating stable chromosome-substitution lines, earlier studies were able to show that the two major autosomes have very large effects on both male mating success and female mating preference. In this study, we genetically dissect this sexual isolation by recombination analysis between a whole-chromosome substitution line (which carries a Zimbabwe-derived third chromosome) and a strain with seven visible markers on that chromosome. Four loci are responsible for male mating success and three others are found to control female mating preference. Because male and female traits are not closely linked, their strong association among isofemale lines is most likely a reflection of sexual selection in nature. The results suggest that a large number of behavioral loci may evolve concurrently in the incipient stage of speciation before other aspects of reproductive isolation (such as hybrid sterility) have become evident. The results shed light on the population genetic processes underlying the formation of nascent species, as well as modes of speciation.

Measuring natural and sexual selection on breeding values of male display traits in Drosophila serrata

Fundamental to many theories of sexual selection is the expectation that sexual traits, which males use in an attempt to increase mating success, confer costs as well as benefits to individual males. Although evolution of exaggerated male traits is predicted to be halted, by costs applied by natural selection, there is a lack of empirical work devoted to quantitatively establishing whether natural selection opposes sexual selection generated by the preferences of females. In this study, we quantified natural and sexual selection gradients on breeding values for cuticular hydrocarbon (CHC) components of male contact pheromones in Drosophila serrata. As male sexual traits may often be environmentally condition dependent, breeding values were used in the selection analysis to remove the possibility of environmental correlations between the measured trait and fitness biasing estimates of selection. The direction of natural selection was found to oppose sexual selection on a subset of CHCs examined. Opposing natural and sexual selection suggests that further evolution of the male pheromone may in part be limited by costs associated with attractive male CHC blends.

L’influence des capacités cognitives mâles et femelles sur le choix de partenaire chez le diamant mandarin (Taeniopygia guttata)

S’approvisionner en nourriture est essentiel à la survie et au succès reproducteur. Lorsque les animaux font face à des changements environnementaux brutaux, ils doivent s’ajuster rapidement à leur nouvel environnement et parfois même innover dans leur façon de s’approvisionner. Des processus comportementaux et cognitifs, tels que l’innovation et l’apprentissage, permettent aux animaux d'intégrer de nouveaux comportements à leur répertoire comportemental afin de s'adapter de façon optimale. Les performances cognitives varient entre les individus d’une même population et bien que des études récentes se soient intéressées aux causes de ce phénomène, de convaincantes évidences sont manquantes afin d’expliquer pourquoi ces variations sont maintenues. Au cours de ce mémoire, les questions des pressions de sélection s'exerçant sur les performances d’alimentation par une tâche motrice nouvelle sont abordées afin de mieux comprendre l'évolution des capacités cognitives au sein d'une population captive de diamants mandarins (Taeniopygia guttata). Nous avons tout d'abord testé si les femelles diamants mandarins modifient leurs préférences d'accouplement après avoir observé la performance d'alimentation par une tâche motrice nouvelle des mâles. Afin de déterminer si les femelles sont capables de discriminer entre les mâles sur la base de leur capacité cognitive, nous avons également évalué les performances d’apprentissage de chacune d’elles. En effet, des études ont suggéré qu’il peut être coûteux, spécialement en terme de temps, de discriminer entre des partenaires potentiels sur cette base. La généralisation d’une préférence pour un mâle performant à d’autres mâles possédant le même phénotype permettrait la réduction de ces coûts. Nous avons donc finalement testé si les femelles diamants mandarins peuvent généraliser leur préférence après avoir observé les performances d’alimentation pour une tâche motrice nouvelle d’un mâle. Nos résultats suggèrent que les femelles diamants mandarins ne peuvent évaluer les capacités cognitives d’un mâle par l’intermédiaire de traits indicateurs. Toutefois, nous avons démontré qu’une observation directe des performances d’alimentation d’un mâle guide le choix d’appariement des femelles. Également, nous avons montré que les femelles peuvent généraliser l’apparence du mâle le plus performant et utiliser cette information lors de l’évaluation de nouveaux mâles. La relation entre les performances cognitives et le choix de partenaire pourraient s’expliquer par exemple par une meilleure exploitation de l’habitat, mais nécessite des études plus approfondies.

L’influence des capacités cognitives mâles et femelles sur le choix de partenaire chez le diamant mandarin (Taeniopygia guttata)

S’approvisionner en nourriture est essentiel à la survie et au succès reproducteur. Lorsque les animaux font face à des changements environnementaux brutaux, ils doivent s’ajuster rapidement à leur nouvel environnement et parfois même innover dans leur façon de s’approvisionner. Des processus comportementaux et cognitifs, tels que l’innovation et l’apprentissage, permettent aux animaux d'intégrer de nouveaux comportements à leur répertoire comportemental afin de s'adapter de façon optimale. Les performances cognitives varient entre les individus d’une même population et bien que des études récentes se soient intéressées aux causes de ce phénomène, de convaincantes évidences sont manquantes afin d’expliquer pourquoi ces variations sont maintenues. Au cours de ce mémoire, les questions des pressions de sélection s'exerçant sur les performances d’alimentation par une tâche motrice nouvelle sont abordées afin de mieux comprendre l'évolution des capacités cognitives au sein d'une population captive de diamants mandarins (Taeniopygia guttata). Nous avons tout d'abord testé si les femelles diamants mandarins modifient leurs préférences d'accouplement après avoir observé la performance d'alimentation par une tâche motrice nouvelle des mâles. Afin de déterminer si les femelles sont capables de discriminer entre les mâles sur la base de leur capacité cognitive, nous avons également évalué les performances d’apprentissage de chacune d’elles. En effet, des études ont suggéré qu’il peut être coûteux, spécialement en terme de temps, de discriminer entre des partenaires potentiels sur cette base. La généralisation d’une préférence pour un mâle performant à d’autres mâles possédant le même phénotype permettrait la réduction de ces coûts. Nous avons donc finalement testé si les femelles diamants mandarins peuvent généraliser leur préférence après avoir observé les performances d’alimentation pour une tâche motrice nouvelle d’un mâle. Nos résultats suggèrent que les femelles diamants mandarins ne peuvent évaluer les capacités cognitives d’un mâle par l’intermédiaire de traits indicateurs. Toutefois, nous avons démontré qu’une observation directe des performances d’alimentation d’un mâle guide le choix d’appariement des femelles. Également, nous avons montré que les femelles peuvent généraliser l’apparence du mâle le plus performant et utiliser cette information lors de l’évaluation de nouveaux mâles. La relation entre les performances cognitives et le choix de partenaire pourraient s’expliquer par exemple par une meilleure exploitation de l’habitat, mais nécessite des études plus approfondies.