Variable assortative mating in replicate mating trials using Drosophila melanogaster populations derived from contrasting opposing slopes of 'Evolution Canyon', Israel

Significant assortative mating in laboratory studies has been previously shown between two populations of Drosophila melanogaster collected from micro-climactically contrasting and opposing slopes of 'Evolution Canyon' (Lower Nahal Oren, Israel; Korol et al., 2000). Coupled with evidence that the two populations are adapted to their respective environments, this has been suggested as a rare example of ongoing behaviourally mediated speciation occurring in the face of gene flow. Reproductive isolation between these populations, however, has never been confirmed by replicate experiments in an independent laboratory. For this reason, we tested recent collections of these populations for premating isolation in both the original (Haifa) and a new (Burnaby) laboratory under a variety of experimental protocols. Although non-random mating was found in the majority of trials conducted in Haifa, we were unable to replicate these strong results in Burnaby. Most notably, we failed to detect assortative mating in four separate double choice experiments. Significant non-random mating was detected, however, in three of six single choice experiments in Burnaby, suggesting that the populations are behaviourally differentiated in some manner. Why nonrandom mating was weaker in Burnaby than Haifa is not understood, but suggests that assortative mating may be sensitive to unknown environmental factors.     

Selection and gene flow between microenvironments: the case of Drosophila at Lower Nahal Oren,Mount Carmel,Israel

Drosophila melanogaster and D. simulans from contrasted microenvironments (south- and north-facing slopes of Lower Nahal Oren Canyon in Israel) were tested for genetic differentiation at microsatellite loci, which might be linked to differential adaptation to local ecological factors. No overall genetic differentiation was observed in either species. This indicates that the contrasted selective pressures on the two sides of the valley are not strong enough to cause population subdivision in highly mobile species such as Drosophila. Significant differences in allele frequencies were observed at two microsatellite loci, but on the whole the level of divergence we observed is far lower than has been reported previously.     

Assortative mating between two distinct micro-allopatric populations of Littorina saxatilis (Olivi) on the northeast coast of England

Size assortative mating is a common invertebrate mating pattern and is usually accompanied by male and female sexual selection, and these three behaviours can contribute to reproductive isolation. Two distinct populations of the marine prosobranch Littorina saxatilis, H and M, occur within 15 m of each other on the same shore. Previous studies have demonstrated that these two forms have different reproductive strategies and that the rare hybrids between the two forms show evidence of reproductive dysfunction and hence are less fit than the assumed parental forms. In both populations, female shell height was shown to be a predictor of the number of embryos contained within the brood pouch. The mean shell height of the M population was significantly larger than that of the H population, and the M population matures at a larger shell size than the H population. The two populations show complete assortative mating to type in the field, and occupy different microhabitats on the same shore. Therefore, laboratory-based experiments were performed to determine if assortative mating was maintained in sympatry and also to determine the effect of population density on mate choice. The males of both populations showed sexual selection for female size, choosing to mate with females approximately 10% larger than themselves from an assortment of female sizes. The M population showed complete assortative mating to type, irrespective of the density of H and M females, whereas at low densities the H males did occasionally mate with M females. The role of assortative mating and reinforcement (due to natural selection acting against the less fit hybrids), in maintaining the partial reproductive barrier between the two populations is discussed.     

Reproductive isolating barriers between colour‐differentiated populations of an African annual killifish,Nothobranchius korthausae (Cyprinodontiformes)

Allopatric populations separated by vicariance events are expected to evolve reproductive isolating mechanisms as a result of disparate selection pressures and genetic drift. The appearance of reproductive isolating mechanisms may vary across taxa with differences in the opportunity for mate choice, and may be asymmetrical. In addition, premating barriers may be affected by individual mating experience. We used choice and no‐choice experiments to investigate reproductive isolation between two allopatric (island and mainland) and colour‐differentiated populations of an African annual fish, Nothobranchius korthausae. Assortative mating under experimental conditions was limited and asymmetrical. Preference for sympatric males was only expressed in nonvirgin females from one population. Virgin fish from both populations mated indiscriminately. No difference in the number of eggs laid, fertilization rate and hatching success was detected in no‐choice experiments. All mating combinations produced viable offspring and no postmating barriers were detected in terms of the performance and fertility of F1 hybrids. Overall, we found little evidence for significant reproductive isolation, which is in contrast with the related killifish taxa in which assortative mating can be strong, even among allopatric populations with no colour differentiation. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 62–72.     

Male pheromone polymorphism and reproductive isolation in populations of Drosophila simulans

The dominant cuticular hydrocarbons (HC) in Drosophila simulans are 7‐tricosene (7‐T) and 7‐pentacosene (7‐P). The 7‐T is the major HC in East Africa and in other continents. In West Africa, D. simulans is very rare and displays 7‐P as the major compound. We studied three D. simulans strains from Egypt (Eg), Sao‐Tome (ST), and Cameroon (Cam), with 7‐T, intermediary or 7‐P phenotypes. HC profiles of ST and Cam female differed slightly from corresponding male profiles; females had more 7‐T and less 7‐P. Varying temperature affected all HCs (even those with 27 and 29 carbons)‐not just 7‐T and 7‐P; there was no clear relationship between HC phenotype and resistance to desiccation. We report reproductive isolation between Eg and ST and Eg and Cam (but not between ST and Cam), which is due to Eg and Cam female preferences for their own males. In conclusion, our findings do support divergence of D. simulans populations from West Africa for both pheromonal profile and mating preference.     

Reproductive isolation in natural populations of Drosophila melanogaster from Brazzaville (Congo)

The aim of this work is to analyze the homogamy previously detected between two natural populations of Drosophila melanogaster from Brazzaville. It is shown that mating isolation was still maintained under laboratory conditions 10 years after the populations samples were trapped. Isolation seemed to be due mainly to pre-mating isolation and we checked for any suggestion of post-mating mortality of hybrids. Pre-mating isolation was not symmetrical, and significant ² values were found in 3/4 possible 3-way mating choice experiments. The only exception involved a male from the countryside and two females (one from each population) for which no significant mating preference was detected. Mortality of hybrids was intermediate between those of the parental strains showing a clear maternal effect and the existence of partial dominance. Major differences in the cuticular hydrocarbons were also found and they could account for the isolation. These findings in populations from African breweries indicate that they are closely related to European ones, suggesting that this phenomenon is not a case of sympatric speciation, but probably attributable to the reintroduction of an allopatric population.     

Size Assortative Mating and Reproductive Success of the Funnel-Web Spider, Agelena limbata (Araneae; Agelenidae)

Field observations and laboratory experiments were conducted to assess the relation between male size and reproductive success in the funnel-web spider, Agelena limbata Thorell (Agelenidae), in 2 years. In this species, the body size of males is similar to that of females. In the field, size assortative mating occurred in both years. In 1 year, partial correlation coefficient analysis indicates that male cephalothorax width is a beter predictor of the copulated female cephalothorax width than of the date of pairing. In laboratory experiments, females tended to reject courting males that were smaller in relative body size, and males that were larger in relative body size had greater copulation success. Consequently female rejection of smaller courting males has some contribution to size assortative mating. Since larger females deposited more numerous eggs in the field, larger males are expected to have a higher reproductive success.     

Male emergence timing and mating success in the funnel-web spider,Agelena limbata (Araneae: Agelenidae)

Field observations on the relationship between male mating success and emergence timing in the funnel-web spider,Agelena limbata, were conducted.Agelena limbata is an annual species and adult males appear slightly earlier than adult females in July. As males deposit a copulatory plug at the female epigynum after copulation, copulation with virgin females is important to males. The number of copulations in males with virgin females, which strongly correlates with the longevity of males and the number of females that males courted, did not correlate with the emergence timing of males. Early emerged males and females were significantly larger in size than later ones, but the correlation coefficient between the emerged date and the cephalothorax width was not strong. Males that emerged earlier did not have any advantage in copulating with larger and more fecund females. Furthermore, virgin females first copulated on average 7.9 days after their final molt and the mortality rate of adult males increased after the final molt. These factors may favor the smaller degree of protandry in male emergence timing inA. limbata.     

Wax,sex and the origin of species: Dual roles of insect cuticular hydrocarbons in adaptation and mating

Evolutionary changes in traits that affect both ecological divergence and mating signals could lead to reproductive isolation and the formation of new species. Insect cuticular hydrocarbons (CHCs) are potential examples of such dual traits. They form a waxy layer on the cuticle of the insect to maintain water balance and prevent desiccation, while also acting as signaling molecules in mate recognition and chemical communication. Because the synthesis of these hydrocarbons in insect oenocytes occurs through a common biochemical pathway, natural or sexual selection on one role may affect the other. In this review, we explore how ecological divergence in insect CHCs can lead to divergence in mating signals and reproductive isolation. We suggest that the evolution of insect CHCs may be ripe models for understanding ecological speciation.     

Sexual selection and assortative mating by size and their roles in the maintenance of a polymorphism in Swedish Littorina saxatilis populations

Two independent components of mating behaviour, sexual selection and assortative mating, were studied in two allopatric morphs, one sheltered boulder shore form (S-morph) and one exposed cliff shore form (E-morph), of Littorina saxatilis from the west coast of Sweden. Sexual selection was studied by comparing the sizes of copulating and non-copulating snails in the field. Size assortative mating was studied by collecting copulating pairs in the field, while assortative mating between morphs was investigated by bringing the pure morphs together in intermediary habitats and then noting the matings. The S-morph mated randomly in relation to size in two of the studied populations and exhibited a trend towards size assortative mating in a third, while the E-morph showed size assortative mating in both studied populations. The microdistribution of sizes of snails on the shores could not explain all the size assortative mating found, and instead it is argued that a size-based mate rejection behaviour also contributes to the assortative mating in at least some of these populations. There was sexual selection on size in both males and females in the S-morph, with large individuals being favoured as mates. In contrast, copulating snails of the E-morph were smaller than non-copulating ones. The significantly different sexual selection intensities between the two morphs may help to explain the size differences between them. There was random mating between the E- and the S-morphs of L. saxatilis, which suggests no incipient reproductive isolation between morphs on Swedish rocky shores. This is in agreement with earlier studies of Swedish populations, but is in contrast to the situation found in other geographical areas.     

Sex in a cyclical parthenogen: mating behaviour of Chydorus sphaericus (Crustacea; Branchiopoda; Anomopoda)

1. We describe the interactions during mating in Chydorus sphaericus, a cyclical parthenogenetic anomopod. Mating behaviour is more complex than previously assumed, with evidence for a diffusible chemical to which males react at the onset of mating, for reproductive isolation, and for postcopulatory mate guarding. 2. During mating, the male and female form a ‘mating cross’ that may be maintained for several hours, while copulation itself typically lasts less than a minute. Furthermore, males invariably attach to the right valve of females. Copulation involves intromission of the postabdomen between the valves, so that the gonopores approach the left ovarium. 3. This behaviour is reflected in the morphology of both sexes: males have a specialised anterior valve margin, postabdomen, first limb and rostrum, under selective pressure for successful mate guarding and copulation, while gamogenetic females have asymmetric ovaries, and a species‐specific setulation of the valves. Males of the structurally related Chydorus ovalis react to the presence of C. sphaericus, but fail to dock to females, suggesting a lock‐antilock element in the reproductive isolation of both species. 4. The morphological and ethological adaptations in C. sphaericus suggest that there is a strong selective pressure on mating behaviour in this cyclical parthenogen and specifically towards the formation of the ‘mating cross’.     

Emergence and mating behavior of the oriental fruit moth Cydia molesta (Lepidoptera: Tortricidae) and its potential for reproduction

The Oriental fruit moth, Cydia molesta (Busck, 1916) (Lepidoptera: Tortricidae), is a key pest of fruit and is widely distributed around the world. There are important connections between its behavior and biology and its management in agriculture, but few studies have investigated the associations between adult behaviors and oviposition. In this study, adult emergence, mating, and reproduction were investigated under laboratory and field conditions. The ratio of females to males at eclosion was approximately 1:1. When one virgin female had access to one virgin male, 66% and 34% of the couples copulated just once and twice, respectively; and the infertility rate of eggs (21.39 ± 1.25%) did not vary daily. Males, given access to one new female daily, could copulate multiple times, whereas females seldom mated more than once, indicating a male-biased operational sex ratio, but mating status of the male parent had no effect on progeny egg reproduction. Also, the number of eggs that hatched by all female partners of a male was inversely proportional to copulation duration for the female laying the eggs for total female reproductive success; and the number of eggs laid by all female partners of a male was proportional to their number of matings for total male reproductive success. However, the total number of eggs that hatched did not significantly differ for eggs laid by a female given new virgin males daily for mating (17.75 ± 4.28) versus eggs laid by virgin females (19.17 ± 7.51) presented daily with a male that re-mated daily with the series of females. Therefore, our results showed that females engaged in mate choice and males engaged in mate competition, affecting egg production, a factor that may be used to enhance mating disruption technology against Cydia molesta.     

The impact of Wolbachia,male age and mating history on cytoplasmic incompatibility and sperm transfer in Drosophila simulans

Most insects harbour a variety of maternally inherited endosymbionts, the most widespread being Wolbachia pipientis that commonly induce cytoplasmic incompatibility (CI) and reduced hatching success in crosses between infected males and uninfected females. High temperature and increasing male age are known to reduce the level of CI in a variety of insects. In Drosophila simulans, infected males have been shown to mate at a higher rate than uninfected males. By examining the impact of mating rate independent of age, this study investigates whether a high mating rate confers an advantage to infected males through restoring their compatibility with uninfected females over and above the effect of age. The impact of Wolbachia infection, male mating rate and age on the number of sperm transferred to females during copulation and how it relates to CI expression was also assessed. As predicted, we found that reproductive compatibility was restored faster in males that mate at higher rate than that of low mating and virgin males, and that the effect of mating history was over and above the effect of male age. Nonvirgin infected males transferred fewer sperm than uninfected males during copulation, and mating at a high rate resulted in the transfer of fewer sperm per mating irrespective of infection status. These results indicate that the advantage to infected males of mating at a high rate is through restoration of reproductive compatibility with uninfected females, whereas uninfected males appear to trade off the number of sperm transferred per mating with female encounter rate and success in sperm competition. This study highlights the importance Wolbachia may play in sexual selection by affecting male reproductive strategies.     

The effects of body size on mating and reproduction in Brachinus lateralis (Coleoptera: Carabidae)

Abstract. 1. The causes and reproductive consequences of body size variation of Brachinus lateralis Dejean, a parasitic carabid beetle, were investigated.
2. Body size variation occurs within and between sites. Host size has a major influence on body size of the adult.
3. Fecundity is positively correlated with body size. Egg size is not correlated with body size.
4. Mating males tend to be larger than non-mating males. There is a positive correlation of body sizes in mating pairs.
5. Limited opportunity for host choice may maintain size variation despite the advantages of large size.
6. The non-random patterns of mating for a species without obvious intrasexual aggression suggest that subtle means of male-male competition or female choice may be important.     

Clinal variation and laboratory adaptation in the rainforest species Drosophila birchii for stress resistance, wing size, wing shape and development time

Clinal variation has been described in many invertebrates including drosophilids but usually over broad geographical gradients. Here we describe clinal variation in the rainforest species Drosophila birchii from Queensland, Australia, and potential confounding effects of laboratory adaptation. Clinal variation was detected for starvation and development time, but not for size or resistance to temperature extremes. Starvation resistance was higher at southern locations. Wing shape components were not associated with latitude although they did differ among populations. Time in laboratory culture did not influence wing size or heat knockdown resistance, but increased starvation resistance and decreased recovery time following a cold shock. Laboratory culture also increased development time and altered wing shape. The results indicate that clinal patterns can be detected in Drosophila over a relatively narrow geographical area. Laboratory adaptation is unlikely to have confounded the detection of geographical patterns.     

Multiple mating and its effect on the reproductive success of female Epiphyas postvittana (Lepidoptera: Tortricidae)

Abstract.

• 1 Multiple mating and its effect on reproductive performance of female Epiphyas postvittana (Walker) moths were studied under controlled conditions.
• 2 The age at which the moths mated for the first time ranged from the first to the tenth day after emergence, but 71% of first matings were during the first 3 days.
• 3 The majority (63%) of females had one or two spermatophores in the bursa copulatrix. Some (24%) were found with three to five spermatophores, whereas no successful mating occurred among 13% of individuals. The number of matings was partly dependent on the number of mates available to the female. Between the range of sex ratios of one male to one female and four males to one female maximal mating success occurred at the ratio of three males to one female.
• 4 Virgin females were capable of egg-laying, but mating stimulated and accelerated oviposition. Mated individuals laid twice as many eggs as unmated ones.
• 5 The level of copulatory activity did not influence the longevity of females irrespective of the number of males available to them.
• 6 Sex ratios with greater than one male to a female improved the reproductive success by marginally increasing fecundity and fertility.
• 7 It is concluded that multiple mating would enhance population growth, and is of particular benefit to populations with a preponderance of females, as is known to occur naturally in this species.

     

The effects of temperature and body size on the mating pattern of a gregariously nesting bee, Colletes cunicularius (Hymenoptera: Colletidae)

Abstract. 1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.     

The evolution of intrinsic reproductive isolation in the genus Cakile (Brassicaceae)

In theory, adaptive divergence can increase intrinsic post‐zygotic reproductive isolation (RI), either directly via selection on loci associated with RI, or indirectly via linkage of incompatibility loci with loci under selection. To test this hypothesis, we measured RI at five intrinsic post‐zygotic reproductive barriers between 18 taxa from the genera Cakile and Erucaria (Brassicaceae). Using a comparative framework, we tested whether the magnitude of RI was associated with genetic distance, geographic distance, ecological divergence and parental mating system. Early stages of post‐zygotic RI related to F₁ viability (i.e. initial seed set) tended to be stronger than later stages related to F₁ fecundity (i.e. flower number, fruit number). Mating system significantly influenced early stages of RI, such that RI was lowest when the mother was selfing and father was outcrossing, consistent with an imbalance between sink strength and resistance to provisioning. We found little evidence that adaptive divergence accelerates the evolution of intrinsic post‐zygotic RI, consistent with a nonecological model of evolution that predicts the nonlinear accumulation of RI and RI asymmetry with time (i.e. genetic distance), irrespective of adaptive divergence. Thus, although certain aspects of ecological divergence do not appear to have contributed strongly to the evolution of RI in this system, divergence in mating system actually reduced RI, suggesting that mating system evolution may play a significant role in speciation dynamics.