Parasite infection and the movement of the aquatic snail Potamopyrgus antipodarum along a depth cline.

Parasite species that use two or more host species during their life cycle depend on successful transmission between these species. These successive host species may have different habitat requirements. For example, one host species may be aquatic while the other is terrestrial. To overcome this complicating factor in transmission, a wide diversity of parasite species have adaptations that alter the habitat preference in one host species to facilitate transmission to the next host species.Two common trematode parasites in New Zealand, Atriophallophorus winterbourni and Notocotylus spp., both have a life cycle with two host species. The aquatic snail Potamopyrgus antipodarum is the intermediate host, from which the parasites require transmission to dabbling ducks or other waterfowl. Of these parasites, A. winterbourni is most frequently found in snails from the shallow-water margin. This may indicate parasite-induced movement of infected snails into the foraging habitat of dabbling ducks.To test whether the parasites manipulate the snails to move into shallow water, we stretched tubular mesh cages across depth-specific ecological habitat zones in a lake. Both infected and healthy snails were released into the cages. After 11 days, significantly higher infection frequencies of A. winterbourni were retrieved from the shallowest end of the cages, while Notocotylus spp. frequencies did not vary with depth.The hypothesis that A. winterbourni induces its snail host to move into the shallow-water habitat cannot be rejected based on the experimental results. Although further research is needed to address alternative explanations, the depth preference of infected snails may be due to a parasite adaptation that facilitates trophic transmission of parasites to dabbling ducks.

Sexual conflict over the duration of copulation in Drosophila montana: why is longer better?

BACKGROUND: Conflicts of interest between the sexes are increasingly recognized as an engine driving the (co-)evolution of reproductive traits. The reproductive behaviour of Drosophila montana suggests the occurrence of sexual conflict over the duration of copulation. During the last stages of copulation, females vigorously attempt to dislodge the mounting male, while males struggle to maintain genital contact and often successfully extend copulations far beyond the females' preferred duration. RESULTS: By preventing female resistance, we show that females make a substantial contribution towards shortening copulations. We staged matings under different sex ratio conditions, and provide evidence that copulation duration is a form of male reproductive investment that responds to the perceived intensity of sperm competition as predicted by game theoretical models. Further, we investigated potential benefits to persistent males, and costs to females coerced into longer matings. While males did not benefit in terms of increased progeny production by protracting copulation, female remating was delayed after long first copulations. CONCLUSION: Copulation time is a trait subject to sexual conflict. Mating durations exceeding female optima serve males as a form of 'extended mate guarding': by inducing mating refractoriness in the female, a male extends the time over which its sperm is exclusively used to sire progeny and reduces the likelihood of the female being reinseminated by a competitor.

Parasite resistance predicts fitness better than fecundity in a natural population of the freshwater snail Potamopyrgus antipodarum.

The cost of males should give asexual females an advantage when in competition with sexual females. In addition, high-fecundity asexual genotypes should have an advantage over low-fecundity clones, leading to reduction in clonal diversity over time. To evaluate fitness components in a natural population, we measured the annual reproductive rate of individual sexual and asexual female Potamopyrgus antipodarum, a New Zealand freshwater snail, in field enclosures that excluded competitors and predators. We used allozyme genotyping to assign the asexual females to particular clonal genotypes. We found that the most fecund asexual clones had similar or higher fecundity as the top 10% of sexual families, suggesting that fecundity selection, even without the cost of males, would lead to replacement of the sexual population by clones. Consequently, we expected that the clones with the highest fecundity would dominate the natural population. Counter to this prediction, we found that high annual reproductive rates did not correlate with the frequency of clones in the natural population. When we exposed the same clones to parasites in the laboratory, we found that resistance to infection was positively correlated with the frequency of clones in the population. The correlation between fecundity and parasite resistance was negative, suggesting a trade-off between these two traits. Our results thus suggest that parasite resistance is an important short-term predictor of the success of asexual P. antipodarum in this population.

Male courtship song and female preference variation between phylogeographically distinct populations of Drosophila montana.

Understanding the variation within and between populations in important male mating traits and female preferences is crucial to theories concerning the origin of sexual isolation by coevolution or other processes. There have been surprisingly few studies on the extent of variation and covariation within and between populations, especially where the evolutionary relationships between populations are understood. Here we examine variation in female preferences and a sexually selected male song trait, the carrier frequency of the song, within and between populations from different phylogeographic clusters of Drosophila montana. Song is obligatory for successful mating in this species, and both playback and field studies implicate song carrier frequency as the most important parameter in male song. Carrier frequency varied among three recently collected populations from Oulanka (Finland), Vancouver (Canada), and Colorado (central United States), which represent the main phylogeographic groups in D. montana. Males from Colorado had the most distinct song frequency, which did not follow patterns of genetic differentiation. There was considerable variation in preference functions within, and some variation between, populations. Surprisingly, females from three lines from Colorado seem to have preferences disfavoring the extreme male trait found in this population. We discuss sources of selection on male song and female preference.

Indirect benefits for choosy female grasshoppers (Chorthippus biguttulus)?

Since direct benefits are likely to be absent in the grasshopper Chorthippus biguttulus, indirect genetic benefits are a potential explanation for costly female preference. Choosy females may improve their fitness in terms of enhanced attractiveness of sons alone or additionally by improved viability of offspring. We tested the predictions of these two hypotheses by comparing attractiveness-related song traits and viability in offspring of attractive and unattractive grasshopper males. The experiment was conducted with larvae reared under semi natural lab conditions in one year and under natural conditions in the field in the following year. If reared under natural conditions no significant differences in viability and song traits between offspring of attractive and unattractive males could be found. Offspring reared in the lab produced calling songs with a significantly more exact song rhythm when sired by attractive males than offspring of unattractive males. Offspring of attractive males should thus have a theoretical advantage in mate choice, which, however, did not translate into higher attractiveness values in acoustic female choice experiments. Therefore our experiments could not resolve whether female choice in C. biguttulus evolved according to the sexy son hypothesis. Since viability in offspring of attractive males did not differ from offspring of unattractive males, "good genes" seems unlikely to be the underlying mechanism of female choice.

The attractiveness fragment--AFLP analysis of local adaptation and sexual selection in a caeliferan grasshopper, Chorthippus biguttulus.

Genetic variability among males is a necessary precondition for the evolution of female choice based on indirect genetic benefits. In addition to mutations and host-parasite cycles, migration of locally adapted individuals offers an explanation for the maintenance of genetic variability. In a previous study, conducting a reciprocal transplant experiment on a grasshopper, Chorthippus biguttulus, we found that environmental conditions significantly influenced not only body condition but also an important trait of male calling song, the amplitude of song. Although not significant, all other analysed physical and courtship song traits and attractiveness were superior in native than in transferred males. Thus, we concluded that local adaptation has a slight but consistent influence on a range of traits in our study populations, including male acoustic attractiveness. In our present study, we scanned male grasshoppers from the same two populations for amplification fragment length polymorphism (AFLP) loci connected with acoustic attractiveness to conspecific females. We found greater differences in allele frequencies between the two populations, for some loci, than are expected from a balance between drift and gene flow. These loci are potentially connected with locally adapted traits. We examined whether these alleles show the proposed genotype environment interaction by having different associations with attractiveness in the two populations. One locus was significantly related to sexual attractiveness; however, this was independent of the males' population affiliation. Future research on the evolution of female choice will benefit from knowledge of the underlying genetic architecture of male traits under intraspecific sexual selection, and the 'population genomics' approach can be a powerful tool for revealing this structure.

Factors affecting male song evolution in Drosophila montana.

D. montana (a species of the D. virilis group) has spread over the northern hemisphere, populations from different areas showing both genetic and phenotypic divergence. The males of this species produce an elaborate courtship song, which plays a major role both in species recognition and in intraspecific mate choice. The genetic architecture and physical constraints, as well as the importance of the signal for species recognition, set boundaries within which this signal can vary. Within these limits, courtship song parameters may change, depending on the males' physical condition and on the environment they inhabit. Females are likely to affect song evolution by exerting directional selection toward higher carrier frequencies. Given this complexity, only a comprehensive, multidisciplinary approach, starting with traditional field observation and combining controlled behavioral experiments, biometric measurements, and sophisticated molecular techniques, has the potential of shedding light on the past history and the evolution of this signal, and, eventually, adding to our understanding of the mechanisms, functions, and outcomes of sexual selection in acoustic communication systems.