Population structure in relation to host-plant ecology and Wolbachia infestation in the comma butterfly.

Experimental work on Polygonia c-album, a temperate polyphagous butterfly species, has shown that Swedish, Belgian, Norwegian and Estonian females are generalists with respect to host-plant preference, whereas females from UK and Spain are specialized on Urticaceae. Female preference is known to have a strong genetic component. We test whether the specialist and generalist populations form respective genetic clusters using data from mitochondrial sequences and 10 microsatellite loci. Results do not support this hypothesis, suggesting that the specialist and generalist traits have evolved more than once independently. Mitochondrial DNA variation suggests a rapid expansion scenario, with a single widespread haplotype occurring in high frequency, whereas microsatellite data indicate strong differentiation of the Moroccan population. Based on a comparison of polymorphism in the mitochondrial data and sequences from a nuclear gene, we show that the diversity in the former is significantly less than that expected under neutral evolution. Furthermore, we found that almost all butterfly samples were infected with a single strain of Wolbachia, a maternally inherited bacterium. We reason that indirect selection on the mitochondrial genome mediated by a recent sweep of Wolbachia infection has depleted variability in the mitochondrial sequences. We also surmise that P. c-album could have expanded out of a single glacial refugium and colonized Morocco recently.

Genetics of host plant use and life history in the comma butterfly across Europe: varying modes of inheritance as a potential reproductive barrier.

Comma butterflies (Nymphalidae: Polygonia c-album L.) from one Belgian site and three Spanish sites were crossed with butterflies from a Swedish population in order to investigate inheritance of female host plant choice, egg mass and larval growth rate. We found three different modes of inheritance for the three investigated traits. In line with earlier results from crosses between Swedish and English populations, the results regarding female oviposition preference (choice between Urtica dioica and Salix caprea) showed X-linked inheritance to be of importance for the variation between Sweden and the other sites. Egg mass and growth rate did not show any sex-linked inheritance. Egg mass differences between populations seem to be controlled mainly by additive autosomal genes, as hybrids showed intermediate values. The growth rates of both hybrid types following reciprocal crossings were similar to each other but consistently higher than for the two source populations, suggesting a nonadditive mode of inheritance which is not sex-linked. The different modes of inheritance for host plant preference vs. important life history traits are likely to result in hybrids with unfit combinations of traits. This type of potential reproductive barrier based on multiple ecologically important traits deserves more attention, as it should be a common situation for instance in the early stages of population divergence in host plant usage, facilitating ecological speciation.

Dynamics of host plant use and species diversity in Polygonia butterflies (Nymphalidae).

The ability of insects to utilize different host plants has been suggested to be a dynamic and transient phase. During or after this phase, species can shift to novel host plants or respecialize on ancestral ones. Expanding the range of host plants might also be a factor leading to higher levels of net speciation rates. In this paper, we have studied the possible importance of host plant range for diversification in the genus Polygonia (Nymphalidae, Nymphalini). We have compared species richness between sistergroups in order to find out if there are any differences in number of species between clades including species that utilize only the ancestral host plants ('urticalean rosids') and their sisterclades with a broader (or in some cases potentially broader) host plant repertoire. Four comparisons could be made, and although these are not all phylogenetically or statistically independent, all showed clades including butterfly species using other or additional host plants than the urticalean rosids to be more species-rich than their sisterclade restricted to the ancestral host plants. These results are consistent with the theory that expansions in host plant range are involved in the process of diversification in butterflies and other phytophagous insects, in line with the general theory that plasticity may drive speciation.

Quantitative genetic variation in an island population of the speckled wood butterfly (Pararge aegeria).

Evidence of changes in levels of genetic variation in the field is scarce. Theoretically, selection and a bottleneck may lead to the depletion of additive genetic variance (V(A)) but not of nonadditive, dominance variance (V(D)), although a bottleneck may converse V(D) to V(A). Here we analyse quantitative genetic variation for the Speckled Wood butterfly Pararge aegeria on the island of Madeira about 120 generations after first colonisation. Colonisation of the island involved both a bottleneck and strong natural selection, changing the average value of traits. Several life history and morphological traits with varying levels of change since colonisation were analysed. In accordance with expectations, all traits except one showed relatively low levels of V(A), with an average heritability (h(2)) of 0.078. Levels of V(D) for these traits were relatively high, 20-94% of total variance and on average 80% of V(G). The exception was a morphological trait that probably had not experienced strong natural selection after colonisation, for which a h(2) of 0.27 was found. Another interesting observation is that the population seems resistant to inbreeding effects, which may be the result of purging of deleterious alleles.

Genetics of fluctuating asymmetry in pupal traits of the speckled wood butterfly (Pararge aegeria).

Fluctuating asymmetry (FA), small random differences between left and right, has been extensively used as a measure of individual quality, though its usefulness in that respect is controversial. Whether FA is heritable has implications for sexual selection theory and for its usefulness as an indicator of stress. Heritability (h(2)) of FA is, however, difficult to estimate precisely and reliably. Here we report h(2)s of FA for two pupal traits in the speckled wood butterfly (Pararge aegeria). We used a restriction error maximum likelihood (REML) analysis in combination with a jackknife procedure to analyse a large mixed offspring-parent/half-sib/full sib data set. A five-generation selection experiment provided a second set of narrow sense h(2)s. Narrow sense h(2)s were not significant and on average -0.029 (REML-analysis) and 0.031 (selection experiment) for the pupal segment covering the fore leg (LEG) and 0.057 and 0.004 for a SPOT on that segment. Estimated percentage dominance variances were 0.057 (LEG) and 0.027 (SPOT) and not significantly different from 0. The h(2) estimates had been slightly increased by cage effects. Average FA for LEG after five generations of selection were higher in the high lines than in the low lines, and the control lines were in between. No difference in FA between lines was found for SPOT. Although differences between lines were not significant, a slight h(2) (<3%) for LEG could not be excluded. The genetic effect was, however, small compared with the effect of foodplant quality. Larvae grown on foodplants that were not watered enough for good growth showed significantly higher FA for LEG, but not for SPOT, compared with larvae grown on good foodplants.

Evolutionary dynamics of host-plant specialization: a case study of the tribe Nymphalini.

Two general patterns that have emerged from the intense studies on insect-host plant associations are a predominance of specialists over generalists and a taxonomic conservatism in host-plant use. In most insect-host plant systems, explanations for these patterns must be based on biases in the processes of host colonizations, host shifts, and specialization, rather than cospeciation. In the present paper, we investigate changes in host range in the nymphalid butterfly tribe Nymphalini, using parsimony optimizations of host-plant data on the butterfly phylogeny. In addition, we performed larval establishment tests to search for larval capacity to feed and survive on plants that have been lost from the female egg-laying repertoire. Optimizations suggested an ancestral association with Urticaceae, and most of the tested species showed a capacity to feed on Urtica dioica regardless of actual host-plant use. In addition, there was a bias among the successful establishments on nonhosts toward plants that are used as hosts by other species in the Nymphalini. An increased likelihood of colonizing ancestral or related plants could also provide an alternative explanation for the observed pattern that some plant families appear to have been colonized independently several times in the tribe. We also show that there is no directionality in host range evolution toward increased specialization, that is, specialization is not a dead end. Instead, changes in host range show a very dynamic pattern.

Individual state controls temperature dependence in a butterfly (Lasiommata maera).

In ectotherms there is typically a strong and positive correlation between growth rate and ambient temperature when food is not limiting. However, the exact relationship between growth rate and temperature varies among populations in many species. As a consequence, it has been suggested that selection for a rapid increase in growth rate with temperature should be stronger in populations experiencing a high degree of time-stress, compared with populations experiencing little time-stress. In the present study we take this adaptive hypothesis further and investigate if variation in time-stress among individuals of a single population may affect the relationship between growth rate and ambient temperature. Time-stress was manipulated by rearing larvae of the butterfly Lasiommata maera in different day-length regimes. The results show that individuals experiencing a higher degree of time-stress increase their growth rates more in higher temperatures compared with individuals under less time-stress. Hence, the adaptive hypothesis was supported and the relationship between growth rate and temperature was highly state dependent. These findings may be of general importance for understanding the evolution of life histories in seasonal environments.

Plasticity in life-history traits.

We describe the impact of recent life-history plasticity theory on insect studies, particularly on the interface between genetics and plasticity. We focus on the three-dimensional relationship between three key life-history traits: adult size (or mass), development time and growth rate, and the connections to life cycle regulation, host plant choice, and sexual selection in seasonal environments. The review covers fitness consequences of variation in size, development time and growth rate, and effects of sex, photoperiod, temperature, diet, and perceived mortality risk on these traits. We give special attention to evidence for adaptive plasticity in growth rates because of the important effects of such plasticity on the expected relationships between development time and adult size and, hence, on the use of life-history, fitness, and optimality approaches in ecology, as well as in genetics.