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.     

Implications of a temperature increase for host plant range: predictions for a butterfly

Although changes in phenology and species associations are relatively well‐documented responses to global warming, the potential interactions between these phenomena are less well understood. In this study, we investigate the interactions between temperature, phenology (in terms of seasonal timing of larval growth) and host plant use in the polyphagous butterfly Polygonia c‐album. We found that the hierarchy of larval performance on three natural host plants was not modified by a temperature increase as such. However, larval performance on each host plant and temperature treatment was affected by rearing season. Even though larvae performed better at the higher temperature regardless of the time of the rearing, relative differences between host plants changed with the season. For larvae reared late in the season, performance was always better on the herbaceous plant than on the woody plants. In this species, it is likely that a prolonged warming will lead to a shift from univoltinism to bivoltinism. The demonstrated interaction between host plant suitability and season means that such a shift is likely to lead to a shift in selective regime, favoring specialization on the herbaceous host. Based on our result, we suggest that host range evolution in response to temperature increase would in this species be highly contingent on whether the population undergoes a predicted shift from one to two generations. We discuss the effect of global warming on species associations and the outcome of asynchrony in rates of phenological change.     

Intraspecific variability in number of larval instars in insects

The number of larval instars varies widely across insect species. Although instar number is frequently considered to be invariable within species, intraspecific variability in the number of instars is not an exceptional phenomenon. However, the knowledge has remained fragmentary, and there are no recent attempts to synthesize the results of relevant studies. Based on published case studies, we show that intraspecific variability in the number of larval instars is widespread across insect taxa, occurring in most major orders, in both hemimetabolous and holometabolous insects. We give an overview of various factors that have been observed to affect the number of instars. Temperature, photoperiod, food quality and quantity, humidity, rearing density, physical condition, inheritance, and sex are the most common factors influencing the number of instars. We discuss adaptive scenarios that may provide ultimate explanations for the plasticity in instar number. The data available largely support the compensation scenario, according to which instar number increases in adverse conditions when larvae fail to reach a species-specific threshold size for metamorphosis. However, in Orthoptera and Coleoptera, there are some exceptional species in which instar number is higher in favorable conditions. In more specific cases, the adaptive value of the variability in instar number may be in reaching or maintaining the developmental stage adapted to hibernation, producing additional generations in multivoltine species, or increasing the probability of surviving in long-lasting adverse conditions.     

HOST PLANT UTILIZATION,HOST RANGE OSCILLATIONS AND DIVERSIFICATION IN NYMPHALID BUTTERFLIES: A PHYLOGENETIC INVESTIGATION

It has been suggested that phenotypic plasticity is a major factor in the diversification of life, and that variation in host range in phytophagous insects is a good model for investigating this claim. We explore the use of angiosperm plants as hosts for nymphalid butterflies, and in particular the evidence for past oscillations in host range and how they are linked to host shifts and to diversification. At the level of orders of plants, a relatively simple pattern of host use and host shifts emerges, despite the 100 million years of history of the family Nymphalidae. We review the evidence that these host shifts and the accompanying diversifications were associated with transient polyphagous stages, as suggested by the “oscillation hypothesis.” In addition, we investigate all currently polyphagous nymphalid species and demonstrate that the state of polyphagy is rare, has a weak phylogenetic signal, and a very apical distribution in the phylogeny; we argue that these are signs of its transient nature. We contrast our results with data from the bark beetles Dendroctonus, in which a more specialized host use is instead the apical state. We conclude that plasticity in host use is likely to have contributed to diversification in nymphalid butterflies.     

Investigating Concordance among Genetic Data, Subspecies Circumscriptions and Hostplant Use in the Nymphalid Butterfly Polygonia faunus

Subspecies are commonly used taxonomic units to formally describe intraspecific geographic variation in morphological traits. However, the concept of subspecies is not clearly defined, and there is little agreement about what they represent in terms of evolutionary units, and whether they can be used as reliably useful units in conservation, evolutionary theory and taxonomy. We here investigate whether the morphologically well-characterized subspecies in the North American butterfly Polygonia faunus are supported by genetic data from mitochondrial sequences and eight microsatellite loci. We also investigate the phylogeographic structure of P. faunus and test whether similarities in host-plant use among populations are related to genetic similarity. Neither the nuclear nor the mitochondrial data corroborated subspecies groupings. We found three well defined genetic clusters corresponding to California, Arizona and (New Mexico+Colorado). There was little structuring among the remaining populations, probably due to gene flow across populations. We found no support for the hypothesis that similarities in host use are related to genetic proximity. The results indicate that the species underwent a recent rapid expansion, probably from two glacial refugia in western North America. The mitochondrial haplotype network indicates at least two independent expansion phases into eastern North America. Our results clearly demonstrate that subspecies in P. faunus do not conform to the structuring of genetic variation. More studies on insects and other invertebrates are needed to better understand the scope of this phenomenon. The results of this study will be crucial in designing further experiments to understand the evolution of hostplant utilization in this species.     

Life history perspectives on pest insects: What’s the use?

In his seminal 1954 paper on the ‘population consequences of life history phenomena’, Cole noted that ‘these computations may have practical value in dealing with valuable or noxious species’. In the present paper, the question is asked: ‘is research based on evolutionary perspectives in general, and life history theory specifically, really useful for dealing with insect pests?’ Perhaps such theory‐based research is rather a luxury: time and resources would be better spent on entirely applied aspects of the problem. The conclusion of the present discussion is that having an evolutionary perspective guiding research is actually a very cost‐effective way of dealing with applied problems, as it provides a clear basis for interpretations, generalizations and predictions. Life history theory is a very central and necessary part of both population ecology and general evolutionary theory, and its specific usefulness in pest forecasting and management are discussed. Nevertheless, our ability to predict insect population dynamics is still limited, and so is our ability to make use of an insect’s life history traits to predict its propensity to become a pest. I suggest that the former shortcoming is largely due to poor understanding of insect life history plasticity. This, in turn, may partly be due to a paucity of studies where reaction norms are investigated as putative adaptations. I suggest that the latter shortcoming is due to problems inherent with studying life history traits as adaptations, for example the lack of an independent fitness model and the fact that life histories tend to form syndromes of coadapted traits. These points are illustrated with examples from my own work on non‐pest butterflies and from insect–Eucalyptus systems.     

Adaptation to fluctuating environments in a selection experiment with Drosophila melanogaster

A fundamental question in life‐history evolution is how organisms cope with fluctuating environments, including variation between stressful and benign conditions. For short‐lived organisms, environments commonly vary between generations. Using a novel experimental design, we exposed wild‐derived Drosophila melanogaster to three different selection regimes: one where generations alternated between starvation and benign conditions, and starvation was always preceded by early exposure to cold; another where starvation and benign conditions alternated in the same way, but cold shock sometimes preceded starvation and sometimes benign conditions; and a third where conditions were always benign. Using six replicate populations per selection regime, we found that selected flies increased their starvation resistance, most strongly for the regime where cold and starvation were reliably combined, and this occurred without decreased fecundity or extended developmental time. The selected flies became stress resistant, displayed a pronounced increase in early life food intake and resource storage. In contrast to previous experiments selecting for increased starvation resistance in D. melanogaster, we did not find increased storage of lipids as the main response, but instead that, in particular for females, storage of carbohydrates was more pronounced. We argue that faster mobilization of carbohydrates is advantageous in fluctuating environments and conclude that the phenotype that evolved in our experiment corresponds to a compromise between the requirements of stressful and benign environments.     

Molecular phylogenetic and morphological studies on the systematic position of Heracula discivitta reveal a new subfamily of Pseudobistonidae (Lepidoptera: Geometroidea)

Heracula discivitta Moore is an uncommon moth species currently recorded from India, Nepal and China. Although this species has traditionally been placed in Lymantriinae, its systematic position in Macroheterocera has been enigmatic due to its unique morphological features. Here we used molecular and morphological data to explore the systematic position of H. discivitta. Our molecular phylogenetic analyses indicate that this species is sister to Pseudobiston pinratanai Inoue, a member of a recently established monotypic family Pseudobistonidae. The examinations of morphological features further show that H. discivitta shares synapomorphies with Pseudobistonidae. Based on the analysis results, we propose a new subfamily of Pseudobistonidae (Heraculinae subfam.n. ) to accommodate H. discivitta. The resemblance of the habitus to that of the brahmaeid genus Calliprogonos Mell & Hering is discussed. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:urn:lsid:zoobank.org:pub:63D17850‐6D51‐4E03‐A5D6‐F9EF6E7AF402 .     

A SEX DIFFERENCE IN THE PROPENSITY TO ENTER DIRECT/DIAPAUSE DEVELOPMENT: A RESULT OF SELECTION FOR PROTANDRY

In monandrous mating systems with discrete nonoverlapping generations males should maximize the expected number of matings by starting to emerge before females. This is known as protandry. Moreover, Evolutionarily Stable Strategies (ESS) models show that the male emergence curve should be abruptly truncated before female emergence has ceased. In temperate areas where many insects have partial second generations, we accordingly predict that males should enter diapause development at an earlier date than should females, as a result of late-emerging males being penalized in terms of fewer mating opportunities. The decision to diapause or to develop directly is usually mediated by response to environmental stimuli of which day length is the most important. Hence we predict that the mechanism by which males enter diapause at an earlier date than females will be that of the male reaction norm for diapause development being shifted towards longer day lengths when compared to that of females. As a result of the greater tendency of males to enter diapause development, partial second generations that develop directly should be female biased. As a corollary, first generations should be male biased because some males of the first generation are from the previous year. The prediction that males should enter diapause development earlier in the season, i.e., at longer day lengths, as compared to females was corroborated by rearing Pieris napi under a variety of critical day length regimes producing mixed broods of directly developing and diapausing individuals, and by outdoor rearings of cohorts of larvae of P. napi and P. rapae initiated throughout the season. The prediction that partial second generations should be female biased was corroborated by laboratory rearings at constant temperature of P. napi (Pieridae), Polygonia c-album (Nymphalidae), and Pararge aegeria (Satyridae) under critical day length conditions, producing female-biased sex ratio under direct, and male-biased sex ratio under diapause development.