The biology of Euphranta connexa was investigated to assess its potential as a biological control agent for introduction in North America against the invasive weeds Vincetoxicum rossicum (Kleopow) and V. nigrum (L.) (Apocynaceae). A range of temperatures suitable for development was determined for the pupal and egg stages of E. connexa. The pupa is the overwintering stage of E. connexa and does not exhibit a diapause, but undergoes a cold induced quiescence. The relationship of temperature to pupal developmental was investigated and did not differ among insects from sites over a range of altitude of 1300 m, indicating E. connexa may not be locally adapted. When adult female E. connexa emerged from pupae they had not developed eggs but 10 to 15 days later they had a full complement of developed eggs. Mating of E. connexa did not affect the egg load of females but egg load was reduced in females that had spent longer as pupae in cold conditions. There was evidence that females could reabsorb previously developed eggs. In surveys for larval and larval-pupal parasitoids of E. connexa infested seedpods of V. hirundinaria from three countries, there were a total of 1599 parasitoids from eight Hymenoptera families,and most parasitoids were Braconidae. Prior to the survey, the fruit midge, Contarinia asclepiadis (Diptera: Cecidomyiidae) was considered rare, but it was found in seedpods from all collection sites in Switzerland and infested about 9.2% of seedpods at two localities in 2013. Of 13 parasitoid morphospecies found in the survey, two braconid and one ichneumonid morphospecies were frequent and were parasitoids of E. connexa, three platygastrid morphospecies were probably parasitoids of C. asclepiadis, and three morphospecies were probably hyperparasitoids. The thesis research will allow development of effective methods for studying impact and host range of E. connexa, studies that are required to assess the potential as a biological control agent against V. rossicum and V. nigrum.

Animals and plants in temperate regions must adapt their life cycle to pronounced seasonal variation. The research effort that has gone into studying these cyclical life history events, or phenological traits, has increased greatly in recent decades. As phenological traits are often correlated to temperature, they are relevant to study in terms of understanding the effect of short term environmental variation as well as long term climate change. Because of this, changes in phenology are the most obvious and among the most commonly reported responses to climate change. Moreover, phenological traits are important for fitness as they determine the biotic and abiotic environment an individual encounters. Fine-tuning of phenology allows for synchronisation at a local scale to mates, food resources and appropriate weather conditions. On a between-population scale, variation in phenology may reflect regional variation in climate. Such differences can not only give insights to life cycle adaptation, but also to how populations may respond to environmental change through time. This applies both on an ecological scale through phenotypic plasticity as well as an evolutionary scale through genetic adaptation. In this thesis I have used statistical and experimental methods to investigate both the larger geographical patterns as well as mechanisms of fine-tuning of phenology of several butterfly species. The main focus, however, is on the orange tip butterfly, Anthocharis cardamines, in Sweden and the United Kingdom. I show a contrasting effect of spring temperature and winter condition on spring phenology for three out of the five studied butterfly species. For A. cardamines there are population differences in traits responding to these environmental factors between and within Sweden and the UK that suggest adaptation to local environmental conditions. All populations show a strong negative plastic relationship between spring temperature and spring phenology, while the opposite is true for winter cold duration. Spring phenology is shifted earlier with increasing cold duration. The environmental variables show correlations, for example, during a warm year a short winter delays phenology while a warm spring speeds phenology up. Correlations between the environmental variables also occur through space, as the locations that have long winters also have cold springs. The combined effects of these two environmental variables cause a complex geographical pattern of phenology across the UK and Sweden. When predicting phenology with future climate change or interpreting larger geographical patterns one must therefore have a good enough understanding of how the phenology is controlled and take the relevant environmental factors in to account. In terms of the effect of phenological change, it should be discussed with regards to change in life cycle timing among interacting species. For example, the phenology of the host plants is important for A. cardamines fitness, and it is also the main determining factor for oviposition. In summary, this thesis shows that the broad geographical pattern of phenology of the butterflies is formed by counteracting environmental variables, but that there also are significant population differences that enable fine-tuning of phenology according to the seasonal progression and variation at the local scale. / <p>At the time of the doctoral defense, the following paper was unpublished and had a status as follows: Paper 4: Manuscript.</p>