Ecology of Veliidae and Mesoveliidae (Hemiptera: Heteroptera: Gerromorpha) was studied in selected European species. The research of these non-gerrid semiaquatic bugs was especially focused on voltinism, overwintering with physiological consequences and wing polymorphism with dispersal pattern. Hypotheses based on data from field surveys were tested by laboratory, mesocosm and field experiments. New data on life history traits and their ecophysiological consequences are discussed in seven original research papers (four published journal papers, two submitted papers and one communication in conference proceedings), creating core of this thesis.

Abstract Females may either mate with one (monandry) or several (polyandry) males during a single breeding season. The polyandrous mating system has prompted numerous studies since the recognition of a widespread occurrence of mixed paternity among animals. Consequently, the benefits of polyandry have become well-established, and the female role in sexual selection upgraded. Females may gain both material and genetic benefits from multiple mating. Hence, the occurrence of polyandry is understandable, whereas monandry remains an evolutionary puzzle especially among species with male nutrient provisioning. I studied both the life history variation among female mating tactics and the adaptive significance of monandry in varying environmental conditions in the green-veined white butterfly [Pieris napi, (L. 1758)], which is a predominantly polyandrous species with nuptial feeding. I used a combination of explicit laboratory experiments and field studies. My results show that monandry and degrees of polyandry are distinct strategies with life history differences reaching beyond mating frequencies. Polyandry corresponded with a higher lifetime fecundity than monandry in P. napi. Polyandry was, however, associated with relatively low fecundity during the early days of reproduction. Thus, monandry is beneficial if time for reproduction is limited severely enough or other female traits or behaviours associated with polyandry are traded off against longevity. Due to temporal variation in reproductive rate among mating tactics, offspring of polyandrous females have less time to complete development. Accordingly, polyandrous females developed at a faster rate as larvae than monandrous ones under optimal conditions. Despite growth rate variation, monandrous females were more likely to contribute to additional summer generation in conditions that allow production of only a partial second generation, and thus monandry is favoured under these conditions. Genetic variation in female mating tactics will not only prevail if environmental conditions do not allow all individuals to contribute evenly to the directly breeding generation in bivoltine populations, but also if even the production of a single generation per year is time-limited. A general conclusion would be that seasonality and unpredictability of fitness in the wild drives the evolution of optimal female mating tactics and promotes the maintenance genetic variation in mating frequencies, regardless of the direct benefits of nuptial feeding. Even if a high degree of polyandry would be the most profitable mating tactic in an average year, strong annual variation in weather conditions and the duration of summer may create possibilities for a temporally fluctuating selection that promotes a co-existence of different mating tactics because variance of fitness is likely increase with an increasing mating frequency.

Native bees are only recently gaining attention for the extent to which they aid in pollination and ecosystem services. These services are threatened by predictions of warming temperatures if bees are not able to respond. Voltinism - the number of generations produced annually- can strongly influence thermal conditions experienced by both developing and adult bees based on emergence strategies for each voltinism type. Differences in experienced thermal conditions brought on by climate change could therefore affect upper thermal limits (UTL) in bees. This study observes UTLs across a foraging season within and among native bee species vi to elucidate the potential response by bees to a warming climate. Bees were collected across the field season in Central Virginia and subjected to dynamic ramping trials to determine the fatal knockdown point (FKP). Results show in both univoltine and multivoltine bees an increase in upper thermal tolerance as the foraging season progresses (in quadratic and linear fashions respectively). Within multivoltine bees, FKP was related to nest type; with stem-nesting bees having the highest FKP. All average FKPs were higher than historic air temperatures of the study region, but within several degrees of the highest recorded maximum temperature. The diversity of responses in native bees provides evidence for both genetic and plastic responses to extreme temperature. While bees still face a potential myriad of other issues brought on by climate change, the observed increases in FKP across warming temperatures offer tentatively hopeful predictions for limited physiological responses in native bees to a warming climate

Abstract Both the length of the season that is favourable for insect growth and reproduction and the number of generations emerging per season (voltinism) increase with decreasing latitude. Thus, time constraints on reproduction and juvenile development decrease with decreasing latitude, except where voltinism changes and time constraints suddenly increase as the season must be shared with one more generation. I studied the evolution of insect life histories in relation to time constraints from two perspectives: polymorphism and clinal variation. Life history polymorphism in seasonal environments was studied with the butterfly Pieris napi that has discrete life history strategies, and polymorphic natural populations. Experimental studies showed that asymmetric intraspecific larval competition and divergent timing of reproduction between the strategies may promote the maintenance of polymorphism. A simulation model showed that the divergent timing of reproduction between the strategies is sufficient to maintain polymorphism even in the absence of intraspecific competition. Clinal variation was studied empirically with four geometrid moths (Cabera exanthemata, Cabera pusaria, Chiasmia clathrata and Lomaspilis marginata) and generally in theory. Due to latitudinal saw-tooth variation in time constraints, traditional theory predicts a saw-tooth cline in body size and development time. A common garden experiment with the four geometrid moths did not support the traditional theory even when a saw-tooth cline in body size was found, suggesting that the theory is based on unrealistic assumptions. A theoretical analysis showed that reproductive effort should be high in populations under intense time constraints and low in populations experiencing no time constraints, resulting in a saw-tooth cline. In the four geometrid moths, support for these predictions was found, although the observed clinal variation deviated from the predicted pattern. The results imply that clinal variation is expected in almost any continuous life history trait, whereas polymorphism of different strategies may emerge when life histories fall into discrete categories. When inferring the evolution of a single trait, complex interdependencies among several traits should be considered, as well as the possibility that the time constraints are not similar for each generation in multivoltine populations. / Tiivistelmä Hyönteisten kasvu ja kehitys ovat pääsääntöisesti mahdollisia vain kesän aikana. Etelään päin mentäessä kesä pitenee, ja saman kesän aikana kehittyvien hyönteissukupolvien määrä kasvaa. Kesän pituus aiheuttaa lisääntymiseen ja toukkien kasvuun kohdistuvan aikarajoitteen, joka heikkenee etelään päin siirryttäessä. Aikarajoite kuitenkin tiukkenee siellä, missä yksi uusi sukupolvi ehtii juuri kehittymään saman kesän aikana, sillä kesä on nyt jaettava useamman sukupolven kesken. Väitöstyössä tarkastelin hyönteisten elinkierto-ominaisuuksien evoluutiota suhteessa aikarajoitteisiin sekä diskreetin että jatkuvan muuntelun näkökulmista. Diskreettiä muuntelua tutkin lanttuperhosella (Pieris napi), jolla esiintyy diskreettejä elinkiertostrategioita. Kokeellisesti osoitin, että toukkien välinen kilpailu on epäsymmetristä, mikä yhdessä eri elinkiertostrategioiden erilaisen lisääntymisen ajoittumisen kanssa voi ylläpitää diskreettiä muuntelua. Simulaatiomalli osoitti, että erilaiset elinkiertostrategiat voivat säilyä populaatiossa pelkästään niiden erilaisen lisääntymisen ajoittumisen ansiosta. Elinkierto-ominaisuuksien jatkuvaa muuntelua tutkin neljän mittariperhosen (Cabera exanthemata, Cabera pusaria, Chiasmia clathrata ja Lomaspilis marginata) avulla ja teoreettisesti yleisellä tasolla. Aikaisempi teoria ennustaa ruumiinkoon ja kehitysajan muuntelevan sahalaitakuvion mukaisesti siirryttäessä pohjoisesta etelään, koska aikarajoitteet muuntelevat samalla tavalla. Tämä teoria perustunee epärealistisiin oletuksiin, koska kokeelliset tulokset eivät tukeneet teoriaa silloinkaan, kun mittariperhosten ruumiinkoko muunteli ennustetulla tavalla. Teoreettinen tutkimus osoitti, että myös lisääntymispanostuksen tulisi muunnella sahalaitakuvion mukaisesti suhteessa kesän pituuteen siten, että se on korkeimmillaan siellä, missä aikarajoitteet ovat tiukat. Mittariperhosten tutkiminen antoi jossain määrin tukea tälle ennusteelle. Tulosten perusteella jatkuvaa maantieteellistä muuntelua ennustetaan elinkierto-ominaisuuksille, jotka muuntelevat jatkuvalla asteikolla. Erilaiset elinkiertostrategiat voivat sen sijaan säilyä populaatiossa, jos elinkierto-ominaisuuksien muuntelu on diskreettiä. Eri ominaisuuksien monimutkaiset vuorovaikutukset sekä eri sukupolvien mahdollisesti kokemat erilaiset aikarajoitteet olisi syytä tuntea, kun tarkastelun kohteena on yksittäisen ominaisuuden evoluutio.

There is much evidence that herbivory is a key innovation for the tremendous success of insect. In this thesis I have investigated different aspects of host plant utilization and phenotypic plasticity using the polyphagous comma butterfly, Polygonia c-album. Even though external conditions affect a phenotypic plastic response, the outcome is often influenced by a genetic background which may differ among populations. In Paper I we suspected the genetic background to seasonal polymorphism to be X-linked. However, results from interspecific hybridization between two populations suggested that diapause response is instead inherited in a mainly autosomally additive fashion, with a possible influence of sexual antagonism on males. In Paper II we showed that female oviposition preference is not a plastic response influenced by larval experience, but has a genetic background coupled to host plant suitability. Further, there is a strong individual correlation between larval host plant acceptance and female host plant specificity (Paper III). We believe this to be a larval feed-back genetically linked to female host specificity: offspring to ‘choosy’ specialist mothers benefit by remaining on the original host while offspring to less discriminating generalist mothers should risk inspecting the surroundings, thus compensating for potential poor female choice. In the larval mid-gut, genes are differentially expressed depending on host plant diet (Paper IV). Therefore, we expected to find fitness consequences of host plant switch. However, although growth rate was affected in a few treatments, larvae were generally surprisingly good at adjusting to new diets (Paper V). To conclude, host plant choice in both female and larval life stage is connected to performance. Combined with increased understanding about the plastic response to diet intake and seasonal polymorphism we have gained further insights into the processes of local adaptations and speciation in the Lepidoptera. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Submitted Manuscript; Paper 5: Manuscript