[Truncated abstract] Despite the costs associated with mating, females of many taxa solicit multiple mates during a single reproductive event (polyandry). Polyandry is clearly adaptive when females gain direct benefits from males at mating. However, polyandry has also been shown to increase female fitness in the absence of direct benefits. Thus, a number of genetic benefit hypotheses have been developed to account for the origin of this behaviour. Although not mutually exclusive, a distinction lays between genetic benefits that propose defense against reproductive failure (nonadditive genetic effects), and those that propose benefits from intrinsic sire effects (additive genetic effects). Nonadditive genetic benefits of polyandry have been documented in a number of species; by soliciting multiple mates females can avoid inbreeding and other forms of incompatibility between parental genotypes. Polyandry may also increase female reproductive success when genetically superior males have greater success in sperm competition, and produce better quality offspring. An inevitable consequence of polyandry is that sperm from rival males will overlap in the female reproductive tract and compete to fertilise the ova. The outcome of sperm competition is typically determined by bias in sperm use by the females, interactions between parental genotypes, and ejaculate characteristics that provide a fertilisation advantage. Thus, sperm competition is recognised as a persuasive force in the evolution of male reproductive traits. Comparative analyses across species, and competitive mating trials within species have suggested that sperm competition can influence the evolution of testis size and sperm production, and both sperm form and sperm function. ... After six generations of selection I observed phenotypic divergence in litter size - litter size increased in the polyandrous lines but not in the monandrous lines. This result was not attributable to inbreeding depression, or environmental/maternal effects associated with mating regime. Genetic benefits associated with polyandry could account for this result if increased litter size were attributable to increased embryo survival. However, males from the polyandrous lineages were subject to sperm competition, and evolved ejaculates with more sperm, suggesting that evolutionary increases in litter size may in part be due to improved male fertility. Finally, Chapter Five is an investigation of the natural variation in levels of polyandry in the wild, and the potential for sperm competition to drive macroevolutionary changes in male reproductive traits among geographically isolated island populations of house mice. I sampled seven island populations of house mice along the coast of Western Australia and, by genotyping pregnant females and their offspring, determined the frequency of multiply sired litters within each population. I applied the frequency of multiple paternity as an index of the risk of sperm competition, and looked for selective responses in testis size and ejaculate traits. I found that the risk of sperm competition predicted testis size across the seven island populations. However, variation in sperm traits was not explained by the risk of sperm competition. I discuss these results in relation to sperm competition theory, and extrinsic factors that influence ejaculate quality.

This thesis presents two studies where natural and sexual selection have interacted to evolve sexual behaviours. The thesis uses mathematical modelling to understand how these forces have caused each behaviour to evolve. This is useful because the results allow for reflection on the potential role of sexual selection in adaptation of these species to a changing environment. The first study is of early male arrival to spring breeding grounds in migratory avian species, this is termed protandry. The study explores the main hypotheses for avian protandry and then tests the susceptibility of each hypothesis to changing environment. The second study is of convenience polyandry in species where there is conflict over mating rate. Females have multiple strategies to avoid harassive males but strategies vary in cost and success rate; she must balance her strategy use to minimise her fitness depreciation. The study identifies the main factors that cause convenience polyandry to evolve and paves the way for future studies to investigate if sexual selection over resistance strategy provides these species a future advantage in adaptation to a changing environment.

Estudos relacionados ao sistema de acasalamento nos insetos permite a compreensão dos aspectos evolutivos do comportamento reprodutivo de uma espécie. Esses sistemas são classificados, geralmente, pelo número de parceiros sexuais de um indivíduo durante um período de acasalamento. Em fêmeas, pode ocorrer monogamia, quando a cópula for realizada com somente um macho e, poliandria, quando há ocorrência de múltiplas cópulas com machos diferentes ou com o mesmo macho (cópulas repetidas). Em espécies como a broca-do-café, Hypothenemus hampei, que passam a maior parte de sua vida (ovo a adulto) no interior do fruto de café e possuem ciclo multivoltino os estudos envolvendo o sistema de acasalamento são escassos. Portanto, este trabalho visou estudar o acasalamento da broca-do-café, avaliando se há ocorrência de poliandria e os possíveis efeitos de múltiplas cópulas e cópulas repetidas sobre a bioecologia da fêmea e sua prole. Os resultados revelaram pela primeira vez a ocorrência de poliandria em H. hampei. Foi observado um maior número de recópulas em fêmeas colonizadoras, quando comparado com as fêmeas em oviposição, sugerindo que a condição fisiológica da fêmea foi um fator decisivo no momento da recópula. Além disso, a duração da recópula foi menor nas fêmeas em oviposição do que em fêmeas colonizadoras. A fecundidade foi afetada negativamente pela poliandria, sugerindo a presença de um custo associado a realização de múltiplas cópulas. A longevidade não foi influenciada pelo sistema de acasalamento, demonstrando que a poliandria não influenciou o tempo de vida das fêmeas. A recuperação larva:adulto em H. hampei não foi influenciada pelo sistema de acasalamento. Por outro lado, a taxa de eclosão de larvas foi superior na condição de poliandria, comparativamente à monogamia. / Researches about insects mating systems are important to understand the aspects of reproductive behavior evolution. Mating systems are usually grouped according to the number of mates during mating. Insect females mating system are classified as monogamy when there is only one copulation per male while polyandry refers to multiple mating with different males or with the same male (repeated mating). There is a lack of researches dealing with mating systems in species like the coffee berry borer, Hypothenemus hampei, which spend most of its life cycle (egg to adult) inside of the coffee berry and has a multivoltine cycle. This research aimed to study the mating system of H. hampei females, verifying the existence of polyandry and evaluating the effects of the multiple and repeated mating on the bioecology of the female and its offspring. The current study showed for the first time the occurrence of polyandry on females of H. hampei. The number of remating in colonizing females was higher than in the females under oviposition, thus the female physiological condition is crucial on remating behavior. The copula duration was shorter in females under oviposition than in the colonizing females. Fecundity was negatively affected by polyandry suggesting the existence of costs associated with multiple mating by females. Longevity were not influenced by the mating systems, showing that polyandry does not affect female lifetime. Although egg hatching was superior in polyandry compared to monogamy, the larva:adult recovery rate in H. hampei was not affected by the mating system.

Thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg in fulfilment of the requirements for the degree of Master of Science. / Breeding was observed in the foam nest frog, Chiromantis xerampelina, over three breeding seasons The mating pattern was characterized by an extended breeding season with a male-biased operational sex ratio and asynchronous and unpredictable female arrival. At more than 90% of nests, from one to seven unpaired males ('peripheral males') gathered around the amplexing pair during nest construction. Those peripheral males closest to the pair competed 'With each other, and with the amplexing male, to position their cloacae against the female's cloaca during oviposition bouts. In a detailed study of a single population, over 80% of males were observed as peripheral males, and 57% of males were observed both in amplexus and as peripheral males. Male mating success and participation at nests was unrelated to size or weight. Chorus participation was the best predictor of male, mating success and participation at nests. The most plausible explanation for the presence of peripheral males was a sperm competition hypothesis; namely that peripheral males compete with the amplexing male for fertilizations by shedding sperm into the nest. I collected data on body mass and testis mass for 13 African anurans. Using additional published data on 19 Japanese anurans, an allometric relationship between body mass and testis mass was calculated. using 16 genera as independent data points. This revealed that C. xerampelina have testes fourteen times heavier than predicted on the basis of body mass. This is consistent with a trend seen in several taxa where testis size is related to the intensity of sperm competition. An additional experiment, in which the arnplexing male was prevented from shedding sperm into the nest, showed that peripheral males are capable of fertilizing eggs. I conclude that peripheral males are engaged in an opportunistic alternative mating tactic involving sperm cosnpetition. More than half the observed females bred polyandrously, some mating with up to three males, This was the result of amplexing males dismounting between nesting sessions, and males displacing one another from amplexus. / Andrew Chakane 2018

The thesis underpinning this dissertation is based on the Zulu people’s belief in the living-dead and the fact that the latter control the lives of their living descendants. The living descendants use names to express their dissatisfaction with one another. The families perform rituals to appease the living-dead. The living-dead are perceived as guardian angels who are closer to God. They are believed to be able to reward good behavior and reprimand those who are not behaving in an acceptable manner. Names, as Bhengu (1975:52) states, connect people to their living-dead. It is therefore important that this relationship with the living-dead is maintained. Friction is always going to be a problem in polygynous families. Avoiding confrontation is important to people who want to appease their living-dead, who control their lives. Personal names act as a deterrent to angering the living-dead. In a situation where getting even is not an option, opting for a name to voice your disapproval is the easy way out. Names become communication channels between members of the family and the community at large. This study is done from an ethnographic perspective with an attempt to fully describe a variety of aspects of a cultural group to enhance understanding of the people being studied. Spradley (1980:3) states that “The essential core of ethnography is the concern with the meaning of actions and events to the people we seek to understand”. This understanding may be seen as the basis of the method; through ethnographic study, the researcher comes to comprehend, through detailed observation, the existences of peoples and their cultures. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2012.

Elaborate sexual ornaments evolve because mate choice exerts strong sexual selection favouring individuals with high levels of ornament expression. Consequently, even at evolutionary equilibrium, life history theory predicts that ornamental traits should be under directional sexual selection that opposes contrasting selection to reduce the costs associated with their maintenance. Otherwise, the resources used to maintain ornaments should be used to improve other life history functions. Elaborate female ornaments have only evolved in a few species, despite females commonly experiencing strong sexual selection. One explanation for this rarity is that male preferences for female ornaments may be self-limiting: females with higher mating success become less attractive because of the lower paternity share they provide to mates with every additional sperm competitor. The unusual species in which female ornaments do occur can provide rare insight into how selection can favour the expression of expensive characters in females despite their costs. The main goal of my thesis was to determine how sexual selection acts on exaggerated sexual ornaments, and give new insight into how these ornaments may have evolved, in spite of the self-limiting nature of selection on male preferences. To determine the strength of sexual selection acting on female ornamentation in dance flies, we developed new microsatellite markers to assess polyandry rates by genotyping stored sperm in wild female dance flies. We first used polyandry rates to determine whether ornament expression was associated with higher mating success in female Rhamphomyia longicauda, a species that has evolved two distinct and exaggerated female ornaments. Contrary to our predictions, we found no evidence that females with larger ornaments enjoy higher mating success. We then compared polyandry rates in R. longicauda to those of two other species of dance fly, one (Empis aestiva) that has i independently evolved female ornaments on its legs, and another (E. tessellata) that does not possess any discernable female ornaments. We also estimated the opportunity for sexual selection, which we found to be similar and relatively low in all three species. Moreover, the standardized sexual selection gradients for ornaments were weak and non-significant in all three species. Females with more elaborate ornaments, in both within- and cross-species comparisons, therefore did not enjoy higher mating success. Overall, these results suggested that sexual selection operates rather differently in females compared to males, potentially explaining the general rarity of female ornaments. Our amplifications of stored sperm were able to reveal more than just mate numbers. We developed new methods to study patterns of sperm storage in wild female dance flies. We investigated how the skew in sperm genotypes from mixed sperm stores changed with varying levels of polyandry. Our data suggested that sperm stores were dominated by a single male in R. longicauda, and that the proportion of sperm contributed by this dominant male was largely independent of the number of rival males’ sperm present in the spermatheca. These results were consistent with the expectation of males using sperm ‘offence strategies’ in sperm competition and that the most successful male is likely to be the female’s last partner before oviposition. As a whole, my thesis contributed new molecular resources for an understudied and fascinating group of organisms. It exploited these new resources to provide the first estimates of lifetime mating success in several related species, and suggested that the general prediction that ornament expression should covary with sexual selection intensity does not seem to hold in this group. Instead, both the unusual prevalence of ii ornaments and the inconsistent evidence for sexual selection that sustains them in dance flies may owe their existence to the confluence of two important factors. First, the conditions under which sperm competition occurs: as last male precedence is likely, males are selected to prefer the most gravid females to secure a high fraction of her offspring’s paternity as they are unlikely to mate again before oviposition. Second, potent sexually antagonistic coevolution between hungry females and discerning males: females have evolved ornaments to disguise their stage of egg maturity to receive the benefits of nuptial gifts, while males face the challenge of distinguishing between gravidity and ornamentation in females.

Estudos relacionados ao sistema de acasalamento nos insetos permite a compreensão dos aspectos evolutivos do comportamento reprodutivo de uma espécie. Esses sistemas são classificados, geralmente, pelo número de parceiros sexuais de um indivíduo durante um período de acasalamento. Em fêmeas, pode ocorrer monogamia, quando a cópula for realizada com somente um macho e, poliandria, quando há ocorrência de múltiplas cópulas com machos diferentes ou com o mesmo macho (cópulas repetidas). Em espécies como a broca-do-café, Hypothenemus hampei, que passam a maior parte de sua vida (ovo a adulto) no interior do fruto de café e possuem ciclo multivoltino os estudos envolvendo o sistema de acasalamento são escassos. Portanto, este trabalho visou estudar o acasalamento da broca-do-café, avaliando se há ocorrência de poliandria e os possíveis efeitos de múltiplas cópulas e cópulas repetidas sobre a bioecologia da fêmea e sua prole. Os resultados revelaram pela primeira vez a ocorrência de poliandria em H. hampei. Foi observado um maior número de recópulas em fêmeas colonizadoras, quando comparado com as fêmeas em oviposição, sugerindo que a condição fisiológica da fêmea foi um fator decisivo no momento da recópula. Além disso, a duração da recópula foi menor nas fêmeas em oviposição do que em fêmeas colonizadoras. A fecundidade foi afetada negativamente pela poliandria, sugerindo a presença de um custo associado a realização de múltiplas cópulas. A longevidade não foi influenciada pelo sistema de acasalamento, demonstrando que a poliandria não influenciou o tempo de vida das fêmeas. A recuperação larva:adulto em H. hampei não foi influenciada pelo sistema de acasalamento. Por outro lado, a taxa de eclosão de larvas foi superior na condição de poliandria, comparativamente à monogamia. / Researches about insects mating systems are important to understand the aspects of reproductive behavior evolution. Mating systems are usually grouped according to the number of mates during mating. Insect females mating system are classified as monogamy when there is only one copulation per male while polyandry refers to multiple mating with different males or with the same male (repeated mating). There is a lack of researches dealing with mating systems in species like the coffee berry borer, Hypothenemus hampei, which spend most of its life cycle (egg to adult) inside of the coffee berry and has a multivoltine cycle. This research aimed to study the mating system of H. hampei females, verifying the existence of polyandry and evaluating the effects of the multiple and repeated mating on the bioecology of the female and its offspring. The current study showed for the first time the occurrence of polyandry on females of H. hampei. The number of remating in colonizing females was higher than in the females under oviposition, thus the female physiological condition is crucial on remating behavior. The copula duration was shorter in females under oviposition than in the colonizing females. Fecundity was negatively affected by polyandry suggesting the existence of costs associated with multiple mating by females. Longevity were not influenced by the mating systems, showing that polyandry does not affect female lifetime. Although egg hatching was superior in polyandry compared to monogamy, the larva:adult recovery rate in H. hampei was not affected by the mating system.

La sélection de la parentèle est le concept actuellement le plus fréquemment avancé pour justifier l’évolution et le maintien d'une caste ouvrière stérile chez les Hyménoptères sociaux. La fourmi méditerranéenne Cataglyphis cursor possède plusieurs traits biologiques qui font de cette espèce un modèle particulièrement intéressant pour tester les prédictions de la théorie de la sélection de la parentèle, le plus important d'entre eux étant la capacité des ouvrières, qui ne s'accouplent jamais, à produire une descendance mâle (haploïde) par parthénogenèse arrhénotoque, ou femelle (diploïde) par parthénogenèse thélytoque. Nos analyses génétiques, basées sur des marqueurs microsatellites développés au préalable pour cette espèce, ont révélé que les reines utilisent la reproduction sexuée et asexuée respectivement pour la production de la caste ouvrière et reproductrice. L'analyse du pedigree des reproductrices issues de la reproduction asexuée nous a permis d'identifier le mécanisme cytologique de la parthénogenèse thélytoque et d'estimer la proportion de reines issues de la reproduction des ouvrières au sein de la population. De plus, bien que les reines soient capables de produire une descendance diploïde (femelle) par parthénogenèse thélytoque, elles ont conservé la reproduction sexuée pour la caste ouvrière et s'accouplent avec plusieurs mâles. Ceci indique que la reproduction sexuée a une fonction importante au niveau de la colonie, et nous avons testé certaines des hypothèses avancées pour justifier l'évolution de la polyandrie. Finalement, nous avons étudié l'impact des stratégies de dispersion de cette espèce sur le sex-ratio de la descendance sexuée. Ces résultats confirment l'intérêt que représente l'étude des stratégies reproductrices chez les Hyménoptères sociaux pour tester les prédiction de diverses théories en biologie évolutives et ouvrent également de nouvelles perspectives de recherche, tant chez C. cursor que chez d'autres espèces appartenant au genre Cataglyphis. / Kin selection is, to date, the most widely accepted theory to justify the evolution of a sterile worker caste among social Hymenoptera. The Mediterranean ant Cataglyphis cursor represents an interesting biological model for several reasons, the most important of them being the ability for unmated workers to produce haploid (male) offspring, through arrhenotokous parthenogenesis, and diploid (female) offspring, through thelytokous parthenogenesis. Our genetic analyses, based on microsatellite loci developed for this purpose, revealed that queens selectively use sexual and asexual reproduction to produce workers and sexuals, respectively. Pedigree analyses allowed us to identify the cytological mechanism involved in thelytokous parthenogenesis and to estimate the proportion of worker-produced queens in the study population. Although C. cursor queens do not require mating to produce diploid offspring, they have retained sexual reproduction and mate multiply with up to 8 males. This suggests that sexual reproduction has important benefits for colony function, and we tested several hypotheses accounting for the evolution of polyandry. Eventually, we studied the effect of dispersal strategies on sex-ratio of the sexual brood. These results confirm the interest of investigating the reproductive strategies of social Hymenoptera to test the predictions of diverse theories in the field of evolutionary biology, and open new research perspectives in C. cursor and other ants of the Cataglyphis genera. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Abstract Sexual selection results from competition between individuals over access to gametes of the opposite sex. Starting with anisogamy, with females producing larger and fewer gametes than males, females usually invest more in reproduction and thus have lower potential reproductive rates than males. Therefore males are traditionally suggested to compete over females whereas females are choosy. The different evolutionary interests between the sexes result in different selection regimes considering e.g. mating rates. In some species, this conflict is even seen as apparent struggles between males and females over mating decision and/or duration, resulting in antagonistic coevolution with both sexes evolving adaptations to gain more control over the other sex. In this thesis, I studied the role of A. paludum female abdominal spines as an adaptation to reject male mating attempts. I studied both the effect of polyandry and multiple matings on female fecundity to distinguish between the potential material and genetic benefits from mating to females. I also explored the role of various male and female morphological traits on precopulatory sexual selection and the consequent female reproductive output. My results show that female abdominal spines are likely evolved through arms races between the sexes to increase female control over mating decision but according to my studies, female spines are not under active selection. Instead, certain male morphological types were more successful in achieving matings, and mating with these males also increased female fecundity. I found that multiple matings increase female fecundity up to a point, supporting a theory of optimal female mating rate. According to my results, the benefits from multiple mating to A. paludum females result from material origin whereas polyandry per se is even detrimental to female fecundity. / Tiivistelmä Seksuaalivalinta on pohjimmiltaan seurausta yksilöiden välisestä, lisääntymiseen liittyvästä kilpailusta toisen sukupuolen sukusoluista. Seksuaalivalinta on evoluutiomekanismi, joka osaltaan on johtanut sukupuolten välisiin eroihin monissa morfologisissa ja käyttäytymiseen liittyvissä ominaisuuksissa. Sukupuolet eroavat jo siinä, että naaraat tuottavat vähemmän mutta suurempia sukusoluja kuin koiraat. Koska sukupuolet siis investoivat lisääntymiseen jo alkuvaiheessa eri tavalla, niiden välillä on aina jonkin asteinen seksuaalikonflikti. Konfliktin taustalla on sukupuoliin kohdistuva erilainen valintapaine; koiraat saavuttavat usein optimaalisen hedelmällisyytensä suuremmilla parittelujen määrällä kuin naaraat. Joillakin eläinlajeilla seksuaalikonflikti on niin ilmeinen, että se ilmenee koiraan ja naaraan välisinä kamppailutilanteina parittelupäätöksen ja/tai parittelun keston suhteen. Seksuaalikonflikti voi johtaa molemmilla sukupuolilla erilaisiin sopeumiin, jotka lisäävät yksilön kontrollia parittelujen suhteen. Tässä väitöskirjatyössä tutkin naaraan takapäässä sijaitsevien abdominaalisten piikkien merkitystä järvivesimittari A. paludum-naaraiden kyvyssä torjua parittelemaan pyrkiviä koiraita. Tutkin myös polyandrian ja toistuvien parittelujen vaikutusta naaraan hedelmällisyyteen saadakseni selville, lisäävätkö koiraasta johtuvat mahdolliset materiaaliset tai geneettiset tekijät naaraan hedelmällisyyttä. Lisäksi selvitin useiden koiraan ja naaraan morfologisten ominaisuuksien yhteyttä parittelua edeltävään seksuaalivalintaan sekä naaraan hedelmällisyyteen. Tulokseni osoittavat, että naaraan abdominaaliset piikit ovat todennäköisesti kehittyneet seksuaalikonfliktin seurauksena lisäämään naaraan kontrollia parittelupäätöksen suhteen. Niihin ei kuitenkaan nykyisellään näytä kohdistuvan merkittävää valintapainetta. Sen sijaan morfologialtaan tietynlaiset koirastyypit vaikuttavat olevan aktiivisen valinnan kohteena ja parittelu tällaisten koiraiden kanssa lisää myös naaraan hedelmällisyyttä. Toistuvat parittelut saman koiraan kanssa lisäävät naaraan hedelmällisyyttä tiettyyn optimiin asti, mikä tukee teoriaa optimaalisesta parittelujen määrästä. Sen sijaan polyandria itsessään vaikuttaa naaraan hedelmällisyyteen jopa heikentävästi. Niinpä A. paludum-naaraiden saama hyöty useista paritteluista näyttää olevan materiaalista, kun taas polyandriasta saatavia geneettisiä etuja ei tässä tutkimuksessa tullut ilmi.

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.