Dispersal and related life history traits in the Glanville fritillary butterfly

Most studies of life history evolution are based on the assumption that species exist at equilibrium and spatially distinct separated populations. In reality, this is rarely the case, as populations are often spatially structured with ephemeral local populations. Therefore, the characteristics of metapopulations should be considered while studying factors affecting life history evolution. Theoretical studies have examined spatial processes shaping the evolution of life history traits to some extent, but there is little empirical data and evidence to investigate model predictions. In my thesis I have tried to bridge the gap between theoretical and empirical studies by using the well-known Glanville fritillary (Melitaea cinxia) metapopulation as a model system. The long-term persistence of classic metapopulations requires sufficient dispersal to establish new local populations to compensate for local extinctions. Previous studies on the Glanville fritillary have shown that females establishing new populations are not a random sample from the metapopulation, but they are in fact more dispersive than females in old populations. Many other life-history traits, such as body size, fecundity and lifespan, may be related to dispersal rate. Therefore, I examined a range of correlated traits for their evolutionary and ecological consequences. I was particularly interested in how the traits vary under natural environmental conditions, hence all studies were conducted in a large (32 x 26 m) outdoor population cage built upon a natural habitat patch. Individuals for the experiments were sampled from newly-established and old populations within a large metapopulation. Results show that females originating from newly-established populations had higher within-habitat patch mobility than females from old populations. I showed that dispersal rate is heritable and that flight activity is related to variation in a gene encoding the glycolytic enzyme phosphoglucose isomerase. Both among-individual and among-population variation in dispersal are correlated with the reproductive performance of females, though I found no evidence for a trade-off between dispersal and fecundity in terms of lifetime egg production or clutch size. Instead, the results suggest that highly dispersive females from newly-established populations have a shorter lifespan than females from old populations, and that dispersive females may pay a cost in terms of reduced lifetime reproductive success due to increased time spent outside habitat patches. In summary, the results of this thesis show that genotype-dependent dispersal rate correlates with other life history traits in the Glanville fritillary, and that the rapid turnover of local populations (extinctions and re-colonisations) is likely to be the mechanism that maintains phenotypic variation in many life history traits at the metapopulation level.

Climate forcing on avian life history

In the 21st century, human-induced global climate change has been highlighted as one of the most serious threats to ecosystems worldwide. According to global climate scenarios, the mean temperature in Finland is expected to increase by 1.8 4.0°C by the end of the century. The regional and seasonal change in temperature has predicted to be spatially and temporally asymmetric, where the High-Arctic and Antarctic areas and winter and spring seasons have been projected to face the highest temperature increase. To understand how species respond to the ongoing climate change, we need to study how climate affects species in different phases of their life cycle. The impact of climate on breeding and migration of eight large-sized bird species was studied in this thesis, taking food availability into account. The findings show that climatic variables have considerable impact on the life-history traits of large-sized birds in northern Europe. The magnitude of climatic effects on migration and breeding was comparable with that of food supply, conventionally regarded as the main factor affecting these life-history traits. Based on the results of this thesis and the current climate scenarios, the following not mutually exclusive responses are possible in the near future. Firstly, asymmetric climate change may result in a mistiming of breeding because mild winters and early spring may lead to earlier breeding, whereas offspring are hatching into colder conditions which elevate mortality. Secondly, climate induced responses can differ between species with different breeding tactics (income vs. capital breeding), so that especially capital breeders can gain advantage on global warming as they can sustain higher energy resources. Thirdly, increasing precipitation has the potential to reduce the breeding success of many species by exposing nestlings to more severe post-hatching conditions and hampering the hunting conditions of parents. Fourthly, decreasing ice cover and earlier ice-break in the Baltic Sea will allow earlier spring migration in waterfowl. In eiders, this can potentially lead to more productive breeding. Fifthly, warming temperatures can favour parents preparing for breeding and increase nestling survival. Lastly, the climate-induced phenological changes in life history events will likely continue. Furthermore, interactions between climate and food resources can be complex and interact with each other. Eiders provide an illustrative example of this complexity, being caught in the crossfire between more benign ice conditions and lower salinity negatively affecting their prime food resource. The general conclusion is that climate is controlling not only the phenology of the species but also their reproductive output, thus affecting the entire population dynamics.

Cultural Trauma and Life Stories

The purpose of this work is to use the concepts of human time and cultural trauma in a biographical study of the turning points in the recent history of Estonia. This research is primarily based on 148 in-depth biographical interviews conducted in Estonia and Sweden in 1995-2005, supplemented by excerpts from 5 collections and 10 individually published autobiographies. The main body of the thesis consists of six published and of two forthcoming separate refereed articles, summarised in the theoretical introduction, and Appendix of the full texts of three particular life stories. The topic of the first article is the generational composition and the collective action frames of anti-Soviet social mobilisation in Estonia in 1940-1990. The second article details the differentiation of the rites of passage and the calendar traditions as a strategy to adapt to the rapidly changed political realities, comparatively in Soviet Estonia and among the boat-refugees in Sweden. The third article investigates the life stories of the double-minded strategic generation of the Estonian-inclined Communists, who attempted to work within the Soviet system while professing to uphold the ideals of pre-war Estonia. The fourth article is concentrated on the problems of double mental standards as a coping strategy in a contradictory social reality. The fifth article implements the theory of cultural trauma for the social practice of singing nationalism in Estonia. The sixth article bridges the ideas of Russian theoreticians concerning cultural dialogue and the Western paradigm of cultural trauma, with examples from Estonian Russian life stories. The seventh article takes a biographical look at the logic of the unraveling of cultural trauma through four Soviet decades. The eighth article explores the re-shaping of citizen activities as a strategy of coping with the loss of the independent nation state, comparatively in Soviet Estonia and among Swedish Estonians. Cultural trauma is interpreted as the re-ordering of the society s value-normative constellation due to sharp, violent, usually political events. The first one under consideration was caused by the occupations of the Republic of Estonia by the Soviet army in 1940-45. After half a century of suppression the memories of these events resurfaced as different stories describing the long-term, often inter-generational strategies of coping with the value collapse. The second cultural trauma is revealed together with the collapse of the Soviet power and ideology in Estonia in 1991. According to empirical data, the following three trauma discourses have been reconstructed: - the forced adaptation to Soviet order of the homeland Estonians; - the difficulty of preserving Estonian identity in exile (Sweden); - the identity crisis of the Russian population of Estonia. Comparative analyses of these discourses have shown that opposing experiences and worldviews cause conflicting interpretations of the past. Different social and ethnic groups consider coping with cultural trauma as a matter of self-defence and create appropriate usable pasts to identify with. Keywords: human time, cultural trauma, frame analysis, discourse, life stories

Environmental change and human history in the Jabal Harun area, Jordan

The research is related to the Finnish Jabal Harun Project (FJHP), which is part of the research unit directed by Professor Jaakko Frösén. The project consists of two interrelated parts: the excavation of a Byzantine monastery/pilgrimage centre on Jabal Harun, and a multiperiod archaeological survey of the surrounding landscape. It is generally held that the Near Eastern landscape has been modified by millennia of human habitation and activity. Past climatic changes and human activities could be expected to have significantly changed also the landscape of the Jabal Harun area. Therefore it was considered that a study of erosion in the Jabal Harun area could shed light on the environmental and human history of the area. It was hoped that it would be possible to connect the results of the sedimentological studies either to wider climatic changes in the Near East, or to archaeologically observable periods of human activity and land use. As evidence of some archaeological periods is completely missing from the Jabal Harun area, it was also of interest whether catastrophic erosion or unfavourable environmental change, caused either by natural forces or by human agency, could explain the gaps in the archaeological record. Changes in climate and/or land-use were expected to be reflected in the sedimentary record. The field research, carried out as part of the FJHP survey fieldwork, included the mapping of wadi terraces and cleaning of sediment profiles which were recorded and sampled for laboratory analyses of facies and lithology. To obtain a chronology for the sedimentation and erosion phases also OSL (optically stimulated luminescence) dating samples were collected. The results were compared to the record of the Near Eastern palaeoclimate, and to data from geoarchaeological studies in central and southern Jordan. The picture of the environmental development was then compared to the human history in the area, based on archaeological evidence from the FJHP survey and the published archaeological research in the Petra region, and the question of the relationship between human activity and environmental change was critically discussed. Using the palaeoclimatic data and the results from geoarchaeological studies it was possible to outline the environmental development in the Jabal Harun area from the Pleistocene to the present.It is appears that there was a phase of accumulation of sediment before the Middle Palaeolithic period, possibly related to tectonic movement. This phase was later followed by erosion, tentatively suggested to have taken place during the Upper Palaeolithic. A period of wadi aggradation probably occurred during the Late Glacial and continued until the end of the Pleistocene, followed by significant channel degradation, attributed to increased rainfall during the Early Holocene. It seems that during the later Holocene channel incision has been dominant in the Jabal Harûn area although there have been also small-scale channel aggradation phases, two of which were OSL-dated to around 4000-3000 BP and 2400-2000 BP. As there is no evidence of tectonic movements in the Jabal Harun area after the early Pleistocene, it is suggested that climate change and human activity have been the major causes of environmental change in the area. At a brief glance it seems that many of the changes in the settlement and land use in the Jabal Harun area can be explained by climatic and environmental conditions. However, the responses of human societies to environmental change are dependent on many factors. Therefore an evaluation of the significance of environmental, cultural, socio-economic and political factors is needed to decide whether certain phenomena are environmentally induced. Comparison with the wider Petra region is also needed to judge whether the phenomena are characteristic of the Jabal Harun area only, or can they be connected to social, political and economic development over a wider area.

Mutations as molecular tools : The metabolic-rate dependent molecular clock and DNA barcoding of allied species

Mutation and recombination are the fundamental processes leading to genetic variation in natural populations. This variation forms the raw material for evolution through natural selection and drift. Therefore, studying mutation rates may reveal information about evolutionary histories as well as phylogenetic interrelationships of organisms. In this thesis two molecular tools, DNA barcoding and the molecular clock were examined. In the first part, the efficiency of mutations to delineate closely related species was tested and the implications for conservation practices were assessed. The second part investigated the proposition that a constant mutation rate exists within invertebrates, in form of a metabolic-rate dependent molecular clock, which can be applied to accurately date speciation events. DNA barcoding aspires to be an efficient technique to not only distinguish between species but also reveal population-level variation solely relying on mutations found on a short stretch of a single gene. In this thesis barcoding was applied to discriminate between Hylochares populations from Russian Karelia and new Hylochares findings from the greater Helsinki region in Finland. Although barcoding failed to delineate the two reproductively isolated groups, their distinct morphological features and differing life-history traits led to their classification as two closely related, although separate species. The lack of genetic differentiation appears to be due to a recent divergence event not yet reflected in the beetles molecular make-up. Thus, the Russian Hylochares was described as a new species. The Finnish species, previously considered as locally extinct, was recognized as endangered. Even if, due to their identical genetic make-up, the populations had been regarded as conspecific, conservation strategies based on prior knowledge from Russia would not have guaranteed the survival of the Finnish beetle. Therefore, new conservation actions based on detailed studies of the biology and life-history of the Finnish Hylochares were conducted to protect this endemic rarity in Finland. The idea behind the strict molecular clock is that mutation rates are constant over evolutionary time and may thus be used to infer species divergence dates. However, one of the most recent theories argues that a strict clock does not tick per unit of time but that it has a constant substitution rate per unit of mass-specific metabolic energy. Therefore, according to this hypothesis, molecular clocks have to be recalibrated taking body size and temperature into account. This thesis tested the temperature effect on mutation rates in equally sized invertebrates. For the first dataset (family Eucnemidae, Coleoptera) the phylogenetic interrelationships and evolutionary history of the genus Arrhipis had to be inferred before the influence of temperature on substitution rates could be studied. Further, a second, larger invertebrate dataset (family Syrphidae, Diptera) was employed. Several methodological approaches, a number of genes and multiple molecular clock models revealed that there was no consistent relationship between temperature and mutation rate for the taxa under study. Thus, the body size effect, observed in vertebrates but controversial for invertebrates, rather than temperature may be the underlying driving force behind the metabolic-rate dependent molecular clock. Therefore, the metabolic-rate dependent molecular clock does not hold for the here studied invertebrate groups. This thesis emphasizes that molecular techniques relying on mutation rates have to be applied with caution. Whereas they may work satisfactorily under certain conditions for specific taxa, they may fail for others. The molecular clock as well as DNA barcoding should incorporate all the information and data available to obtain comprehensive estimations of the existing biodiversity and its evolutionary history.

Women and Marital Breakdown in South India : Reconstructing Homes, Bonds and Persons

Women and Marital Breakdown in South India: Reconstructing Homes, Bonds and Persons is an ethnographic analysis of the situation of divorced and separated women and their families in the South Indian city of Bangalore. The study is based on 16 months of anthropological fieldwork, i.e., participant observation and life history interviews among 50 divorced and separated women from different socio-religious backgrounds in their homes, in the women s organisations and in the Family Court. The study follows the divorced and separated women from their natal homes to their affinal homes through homelessness and legal battles to their reconstructed natal, affinal or single homes in order to find out what it means to be a person within hierarchical gender and kinship relations in South India. Marital breakdown impacts on kin relations and discloses the existing gender relations and power structure through its consequences. It makes the transformability of relational personhood as well as the transformability of relational society and culture visible. Although the study reveals the painful history of women s ill-treatment in marriage, family and kinship systems, it also demonstrates the women s rejection of the domination; and shows their ability to re-negotiate and promote changes not only to their own positions but to the whole hierarchical system as well. The study explores the divorced and separated women s manifold dilemmas, complicated legal battles, and endless arrangements when they have to struggle with the very practical problems of supporting themselves financially, finding and making a new home for themselves, and re-arranging relationships with their kin and friends. As marital breakdown fundamentally transforms the women s relational field, it forces them to recreate substitutive relations in a flexible way and, simultaneously, to re-construct themselves and their lives without a ready or positive cultural or behavioural template. This process reveals the agency of the divorced and separated women as well as shedding light on issues of gender and the cultural construction of the person in South India. This topical study explores the previously neglected subject of marital breakdown in India and shows the new meaning of kinship in South India.

Host-parasitoid relationship in different Cotesia melitaearum and Melitaea cinxia populations around the Baltic Sea

The relationship between hosts and parasites is one of the most studied interactions between living organisms, and it is both universal and common in nature. Parasitoids are special type of parasites whose offspring develop attached to or within a single host organism that it ultimately consumes and kills. Hosts are arthropods and most parasitoids belong to the insect order Hymenoptera. For almost two decades metapopulation research on the Glanville fritillary butterfly (Melitaea cinxia) has been conducted in the Åland Islands, Finland. The studies have been concerned with the population dynamics, evolution, genetics, behavior, natural history and life history characteristics of M. cinxia, as well as with species interacting with the butterfly. The parasitoids of M. cinxia have been under long term studies and much has been learned about specific host-parasitoid interactions during the past decade. The research for this Master s thesis was done in the Åland Islands during summer 2010. I conducted a reciprocal transplant style experiment in order to compare the performance of host butterflies (M. cinxia) under attack by different parasitoid wasps (C. melitaearum). I used hosts and parasitoids from five origins around the Baltic Sea: Öland, Uppland, Åland, Saaremaa and Pikku-Tytärsaari. The host-parasitoid relationship was studied in terms of host susceptibility and parasitoid virulence, addressing specifically the possible effects of inbreeding and local adaptation of both parasitoids and their hosts. I compared various factors such as host defence ratio, parasitoid development rate, cocoon production rate etc. I also conducted a small scale C. melitaearum egg development experiment and C. melitaearum external morphology comparison between different parasitoid populations. The results show that host resistance and parasitoid virulence differ between both host and parasitoid populations. For example, Öland hosts were most susceptible to parasitoids and especially vulnerable to Pikku-Tytärsaari wasps. Pikku-Tytärsaari wasps were most successful in terms of parasitoids virulence and efficiency except in Saaremaa hosts, where the wasp did not succeed. Saaremaa hosts were resistant except towards Åland parasitoids. I did not find any simple pattern concerning host resistance and parasitoid virulence between inbred and outbred populations. Also, the effect of local adaptation was not detected, perhaps because metapopulation processes disturb local adaptation of the studied populations. Morphological comparisons showed differences between studied wasp populations and sexual dimorphism was obvious with females being bigger that males. There were also interesting differences among populations in male and female wing shapes. The results raise many further questions. Especially interesting were Pikku-Tytärsaari wasps that did well in terms of efficiency and virulence. Pikku-Tytärsaari is a small, isolated island in the Gulf of Finland and both the host and parasitoids are extremely inbred. For the host and parasitoid to persist in the island, the host has to have some mechanisms to escape the parasitoid. Further research will be done on the subject to discover the mechanisms of Pikku-Tytärsaari host s ability to escape parasitism. Also, genetic analyses will be conducted in the near future to determine the relatedness of used C. melitaearum populations.

Phylogeography and demographic history of Lacerta lepida in the Iberian Peninsula

The Iberian Peninsula is recognized as an important refugial area for species survival and diversification during the climatic cycles of the Quaternary. Recent phylogeographic studies have revealed Iberia as a complex of multiple refugia. However, most of these studies have focused either on species with narrow distributions within the region or species groups that, although widely distributed, generally have a genetic structure that relates to pre-Quaternary cladogenetic events. In this study we undertake a detailed phylogeographic analysis of the lizard species, Lacerta lepida, whose distribution encompasses the entire Iberian Peninsula. We attempt to identify refugial areas, recolonization routes, zones of secondary contact and date demographic events within this species. Results support the existence of 6 evolutionary lineages (phylogroups) with a strong association between genetic variation and geography, suggesting a history of allopatric divergence in different refugia. Diversification within phylogroups is concordant with the onset of the Pleistocene climatic oscillations. The southern regions of several phylogroups show a high incidence of ancestral alleles in contrast with high incidence of recently derived alleles in northern regions. All phylogroups show signs of recent demographic and spatial expansions. We have further identified several zones of secondary contact, with divergent mitochondrial haplotypes occurring in narrow zones of sympatry. The concordant patterns of spatial and demographic expansions detected within phylogroups, together with the high incidence of ancestral haplotypes in southern regions of several phylogroups, suggests a pattern of contraction of populations into southern refugia during adverse climatic conditions from which subsequent northern expansions occurred. This study supports the emergent pattern of multiple refugia within Iberia but adds to it by identifying a pattern of refugia coincident with the southern distribution limits of individual evolutionary lineages. These areas are important in terms of long-term species persistence and therefore important areas for conservation.