This thesis deals with the question of whether planktonic protits of the genus Dinophysis have permanent plastids (=chloroplasts) or practice kleptoplasty, i.e. acquire plastids via predation on other microorganisms. Sequencing the plastid 16S rDNA of Dinophysis spp. collected from 4 different geographical regions unveiled two different plastid genotypes within this genera: one that was found at all locations investigated, identical to that of the free-living cryptophyte Teleaulax amphioxeia, and another found only in the Greenland Sea, closely related to that of the cryptophyte Geminigera cryophila. Both types were found within the species D. acuminata. These findings imply that the plastids in Dinophysis spp. were not inherited from a common ancestor, but acquired from feeding. By using flow cytometry in combination with an acidotrophic probe, it was shown that 71 % of the cells in a D. norvegica population in the aphotic zone of the Baltic Sea had food-vacuoles. Dinophysis used to be regarded as a primarily phototrophic organism, and this was a higher proportion of cells with food-vacuoles than reported earlier. To further study if Dinophysis needs constant refill of new plastids from the environment, a new method combining flow-cytometry and quantitative real-time PCR was developed to compare the levels of nuclear and plastid DNA in different phases of the cell-cycle. Results showed that plastid acquisition in Dinophysis was uncoupled with the cell-cycle, which is different than the pattern seen in microalgal species with permanent plastids. Furthermore, when quantitative real-time PCR combined with flow-cytometry was used to follow D. caudata cultures during a 65 days starvation/feeding experiment, the cells first went through a steady decrease in plastid DNA during starvation. In contrast, after feeding on the ciliate Myrionecta rubra, plastid DNA in starved cells increased 7-fold, thereby directly revealing the kleptoplastic behavior. The main conclusion from this thesis is that Dinophysis cells are actively taking up kleptoplastids from the ciliates on which they feed, and that kleptoplasty is an important key to understand Dinophysis ecology. Part of this thesis work has also been dedicated to the application and optimization of new methods, and it shows how quantitative real-time PCR, flow cytometry and molecular methods in different combinations can be used as powerful tools for the study of plankton ecology.

Coral reefs belong to the most diverse and the most threatened ecosystems on earth. Anthropogenic stressors and climate change have led to mortalities at levels unprecedented in modern times. The aims of this thesis are to investigate aspects of the corals’ ability to reproduce, disperse, adapt and survive. Papers I-III study reproduction in a common soft coral species, Sarcophyton elegans, with previously unknown reproductive modes. Paper IV investigates genetic distribution of coral-symbiont associations in Galaxea fascicularis focusing on adaptation to the environment along the coastline of Vietnam. Sarcophyton  elegans is a gonochoric broadcast spawner with a 1:1 sex ratio. Reproduction is strictly size dependent. Oogenesis takes 19-24 months, with a new cycle commencing every year. Spermatogenesis takes 10-12 months. The majority of gametes were released during the annual austral mass spawning event after full moon in November, but spawning also occur between August and February. The polyps at the outer edge of the colonies released their gametes first, followed by polyps situated closer to the center during subsequent months. Colonies upstream in the prevailing current spawn earlier than those downstream. The colonies were arranged in clusters of alternating males and females, which spawned simultaneously and were of the same genotype. Fission and buddying is a common mode to expand locally. Additionally, females undergoing fission divided into the most fecund size classes. The G. fascicularis and their associated symbionts were not genetically coupled to each other but to environmental factors. The host displayed an inshore-offshore zonation, with higher diversity offshore. The D1a symbiont exhibited an inshore- offshore zonation. In contrast; the 5 different C symbiont types showed a latitudinal distribution gradient, which shifted in dominance north to south. The study highlights the importance of protecting resilient coral and algal genotypes in stressed areas and the need to understand reproductive modes for coral conservation. / Các rạn san hô là một trong những hệ sinh thái có tính đa dạng và bị đe dọa cao nhất trên trái đất. Các áp lực từ con người và nhiệt độ nước biển tăng (SSTs) đã gây ra hiện tượng “tẩy trắng” gây chết san hô ở mức độ cao chưa từng thấy trong thời điểm hiện tại. Mục tiêu của nghiên cứu này là tìm hiểu khả năng của san hô trong thích nghi, phân tán và sống sót nhằm duy trì quần thể. Bài báo số II-III là những nghiên cứu đầu tiên về đặc điểm sinh sản của loài san hô mềm phổ biến, Sarcophyton elegan tại Australia. Bài báo số IV nghiên cứu về phân bố nguồn gen của tảo cộng sinh trong loài san hô Galaxea fascicularis, tập trung vào sự thích nghi với môi trường dọc theo vùng biển Việt Nam, khu vực bị ô nhiễm từ lục địa. Sarcophyton elegans được biết với đặc điểm sinh sản cả vô tính và hữu tính. Loài này là loài sinh sản bằng cách phân tán trứng, với tỷ lệ giới tính là 1:1 và sự sinh sản hữu tính bị khống chế nghiêm ngặt bởi kích cỡ của tập đoàn (Bài báo II, phần phương pháp của Bài báo I). Quá trình tạo trứng kéo dài từ 19 đến 24 tháng với chu kỳ sinh sản lặp lại hàng năm, và sự sinh tinh kéo dài từ 10 đến 12 tháng. Phần lớn giao tử được giải phóng trong một thời gian ngắn sau ngày trăng tròn của tháng 11, nhưng giao tử vẫn được giải phóng trong ngày trăng tròn của các tháng từ tháng 8 đến tháng 1 năm sau. Các polyp autozooid nằm phía ngoài của tập đoàn giải phóng giao tử trước, sau đó là các polyp nằm gần lõi trong các tháng tiếp theo. Các tập đoàn ngược lên trong dòng chảy thịnh hành đẻ trứng sớm hơn các tập đoàn xuôi dòng khoảng một tháng (Bài báo II). Các tập đoàn được sắp xếp thành từng đám từ 7 đến hàng trăm tập đoàn trong mỗi nhóm, bao gồm cả đực và cái. Các tập đoàn trong cùng một nhóm sinh sản cùng một thời điểm. (Bài báo II) và mỗi nhóm có cùng một kiểu di truyền (Bài báo III) có đầy đủ 13 (có thể là 22) kiểu di truyền  khác nhau. Sự phân đôi và kết đôi phụ thuộc hoàn toàn vào kích thước và có lẽ là phương thức mở rộng phổ biến nhất. Sự phân đôi phải mất 2 năm hoặc hơn mới hoàn thành. Thêm vào đó, con cái trải qua quá trình phân đôi thành kích cỡ có khả năng sinh sản cao nhất (Bài báo III). Có 6 nhóm haplotypes (mtDNA) của loài G. fascicularis và tảo cộng sinh Symbiodinium (ITS2 rDNA) không đóng cặp với nhau nhưng lại gắn với các yếu tố môi trường, có thể như kết quả của phương thức sinh sản của vật chủ (Bài báo IV). Vật chủ có sự phân vùng rõ rệt giữa gần bờ và xa bờ, với sự đa dạng cao hơn hẳn của các rạn xa bờ so với các rạn gần bờ, khu vực thường xuyên bị độ đục, ô nhiễm và lắng đọng trầm tích tác động. Tảo cộng sinh Symbiodinium D1a ITS2 điểm hình của sự phân vùng gần bờ và xa bờ. Ngược lại, 5 loại C khác lại có sự phân vùng theo vĩ tuyến, với sự tăng lên rõ rệt theo chiều Bắc-Nam, cùng với sự ổn định SST và sự tăng lên của các SST. Nghiên cứu này đã chỉ rõ tầm quan trọng trong bảo vệ các loài san hô và tảo biển bản địa tại các khu vực bị đe dọa (Bài báo IV) và sự cần thiết phải hiểu các phương thức sinh sản (Bài báo II-III) và các thông số môi trường trong việc xác định mức độ đa dạn sinh học và sự hấp thụ của sinh vật cộng sinh trong san hô cứng và san hô mềm. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: Manuscript.

The colour choice of spiders and the effects of competition on this phenomenon were examined in the laboratory using the sit-and-wait predator goldenrod crab spider (Misumena vatia, (Thomisidae)). This species can  be both white and yellow (could also change colour)  and it was here examined whether individuals preferred to sit on flowers which has the same colouration as their bodies. I found that M.vatia showed preference for flowers that matched their own body colour, but when competition was introduced this preference disappear. It seems, however, that position is a factor that can influences the substrate decisions of M. vatia, where the edge of the flower is preferred.

This research aims to investigate the effects that dredging may have on benthic fauna. The dredging took place in the stream Ireån on northern Gotland during the early summer 2010. My area of research was taking place in a scenic area downstream in Ireån close to the Baltic Sea. I chose this area due to the dredging that started just upstream of this area and continued further upstream. Thus it is not the direct impact of the dredging that is the focus for this paper but the side-effect the dredging may have on the area downstream. In this research I used the M42-method and BQI- index in order to evaluate the results. The effects on the dredging were weaker than expected, with only minor changes occurring within the benthic fauna (BQI-value before dredging: 3.35, BQI-value after dredging: 3.51).

<p>Biological invasions are an important issue of global change and an increased understanding of invasion processes is of crucial importance for both conservation managers and international trade. In this thesis, I have studied the invasion of the brown seaweed <i>Fucus evanescens</i>, to investigate the fate and effect of a perennial, habitat-forming seaweed introduced to a coastal ecosystem. A long-term study of the spread of <i>F. evanescens</i> in Öresund (southern Sweden) showed that the species was able to expand its range quickly during the first 20 years after the introduction, but that the expansion has been slow during the subsequent 30 years. Both in Öresund and in Skagerrak, the species is largely restricted to sites where native fucoids are scarce. Laboratory experiments showed that the restricted spread of <i>F. evanescens</i> cannot be explained by the investigated abiotic factors (wave exposure and salinity), although salinity restricts the species from spreading into the Baltic Sea. Neither did I find evidence for that herbivores or epibiota provide biotic resistance to the invader. On the contrary, <i>F. evanescens</i> was less consumed by native herbivores, both compared to the native fucoids and to <i>F. evanescens</i> populations in its native range, and little overgrown by epiphytes. Instead, the restricted spread may be due to competition from native seaweeds, probably by pre-occupation of space, and the establishment has probably been facilitated by disturbance. </p><p>The studies provided little support for a general enemy release in introduced seaweeds. The low herbivore consumption of <i>F. evanescens</i> in Sweden could not be explained by release from specialist herbivores. Instead, high levels of chemical anti-herbivore defence metabolites (phlorotannins) could explain the pattern of herbivore preference for different fucoids. Likewise, the low epibiotic colonisation of <i>F. evanescens </i>plants could be explained by high resistance to epibiotic survival. This shows that colonisation of invading seaweeds by native herbivores and epibionts depends on properties of the invading species. The large differences between fucoid species in their quality as food and habitat for epibionts and herbivores imply that invasions of such habitat-forming species may have a considerable effect on a number of other species in shallow coastal areas. However, since <i>F. evanescens</i> did not exclude other fucoids in its new range, its effect on the recipient biota is probably small.</p>

<p>Sexual selection arises through variation in reproductive success. This thesis investigates different aspects important in sexual selection, namely nest building, sperm competition, paternity and paternal care, and their mutual interrelationships. In the studied species, the sand goby (<i>Pomatoschistus minutus</i>) and the common goby (<i>Pomatoschistus microps</i>), sperm competition did arise when small males, so called sneakers, sneaked into other males nests and released sperm. They seemed to use female behaviour as their prime cue for a sneaking opportunity. However, also nest-holders, both with and without eggs, were found to fertilize eggs in the nests of other males.</p><p>Clearly, nest-holding males tried to prevent other males from spreading their sperm in their nests, since they showed aggression towards such males. A nest building experiment indicated that the small nest-openings found in the sneaker male treatment were sexually selected through protection against sneaking or by female choice. Yet, no behavioural or genetical support for the hypothesis that the nest functions as a physical or visual defence, or that sneaker males prefer to sneak upon nests with wide nest-openings, were found in the other studies. Still, individual nest-holding males showed a higher mucus preparation effort inside the nest in the presence of a sneaker male than when alone. In close relatives, such mucus contains sperm, suggesting an importance in sperm competition. However, the mucus may also have pheromone and anti-bacterial functions and may constitute a mating effort, as found in other gobies. Both a behavioural and a mate choice experiment suggested that the males were not less eager to spawn in the presence of a sneaker male.</p><p>Sneak intrusion did not affect nest defence, fanning or filial cannibalism, nor had paternity an effect on filial cannibalism. This and various life history aspects, together with the fact that the parasitic male only fertilized a fraction of the clutches, would predict females to ignore sneaker males. This was also the case, as the presence of sneaker males was found not to affect female spawning decision. Still, several females spawned in two nests, which coincided with parasitic spawnings, suggesting a cost of disturbance for the females and thus a substantial cost to the nest-holding males in terms of lost mating success. However, females paid attention to other traits in their choice of mate since spawning was associated with sand volume of the nest, but not with nest-opening width. Also, female (but not male) courtship was correlated with partial clutch filial cannibalism, indicating that females are able to anticipate future male cannibalism.</p><p>In a partial correlation of nest opening, sand volume, male courtship display, displacement fanning and male size, a large number of traits were correlated both positively and negatively with regard to how we may expect them to be appreciated by females. For instance, males which fan well also build large nests or display intensely (but not both). Together with all the other results of this thesis, this shows the entangled selection pressures working on breeding animals, as well as the different male and female tactics employed to maximize their reproduction.</p>

<p>An influence at one trophic level can result in dynamic impacts also on other components of a food web. These dynamics are known as trophic cascades, and can be both top-down and bottom-up. After a near-collapse of the Baltic cod <i>Gadus morhua</i> stock in the 1980s, its main prey sprat <i>Sprattus sprattus</i> increased dramatically. The main food of sprat, marine copepods, decreased during the same time period, likely a combined effect of increased predation pressure from sprat and decreasing salinities. This shortage of food for sprat resulted in decreasing quality of sprat as a food source for common guillemots <i>Uria aalge</i>. However, a recent increase in fishing for sprat has again resulted in better feeding conditions for guillemots.</p><p>Human impacts on this simple food web can be complex. In the early 20th century, marine mammals were abundant and nutrient levels were low in the Baltic Sea. This thesis illustrate that this situation corresponded to lower fish biomass. A reduction of seals early in the century led to reduced top-down control, which resulted in increasing fish stocks. Later, in the 1950s, the largest inflow of salt water during the century mobilized accumulated phosphorus from the deep sediments, which stimulated nitrogen fixation. Combined with increasing anthropogenic nutrient loads, this led to increased primary production and a rapid change from an oligotrophic to a eutrophicated state. This change can be termed a regime shift, which also stimulated fish production. Subsequent over-fishing of cod likely caused a second regime shift, from a cod- to a clupeid- dominated state, which led to the described effects on the common guillemots.</p><p>Several factors affect the life-history of this long-lived seabird. Bycatches in gillnet fisheries is one factor directly affecting guillemot survival, and the proportion of bycatches increased during a period of increasing fishing effort. Surprisingly, avian cholera, a previously undocumented disease in common guillemots, was found at times to cause considerable adult mortality. Common guillemot life-history information can communicate the diversity of factors influencing marine ecosystems – hopefully this can increase our understanding of how complex even "simple" food webs are.</p>

<p>Lichens are fascinating symbiotic systems, where a fungus and a unicellular alga, most often green (bipartite green algal lichens; 90% of all lichens), or a fi lamentous cyanobacterium (bipartite cyanobacterial lichens; 10% of all lichens) form a new entity (a thallus) appearing as a new and integrated organism: in about 500 lichens the fungus is associated with both a cyanobacterium and an alga (tripartite lichens). In the thallus, the lichen bionts function both as individual organisms, and as a symbiont partner. Hence, in lichens, the participating partners must both be able to receive and acquire resources from the other partner(s) in a controlled way.</p><p>Lichens are particularly successful in harsh terrestrial environments. In part this is related to their poikilohydric nature and subsequent ability to repeatedly become desiccated and hydrated. Metabolic activity, i.e. photosynthesis, respiration, and for cyanobacterial lichens N2-fixation, is limited to periods when the thallus is suffi ciently hydrated. Mineral nutrients are mainly acquired from dry or wet deposition directly on the thallus. Taken together it then appears that lichens are to a large extent passively controlled by their environment, making their control over resource allocation and acquisition particularly challenging.</p><p>The aim of this thesis was to investigate resource acquisition and allocation processes in different lichens, and to see how these respond to changes in resource availability. This was done by following lichen growth in the fi eld during manipulation of water, light, and nutrient supply, and by assessing the responses of both the integrated thallus as well as the individual bionts. As a fi rst step, resource allocation and acquisition was investigated for a broad range of lichens aiming to determine the magnitude of metabolic variation across lichens. Seventy-fi ve lichen species were selected to cover as broad a spectrum as possible regarding taxonomy, morphology, habitat, and nitrogen requirements. The lichens had invested their nitrogen resources so that photosynthetic capacity matched respiratory carbon demand around a similar equilibrium across the contrasting species. Regulation of lichen growth was investigated in another study, using the two tripartite species <i>Nephroma arcticum</i> and <i>Peltigera aphthosa</i>, emphasizing the contribution of both internal and external factors. The empirical growth models for the two lichens were similar, showing that weight gain is to a higher extent dependent on those external factors that regulate their photosynthesis, whilst area gain is more controlled by internal factors, such as their nitrogen metabolism. This might be inferred from another study of the same species, where nitrogen manipulations resulted in an undisturbed weight gain, a similar resource allocation pattern between the bionts, but a distorted area gain. </p><p>Aiming to investigate lichen nitrogen relations even further, lichens’ capacities to assimilate combined nitrogen in the form of ammonium, nitrate and amino acids were assessed using 14 contrasting boreal species. All these had the capacity to assimilate all the three nitrogen forms, with ammonium absorption being more passive, and nitrate uptake being low in bipartite cyanobacterial lichens. Differences in uptake capacities between species were more correlated to photobiont than to morphology or substrate preferences. Finally, to investigate intra-specifi c plasticity in relation to altered nutrient supply, resource investments between photo- and mycobiont were investigated in the two bipartite green algal lichens <i>Hypogymnia physodes </i>and and <i>Platismatia glauca</i> in a low and a high nutrient environ- in a low and a high nutrient environ- ment. In both species, more of the resources had been directed to the photobiont in the high nutrient environment also increasing their overall carbon status. Taken together, my studies indicate that in spite of the apparent passive environmental control on lichen metabolism, these symbiotic organisms are able to both optimize and control their resource acquisition and allocation processes.</p>

<p>Multiple mating by females is common in nature. Yet, the evolution and maintenance of polyandry remains a bit of an evolutionary puzzle. It was my aim in this thesis to reach a greater understanding of this phenomenon as well as to investigate the consequences of polyandry on the evolution of reproductive divergence in insects. In an extensive meta analysis addressing the direct effects of multiple mating on female fitness in insects, I found that insects gain from multiple matings in terms of increased lifetime offspring production. In species without nuptial feeding, increased mating rate leads to decreased female lifespan and my results strongly support the existence of an intermediate optimal female mating rate. However, results from an experimental study where I examined the relationship between female fitness and mating rate in the bean weevil (Callosobruchus maculatus) showed that female fitness was maximized at two alternative mating rates, indicating that some species may exhibit a more complex relationship between the costs and benefits of mating. In the meta analysis on species with nuptial feeding, I found only positive effects of increased mating rate and the puzzle is rather what constrains the actual mating rates of females in these groups.</p><p>Sexual selection is a very potent driver of rapid evolutionary change in reproductive characters. Most research has focussed on precopulatory sexual selection, but in promiscuous species sexual selection continues after copulation and variance in male fertilization success gives rise to postcopulatory sexual selection. In this thesis I found that three allopatric populations of the red flour beetle (Tribolium castaneum) have diverged in traits related to reproduction. Male genotype affected all aspects of female reproduction, but more interestingly, males and females interacted in their effect on offspring production and reproductive rate, showing that the divergence was due to the evolution of both male and female reproductive traits.</p><p>When studying postcopulatory sexual selection, sperm competition has been put forward as the main source of variance in fertilization success. The results from a set of double-mating experiments, using the same populations of flour beetles, provided strong evidence that cryptic female choice is also important in generating variance in male fertilization success. I found not only main effects of female genotype on male fertilization success but also male-female interactions which provide more unambiguous evidence for cryptic female choice. Finally, I attempted to uncover which male signals-female receptors are involved in the reproductive divergence observed in the Tribolium populations. In a double-mating experiment I manipulated female perception of two male reproductive signals, copulatory courtship and cuticular hydrocarbons, and the results indicate that, within populations, both signals are sexually selected. However, only male cuticular hydrocarbons seem to be involved in the reproductive divergence between the populations. </p><p>In conclusion, multiple mating by female insects can be understood solely in terms of direct fitness benefits resulting from increased offspring production. I have shown that postcopulatory sexual selection can lead to rapid divergence in reproductive traits related to mating and that cryptic female choice plays an important role in this divergence.</p>

<p>Temporal and spatial variations of pelagic microorganisms in the northern Baltic Sea were studied, as well as factors influencing their abundance and growth rates. Three main questions were asked 1) How does increased productivity influence the structure of the microbial food web? 2) Does predation limitation vary between trophic levels? 3) What is the relative importance of resource and predation limitation at different trophic levels?</p><p>A field study in the northern Baltic Sea showed that dominating protozoa, flagellates and ciliates, increased with increasing primary productivity from north to south. Furthermore, relatively small protozoan cells dominated in the low productive north, while larger cells became more dominant in the south. The relationship between plankton size structure and productivity was further studied in an experimental system. In agreement with present theories regarding nutrient status of pelagic food webs, increased productivity caused a lengthening of the food chain as well as a change in plankton size structure. While microplankton dominated in nutrient rich treatments pico- and nanoplankton dominated during nutrient poor treament. The flagellate community was dominated by a potentially mixotroph, <i>Chrysochromulina</i> sp., at low nutrient concentrations. To our knowledge this is the first experimental study showing that <i>Chrysochromulina</i> sp. in resemblance with other mixotrophs is favoured by nutrient poor conditions compared to strict autotrophs and heterotrophs.</p><p>During a stratified summer period autotrophic microorganisms in the northern Baltic Sea did not respond to removal of potential predators, indicating that they were primarily limited by inorganic nutrients. An exception was small eucaryotic picoplankton that showed a large response to predator removal. Among the heterotrophic microorganisms direct effect of predation seemed to increase from ciliates, heterotrophic bacteria, small heterotrophic flagellates, medium flagellates to large flagellates. No quick indirect effect was observed, but after four days trophic cascades were detected.</p><p>The relative importance of resource and predation limitation was studied among heterotrophic bacteria, flagellates and ciliates in the northern Baltic Sea. For all these groups, resource limitation seemed to prevail during the summer period. The results also indicated that the relative importance of predation increased with the productivity of the system. To our knowledge there are no earlier measurements on the relative importance of resource and predation limitation for micoorganisms in the pelagic environment.</p>