During the last few decades, coral reefs have become a disappearing feature of tropical marine environments, and those reefs that do remain are severely threatened. It is understood that humans have greately altered the environment under which these ecosystems previously have thrived and evoloved. Overharvesting of fish stocks, global warming and pollution are some of the most prominent threats, acting on coral reefs at several spatial and temporal scales. Presently, it is common that coral reefs have been degraded into alternative ecosystem regimes, such as macroalgae-dominated or sea urchin-barren. Although these ecosystems could potentially return to coral dominance in a long-term perspective, when considdering current conditions, it seems likely that they will persist in their degraded states. Thus, recovery of coral reefs cannot be taken for granted on a human timescale. Multiple stressors and disturbances, which are increasingly characteristic of coral reef environments today, are believed to act synergistically and produce ecological surprises. However, current knowledge of effects of compounded disturbances and stress is limited. Based on five papers, this thesis investigates the sublethal response of multiple stressors on coral physiology, as well as the effects of compounded stress and disturbance on coral reef structure and function. Adaptive responses to stress and disturbance in relation to prior experience are highlighted. The thesis further explores how inherent characteristics (traits) of corals and macroalgae may influence regime expression when faced with altered disturbance regimes, in particular overfishing, eutrophication, elevated temperature, and enhanced substrate availability. Finally, possibilities of affecting the resilience of macroalgae-dominaed reefs and shifting the community composition towards a coral-dominated regime are explored.

Chemical signals are the most widely used form of sexual communication throughout the living world. However, there is in general little knowledge about what these signals actually communicate. The role of chemical signals, i.e. pheromones, in sexual behaviour has traditionally been seen as restricted to mate attraction and species recognition. This thesis reviews the evidence for pheromones as indicators of mate quality, and then investigates three important factors for mate quality signals – individual variation, heritability and cost – by using the male pheromone of the lekking fruit fly Drosophila grimshawi as a model. The experiments presented indicate that the pheromone of D. grimshawi has multiple functions, and that these functions vary with social context. Thus, with regard to females, the pheromone seems to act mainly as a species/mate recognition signal, since females show little preference for the amount of pheromone deposited by a male. Moreover, males invest less in pheromone production when subjected to females as compared to when subjected to rival males. However, the pheromone seems to be costly in production since males that invest much in pheromone deposition has a shorter lifespan. This suggests a function for the pheromone in male-male interactions. Males can distinguish their own pheromone depositions from those of a strange male, and also discriminate between pheromone depositions from one and two strange males. This might give them the ability to assess the size of a lek and the competitive capacities of rivals, information that should be useful when optimizing sexual behaviour. In conclusion, the pheromone seems to act as an honest mate/competitor quality signal in some social contexts, and as a non-costly species/mate recognition signal in other. In addition, I show for the first time that a chemical signal has differential fitness costs, and that an insect is able to distinguish between individual odour signatures.

For many years, the importance of microorganisms attached to surfaces in littoral zones and wetlands has been disregarded when describing aquatic ecosystem dynamics. Supporting evidence is scarce but convincing that these microbial communities are not only very productive but can often serve as major regulators of nutrient and carbon dynamics in many freshwaters. In order to determine the quantitative importance of epiphytic bacteria for the overall carbon turnover, I compared the relative contribution of epiphytic bacteria on the submerged macrophyte Ranunculus circinatus, sediment and free-living bacteria to the total bacterial production. Sediment bacteria generally dominated total bacterial biomass in the littoral zone. Although the epiphytic biomass on R. circinatus was ten times lower than the biomass of sediment bacteria, it often contributed at least equally to the total bacterial production. Thus, the results presented in this thesis confirm that most bacterial biomass and production in shallow lakes is associated with surfaces, and that in littoral zones with dense macrophyte stands, epiphytic bacteria can contribute significantly to the overall carbon turnover. There is increasing evidence that not all cells in natural bacterial communities are metabolically active. In Lake Erken, there were large differences in the fraction of active bacteria between different habitats, while the within-habitat differences were small. The sediments had the largest bacterial fraction, followed by epiphytic bacteria, while in the water column only a few percent of the bacteria were active. In this thesis the fraction of active bacteria is connected to environmental fluctuations. I hypothesize that smaller fluctuations in chemical, biological or physical factors result in large active bacterial fractions. Thus, small environmental fluctuations within a habitat allow large active bacterial fractions, while the active fraction is constrained when the environmental fluctuations are large.

Wetlands contain both aquatic and terrestrial environments which generates high biodiversity. However, they are commonly associated with mosquitoes (Diptera: Culicidae), and mosquitoes are usually regarded as negative by humans because they can cause nuisance and transmit diseases. This thesis aimed to clarify the association between mosquitoes and wetlands and to achieve a more balanced view of biodiversity in wetlands by including mosquito diversity. Studies on adult mosquito diversity and assemblages were performed in 18 wetlands spread over Sweden. The Swedish mosquito species were organized in ten functional groups based on four life-history characteristics. This classification was used as an additional diversity measurement and as a tool for presentation of mosquito data. Mosquito diversity showed several of the well-established diversity patterns such as a latitudinal gradient, a species-area relationship and a distribution-abundance relationship. In a landscape perspective, diversity of both mosquitoes and dytiscids (Coleoptera: Dytiscidae) were positively influenced by a high proportion of permanent water and a high amount of open areas, indicating co-varying diversity patterns. Mosquito assemblages in the Nedre Dalälven region were mainly structured by the extent of flooded areas and wetland type (wet meadow, swamp and bog). In addition to the influence of the proportion of temporary wetlands at a local scale, the proportion of forest gained importance at larger spatial scales and in relation to dispersal distances of species. In southern Sweden, mosquito faunas differed between natural and constructed wetlands, partly reflecting differences in wetland size. In an experiment, different responses of two co-occurring mosquito species to rapid larval habitat desiccation indicate that weather conditions after a flood could influence mosquito assemblages. The conclusions of this thesis provide suggestions on how to construct and position wetlands for increased insect diversity, and indicate that low abundance of major nuisance species might be crucial for acceptance of wetlands near human settlements.

In the scope of future population support, agricultural productivity, in particular in sub-Saharan Africa, has to increase drastically to meet the UN’s millennium development goals of eradicating extreme poverty and hunger by 2015. Water availability in the root-zone limits crop production in large parts of the developing world. As competition for fresh water increases, water of lower quality, for example saline or polluted water, is often used for irrigation. Low-cost drip systems are suitable for saline water irrigation because they effectuate a minimisation of salt accumulation, leaf burn and peaks in salt concentration. Nonetheless, all types of saline water irrigation contain the risk for causing soil salinisation. Thus, in order to achieve long-term sustainability of these systems, appropriate management strategies are needed. The choice of management practices may be influenced by local conditions such as climate, soil and irrigation water salinity. A litera-ture review showed that there is a potential for saline water irrigation in sub-Saharan Africa in water scarce areas. Low-cost drip irrigation with saline water (6 dS m-1) was successfully used to irrigate two consecutive crops of tomato in semi-arid South Africa. An integrated ecosystems model was developed to simulate long-term yield and salt accumulation in a drip-irrigated agricultural system for a range of salinities, climates and management techniques. Crop, salt and water balance data from two field experiments conducted in Israel and South Africa, respectively, were used to parameterise and test the model. Emphasis was placed on testing the usability of the model as a tool for evaluating the importance of certain plausible management options of low-cost, drip-irrigation systems. Therefore, particular focus was directed towards correctly describing soil salinity stress on plant growth and soil evaporation from a distributed (wetted and dry) surface. In addition, the model was developed to function for different climates without having to change any other parameters or variables except for the actual climatic data. Simulations were subsequently run over a 30-year period to study long-term yield and salt accumulation in the soil profile for two sites in South Africa, demonstrating the applicability of the model. Model simulations showed that high soil salinities reduced crop growth and thus increased both drainage and soil evaporation. Further, covering the soil with a plastic sheet led to a reduction of soil evaporation and a subsequent increase in both transpiration and drainage. Rainfall was crucial for the leaching of salts from the soil, and thus in regions with low levels of rainfall, a higher leaching fraction of supplied saline irrigation water has to compensate for the lack of rain. However, a high leaching fraction also causes large amounts of salt leaching, which could potentially pollute underlying groundwater and downstream ecosystems. This risk can be mitigated using mulching, which minimises non-productive water losses, thereby lowering irrigation water needs. The choice of irrigation water salinity, frequency of irrigation and soil coverage may differ between the farmer and the regional water manager due to different preferences. Furthermore, the study highlighted how environmental variables such as water use efficiency and radiation use efficiency can be used as indicators of system performance. Whereas the latter is first and foremost a general stress indicator, water use efficiency more precisely describes specific factors such as plant size, allocation patterns and evaporative demand, which will affect the exchange of carbon dioxide and water through the stomata. / QC 20101102

Most urban areas of developing countries were hitherto experiencing unprecedented growth in their population, the phenomenon commonly referred to as urbanization, which in this study can be said to be the proportion of urban population relative to the total population of a region. This phenomenon has opportunities vis-à-vis challenges, whose impact on natural resources in general and water resources in particular, cannot be over- emphasized. It is within the context of these urban challenges that this study attempts to look into and possibly assess the situation in the urban and peri- urban areas of Kumasi Metropolitan Area (KMA) in Ghana, whose urbanization processes is prejudiced by this study to be increasing rapidly. The study specifically attempted to assess thechanges in the urbanization patterns and the possible future urbanization trend of the area within a specific time frame. The study also attempts to look at the causes of this urbanization, and its impact on water resources in the Area, both qualitatively and quantitatively, envisaged to be the results of socio-economic activities taking place in the Area. Finally, the study attempts to look into the measures put in place to curb these challenges. Amid paucity of data, however, the study reveals that the most single contributor of urbanization processes in the area has been migration from the countryside, and in particular from the northern part of the country. The study also reveals the fact that even though the general quality standard of some water resources is better, the largely uncontrolled socio-economic activities, coupled with the deplorable sanitary conditions in the KMA, has potential to degrade the water resources in the KMA. In addition, the study reveals that little attention is given, in terms of policy formulation to curb this urbanization processes and to protect water resources in the area.

An understanding of the impacts on freshwater systems from water management and development practices and the recognition that freshwater ecosystems are dependent on various aspects of the natural hydrological variability has given rise to a relatively new field of science commonly referred to as environmental flows. The assessment and application of environmental flows has advanced considerably in the last ten years. To coincide with the emergence and expansion of the environmental flows concept around the world, this survey study was undertaken aiming to identify people’s perceptions and interpretations of the environmental flows concept and its role in water management. It also aims to add to existing knowledge of the extent to which the concept is being applied, how it is being translated into policy and practice and the major challenges and opportunities that exist for continued understanding and implementation. The survey was distributed to a range of people representing different water-related sectors and regions around the world. A total of 272 responses representing 64 countries in the six major regions of the world was received. The responses were compiled and analysis of aspects of the respondents backgrounds as well as the questions were conducted using the computer statistical program SPSS. Representation of specific groups, particularly water user groups, and specific regions in the survey however was low, demonstrating possible limitations of the survey distribution method as well as the lack of concept awareness and application in many parts of the world. The ways in which people define and interpret the concept varied widely. The degree to which the concept was applied shows the growing recognition around the world of the need to consider the environmental water requirements when making decisions on water allocations. Despite the growing recognition many areas do not yet apply the concept. The survey allowed the opportunity for respondents to highlight what they perceived asthe major obstacles and difficulties for the concept within their respective areas. Lack of understanding among stakeholders of the socio-economic costs and benefits associated with concept implementation and a lack of political will were the two most common obstacles for the continued adoption and application of environmental flows around the world. Overall, the survey delivered promising signs for the continued evolution of environmental flows within water management. There was widespread opinion that the concept of environmental flows was an essential element in the efforts to achieve sustainable management of water resources.

Female reproductive decisions have been suggested to be highly influenced by mate quality. I have studied whether offspring quality may be adjusted by females to match the attractiveness of males and how strong control females have over their reproductive investment focusing on egg size. This was done in the Banggai cardinalfish (Pterapogon kauderni), a sex-role reversed obligate paternal mouthbrooder where males invest heavily into reproduction. As this species is suitable for both laboratory experiments and field studies it is an ideal candidate for the study of reproductive investment. Mating was size-assortative and both males and females benefited from pairing with large partners. However, male size determined the reproductive output of a pair. Females courted large males more intensively and produced larger, but not fewer eggs when mated to large males as compared to small males. Further, this matching of egg size to mate attractiveness may be fast. Female courtship behaviours contained honest information regarding both clutch weight and egg maturity, traits that may be highly important for male mate choice. Surprisingly, males played an important part in territory defence suggesting relatively equal sex-roles in this species. Also, this species showed stable group structures which may be important for the evolution of female plasticity in reproductive investment due to high variance in quality of available mates. This thesis suggest that females have a remarkable control over their reproductive investments and that male quality may be highly influential on reproductive decisions regarding offspring quality. Furthermore, it suggest that sexual selection may have strong effects on the evolution of egg size and parental care on a whole.

<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>