Environmental factors influence species and habitats on multiple scales creating a mosaic of distribution patterns. Studying factors shaping these patterns are central to our understanding of population dynamics and ultimately ecosystem functioning. Information on the distribution of resources and conservation values are also highly needed in marine management as coastal areas are increasingly influenced by human activities. In this thesis, large-scale field data is used to explore how strong environmental gradients found on multiple scales in the coastal areas of the Baltic Sea influence fish habitats. The underlying concepts are based in the field of species distribution modelling, whereby habitat maps can be produced using environmental layers in a geographic information system. Distribution modelling is further used to address both ecological and applied questions by examining effects of habitat limitation on fish population sizes and to evaluate management actions aimed at habitat conservation. I show that specific habitat requirements for fish species of both freshwater and marine origin can be described using environmental variables and that species-environment relationships can be used to predict the distribution of early life-stages of fish in the Baltic Sea archipelagos. Further, predicted habitat availability of a specific life-stage was directly related to adult population size of Eurasian perch Perca fluviatilis, signifying that the abundance of large predatory fish can be limited by specific recruitment habitats. Lastly, by predicting the distribution of an assemblage of coastal fish species and their associated habitats, an assessment of a network of marine protected areas was performed. Results revealed large gaps in the current network and identified areas suitable for future protection. By demonstrating how current habitat protection can be improved by including critical habitats for coastal fish population sizes this thesis points to the benefits of integrating nature conservation and fisheries management. Based on these findings I conclude that species distribution modelling provides a suitable analytical framework for assessing the habitat requirements of organisms. An increased understanding of habitat-population relationships and an ability to accurately map ecologically important features will be of great value for an ecosystem-based marine management. ­ / Felaktigt tryckt som Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 709

This doctoral dissertation was produced in a cooperation between Halmstad University (Wetland Research Centre, School of Business and Engineering) and Lund University (Limnology &amp; Marine Biology, Department of Ecology). Abstract . Wetland creation at large, regional scales is implemented as a measure to abate the biodiversity loss in agricultural landscapes and the eutrophication of watersheds and coastal areas by non-point source nutrient pollution (mainly nitrogen). The consequences of creating many new wetlands for biodiversity conservation and nutrient reten- tion (ecosystem functioning) in agricultural landscapes are still relatively unknown, both on local (per wetland) and regional (per landscape) scales. In Sweden, wetland creation has progressed already since the 1990s, and by now larger numbers of created wetlands are present, mainly in the intensively farmed landscapes of southwestern Sweden. This thesis aimed to investigate the following aspects in these systems: (i) their large-scale effects on biodiversity, (ii) their functional diversity of bacterial denitrifiers, (iii) the abiotic and biotic influences on wetland ecosystem functioning, (iv) the potential for biodiversity-function links, and (v) the potential for functional links and joint functioning.(i) Created wetlands hosted diverse assemblages of macroinvertebrates and plants. They maintained a similar com- position and diversity as natural ponds in agricultural landscapes. The environmental conditions per wetland did hardly affect macroinvertebrate and plant assemblages, and the prerequisites for nutrient retention did neither. In landscapes were wetland creation efforts had increased the total density of small water bodies by more than 30%, macroinver- tebrate diversity of created wetlands was facilitated on both local and regional scales. (ii) Diverse communities of denitrifying bacteria with the capacity for conducting different denitrification steps (functional types) were present in all investigated wetlands. The richness of denitrifying bacteria communities was affected by nitrate concentration and hydraulic loading rate, which may potentially be relevant for the nitrogen retention function of created wetlands. The diversity across different functional types of bacterial denitrifiers increased with nitrate concentration. (iii) Both abiotic and biotic factors influenced ecosystem functions of created wetlands. Variation in nitrogen retention was associated to nitrate load, but even to vegetation parameters. In wetlands with constant nitrate load, planted emergent vegetation facilitated nitrogen retention compared to other vegetation types. In wetlands with variable loads, nitrogen retention was facilitated if nitrate load was high and many different vegetation types were present; nitrogen load could explain the majority of the variation in nitrogen retention compared to vegetation parameters. Phosporus retention of created wetlands was best explained by vegetation parameters. Litter decomposition was inhibited at high nitrate to phosphorus ratios. Methane production increased with age and decreased with plant cover. (iv) Biodiversity may facilitate wetland ecosystem functions, particularly in dynamic wetland ecosystems. Nitrogen retention increased with vegetation type diversity, phosphorus retention capacity with plant richness, and litter decomposition with macroinvertebrate diversity. (v) Created wetlands have the capacity of sustaining several parallel ecosystem services. Some wetland functions were coupled; nitrogen retention increased with fast litter decomposition. On the other hand, methane emission and nitro- gen retention were independent of each other, as were nitrogen and phosphorus retention.In conclusion, created wetlands have the potential to at least partly abate the lost biodiversity and multifunctionality caused by the past extensive destruction of natural wetlands in agricultural landscapes. / <p>[Paper II] Milenkovski S., Thiere G., Weisner S.E.B., Berglund O. &amp; Lindgren P.-E. Variation of eubacterial and denitrifying bacterial biofilm communities among constructed wetlands. Submitted manuscript. [Paper V] Thiere G. &amp; Weisner S.E.B. Influence of biotic and abiotic parameters on ecosystem functioning of created wetlands. Manuscript.</p>

Benthic diatoms are valuable tools for biological monitoring and paleo-ecological reconstruction of past environmental conditions. This thesis aims at describing size-related properties of benthic diatoms and suggests that data assessment for community analysis can be improved by considering the importance of scale. It investigates which environmental factors structure epilithic diatom communities on the coast of the Gulf of Bothnia and identifies environmental factors correlated with phytobenthic biomass. It also contributes to the floristic knowledge of the Baltic Sea. Diatom species show large variation in size. The responses of large species (≥1000 μm3) in diatom communities to environmental factors are underestimated if solely measured as relative abundance, as is the common practice. However, relative abundance gives the best gradient resolution, as compared to surface area and biovolume, if species are counted separately in two biovolume classes. Small and large species in the same community may respond differently to the same environmental factors. To assess the principal environmental factors structuring diatom communities in the Gulf of Bothnia, 270 quantitative samples were collected from submerged stones. Sampling was carried out in spring in four areas of the Bothnian Bay, characterized by a stable north-south salinity gradient (0.4-3.3 psu), and in three areas of the Bothnian Sea which has a rather uniform salinity of ca. 5 psu. Canonical correspondence analysis (CCA) identified salinity and exposure to wave action as the principal factors structuring the diatom communities of the Bothnian Bay, whereas exposure to wave action was the principal factor in the Bothnian Sea. Measurements of relative ignition loss suggested that the cover of macroalgae, and thereby the higher abundance of epiphytic diatoms in the epilithic samples, was positively correlated with salinity in the Bothnian Bay and with water movement in the Bothnian Sea. Two new brackish water species are described, Navicula sjoersii S. Busse &amp; Snoeijs and N. bossvikensis S. Busse &amp; Snoeijs. The new species are compared with N. perminuta Grunow, a common brackish-water species.

The work in this thesis deals with (1) the effects of woody debris on stream channel morphology and retention of organic material, and (2) the dynamics of woody debris and its relation to riparian forest history and composition. The studied stream reaches are situated in mature, productive forests in the boreal zone of Sweden. Wood variables were important predictors of the frequency of debris dams, pool area, the proportion of pools formed by wood, and variation in the bankfull channel width. Pools formed by woody debris were mainly created by damming and had larger surface areas and residual depths than pools formed by other agents. Stream reaches intersecting old-growth forest (with minor influence of forest management) had coarser and longer woody debris pieces, greater amounts of wood, more debris dams, and wood-formed pools compared to streams surrounded by forests influenced by selective logging. The influence of past forest management on the quality and quantity of woody debris in streams were analyzed by using dendrochrnological methods. Selective loggings and absence of forest fires after 1831 resulted in lower input rates and a gradual replacement of pine by sruce over time. Residence times in stream channels of woody debris (&gt;10 cm in basal diameter) were long and the oldest dated pieces of pine and spruce were over 300 and 100 years, respectively. Dynamics of woody debris were explored by comparing wood volumes and characteristics between stream channels and their riparianforests and between old growth and managed sites. Wood volumes recorded in the stream channels exceeded, but were related to, the volumes found in the riparian forests. Limited input of woody debris by bank cutting and absence of slope processes suggest that recruitment processes of woody debri to stream channels are similar as in riparian forests and slow decay in channels results in greater volumes. The retentiveness of organic material in stream channels was examined by using release and capture experiments in multiple reaces during varying discharges using different sizes of leaf mimics. Sixty eight percent of the variation in retention was explained by a multiple regression model including discharge and leaf mimic siz. Between 44 and 80% of the variation in retention among reaches was explained by channel constraint, gravel coverage, and woody debris variables as the most important. Estimates from a partial least squares (PLS) model suggest an increase in mean transport distances by 22 to 53% in managed forest streams compared to old growth conditions and in a low wood scenario, mean transport distances increased by 38 to 99% with larger increases for higher discharges and larger particle sizes. To regain more pristine conditions of stream channels, management and restoration are needed to increase the amount of woody debris that recreates lost channel structures and increaes the retention of organic material.

The brood-nosed pipefish Syngnathus typhle is a pipefish with reversed sex roles. Males prefer to mate with large and ornamented females and females compete for partners. It has previously been shown that males mate more randomly when water becomes turbid. In the Baltic Sea eutrophication has become a severe problem with turbid waters as one consequence. Turbidity makes visual cues less effective and thus weakens sexual selection in species using vision to discriminate between partners. It also affects mate encounter rates in species using vision to find each other. A lower mate encounter rate means a weakening of sexual selection and that individuals mate more randomly. One way to compensate for lower visibility would be to use olfactory cues instead. In this study I investigated whether S. typhle could use olfactory cues to discriminate between sexes. I found no support that they could use olfactory cues to find a partner.

<p>The purpose of this thesis is to analyse interactions between freshwater flows, terrestrial ecosystems and human well-being. Freshwater management and policy has mainly focused on the liquid water part (surface and ground water run off) of the hydrological cycle including aquatic ecosystems. Although of great significance, this thesis shows that such a focus will not be sufficient for coping with freshwater related social-ecological vulnerability. The thesis illustrates that the terrestrial component of the hydrological cycle, reflected in vapour flows (or evapotranspiration), serves multiple functions in the human life-support system. A broader understanding of the interactions between terrestrial systems and freshwater flows is particularly important in light of present widespread land cover change in terrestrial ecosystems. </p><p>The water vapour flows from continental ecosystems were quantified at a global scale in Paper I of the thesis. It was estimated that in order to sustain the majority of global terrestrial ecosystem services on which humanity depends, an annual water vapour flow of 63 000 km3/yr is needed, including 6800 km3/yr for crop production. In comparison, the annual human withdrawal of liquid water amounts to roughly 4000 km3/yr. A potential conflict between freshwater for future food production and for terrestrial ecosystem services was identified. </p><p>Human redistribution of water vapour flows as a consequence of long-term land cover change was addressed at both continental (Australia) (Paper II) and global scales (Paper III). It was estimated that the annual vapour flow had decreased by 10% in Australia during the last 200 years. This is due to a decrease in woody vegetation for agricultural production. The reduction in vapour flows has caused severe problems with salinity of soils and rivers. The human-induced alteration of vapour flows was estimated at more than 15 times the volume of human-induced change in liquid water (Paper II). </p>

<p>The thesis analyses relationships between ecological and social systems in the context of coastal ecosystems. It examines human impacts from resource extraction and addresses management and governance behind resource exploitation. The main premises are that a lack of ecological knowledge leads to poor ecosystem management and that the dichotomy between social and natural systems is an artificial one. The thesis illustrates the importance of basing resource management on the ecological conditions of the resource and its ecosystem. It also demonstrates the necessity of accounting for the human dimension in ecosystem management and the challenges of organising human actions for sustainable use of ecosystem services in the face of economic incentives that push users towards short-term extraction.</p><p>Many Caribbean coral reefs have undergone a shift from coral to macroalgal domination. An experiment on Glovers Reef Atoll in Belize manually cleared patch reefs in a no-take zone and a fished zone (Papers I and II). The study hypothesised that overfishing has reduced herbivorous fish populations that control macroalgae growth. Overall, management had no significant effect on fish abundance and the impacts of the algal reduction were short-lived. This illustrated that the benefits of setting aside marine reserves in impacted environments should not be taken for granted. </p><p>Papers III and IV studied the development of the lobster and conch fisheries in Belize, and the shrimp farming industry in Thailand respectively. These studies found that environmental feedback can be masked to give the impression of resource abundance through sequential exploitation. In both cases inadequate property rights contributed to this unsustainable resource use. </p><p>The final paper (V) compared the responses to changes in the resource by the lobster fisheries in Belize and Maine in terms of institutions, organisations and their role in management. In contrast to Maine’s, the Belize system seems to lack social mechanisms for responding effectively to environmental feedback. The results illustrate the importance of organisational and institutional diversity that incorporate ecological knowledge, respond to ecosystem feedback and provide a social context for learning from and adapting to change.</p>

<p>Modern food production is a complex, globalized system in which what we eat and how it is produced are increasingly disconnected. This thesis examines some of the ways in which global trade has changed the mix of inputs to food and feed, and how this affects food security and our perceptions of sustainability. One useful indicator of the ecological impact of trade in food and feed products is the Appropriated Ecosystem Areas (ArEAs), which estimates the terrestrial and aquatic areas needed to produce all the inputs to particular products.</p><p>The method is introduced in Paper I and used to calculate and track changes in imported subsidies to Swedish agriculture over the period 1962-1994. In 1994, Swedish consumers needed agricultural areas outside their national borders to satisfy more than a third of their food consumption needs. The method is then applied to Swedish meat production in Paper II to show that the term “Made in Sweden” is often a misnomer. In 1999, almost 80% of manufactured feed for Swedish pigs, cattle and chickens was dependent on imported inputs, mainly from Europe, Southeast Asia and South America. Paper III examines ecosystem subsidies to intensive aquaculture in two nations: shrimp production in Thailand and salmon production in Norway. In both countries, aquaculture was shown to rely increasingly on imported subsidies. The rapid expansion of aquaculture turned these countries from fishmeal net exporters to fishmeal net importers, increasingly using inputs from the Southeastern Pacific Ocean.</p><p>As the examined agricultural and aquacultural production systems became globalized, levels of dependence on other nations’ ecosystems, the number of external supply sources, and the distance to these sources steadily increased. Dependence on other nations is not problematic, as long as we are able to acknowledge these links and sustainably manage resources both at home and abroad. However, ecosystem subsidies are seldom recognized or made explicit in national policy or economic accounts. Economic systems are generally not designed to receive feedbacks when the status of remote ecosystems changes, much less to respond in an ecologically sensitive manner. Papers IV and V discuss the problem of “masking” of the true environmental costs of production for trade. One of our conclusions is that, while the ArEAs approach is a useful tool for illuminating environmentally-based subsidies in the policy arena, it does not reflect all of the costs. Current agricultural and aquacultural production methods have generated substantial increases in production levels, but if policy continues to support the focus on yield and production increases alone, taking the work of ecosystems for granted, vulnerability can result. Thus, a challenge is to develop a set of complementary tools that can be used in economic accounting at national and international scales that address ecosystem support and performance.</p><p>We conclude that future resilience in food production systems will require more explicit links between consumers and the work of supporting ecosystems, locally and in other regions of the world, and that food security planning will require active management of the capacity of all involved ecosystems to sustain food production.</p>

<p>Conservatism is a central theme of organismic evolution. Related species share characteristics due to their common ancestry. Some concern have been raised among evolutionary biologists, whether such conservatism is an expression of natural selection or of a constrained ability to adapt.</p><p>This thesis explores adaptations and constraints within the plant reproductive phase, particularly in relation to the evolution of fleshy fruit types (berries, drupes, etc.) and the seasonal timing of flowering and fruiting. The different studies were arranged along a hierarchy of scale, with general data sets sampled among seed plants at the global scale, through more specific analyses of character evolution within the genus <i>Rhamnus </i>s.l. L. (Rhamnaceae), to descriptive and experimental field studies in a local population of <i>Frangula alnus</i> (Rhamnaceae). Apart from the field study, this thesis is mainly based on comparative methods explicitly incorporating phylogenetic relationships. The comparative study of <i>Rhamnus</i> s.l. species included the reconstruction of phylogenetic hypotheses based on DNA sequences.</p><p>Among geographically overlapping sister clades, biotic pollination was not correlated with higher species richness when compared to wind pollinated plants. Among woody plants, clades characterized by fleshy fruit types were more species rich than their dry-fruited sister clades, suggesting that the fleshy fruit is a key innovation in woody habitats. Moreover, evolution of fleshy fruits was correlated with a change to more closed (darker) habitats.</p><p>An independent contrast study within <i>Rhamnus</i> s.l. documented allometric relations between plant and fruit size. As a phylogenetic constraint, allometric effects must be considered weak or non-existent, though, as they did not prevail among different subclades within <i>Rhamnus</i> s.l. Fruit size was correlated with seed size and seed number in <i>F. alnus</i>. This thesis suggests that frugivore selection on fleshy fruit may be important by constraining the upper limits of fruit size, when a plant lineage is colonizing (darker) habitats where larger seed size is adaptive.</p><p>Phenological correlations with fruit set, dispersal, and seed size in <i>F. alnus</i>, suggested that the evolution of reproductive phenology is constrained by trade-offs and partial interdependences between flowering, fruiting, dispersal, and recruitment phases. Phylogenetic constraints on the evolution of phenology were indicated by a lack of correlation between flowering time and seasonal length within <i>Rhamnus cathartica</i> and <i>F. alnus</i>, respectively. On the other hand, flowering time was correlated with seasonal length among <i>Rhamnus</i> s.l. species. Phenological differences between biotically and wind pollinated angiosperms also suggested adaptive change in reproductive phenology.</p> / <p>Äpplet faller inte långt från trädet. Men varför? Den biologiska mångfalden präglas i stor utsträckning av fylogenetiskt bevarade karaktärsdrag; närbesläktade arter är lika. Det pågår en diskussion bland evolutionsbiologer om i vilken utsträckning denna konservatism är ett resultat av naturlig selektion eller av en begränsad anpassningsförmåga.</p><p>Denna avhandling diskuterar begreppet evolutionära begränsningar i relation till den reproduktiva fasen hos växter. I fokus ligger särskilt evolutionen av bärliknande (endozoochora) frukter respektive evolutionen av säsongsmässiga mönster (fenologi) för blomning och fruktsättning. Avhandlingen är hierarkiskt organiserad så att olika delstudier gjordes på olika skalnivåer: fenologi- och fruktevolution analyserades för fröväxter respektive gömfröiga växter; inom brakvedssläktena Rhamnus och Frangula (Rhamnaceae); samt för en lokal population av brakved (Frangula alnus). Populationsstudien baserades på såväl experimentella som deskriptiva data, medan övriga studier i huvudsak genomfördes med fylogenetisk komparativ metodik baserade på litteraturdata. Som en del av de komparativa studierna rekonstruerades fylogenetiska hypoteser för Rhamnus s.l. utifrån DNA-sekvenser (ITS, trnL-F), vilka gav stöd för att Frangula och Rhamnus är monofyletiska systersläkten.</p><p>I en biogeografiskt kontrollerad fylogenetisk kontrastanalys upptäcktes inga skillnader i artrikedom mellan djur- och vindpollinerade växter. Bärliknande frukter verkar dock vara en betydelsefull karaktär (key innovation) i skogsmiljöer. För det första var fylogenetiska klader med bärliknande frukter mer artrika än systerklader med torra frukter. Dessutom var uppkomster av bärliknande frukt korrelerad till habitat med mer sluten vegetation.</p><p>En fylogenetisk kontrastanalys av allometriska effekter visade på en positiv korrelation mellan växtindividens och fruktens storlek inom Rhamnus s.l. Upprepade analyser av dessa allometriska samband i olika monofyletiska subklader inom Rhamnus s.l. indikerar dock att allometri, såsom evolutionär begränsning, är svag. I populationsstudien av F. alnus var fruktstorlek positivt korrelerad till såväl fröantal som frövikt. En generell hypotes utifrån denna avhandling blir därför att frugivorer (de djur som äter bären och därmed sprider växtens frön) utövar ett starkt selektionstryck på växter som koloniserar ett habitat där fröstorlek är adaptivt, eftersom fruktstorlek påverkar frugivorernas (ssk. fåglars) sätt att hantera frukten.</p><p>Reproduktiv fenologi var korrelerad till fruktsättning, fröspridning och fröstorlek hos F. alnus. Detta stödjer tanken att evolutionen av blomnings- och fruktsättningstider begränsas av avvägningar (trade-offs) och partiella beroenden mellan blomning-, fruktsättnings-, spridnings och rekryteringsfaserna. Inomartsvariation i blomningstid för F. alnus respektive getapel (Rhamnus cathartica) över en nordeuropeisk latitudinell transekt var inte korrelerad till säsongslängd, vilket antyder att utvecklingen av blomningstider är evolutionärt begränsad. Å andra sidan påvisades en sådan korrelation mellan blomningstid och säsongslängd i en mellanartsstudie inom Rhamnus s.l. Den fylogenetiska kontrastanalysen mellan djur- och vindpollinerade växter visade också på adaptiva skillnader i fenologi. Blomningstiderna inom en klad med biotiskt pollinerade arter var mer åtskilda än inom den vindpollinerade systerkladen, och den välkända iakttagelsen att vindpollinerade träd blommor tidigare på säsongen fick även stöd i ett fylogenetiskt komparativt perspektiv.</p>

<p>Patterns of site occupancy among boreal stream insects were studied in central Sweden with focus on sparsely distributed species and the role of dispersal and niche limitations.</p><p>In the study of dispersal limitation, I found that effects of an extraordinarily harsh winter in small to medium-sized streams were strongest in sites located in small streams and far from lake outlets. Species richness and the total abundance of macroinvertebrates and trout returned to pre-disturbance levels after three years. However, some species showed slow recolonization and the proportion of holoaquatic taxa was still reduced after three years. In a second study, I found a positive correlation between site occupancy in stream caddisflies and morphological traits associated with fast and energy-efficient flight, whereas specialized spring caddisflies showed a negative correlation to these traits compared to stream species. This suggested that streams, but not springs, select for strong dispersal ability in caddisflies. In a survey of springs in central Sweden, hydrogeology was found to be a useful predictor of the occurrence of spring specialists. Two of these, <i>Crunoecia irrorata</i> Curtis and <i>Parachiona picicornis</i> (Pictet), were found exclusively in glaciofluvial springs, characterized by a stable discharge and temperature. Less specialized members of the spring fauna (i.e. species also occurring in streams, ponds or lakes) also occurred in moraine and limestone springs characterized by more unstable conditions. </p><p>Niche limitations were studied by contrasting large-scale distributions of closely related rare and common stoneflies. Differences in temperature requirements in the juvenile stages and life cycles suggested that the rare species, <i>Isogenus nubecula</i> Newman, was restricted by a limited tolerance to low stream temperatures, whereas the two common species, <i>Isoperla grammatica</i> (Poda) and <i>Diura nanseni</i> (Kempny), appeared to have a broader tolerance to climatic conditions in the study area. In a second study of niche limitations, macroinvertebrate assemblages in 88 streams in Central Sweden showed a nested distribution pattern. Most species deviating from expected distributions occurred in small streams, indicating competitive exclusion from species-rich sites, predator avoidance, or specialization to unique habitat features of small streams. In the last paper, the longitudinal distribution of filter-feeding caddisflies in a lake-outlet stream demonstrated patterns concordant to feeding specialization. </p>