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. 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. 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. 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.

Human-mediated introductions of non-indigenous species now threaten to homogenize the biota of the Globe, causing huge economic and ecological damage. This thesis studies the ecological role of 3 invasive planktonic crustaceans, the omnivorous copepod Acartia tonsa (western Atlantic and Indo-Pacific) and the predatory cladocerans, Cercopagis pengoi (Ponto-Caspian) and Bythotrephes longimanus (Eurasian). B. longimanus invaded the North American Great Lakes in 1982, C. pengoi the Baltic in 1992 and the Great Lakes in 1999, while A. tonsa has an extensive invasion history that includes the Baltic. We review current knowledge on feeding biology of the predatory cladocerans. A study of stable C and N isotope ratios indicated mesozooplankton as the main food source of C. pengoi in the northern Baltic Sea proper, with young C. pengoi also eating microzooplankton, such as rotifers. Young-of-the-year herring did eat C. pengoi and herring trophic position shifted from 2.6 before the invasion to 3.4 after, indicating that C. pengoi had been “sandwiched” into the modified food web between mesozooplankton and fish. Salinity tolerance experiments on Acartia tonsa and co-occurring Acartia clausi showed the formers euryhaline character and high grazing potential. Energy partitioning between ingestion, production and respiration was rather constant over the tested salinity range of 2 to 33, with small differences in gross growth efficiency and cost of growth, but maximum ingestion at 10-20. Egg hatching in A. tonsa was only reduced at the lowest salinity. Extreme changes in salinity were needed to cause significant mortality of A. tonsa in the field, but its feeding activity could be severely reduced by salinity changes likely to occur in estuaries. A study of a hypertrophic estuary showed that A. tonsa can sustain a population despite very high mortality rates, caused by predation, high pH and low oxygen, helping explain the success of A. tonsa as an invader of estuaries.

Aquatic sediments are, by surface, the largest habitat on Earth. A wide diversity of organisms inhabit these sediments and by their actions they have a large influence on and also mediate many ecosystem processes. Several of these processes, such as decomposition and remineralisation of organic matter are important on a global scale and are essential to sustain life on Earth. The main aim of this thesis was to use an experimental ecosystem ecology approach in order to study some of these ecosystem processes in marine sediments and how they are linked to biodiversity. Paper I and II found that an increased species richness of sediment deposit feeders increases the processing of organic matter from phytoplankton settled on the sea-floor, and that species-rich communities have a more efficient resource utilization of deposited organic matter. The results in paper IV and V also suggest that there is a link between microbial diversity in sediments and the degradation of organic contaminants. Paper V also shows that antibiotic pollution is a potential threat to natural microbial diversity and microbially mediated ecosystem services. The introduction of invasive species to ecosystems is another major threat to biodiversity and was studied in Paper II and III, by investigating the ecology of Marenzelleria arctia, a polychaete worm recently introduced in the Baltic Sea. Paper II suggests that M. arctia mainly utilize food resources not used by native deposit feeders, thus potentially increasing the benthic production in the Baltic Sea by increasing resource use efficiency. Paper III, however, show that M. arctia is protected from predation by the native benthic invertebrate predators, due to its ability to burrow deep in the sediment, suggesting that predation on M. arctia by higher trophic levels is restricted, thereby limiting trophic transfer. In conclusion, this thesis gives some examples of the importance of marine biodiversity for the generation of a few key ecosystem processes, such as organic matter processing and the degradation of harmful contaminants. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 4: In press.

There are large gaps in our understanding how fish populations are affected by the anthropogenic noise and the alteration of habitat caused by the construction and operation of offshore wind farms. These issues are of great importance as the construction of offshore wind farms will increase all over the world in the near future. This thesis studies these effects with a focus on fish. The wind turbine foundations function as artificial reefs and are colonized by invertebrates, algae and fish. The epibenthic assemblages are influenced by factors such as hydrographical parameters, time of submergence, distance to natural hard bottom, material and texture (PAPER I, II). Once an epibenthic assemblage has been developed, fish utilize it for different ecosystem services such as food, shelter, and spawning and nursery area. Benthic and semi-pelagic species show a stronger response to the introduced foundation than pelagic species, as it is the bottom habitat that has mainly been altered (PAPER I, II). Pelagic species could be positively affected by the increased food availability - but it takes time and the effect is local. Construction noise like pile driving creates high levels of sound pressure and acoustic particle motion in the water and seabed. This noise induces behavioural reactions in cod (Gadus morhua) and sole (Solea solea). These reactions could occur up to tens of kilometres distance from the source (PAPER III). During power production, the wind turbines generate a broadband noise with a few dominating tones (PAPER IV, V), which are detectable by sound pressure sensitive fish at a distance of several kilometres even though intense shipping occurs in the area. Motion sensitive species will only detect the turbine noise at around a ten meter distance. Sound levels are only high enough to possibly cause a behavioural reaction within meters from a turbine (PAPER IV, V). / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3, 4 and 5: Manuscripts.

The development of human societies rests on functioning ecosystems. This thesis builds on integrated theories of linked social-ecological systems and complex adaptive systems to increase the understanding of how to strengthen the capacity of ecosystems to generate services that sustain human well-being. In this work, I analyze such capacity in human-dominated production ecosystems in Tanzania and Madagascar, and how this capacity is related to local management practices. Resilience of social-ecological systems refers to the capacity to buffer change, to re-organize following disruption, and for adaptation and learning. In Papers I and II, qualitative interview methods are used for mapping and analyses of management practices in the agroecosystem of the Mbulu highlands, Northern Tanzania. Practices such as soil and water conservation, maintenance of habitats for pollinators and predators of pests, intercropping, and landscape diversification, act to buffer food production in a variable environment and sustain underlying ecological processes. The practices are embedded in a decentralized but nested system of institutions, such as communal land rights and social networks, that can buffer for localized disturbances such as temporary droughts. Paper II compares these findings with practices in a farming system in Sweden, and suggests that similar mechanisms for dealing with uncertainty and change can exist in spite of different biophysical conditions. In Papers III and IV, interviews are combined with GIS tools and vegetation sampling to study characteristics and dynamics of the dry forests of Androy, southern Madagascar. Paper III reports on a previously underestimated capacity of the dry forest of southern Madagascar to regenerate, showing areas of regeneration roughly equal areas of degenerated forest (18 700 ha). The pattern of forest regeneration, degradation, and stable cover during the period 1986-2000 was related to the enforcement of customary property rights (Paper III). Paper IV reports on a network of locally protected forest patches in Androy that is embedded in a landscape managed for agricultural or livestock production and contributes to the generation of ecosystem services and ecosystem resilience at a landscape scale. Forest protection is secured by local taboos that provide a well-functioning and legitimate sanctioning system related to religious beliefs. In Paper V, two spatial modeling tools are used to assess the generation of two services, crop pollination and seed dispersal, by the protected forest patches in southern Androy. The functioning of these services is dependent on the spatial configuration of protected patches in the fragmented landscape and can be highly vulnerable to even small changes in landscape forest cover. In conclusion, many of the identified practices are found to make ecological sense in the context of complex systems and contribute to the resilience of social-ecological systems. The thesis illustrates that the capacity of human-dominated production ecosystems to sustain a flow of desired ecosystem services is strongly associated with local management practices and the governance system that they are embedded in, and that, contrary to what is often assumed, local management can and does add resilience for desired ecosystem services. These findings have substantial policy implications, as insufficient recognition of the dynamics of social-ecological interactions is likely to lead to failure of schemes for human development and biodiversity conservation.