For over three decades, amphibian populations have been declining across the globe. Emerging infectious diseases are responsible for some of these declines. Ranaviruses have caused die-offs in wild amphibian populations on 4 continents, in 5 Canadian provinces, and in over 25 U.S. states. In order to understand host-pathogen dynamics, it is critical to establish baseline information on species susceptibility and the effects of natural stressors. The goal of my thesis research was to quantify the effects of anuran development and exposure to invertebrate predators on species-specific susceptibility to ranavirus. My experiments were designed in factorial arrangements, and consisted of exposure to ranavirus during different developmental stages or with and without predator cues in a controlled environment. I found that exposure to invertebrate predator cues did not increase susceptibility to ranavirus for 4 anuran species tested. Susceptibility differed among embryo, hatchling, larval and metamorph stages, but trends differed among species and did not follow predictions based on Xenopus laevis immune function. Low susceptibility during the embryo stage was the only consistent development result among species, perhaps owing to protective qualities of the vitelline membrane or mucoidal capsules surrounding the embryo. Across 7 anuran species tested, mean mortality rates ranged from 5 – 100%, with Lithobates sylvaticus and Scaphiopus holbrookii most susceptible. I found that infection rates and viral load were correlated with mortality rates, thus these variables are good indicators of susceptibility to ranavirus. My results indicate that ranaviruses can cause catastrophic natural mortality in some anuran species, and likely play a significant role in local population dynamics. For highly susceptible species, ranaviruses could cause local extirpations that lead to species declines. More information is needed on the role of natural (e.g., co-infection, competition) and anthropogenic stressors in driving ranavirus epizootic events. I encourage natural resource agencies to initiate ranavirus surveillance programs, especially for rare species and fragmented populations. Future studies should take an immunogenetic approach to identifying mechanisms driving susceptibility. Identifying mechanisms associated with ranavirus emergence is fundamental to developing science-based conservation strategies.

The strong increase in number of Great cormorants Phalacrocorax carbo in Sweden in recent years has led to conflicts - particularly with fishery. This thesis focuses on the possible effects of cormorant predation on fish populations. In total, data from 15 lakes in South Sweden were included in this study while most studies were carried out in Lake Ymsen. The results suggest that the impact of cormorant predation on natural fish populations was small, and I observed no decline in fish mass after cormorants established. Cormorant predation on eel was difficult to evaluate because of several confounding factors. Ruffe, roach and perch were the most important prey species to the cormorants and most fish taken were small. Cormorants do not seem to catch species and sizes in proportion to their occurrence in the fish community. Total fish removal by cormorants varied considerably among lakes (0.2-15.0 kg/ha) and cormorant population sizes at the different lakes were significantly positively correlated with fishery catches, which in turn was significantly positively correlated with total phosphorous levels. Thus, cormorant densities in lakes, and perhaps elsewhere, seem to be governed chiefly by fish densities. The fact that cormorant predation appears not to reduce fish densities suggest cormorants to be regulated by other means than prey depletion. The mechanism behind population regulation could be a behavioural response of fish, making fish more difficult to catch for the cormorants. In recent years, cormorant populations have been subjected to intensive legal and illegal actions with the aim to reduce cormorant numbers. However, the actions currently carried are well below the efforts needed to limit population sizes. To conclude, cormorants appear to compete little with fishery, with regards to free-living fish. The main problem is that cormorants sometimes damage and take away fish in fishing gears.

This thesis examines risk management in breeding Siberian jays (Perisoreus infaustus), which is indigenous to the northern taiga. Parent behaviour and the nest are cryptic. A new nest is built each year. It is placed on spruce or pine branches close to the trunk and well insulated with lichens, feathers and reindeer hair. Nest failure rate was the main factor driving annual variations in jay numbers. The probability for nesting attempts to be successful ranged annually between 0.08 and 0.70. Nest predation was rampant and a main cause of nest failure. Nest predators were mainly other corvids (primarily the Eurasian jay Garrulus glandarius). Habitat quality was the main factor determining the risk of predation. The risk for nest failure due to predation was higher in thinned forests with an open structure and with a high abundance of man-associated corvid species (jays, crows, raven). Siberian jay parents show several strategic adjustments in life-history and behaviour to the risk of nest predation. Parents traded reduced feeding rates for a lower predation risk and allocated feeding to low risk situations. Chick provisioning imposes a cost by drawing the attention of visually hunting predators to the location of nests, and parents adjusted their daily routines and avoided exposure by allocating provisioning to times of low activity among nest predators. These strategic adjustments of feeding efforts were estimated to reduce the exposure to nest predators by 26 percent. Also, parents adjusted their reproductive efforts to the perceived presence of predators in a playback experiment. Siberian jays reduced their reproductive investment by laying a smaller clutch size when high risk of nest predation reduced the value of current reproduction, as predicted from life-history theory.

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? 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, Chrysochromulina sp., at low nutrient concentrations. To our knowledge this is the first experimental study showing that Chrysochromulina sp. in resemblance with other mixotrophs is favoured by nutrient poor conditions compared to strict autotrophs and heterotrophs. 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. 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.

The abundance of predators and their impact on ecosystem dynamics is a vividly discussed topic in current ecology. In my studies, incorporating field observations, field experiments and theoretical modeling, I explored the importance of predators and predation in a low arctic tundra ecosystem in northern Norway. This involved observing the abundance and spatial activity of predators (raptors and small mustelids); manipulating the abundance of predators (spiders and birds) in an arthropod community; and exploring the theoretical consequences of intraguild predation on the coexistence among predators. The results show that predation is important both in the arthropod assemblage and, depending on the productivity of the community, in the vertebrate assemblage. In arthropod communities predators are at least as abundant as their prey, whereas in the vertebrate part of ecosystem, predators are substantially less abundant than their prey. Still, in both cases predators had strong impact on their prey, influencing the abundance of prey and the species composition of prey assemblages. The impact of predation cascaded to the plant community both in the reticulate and complex arthropod food web and in the linear food chain-like vertebrate community. In the vertebrate-based community we could even observe the long time scale effect on plant community composition. Within the predator community, exploitation competition and intraguild predation were the structuring forces. As the arthropod communities consist of predators with different sizes, intraguild predation is an energetically important interaction for top predators. As a consequence, they reduce the abundance of intermediate predators and the impact of intermediate predators on other prey groups. Moreover, being supported by intermediate predators, top predators can have stronger impact themselves on other prey groups. In vertebrate communities, intraguild predation seems to be unimportant as energetic link, instead it manifests as an extreme version of interference competition. Therefore intraguild predation reduces the likelihood of coexistence, as it is due limited prey diversity and intense exploitative competition already precarious in the low arctic tundra. In conclusion, predators have strong impact on their prey, especially in the more productive parts of the low arctic tundra. This applies even to the food webs with complex and reticulate structure, and these effects carry through the community both in the short time scale of population growth and on the long time scale of population generations.

I combined comparative and experimental studies in the field and in the greenhouse to examine factors influencing reproductive success in two insect-pollinated herbs, the tristylous, selfincompatible perennial Lythrum salicaria and the self-compatible, biennial Pedicularis palustris. More specifically, I explored (i) the effects of plant population characteristics on the intensity and outcome of interactions with pollinators and seed predators, (ii) whether flower morphology affects duration of stigma receptivity, and (iii) whether damage-induced reduction in floral display reduces pollinator visitation and increases pollen limitation. As predicted, the rate of pollinator visitation tended to increase and the number of flowers probed per plant tended to decrease with increasing population size in L. salicaria, but these relationships only approached statistical significance. By taking advantage of the pollen size polymorphism that is typical of many heterostylous plants, I could show that the number of compatible pollen grains received increased with population size, and that this was associated with a reduction in pollen limitation and increased seed output per flower. The deposition of high numbers of incompatible conspecific and heterospecific pollen grains did not appear to reduce seed set. In P. palustris, fruit set and seed predation varied markedly among populations and years, but this variation could only partly be explained by variation in population size. Fruit set was positively related to population size, and seed predation was negatively related to population size, in one of three years. Similarly, the level of pollen limitation, which was quantified in two years, varied among populations, but was not related to population size, density or isolation. In L. salicaria, both the duration of stigma receptivity and the effect of prior self-pollination on seed output varied among style morphs. These differences may contribute to morph-specific differences in pollen limitation and seed production documented in the field. The results of a field experiment demonstrated that damage to the shoot apex may markedly reduce fruit production in L. salicaria, and suggested that this is mainly because damage reduces flower number. I found no evidence that a reduced floral display increased the severity of pollen limitation. Taken together, the results show that interactions with both pollinators and herbivores may markedly affect reproductive output in the two plant species, and that the intensity of both mutualistic and antagonistic interactions vary considerably in space and time. Moreover, they are consistent with the hypothesis that pollination success should depend less on population size in selfcompatible than in self-incompatible plants.

Plant-animal interactions are affected by both abundance and distribution of interacting species and the community context in which they occur. However, the relative importance of these factors is poorly known. I examined the effects of predator host range, environmental factors, host plant populations, plant traits and fruit abortion on the intensity of pre-dispersal seed predation in 46 host populations of the perennial herb Lathyrus vernus. I recorded damage by beetle pre-dispersal seed predators, mainly Apion opeticum and Bruchus atomarius with different host ranges on L. vernus as well as on two additional host plants. Local seed predator population size was mainly influenced by plant population size, current seed production and beetle population size in the previous year, but was not strongly affected by connectivity. The monophagous seed predator was less abundant and had lower densities than the oligophagous. Both predator species had a strong ability to track fluctuations in seed production; intensity of predation increased with relative increases in seed production. Oligophagous predation on L. vernus increased with the abundance of alternative hosts, but presence of L. vernus did not affect predation on alternative hosts. Abundances and trait preferences differed among three co-occurring seed predators, but were also associated with the abundance of the other species. Overall, seed predation influenced selection on flower number. I found clear indications of seed predator offence but no obvious plant defence. The pattern of fruit abortion was associated with reduced plant fitness since the seed predator had an advanced ability to locate fruits with high probability of retention. Taken together, different factors influencing abundance of the seed predator species, different preferences, and context dependent trait selection are likely to result in complex spatio-temporal variation in overall seed losses and trait selection in the common host plant.

Hard clams, Mercenaria mercenaria, are slow-moving organisms that are heavily preyed upon by both blue crabs and knobbed whelks in coastal Georgia. Hard clams are unable to escape from these predators, and when found, are commonly injured and/or consumed. Thus, their best survival strategy is to avoid their predators. In this study, we compared changes in clam behavior when exposed to blue crab and knobbed whelk predators. Clams reduced their feeding time when exposed to crabs and whelks, exudates from these predators, and to injured conspecifics. In a field experiment, we compared clam survival when caged predators where near clam beds vs. controls with empty cages. Clam survival was significantly higher when caged crabs or whelks were near, suggesting that clams detected these predators, reduced their feeding time, and were less apparent to ambient consumers. In lab behavioral assays, clams were less responsive to blue crabs in turbulent flows, and in the field, turbulence reduced the distance clams reacted to blue crabs. Previous studies have shown that blue crabs turbulence also diminishes blue crab foraging efficiency, and we conducted a field experiment to determine how turbulence affected clam-crab interactions. Our results suggest that predation intensity is greatest at intermediate turbulence levels, and lowest in flows with low and high turbulence levels. We attribute this pattern of predation intensity to differential effects of turbulence on the sensory abilities of crabs and clams. That is, in low turbulent flows, clams have a sensory advantage over crabs, and initiate avoidance behaviors before they are detected. However, as turbulence increases, clam perception diminishes faster than crabs, switching the sensory advantage to crabs, and making clams more vulnerable to consumers. In highly turbulent flows, crab perception declines at a rate faster than clams, and the sensory advantage returns to clams.

Frogs, toads, and salamanders are well known for harboring an array of distasteful (and poisonous) secondary metabolites, presumably as antipredator defenses; yet few experiments have rigorously demonstrated the efficacy of amphibian chemical defenses against ecologically relevant consumers. For example, despite an absence of rigorous statistical evidence showing their distastefulness to predators, eastern newts (Notophthalmus viridescens (Rafinesque))--a common salamander in lentic North American habitats--are assumed to tolerate diverse predator assemblages because newts secrete tetrodotoxin (TTX), a neurotoxin. Here we combine laboratory and field-based ecology with bioassay-guided separation of chemical extracts to show that eastern newts--although chemically protected against ecologically important consumers in lentic systems--nonetheless suffer substantial predation when tethered in the field. When offered newts with alternative prey (paedomorphic Ambystoma talpoideum), red swamp crayfish (Procambarus clarkii) and largemouth bass (Micropterus salmoides) were 9-10x as likely to feed on A. talpoideum as newts. Additionally, juvenile bluegill (Lepomis machrochirus) were 70% less likely to consume newt eggs compared to control food pellets. We also show that different newt tissues were differentially palatable to predatory fish. All bluegill tested consumed a palatable control food, but only 20% consumed dorsal skin, only 35% ate ventral skin, but 75% fed on newt viscera, suggesting that deterrent metabolites are concentrated in the skin. Bioassay-guided fractionation revealed that crude and water-soluble newt chemical extracts inhibited bluegill feeding, definitively establishing the chemical nature of newt antipredator defenses, although we were unsuccessful at isolating the chemical compounds responsible for unpalatability. Yet, deterrent activity in the polar but not the lipophilic chemical fraction and bioassay results demonstrating that naıve predators rapidly learn to avoid natural concentrations of TTX support the possible role of TTX in suppressing predation on newts. However, when tethered in the field, newt mortality was 55% higher in ponds with predatory fishes than in ponds lacking fishes (62% vs. 40% respectively), indicating the possible existence of other predators that are resistant to (or tolerant of) newt chemical defenses. Together, these results stress the importance of rigorous, ecologically relevant, and hypothesis-driven experimentation to better understand the complexity of chemically- mediated predator-prey interactions, even for well-studied species like N. viridescens.

This collection of research examines how changes in the sensory landscape, mediated by both odor and hydrodynamic properties, impact odor-mediated predator-prey interactions in salt marsh communities. I approached this research using an interdisciplinary framework that combined field and laboratory experimentation to address issues of scale and make connections between predator behavior and patterns of predation in the field. I explored a variety of interactions mediated by changes in the sensory landscape including; indirect effects of biotic structure on associated prey, predator responses to patches of prey with differing density and distribution, and dynamic interactions between predators and prey distributions. I found that biotic structure (oyster reefs [Crassostrea virginica]) has negative indirect effects on associated hard clam prey (Mercenaria mercenaria) through the addition of oyster reef odor cues that attract predators (blue crabs [Callinectes sapidus] and knobbed whelks [Busycon carica])and increase foraging success near the structural matrix. Variation in the structure of patch-scale prey odor plumes created by multiple prey results in predator-specific patterns of predation as a function of patch density and distribution which are mediated by differences in predator sensory ability. There is a potential negative feedback loop between blue crab predators and hard clam prey distributions; clam patches assume random within-patch distributions after exposure to blue crab predators, making the detection of patches by future blue crab predators more difficult. Sensory landscapes are also mediated by water flow, which transports prey odor plumes downstream to predators. Characterization of water flow in small-scale estuary systems indicates that values of turbulent flow parameters are highly context specific and depend on both tidal type (spring, neap, normal) and site. Wind and tidal range seem to be good predictors for wave components and turbulent components of fluctuating flow parameters, respectively, although the strength of their predictive ability is dependent on time scale. Modifications of the sensory landscape through changes in structurally-induced turbulence, mixing of individual plumes from multiple prey, and bulk velocity and turbulence characteristics need to be considered when formulating predictions as to the impact of predators on naturally occurring prey populations in the field.