For many species, distribution and abundance is driven by a trade-off between abiotic and biotic stress tolerance (i.e. physical stress versus competition or predation stress). This trade- off may be caused by metabolic rate differences in species such that slow metabolic rates increase abiotic tolerance but decrease biotic tolerance. I investigated how metabolic rate differences were responsible for an abiotic-biotic tolerance trade-off in brown mudfish (Neochanna apoda) and banded kokopu (Galaxias fasciatus), that drives the allopatric distribution of these fish in podocarp swamp-forest pools. Brown mudfish and banded kokopu distribution across 65 forest pools in Saltwater forest, Westland National Park, New Zealand was almost completely allopatric. Mudfish were restricted to pools with extreme abiotic stress including hypoxia, acidity and droughts because of kokopu predation in benign pools. This meant the mudfish realised niche was only a small fraction of their large fundamental niche, which was the largest out of sixteen freshwater fish species surveyed in South Island West Coast habitats. Thus mudfish had a large fundamental to realised niche ratio because of strong physiological stress tolerance but poor biotic stress tolerance compared to other fish. A low metabolic capacity in mudfish compared to kokopu in terms of resting and maximum metabolic rates and aerobic scope explained the strong mudfish tolerance to extreme abiotic stress, but also their sensitivity to biotic stress by more powerful kokopu in benign pools, and hence their allopatric distribution with kokopu. Despite being restricted to extreme physical stress, mudfish populations were, in fact, more dense than those of kokopu, because of low individual mudfish resting metabolic rates, which would cause resources to be divided over more individuals. Distribution and abundance in mudfish and kokopu were therefore driven by an abiotic-biotic tolerance trade-off caused by a physiological trade-off between having slow or fast metabolic rates, respectively. The negative relationship between species resting metabolic rates and their tolerance to abiotic stress provides a way of estimating the impact of human induced environmental change that can either increase or decrease habitat harshness. Thus species with low metabolic rates, like mudfish, will be negatively affected by human induced environmental change that removes abiotic habitat stress and replaces it with benign conditions. My evidence shows that extreme stressors provide a protective habitat supporting high mudfish biomass with significant conservation value that should be maintained for the long-term persistence of mudfish populations.

Species have specific tolerances to a variety of environmental variables including temperature, dissolved oxygen (DO) and turbidity. Changes in either of these variables can therefore be expected to affect predator-prey interactions in shallow water ecosystems. Temperature drives the metabolic rates of poikilotherms, including fish. Hypoxic conditions generally affect larger fishes to a greater degree than smaller fishes, though the presence of physostomous swim bladders in certain species can alter that relationship. Finally there are species of fish that rely on vision for food acquisition while other species rely on other senses such as chemical cues. Changes in turbidity levels could therefore affect foraging efficiency of visual foragers. This thesis examines the role that each of these environmental variables (temperature, DO and turbidity) can have on community composition and therefore predator prey interactions, with a specific focus on the role of temperature in structuring predator-prey interactions. Laboratory, field and theoretical studies suggest that as temperature increases, encounter rates between predators and prey will increase. Prey are more active, spend more time foraging, and increase their use of risky habitats in warmer environments in laboratory experiments. In the field, prey and predator activity and/or abundance is positively related to temperature. These laboratory and field studies suggest that temperature increases should result in increased predation rtes of prey. Finally, the results of a dynamic state dependent optimization model also suggest that periods of warming will result in a lowering of the probability of survival of the fathead minnow, Pimephales promelas, a prey species, over the-ice free season. A reduction in DO levels in aquatic ecosystems results in a reduction in the number of and/or activity of predators present. This should result in a reduction in predation risk to prey. However, when endothermic predators are factored in to this equation, this reduction in risk may not occur. The presence of avian predators of small forage fish are directly related to the level of DO in the water, regardless of the abundance of prey fish present. This relationship is likely a result of behavioural decisions of prey that occurs in hypoxic conditions. In periods of low DO, prey fishes may exploit areas of higher DO that are closer to the surface of the waters. While their piscine predators may not be able to tolerate the low DO levels regardless of the position of prey in the water column, avian predators appear to be able to cue in to this increase in availability of potential prey, reducing any benefits that might occur by occupying surface areas where DO levels might be slightly higher than lower in the water column. As compared to temperature and DO, turbidity does not appear to affect the potential risk of predation to forage fish. The catch per unit effort (CPUE) of foragers who rely on vision and those that rely on chemical cues to forages, were not related to turbidity levels. Turbidity levels were also not related to the abundance of avian predators. This suggests that in this generally turbid, shallow water ecosystem, changes in turbidity do not affect the overall species composition of the system. Predator-prey interactions in the system are also not likely to be affected by turbidity. In contrast to this, temperature and DO are likely to influence the interactions between predators and their prey in a shallow water ecosystem. Both increases in temperature and decreases in DO may result in increases in predation pressure on prey. While temperature increases will likely result in increased predation on prey by piscine predators, a reduction in DO, which often occurs as temperature increases, will likely result in increased predation on prey by avian predators, even as predation pressure by piscine predators decrease.

The purpose of this thesis is to investigate if the application of the ecosystem services concept can provide decision makers and stakeholders with additional relevant information for decisions regarding establishing of aquaculture on a local level, using the Lake Storsjön as a case study. In order to provide this knowledge, three different tasks are carried out within this thesis: the identification of the main ecosystem services provided by Lake Storsjön, the understanding of ecosystem services that will be affected by the expansion of aquaculture and the identification of the services that will be used for a possible expansion of the aquaculture sector. The CICES methodology (Common International Classification of Ecosystem Services) is used to identify the ecosystem services provided by Lake Storsjön. The understanding of the affected ecosystem services is carried out with the use of a Delphi-inspired approach. The identification of the services required for the establishment of aquaculture is driven by a cross-reference matrix. Lake Storsjön provides thirty-nine out of the fifty-nine ecosystem services included in the CICES. About half of these will be effected by environmental impacts connected to operations of ecosystem management required by aquaculture expansion, mostly with a low or medium degree of impact. An increase in the implementation of aquaculture might cause detrimental trade-offs with these ecosystem services: “Wild animals”, “Genetic materials from all biota“, “Disease control”, “Maintaining nursery populations and habitats” and “Filtration/dilution/sequestration/storage/accumulation by ecosystem”. An expansion of the aquaculture sector requires seven of the ecosystem services provided by Lake Storsjön. The vital services for this process are: “Surface water for non-drinking purposes”, “Filtration/dilution/sequestration/storage/accumulation by ecosystem” and “Chemical condition of freshwater”. These services have to be safeguarded and maintained in order to guarantee adequate conditions for an expansion of this sector. At the same time the expansion of aquaculture supports the service “Animals from in situ aquaculture”. The benefits connected to the implementation of aquaculture shall be economically assessed and compared to the actual economic value delivered by the other ecosystem services in order to further understand the positives and negatives outcomes of aquaculture expansion in an ecosystem services perspective. This is considered to be a relevant step for strategic and decision making processes concerning aquaculture expansion in Lake Storsjön. In addition to the economic perspective, it seems that the most relevant factor when discussing planning and development processes towards ecosystem services is the importance of not overshooting the resilience ability of the ecosystem in order to ensure the accessibility of the services to future generations. This belief is recommended to be applied to Lake Storsjön in order to guarantee a conscious expansion of the aquaculture sector on a social, economic and environmental level.

Species have specific tolerances to a variety of environmental variables including temperature, dissolved oxygen (DO) and turbidity. Changes in either of these variables can therefore be expected to affect predator-prey interactions in shallow water ecosystems. Temperature drives the metabolic rates of poikilotherms, including fish. Hypoxic conditions generally affect larger fishes to a greater degree than smaller fishes, though the presence of physostomous swim bladders in certain species can alter that relationship. Finally there are species of fish that rely on vision for food acquisition while other species rely on other senses such as chemical cues. Changes in turbidity levels could therefore affect foraging efficiency of visual foragers. This thesis examines the role that each of these environmental variables (temperature, DO and turbidity) can have on community composition and therefore predator prey interactions, with a specific focus on the role of temperature in structuring predator-prey interactions. Laboratory, field and theoretical studies suggest that as temperature increases, encounter rates between predators and prey will increase. Prey are more active, spend more time foraging, and increase their use of risky habitats in warmer environments in laboratory experiments. In the field, prey and predator activity and/or abundance is positively related to temperature. These laboratory and field studies suggest that temperature increases should result in increased predation rtes of prey. Finally, the results of a dynamic state dependent optimization model also suggest that periods of warming will result in a lowering of the probability of survival of the fathead minnow, Pimephales promelas, a prey species, over the-ice free season. A reduction in DO levels in aquatic ecosystems results in a reduction in the number of and/or activity of predators present. This should result in a reduction in predation risk to prey. However, when endothermic predators are factored in to this equation, this reduction in risk may not occur. The presence of avian predators of small forage fish are directly related to the level of DO in the water, regardless of the abundance of prey fish present. This relationship is likely a result of behavioural decisions of prey that occurs in hypoxic conditions. In periods of low DO, prey fishes may exploit areas of higher DO that are closer to the surface of the waters. While their piscine predators may not be able to tolerate the low DO levels regardless of the position of prey in the water column, avian predators appear to be able to cue in to this increase in availability of potential prey, reducing any benefits that might occur by occupying surface areas where DO levels might be slightly higher than lower in the water column. As compared to temperature and DO, turbidity does not appear to affect the potential risk of predation to forage fish. The catch per unit effort (CPUE) of foragers who rely on vision and those that rely on chemical cues to forages, were not related to turbidity levels. Turbidity levels were also not related to the abundance of avian predators. This suggests that in this generally turbid, shallow water ecosystem, changes in turbidity do not affect the overall species composition of the system. Predator-prey interactions in the system are also not likely to be affected by turbidity. In contrast to this, temperature and DO are likely to influence the interactions between predators and their prey in a shallow water ecosystem. Both increases in temperature and decreases in DO may result in increases in predation pressure on prey. While temperature increases will likely result in increased predation on prey by piscine predators, a reduction in DO, which often occurs as temperature increases, will likely result in increased predation on prey by avian predators, even as predation pressure by piscine predators decrease.

Most eukaryotes harbor a diverse community of parasitic, mutualistic, and commensal microbial symbionts. Although the diversity of these microbial symbiotic communities has recently drawn considerable attention, theory regarding the evolution of interactions among symbionts and with the host is still in its nascent stages. Here we evaluate the role of interactions among coinfecting symbionts in the evolution of symbiont virulence toward the host. To do so, we place the virulence-transmission trade-off into a community context and model the evolution of symbiont trophic modes along the continuum from parasitism (virulence) to mutualism (negative virulence). We establish a framework for studying multiple infections of a host by the same symbiont species and coinfection by multiple species, using a concept of shared costs, wherein the negative consequences of virulence (or harm) toward the host are shared among symbionts. Our results show that mutualism can be maintained under infection by multiple symbionts when shared costs are sufficiently low, while greater virulence and parasitism toward the host are more likely when shared costs are high. Last, for coinfection by more than one species, we show that if the presence of a mutualist ameliorates some of the costs of pathogen virulence, then the symbiotic community may more often evolve to a more commensal state and maintain mutualisms.

No description available.

No description available.

Amphibians are influenced by climate change, but we do not have a clear understanding of how changes in temperature, precipitation, or both, may influence amphibian larvae in temperate regions. Do amphibian larvae have similar developmental responses to increased temperature and increased drying rates of wetlands - both plausible effects of climate change? What influence do the interactive effects of temperature and drying have on the relationships between the larval responses themselves (e.g., survival, body size, and time to metamorphosis)? To address these questions, we studied larval responses of two amphibian species, wood frogs (<em>Lithobates sylvaticus</em>) and spring peepers (<em>Pseudacris crucifer</em>), to simulated warming and drying in experimental ponds. Over 13 weeks, we manipulated temperature and water levels in ponds to produce 4 treatments: control, drying, warming, and drying + warming. Our manipulations created warming treatments that were on average 2 ° C higher than controls, and our drying treatments decreased in water depth by 2.5 cm each week compared to warming and control treatments that held a consistent amount of water. In both species, warming treatments resulted in significantly earlier timing of metamorphosis, and drying treatments resulted in significantly reduced body size. We saw a negative relationship between body size and time to metamorphosis (i.e., individuals that metamorphosed faster generally had larger body sizes), indicating an unexpected decoupling of the typical positive relationship between time to- and size at metamorphosis. The strength of the relationship between responses also varied by treatment for wood frogs but not spring peepers, indicating that the responses of larval amphibians to climate change may vary among species. Our study reveals complex relationships among larval survival, body size, and time to metamorphosis and highlights the need for considering not only the role of interacting climate-related pressures on amphibians but also the mechanisms underlying coupling of larval responses to these pressures. We encourage future research and discussion on a better understanding of why different climate pressures caused different responses, and if these patterns may be consistent in other aquatic species. / Master of Science / Across the globe, shifts in temperatures and the availability of freshwater habitats due to climate change are presenting challenges as well as opportunities for many species, particularly those that rely on freshwater habitats to complete their life cycle. Climate change is leading to warmer water temperature and accelerated drying of wetlands and ponds. Warming and drying often occur simultaneously, yet our understanding of how warming and drying may interact and affect sensitive aquatic species is limited. Amphibians with an aquatic life stage (for example, frog tadpoles) are particularly vulnerable to the effects of climate change on wetlands and ponds because they must transform from swimming larvae to land-dwelling adults before aquatic habitats dry out. Warming and drying help amphibian larvae determine when to start that process, called metamorphosis. For this reason, amphibian larvae in aquatic habitats are especially vulnerable to shifts in water temperature and the timing of drying. In this thesis, I explore how warming and drying influence amphibian survival, body size, and time to metamorphosis. To better understand the responses of amphibian larvae to warming and drying, I tested the effects of warming and drying on three response variables: amphibian survival, body size, and time to metamorphosis. I used two different species, wood frogs and spring peepers, to determine whether frogs' responses vary among species. I created an artificial pond experiment where I filled large tanks to represent natural ponds in a controlled, outdoor setting. In these artificial ponds, I measured wood frog and spring peeper growth under experimentally increased water temperatures and accelerated drying levels over 13 weeks. I found a negative relationship between body size and time to metamorphosis, suggesting individuals who spent less time in the water as larvae were more likely to be larger than individuals who spent more time in the water as larvae. Additionally, ponds with higher larval survival were associated with larger body size and a shorter time to metamorphosis. Warmer water temperatures led to a shortened time to metamorphosis but did not always lead to higher body sizes. Accelerated drying did not lead to a shortened time to metamorphosis, but it did lead to smaller body sizes in both species compared to control and warming ponds. Overall, I found complex relationships among larval responses with the directions of responses varying between treatments and species. This highlights the need for considering the role of climate-related changes in the environment (warming and drying) as well as the interactions between specific larval responses to those environmental changes. By understanding how warming and drying influence amphibian larval success, we can make a more direct link between climate change and its effects on aquatic larvae. Incorporating the responses between survival, body size, and time to metamorphosis to gain a more complete understanding of amphibian larval responses to the changing climate is an important step toward conserving and protecting freshwater aquatic species.

Stress can be defined as factors which reduce an individual’s survival and/or reproductive ability. Certain stressors strike harder against individuals the more harmful mutations they carry, thus increasing selection against harmful mutations. The aim of this project is to test if exposure to stress, during many generations, decreases the frequency of harmful mutations and lead to more adapted individuals, or if specific adaptations to the stressor override this effect and results in reduced adaptation in the original environment through trade-offs. To test these hypotheses, I use selection lines of Drosophila melanogaster, where the flies during the larval state either have been exposed to media with reduced nutritional value or a heat shock treatment over 22 generations. The results from this experiment show that the selection lines have adapted to their novel environments, since survival of the larvae had increased in the novel environment they had been exposed to for 22 generations. However, survival of selection lines were if anything decreased rather than elevated in the original environment. A plausible explanation to these results is that adaptations to stressors occur both through trade-offs and reduced frequency of generally harmful mutations, but that the effect of the former possibly is larger than the latter. / Stress kan definieras som faktorer som minskar en individs överlevnad och/eller reproduktiva förmåga. Vissa stressmiljöer slår relativt hårdare mot individer ju fler skadliga mutationer de bär, och ökar därför selektionstrycket mot skadliga mutationer. Detta projekt har som mål att testa om exponering mot sådana stressmiljöer, under flera generationer, minskar frekvensen av skadliga mutationer och leder till generellt bättre anpassade individer, eller om specifika anpassningar till stressmiljön överskuggar denna effekt och via trade-offs leder till individer som är sämre anpassade till ursprungsmiljön. För att testa dessa hypoteser använder jag mig här av selektionslinjer av Drosophila melanogaster, där flugorna under larvstadiet antingen utsatts för en näringsfattig miljö eller en värmechock under 22 generationer. Resultaten från detta experiment visar en tydlig evolutionär respons, i och med att larvöverlevnaden ökat för de selekterade linjerna i den stressmiljö de utsatts för efter 22 generationer. Test av överlevnad i ursprungsmiljön visar dock ingen signifikant skillnad mot kontrollinjerna, men om något att de selekterade linjerna klarade sig något sämre. Dessa resultat tyder om något på att anpassningar som skett till den nya miljön på bekostnad av anpassningar i ursprungsmiljön (via trade-offs) överskuggar ökad anpassning via en minskad frekvens av generellt skadliga mutationer.

Maternal effects are important mechanisms by which mothers’ may influence the phenotype of their offspring. Females may vary in the resources they can provide during offspring development and understanding the factors responsible for this variation is key to understanding offspring success- in early life as well as later life. Differential allocation has been reported to occur, however how it impacts on offspring and mother’s future reproduction still remains unclear. This is also true for maternal transferred substances like maternally transferred immunity. Contributions to date have been limited to snapshots in time, mean level of transfer and/or limited information regarding other maternal traits. For my thesis, I aim to further the understanding of maternal allocation effects and explore the transfer of maternal antibodies over an immune response of a mother, across multiple breeding attempts and accounting for embryo, maternal and paternal traits. Furthermore, I determine the effect of key male traits on general egg traits along with maternal antibodies. I examine this at the individual level using Chinese painted quail (Coturnix chinensis) who are prolific layers and sexually dimorphic. To date the majority of differential allocation studies have not necessarily addressed the assumptions of differential allocation theory. In Chapter 2 of this thesis I attempt to address some of these assumptions and explore the impact of male characteristics across a number of clutches and find separate effects of initial pairing and subsequent pairings. I found that mothers can create, by differential allocation, clutches of varying size but egg components (egg mass) appears to be largely influenced by initial clutch pairing and not by paternal traits. Furthermore, the effect on egg mass appears to be a secondary effect mediated by females adjusting their condition based on their initial pairing. I demonstrate that unlike general clutch traits (clutch size, egg mass) maternal antibodies are not affected by male characteristics (Chapter 3) carry-over effects of egg size means antibody levels may be influenced throughout life by early experiences. However, maternal immune response may be detrimentally linked to viability of offspring. Whereas maternally transferred antibodies appear to have no relationship with maternal or paternal traits, oocyte yolk antibodies during development were found to correlate with female antibodies up to 48hr prior to lay. In Chapter 4, I examine a neglected area regarding maternal effect- exploring variation between female in their transfer of antibodies. Individual females were highly consistent in the relative level of specific blood antibodies transferred to eggs across different phases of their immune response, across challenge types (bacterial and viral) and that some females consistently transfer significantly more than others. The relative level of circulating antibody transferred was independent of the individual’s overall strength of antibody response and related to the female’s body condition (while the individual’s own antibody responses were not). We found no evidence for any trade-offs between the amount transferred and overall reproductive investment in this chapter. In Chapter 5, I discuss the wider implications of my findings and suggest future research directions.