Many prey species lack innate recognition of their potential predators. Hence, learning is required for them to recognize and respond to predation threats. When wild-caught, these same species may show amazing sophistication in their responses to predator cues. They are able to adjust the intensity of their antipredator responses to a particular predator according to the degree of threat posed by that predator. This ability is therefore acquired through learning. While many studies have shown that prey can learn to respond to predator cues through different learning modes, little is known about what the prey are actually learning. The results presented in this thesis show that learned predator recognition goes beyond the simple labelling of predators as dangerous. Using fathead minnows (Pimephales promelas), woodfrog (Rana sylvatica) tadpoles and boreal chorus frog (Pseudacris maculata) tadpoles, I demonstrated that a one time learning event, either through pairing with alarm cues or through social learning, was enough for prey to learn the level of threat associated with the novel predator cues. I showed that the level of danger associated with the predator cues was determined by the concentration of alarm cues when learning through pairing of alarm cues, or by the intensity of antipredator response displayed by the tutors and by the tutor-to-observer ratio when learning occurred through cultural transmission. Moreover, when subsequently exposed to predator cues, prey adjusted their antipredator responses according to the change in concentration of predator cues between the learning event and the subsequent exposure. Prey displayed stronger antipredator responses when exposed to higher concentrations of predator cues and vice versa. When minnows were provided with conflicting information about the danger level associated with a predator, they displayed a safety strategy and used the most recent information available to respond to predation threats. On a longer time scale, the data also suggest that woodfrog tadpoles are able to learn to respond to predation threats according to the risk posed by the predator at different times of day. Finally, I showed that prey learn to recognize particular characteristics of predators and can generalize their antipredator responses to novel species sharing those characteristics. However, generalization of predator recognition is dependent on the level of risk associated with the predator. Threat-sensitive learning is an extremely complex process shaped by the millions of years of selection imposed by predators on prey.

Many prey species lack innate recognition of their potential predators. Hence, learning is required for them to recognize and respond to predation threats. When wild-caught, these same species may show amazing sophistication in their responses to predator cues. They are able to adjust the intensity of their antipredator responses to a particular predator according to the degree of threat posed by that predator. This ability is therefore acquired through learning. While many studies have shown that prey can learn to respond to predator cues through different learning modes, little is known about what the prey are actually learning. The results presented in this thesis show that learned predator recognition goes beyond the simple labelling of predators as dangerous. Using fathead minnows (Pimephales promelas), woodfrog (Rana sylvatica) tadpoles and boreal chorus frog (Pseudacris maculata) tadpoles, I demonstrated that a one time learning event, either through pairing with alarm cues or through social learning, was enough for prey to learn the level of threat associated with the novel predator cues. I showed that the level of danger associated with the predator cues was determined by the concentration of alarm cues when learning through pairing of alarm cues, or by the intensity of antipredator response displayed by the tutors and by the tutor-to-observer ratio when learning occurred through cultural transmission. Moreover, when subsequently exposed to predator cues, prey adjusted their antipredator responses according to the change in concentration of predator cues between the learning event and the subsequent exposure. Prey displayed stronger antipredator responses when exposed to higher concentrations of predator cues and vice versa. When minnows were provided with conflicting information about the danger level associated with a predator, they displayed a safety strategy and used the most recent information available to respond to predation threats. On a longer time scale, the data also suggest that woodfrog tadpoles are able to learn to respond to predation threats according to the risk posed by the predator at different times of day. Finally, I showed that prey learn to recognize particular characteristics of predators and can generalize their antipredator responses to novel species sharing those characteristics. However, generalization of predator recognition is dependent on the level of risk associated with the predator. Threat-sensitive learning is an extremely complex process shaped by the millions of years of selection imposed by predators on prey.

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 dissertation demonstrates the construction of the Panthera Population Persistence (PPP), an individual-based model for the Sumatran tiger (Panthera tigris sumatrae) which provides proper theoretical and application frameworks for the conservation of this tiger sub-species in central Sumatra. The PPP model was developed to gain insight into tiger-preyhabitat relationships as well as the effect of human impacts on the persistence of tiger populations. The model addresses three main problems for the survival of the Sumatran tiger: tiger poaching, prey depletion, and habitat loss. The description of the PPP model serves as an in-depth study of existing literature and covers the most important factors of existing models for tiger conservation. Existing modelling approaches have been improved by the inclusion of finer description of individual-level traits and behaviours in the PPP model. The modelling approach allows a direct inter-relationships between individuals and their environment. The relationship between individual behaviours, intrinsic states, and external factors are simulated spatially explicitly in a bottom-up approach where the emergence of the population dynamics of tiger and prey can be observed under different scenarios. The integration between the PPP model and geographical information system (GIS) has provided a much more meaningful spatial data by revealing the mechanism of the response of individuals to the present land-use types. The relative importance of the parameters within the PPP model was tested using two modes of sensitivity analysis: The Morris Method and the traditional One-factor-at-a-time method. The results provided guidance for the application of reasonable sensitivity analysis during the development of individual-based models. The Morris Method suggested that the overall output of the PPP model showed a high sensitivity on the change of time required by a tigress to take care of cubs. The analysis also revealed that the number of dispersers was sensitive toward perceptual distance of individuals to detect the presence of prey. Comparison with a similar predator-prey models provided insight into the predator-prey relationship. The comparison also suggested that perceptual distance of the individual is important for any spatially explicit individual-based model involving predator-prey relationships. The parameterization of the individual perceptual distance of tigers was tested by using existing literature on prey consumption by tigers as a benchmark. The simulation results were within the range of scientific acceptance for the number of prey killed by a tiger. Thus, further use of the set of parameters for a tiger’s perceptual distance is less uncertain for the output of the PPP model. The effect of habitat quality and landscape configuration on the mortality and migration of prey were evaluated through the use of virtual habitats and landscapes. The findings suggested that a good habitat quality enables prey survival, increases the population available for predation by tigers. When a low-quality habitat is combined with a high-quality habitat, the number of migrating prey was high, reducing resources for tigers. This suggested that landscape composition should be considered when predicting population persistence of the Sumatran tiger. Optimal movement of two different prey resulted in a high density of prey in high-quality habitat, providing a concentration of prey in a tiger’s habitat, but resulted in a lower tiger predation rate than random movement and species specific movement. The PPP model has been applied to evaluate the effect of poaching, prey depletion, and their combination for the probability of extinction of a tiger population. The results from the evaluation showed that prey depletion, tiger poaching, and a combination of both, created a 100% probability of extinction within 20 years if the density and frequency of those threats at high rates. However, the duration of those threats in the system caused a 100% probability of extinction from tiger poaching. The results are able to contribute to optimize anti-poaching programs in future, to reduce significantly the probability of total extinction of Sumatran tiger. Furthermore, various landscape configurations have been tested against the probability and time of extinction for the Sumatran tiger population. The integration of spatial GIS-data in the model provides an insight into the relationship between tiger-prey-habitat. The results suggested that habitat quality surrounding a protected area plays an important role for the persistence of the Sumatran tiger population. This study also recommends agroforestry systems as reasonable land-use type in the vicinity of protected areas. They provide not only positive effects for tiger conservation purpose but they also appear as adaptable to the current land-use situation in Sumatra island. / Die vorliegende Dissertation beschreibt die Entwicklung des Panthera Populations Persistence (PPP) Modells, eines individuenbasierten Simulationsmodells für den Sumatra-Tiger (Panthera tigris sumatrae). Dieses stellt einen geeigneten theoretischen und anwendungsbezogenen Rahmen für den Schutz dieser Tiger-Unterart in Zentralsumatra bereit. Das PPP-Modell wurde entwickelt, um Einblicke in die Tiger-Beute-Habitat-Beziehungen zu gewinnen, sowie um den Effekt anthropogener Einflüsse auf den Fortbestand von Tigerpopulationen abzuschätzen. Dabei werden die drei Hauptprobleme für das Überleben des Sumatra-Tigers analysiert: die Wilderei, der Rückgang von Beutetieren und der Verlust von geeigneten Habitaten. Die Beschreibung des PPP-Modells gibt zunächst einen umfassenden Überblick zum aktuellen Wissensstand auf dem Gebiet des Tigerschutzes und integriert die wichtigsten Faktoren bereits existierender Modellansätze. Diese konnten durch die Einbeziehung einer detaillierten Beschreibung von individuellen Merkmalen und Verhalten verbessert werden. Das PPPModell stellt somit das Individuum in einen direkten Zusammenhang mit dessen Umwelt. Die Beziehung zwischen individuellem Verhalten, intrinsischen Merkmalen und externen Faktoren werden räumlich-explizit in einem bottom-up Ansatz simuliert. Damit kann sowohl die Populationsdynamik des Tigers als auch die seiner Beutetiere unter verschiedenen Annahmen beobachtet werden. Die Verknüpfung des PPP-Modells mit Geographischen Informationssystemen (GIS) bietet die Möglichkeit, die Reaktionsmechanismen der Individuen basierend auf der gegenwärtigen Landnutzungssituation zu simulieren und somit realitätsnahe räumliche Daten zu generieren. Die relative Bedeutung der Modell-Parameter auf die Simulationsergebnisse kann durch Sensitivitätsanalysen ermittelt werden. Hier wurden zwei verschiedene Ansätze verwendet: die Morris-Methode und die herkömmliche One-factor-at-a-time Methode. Der Vergleich beider methodischen Ansätze zeigte somit beispielhaft die Eignung unterschiedlicher Sensitivitätsanalysen für individuenbasierte Modelle auf. Die Morris-Methode zeigte, dass das Gesamtergebnis des PPP-Modells eine hohe Sensitivität gegenüber der Veränderung der Zeit aufweist, die ein Tigerweibchen braucht, um ihre Jungen aufzuziehen. Die Analyse zeigt auch, dass die Anzahl an abwandernden Tigern sensitiv gegenüber der IndividuellenWahrnehmungsdistanz von Beute ist. Der Vergleich mit einem ähnlichen Räuber-Beute-Modell lässt vermuten, dass diese Wahrnehmungsdistanz eines Individuums generell als ein entscheidender Faktor für Räuber-Beute-Beziehungen in räumlich-expliziten Individuenmodellen an- gesehen werden kann. Die Parametrisierung der IndividuellenWahrnehmungsdistanz des Tigers wurde so gewahlt, dass die damit ermittelten Simulationsergebnisse den Beutekonsum des Tigers, wie in der Literatur beschrieben, weitgehen widerspiegeln. Sie ist somit für die weitere Anwendung im PPP-Modell ausreichend gut beschrieben. Simulationsszenarien, welche verschiedene Habitatqualitäten sowie Landnutzungsmuster berücksichtigen, zeigen auch deren Bedeutung für die Mortalität und Migration der Beutetiere. Eine gute Habitatqualität hat eine geringe Mortalität der Beutetiere zur Folge, welche dann wiederum für den Tiger in ausreichender Zahl zur Verfügung stehen. Treten geringe Habitatqualitäten angrenzend an ein Habitat mit hoher Qualität auf, führte dies zu einer hohen Anzahl an abwandernden Beutetieren, womit sich die Ressourcen für den Tiger verringern. Die Landschaftsmerkmale sollten also bei der Vorhersage des Populationsfortbestandes des Sumatra-Tigers berücksichtigt werden. Die optimale Bewegung von zwei verschiedenen Beutetieren ergab eine hohe Beutedichte in einem Habitat mit hoher Qualität und stellte konzentriert Beute in einem Tigerhabitat bereit. Allerdings resultierte dies auch in einer geringeren Prädationsrate des Tigers, verglichen mit zufälligen oder artenspezifischen Bewegungen. Das PPP-Modell wurde angewandt, um die Auswirkungen von Wilderei, Beutetierrückgang sowie die Kombination beider Faktoren auf die Aussterbewahrscheinlichkeit einer Tigerpopulation zu bewerten. Die Ergebnisse zeigen, dass die genannten Faktoren eine 100-prozentige Aussterbewahrscheinlichkeit innerhalb von 20 Jahren zur Folge haben, wenn die Dichte und Häufigkeit dieser Bedrohungen hoch sind. Die Dauer dieser Bedrohungen im System verursachte allerdings eine 100-prozentige Aussterbewahrscheinlichkeit nur für die Wilderei von Tigern. Betrachtet man unabhängig von Dichte und Häufigkeit einzig die Dauer der Bedrohung, führt lediglich die Wilderei zum 100%-igen Aussterben. Diese Ergebnisse können maßgeblich dazu beitragen, zukünftig Schutzprogramme gegen die Wilderei zu optimieren, um das Aussterben des Sumatra-Tigers zu verhindern. DesWeiteren wurde der Einfluss von unterschiedlichen Landnutzungsmustern auf die Aussterbewahrscheinlichkeit und -zeit einer Sumatra-Tigerpopulation aufgezeigt. Die Integration von räumlichen GIS-Daten in das Modell ermöglichte einen Einblick in die Beziehungen zwischen Tiger, Beutetieren und Habitat. Die Ergebnisse zeigen, dass die Habitatqualität um Schutzgebiete herum eine wichtige Rolle für den Fortbestand der Population spielt. Die vorliegende Arbeit empfiehlt Agroforstsysteme als eine geeignete Landnutzungsform in der Nähe von Schutzgebieten, welche sowohl positive Effekte für den Tigerschutz bietet als auch mit den gegenwärtigen Landnutzungsmustern in Sumatra vereinbar erscheint.

No description available.

This thesis explores the methodology of state, and in particular, parameter estimation for time series datasets. Various approaches are investigated that are suitable for nonlinear models and non-Gaussian observations using state space models. The methodologies are applied to a dataset consisting of the historical lynx and hare populations, typically modeled by the Lotka- Volterra equations. With this model and the observed dataset, particle filtering and parameter estimation methods are implemented as a way to better predict the state of the system. Methods for parameter estimation considered include: maximum likelihood estimation, state augmented particle filtering, multiple iterative filtering and particle Markov chain Monte Carlo (PMCMC) methods. The specific advantages and disadvantages for each technique are discussed. However, in most cases, PMCMC is the preferred parameter estimation solution. It has the advantage over other approaches in that it can well approximate any posterior distribution from which inference can be made. / Master's thesis

The seasonal and diel feeding habits of the goosander, Mergus merganser, the red-breasted merganser, M. serrator, gulls (Larus canus, L. argentatus and L. fuscusj and terns, Sterna hirundo/paradisaea were studied at 64V05'N. Birds' activity patterns were influenced by the nocturnal spawning of the river lamprey (Lampetra fluviatilis). Food selection and food consumption by hand-raised mergansers together with aquarium studies of the diel activity patterns of their most important prey supplemented the field data. River lamprey dominated the diet of the goosander by weight and the sculpin Cottus gobio by number. The fish consumption of the goosanders was found to be about 12% of the available river lamprey biomass and about 17% of the sculpin biomass during the breeding season. In experimental situations the river lamprey was a low- pritority species compared with salmon , Salmo salar, brown trout, 53. trutta, and minnow, Phoxinus phoxinus, when presented to satiated birds of both Mergus species. When hungry, however, the birds caught the available prey irrespective of species, but they selected larger prey when two size classes were present. Experimental results were compared with field data on availability, consumption and the escape behaviour of the various fish species. The rivers emptying in the Bothnian Bay are regarded as important feeding areas for birds breeding along the coast. In early spring they get access to abundant and reliable food resources, e.g the river lamprey. Later on the three-spined stickleback, Gasterosteus aculeatus, supply both Mergus species with food in the river and also along the coast. From the middle of June potential food supply available to birds decreases in the rivers and becomes more abundant in the coastal area. / <p>S. 1-41: sammanfattning, s. 43-227: 7 uppsatser</p> / digitalisering@umu

No description available.

I investigated the effects ofhabitat loss and subsequent restoration on invertebrate community structure and ecosystem functioning in a mined peat bog in the North Island, New Zealand. In an experimental trial, the impact of peat bog habitat loss and isolation on the invertebrate community associated with Sporadanthus ferrugineus (Restionaceae) was investigated. Potted S. ferrugineus plants were exposed to invertebrates at various distances up to 800 m from an intact habitat (the presumed source population) over 18 weeks. Invertebrates rapidly colonised the experimental plants, with all major Orders and trophic groups present on Sc ferrugineus within 6 weeks. However. with increasing distance away from the undisturbed habitat, there was a significant decrease in total richness and abundance of invertebrates associated with the potted plants. Additional tests showed that even a moderate degree of isolation (i.e. greater than 400 m) from the intact habitat caused an almost complete failure of 'Batrachedra' sp. to colonise its host plant, at least in the short-term, The density of eggs and larvae, and the average larval size of 'Batrachedra' sp. (Lepidoptera: Coleophoridae) colonising Si ferrugineus plants, as well as the proportion of Si ferrugineus stems damaged by 'Batrachedra' sp. herbivory, all decreased logarithmically with increasing distance from the intact habitat. Surprisingly, though, the rate of recovery of the insect-plant interaction following experimental habitat restoration was remarkably rapid (i.e. between 3Y2 and 6 years). After just 6 years there was no significant difference in insect-plant interactions between the intact peat bog sites and any of the experimentally restored sites up to 800 m away. These results suggest that the degree of isolation from undisturbed habitat has a major impact on the rate and patterns of restoration recovery in the invertebrate community and that some insect-plant interactions can recover rapidly from habitat loss with restoration management. Restoration of mined peat bogs in northern New Zealand is initiated by establishing a native vegetation cover to minimize further peat degradation. The effects of various restoration techniques on litter decomposition, microbial community activity and beetle community composition were investigated within an experimental trial, These treatments included translocation ofpeat bog habitat (direct transfer of islands), milled peat islands with no seed and milled peat islands with seed, and were compared with an unrestored mined site and an undisturbed peat bog. In all the response variables measured, the undisturbed peat bog sites had significantly higher decomposition rates and microbial respiration rates, and significantly higher abundance and species richness of beetles than any of the restoration treatments. Inaddition, the technique used to restore mined peatlands had a significant effect on the beetle community composition and litter decomposition processes. Despite a rapid initial change in the beetle community following habitat translocation, the direct transfer islands were still the most similar in beetle species composition to the undisturbed peat bog. Microbial activity and decomposition rates were higher in the direct transfer and mined peat surface after 6 months. However, even after 12 months, decomposition rates in the restored habitats were still far from reaching the levels recorded in the undisturbed peat bog. The results suggest that beetle community structure and ecosystem processes such as decomposition and microbial activity rates may be able to recover faster with certain restoration techniques, such as direct transfer of intact habitat islands. Subsequently, I examined long-term beetle community reassembly on islands that had been restored by creating raised areas ofprocessed peat with the addition of Leptospermum scoparium seed. Monitoring of different-aged restored islands representing the full range of restoration ages (up to 6 years) available at the peat mine, indicated that as the peat islands became older and the vegetation structure became more complex, the abundance, species richness and composition of the beetle community became increasingly similar to the community in the undisturbed peat bog. Despite this, distinct differences between the intact peat bog and older restored peat islands still persisted, even after 6 years, particularly at an individual species level. However, it is predicted that within 12 years the restored peat islands will share 100% ofbeetle species in common with the undisturbed peat bog. Taken together, these results indicate that restoration is effective in initiating the recovery of beetle assemblages and ecosystem processes (such as litter decomposition and microbial community activity) in cut-over peat bogs. However, it is estimated to take at least 12 years before pre-mining communities and functions are attained, and ongoing monitoring to develop an understanding of the longer-term dynamics of such ecosystems and processes is clearly required.