Predation by olfactory meso-predators has a large impact on avian nest success, particularly for ground-nesting waterfowl. Olfactory predators rely on odors to locate their prey. Weather conditions (e.g. wind speed, humidity, and temperature), vegetation, and landscape features affect the dissipation rate of odors and could affect the foraging efficiency of olfactory predators. I conducted 2 studies to determine if weather and landscape impact predator foraging ability and behavior: a predator survey study and an artificial nest study. The objective of the predator survey was to investigate how landscape and weather conditions interact to influence the distribution of olfactory meso-predators [e.g. red foxes (Vulpes vulpes), skunks (Mephitis mephitis), and raccoons (Procyon lotor)] in their nightly foraging on the dike. Specifically, I examined how wind speed, wind orientation, temperature, and humidity affect the distribution, number, and species of olfactory meso-predators foraging on the Arthur V. Watkins Dike at Willard Bay State Park and Reservoir. The objective of the artificial nest study was to determine if weather, vegetation, or nest location relative to a large-scale surface feature have an effect on survival of artificial ground-nests in an area dominated by olfactory meso-predators. Artificial nests were placed on the dike throughout the summer of 2009. Spotlighting surveys for predators were conducted from August 2008 to August 2009. I found that section of the dike, time since study initiation, terrain type on the dike, wind speed, and vegetation height during daylight hours affected nest survival. The results indicated that predators formed olfactory search images in that nest survival decreased over the summer, while predator populations remained constant. I observed foxes, skunks, and raccoons while spotlighting for predators. After accounting for time, wind speed and direction were significant predictors of predators' nightly foraging activity with most predators observed when wind speeds were 2 to 4 m/s and winds were blowing from the northwest. Overall the model accounted for 75% of the nightly variation in predator numbers. Additionally, wind speed and direction impacted where predators were foraging. There were interspecific differences among predators in their responses to wind speed with raccoons being observed more than skunks and foxes when the wind was calm and blowing from the south. The results of the spotlighting data indicate that wind speed and direction have a strong effect on foraging activity. Overall, I concluded that wind speed affects predator foraging ability and behavior.

The influence of primary compounds (energy, protein, minerals, and vitamins) in animal nutrition and foraging behavior has been studied for years. The roles of secondary compounds (terpenes, alkaloids, and phenolic compounds) are equally important, yet they have been ignored until recently. Where secondary compounds were once considered toxic by-products of plant metabolism, we now know they are actively involved in plant and animal behavior, health, and productivity. Though often appreciated historically for their negative impacts on food intake and animal production, we are becoming increasingly aware of their beneficial roles in the health of plants, animals, and humans. When animals can ingest an array of plants that contain different kinds and amounts of secondary compounds, they can mix different foods in their diet to better use both primary and secondary compounds, enhancing their health and production, as well as economic and ecological characteristics of landscapes. Endophyte-infected tall fescue contains the alkaloids perlolidine, perloline, ergotamine, and ergovaline, which are all steroidal or protein-like in nature. Tannins and saponins have a high affinity for binding proteins and lipids in the gastro-intestinal tract of animals, and facilitating their excretion from the body. These findings suggest animals may increase their use of forages with alkaloids when they are also provided forages that contain tannins or saponins. The sequence in which forages with different secondary compounds are ingested may influence any potential interactions because different compounds have different residence times in the gastrointestinal tract. I tested the hypothesis that cattle and sheep foraging behavior is influenced by eating different combinations of forages containing secondary compounds in different sequences. In pen and pasture trials, I showed that 1) cattle grazing pastures of endophyte-infected tall fescue (TF) grazed more often on TF when first allowed to graze legumes containing either tannins or saponins, and they grazed TF much more when they first grazed legumes as opposed to the reverse sequence; 2) sheep fed basal diets high in the alkaloid ergotamine d tartrate (EDT) ate more when supplemented with food containing either tannins or saponins, but in contrast to the trials on pasture with cattle, their behavior was not dramatically influenced by sequence; 3) cattle fed fresh cut endophyte-infected tall fescue were not influenced by the addition of tannin to their drinking water, as tannins limited both water and forage intake; 4) sheep fed food containing EDT ate more when supplemented with food containing tannins or when given a choice of foods containing tannins or saponins, than sheep supplemented with food containing saponins or no additional secondary compound. Results from these studies suggest that secondary compounds interact with one another to influence foraging behavior, and that sequence of food ingestion can be influential when animals graze on pastures.

Au cours de cette thèse, des outils théoriques (simulation) et expérimentaux (tracking) ont été développés afin d’établir les mécanismes sous-jacents aux phénomènes de choix collectifs d’exploitation de sources de nourriture et à la gestion collective des stocks de nourriture constitués au sein d’une colonie de fourmis. Le premier volet de mon travail révèle l’influence de caractéristiques physiques (telles que l’angularité et la topographie) des pistes entre le nid et les sources de nourritures, sur les comportements des fourmis au niveau individuel et leur résultante au niveau collectif en termes d’exploitation de ressources alimentaires. Le second volet de ma thèse est consacré aux dynamiques et régulations à l’œuvre au sein des phénomènes de dissémination collective et d’accumulation de nourriture au niveau intranidal. Les réseaux de distributions et d’échanges, reconstitués chez différentes espèces grâce à un marquage individuel et dans différents contextes montrent des similarités inattendues, notamment en termes d’hétérogénéités interindividuelles et de robustesse face à la variabilité des ressources. / During my PhD I have developed theoretical (simulation) and experimental (tracking) tools to investigate the mechanisms underlying the collective food source choice as well as collective food management at the intranidal level in ants. A first part of my thesis reveals the influence of the physical (and in particular the angularity and the topography) properties of the trail to the food source on the ant behavior at the individual level and its resulting food source choice/exploitation at the collective level. The second part of my thesis is dedicated to the dynamics of the food flow entering the nest, its regulation and the food dissemination/accumulation processes at the intranidal level through trophallaxis (mouth-to-mouth fluid exchange). The networks of food distributions and exchanges were built for two ant species in various experimental conditions. Interestingly, all theses networks show several similar properties, such as a high degree of heterogeneity in the individual participation to the food exchange activity as well as a robustness facing environmental variabilities.

Herbivory is one major force accelerating aspen decline in North America, but it is unclear why herbivores prefer certain aspen stands over others, or over other plant species in the understory. In this dissertation, I determined the influence of nutrients and plant secondary compounds (PSC), physiological state, chemical composition, and prior experience on aspen preference by sheep in controlled pen experiments. In addition, I explored the relationship between herbivory, regeneration, recruitment, and other landscape elements for specific aspen stands within Wolf Creek Ranch in northern Utah using biomass and chemical composition of the understory and chemical defenses of juvenile aspen trees (i.e., the foodscape). Aspen intake was enhanced when lamb diets contained a high crude protein to energy ratio or when the basal diet contained a low density of energy. Intake was depressed as concentrations of PG (phenolic glycosides) increased in aspen leaves or when lambs were fed a high energy to protein ratio. The effects of nutrients on aspen intake were greater when phenolic glycosides in aspen were present at low concentrations. However, when given a choice between aspen leaves of high or low PG content, lamb preference depended more on aspen nutrient and mineral availability, or on prior diet, than on defense chemistry. On the landscape, I found that stands at low elevations with low abundance of nutrients in the understory are more likely to experience less regeneration and recruitment than those growing within nutrient-rich sites. Aspen browsing was negatively correlated with PG content in aspen stands, and elk presence (measured via fecal pellets) was negatively correlated with abundance of understory protein. In conclusion, aspen herbivory appears to be controlled by the interplay between types and amounts of nutrients offered by the landscape and the chemical composition of aspen stands. A clear assessment of these variables on the landscape, i.e., the foodscape, will aid in the development of novel management strategies aimed at providing nutrients (e.g., through supplements, introduced forages) at strategic locations in order to reduce aspen herbivory within at-risk aspen stands.

<p>The effect of water temperature prey capture success and foraging behavior of drift-feeding juvenile brown trout (<em>Salmo trutta</em>) was examined in a laboratory stream. Water temperature treatments were 5.7, 6.7, 8.0, 10, 12 and 14°C. Five wild brown trout, age 1+ and collected by electrofishing from a stream in Western Sweden, were used in the experiments. There was a significant effect of water temperature on both prey capture probability and the percentage of time spent resting on the substrate while drift-feeding. At low water temperatures the fish suffered a reduced prey capture capability and spent more time resting on the substrate in between the excursions to capture drifting prey. Temperature did not significantly affect the amount of time fish spent foraging holding a station in the current versus active searching. Significant positive correlations were found between holding a station and prey capture probability at four out of the six different water temperature treatments.</p><p> </p>

A ubiquitous problem for all foragers is the trade-off between acquiring food energy while simultaneously avoiding the risk of predation. In central montane Alberta I modelled how ungulate forage changes with succession within cutblocks and the implications for forage availability to ungulates under current harvest regimes. Because cutblocks are discrete patches in space, I developed a dynamic state variable model for an ungulate to explore under what conditions an individual forager could (1) behaviourally avoid predation within a patch through inactivity, (2) overcome patch isolation when confronted with predation during transit between patches, and (3) alter patch use across a home range to optimize fitness. The model includes the requirement to process forage into energy through rumination behaviour that constrains foraging, and compares outcomes under a time-minimizing (sigmoid) and energy maximizing (linear) fitness functions. When an ungulate is in high energetic state, inactivity provides an effective behavioural refuge, or animals prioritize safety over energy gain, individuals avoid predation within patches reducing the need to move between patches. When energy acquisition is prioritized, individuals are at a low energetic state, or within patch anti-predator behaviours are ineffective, individuals move among patches to avoid predators in space, and configuration of the patches influences fitness. When model results were qualitatively compared to activity patterns and cutblock use of female, GPS-collared elk appear to follow a time minimizing strategy in their patch use across the home range and with their activity within riskier patches. I discuss the implications of these findings for the management of elk and cutblocks in west central Alberta.

Understanding the ways in which animals utilize space and obtain food are central themes in modern ecology. Formulating broad principles and elucidating the factors explaining such patterns are limited, however, by the availability of data from a broad range of species and systems. This problem especially true of snakes, a predator group about which even the most basic natural history data are often entirely lacking, even among abundant, widespread, and ecologically-important species. I studied the natural history and ecosystem role of one such species, the fer-de-lance (Bothrops asper) in lowland rainforest in Costa Rica. B. asper is a large, cryptic pitviper that is highly abundant in many Central American ecosystems and is strongly relevant to human health due to high incidence of snakebite, yet its biology under natural conditions is almost entirely undocumented. I used radiotelemetry to quantify home range, movement patterns, habitat usage, and foraging behavior. B. asper was found to have smaller home ranges and reduced movement patterns than similarly-sized temperate pitvipers, likely due to a greater reliance upon ambush foraging in patches of high prey density. Snakes also demonstrated strong selection for swamp habitat, which may reflect efforts to exploit frogs as a primary food source due to low availability of small mammals at the study site. I subsequently addressed the trophic status of this B. asper population using a supplemental-feeding experiment. In comparison to control snakes, individuals receiving supplemental food had smaller home ranges, shorter and less frequent movements, increased mass acquisition, and shifted to primarily forest rather than swamp habitat. These results support the suggestion that B. asper at the study site are strongly food-limited. Finally, I tested the hypothesis that fer-de-lance mediate local seed-predation rates by influencing habitat usage and foraging behavior of rodents. A series of behavioral experiments conflicted with many existing studies in failing to support this idea, as three rodent species demonstrated little snake avoidance, and none of likely ecological relevance. Collectively, this dissertation represents the first comprehensive field study of Bothrops asper and is among the first for any tropical snake, and suggests several avenues for future research.

All animals are adapted to the environmental conditions of the habitat they chose to live in. It was the aim of this PhD-project, to show which behavioral strategies are expressed as mechanisms to cope with the constraints, which contribute to the natural selection pressure acting on individuals. For this purpose, small mammals were exposed to different levels and types of predation risk while actively foraging. Individuals were either exposed to different predator types (airborne or ground) or combinations of both, or to indirect predators (nest predators). Risk was assumed to be distributed homogeneously, so changing the habitat or temporal adaptations where not regarded as potential options. Results show that wild-caught voles have strategic answers to this homogeneously distributed risk, which is perceived by tactile, olfactory or acoustic cues. Thus, they do not have to know an absolut quality (e.g., in terms of food provisioning and risk levels of all possible habitats), but they can adapt their behavior to the actual circumstances. Deriving risk uniform levels from cues and adjusting activity levels to the perceived risk is an option to deal with predators of the same size or with unforeseeable attack rates. Experiments showed that as long as there are no safe places or times, it is best to reduce activity and behave as inconspicuous as possible as long as the costs of missed opportunities do not exceed the benefits of a higher survival probability. Test showed that these costs apparently grow faster for males than for females, especially in times of inactivity. This is supported by strong predatory pressure on the most active groups of rodents (young males, sexually active or dispersers) leading to extremely female-biased operative sex ratios in natural populations. Other groups of animals, those with parental duties such as nest guarding, for example, have to deal with the actual risk in their habitat as well. Strategies to indirect predation pressure were tested by using bank vole mothers, confronted with a nest predator that posed no actual threat to themselves but to their young (Sorex araneus). They reduced travelling and concentrated their effort in the presence of shrews, independent of the different nutritional provisioning of food by varying resource levels due to the different seasons. Additionally, they exhibited nest-guarding strategies by not foraging in the vicinity of the nest site in order to reduce conspicuous scent marks. The repetition of the experiment in summer and autumn showed that changing environmental constraints can have a severe impact on results of outdoor studies. In our case, changing resource levels changed the type of interaction between the two species. The experiments show that it is important to analyze decision making and optimality models on an individual level, and, when that is not possible (maybe because of the constraints of field work), groups of animals should be classified by using the least common denominator that can be identified (such as sex, age, origin or kinship). This will control for the effects of the sex or stage of life history or the individual´s reproductive and nutritional status on decision making and will narrow the wide behavioral variability associated with the complex term of optimality. / Das Verhalten von Tieren ist das Ergebnis eines kontinuierlichen Anpassungsprozesses im Laufe der Evolution einer Art und damit der Veränderung der Umgebung in der es lebt und der Interaktion mit anderen Arten. Dies wird besonders deutlich im Verhalten von potentiellen Beutetieren, ihre Strategien beinhalten meist ein möglichst unauffälliges Verhalten im Zusammenspiel mit reduzierter Bewegung und möglichst guter Tarnung. Dementgegen stehen essentielle Bedürfnisse, wie zum Beispiel die Nahrungssuche, die Verteidigung von Ressourcen (zum Beispiel Territorien, Futterstellen) und die Suche nach Paarungspartnern. Beutetiere leben also in einem Spannungsfeld indem sie Ihr Verhalten optimieren müssen. Hierbei stehen die Ernährung, erfolgreiche Verpaarung und andere Chancen auf der einen Seite, die Vermeidung von Begegnungen mit Prädatoren auf der anderen. Vor allem Kleinsäuger sind häufig als Beutetiere mit einer Vielzahl von Prädatoren aus der Luft und auf dem Boden konfrontiert. Sie müssen für die verschiedenen Bedrohungen adaptive Verhaltensanpassungen bereit haben und in der Lage sein, auf die optischen, olfaktorischen oder akustischen Signale, die die Gefahr durch Prädatoren anzeigen, mit plastischen Verhaltensmustern zu reagieren. Die vorliegende Dissertation beschäftigt sich mit bisher als Konstanten behandelten Faktoren und untersucht anhand von Verhaltensexperimenten mit wilden Wühlmäusen (Microtus arvalis) folgende Fragestellungen: - Wie verhalten sich Tiere, die einer homogenen Risikoverteilung ausgesetzt sind, zum Beispiel weil ihr Prädator genauso gross ist wie sie, im gleichen Habitat lebt und es keinen sicheren Ort gibt? - Mit welchen Anpassungen reagieren Tiere, wenn sie gleichzeitig verschiedenen Prädatoren ausgesetzt sind? - Wie unterscheiden sich die Nahrungssuchstrategien von Männchen und Weibchen? - Wie verhalten sich laktierende Weibchen, die einer permanenten, indirekten Gefahr, z.B. durch einen Nestprädator ausgesetzt sind? Die Ergebnisse der verschiedenen Versuche in künstlichen Arenen und Aussengehegen zeigen, dass die Mäuse in der Lage sind, adaptive Verhaltensanpassung an homogenes Risiko und verschiedene Prädationstypen abzurufen. So sind sie in der Lage, Luft- von Bodenprädatoren zu unterscheiden und jeweils das Verhalten zu zeigen, dass die größtmögliche Sicherheit mit sich bringt. Die simultane Kombination von verschiedenen Prädatoren bewirkt hierbei additive Effekte. Gibt es keine Auswahl zwischen Habitaten, sondern nur unterschiedliche homogene Risikolevel, reagieren sie auf steigendes Risiko immer mit verminderter Aktivität und konzentrieren ihre Nahrungssuche auf weniger Futterstellen, beuten diese dafür jedoch länger aus. Die Wertigkeit von Futterstellen und alternativen Optionen verändert sich also mit dem Risikolevel. Ähnliches zeigt sich auch in den unterschiedlichen Futtersuchstrategien von Männchen und Weibchen. Die untersuchte Art ist polygyn und multivoltin, dementsprechend verbinden die Männchen mit ihrer Nahrungssuche Aktivitäten wie die Suche nach Paarungspartnern und unterscheiden sich die Aktivitätsmuster zwischen Männchen und Weibchen. Zusätzlich zeigen die Ergebnisse, das laktierende Weibchen in der Lage sind, das Risiko für sich und für Ihre Jungen abzuschätzen, wenn sie mit einem Nestprädator (Sorex araneus) konfrontiert werden. Für die Interaktion zwischen diesen beiden Arten ist jedoch die Saison (und damit die Ressourcenlage), in der sie sich begegnen, von entscheidender Bedeutung. Wühlmäuse reagieren mit entsprechenden Verhaltensanpassungen zum Schutz des Nestes um die Überlebenschancen ihrer letzten Würfe im Herbst zu erhöhen. Die vorliegende Arbeit konnte grundsätzliche Probleme der antiprädatorischen Verhaltensanpassung von Beutetieren klären und wichtige Faktoren der Entscheidungsfindung unter Prädationsdruck analysieren. Sie zeigt, dass Tiere das Risiko in ihrer Umgebung nicht unbedingt über direkt Signale wahrnehmen, sondern ihre Verhaltensstrategien einem empfundenen Gesamtrisikolevel anpassen. Dies ermöglicht ihnen, adaptive Strategien zu verfolgen, auch wenn sie keine Auswahl an sicheren Habitaten haben. Sie zeigt auch die unterschiedliche Wahrnehmung von Risiken durch Männchen und Weibchen, die durch die unterschiedlichen mit der Aktivität zusätzlich wahrgenommenen Chancen verknüpft zu sein scheint. Zusätzlich wurde der Einfluss des reproduktiven Status (z.B. laktierend), sowie der Ressourcenlage (z. B. je nach Saison) nachgewiesen.

Planktivorous fish affect the species composition, size and morphology of their zooplankton prey by size selective predation. Remains of cladocerans are deposited in the lake sediments and this microfossil community hence reflects the historical changes in the planktivore fish community. Most earlier work on quantitatively inferring past planktivory deals with ephippia (resting eggs) of Daphnia, however these remains are very scarce in northern Swedish lakes. In these lakes, often dominated by fish that are not obligate planktivores, such as perch (Perca fluviatilis), the individual size to a large degree determines the type of resource consumed. Therefore, in these lakes, the size distribution as well as the density of the fish community determines the overall planktivoryThis thesis deals with evaluating the use of Bosmina remains in the sediment as indicators of past planktivory. I developed a transfer function based on Bosmina remains to be used in oligotrophic lakes with omnivorous species to reconstruct past planktivory. I test the transfer function in a lake where the history is to a large extent known. Furthermore, in this thesis a novel quantitative measure of planktivore pressure, the PCC (Planktivore Community Capacity) was developed. It was also possible to compare the morphology measurements of carapace length and mucro length of contemporary Bosmina sampled in the lake, with the same measurements on microfossils of Bosmina sedimented during the corresponding time period. By using the resurrection approach this thesis also evaluates the use of Ceriodaphnia clones as indicators of micro-evolutionary responses to past planktivory.From the studies in this thesis I can draw some major conclusions; the morphological measurements carapace length and mucro length of Bosmina can be used to infer past planktivory in northern Swedish lakes. PCC has a higher explanatory power than CPUE in these lakes dominated by omnivorous fish. The morphology of the contemporary Bosmina corresponds well with the remains found in the sediment dating from the same time period. Both the contemporary and microfossil Bosmina changes in body size characteristics was in turn strongly related to PPC. Finally, the hatching of Ceriodaphnia ephippia is promising for studying past planktivory, both because of the high viability of the ephippia and the relationship between eye area and inferred PCC.

Neighbouring plants can interact strongly, competing for resources including light, water, animal mutualists, and local germination sites. From an evolutionary perspective, this implies that a plant’s best resource acquisition strategy will usually depend on the traits of its neighbours, and for plants in particular, neighbours are often genealogical relatives. Here, I use a combination of theory and experiments to expose some important consequences of social interactions among plants. The first model analyzes selection on traits used to attract pollinators, showing that competitive interactions (in the absence of local relatedness) can select for exaggerated secondary sexual characters. To complement this model, I performed experiments that confirm the mechanisms by which adaptive pollinator foraging naturally leads to interactions among plants. The observed foraging behaviour (of bumble bees) also provides unique evidence for ‘Bayesian foraging’, a sophisticated type of resource assessment that depends on prior experience in a particular environment. A second model considers how selection on the sex allocation of cosexual, animal-dispersed plants leads to competition and cooperation over local germination sites, sometimes leading to the origin of gender dimorphism. The model reveals novel ecological contexts in which disruptive selection on sex allocation can arise, and in general, illustrates how selection for cooperation can facilitate or inhibit evolutionary diversification. In the models considered here, cooperation is indiscriminant, but plants might also assess the relatedness of neighbours and cooperate with kin over non-kin. In the final chapter, I present experimental evidence that is consistent with preferential cooperation over soil resources among sibling plants. This study is the first to link a potentially cooperative resource allocation strategy with an increase in the mean fitness of related plants.