Northern environments are often characterised by large seasonal and annual fluctuations in food abundance. In this thesis, I examined how arctic foxes (</i>Alopex lagopus</i>) used seasonally superabundant foods (geese and their eggs) and how access to these foods influenced population dynamics of arctic foxes. I addressed this against a backdrop of variation in lemming and vole abundance (small mammals hereafter) the main foods of arctic foxes throughout most of their range. Field work was done at the large goose colony at Karrak Lake and surrounding areas in the Queen Maud Gulf Bird Sanctuary in Nunavut, Canada, in the spring and summers of 2000 to 2004. <p> Behavioural observations of individually-marked arctic foxes showed that they took and cached 2,000-3,000 eggs per fox each year and that the rate at which they took eggs was largely unrelated to individual attributes of foxes (e.g. sex, size, and breeding status) and nesting distribution of geese. Further, the rate at which foxes took eggs varied considerably within individuals in that foxes were efficient at taking eggs at times and inefficient at other times. This may have resulted from foxes switching between foraging actively and taking eggs opportunistically while performing other demands such as territorial behaviours. <p>Comparison of stable isotope ratios (13C and 15N) of fox tissues and those of their foods showed that the contribution of cached eggs to arctic fox diets was inversely related to collared lemming (<i>Dicrostonyx torquatus</i>) abundance. In fact, the contribution of cached eggs to overall fox diets increased from <28% in years when collared lemmings were abundant to 30-74% in years when collared lemmings were scarce. Furthermore, arctic foxes used cached eggs well into the following spring (almost 1 year after eggs were acquired) a pattern which differs from that of carnivores generally storing foods for only a few days before consumption. <p>A field-study of experimental caches showed that survival rate of these caches was related to age of cache sites in the first year of the study (e.g. 0.80 and 0.56 per 18-day period for caches from new and 1 month old cache sites, respectively) and departure by geese after hatch in the second year of the study (e.g. 0.98 and 0.74 per 18-day period during and after goose nesting, respectively). Food abundance and deterioration of cache sites (e.g. loss of soil cover and partial exposure of caches) were, thus, important factors affecting cache loss at Karrak Lake. Further, annual variation in the importance of these factors suggests that strategies to prevent cache loss are not fixed in time but vary with existing conditions. Evolution of caching behaviours by arctic foxes may, thus, have been shaped by multiple selective pressures. <p>Comparisons of reproductive output and abundance of arctic foxes inside and outside the goose colony at Karrak Lake showed that (i) breeding density and fox abundance were 2-3 times higher inside the colony than they were outside the colony and (ii) litter size, breeding density, and annual variation in fox abundance followed that of small mammal abundance. Small mammal abundance was, thus, the main governor of population dynamics of arctic foxes whereas geese and their eggs elevated fox abundance and breeding density above that which small mammals could support. These results highlight both the influence of seasonal and annual variation on population dynamics of consumers and the linkage between arctic environments and wintering areas by geese thousands of kilometres to the south.

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.

The contagious nature of the black-tailed prairie dog (Cynomys ludovicianus) jump-yip display may provide information to signalers and receivers regarding the vigilance of neighbours. Videotaping jump-yip bouts and the behaviour of both bout initiators and respondents within those bouts provided evidence that: 1) individuals became vigilant immediately following jump-yip production, but exhibited minimal changes in their immediate post-jump-yip behaviour with changes in the characteristics of the preceding bout, 2) bout initiators spent more time actively foraging and exhibited vigilance behaviours less frequently with greater levels of response in the preceding jump-yip bout, 3) respondents spent more time actively foraging and less time vigilant following bouts with greater response. These results suggest that black-tailed prairie dogs base behavioural decisions at least in part on the characteristics of their jump-yip bouts and thus the jump yip display may provide information about the vigilance of group members. / October 2007

Animals in zoos live in more barren environments than their conspecifics in nature. In nature, elephants spend more than 75 % of the day foraging and have been observed lying down two hours during night. Stereotypies are common in many different species in zoos. Stereotypic behaviors may constitute up to 50 % of the daily activity budget of zoo elephants. Modern zoos try to reduce stereotypic behaviors by means of environmental enrichment. Three elephant cows at Kolmårdens Djurpark were observed during night to map their nocturnal behaviors and see how they changed with feeding enrichment. The elephants were lying down on their sides 25 % of the time observed. With feeding enrichment offered in the morning, the stereotypies was reduced in one individual from 43 % to 13 % (P&lt;0.01) and the foraging was increased in two individuals from 45 % to 80 % (P&lt;0.01) and from 45 % to 64 % (P&lt;0.05). Increased food access increased foraging behavior and as environmental enrichment was able to reduce the stereotypic behaviors in these Asian elephant.

Northern environments are often characterised by large seasonal and annual fluctuations in food abundance. In this thesis, I examined how arctic foxes (</i>Alopex lagopus</i>) used seasonally superabundant foods (geese and their eggs) and how access to these foods influenced population dynamics of arctic foxes. I addressed this against a backdrop of variation in lemming and vole abundance (small mammals hereafter) the main foods of arctic foxes throughout most of their range. Field work was done at the large goose colony at Karrak Lake and surrounding areas in the Queen Maud Gulf Bird Sanctuary in Nunavut, Canada, in the spring and summers of 2000 to 2004. <p> Behavioural observations of individually-marked arctic foxes showed that they took and cached 2,000-3,000 eggs per fox each year and that the rate at which they took eggs was largely unrelated to individual attributes of foxes (e.g. sex, size, and breeding status) and nesting distribution of geese. Further, the rate at which foxes took eggs varied considerably within individuals in that foxes were efficient at taking eggs at times and inefficient at other times. This may have resulted from foxes switching between foraging actively and taking eggs opportunistically while performing other demands such as territorial behaviours. <p>Comparison of stable isotope ratios (13C and 15N) of fox tissues and those of their foods showed that the contribution of cached eggs to arctic fox diets was inversely related to collared lemming (<i>Dicrostonyx torquatus</i>) abundance. In fact, the contribution of cached eggs to overall fox diets increased from <28% in years when collared lemmings were abundant to 30-74% in years when collared lemmings were scarce. Furthermore, arctic foxes used cached eggs well into the following spring (almost 1 year after eggs were acquired) a pattern which differs from that of carnivores generally storing foods for only a few days before consumption. <p>A field-study of experimental caches showed that survival rate of these caches was related to age of cache sites in the first year of the study (e.g. 0.80 and 0.56 per 18-day period for caches from new and 1 month old cache sites, respectively) and departure by geese after hatch in the second year of the study (e.g. 0.98 and 0.74 per 18-day period during and after goose nesting, respectively). Food abundance and deterioration of cache sites (e.g. loss of soil cover and partial exposure of caches) were, thus, important factors affecting cache loss at Karrak Lake. Further, annual variation in the importance of these factors suggests that strategies to prevent cache loss are not fixed in time but vary with existing conditions. Evolution of caching behaviours by arctic foxes may, thus, have been shaped by multiple selective pressures. <p>Comparisons of reproductive output and abundance of arctic foxes inside and outside the goose colony at Karrak Lake showed that (i) breeding density and fox abundance were 2-3 times higher inside the colony than they were outside the colony and (ii) litter size, breeding density, and annual variation in fox abundance followed that of small mammal abundance. Small mammal abundance was, thus, the main governor of population dynamics of arctic foxes whereas geese and their eggs elevated fox abundance and breeding density above that which small mammals could support. These results highlight both the influence of seasonal and annual variation on population dynamics of consumers and the linkage between arctic environments and wintering areas by geese thousands of kilometres to the south.

Despite decades of protection, the endangered North Atlantic right whale (Eubalaena glacialis) has failed to recover, primarily due to interactions with fishing gear and ship strikes. Right whales range along the U.S. east coast, foraging year round in the Gulf of Maine while a subset of the population travels to the South Atlantic Bight each year to calve. The habitat requirements of the right whale are poorly understood. I investigated the relationship between the distribution of right whales and physical oceanographic conditions in an effort to create predictive models of essential right whale habitats. Additionally, the distribution of right and humpback whales (Megaptera novaeangliae) relative to fixed fishing gear was examined to assess spatio-temporal overlap. Habitat preferences were assessed using aerial survey data of whale locations and a range of topological and satellite derived physical parameters including bathymetry, sediment type, sea surface temperature, thermal gradients and surface roughness. A suite of non-parametric quantitative techniques including Mantel tests, log likelihood functions, Generalized Additive Models, Spearman Rank Correlations and the Williamson's spatial overlap index were used to assess relationships between whales and habitat variables. Our findings indicate that suitable calving habitat along the east coast may extend much farther to the north than is currently recognized. Our model correctly identified several well documented current and historic calving grounds in the eastern Atlantic but failed to fully identify a heavily used calving area off Argentina, which is characterized by lower surface water temperatures than the other calving regions. In the Gulf of Maine, right whale distribution was correlated primarily with sea surface temperature, sediment type and bathymetry. Predictive models offered insights into right whale habitat preferences for foraging but failed to wholly capture the physical factors underlying right whale distribution. I found the relative density of right and humpback whales and fixed fishing gear in the Gulf of Maine to be negatively correlated in most seasons and areas. These findings demonstrate that the regular co-occurrence of high densities of whales and gear is not a prerequisite for entanglement. Prohibiting entangling lines in areas where whales are known to forage could substantively reduce entanglement. / Dissertation

<p>Abstract</p> <p>Risky foraging is a male reproductive tactic in most polygynous mammals. It is speculated to result from intense intra-sexual reproductive competition. Consequently this behavior has been speculated to increase a male's reproductive competitiveness. However, individual males may differ in their propensity to take foraging risks. </p> <p>We therefore conducted a study on crop raiding behavior (a risky foraging strategy) in African elephants from the greater Amboseli ecosystem, in southern Kenya. We specifically examined the population sizes, gender and patterns of raiding elephants and investigated the effect of crop-raiding and genetic heterozygosity on male body size. We also examined the influence of age and genetic relatedness on observed patterns of association. Finally, we examined the role of life history milestones, association patterns and social structure on the acquisition of crop raiding behavior among wild free ranging male African elephants. With regard to the influence of association patterns on crop raiding behavior, we were specifically interested in understanding the mechanisms by which social learning might occur among male elephants.</p> <p>Our results showed that 241elephants from different populations in the ecosystem converged to raid farms. Approximately 35% of raiders were from Amboseli National Park, and the rest were other populations in the ecosystem. We observed only post-pubertal males but not females to raid. About one third of post-pubertal males from the Amboseli population were raiders. We found evidence of habitual raiding by some individuals. Crop raiding predicted post-pubertal male size, with raiders being larger than non-raiders. This result suggests that taking risks pays off for males. Our results also showed that other variables known to influence growth like genetic heterozygosity had no effect on size-for-age in male elephants, because low-heterozygosity males were rare. The probability that an individual male is a crop raider was greater for older individuals than young males. The probability that a male is a raider was greater when his two closest associates were raiders versus when they were not raiders and when a male's second closest associate was older, versus when his second closest associate was of similar age or younger. These results suggest that increasing energetic demands associated with life history milestones and social learning play a significant role in the initiation of crop raiding behavior. Raiders did not cluster into separate social units from non-raiders, probably due to the nature of social learning exhibited by this species and due to the diffuse nature of male elephant social units.</p> <p>These results have implications for understanding the evolution of risky foraging behavior in males, and for understanding the role of kin selection, dominance hierarchies and social learning in male elephant social systems. Results also have implications for understanding the spread of adaptive complex behavior in natural populations.</p> / Dissertation

<p>Oceanographic features, such as fronts, eddies, and upwellings, provide important foraging areas for marine predators. These areas serve as important "hotspots" of marine life, by aggregating weakly swimming lower and mid-trophic level species which, in turn, attract foraging predators. Despite the importance of these dynamic features, we lack a comprehensive understanding of how they create foraging habitat for seabirds and other marine predators. In the first part of this dissertation, I review current knowledge of how seabirds use oceanographic features with an emphasis on developing a more mechanistic understanding of these features, and identify important considerations for future studies. I use the findings of this review to inform two field research projects in the Bay of Fundy, Canada and Onslow Bay, North Carolina. In these two projects, I examined seabird abundance and distribution in relation to oceanographic features that occur at different spatial and temporal scales. In the first project, I examined foraging habitat of red-necked phalaropes (<italic>Phalaropus lobatus</italic>) in relation fine-scale tidal forcing near the Brier Island ledges in the Bay of Fundy. This research demonstrated the importance of biophysical interactions in creating phalarope habitat, and characterized red-necked phalarope habitat in both space and time. In Onslow Bay, I investigated the effects of Gulf Stream fronts and eddies on the abundance and distribution of seabirds using both remotely sensed and in situ data. I used fisheries acoustics surveys to investigate prey distribution within Gulf Stream frontal eddies. I then developed habitat models for the six most commonly sighted species or species groups (Cory's shearwaters, <italic>Calonectris diomedea</italic>; greater shearwaters (<italic>Puffinus gravis</italic>; Wilson's storm petrel, <italic>Oceanites oceanicus</italic>; Audubon's shearwaters, <italic>Puffinus lherminieri</italic>; black-capped petrels, <italic>Pterodrama hasitata</italic>; and red and red-necked phalaropes, grouped together as <italic>Phalaropus</italic> spp.) using multivariate modeling techniques. Gulf Stream frontal eddies influenced the abundance and distribution of seabirds in Onslow Bay, although frontal features were not as important in predicting seabird habitat as demonstrated in previous studies in the South Atlantic Bight. Prey availability in Gulf Stream frontal eddies was highest in eddy cold core regions, particularly in those regions close to the Gulf Stream. Taken together, the results of my dissertation: underscore the importance of conducting standardized surveys to assess dynamic environmental variables; demonstrate the use of multivariate methods to examine seabird foraging in relation to oceanographic features; emphasize the need to evaluate both prey distributions and physical regimes within oceanographic features at depth; and highlight the importance of temporal aspects of oceanographic features, such as the persistence and age of the features, when assessing the role that these features play in creating seabird foraging habitat.</p> / Dissertation

The northern fur seal (NFS) population on the Pribilof Islands (PI) is currently declining while the population on the Commander Islands (CI which includes Bering and Medny Islands) is stable. The reasons for the different population trajectories remain unknown. Comparing differences in behavioral ecology and predation pressure between these two populations could provide an explanation. This study examined lactating NFS female behavior to determine: 1) summer foraging patterns (trip duration, trip direction, dive depth) of animals from two nearby rookeries on Bering Island, 2) winter migration from Medny and Bering Islands relative to patterns of ocean productivity, and 3) the potential impact of killer whale predation on population dynamics. Data were collected from 2003 to 2010 using visual observations and telemetry. Twenty-one satellite transmitters, 29 time-depth recorders and 17 geolocation recorders were deployed. Shore-based observations of killer whale predation and photo-identification were conducted near the CI rookeries in 19992010. During lactation, both mean foraging trip duration and mean maximum diving depth (3.4 plus/minus 1.3 days and 17.7 plus/minus 6.8 m, respectively) for NFS adult females (n = 28) did not significantly change among years. Although foraging areas of NFS from the two rookeries on Bering Island overlapped, the mean direction of travel from Severo-Zapadnoe rookery was significantly (p<0.01) different compared with Severnoe rookery. The foraging patterns suggested that these females had a reliable food source that did not change despite potential environmental changes or the effects of fisheries. During their winter migration, NFS females from the CI traveled to the Transition Zone Chlorophyll Front (32° N-42° N) in the North Pacific Ocean. Their winter migration routes and the location of overwinter foraging areas were positively correlated with high ocean productivity (near surface chlorophyll a concentration). Over 82 percent (n=17) of these females spent 38 months near the eastern coast of Hokkaido, Japan and followed the coastal high productivity areas on their way back to the CI. Transient killer whales in groups of 2-12 individuals were repeatedly observed preying mostly on NFS males during the summer. The simulation model showed little impact on population dynamics as long as male fur seals were the primary prey. However, if the number of killer whales increased or they changed their diet to include females and pups, then the NFS population on the CI could decline. The winter migration of NFS from CI and PI are similar. Lactating NFS from the PI exhibit greater summer foraging effort (longer average trip duration and bout duration; greater number of deep dives) compared with females from the CI.

In 2002, a proposal was submitted to divert water from the confluence of the Guadalupe and San Antonio Rivers to San Antonio, Texas. To investigate the potential impacts of diverting water from the Guadalupe Estuary, my research focused on the foraging ecology of the crane and population ecology of the blue crab, a documented crane food. During winters 2004-2005 and 2005-2006, I examined diets and optimal foraging patterns of territorial adult cranes at Aransas National Wildlife Refuge, Texas. To identify foods of greatest benefit to cranes, I used currencies of optimization to evaluate foraging gain, cost, and efficiency. Foraging benefit differed among foods, depending on the foraging currency used and resource targeted (e.g., energy). Wolfberry fruit, snails, and insects were consumed in the highest quantities, required the least foraging effort, and were generally associated with the highest foraging efficiency. Blue crabs and clams were important sources of protein and biomass. During September 2003-October 2005, I used novel artificial settlement substrates and modified methods of standard deployment to investigate the spatio-temporal patterns of blue crab terminal settlement and recruitment rates. Monitoring rates in shallow bay habitat and ponds of the interior salt-marsh revealed megalopal crabs were developmentally advanced when arriving to study sites and the extent to which young crabs infiltrated the salt-marsh increased with age. Such findings suggested sites represented optimal terminal settlement habitat and consequently critical nursery habitat. Model selection indicated water temperature before and during embryonic development was the best predictor of megalopal settlement, whereas juvenile recruitment was most influenced by recent precipitation. I studied the size-specific abundance patterns of blue crabs in and around mature salt-marsh. Using drop-trapping and throw-trapping methods, I collected monthly samples in several habitats during October 2004-March 2006. Interior-marsh habitats contained fewer but larger crabs than bay habitat. Crabs contributing greatest to biomass were smallest (11-30 mm carapace-width) in bay habitat, larger (31-80 mm) along interior-marsh pond edges, and largest (41-130 mm) in interior-marsh open water. Model selection revealed crab density was most influenced by micro-site characteristics (habitat, water column structure type and structural complexity). Overall, shallow bay provided important nursery habitat for young blue crabs and interior marsh ponds were important for dispersing juvenile and adult crabs.