Understanding strategies of resource partitioning within populations is a fundamental aspect of the study of ecology, and one which has been well documented across various taxa. In seabirds, sex-specific differences in foraging behaviour are often thought to reflect strategies of resource partitioning to avoid intra-specific competition. At present Bird Island in Algoa Bay hosts the world’s largest breeding population of gannets, with recent population estimates numbering well over 90 000 breeding pairs. Given the large size of this colony, intense levels of intra-specific competition are expected during the breeding season, particularly in years of poor feeding conditions. In addition, continuous foraging around the breeding colony may lead to the local depletion of prey as the season progresses, calling for flexibility in the foraging behaviour and diet of breeding individuals. Therefore, the aim of this dissertation was to investigate resource partitioning in Cape gannets breeding at Bird Island through strategies of sex-specific foraging and intra-seasonal variability in foraging behaviour and diet. To achieve this, the spatial and temporal aspects of Cape gannet foraging behaviour were assessed during the early stages of chick rearing, over two consecutive breeding seasons (2011/12, 2012/13) using GPS tracking and VHF monitoring. In addition, intra-seasonal variability in foraging behaviour and diet were investigated, taking into account the influence of sex and chick age, to provide a comprehensive assessment of foraging behaviour throughout the chick rearing period. GPS tracking data for 79 birds revealed limited evidence for sex-specific differences over spatial dimensions. An extension in the foraging range of females during a year of lower prey availability (2012/13) was, however, evident. This suggests a possible sex-specific response to prey limitation, likely reflecting resource partitioning strategies or differences in nutritional requirements. VHF data for 95 additional birds revealed a clear pattern in temporal foraging behaviour. In general, females were more likely to be on a foraging trip during the early and mid-day hours, with the probability of males being on a foraging trip increasing towards the late afternoon. Continuous tracking of individuals throughout the 2015/16 breeding season further revealed little evidence for sex-specific differences in spatial distribution and forging effort during both the guard and post-guard stages of chick rearing. A clear increase in foraging range was evident during the early stages of chick rearing as the season progressed. However, birds attending chicks older than 50 days showed an unexpected contraction in foraging range, which appeared to coincide with a dietary shift. During the early stages of breeding the diet was comprised almost exclusively of anchovy, the proportion of which decreased progressively in the diet throughout the breeding season. This was mirrored by a substantial increase in the proportion of saury in the diet. These results suggest that Cape gannets increase their foraging range in response to local prey depletion as the breeding season progresses. However, when alternative prey becomes available they may shift their diet, allowing them to access prey closer to the breeding colony. Overall, this dissertation provides insight into sex-specific behaviour in a monomorphic seabird, suggesting a marginal degree of spatial segregation. The results provide the first support for sex-related temporal foraging segregation in gannets. Such separation could potentially be driven by resource partitioning, but could also relate to differential nutritional requirements. In addition, this study also provided the first comprehensive assessment of foraging behaviour throughout the guard and post guard stages of chick rearing in gannets. These results suggest that a combination of foraging and dietary flexibility may allow Cape gannets to buffer the effects of prey depletion during the breeding season. In conclusion, resource partitioning through sex-specific foraging appears to be limited in the Cape gannet population at Bird Island. Segregation via alternative pathways may however be possible and as such, future studies should consider the influence of individual level segregation and habitat specialization.

Two congeneric damselfly species, Enallagma traviatum and E. aspersum, dominate the littoral macroinvertebrates of Bays Mountain Lake and of the adjacent fish-free Ecology Pond, respectively (northeastern Tennessee, USA). Extending previous experimental studies, we test seven hypotheses concerning the role of fish (bluegill sunfish, Lepomis macrochirus) and larvaldragonfly (Anax junius) predation, competitive effects on damselflies, and the interaction between competition and predation, in determining invertebrate dominance in these communities. Three types of experiments were conducted: an enclosure experiment within Ecology Pond, an outdoor replicated tub experiment, and a laboratory behavior experiment. The in-situ enclosure experiment showed that E. traviatum larvae were more susceptible to Anax predation than were E. aspersum larvae; a tendency toward greater vulnerability to fish of E. aspersum compared with E. traviatum was not statistically significant. The outdoor tub experiment confirmed both of these trends with statistically significant results. In the tubs, both predators inhibited feeding of both zygopterans (as indicated by reduced fecal mass), particularly for E. aspersum in the presence of fish. This effect appears to have been primarily indirect, mediated through exploitation of the zooplankton. We also detected competitive effects of E. traviatum on E. aspersum: E. traviatum reduced the emergence and increased the exposure above the substrate of E. aspersum. In the absence of predators, E. traviatum inhibited feeding of E. aspersum via interference. In the laboratory behavior experiment, predators inhibited crawling by E. aspersum. E. aspersum was more exposed than was E. traviatum; it swam and crawled more than did E. traviatum, considerably increasing these movements at night. Over all, E. traviatum consistently appeared to be the more cryptic of the two species, and E. aspersum appeared to be much more active. Our results suggest an explanation for the clear difference in structure between communities like Bays Mountain Lake and Ecology Pond: predaceous fish eliminate large invertebrate predators and shift the community toward cryptic forms at relatively low densities, reflecting the effects of both predation and exploitation competition. In the absence of fish, large invertebrate predators are less able to deplete littoral invertebrates but may favor the more active forms, perhaps because these are better able to avoid invertebrate predators.

The Cape gannet has undergone considerable population change and redistribution over the past 50 years. This has been linked to shifts in the abundance and distribution of their dominant prey, sardine and anchovy. Five breeding colonies, along the west coast of Southern Africa, have shown rapid population declines as a result of reduced prey availability. In contrast, a single colony (Bird Island, Algoa Bay) on the south coast of South Africa has, over the same period, grown fivefold and now supports approximately two thirds of the total population. Due to its conservation importance, and isolation from other breeding localities, it is important to assess the health of the Bird Island colony, and determine how foraging distribution relates to the environment to evaluate current measures of protection. This was achieved through two related studies; a long-term dietary analysis spanning 34 years, and a spatial foraging study, which related three years of tracking data to estimates of prey availability, oceanographic features and marine protected areas (MPAs). The results of the dietary study showed that the dietary constituents of Cape gannets breeding at Bird Island have remained similar over the last three decades, but the importance of sardine and anchovy has increased significantly. For sardine, in particular, this reflects an increased availability of this species (as deduced from hydroacoustic surveys) within the foraging range of the Bird Island colony. The dietary abundance of anchovy was found to be negatively correlated with that of sardine. Surprisingly,.the dietary abundance of anchovy was also negatively correlated with estimates of its availability based on acoustic surveys. The latter is likely to be due to sardine being a preferred prey item. Recent decreases in the dietary contribution of sardine (since 2005) suggest that this species is becoming less available to gannets, with profound implications in terms of nutrient gain associated with foraging. However, this has been mediated by an increase in the dietary contribution of anchovy, which now accounts for the vast majority of prey taken. Spatially, the foraging range of the Bird Island colony expanded during the three years of study, indicating an increase in effort. This increase was likely in an effort to track a distributional change of sardine and anchovy, which showed an apparent westward shift during the study period. There was, however, no evidence of birds tracking features associated with high productivity. This may have been partly due to anomalously warm conditions during the summer of 2012/2013, in which an absence of coastal upwelling prevented surface cooling. Nonetheless, low sea surface temperatures and high chlorophyll a concentrations do not seem to be reliable indicators of important Cape gannet foragingareas. Foraging effort was largely concentrated outside of MPAs, indicating that the current MPA network provides little protection for foraging gannets. This could change with the additional protection of the proposed Greater Addo Elephant National Park MPA expansion, as prey species are allowed to recover following the removal of fishing pressure. Overall, the colony appears to be in good condition as the diet is still dominated by live prey items, and the foraging range remains smaller than many of the colonies along the west coast. However, it is important that monitoring be continued, in particular with respect to changes in the availability of sardine and anchovy. Long-term shifts of these species out of the colony‘s foraging range could negatively influence the population in the future. This might be worsened by interspecific competition for prey resources. Considering the conservation importance of this population, maintenance of healthy prey stocks within the home range of breeding Cape gannets should be prioritised in order to prevent declines similar to those observed at west coast colonies.

For several decades now, seabirds have been considered to be useful indicators of the state of their prey resources because of how they reflect lower-level trophic variability through proxies such as diet or behaviour. However, collection of appropriate data is often challenging in the marine environment because of logistical or financial constraints. In this study, time-activity budgets were studied in the Cape gannet (Morus capensis), a seabird which has been advocated as a potential bioindicator for local epipelagic prey. VHF transmitters attached to PVC leg-rings were fitted to 50 adult breeding pairs during the 2011/2012 and 2012/2013 breeding seasons at Bird Island, Algoa Bay, to determine whether an automated method of continuously collecting time-activity budget data can replace conventional laborious direct observations. To validate that the foraging trip duration data generated from the automated method was a reflection of foraging effort, Time-Depth Recorders (TDRs) were simultaneously equipped to birds with VHF transmitters for three weeks. In order to assess the influence of factors other than prey availability on parent time-activity budgets, sex, chick age and body condition were measured. Additionally, chick growth and survival were recorded in order to investigate the gender-specific effects of parental time-activity budget variability on these parameters. Attachment of VHF transmitters to leg-rings of adult Cape gannets had no observable negative effects on the adults or their chicks in terms of adult body condition, nest attendance and foraging trip durations, or chick growth and survival. Furthermore, the frequency distribution of behaviour bout (foraging trip and nest attendance) durations was no different between automated and directly observed data. However, the automated method did record shorter behaviour bouts, largely attributed to the increased likelihood of direct observations missing birds returning briefly to their nests during older chick provisioning. Additionally, foraging trip duration was highly correlated to foraging effort in terms of time spent resting on the sea surface, flying and diving. The automated method therefore appears to be a good reflection of direct nest attendance observations and foraging effort. Cape gannet time-activity budgets were related to chick age and parental sex. Especially as chicks neared fledging, females spent significantly longer periods of time foraging than males, with males consequently provisioning their chicks more often. Furthermore, adults departing their nests earlier spent more time away from the nest foraging as more same-day daylight hours were available. Chick growth was a function of parent foraging trip duration and associated prey delivery rates. Chick survival was most strongly affected by the amount of time which chicks were left unattended by both parents and consequently exposed to predation by kelp gulls (Larus dominicanus) or to adverse weather conditions. Additionally, for females (but not males) there appeared to be a trade-off between foraging trip duration and chick survival. The Cape gannet appears to demonstrate a strategy whereby the costs of reproduction to the female are shifted towards male-dominated chick provisioning as the chick nears fledging. Drivers of time-activity budget variability such as chick age and parental sex therefore need to be considered if using data on foraging trip duration as a proxy of foraging effort and prey availability.

Predators can have large effects on prey populations and on the structure and function of communities. In addition to direct consumption of prey, predators often cause prey to alter their foraging behavior, habitat selection, and morphology. These non-lethal effects of predators can propagate to multiple trophic levels and often exert equal or larger effects upon communities than those of direct consumption. For non-lethal predatory effects to occur, prey must detect and respond to predation risk. While the importance of information transfer in this process has been realized, few studies explore how prey responses are influenced by predator characteristics and environmental conditions that influence the transmission of cues indicative of predation risk. In this dissertation I investigate factors that influence how a single prey species evaluates and responds to predation risk. Here, I examined: 1) the type and nature of cues prey use to evaluate predator risk; 2) how predator identity, predator diet, and the relative risk of predators influence prey response to predation risk; 3) how hydrodynamic conditions influence the delivery of predator cues; 4) how biogeographic trends in predator distribution influence prey response to predation risk; and 5) how genetic structure might vary according to prey geographic location and habitat. To address these questions, I used a common intertidal model system consisting of the rocky intertidal whelk Nucella lapillus (Linnaeus, 1758) and a suite of its predators, the native rock crab Cancer irroratus (Say, 1817), Jonah crab Cancer borealis (Stimpson, 1859), and the invasive green crab Carcinus maenas (Linnaeus, 1758). Nucella use chemical cues emanating from their most common predator (Carcinus maenas) and crushed conspecifics to evaluate predation risk. Nucella from different habitats experience different levels of predation risk, and Nucella from habitats with high levels of predation had larger antipredatory responses to predator risk cues than Nucella that experienced less predation. These chemical cues indicative of predation risk are influenced by hydrodynamic conditions, and Nucella have the strongest anti-predatory response in flow velocities of u= ~4- 8 cm s^-1. Furthermore, Nucella from geographic regions where green crabs are historically absent did not elicit anti-predatory responses, while Nucella from regions where green crabs are common frequently responded. Findings from my dissertation research demonstrate that prey detection and response to predation risk is highly dependent upon predator identity, predator diet, environmental forces, and biogeographic patterns in predator and prey distributions.