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Predation is considered a powerful ecological force influencing community structure, diversity, and abundance. Paleoecology offers a unique perspective, allowing us to examine ecological processes such as predation over evolutionary timescales. The three studies summarized below include two case studies testing the role of predation in evolution and one method-oriented project, which explores new tools with which to examine predator-prey interactions in the fossil record. Considering the importance of community interactions in the current global biodiversity crisis, understanding ecological and evolutionary dimensions of predation is critical to conservation biology and ecology, as predators are thought to play a vital role in maintaining ecosystem health and biodiversity. Predation has been dismissed as a causal mechanism for some major prey groups. For example, the evolutionary decline and cryptic or antitropical distribution of brachiopods is often explained as due to the potentially low energetic value and suspected non-palatability or toxicity of brachiopod tissues. Here we demonstrate that multiple invertebrate marine predators (crustaceans, echinoderms, and gastropods) are willing and able to consume brachiopods, and that predation pressure on the living brachiopod population may be consequential. Examination of the fossil record is consistent with this interpretation: evidence for drilling and repair of brachiopod shells is found throughout the fossil record in multiple orders. This suggests that although brachiopods may be unwanted prey in the presence of energetically more desirable targets, they do appear to be edible and are subject to intense predator-prey interactions. Limpets are important prey for some crab species, yet little is known about the role of durophagy in the evolution of the limpet shell. Feeding trials using three common species of Pacific Northwest limpets (Lottia digitalis, L. pelta and Tectura scutum) were conducted to assess how different shell morphologies affect mortality and handling time. We predicted that large size, shell ornament, and low-spires would result in either increased survivorship, and/or longer handling times. Contrary to our expectation that ridges resist predation, individuals with smooth morphologies experienced significantly lower mortality, as did those with low-spires. As species possessing high-spires and ridges typically occur high in the intertidal where predation risk due to crabs is relatively low, these morphologies are likely adaptions to physical factors such as thermal stress. One of the major caveats of using gastropod drill holes to assess predator-prey interactions in both the modern and the ancient is the correct identification of drill holes of predatory origin. By examining known predatory drill holes using environmental scanning electron microscopy, we aim to refine the development of a novel technique for augmenting their identification, and to explore the relationship between predator body size, predatory radula dentition, and radular microrasping marks observed on the shells of prey organisms. Electron micrographs were used to measure the spacing of microrasping marks produced by the radula, and the intercusp spacing of the radula dentition. A relationship between predator body size and microrasping marks makes it possible to infer predator size from these microtraces in both modern and fossil specimens, augmenting our ability to examine predator-prey interactions throughout the history of this important ecological interaction. Proxies for predation intensity such as predation traces or antipredatory morphologies provide an invaluable method to examine predation in both modern communities, and the fossil record. Our understanding of the importance of predation in regulating biodiversity and in evolution will continue to grow with the development of new methodologies, and a comprehensive understanding of predatory defenses. / Ph. D.

Due to high mortality encountered by marine fish larvae during their first weeks of life, small changes in the number of individuals surviving through this period can cause large fluctuations in year-class strength. Larval Red Drum (Sciaenops ocellatus) are dependent upon structured estuarine habitat to avoid predation. A study of post-settlement larval Red Drum distribution in a subtropical seagrass meadow in Mission-Aransas Estuary, Texas, USA indicates that larvae settle over approximately two months. Abundance of larger settled larvae was significantly different among sites. The areas of highest larval abundance varied temporally, indicating that the entire extent of the seagrass bed is utilized. Regression analysis of abiotic environmental factors did not explain why larvae were more abundant at particular sites. To characterize the structure and variability of the fish species assemblage that Red Drum encounter upon settlement, larvae and juveniles were captured in the seagrass meadow during weekly collections. Of the 32 fish species collected, seven represented 92% of the assemblage. Multivariate species analysis indicated that collections widely separated in time and space shared the lowest Bray-Curtis similarity. Because Red Drum settle over a relatively long period and co-occur at body sizes known to cause cannibalism under laboratory conditions, I tested combinations of small and large Red Drum larvae at various field-realistic densities and at different levels of seagrass habitat structure to determine potential for cannibalism. Artificial seagrass did not protect small (5 – 6 mm SL) larvae from cannibalism, but natural dense seagrass had a protective effect relative to edge habitat. The final component of this research examined the emergent impacts of a common predator pinfish (Lagodon rhomboides) on mortality and cannibalistic interactions between small and large Red Drum larvae. Both pinfish and large Red Drum larvae alone readily consumed small Red Drum in all seagrass habitat structures tested. However, the combined treatment of pinfish and large Red Drum together led to reduced mortality of small Red Drum. Predation can significantly affect Red Drum survival during the post-settlement period, and multiple predators may have a protective effect on the smallest settlers if predation pressure is re-directed towards a larger size class. / text

Female choice is an important selective force shaping the evolution of communication and speciation in animals. However, predation risk can impose severe costs on longer searches and choosiness, thereby limiting the expression of female preferences for specific male traits. The work detailed in this dissertation explores how mate choice and sexual selection can be influenced by predation risk in túngara frogs. I begin by examining the effects of multiple simulated cues of predation risk on female search behavior and mate choice, taking a departure from the standard presence/absence paradigm used in similar studies to explore responses to quantitative variation in perceived predation risk. I demonstrate that light, longer travel times, and acoustic cues of predators are all sufficient to sway females away from otherwise more attractive conspecific males. Next, I explore the role of predation risk in altering female permissiveness, or the range of signals females will respond to. Using an artificial series of calls intermediate between heterospecific and conspecific, I demonstrate that predation risk dramatically increases the range of signals females will respond to, including a small number of females choosing pure heterospecific calls. Next I attempt to bridge a logical gap with our understanding of search costs, testing questions about how female search paths change with increasing distance. I demonstrate that females use more direct paths and move faster under higher light conditions, potentially reducing sampling but also reducing encounter rates with predators. Lastly, I examine factors that influence how individual females vary in their response to perceived risk, particularly hormonal state and experience. I demonstrate that naïve, captive-bred females respond to acoustic cues produced by natural predators in a manner similar to wild females and that, while hormonal state is obviously important in determining female receptivity, it has little effect directly on how females respond to predators. Together, these studies demonstrate that predation risk not only changes how females respond to conspecific males, but also increases female permissiveness and constrains search behavior. Predation risk can strongly influence and potentially even negate the expression of female preferences, having profound consequences for communication and the evolution of reproductive isolation between populations. / text

Nest success, feeding behavior and anti-predator strategies, were studied to determine if bellbirds (Anthornis melanura) are limited by predation and if pest control can relieve the pressures of predation so that food supply may become a limiting factor for bellbirds. The nest success of bellbirds at the Rotoiti Nature Recovery Project (RNRP), Nelson Lakes, New Zealand (2002-2004 breeding seasons) was compared with data from other bellbird nest success studies in New Zealand. Two years of nest monitoring resulted in a low overall nest success at the RNRP (39 %), an area low in predator density, compared with 16.4 - 67.2 % found for other sites. One-way anova comparisons between the nest success of bellbirds in areas grouped by predator density (absent, low and high) were not significantly different. Regression analyses of nest success versus predator density (stoat or rat tracking indicies) were also not significant. A test of change in bellbird numbers (numbers in February/numbers in November in the same breeding season) versus rat density (tracking index in November) showed no significant trend at the RNRP. A test of density dependence (change in bellbird numbers versus numbers in November) was also not significant. The percent of time bellbirds spent feeding was significantly higher at the RNRP compared to Mt. Misery (Nelson Lakes, New Zealand) during the breeding season only (RNRP = 34 % and Mt. Misery = 23 %). The percent of time bellbirds spent foraging for food was also significantly higher at the RNRP compared to Mt. Misery during the breeding seasons (RNRP = 47 % and Mt. Misery = 36 %). This suggests that predator control at the RNRP has allowed bellbird numbers to increase so that nesting success may be partly limited by food supply during the breeding season. However, despite higher densities of bellbird and other bird species at the RNRP, there was no difference in the percent of time spent feeding (RNRP = 36.7 % and Mt. Misery = 36.3 %) or foraging (RNRP = 49 % and Mt. Misery = 51 %) between the RNRP and Mt. Misery during winter months The nest height of bellbirds was not significantly related to the nest outcome (excluding nests that failed due to non-predation causes) at either the RNRP or Kowhai Bush. Nest predation occurred at a range of heights, which suggests introduced predators are not specialists at any nest height. Bellbirds at the RNRP 111 visited their nests more frequently and stayed for shorter periods at the nest compared with bellbirds at the Cheeseman Valley, consistent with the hypothesis bellbirds alter their behavior to avoid disclosing their nest position. The results of this study, although not significant, do indicate that bellbird nest success is limited by high densities of predators. One-way anova analysis on nest success during the feeding stage was nearly significant (P = 0.096). Regression analysis trend lines show an increased negative effect on nest success at higher predator densities. Few replicates and large variation in nesting success within treatments affect the power of the results. Bellbirds may persist in areas with high predator densities due to anti-predator strategies when visiting their nests. With pest control, ecosystem interactions may be revitalized so that limiting factors other than predation become important for bellbird nest success.

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The life-history traits of two sympatric seabird species, the Common Sterna hirundo and Arctic Tern S. paradisaea, and the flexibility of these traits in relation to short and longer term changes in environmental conditions were measured at Coquet Island, N.E. England. The study focused primarily on inter and intraspecific differences in annual productivity and chick growth, adult time budgets and provisioning rates, and the relationships between these different aspects of reproductive performance. Of the two species, Common Terns delivered larger food items, delivered food at a higher rate to the nest and attended the nest more frequently, indicating that they made trips of shorter average duration. Daily metabolizable energy intake of chicks was about 30% higher in Common Terns than in Arctic Terns, yet the size-specific growth rates of the two species were almost identical, indicating a major difference between species in nestling energy budgets. Brooding appeared to play a less important role in the energy budgets of Common Terns, and the number of chicks that Arctic Terns could raise was probably limited not only by the rate at which parents could supply food to the nest but also by the requirements of chicks for brooding. Increased brood size, low annual food abundance and extreme weather conditions had a negative effect on chick mass development in both tern species. Flexibility of mass growth rates in Arctic and Common Terns may act as a fine-tuning mechanism to regulate provisioning in these species. By maintaining structural growth rates, final fledging mass and final fledging size of nestlings at the expense of retarded mass development rates, these species seem to be able to maximize annual reproductive output and possibly, for parents and nestlings, future survival. Predation of eggs and chicks was generally infrequent and affected mostly very young nestlings. However, Black-headed Gulls took many tern eggs in a year when inshore food supplies were particularly low.

Long-term insect abundances have been studied to investigate factors structuring the insect communities of the temporary wetlands around the River Dalälven floodplain. In the wet meadows and swamps in the River Dalälven floodplains recurrent floods are the ultimate prerequisite for high production of floodwater mosquitoes. In the larval stages these make up a superabundant potential prey for aquatic predatory insects while as adults they become nuisance for people and animals. Mosquito control with Bacillus thuringiensis israelensis (Bti) has been carried out in parts of the floodplain wetlands since 2002. This study investigate potential long-term effects of Bti on non-target insects in general, and non-biting midges (Diptera: Chironomidae) and aquatic predator insects in special in 10 temporary wetlands. Diving-beetles (Coleoptera: Dytiscidae) were the most common aquatic predatory insects in temporary flooded wetlands, and we found no indications that diving-beetles were negatively affected by mosquito larvae reduction with Bti-treatments. Non-target Bti-effects on the insect community in general, measured as increase or decrease of insect abundances, in general showed no effects of Bti except for Coleoptera. The non-biting midges were studied on species level, since this taxon previously has been associated with negative Bti non-target effects in the short-term. We found no reduced production of chironomids at neither family, nor subfamily level in Bti-treated as compared to untreated wetlands. We conclude that other factors than the occasional Bti-treatments dominate in structuring the chironomids fauna. For example, hydrology measured as floods and hydroperiod had substantial impact on insect emergence Increased hydroperiod was associated with lower insect emergence for the majority of the studied taxa, indicating an insect fauna adapted more to terrestrial conditions. In the final study, we developed and applied a molecular method to study interspecific predator – prey relationships between medium-sized diving beetles and floodwater mosquito larvae. Gut content analyses showed that floodwater mosquito larvae are a regular, but limited, part of the diet of medium-sized diving beetles. This thesis is one of the first long-term studies of insects of temporary wetlands in relation to mosquito control actions. The results indicate that hydrology is one of the major factors influencing and structuring the insect communities of the temporary flooded wetlands in the River Dalälven floodplains, and that mosquito control actions with Bti only have marginal effect on insect abundances.

Sessile communities provide an ideal opportunity to understand how population interactions are structured in space, and in turn, how this structure influences population dynamics. The yellow tube sponge, Aplysina fistularis is common on Caribbean coral reefs and is involved in four main types of ecological interactions: symbiosis, predation, disease, and competition. We used spatial and multivariate analysis to describe the population structure of A. fistularis on the Belizean barrier reef, and consequently, how the observed spatial structure influenced sponge morphology and the frequency of population interactions. We found that sponges were non-randomly distributed across the reef, with both local density and tendency towards a clustered spatial distribution increasing with depth. Sponge morphology also varied with depth; deeper sponges were larger and had fewer tubes then their shallower conspecifics. Local density and spatial autocorrelational patterns were not significant predictors of population interactions. The frequency of symbiosis, characterized by the presence of the fish Elacatinus lori, increased with depth, sponge size, and number of sponge tubes. The incidence of predation increased with depth, number of tubes, and the interaction between these two variables. The chance of disease decreased with increasing sponge size. Lastly, the frequency of spatial competition decreased with depth. These results highlight the unexpected finding that in this system, symbiosis, predation, disease, and competition, are density independent, specifically they are not predicted by local density or spatial autocorrelational patterns. Overall, this study provides an essential framework that will greatly enhance our knowledge of sponge ecology on coral reefs.

To evaluate preventive aerial coyote hunting as a depredation management technique, I compared sheep losses to coyote (Canis latransl predation and the hours of corrective predation management required on summer grazing areas with and without hunting the prior winter from helicopters. Correlations were used to test for relationships between the extent, intensity, and timing of aerial hunting and lamb losses to coyote predation. Data on the age, sex, and reproductive status of coyotes killed using aerial hunting, traps, snares, and calling-and-shooting were used to test for differential coyote vulnerability to damage management tools, and to assess the impact of aerial hunting on coyote populations. Winter aerial hunting reduced confirmed and estimated lamb losses to coyote predation and the hours of effort required for corrective predation management the subsequent summer. Aerial hunting increased the number of coyotes killed annually per grazing area, but did not reduce summer coyote removal. There were no consistent relationships between the extent, intensity, or timing of aerial hunting and sheep losses to coyote predation. The male: female ratio for coyotes captured with calling-and-shooting was higher than that for traps or aerial hunting. More juvenile coyotes were killed with aerial hunting than with traps or shooting. However, there was no difference in the age of adult coyotes {>1.5 years old) removed using any control method or between the age of coyotes from areas with and without consistent aerial hunting. Confounding factors in the data and the high number of uncontrolled variables prohibited clear identification of the mechanism making aerial hunting effective. I also examined financial compensation programs as an alternative to lethal control. Nineteen states and 7 Canadian provinces had compensation programs. Compensation programs appeared to be established when wildlife problems were of recent origin, resulted from government actions, and/or were caused by highly valued species. Compensation programs for coyote damage had been established in 4 states/provinces in eastern North America where coyotes are a new problem, but are unlikely to be a acceptable tool for the western U.S.