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Drilling predation presents a rare opportunity to quantify ecological and evolutionary interactions in the fossil record. To date, most of this research has been done on Late Mesozoic and Cenozoic deposits, and large-scale studies have focused on temporal rather than spatial patterns. However, drilling predation occurs throughout the entire Phanerozoic, and patterns in spatial variability may mask secular trends. These issues are addressed in a series of projects presented here. An extensive survey of museum specimens and bulk materials indicate that drilling predation in Late Paleozoic brachiopod prey is relatively rare (<1% of fossil specimens are drilled) but widespread and continuously present. The intensity of drilling predation on Late Paleozoic bivalve mollusks (this is the first quantitative report of this kind) is much higher than that seen for contemporaneous brachiopod prey, but lower than what is common for Late Mesozoic and Cenozoic mollusks. Drilling intensity varies significantly between taxa and across localities, (e.g., a sample of the Pennsylvanian brachiopod Cardiarina cordata produced an estimate of 32.7%, which is an intensity similar to that seen in Cenozoic mollusks and the highest yet reported for any brachiopod). However, data for the brachiopod genus Composita, which appears to be a preferred brachiopod prey in many Late Paleozoic assemblages, show that although this genus is subject to drilling predation continuously throughout its geologic range, the over all intensity is very low (less than 1%) and at no time does the intensity ever exceed 10%. Spatial variation in Miocene assemblages from Europe is shown to be on the same order as temporal variation throughout the Cenozoic. Significant variation in drilling intensity is also documented for the Paleozoic. This emphasizes the point that to fully understand patterns of predation through time, both spatial and temporal distribution must be considered. / Ph. D.

Coastal environments are important in many ways: they provide food and energy for coastal communities, have a variety of unique biologic habitats, and influence global climate. These environments can change due to either anthropogenic or natural phenomena over a wide range of time scales. However, the often overlooked long-term (centennial to millennial) processes may be hidden behind short-term fluctuations observed today. The need for a reference baseline of coastal habitats provides a new opportunity for paleontology, which is ideally equipped to document the long-term trends and reconstruct historical and ancient environments and communities. However due to taphonomic processes such as decay and diagenesis, there will always be some bias inherent to the fossil record. This bias has been often viewed as a negative aspect, undermining the utility of paleontological data for retrieving ecological and environmental records. Yet, fossils are still one of the best sources of direct data about the past faunas and their ecosystems. To overcome the biases introduced by the fossilization process, researchers have used observations and experiments in modern systems as models to compare fossil deposits, a method called actualistic paleontology. Over recent years, such actuopaleontological approaches have been used increasingly to exploit distinct signatures of fossil deposits in environmental reconstructions, quantify effects of anthropogenic processes on invertebrate faunas and regional ecosystems, and even augment archaeological studies. Three studied included in this dissertation exploit different aspects of paleontological techniques to study coastal systems, including both Recent and fossil settings. Two of the studies use modern environments to serve as models for investigating preservation potential and potential biases that affect subfossil and fossil assemblages forming in intertidal environments, from whether certain fauna will be preserved to the biases inherent in a shell deposit. The final study, conducted at a fine geologic resolution, focuses on the morphology of a fossil ancestor of an extant mactrid bivalve that is ecologically important in many present-day coastal habitats and well-studied by ecologists and malacologists. The study attempts to quantify morphological effects of environmental changes that occur over geological time scales. / Ph. D.

The Castle Hayne Limestone (Middle Eocene, North Carolina), noted for its diverse macro-invertebrate fossils, was sampled to assess if early Cenozoic brachiopods from eastern North America record any traces of biotic interactions. Systematic surveys of two North Carolina quarries yielded 494 brachiopods, dominated by one species: Plicatoria wilmingtonensis (Lyell and Sowerby, 1845). Despite subtle variations in taphonomy, taxonomy, and drilling patterns, the two sampled quarries are remarkably similar in terms of quantitative and qualitative paleoecological and taphonomic patterns. Ninety-two brachiopod shells (18.6% specimens) contained a single circular hole. Majority of drillholes were singular, perpendicular to shell surface, and drilled from the outside. In addition, ventral valves were drilled slightly more frequently than dorsal ones and larger brachiopods contained more drillholes than smaller ones. However, the size of drillholes did not correlate with the size of brachiopods. The drillholes record "live-live" biotic interactions, which may represent either predatory attacks or parasitic infestations or combination of those two types of interactions. A notable fraction of specimens bears multiple drillholes, which is consistent with either parasitic nature of interactions or frequent failed predatory events. Drilling frequency was high in both quarries (24.5%); this high frequency reinforces other recent reports (from other continents and Cenozoic epochs) that drilling organisms may be a frequent predator or parasite of brachiopod prey or hosts. The number of case studies reporting high frequencies of drilling in brachiopods is still limited and thus insufficient to draw reliable generalizations regarding the causes and consequences of these occasionally intense ecological interactions. / Master of Science

The costal populations of perch (Perca fluviatilis) in some parts of the Baltic Sea have been in decline for about two decades. Recruitment failure in the early larval stages has been put forward as a possible cause and the decline has also been suggested to coincide with increases in three-pined stickleback (Gasterosteus aculeatus) densities. The aim of this study was to study the effects of growth and survival of newly hatched perch larvae in the presence of the three-pined stickleback, and if possible determine the main mechanism behind any negative effects the perch may suffer under such conditions. Using large scale experimental ponds as a controlled habitat, an experiment was conducted where perch larvae were being exposed to sticklebacks under four different stages of their development. Results showed that the three- spine stickleback can have a strong negative effect on the survival of young perch. This effect was strongest in the earliest stage of perch development and decreased as they grew bigger. The zooplankton densities didn't differ between the controls and stickleback treatments, suggesting that the young perch didn't suffer from food limitation. Instead predation was identified as the main mechanism behind the high mortality. The results of this study highlight the potential danger of the observed patterns of decreasing predator populations in conjunction with increasing populations of smaller prey species in the Baltic Sea.

Predation is a key factor in evolutionary dynamics. It disrupts the potential of fossilisation in prey items and is poorly recorded in the fossil record; failed predation in conical marine gastropods is recorded in scars. Quantifying the scar distribution and collection and taphonomic biases present in the fossil record of the gastropod Neptunea contraria, of the Red Crag Formation, Gelasian, Pleistocene, UK is necessary to approach this dynamic. Neptunea contraria is highly abundant in the Red Crag Formation which is easily accessed. The size and scarring on a large number (450+) of individuals was collected, recorded and measured from pre-existing and new material. The size distribution of Neptunea contraria is non-normal and is enriched in larger individuals, the scar distribution – expected to be Poisson – is not so. Taphonomic and Collection bias had a large influence over the size and scar distributions of Neptunea contraria. Material from the same localities shows very different size distributions. The lack of Poisson distribution suggests different rates of unsuccessful predation over life history of Neptunea contraria, assuming the data is valid. / Predation anses vara en viktig faktor inom ekologi och evolution men till vilken grad har effekterna av predation förändrats genom geologisk tid? Det centrala fokuset i denna studie ligger i att frambringa en förståelse av både population- och predationsfördelningen bland marina snäckor av arten Neptunea contraria av Pleistocen ålder från Red Crag-formationen, East Anglia, Storbritannien.Framgångsrik predation resulterar i förstörelsen av snigeln och dess livshistoria registreras i deras skal. Misslyckad predation bevaras i skalen bland individer som överlevt genom ärrbildningar. Det finns ett okänt samband mellan misslyckad och framgångsrik predation.Samlingen av fossilt material från Sedgwick-museet i Cambridge, Storbritannien, ligger till grund för denna studie. Detta material är ofullständigt (d.v.s. material saknas) och noterbart fragmenterat vilket orsakats av nedbrytande processer, därav tafonomi. Denna studie belyser flertalet källor som ger upphov till ett ofullständigt fossilt register, därav processer direkt relaterade till fossilisering och antropogen insamling. Genom att jämföra flertalet uppsättningar av fossilt material som insamlats av olika personer så kan graden av bias i förhållande till insamlingen undersökas. Resultatet av denna studie visar att samlingen av fossila sniglar som för närvarande finns på Sedgwick-museet är ofullständig. Detta är ett tillstånd som uppkommit delvis på grund av inkomplett insamling.Fördelningen av ärr orsakade av misslyckad predation förväntades följa en poissonfördelning. Denna förutsägelse motsägs sannerligen av nuvarande data. Troligtvis har detta förorsakats av en låg ”miss-lyckad predationsfrekvens”, vilket antyder att graden av predation inte är konstant.Sniglar av en större storlek saknar ärr på den övre delen av sina skal, vilket tyder på att frekvensen av misslyckad predation var låg i de juvenila stadierna. (

Dans le contexte des plans Ecophyto, il est primordial de trouver des alternatives aux traitements phytosanitaires. La lutte biologique par conservation fait partie de ces alternatives et, dans ce cadre, nous nous sommes intéressés à déterminer les rôles possibles des araignées en tant que prédateurs généralistes dans les vergers de pommiers. Pour cela, trois axes de travail ont été définis : (i) d’abord étudier les facteurs locaux (protection phytosanitaire et environnement des vergers) expliquant l’abondance et la diversité des communautés d’araignées; (ii) ensuite déterminer si les araignées sont actives en hiver dans les vergers dans le but d’estimer leur rôle sur les stades hivernants de ravageurs; (iii) et enfin étudier la faisabilité et l’efficacité d’un lâcher d’araignées contre le carpocapse des pommes. Dans le premier axe, nous avons mis en évidence des effets significatifs (abondance, diversité, composition) entre les communautés d’araignées issues des vergers en AB et celles issues des vergers PFI, par contre les effets de l’environnement proche des vergers (haies et parcelles alentour) ne semblent pas prépondérants. Dans le second axe, nous avons démontré que dans les vergers du Sud-est de la France, on trouve des araignées actives même au cœur de l’hiver, les plus abondantes furent les Philodromus sp, les Trachelas sp. et les Clubiona sp. Enfin dans le dernier axe, nous avons montré qu’il était possible de transférer et lâcher des araignées d’intérêt (C. mildei et C. leucapsis) et que l’abondance du carpocapse diminuait quand celle de C. mildei augmentait à l’échelle de l’arbre, au moins la première année après le lâcher. L’ensemble de ces résultats permet de poser les bases d’une gestion active des communautés d’araignée en vue d’augmenter leur contribution à la lutte biologique en verger. / It is now crucial to develop new control tools to replace the use of pesticides. Biological control is one of this tool and we thus studied the possible roles of spiders as generalist predators in apple orchards. Three studies were carried out: the first one to determine which local factors (protection management and characteristics of the surroundings) govern spider communities (abundance, diversity) in apple orchards; the second to determine which spiders remain active in winter to control resting stages of some pests; and the last to assess the feasibility of a transfer and release of some spider species into an orchard. We first showed that organic and IPM orchards hosted very different spider community and that, in contrast, local characteristics (hedges and percentage of orchards in the vicinity) had only minor effects. We further demonstrated that some spiders were active winter-long in apple orchards in South-east of France with high abundances for some genus like Philodromus, Trachelas and Clubiona. Lastly we showed that transferring and releasing some spider species of interest (C. mildei and C. leucapsis) is possible and greatly increased the abundance of the released species for one year in the target orchard. We moreover observed a significantly negative relationship between C. mildei and codling moth abundance at the tree scale. These results proved that spider populations can be managed to increase their role in the biological control of important pests in pomefruit orchards.

It is a widely accepted assumption that fish predation controls structure and functioning of aquatic food webs. In the past, however, a large part of effort was concentrated on lakes and reservoirs. Thus, the knowledge about stream ecosystems is much more rudimentary than that for lakes in this respect. The aim of this thesis, therefore, was to describe and assess the effects of fish predation in natural stream ecosystems. For that purpose a reach scale field experiment was set up using an experimental stretch with benthivorous fish and a fishless reference stretch. A wide range of effects of the fish predators on their stream invertebrates prey was studied. To discriminate between lethal and sublethal predation effects, measuring the physiological status of the organisms seemed promising. However, before it was possible to decide whether or not environmental stress, such as predation, might affect the physiological status, the internal control as well as the seasonal and species-specific variability of the energy amount stored had to be assessed. Thus, the concentration and seasonal dynamics of the major energy storage components triglycerides and glycogen were measured in two species of mayflies (Rhithrogena semicolorata and Ephemera danica) with contrasting life cycle strategies. E. danica is a burrowing, semivoltine collector-gatherer, R. semicolorata is univoltine and scrapes periphyton from stones. Although triglycerides are the major energy reserve in both species throughout the whole larval development (&amp;gt; 84 % of total energy storage) their seasonal dynamic differed considerably. In R. semicolorata the triglyceride concentration declined during the last weeks prior to emergence in both sexes. The same pattern was found in female larvae of E. danica, but not in male E. danica. It is suggested that females use triglycerides in the last larval stages for egg maturation, which is completed in the last larval instar. In male E. danica the triglyceride concentrations remained high until emergence, presumably due to their high energy demands as adults for their swarming flights and mating. The difference in seasonal variation of triglycerides between E. danica and R. semicolorata shows the influence of environmental factors on the dynamics of storage components. E. danica lived in a very stable environment (within the substratum). Therefore the dynamic of energy storage components was optimised with respect to maximal reproduction. R. semicolorata on the other hand, suffered from hostile environmental factors such as predation or food limitation due to low periphyton biomass after leaf sprout and following light limitation in spring. Consequently, the concentration of storage components decreased during spring. One conclusion from this study was that the measurement of storage components might reveal sublethal predation effects. However, season and sex of the organisms are important factors as well and have to be considered in the sampling design. To analyse sublethal predation effects behavioural changes due to the presence of benthivorous fish were measured. Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or may even be a direct escape reaction after a predator encounter. While the control of drift activity by predators has received considerable interest from many researchers, it remains still unclear whether predators reduce or increase drift activity. Drift activity of stream invertebrates was influenced significantly by the presence or absence the two benthivorous fish species gudgeon (Gobio gobio) and stone loach (Barbatula barbatula). Contrary to previous studies gudgeon and stone loach reduced invertebrate drift density and drift activity of Baetis rhodani rather than inducing higher night-time drift. Further, species composition of the invertebrate drift differed significantly between the two stretches. A further conclusion from this study is therefore that drift is not generally a mechanism of active escape from benthos-feeding fish, as previously assumed. In addition, the reduced drift activity in the fish stretch might result in a compensation of the consumptive losses due to fish predation. Thus, in this study design the effects of fish predation on invertebrate community might be underestimated. To detect predation effects on the food web structure the reactions of the grazing mayfly Rhithrogena semicolorata and the shredding amphipod Gammarus pulex to strong predation by benthivorous fish were compared. It has been hypothesised that shredders are generally less vulnerable to fish predation and therefore less likely to be predation-controlled than grazers, because the latter are visible to the predators during their feeding on stone surfaces, while shredders may hide between leaves during foraging. Biomass of G. pulex was significantly reduced in the fish stretch while that of R. semicolorata was not. Since approximately 91 % of the annual production of G. pulex but only 12 % of R. semicolorata production was consumed by benthivorous gudgeon, the observed difference of G. pulex biomass between the fish and reference reach is likely due to a lethal predation effect. However, no sublethal predation effects such as reduced concentration of storage components (triglycerides, glycogen) or reduced reproductive success were observed for both species. Hence, in contrast to the initial hypothesis, in the studied stream the shredder was top-down-controlled, while the grazer was not. It is concluded that top-down control depends on the ecological characteristics of a specific predator-prey pair rather than on trophic guild of the prey. To assess the predation effects on the life history of merolimnic insects and its consequences on fecundity the larval development and emergence of R. semicolorata was studied. It was possible to show lethal and sublethal effects of predation by benthivorous fish (Gobio gobio, Barbatula barbatula). Predation consequently resulted in changes of larval development and population fitness. The presence of two benthivorous fish species (gudgeon and stone loach) led to slower larval development and a delayed emergence. However, no differences in the adult size and fecundity between the fish reach and the reference were observed. Nevertheless, the longer time spent in the larval phase resulted in a higher mortality and therefore in a lower mean population fitness. The presence of gudgeon alone, however, did not seem to influence larval development, growth or time of emergence and consequently fecundity. Further, strong lethal impact of gudgeon could not be detected. Thus, the population fitness measured as the product of adult density and egg number was not reduced by gudgeon alone. It is assumed that the stronger lethal impact in the combined fish experiment is caused mainly by stone loach because the proportion of mayfly consumption by stone loach to mayfly production shortly before emergence was higher than the proportion related to gudgeon. Thus another conclusion is that 1) the impact of predation seems to differ for the fish species and 2) lethal effects have a stronger impact on the population survival than life history changes. Combining the results mentioned above leads to the assumption that predation by benthivorous fish has the potential to shape invertebrate communities and food webs in streams. It was possible to show reductions of benthic densities and mean population fitness. The strength of trophic interactions seemed to be specific for the single predator-prey pairs here. Finally, it can be stated that contrary to previous assumptions consumption of the fish predators seemed to be more important for the prey populations than sublethal predation effects.

Thesis (M.E.S.)--The Evergreen State College, 2001. / Title from title screen viewed (3/20/2008). Includes bibliographical references (leaves 41-44).