This thesis focuses on a black-tailed godwit (Limosa limosa) population, consisting of 35-40 pairs, that breeds on a grazed shore meadow on SE Gotland, Sweden. The small size of the population makes it more prone to extinction due to chance events, than a larger population. The godwits showed microhabitat preferences when choosing nest sites. Godwit nests had higher vegetation over the nest cup, lower surrounding (1-3 m) vegetation and different plant species composition, as compared to random sites. Breeding near conspecifics or other wader species, especially lapwings (Vanellus vanellus) and further away from potential predator perches were the most important factors in decreasing nest predation. A comparison between different shore meadows along the east coast of the island revealed that large, open areas suffered less from nest predation. Thus, shore meadows suitable for breeding godwits should be large and without trees or other predator perches and have a grazing regime that favours variation in vegetation height. Over 80% of previously ringed adults returned each year, but very few birds ringed as chicks were recovered. Hatching success was 55-60% for all observed nests. To predict the future of the current population, demographic data were used in an ecological risk analysis. The simulations showed that the Gotlandic population will not survive the coming 40 years without immigration. Black-tailed godwits are divided into three subspecies. Genetic analyses (mtDNA) revealed that all subspecies had unique haplotypes and there was a clear geographic structure among subspecies. Within the limosa subspecies, godwits on Gotland and Öland showed a high proportion of rare haplotypes, but no genetic variation was found in Dutch birds. These results imply that black-tailed godwits on Gotland and Öland have high conservation value.

In this thesis I investigate predator-induced effects on behavioural and life-history characteristics of prey. At any moment a given predator is capable of attacking a small number of prey. However, the mere presence of a predator may impact a much larger number of individuals, as prey implement various behavioural and developmental mechanisms to reduce the risk of predation. It has become increasingly clear that predator induced responses have the potential to affect patterns of species abundance and distribution as well as individual fitness of prey. I study these responses by incorporating field surveys, semi-field experiments and laboratory experiments. All experiments were done in an aquatic environment using fish or large odonate larvae as predators and damselfly-or diving beetle larvae as prey. My work highlights the importance of monitoring prey behaviour when studying life-history characteristics. I show that fish presence is an important factor for determining species abundance and distribution of odonates, and that prey behaviour may be a good predictor for fish vulnerability. Larval damselflies react behaviourally to predator presence by reducing activity and/or restricting habitat use. I confirm that such anti-predator responses have positive effects on prey survival in the presence of a predator but negative effects on growth and development of prey. In addition, my results suggest that the increase in per capita food resources for surviving prey following a predation episode (i.e. thinning) can have a stronger positive effect on prey growth and development than the negative effect of anti-predator responses. I also show that the strength of an anti-predator response is dependent on resource availability of the prey, with prey responding less strongly when resources are scarce. My results also indicate that the strength of the anti-predator response of damselfly larvae depends on predator diet and larval age. Predators feeding on prey conspecifics induce a stronger behavioural response in young larva than predators that feed on prey heterospecifics do. This diet-effect was not found in larvae late in ontogeny, due to an increased activity of larva where predators consumed damselflies. Such increased larval activity can be explained as a reaction to a time-constraint. Finally, I found that activity of damselfly larvae is genetically determined and that this has lead to a behavioural syndrome that might limit larval plasticity to a certain activity-range. This phenomenon may have implications for how well larvae are able to react to both biotic and abiotic changes in the environment.

A prey animal can use different strategies to avoid becoming eaten by predators. One such widely recognised strategy is the use of body colouration to decrease the risk of becoming detected, i.e. cryptic colouration. The principles of crypsis that I have studied are background matching, disruptive colouration and distractive markings. Further, I also studied the concealing effect of the visual background habitats. I used artificial prey items and backgrounds, and blue tits (Cyanistes caeruleus) as predators, to investigate prey concealment. In Paper I, I tested if high-contrast markings in prey coloration or in the background would result in a distracting effect. I found that such markings did increase prey search time, even when the prey markings were lighter or darker than the background. In Paper II, I studied the use of chromatic cues by predators when searching for prey. The birds easily detected prey that chromatically deviated from its background. Interestingly, background-matching prey was more difficult to detect when the colour scheme had low ultraviolet and high shortwave reflectance compared to when the reflectance bands were even. In Paper III, I studied optimisation of achromatic contrast within prey colour pattern and also the effect of shape diversity of background pattern elements on prey detection. I found that all prey types were more difficult to detect on the diverse background, but the level of contrast within prey pattern did not influence search times. In Paper IV, I further investigated how a prey should optimise its patterning with respect to background matching. I found that prey with repeated pattern elements was equally hard to detect as prey with more variable pattern. However, prey with a spatially regular pattern (aligned pattern elements) was easier to detect than prey with a spatially irregular pattern. In this paper I also found that high complexity of element shapes in the background, made the search task more difficult. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript. Paper 2: Accepted

The dynamics of natural populations may be influenced by a variety of factors, ranging from feeding interference and territoriality to the risk of predation and climatic effects. The relative influence of these factors may be contingent upon the quality of the habitat in which individuals reside. A factor that can largely affect population dynamics and that often covaries with habitat structure is predation risk. However, the combined effect of habitat and predation risk can vary according to the social context and intrinsic characteristics that affect individual behavioural responses. This thesis investigates the effects of habitat quality at the level of the population and the individual, and examines how it relates to the above factors in Siberian jays (Perisoreus infaustus), a territorial, group-living species in which the main cause of mortality is predation. The results demonstrate a strong effect of habitat on survival, reproduction and behaviour. Mortality was generally higher in open areas and managed forests and reproductive success decreased after forest management. Breeding females were more sensitive to environmental factors than males, possibly because of higher reproductive costs. Estimates of spatial demography suggested that there were more sinks than sources, and that they were located in open, managed forests. Behavioural decisions confirmed that open forests were associated with higher predation risks. However, decisions depended on social context; immigrants took highest risks and were the recipients of most aggression, largely an effect of social subordination. Also, parents provide their offspring with benefits that are withheld from immigrants. As a result, first-year survival was higher in retained offspring. Investigating the effects of multi-scale habitat quality on individual behaviour and population dynamics has generated an increased understanding of the effects of forest management on the dynamics of this population. This approach should facilitate development of an effective conservation management policy for this species.

The effect of environmental modification of predator sensory abilities remains largely unknown, despite the importance of predators to ecosystem function. I conducted a series of experiments to investigate effects of hydrodynamics on the chemosensory search behavior and foraging success of two species of marine gastropods, knobbed whelks (

Biomanipulation im Sinne einer top-down Steuerung des Nahrungsnetzes wurde in Standgewässern intensiv erforscht und wird als Mittel zur Verbesserung der Wasserqualität bzw. Verringerung der Eutrophierungserscheinungen eingesetzt. Ebenso könnte die Idee der Biomanipulation genutzt werden, um die Eutrophierung in kleinen Fließgewässern zu verringern. Eine Förderung benthischer herbivorer Schlüsselorganismen (z.B. Eintagsfliegenlarven) durch die Reduzierung des Fischfraßdruckes könnte helfen, Algenmassenentwicklungen zu vermeiden. Studien zur Biomanipulation von Nahrungsnetzen in Standgewässern zeigten allerdings eine zunehmende Entwicklung von planktischen invertebraten Räubern bei Abwesenheit planktivorer Fische. Die Bedeutung des Fraßdruckes invertebrater Räuber in fischfreien Fließgewässernahrungsnetzen ist dagegen noch unbekannt. Aufgrund von letalen und subletalen Fischeffekten könnte die Konsumtion räuberischer Invertebraten bei Fischabwesenheit so stark gefördert werden, dass das Ziel der Biomanipulation verfehlt wird. Invertebrate Räuber haben insgesamt einen relativ hohen Anteil an der Biomasse der Benthosgemeinschaft in Fließgewässernahrungsnetzen. Sie könnten somit potentiell einen ähnlichen Fraßdruck wie vertebrate Räuber (z.B. benthivore Fische) auf die unteren trophischen Ebenen ausüben. Auch der Anteil der Omnivorie ist in Fließgewässernahrungsnetzen hoch. Ziel dieser Arbeit war es deshalb, die Bedeutung invertebrater Prädation im Nahrungsnetz eines kleinen Fließgewässers (Gauernitzbach) unter Berücksichtigung der obligaten Räuber sowie des räuberischen Potentials Omnivorer abzuschätzen. Außerdem wurde der Einfluss benthivorer Fische auf die Populationsentwicklung und das Ernährungsverhalten der wichtigsten invertebraten Räuber und Omnivoren untersucht. Dazu wurde ein Bachabschnitt, der mit benthivoren Fischen (Gobio gobio, Barbatula barbatula) besetzt war, mit einem fischfreien Abschnitt über zwei Jahre verglichen. Nach der Identifizierung wichtiger Räuber- und Omnivorenarten anhand ihrer Biomasse und ihrer trophischen Positionen mittels Isotopenanalyse wurde ihr Gesamtfraßdruck abgeschätzt. Invertebrate Prädatoren im Nahrungsnetz des Gauernitzbaches waren Rhyacophila fasciata und Plectrocnemia conspersa (Trichoptera), Isoperla grammatica (Plecoptera) sowie Dugesia gonocephala (Turbellaria). Wichtige Omnivore aufgrund ihrer hohen Biomassen waren Gammarus pulex (Amphipoda) und Hydropsyche spp. (Trichoptera). Anhand eines Mischungsmodells (ISOSOURCE), das auf der Basis der stabilen Isotopensignaturen d13C und d15N die einzelnen Ressourcenanteile einer gemischten Signatur berechnet, konnte das ausgeprägte räuberische Potential der Omnivoren gezeigt werden, da sie einen hohen tierischen Anteil in der Nahrung aufwiesen (20-90 %). Im Nahrungsnetz des Gauernitzbaches waren Omnivore als Räuber damit ebenso bedeutsam wie obligat räuberische Invertebrate. Der Gesamtfraßdruck der invertebraten Räuber und Omnivoren auf die Beuteorganismen (bis 16 g TM m-2 a-1) in der Fischstrecke war höher als der Fraßdruck der benthivoren Fische (~3 g TM m-2 a-1). Dieser Unterschied lag hauptsächlich in höheren täglichen Konsumtionsraten bei ähnlich hohen Biomassen der Invertebraten begründet. Durch die Analyse der stabilen Isotope von Kohlenstoff und Stickstoff konnten die trophischen Beziehungen im Nahrungsnetz des Gauernitzbaches identifiziert werden. Dabei stellten allochthone Kohlenstoffquellen im Frühjahr und Herbst die bedeutendste Ressource für die Primärkonsumenten dar, während autochthone Ressourcen nur im Frühjahr für wenige Herbivore relevant waren. Dies spiegelte sich ebenso in der Nahrung der Omnivoren wider, die neben Organismen hauptsächlich Detritus nutzten. Ebenso zeigten die invertebraten Räuber eine deutliche Abhängigkeit vom allochthonen Kohlenstoff über die Nutzung der Primärkonsumenten. Letale Effekte der Fische auf die räuberischen und omnivoren Populationen im Gauernitzbach fanden hauptsächlich in Abhängigkeit von den bewohnten Habitaten statt. Die Pool-Arten (P. conspersa, D. gonocephala, G. pulex) entwickelten geringere Biomassen in der Fischstrecke, während die Biomassen riffle-bewohnender Invertebraten (R. fasciata, I. grammatica, Hydropsyche spp.) nicht deutlich reduziert wurden. Dabei waren allerdings meist hohe Biomassen in der Fischstrecke weniger stark ausgeprägt als in der fischfreien Strecke. Die Adulten der untersuchten Arten wurden oft stärker letal beeinflusst als die Juvenilstadien. So war die Emergenzproduktion der räuberischen Köcherfliegen und die Abundanz großer und eiertragender Weibchen von G. pulex in der Fischstrecke deutlich verringert. Subletale Effekte der Fische traten im Gauernitzbach in Abhängigkeit von den Ernährungstypen auf. Während die adulten räuberischen Köcherfliegenarten R. fasciata und P. conspersa keine negative Beeinflussung der Fitness und Größe zeigten, emergierte die omnivore Köcherfliege H. instabilis in der Fischstrecke kleiner als in der fischfreien. Außerdem unterschieden sich die Antworten der omnivoren Arten vermutlich aufgrund ihrer unterschiedlichen Lebenszyklen. G. pulex zeigte als ausschließlich aquatisch lebende Art keine subletalen Kosten (unveränderte Fitness) bei Fischanwesenheit wie dies für die omnivore merolimnische Insektenart H. instabilis beobachtet wurde. Die mittlere Fitness der Populationen (gemessen an der Eiproduktion pro Zeit- und Flächeneinheit) war für alle untersuchten invertebraten Räuber und Omnivoren bei Fischanwesenheit deutlich geringer als in der fischfreien Strecke. Für die Räuber (R. fasciata, P. conspersa) und G. pulex waren vermutlich letale Fischeffekte für den geringen Reproduktionserfolg verantwortlich. Dagegen wurde die Fitness der Population von H. instabilis durch die Kombination aus der geringeren individuellen Körpergröße (subletal) und den schwachen letalen Effekten auf die Emergenz reduziert. Der Einfluss der Fische auf die räuberischen Larven von R. fasciata und P. conspersa resultierte in einem Wechsel der Beutezusammensetzung, nicht in der Reduzierung der täglichen Fraßaktivität. Dies war vermutlich auf eine Aktivitätsänderung des Räubers R. fasciata bzw. auf die veränderte Beuteverfügbarkeit für P. conspersa unter Fischeinfluss zurückzuführen. Eine grundsätzliche Verringerung der Fraßaktivität oder der tierischen Anteile in der Nahrung der omnivoren Arten unter dem Einfluss benthivorer Fische wurde nicht festgestellt. Dagegen änderten sich die Relationen der Ressourcen Omnivorer zueinander, so konnte in der fischfreien Strecke eine flexiblere Ressourcennutzung nachgewiesen werden. Die jährliche Gesamtkonsumtion der räuberischen und omnivoren Invertebraten wurde durch die Anwesenheit der benthivoren Fische um 20-50 % reduziert, wofür hauptsächlich fischinduzierte Biomasseänderungen verantwortlich waren. In der fischfreien Strecke konnte dagegen die erhöhte Konsumtion der Invertebraten den fehlenden Fischfraßdruck mehr als kompensieren. Allerdings änderte sich die beutespezifische Konsumtion der Räuber in der fischfreien Strecke durch den verhaltensinduzierten Wechsel in der Beutezusammensetzung. Damit fand ein erhöhter Fraßdruck auf detritusfressende Arten (z.B. Diptera) statt, während die vermuteten Schlüsselarten einer potentiellen Biomanipulation (grazende Eintagsfliegen) in der fischfreien Strecke relativ unbeeinflusst blieben. Da die invertebrate Prädation in Fließgewässern sehr bedeutend sein kann, ist Biomanipulation nur unter Berücksichtigung aller Populationen räuberischer Invertebraten möglich. Nur durch Einstellung einer optimalen (relativ hohen) Biomasse geeigneter benthivorer Fischarten kann eine ausreichende Hemmung der invertebraten Räuber erreicht werden, so dass grazende Arten indirekt profitieren. / Biomanipulation (top-down control of the food webs) in lakes was intensively studied and can today be regarded as an important tool for improvement of the water quality and eutrophication. Similarly, the idea of biomanipulation can be used to reduce eutrophication phenomena in streams. The enhancement of benthic herbivorous key species (e.g. mayflies) by reduction of fish predation pressure could prevent exploding benthic algae biomass. Studies dealing with trophic cascades in lakes suggested that biomass of planktic invertebrate predators is increasing in the absence of planktivorous fish. But until now the importance of invertebrate predation pressure in fishless stream food webs is hardly known. Due to lethal and sublethal fish effects the consumption of predatory invertebrates could be enhanced by the absence of fish to such an extent that biomanipulation could be off target. In general, the proportion of invertebrate predators to the biomass of the benthic community in stream food webs is relatively high. Therefore, invertebrate predation on benthic prey can be as important as vertebrate predation (e.g. by benthivorous fish). Furthermore, omnivory occurs frequently in streams. Therefore, the aim of this thesis was to assess the importance of invertebrate predation in a food web of a small stream (Gauernitzbach) including the obligatory predators as well as the predatory potential of omnivores. Furthermore, the impact of benthivorous fish on population dynamics and feeding behaviour of the most important predators and omnivores was studied. Therefore, a reach stocked with benthivorous fish (Gobio gobio, Barbatula barbatula) was compared with an upstream fishless reach over a two-year period. After identification of important predators and omnivores based on their biomass and trophic position by stable isotope analyses their total consumption was estimated. Rhyacophila fasciata and Plectrocnemia conspersa (Trichoptera), Isoperla grammatica (Plecoptera) as well as Dugesia gonocephala (Turbellaria) were the invertebrate predators in the food web of Gauernitzbach. Because of their high biomasses the most important omnivores were Gammarus pulex (Amphipoda) and Hydropsyche spp. (Trichoptera). Using a mixing model (ISOSOURCE) that estimates the proportions of single resources on a mixed signature on the basis of the stable 13C and 15N isotopes, the predatory potential of both omnivores was pronounced by a high animal food proportion (20-90%). In this stream food web the predation impact of omnivores was comparable to this of obligatory predacous invertebrates. The total invertebrate predation pressure (up to 16 g TM m-2 a-1) in the fish reach was higher than the vertebrate predation pressure (~3 g TM m-2 a-1) mainly caused by the higher daily consumption rates and similar biomass compared to fish. Using stable isotope analyses of carbon and nitrogen the trophic pathways in the food web of Gauernitzbach could be identified. The most important resources for primary consumers in spring and autumn were allochthonous carbon sources. Autochtnonous carbon sources were only utilized by a few herbivores in spring. This was also reflected in the food of omnivores which assimilated mainly detritus beside animal material. Similarly, the predators depended distinctly on the allochthonous pathway by feeding primary consumers. Lethal fish effects on the predatory and omnivorous invertebrate populations were mainly depended on their preferred habitats. Pool-dwelling species (P. conspersa, D. gonocephala, G. pulex) showed decreased biomass in the presence of fish. Contrary, the biomass of riffle-dwelling invertebrates (R. fasciata, I. grammatica, Hydropsyche spp.) was not significantly reduced in the fish reach. Thereby, the highest biomass values in the fish reach could not reach the same level as in the fishless reach. Adults of the investigated species were influenced more lethal than the young stages. Hence, the emergence production of the predatory caddisflies and the abundance of the biggest and gravid females of G. pulex were reduced in the fish reach. Sublethal fish effects existed in dependence on the feeding groups of the invertebrates in Gauernitzbach. Whereas the adult predatory caddisflies R. fasciata and P. conspersa did not show disadvantages in fitness and size, the omnivorous caddisfly H. instabilis emerged with smaller size in the fish reach than in the fishless reach. Different responses of both omnivorous species were observed because of their different life cycles. The exclusively aquatic living G. pulex was not influenced sublethal by fish as the merolimnic insect species H. instabilis. The average fitness of population (measured as egg production per unit time and space) of the studied invertebrate predators and omnivores was distinctly reduced in the fish reach compared to the fishless reach. Probably, lethal fish effects were mainly responsible for the low reproductive success of the predatory species (R. fasciata, P. conspersa) and G. pulex. In contrast, the reduction of population fitness of H. instabilis was caused by the reduced individual adult size (sublethal effect) in combination with the weak lethal effect on emergence. The impact of fish on the feeding behaviour of the predatory larvae of R. fasciata and P. conspersa resulted in a switched prey composition, whereas a reduction in the daily feeding activity was not observed. That was supposed because of changes in the activity pattern by the predator R. fasciata or the changed prey availability for P. conspersa in the fish reach. A general reduction in feeding activity or animal food proportion of the omnivores was not assessed in the presence of fish. In contrast, the relations between the resources switched to a more variable utilisation in the fishless reach. The annual total consumption of invertebrate predators and omnivores was decreased by 20-50% in the fish reach, mainly caused by fish induced biomass reduction. The missing fish consumption could be at least balanced by higher invertebrate consumption in the fishless reach. Contrary, the prey specific consumption of invertebrate predators changed by behavioural-induced switch in the prey composition between the stream reaches. Therefore, a higher predation pressure on detritus feeding species (e.g Diptera) was found in the fishless reach, but not on the supposed key species of the potential biomanipulation (e.g. grazing mayflies). Due to the high importance of invertebrate predation in stream food webs, the aim of biomanipulation can only be reached by considering the populations of predatory invertebrates. Controlling the biomass of suitable benthivorous fish to an optimum (relatively high) level could sufficiently reduce the impact of invertebrate predators leading to an indirect enhancement of grazing species.

The ability of prey to detect and respond appropriately to predator risk is important to overall prey fitness. Many aquatic organisms assess risk through the use of chemical cues that can change with predator diet. Two variable characteristics of diet are: 1. prey type and 2. prey mass. To assess the effect of these two characteristics on the assessment of risk by the mud crab Panopeus herbstii, I exposed mud crabs to the urine of the blue crab Callinectes sapidus fed one of 5 diet treatments: 10g of oyster shell free wet mass, 5g of oyster shell free wet mass, 10g crushed mud crabs, 5g crushed mud crabs, and a mix of 5g of oyster shell free wet mass and 5g crushed mud crab. Effects on P. herbstii foraging were tested in a previously developed bioassay by measuring shrimp consumption over a 4 hour period. I hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a diet of crushed mud crabs than to urine from C. sapidus fed a diet of oysters. I further hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a high mass diet relative to a lower mass diet. Contrary to expectations there was no observed effect of urine on P. herbstii foraging in any of the treatments. Results suggest that bioassay protocol may be unreliable suggesting further replication to determine the difference between this study and previous results. Future studies examining how P. herbstii varies with urine concentration will aid in understanding the ecological scale of this predator cue system. Determining the role of other potential cue sources will improve the predictive abilities of these studies.

No description available.

The objectives of this study were: 1) to compare and contrast the effects of variations in natural densities of two common freshwater predators of crustacean zooplankton, Chaoborus spp. and Mysis relicta; and 2) to determine whether the combined impacts of these predators together differed from their effects when alone. In deep (>10m) mesocosms, additions of Chaoborus and Mysis at natural densities did not result in large changes in zooplankton abundances, lengths, or biomass. Significant decreases in abundance were observed only for Bosmina longirostris and Daphnia spp. In small (~20L) enclosures, higher predator densities caused zooplankton declines over three days. Strong interactive effects between Chaoborus and Mysis were not detected in either the large or small enclosures. This suggests that the combined effects of these two predators can be predicted from their effects determined in isolation.

Lake Winnipeg has been subject to intense eutrophication and invasive species such as Rainbow Smelt and Eubosmina coregoni for the last 40 years. This study demonstrated significant increases in total phosphorus, total nitrogen, chlorophyll-a, overall zooplankton abundance, and specifically Cladocera, between 1969 and the first decade of the 21st century. There were specific basin differences in the long-term changes of the Cladocera community, with the species Chydorus sphaericus and Ceriodaphnia quadrangula negatively affected by high levels of phosphorus, nitrogen and chlorophyll-a. Moreover, long-term change in the Cladocera community composition and abundance throughout the years (1969-2011) during summer was correlated with intensifying eutrophication. Additionally, weak diel vertical migration in the zooplankton community was observed for the first time in Lake Winnipeg.