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 / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Manuscript.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.

In our ever more populated world, the rapid expansion and intensification of agriculture is driving worldwide biodiversity loss, and the interactions between production landscapes and wildlife conservation are becoming increasingly important. Farming systems depend on ecosystem services such as biological control, while conservationists are calling for the establishment of conservation initiatives in non-preserve landscapes. Despite this, the goals of agriculture and the goals of predator-conservation are rarely mutual. Here, I demonstrate one of the first examples of a mutually beneficial scenario between agriculture and predator conservation. I used, as a case study, a reintroduction project that translocated individuals of the threatened New Zealand falcon (Falco novaeseelandiae) from the hills of Marlborough into vineyards, to determine if predators can survive within an agricultural landscape while simultaneously providing that landscape with biological control services. Examples of vertebrates providing biological control to agriculture are rare. I show that the presence of falcons in vineyards caused an economically important reduction in grape damage worth over US $230/ ha. Falcon presence caused a 78- 83% reduction in the number of introduced European pest birds, which resulted in a 95% reduction in the damage caused by these species. Falcon presence did not cause a reduction in the abundance of the native silvereye (Zosterops lateralis), but did halve the damage caused by this species. To assess the conservation value of the falcon translocations, I used remote videography, direct observations and prey analysis to measure the behavioural changes associated with the relocation of falcons from their natural habitat in the hills and into vineyards. Falcons in vineyard nests had higher nest attendance, higher brooding rates, and higher feeding rates than falcons in hill nests. Additionally, parents in vineyard nests fed their chicks a greater amount of total prey and larger prey items compared to parents in hill nests. I also found an absence of any significant diet differences between falcons in hill and vineyard habitats, suggesting that the latter may be a suitable alternative habitat for falcons. Because reintroduced juvenile falcons were released in areas devoid of adult falcons, it was possible that they were missing essential training normally provided by their parents. I used direct observations to demonstrate that the presence of siblings had similar effects to the presence of parents on the development of juvenile behaviour, with individuals flying, hunting, and playing more often when conspecifics were present. Finally, through the use of artificial nests and remote videography, I identified that falcons nesting in vineyards are likely to suffer lower predation rates. I also found that falcons in vineyards are predated by a less dangerous suite of animals (such as hedgehogs, Erinaceus europaeus, and avian predators), than their counterparts in the hills, which are predated by more voracious species (such as stoats, Mustela erminea, and feral cats, Felis catus). The work presented in this thesis has also added to the current knowledge of New Zealand falcon breeding behaviour, prey preferences, and behavioural development. Although agricultural regions globally are rarely associated with raptor conservation, and the ability of raptors to control the pests of agricultural crops has not been previously quantified, these results suggest that translocating New Zealand falcons into vineyards has potential for both the conservation of this species, and for providing biological control services to agriculture

This thesis deals with behavioural strategies used by caddis larvae in pairwise contests and when selecting microhabitats. Effects of caddis larvae on survival and habitat selection of other insect taxa have also been studied. The behaviours used by Arctopsyche ladogensis larvae fighting for nets, and Agrypnia pagetana larvae fighting for cases, agreed well with predictions from the sequential assessment game, which is an ESS model of animal fighting behaviour. Establishment by net-spinning Hydropsyche siltalai larvae on artificial substrates was highest at intermediate densities of residents. Emigration/mortality was density independent, and it was higher at a poor site (low food availability) than at a rich site. Establishment was unaffected by site quality. Growth was density dependent because larvae in upstream positions reduced both current velocity and concentration of food particles for larvae in downstream positions. A field experiment involved manipulations of the density of H. siltalai larvae and their nets in a lake outlet stream. H. siltalai larvae affected all abundant taxa, but the mechanism involved varied between taxa. Rhyacophila nubila (Trichoptera) and chironomid larvae benefited from the presence of H. siltalai nets. Negative effects on nymphs of the mayfly Ephemerella ignita were due to predation by H. siltalai larvae, while a combination of predation and increased emigration in response to nets depressed densities of Simulium truncatum blackfly larvae. / <p>Diss. (sammanfattning) Umeå : Umeå universitet, 1992, härtill 5 uppsatser</p> / digitalisering@umu