This study investigates size-selective predation of four groups predaceous aquatic insects: adults and larvae of diving beetles (Dytiscidae), larvae of dragonflies (Odonata) and aquatic true bugs (Heteroptera) using laboratory feeding experiments and relates the results to morphometric data obtained for each predator.

Severe declines in the number of some songbirds over the last 40 years have caused heated debate amongst interested parties. Many factors have been suggested as possible causes for these declines, including an increase in the abundance and distribution of an avian predator, the Eurasian sparrowhawk Accipiter nisus. To test for evidence for a predator effect on the abundance of its prey, we analyse data on 10 species visiting garden bird feeding stations monitored by the British Trust for Ornithology in relation to the abundance of sparrowhawks. We apply Bayesian hierarchical models to data relating to averaged maximum weekly counts from a garden bird monitoring survey. These data are essentially continuous, bounded below by zero, but for many species show a marked spike at zero that many standard distributions would not be able to account for. We use the Tweedie distributions, which for certain areas of parameter space relate to continuous nonnegative distributions with a discrete probability mass at zero, and are hence able to deal with the shape of the empirical distributions of the data. The methods developed in this thesis begin by modelling single prey species independently with an avian predator as a covariate, using MCMC methods to explore parameter and model spaces. This model is then extended to a multiple-prey species model, testing for interactions between species as well as synchrony in their response to environmental factors and unobserved variation. Finally we use a relatively new methodological framework, namely the SPDE approach in the INLA framework, to fit a multi-species spatio-temporal model to the ecological data. The results from the analyses are consistent with the hypothesis that sparrowhawks are suppressing the numbers of some species of birds visiting garden feeding stations. Only the species most susceptible to sparrowhawk predation seem to be affected.

To show how predaceous fish and dragonflies affect benthic community structure, we conducted enclosure experiments in the littoral zone of Bays Mountain Lake, Tennessee. A 'natural' benthic assemblage was subjected to all combinations of 2 densities of 3 predator treatments - small sunfish, 0 or 4/m2; large dragonfly larvae, 0 or 15/m2; and dragonfly eggs at 2 densities, 90 or 900/m2. Treatments were assigned randomly in each of 6 spatiotemporal blocks. Net effects of predation over 4 mo show that small sunfish had 'strong' effects (>50% reduction of densities) on triclads, large daphnid cladocerans, and snails, and 'moderate' effects (50% > reduction > 25%) on small dragonflies and ostracods; all these effects were statistically significant, except the one on ostracods. Large dragonflies had moderate non-significant effects on triclads and ostracods. There were no significant increases in prey density associated with fish predation; but chydorid cladocerans and midge larvae showed 'weak' non-significant increases that might be caused by 'indirect effects' of fish predation on invertebrate predators or grazers. There were only 2 significant interaction terms indicative of 'higher-order interactions': Fish X Dragonfly on the large daphnid Simocephalus, and Fish X Egg Density on snails, which were associated with relatively low probabilities. In both cases, the net effect of dragonflies was to reduce prey densities more when fish were present.

Anthropogenic provisioning of water (water developments) to enhance abundance and distribution of wildlife is a common management practice in arid regions where water is limiting. Despite the long-term and widespread use of water developments, little is known about how they influence distribution, competition dynamics, and behavior of native species. To elucidate the potential influences of water developments on native species, we tested hypotheses concerning the occurrence and behavior of native kit foxes (Vulpes macrotis). First, we tested the indirect effect of water hypothesis (IEWH) which proposes that water developments negatively affect the arid-adapted kit fox by enabling a water-dependent competitor (i.e., coyote; Canis latrans) to expand distribution in arid landscapes. We tested the two predictions of the IEWH (i.e., coyotes will visit areas with water more frequently and kit foxes will avoid coyotes) and evaluated relative use of water by canids in the Great Basin and Mojave Deserts from 2010 to 2012. We established scent stations in areas with (wet) and without (dry) water and monitored visitation by canids to scent stations and water sources using infrared-triggered cameras. There was no difference in the proportions of visits to scent stations in wet or dry areas by coyotes or kit foxes at either study area. There was no correlation between visits to scent stations by coyotes and kit foxes. Visitation to water sources was not different for coyotes between study areas, but kit foxes visited water sources more in Mojave than Great Basin. The intense visitation to water by kit foxes in Mojave challenges our understanding that this species does not readily drink water. Our results did not support the IEWH in the Great Basin or Mojave Deserts for these two canids. Second, we tested three hypotheses that have been proposed to explain spatial variation in vigilance behavior. The predator-vigilance hypothesis (PVH) proposes that prey increase vigilance where there is evidence of predators. The visibility-vigilance hypothesis (VVH) suggests that prey increase vigilance where detection of predators is impeded or visibility is obstructed. The refuge-vigilance hypothesis (RVH) proposes that prey may perceive areas with low visibility (greater cover) as refuges and decrease vigilance. We evaluated support for these hypotheses using the kit fox, a solitary carnivore subject to intraguild predation, as a model. From 2010 to 2012, we used infrared-triggered cameras to record video of kit fox behavior at water developments in the Mojave Desert. The RVH explained more variation in vigilance behavior of kit foxes than the other two hypotheses (AICc model weight = 0.37). Kit foxes were less vigilant at water developments with low overhead cover (refuge) obstructing visibility. Based on our results, the PVH and VVH may not be applicable to all species of prey. Solitary prey, unlike gregarious prey, may use areas with concealing cover to maximize resource acquisition and minimize vigilance.

<p>Diss. (sammanfattning) Umeå : Univ., härtill 6 uppsatser.</p> / digitalisering@umu

A major challenge in agroecosystems is how to manage the systems so that it reduces crop pests and enhances natural pest control. This thesis investigates patterns of crop pests and top-down effects of birds and arthropod predators in relation to land-use composition across spatial scales. In paper (I) I examined the crop distribution and land-use types in relation to the crop raiding patterns in 15 transectsin sites close to and far from forests along with a questionnaire survey at household level. I found severe crop raiding close to forests, but it had no impact on crop composition growing between the two sites. In paper (II) I examined the effect of forest and tree cover, at local and landscape scales, on the abundance of arthropod predators by collecting specimens from 40 home gardens. My result showed higher abundance of arthropod predators when either the home garden or the surroundings had a high tree-cover, compared to when tree-cover at both scales was similarly either high or low. In paper (III) I investigated the effect of excluding birds and arthropod predators on leaf damage on rape seed in 26 home gardens. I found stronger top-down impacts from arthropod predators on crop pests in tree-poor gardens than in tree-rich gardens. There was no effect of birds. In paper (IV) I explored the effect of landscape complexity on bird and arthropod predation using plasticine caterpillars in 36 home gardens across landscapes. The rate of arthropod predation on caterpillars was higher in simple than in complex landscapes. The rate of bird predation did not vary between complex and simple landscapes. In simple landscapes, arthropod predation was higher than that of birds. The overall results suggest that simplified gardens/landscapes still have enough habitat heterogeneity to support arthropod predators for the significant top-down controlling effect on crop pests. However, I did not find clear effect of complexityon the top-down effect of birds. / <p>At the time of the doctoral defence the following papers were unpublished and had a status as follows: Paper 1: Manuscript; Paper 3: Manuscript; Paper 4: Manuscript</p>

The principle of competitive exclusion holds that the strongest competitor for a single resource can exclude other species. Yet in many systems, more similar species appear to stably coexist than the small number of limiting resources. Understanding how and when similar species can stably coexist has taken on new urgency in managing biological invasions and their ecological impacts. Recent theoretical advances emphasize the importance of predators in determining coexistence. The effects of predators, however, can be mediated by behavioral changes induced in their prey as well as by their lethality. In this dissertation, I ask how considering multiple trophic levels changes our understanding of how a grass invasion (Pennisetum ciliare) affects species diversity and dynamics in southeastern Arizona. In considering interactions with plant consumers, and with the predators of those consumers, this research reveals more general ecological processes that determine species diversity across biological communities. I first present evidence from a grass removal experiment in the field that shows increased emergence and short-term survival of native perennial plants without grass. This is consistent with Pennisetum ciliare causing the observed concurrent decline in native plant abundance following invasion. I then present results from greenhouse and field studies consistent with that suppression of native plants being driven primarily through resource competition rather than increased rodent granivory. Granivorous rodents do not solely function as consumers, however, because they cache their harvested seeds in shallow scatter-hoards, from which seeds can germinate. Rodents thus act also as seed dispersers in a context-dependent mutualism. The primary granivores in areas invaded by Pennisetum ciliare are pocket mice (genus Chaetodipus), which have a well-studied tendency to concentrate their activity under plant cover to avoid predation by owls. Because the dense canopy of the grass may provide safer refuge, I hypothesized the pocket mice may be directly dispersing native seeds closer to the base of the invasive grass. Such a behavior could increase the competitive effect of the grass on native plant species, further driving the impacts of the invasion. By offering experimental seeds dusted in fluorescent powder and tracking where the seeds were cached, I show that rodents do preferentially cache experimental seeds under the grass. This dispersal interaction may be more general to plant interactions with seed-caching rodents across semi-arid regions that are experiencing plant invasions. Finally, I ask how the predator avoidance behavior exhibited by these rodents affects their ability to coexist with one another. Not only could their diversity affect that of the plant community, but the effects of plant invasions can cascade through other trophic levels. Theoretical understanding of how similar predator avoidance strategy alters coexistence had not yet been developed, however. Instead of a field study, therefore, I modified a general consumer-resource model with three trophic levels to ask whether avoidance behavior by the middle trophic level alters the ability of those species to coexist. I found that more effective avoidance behavior, or greater safety for less cost, increased the importance of resource partitioning in determining overall niche overlap. Lowering niche overlap between two species promotes their coexistence in the sense that their average fitness can be more different and still permit coexistence. These results provide novel understanding of behavioral modifications to population dynamics in multi-trophic coexistence theory applicable to this invasion and more broadly.

No description available.

Multi-species functional response models are required to model the predation of generalist preda- tors, which consume more than one prey species. In chapter 2, a new model for the multi-species functional response is presented. This model can describe generalist predators that exhibit func- tional responses of Holling type II to some of their prey and of type III to other prey. In chapter 3, I review some of the theoretical distinctions between Bayesian and frequentist statistics and show how Bayesian statistics are particularly well-suited for the fitting of functional response models because uncertainty can be represented comprehensively. In chapters 4 and 5, the multi- species functional response model is fitted to field data on two generalist predators: the hen harrier Circus cyaneus and the harp seal Phoca groenlandica. I am not aware of any previous Bayesian model of the multi-species functional response that has been fitted to field data. The hen harrier's functional response fitted in chapter 4 is strongly sigmoidal to the densities of red grouse Lagopus lagopus scoticus, but no type III shape was detected in the response to the two main prey species, field vole Microtus agrestis and meadow pipit Anthus pratensis. The impact of using Bayesian or frequentist models on the resulting functional response is discussed. In chapter 5, no functional response could be fitted to the data on harp seal predation. Possible reasons are discussed, including poor data quality or a lack of relevance of the available data for informing a behavioural functional response model. I conclude with a comparison of the role that functional responses play in behavioural, population and community ecology and emphasise the need for further research into unifying these different approaches to understanding predation with particular reference to predator movement. In an appendix, I evaluate the possibility of using a functional response for inferring the abun- dances of prey species from performance indicators of generalist predators feeding on these prey. I argue that this approach may be futile in general, because a generalist predator's energy intake does not depend on the density of any single of its prey, so that the possibly unknown densities of all prey need to be taken into account.

The decline in range and density of frugivorous birds worldwide could have consequences for the functioning of ecosystem processes such as seed dispersal. I endeavoured to determine the effects of bird declines on seed dispersal and assess the current status of dispersal in New Zealand. My first aim was to determine whether decreased bird density can reduce competitive interactions between birds, leading to birds concentrating on higher-reward fruit species. I measured fruit removal rates and fruit preferences of birds in Canterbury forest remnants with high (Hay and Prices) and low (Lords and Kaituna) bird densities. Removal rates of Melicytus ramiflorus and Coprosma spp. were lower at low-bird sites, and the size of this effect was greater for low-reward plant species. Coprosma areolata (a low-sugar fruit) had limited dispersal at Kaituna (59% fruit removed by end of 2012 season), compared with 92% removal at Hay, whereas 99-100% of M. ramiflorus and C. robusta (higher-sugar) fruits were removed at both sites. My second aim was to determine whether seed dispersal by introduced possums (Trichosurus vulpecula) can compensate for bird declines. I analysed seeds present in possum and bird faecal samples collected at Kowhai Bush, Kaikoura. Possums dispersed <3% of the total seeds, much less than bellbirds (21%), silvereyes (16%), song thrushes (33%) and blackbirds (28%). Possums also destroyed approximately 15% of seeds found in faeces, reduced the germination of gut-passed C. robusta seed to half of that from bird faeces (30% vs. 60-70%), and did not swallow fruits any larger than those moved by the much smaller birds (c. 7 mm diameter). My third aim was to determine the relationship between percentage of fruits dispersed and distance from parent tree for three large-seeded trees; Beilschmiedia tawa, Elaeocarpus dentatus, and Prumnopitys ferruginea. I fitted dispersal kernels to the observed dispersal distances out to 50 m for both undispersed whole fruits and seeds consumed by a bird, and found that dispersal quantity below the parent tree strongly underestimates total dispersal quantity. The average percentage dispersed overall was 81% for B. tawa, 75% for E. dentatus and 91% for P. ferruginea, and for all species finding only 11-18% clean seeds under the parent tree would correspond to an overall percent dispersed of at least 50% of the whole seed crop. My final aim was to determine the consequences of dispersal failure for recruitment in three plant species; Elaeocarpus hookerianus, Ripogonum scandens, and C. robusta. I compared the fate of dispersed and undispersed seeds in a manipulative field experiment. Exclusion of mammals plus removal of fruit pulp had the greatest effect on survival, while the combined effects of dispersal failure (under parent, high density, whole fruit) and inclusion of mammals decreased the number of live seedlings present at the final count by 75-92%. Overall, most native plants were receiving adequate dispersal and there was little evidence for strong risks to plant regeneration from dispersal failure, despite the reduced bird densities and ongoing negative effects of introduced mammals in New Zealand.