The importance of interaction strength for food web dynamics and ecosystem functioning

Global biodiversity is eroding at an alarming rate, through a combination of anthropogenic disturbance and environmental change. Ecological communities are bewildering in their complexity. Experimental ecologists strive to understand the mechanisms that drive the stability and structure of these complex communities in a bid to inform nature conservation and management. Two fields of research have had high profile success at developing theories related to these stabilising structures and testing them through controlled experimentation. Biodiversity-ecosystem functioning (BEF) research has explored the likely consequences of biodiversity loss on the functioning of natural systems and the provision of important ecosystem services. Empirical tests of BEF theory often consist of simplified laboratory and field experiments, carried out on subsets of ecological communities. Such experiments often overlook key information relating to patterns of interactions, important relationships, and fundamental ecosystem properties. The study of multi-species predator-prey interactions has also contributed much to our understanding of how complex systems are structured, particularly through the importance of indirect effects and predator suppression of prey populations. A growing number of studies describe these complex interactions in detailed food webs, which encompass all the interactions in a community. This has led to recent calls for an integration of BEF research with the comprehensive study of food web properties and patterns, to help elucidate the mechanisms that allow complex communities to persist in nature. This thesis adopts such an approach, through experimentation at Lough Hyne marine reserve, in southwest Ireland. Complex communities were allowed to develop naturally in exclusion cages, with only the diversity of top trophic levels controlled. Species removals were carried out and the resulting changes to predator-prey interactions, ecosystem functioning, food web properties, and stability were studied in detail. The findings of these experiments contribute greatly to our understanding of the stability and structure of complex natural communities.

Strong preference for decapod prey by the western rock lobster Panulirus cygnus

Western rock lobsters, Panulirus cygnus are an abundant benthic consumer distributed along the temperate west coast of Australia and constitute the largest single species fishery in Australia. As a dominant consumer, it is important to understand their predator-prey interactions as they can potentially exert strong trophic effects, and may influence ecosystem function as seen in other spiny lobster species. While previous field studies have focused on the diet composition of P. cygnus, this study investigated their preference for various benthic invertebrate prey to better understand the likely predator-prey interactions of P. cygnus. Prey preferences of small sub-legal juvenile lobsters, as well as medium and large legal-sized mature lobsters were investigated using laboratory feeding trials to identify size-associated differences in lobster prey preference. Handling time and diet quality were investigated to estimate energetic cost and gain from consuming different prey which may explain prey choice by lobsters. It was found that large lobsters preferred crabs and mussels while medium and small lobsters preferred crabs over mussels, gastropods, and sea urchins. This suggests that strong predator-prey interactions between P. cygnus and crabs may occur in the wild.

Scaling of Food-Web Properties with Diversity and Complexity Across Ecosystems

Trophic scaling models describe how topological food-web properties such as the number of predator prey links scale with species richness of the community. Early models predicted that either the link density (i.e. the number of links per species) or the connectance (i.e. the linkage probability between any pair of species) is constant across communities. More recent analyses, however, suggest that both these scaling models have to be rejected, and we discuss several hypotheses that aim to explain the scale dependence of these complexity parameters. Based on a recent, highly resolved food-web compilation, we analysed the scaling behaviour of 16 topological parameters and found significant power law scaling relationships with diversity (i.e. species richness) and complexity (i.e. connectance) for most of them. These results illustrate the lack of universal constants in food-web ecology as a function of diversity or complexity. Nonetheless, our power law scaling relationships suggest that fundamental processes determine food-web topology, and subsequent analyses demonstrated that ecosystem-specific differences in these relationships were of minor importance. As such, these newly described scaling relationships provide robust and testable cornerstones for future structural food-web models.

Functional responses of the intertidal amphipod Echinogammarus marinus: effects of prey supply, model selection and habitat complexity.

The influence of predation in structuring ecological communities can be informed by examining the shape and magnitude of the functional response of predators towards prey. We derived functional responses of the ubiquitous intertidal amphipod Echinogammarus marinus towards one of its preferred prey species, the isopod Jaera nordmanni. First, we examined the form of the functional response where prey were replaced following consumption, as compared to the usual experimental design where prey density in each replicate is allowed to deplete. E. marinus exhibited Type II functional responses, i.e. inversely density-dependent predation of J. nordmanni that increased linearly with prey availability at low densities, but decreased with further prey supply. In both prey replacement and non-replacement experiments, handling times and maximum feeding rates were similar. The non-replacement design underestimated attack rates compared to when prey were replaced. We then compared the use of Holling’s disc equation (assuming constant prey density) with the more appropriate Rogers’ random predator equation (accounting for prey depletion) using the prey non-replacement data. Rogers’ equation returned significantly greater attack rates but lower maximum feeding rates, indicating that model choice has significant implications for parameter estimates. We then manipulated habitat complexity and found significantly reduced predation by the amphipod in complex as opposed to simple habitat structure. Further, the functional response changed from a Type II in simple habitats to a sigmoidal, density-dependent Type III response in complex habitats, which may impart stability on the predator−prey interaction. Enhanced habitat complexity returned significantly lower attack rates, higher handling times and lower maximum feeding rates. These findings illustrate the sensitivity of the functional response to variations in prey supply, model selection and habitat complexity and, further, that E. marinus could potentially determine the local exclusion and persistence of prey through habitat-mediated changes in its predatory functional responses.

Prey field switching based on preferential behaviour can induce Lévy flights

Using the foraging movements of an insectivorous bat, Myotis mystacinus, we describe temporal switching of foraging behaviour in response to resource availability. These observations conform to predictions of optimized search under the Lévy flight paradigm. However, we suggest that this occurs as a result of a preference behaviour and knowledge of resource distribution. Preferential behaviour and knowledge of a familiar area generate distinct movement patterns as resource availability changes on short temporal scales. The behavioural response of predators to changes in prey fields can elicit different functional responses, which are considered to be central in the development of stable predator-prey communities. Recognizing how the foraging movements of an animal relate to environmental conditions also elucidates the evolution of optimized search and the prevalence of discrete strategies in natural systems. Applying techniques that use changes in the frequency distribution of movements facilitates exploration of the processes that underpin behavioural changes. © 2012 The Author(s) Published by the Royal Society. All rights reserved.

Depredación organizacional

El concepto de depredación en el ámbito de la organización es comúnmente asociado al deterioro de sus recursos, pero al abordarlo desde la ecología de poblaciones, el resultado es un enfoque centrado en las relaciones entre los actores de la organización y la interacción de tres variables principales: el poder de los actores, la intimidad entre ellos y la letalidad de sus actuaciones; estas dos últimas solicitadas en préstamo desde la clasificación funcional propuesta por la ecología de poblaciones y claves en la relación presa-depredador. El documento presenta una aproximación para aprovechar estos conocimientos disponibles en la ecología y argumenta que su estudio facilita la comprensión de los estilos de estrategia y dirección en la organización.

Temporal and spatial variability of mobile fauna on a submarine cliff and boulder scree complex: a community in flux

The mobile component of a community inhabiting a submarine boulder scree/cliff was investigated at Lough Hyne, Ireland at dawn, midday, dusk and night over a 1-week period. Line transects (50 m) were placed in the infralittoral (6 m) and circumlittoral (18 m) zones and also the interface between these two zones (12 m). The dominant mobile fauna of this cliff consisted of echinoderms (6 species), crustaceans (10 species) and fish (23 species). A different component community was identified at each time/depth interval using Multi-Dimensional Scaling (MDS) even though both species diversity (Shannon-Wiener indices) and richness (number of species) remained constant. These changes in community composition provided indirect evidence for migration by these mobile organisms. However, little evidence was found for migration between different zones with the exception of the several wrasse species. These species were observed to spend the daytime foraging in the deeper zone, but returned to the upper zone at night presumably for protection from predators. For the majority of species, migration was considered to occur to cryptic habitats such as holes and crevices. The number of organisms declined during the night, although crustacean numbers peaked, while fish and echinoderms were most abundant during day, possibly due to predator-prey interactions. This submarine community is in a state of flux, whereby, community characteristics, including trophic and energetic relationships, varied over small temporal (daily) and spatial (m) scales.

Ratio-dependent response of a temperate Australian estuarine system to sustained nitrogen loading

Classical resource- and the less studied ratio-dependent models of predator–prey relationships provide divergent predictions as to the sustained ecological effects of bottom-up forcing. While resource-dependent models, which consider only instantaneous prey density in modelling predator responses, predict community responses that are dependent on the number of trophic levels in a system, ratio-dependent models, which consider the number of prey per consumer, predict proportional increase in each level irrespective of chain length. The two models are only subtly different for systems with two or three trophic levels but in the case of four trophic levels, predict opposite effects of enrichment on primary producers. Despite the poor discriminatory power of tests of the models in systems with two or three trophic levels, field tests in estuarine and marine systems with four trophic levels have been notably absent. Sampling of phytoplankton, macroinvertebrates, invertebrate-feeding fishes, piscivorous fishes in Kooloonbung Creek, Hastings River estuary, eastern Australia, subject to over 20 years of sewage discharge, revealed increased abundances in all four trophic levels at the disturbed location relative to control sites. Increased abundance of phytoplankton at the disturbed site was counter to the predictions of resource-dependent models, which posit a reduction in the first trophic level in response to enrichment. By contrast, the increase in abundance of this first trophic level and the proportionality of increases in abundances of each of the four trophic groups to nitrogen loading provided strong support for ratio dependency. This first evidence of ratio dependence in an estuarine system with four trophic levels not only demonstrates the applicability of ecological theory which seeks to simplify the complexity of systems, but has implications for management. Although large nutrient inputs frequently induce mortality of invertebrates and fish, we have shown that smaller inputs may in fact enhance biomass of all trophic levels.

Population consequences of a predator-induced habitat shift by trout in whole-lake experiments

In a replicated whole-lake experiment, we (a) tested for the existence of a flexible habitat shift in response to predator presence in age-0 rainbow trout (Oncorhynchus mykiss) at risk of cannibalism and (b) evaluated the population-level consequences of habitat shifts in terms of growth and survival over their first growing season. Daphnid food and adult trout predators were substantially more abundant in pelagic than in littoral habitats. Age-0 trout used all habitats in populations without adult trout predators, whereas age-0 trout were observed only in the less profitable littoral habitat in populations with adult trout. Consequently, mean fall mass of age-0 trout in the presence of predators was almost half that observed in populations without adult trout. Despite the shift in habitat use, age-0 trout experienced 90% mortality when adult trout predators were present, in comparison to only 36% mortality when absent. We conclude that the commonly observed habitat shifts by fish at risk of predation, observed at smaller scales, do in fact occur at the whole-system scale over long time intervals. These results suggest that fish are able to perceive risk at large spatial scales and thus take advantage of profitable (but normally risky) habitats when predators are absent, or move to less profitable refuge habitats when predators are present.

How well do predators adjust to climate-mediated shifts in prey distribution? A study on Australian water pythons

Climate change can move the spatial location of resources critical for population viability, and a species' resilience to such changes will depend upon its ability to flexibly shift its activities away from no-longer-suitable sites to exploit new opportunities. Intuition suggests that vagile predators should be able to track spatial shifts in prey availability, but our data on water pythons (Liasis fuscus) in tropical Australia suggest a less encouraging scenario. These pythons undergo regular long-range (to >10 km) seasonal migrations to follow flooding-induced migrations by their prey (native dusky rats, Rattus colletti). However, when an extreme flooding event virtually eliminated rats for a three-year period, the local pythons did not disperse despite the presence of abundant rats only 8 km away; instead, many pythons starved to death. This inflexibility suggests that some vagile species that track seasonally migrating prey may do so by responding to habitat attributes that have consistently predicted prey availability over evolutionary time, rather than reacting to proximate cues that signal the presence of prey per se. A species' vulnerability to climate change will be increased by an inability to shift its activities away from historical sites toward newly favorable areas.

Preference for feeding at habitat edges declines among juvenile blue crabs as oyster reef patchiness increases and predation risk grows

Both habitat patchiness and behaviorally-mediated indirect effects (BMIEs; predator- induced changes in prey behavior that affect the prey's resources) are important in many food webs, but the relationships between these 2 factors have yet to be investigated. To explore effects of habitat patchiness and variation in perceived risk of predation on food-web dynamics, we conducted a factorial experiment in a model aquatic food chain of predator-prey-resource using 2 contrasting predators (adult blue crab Callinectes sapidus and toad fish Opsanus tau), juvenile blue crab as prey, and mussel Geukensia demissa as resource. Both predator presence and habitat patchiness influenced the prey's preference for consuming resources at patch edges instead of interiors. The preference of prey for consuming resources at habitat edges was 4 times stronger in continuous oyster reef habitat than in smaller habitat patches. This suggests that interior resources in continuous habitat experience a refuge from consumption, but this refuge is largely lost in patchy habitat. The mere presence of predators reduced the prey's preference for consuming resources at habitat edges. This BMIE was significant for the ambush predator (toadfish) and the treatment containing both predators, but not for the actively hunting predator (adult blue crab). We conclude that habitat patchiness and predator presence can jointly affect resource distribution by inducing shifts in prey foraging behavior, revealing a need to incorporate BMIEs into habitat fragmentation studies. This conclusion has broad and growing relevance as anthropogenic factors increasingly modify predator abundances and fragment coastal habitats. © Inter-Research 2012.

Stomach fullness modulates prey size choice in the frillfin goby, Bathygobius soporator

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Foraging behaviour by an intraguild predator blowfly, Chrysomya albiceps (Diptera : Calliphoridae)

Optimal foraging theory assumes that predators use different prey types to maximize their rate of energetic gain. Studies focusing on prey preference are important sources of information to understand the foraging dynamics of Chrysomya albiceps. The purpose of this investigation is to determine the influence of larval starvation in C. albiceps on the predation rate of different prey blowfly species and instars under laboratory conditions. Our results suggest that C. albiceps prefers Cochliomyia macellaria larvae to Chrysomya megacephala under non-starvation and starvation conditions. Nevertheless, predators gained more weight consuming C. macellaria. This result suggests that C. albiceps profit more in consuming C. macellaria rather than C. megacephala. The foraging behaviour displayed by C. abiceps on their prey and the consequences for the blowfly community are also discussed.

Are you experienced? Predator type and predator experience trade-offs in relation to tadpole mortality rates

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)