How does global change affect the strength of trophic interactions?

Recent research has generally shown that a small change in the number of species in a food web can have consequences both for community structure and ecosystem processes. However 'change' is not limited to just the number of species in a community, but might include an alteration to such properties as precipitation, nutrient cycling and temperature, all of which are correlated with productivity. Here we argue that predicted scenarios of global change will result in increased plant productivity. We model three scenarios of change using simple Lotka-Volterra dynamics, which explore how a global change in productivity might affect the strength of local species interactions and detail the consequences for community and ecosystem level stability. Our results indicate that (i) at local scales the average population size of consumers may decline because of poor quality food resources, (ii) that the strength of species interactions at equilibrium may become weaker because of reduced population size, and (iii) that species populations may become more variable and may take longer to recover from environmental or anthropogenic disturbances. At local scales interaction strengths encompass such properties as feeding rates and assimilation efficiencies, and encapsulate functionatty important information with regard to ecosystem processes. Interaction strengths represent the pathways and transfer of energy through an ecosystem. We examine how such local patterns might be affected given various scenarios of 'global change' and discuss the consequences for community stability and ecosystem functioning. (C) 2004 Elsevier GmbH. All rights reserved.

Modelling the influence of biotic factors on species distribution patterns

Biotic interactions can have large effects on species distributions yet their role in shaping species ranges is seldom explored due to historical difficulties in incorporating biotic factors into models without a priori knowledge on interspecific interactions. Improved SDMs, which account for biotic factors and do not require a priori knowledge on species interactions, are needed to fully understand species distributions. Here, we model the influence of abiotic and biotic factors on species distribution patterns and explore the robustness of distributions under future climate change. We fit hierarchical spatial models using Integrated Nested Laplace Approximation (INLA) for lagomorph species throughout Europe and test the predictive ability of models containing only abiotic factors against models containing abiotic and biotic factors. We account for residual spatial autocorrelation using a conditional autoregressive (CAR) model. Model outputs are used to estimate areas in which abiotic and biotic factors determine species’ ranges. INLA models containing both abiotic and biotic factors had substantially better predictive ability than models containing abiotic factors only, for all but one of the four species. In models containing abiotic and biotic factors, both appeared equally important as determinants of lagomorph ranges, but the influences were spatially heterogeneous. Parts of widespread lagomorph ranges highly influenced by biotic factors will be less robust to future changes in climate, whereas parts of more localised species ranges highly influenced by the environment may be less robust to future climate. SDMs that do not explicitly include biotic factors are potentially misleading and omit a very important source of variation. For the field of species distribution modelling to advance, biotic factors must be taken into account in order to improve the reliability of predicting species distribution patterns both presently and under future climate change.

The biology and ecology of the ocean sunfish Mola mola: a review of current knowledge and future research perspectives

Relatively little is known about the biology and ecology of the world's largest (heaviest) bony fish, the ocean sunfish Mola mola, despite its worldwide occurrence in temperate and tropical seas. Studies are now emerging that require many common perceptions about sunfish behaviour and ecology to be re-examined. Indeed, the long-held view that ocean sunfish are an inactive, passively drifting species seems to be entirely misplaced. Technological advances in marine telemetry are revealing distinct behavioural patterns and protracted seasonal movements. Extensive forays by ocean sunfish into the deep ocean have been documented and broad-scale surveys, together with molecular and laboratory based techniques, are addressing the connectivity and trophic role of these animals. These emerging molecular and movement studies suggest that local distinct populations may be prone to depletion through bycatch in commercial fisheries. Rising interest in ocean sunfish, highlighted by the increase in recent publications, warrants a thorough review of the biology and ecology of this species. Here we review the taxonomy, morphology, geography, diet, locomotion, vision, movements, foraging ecology, reproduction and species interactions of M. mola. We present a summary of current conservation issues and suggest methods for addressing fundamental gaps in our knowledge.

Press perturbations and indirect effects in real food webs

The prediction of the effects of disturbances in natural systems is limited by the general lack of knowledge on the strength of species interactions, i.e., the effect of one species on the population growth rate of another, and by the uncertainty of the effects that may be manifested via indirect pathways within the food web. Here we explored the consequences of changes in species populations for the remaining species within nine exceptionally well-characterized empirical food webs, for which, unlike the vast majority of other published webs, feeding links have been fully quantied. Using the inverse of the Jacobian matrix, we found that perturbations to species with few connections have larger net effects (considering both direct and indirect pathways between two species) on the rest of the food web than do disturbances to species that are highly connected. For 40% of predator-prey links, predators had positive net effects on prey populations, due to the predominance of indirect interactions. Our results highlight the fundamental, but often counterintuitive, role of indirect effects for the maintenance of food web complexity and biodiversity.

Global change alters the stability of food webs

Recent research has generally shown that a small change in the number of species in a food web can have consequences both for community structure and ecosystem processes. However 'change' is not limited to just the number of species in a community, but might include an alteration to such properties as precipitation, nutrient cycling and temperature. How such changes might affect species interactions is important, not just through the presence or absence of interactions, but also because the patterning of interaction strengths among species is intimately associated with community stability. Interaction strengths encompass such properties as feeding rates and assimilation efficiencies, and encapsulate functionally important information with regard to ecosystem processes. Interaction strengths represent the pathways and transfer of energy through an ecosystem. We review the best empirical data available detailing the frequency distribution of interaction strengths in communities. We present the underlying (but consistent) pattern of species interactions and discuss the implications of this patterning. We then examine how such a basic pattern might be affected given various scenarios of 'change' and discuss the consequences for community stability and ecosystem functioning.

Plant community assembly at small scales: spatial versus environmental factors in a central European grassland

Dispersal limitation and environmental conditions are crucial drivers of plant species distribution and establishment. As these factors operate at different spatial scales, we asked: Do the environmental factors known to determine community assembly at broad scales operate at fine scales (few meters)? How much do these factors account for community variation at fine scales? In which way do biotic and abiotic interactions drive changes in species composition? We surveyed the plant community within a dry grassland along a very steep gradient of soil characteristics like pH and nutrients. We used a spatially explicit sampling design, based on three replicated macroplots of 15x15, 12x12 and 12x12 meters in extent. Soil samples were taken to quantify several soil properties (carbon, nitrogen, plant available phosphorus, pH, water content and dehydrogenase activity as a proxy for overall microbial activity). We performed variance partitioning to assess the effect of these variables on plant composition and statistically controlled for spatial autocorrelation via eigenvector mapping. We also applied null model analysis to test for non-random patterns in species co-occurrence using randomization schemes that account for patterns expected under species interactions. At a fine spatial scale, environmental factors explained 18% of variation when controlling for spatial autocorrelation in the distribution of plant species, whereas purely spatial processes accounted for 14% variation. Null model analysis showed that species spatially segregated in a non-random way and these spatial patterns could be due to a combination of environmental filtering and biotic interactions. Our grassland study suggests that environmental factors found to be directly relevant in broad scale studies are present also at small scales, but are supplemented by spatial processes and more direct interactions like competition.

Does chemical composition of individual Scots pine trees determine the biodiversity of their associated ground vegetation?

Despite plant secondary metabolites being major determinants of species interactions and ecosystem processes, their role in the maintenance of biodiversity has received little attention. In order to investigate the relationship between chemical and biological diversity in a natural ecosystem, we considered the impact of chemical diversity in individual Scots pine trees (Pinus sylvestris) on species richness of associated ground vegetation. Scots pine trees show substantial genetically determined constitutive variation between individuals in concentrations of a group of secondary metabolites, the monoterpenes. When the monoterpenes of particular trees were assessed individually, there was no relationship with species richness of associated ground flora. However, the chemical diversity of monoterpenes of individual trees was significantly positively associated with the species richness of the ground vegetation beneath each tree, mainly the result of an effect among the non-woody vascular plants. This correlation suggests that the chemical diversity of the ecosystem dominant species has an important role in shaping the biodiversity of the associated plant community. The extent and significance of this effect, and its underlying processes require further investigation.

Ecological communities are vulnerable to realistic extinction sequences

Loss of species will directly change the structure and potentially the dynamics of ecological communities, which in turn may lead to additional species loss (secondary extinctions) due to direct and/or indirect effects (e.g. loss of resources or altered population dynamics). Furthermore, the vulnerability of food webs to repeated species loss is expected to be affected by food web topology, species interactions, as well as the order in which species go extinct. Species traits such as body size, abundance and connectivity might determine a species' vulnerability to extinction and, thus, the order in which species go primarily extinct. Yet, the sequence of primary extinctions, and their effects on the vulnerability of food webs to secondary extinctions, when species abundances are allowed to respond dynamically, has only recently become the focus of attention. Here, we analyse and compare topological and dynamical robustness to secondary extinctions of model food webs, in the face of 34 extinction sequences based on species traits. Although secondary extinctions are frequent in the dynamical approach and rare in the topological approach, topological and dynamical robustness tends to be correlated for many bottom-up directed, but not for top-down directed deletion sequences. Furthermore, removing species based on traits that are strongly positively correlated to the trophic position of species (such as large body size, low abundance, high net effect) is, under the dynamical approach, found to be as destructive as removing primary producers. Such top-down oriented removal of species are often considered to correspond to realistic extinction scenarios, but earlier studies, based on topological approaches, have found such extinction sequences to have only moderate effects on the remaining community. Thus, our result suggests that the structure of ecological communities, and therefore the integrity of important ecosystem processes could be more vulnerable to realistic extinction sequences than previously believed.

Application of low temperature plasmas for restoration/conservation of archaeological objects

The low-temperature low-pressure hydrogen based plasmas were used to study the influence of processes and discharge conditions on corrosion removal. The capacitive coupled RF discharge in the continuous or pulsed regime was used at operating pressure of 100-200 Pa. Plasma treatment was monitored by optical emission spectroscopy. To be able to study influence of various process parameters, the model corroded samples with and without sandy incrustation were prepared. The SEM-EDX analyzes were carried out to verify corrosion removal efficiency. Experimental conditions were optimized for the selected most frequent materials of original metallic archaeological objects (iron, bronze, copper, and brass). Chlorides removal is based on hydrogen ion reactions while oxides are removed mainly by neutral species interactions. A special focus was kept for the samples temperature because it was necessary to avoid any metallographic changes in the material structure. The application of higher power pulsed regime with low duty cycle seems be the best treatment regime. The low pressure hydrogen plasma is not applicable for objects with a very broken structure or for nonmetallic objects due to the non-uniform heat stress. Due to this fact, the new developed plasmas generated in liquids were applied on selected original archaeological glass materials.

Invader-invader interactions in relation to environmental heterogeneity leads to zonation of two invasive amphipods, Dikerogammarus villosus (Sowinsky) and Gammarus tigrinus Sexton: amphipod pilot species project (AMPIS) report 6

As biological invasions continue, interactions occur not only between invaders and natives, but increasingly new invaders come into contact with previous invaders. Whilst this can lead to species replacements, co-existence may occur, but we lack knowledge of processes driving such patterns. Since environmental heterogeneity can determine species richness and co-existence, the present study examines habitat use and its mediation of the predatory interaction between invasive aquatic amphipods, the Ponto-Caspian Dikerogammarus villosus and the N. American Gammarus tigrinus. In the Dutch Lake IJsselmeer, we found broad segregation of D. villosus and G. tigrinus by habitat type, the former predominating in the boulder zone and the latter in the soft sediment. However, the two species co-exist in the boulder zone, both on the short and longer terms. We used an experimental simulation of habitat heterogeneity and show that both species utilize crevices, different sized holes in a plastic grid, non-randomly. These amphipods appear to optimise the use of holes with respect to their 'C-shape' body size. When placed together, D. villosus adults preyed on G. tigrinus adults and juveniles, while G. tigrinus adults preyed on D. villosus juveniles. Juveniles were also predators and both species were cannibalistic. However, the impact on G. tigrinus of the superior intraguild predator, D. villosus, was significantly reduced where experimental grids were present as compared to absent. This mitigation of intraguild predation between the two species in complex habitats may explain the co-existence of these two invasive species.

Role of behaviour in biological invasions and species distributions; lessons from interactions between the invasive Gammarus pulex and the native G-duebeni (Crustacea : Amphipoda)

In attempting to understand the distributions of both introduced species and the native species on which they impact, there is a growing trend to integrate studies of behaviour with more traditional life history/ecological approaches. The question of what mechanisms drive the displacement of the freshwater amphipod Gammarus duebeni by the often introduced G pulex is presented as a case study Patterns of displacement are well documented throughout Europe, but the speed and direction of displacement between these species can be varied. From early studies proposing interspecific competition as causal in these patterns, I review research progress to date. I show there has been no evidence for interspecific competition operating, other than the field patterns themselves, a somewhat tautological argument. Rather, the increased recognition of behavioural attributes with respect to the cannibalistic and predatory nature of these species gave rise to a series of studies unravelling the processes driving field patterns. Both species engage in 'intraguild predation' (IGP), with moulting females particularly vulnerable to predation by congeneric males. G pulex is more able both to engage in and avoid this interaction with G duebeni. However, several factors mediate the strength and asymmetry of this IGP, some biotic (e.g. parasitism) and others abiotic (e.g. water chemistry). Further, a number of alternative hypotheses that may account for the displacement (hybridization; parasite transmission) have been tested and rejected. While interspecific competition has been modelled mathematically and found to be a weak interaction relative to IGP, mechanisms of competition between these Gammarus species remain largely untested empirically. Since IGP may be finely balanced in some circumstances, I conclude that the challenge to detect interspecific competition remains and we require assessment of its role, if any, in the interaction between these species. Appreciation of behavioural attributes and their mediation should allow us to more fully understand, and perhaps predict, species introductions and resultant distributions.

Trophic dynamics within a hybrid zone - interactions between an abundant cyprinid hybrid and sympatric parental species

1. Recent proliferation of hybridisation in response to anthropogenic ecosystem change, coupled with increasing evidence of the importance of ancient hybridisation events in the formation of many species, has moved hybridisation to the forefront of evolutionary theory.

2. In spite of this, the mechanisms (e. g. differences in trophic ecology) by which hybrids co-exist with parental taxa are poorly understood. A unique hybrid zone exists in Irish freshwater systems, whereby hybrid offspring off two non-native cyprinid fishes often outnumber both parental species.

3. Using stable isotope and gut content analyses, we determined the trophic interactions between sympatric populations of roach (Rutilus rutilus), bream (Abramis brama) and their hybrid in lacustrine habitats.

4. The diet of all three groups displayed little variation across the study systems, and dietary overlap was observed between both parental species and hybrids. Hybrids displayed diet, niche breadth and trophic position that were intermediate between the two parental species while also exhibiting greater flexibility in diet across systems.

Possible interactions between environmental factors in determining species optima

Questions: 1. Indicator values, such as those of Ellenberg, for different environmental factors are seen as independent. We tested for the presence of interactions between environmental factors ( soil moisture and reaction) to see if this assumption is simplistic. 2. How close are Ellenberg indicator values (IVs) related to the observed optima of species response curves in an area peripheral to those where they have been previously employed and 3. Can the inclusion of bryophytes add to the utility of IVs?

Location: South Uist, Outer Hebrides, Scotland, UK.

Methods: Two grids (ca. 2000 m x 2000 m) were sampled at 50-m intervals across the transition from machair to upland communities covering an orthogonal gradient of both soil pH ( reaction) and soil moisture content. Percentage cover data for vascular plants, bryophytes and lichens were recorded, along with pH and moisture content of the underlying sand/soil/peat. Reaction optima, derived from species response curves calculated using HOF models, were compared between wet and dry sites, and moisture optima between acidic and basic samples. Optima for the whole data set were compared to Ellenberg IVs to assess their performance in this area, with and without the inclusion of bryophytes.

Results: A number of species showed substantially different pH optima at high and low soil moisture contents (18% of those tested) and different soil moisture optima at high and low pH (49%). For a number of species the IVs were poor predictors of their actual distribution across the sampled area. Bryophytes were poor at explaining local variation in the environmental factors and also their inclusion with vascular plants negatively affected the strength of relationships.

Conclusions: A substantial number of species showed an interaction between soil moisture and reaction in determining their optima on the two respective gradients. It should be borne in mind that IVs such as Ellenberg's may not be independent of one another.

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a benthic marine assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of Sargassum muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of Sargassum muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.