Spatial ecology of food webs : herbivore-parasitoid communities on the pedunculate oak

Herbivorous insects, their host plants and natural enemies form the largest and most species-rich communities on earth. But what forces structure such communities? Do they represent random collections of species, or are they assembled by given rules? To address these questions, food webs offer excellent tools. As a result of their versatile information content, such webs have become the focus of intensive research over the last few decades. In this thesis, I study herbivore-parasitoid food webs from a new perspective: I construct multiple, quantitative food webs in a spatially explicit setting, at two different scales. Focusing on food webs consisting of specialist herbivores and their natural enemies on the pedunculate oak, Quercus robur, I examine consistency in food web structure across space and time, and how landscape context affects this structure. As an important methodological development, I use DNA barcoding to resolve potential cryptic species in the food webs, and to examine their effect on food web structure. I find that DNA barcoding changes our perception of species identity for as many as a third of the individuals, by reducing misidentifications and by resolving several cryptic species. In terms of the variation detected in food web structure, I find surprising consistency in both space and time. From a spatial perspective, landscape context leaves no detectable imprint on food web structure, while species richness declines significantly with decreasing connectivity. From a temporal perspective, food web structure remains predictable from year to year, despite considerable species turnover in local communities. The rate of such turnover varies between guilds and species within guilds. The factors best explaining these observations are abundant and common species, which have a quantitatively dominant imprint on overall structure, and suffer the lowest turnover. By contrast, rare species with little impact on food web structure exhibit the highest turnover rates. These patterns reveal important limitations of modern metrics of quantitative food web structure. While they accurately describe the overall topology of the web and its most significant interactions, they are disproportionately affected by species with given traits, and insensitive to the specific identity of species. As rare species have been shown to be important for food web stability, metrics depicting quantitative food web structure should then not be used as the sole descriptors of communities in a changing world. To detect and resolve the versatile imprint of global environmental change, one should rather use these metrics as one tool among several.

Efficiency, energy and stoichiometry in pelagic food webs; reciprocal roles of food quality and food quantity

In most lakes, zooplankton production is constrained by food quantity, but frequently high C:P poses an additional constraint on zooplankton production by reducing the carbon transfer efficiency from phytoplankton to zooplankton. This review addresses how the flux of matter and energy in pelagic food webs is regulated by food quantity in terms of C and its stoichiometric quality in terms of C:P. Increased levels of light, CO2 and phosphorus could each increase seston mass and, hence, food quantity for zooplankton, but while light and CO2 each cause increased C:P (i.e. reduced food quality for herbivores), increased P may increase seston mass and its stoichiometric quality by reducing C:P. Development of food quality and food quantity in response to C- or P-enrichments will differ between 'batch-type' lakes (dominated by one major, seasonal input of water and nutrients) and 'continuous-culture' types of lakes with a more steady flow-rate of water and nutrients. The reciprocal role of food quantity and stoichiometric quality will depend strongly on facilitation via grazing and recycling by the grazers, and this effect will be most important in systems with low renewal rates. At high food abundance but low quality, there will be a 'quality starvation' in zooplankton. From a management point of view, stoichiometric theory offers a general tool-kit for understanding the integrated role of C and P in food webs and how food quantity and stoichiometric quality (i.e. C:P) regulate energy flow and trophic efficiency from base to top in food webs.From a management point of view, stoichiometric theory offers a general tool-kit for understanding the integrated role of C and P in food webs and how food quantity and stoichiometric quality (i.e. C:P) regulate energy flow and trophic efficiency from base to top in food webs.

Implications of changes in trophic diversity and food webs on fisheries and the environment

Most of the earth's ecosystems are experiencing slight to catastrophic losses of biodiversity, caused by habitat destruction, alien species introduction, climate change and pollution (Wilcove et al., 1998). These human effects have led to the extinction of native fish species, the collapse of their populations and the loss of ecological integrity and ecosystem functioning (Ogutu-Ohwayo & Hecky, 1991; Witte et al. , 1992a; Mills et al., 1994; Vitousek et al., 1996). Food webs are macro-descriptors of community feeding interactions that can be used to map the flow of materials and nutrients in ecosystems (Jepsen & Winemiller, 2002). Comparative food web studies have been used to address theoretical questions such as 'does greater trophic connectivity increase stability?' (Cohen et al., 1990), and 'does the number of trophic levels increase with productivity?' (Briand & Cohen, 1987). Answers to such questions have obvious applications for natural resources management. From a multi-species fisheries standpoint, there is a need to understand consumer-resource dynamics within complex trophic networks.

Trophic interrelationships and food-webs among the fishes in ecosystems of the Victoria and Kyoga lake basin

The Victoria and Kyoga lake basins had a high fish species diversity with many fish species that were found only in these lakes. Two Tilapiines species Oreochromis esculentus and Oreochromis variabilis were the most important commercial species in these lakes and were found nowhere else on earth except in the Victoria and Kyoga lake basins (Graham 1929, Worthington 1929). Lakes Kyoga and Nabugabo also had endemic haplochromine species (Worthington 1929, Trewavas 1933, Greenwood 1965, 1966). As stocks of introduced species increased, stocks of most of the native species declined rapidly or disappeared altogether. The study was carried out on Lakes Victoria and Kyoga, River Nile, some selected satellite lakes from the two basins namely Lakes Mburo, Kachera, Wamala, Kayanja, Kayugi, Nabugabo, Victoria, Victoria nile and River Sio(Victoria lake basin). Lakes Kyoga (Iyingo), Nawampasa, Nakuwa, Gigati, Nyaguo, Agu, Kawi and Lemwa (Kyoga lake basin). Species composillon and relative abundance of fishes were estimated by detennining the overall average total number of each species encountered. A trophic consists of species using the same food category. Shannon-Weaver Index of diversity H (Pielou, 1969) and number of trophic groups, were used to estimate the Trophic diversity of various fish species in the lakes. Food analysis has been done on some fishes in some of the sampled lakes and is still going on, on remaining fishes and in some lakes. Generally fish ingested detritus, Spirulina, Melosira, filamentous algae, Planktolyngbya, Microcysists, Anabaena, Merismopedia, Spirogyra, higher plant material, rotifers, Ostracodes, Chironomid larvae and pupae, Choaborus larvae, Odonata, Povilla, Insect remains, Caridina, fish eggs and fish. Eight trophic groups were identified from thes food items ingestes. These included detritivores, algae eaters, higher plant eaters, zooplanktivores, insectivores, molluscivores, prawn eaters, paedophages and piscivores. Trophic diversity by number of trophic groups was highest in Lake Kyoga (6) followed by lakes Kayugi, Nabugabo, River Nile and Mburo (3) and the lowest number was recorded in kachera (2).

Size-related shifts in reliance on benthic and pelagic food webs by lake anchovy

This paper reports large variations in stable carbon and nitrogen isotope ratios of lake anchovy (Coilia ectenes taihuensis) from Lake Chaohu, China. The lake anchovy exhibited a significant C-13- and N-15- enrichment in relation to increasing fish length, and the isotopic compositions of small lake anchovy (<= 130 mm) were significantly more enriched than those of large lake anchovy (> 130 mm). The significant differences in the isotopic compositions of small and large lake anchovy suggested that their assimilated diets differed over a period of time and reflected the size-related diet shift of this fish. Bellamya aeruginosa and Corbicula fluminea were used to establish the baseline carbon signal of benthic and pelagic food webs, and these data were used to parameterize a 2-source mixing model to estimate in consumers the contribution of carbon derived from benthic versus pelagic food webs. Mixing models showed that small lake anchovy derived only 37% of their carbon from benthic food web, indicating increased reliance on pelagic prey, whereas benthic prey contributed 71% of large lake anchovy diet, suggesting greater use of benthic sources. These data indicate that there was a change in lake anchovy feeding strategy related to their size, suggesting a role in dynamic coupling between pelagic and benthic food chains. The trophic position of small lake anchovy averaged 3.0, indicating a zooplankton-based diet, compared with 3.6 in large lake anchovy, indicative of an increase in piscivorous diet. Overlap in the isotopic compositions of small and large lake anchovy probably indicated that these fish occasionally shared common diets, as suggested by stomach content studies, and/or resulted from the differences in the rate of isotopic turnover depending on differences in growth rate and metabolic turnover between small and large anchovy during diet shift from pelagic to benthic food webs. This study presents the contributions of benthic and pelagic food webs supporting lake anchovy and indicates that the intraspecific isotopic dynamic should be considered when applying stable isotope analyses to infer trophic interactions in aquatic ecosystems.

Sardine cycles, krill declines, and locust plagues: revisiting ‘wasp-waist’ food webs

‘Wasp-waist’ systems are dominated by a mid trophic-level species that is thought to exert top-down control on its food and bottom-up control on its predators. Sardines, anchovy, and Antarctic krill are suggested examples, and here we use locusts to explore whether the wasp-waist concept also applies on land. These examples also display the traits of mobile aggregations and dietary diversity, which help to reduce the foraging footprint from their large, localised biomasses. This suggests that top-down control on their food operates at local aggregation scales and not at wider scales suggested by the original definition of wasp-waist. With this modification, the wasp-waist framework can cross-fertilise marine and terrestrial approaches, revealing how seemingly disparate but economically important systems operate.

Part-whole relations between food webs and the validity of local food-web descriptions

This work analyzes the relationship between large food webs describing potential feeding relations between species and smaller sub-webs thereof describing relations actually realized in local communities of various sizes. Special attention is given to the relationships between patterns of phylogenetic correlations encountered in large webs and sub-webs. Based on the current theory of food-web topology as implemented in the matching model, it is shown that food webs are scale invariant in the following sense: given a large web described by the model, a smaller, randomly sampled sub-web thereof is described by the model as well. A stochastic analysis of model steady states reveals that such a change in scale goes along with a re-normalization of model parameters. Explicit formulae for the renormalized parameters are derived. Thus, the topology of food webs at all scales follows the same patterns, and these can be revealed by data and models referring to the local scale alone. As a by-product of the theory, a fast algorithm is derived which yields sample food webs from the exact steady state of the matching model for a high-dimensional trophic niche space in finite time. (C) 2008 Elsevier B.V. All rights reserved.

Food webs: Experts consuming families of experts

Food webs of habitats as diverse as takes or desert valleys are known to exhibit common

Soil resource supply influences faunal size-specific distributions in natural food webs

The large range of body-mass values of soil organisms provides a tool to assess the ecological organization of soil communities. The goal of this paper is to identify graphical and quantitative indicators of soil community composition and ecosystem functioning, and to illustrate their application to real soil food webs. The relationships between log-transformed mass and abundance of soil organisms in 20 Dutch meadows and heathlands were investigated. Using principles of allometry, maximal use can be made of ecological theory to build and explain food webs. The aggregate contribution of small invertebrates such as nematodes to the entire community is high under low soil phosphorus content and causes shifts in the mass-abundance relationships and in the trophic structures. We show for the first time that the average of the trophic link lengths is a reliable predictor for assessing soil fertility responses. Ordered trophic link pairs suggest a self-organizing structure of food webs according to resource availability and can predict environmental shifts in ecologically meaningful ways.

Body mass-abundance relationships are robust to cascading effects in marine food webs

Body mass has been shown to scale negatively with abundance in a wide range of habitats and ecosystems. It is believed that this relationship has important consequences for the distribution and maintenance of energy in natural communities. Some studies have shown that the relationship between body mass and abundance may be robust to major food web perturbations, fuelling the belief that natural processes may preserve the slope of this relationship and the associated cycling of energy and nutrients. Here, we use data from a long-term experimental food web manipulation to examine this issue in a semi-natural environment. Similar communities were developed in large experimental mesocosms over a six month period. Some of the mesocosms were then subjected to species removals, based on the mean strength of their trophic interactions in the communities. In treatments where the strongest interactors were removed, a community-level trophic cascade occurred. The biomass density of invertebrates increased dramatically in these communities, which led to a suppression of primary production. In spite of these widespread changes in ecosystem functioning, the slope of the relationship between body mass and abundance remained unchanged. This was the case whether average species body mass and abundance or individual organism size spectra were considered. An examination of changes in species composition before and after the experimental manipulations revealed an important mechanism for maintaining the body mass-abundance relationship. The manipulated communities all had a higher species turnover than the intact communities, with the highest turnover in communities that experienced cascading effects. As some species increased in body mass and abundance, new species filled the available size-abundance niches that were created. This maintained the overall body mass-abundance relationship and provided a stabilising structure to these experimental communities.

Interaction strength, food web topology and the relative importance of species in food webs

P>1. We established complex marine communities, consisting of over 100 species, in large subtidal experimental mesocosms. We measured the strength of direct interactions and the net strength of direct and indirect interactions between the species in those communities, using a combination of theoretical and empirical approaches.

Perturbations to trophic interactions and the stability of complex food webs

The pattern of predator-prey interactions is thought to be a key determinant of ecosystem processes and stability. Complex ecological networks are characterized by distributions of interaction strengths that are highly skewed, with many weak and few strong interactors present. Theory suggests that this pattern promotes stability as weak interactors dampen the destabilizing potential of strong interactors. Here, we present an experimental test of this hypothesis and provide empirical evidence that the loss of weak interactors can destabilize communities in nature. We ranked 10 marine consumer species by the strength of their trophic interactions. We removed the strongest and weakest of these interactors from experimental food webs containing >100 species. Extinction of strong interactors produced a dramatic trophic cascade and reduced the temporal stability of key ecosystem process rates, community diversity and resistance to changes in community composition. Loss of weak interactors also proved damaging for our experimental ecosystems, leading to reductions in the temporal and spatial stability of ecosystem process rates, community diversity, and resistance. These results highlight the importance of conserving species to maintain the stabilizing pattern of trophic interactions in nature, even if they are perceived to have weak effects in the system.

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.