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International audience; We discuss the interpretation and dimensions of the population dynamic parameters that are commonly used to quantify the strength of intraspecific and interspecific interactions. The concept of "interaction strength" is not unequivocal. Its theoretical formalization relies on the generalized Lotka-Volterra model. However, four different ways of parameterizing the model have been proposed in the literature, leading to four different definitions of the term "interaction strength." In particular, the dimensions of these four definitions are not identical, some of these incorporating explicitly the dimension used to measure the population size. Using an individual-based simulation model as an illustration, we show that, in the latter case, the interaction strength depends implicitly on the habitat size. As a consequence, it is of crucial importance to quantify the population as a density rather than an absolute population abundance. We insist that the dimension of the interaction strength coefficient should not be overlooked and any quantitative estimation must be given with explicit units.

Urban areas are being affected by rapidly increasing human-made pressures that can strongly homogenize biodiversity, reduce habitat heterogeneity, and facilitate the invasion of alien species. One of the key concerns in invaded urban areas is comparing the trait–environment relationships between alien and native species, to determine the underlying causes of invasiveness. In the current study, we used a trait–environment dataset of 130 native plants and 33 alien plants, recorded in 100 plots covering 50 urban areas and 50 non-urban ones in an urbanization gradient in the arid mountainous Saint-Katherine protected area in Egypt. We measured eleven morphological plant traits for each plant species and ten environmental variables in each plot, including soil resources and human-made pressures, to construct trait–environment associations using a fourth-corner analysis. In addition, we measured the mean functional and phylogenetic distances between the two species groups along an urbanization gradient. Our results revealed strongly significant relationships of alien species traits with human-made pressures and soil resources in urban areas. However, in non-urban areas, alien species traits showed weak and non-significant associations with the environment. Simultaneously, native plants showed consistency in their trait–environment relationships in urban and non-urban areas. In line with these results, the functional and phylogenetic distances declined between the aliens and natives in urban areas, indicating biotic homogenization with increasing urbanization, and increased in non-urban areas, indicating greater divergence between the two species groups. Thereby, this study provided evidence that urbanization can reveal the plasticity of alien species and can also be the leading cause of homogenization in an arid urban area. Future urban studies should investigate the potential causes of taxonomic, genetic, and functional homogenization in species composition in formerly more diverse urbanized areas.

Gaining knowledge of how ecosystems provide essential services to humans is of primary importance, especially with the current threat of climate change. Yet little is known about how increased temperature will impact the biodiversity–ecosystem functioning (BEF) relationship. We tackled this subject theoretically and experimentally. We developed a BEF theory based on mechanistic population dynamic models, which allows the inclusion of the effect of temperature. Using experimentally established relationships between attack rate and temperature, the model predicts that temperature increase will intensify competition, and consequently the BEF relationship will flatten or even become negative. We conducted a laboratory experiment with natural microbial microcosms, and the results were in agreement with the model predictions. The experimental results also revealed that an increase in both temperature average and variation had a more intense effect than an increase in temperature average alone. Our results indicate that under climate change, high diversity may not guarantee high ecosystem functioning.

International audience; Predator-prey interactions in natural ecosystems generate complex food webs that have a simple universal body-size architecture where predators are systematically larger than their prey. Food-web theory shows that the highest predator-prey body-mass ratios found in natural food webs may be especially important as they create weak interactions with slow dynamics that stabilize communities against perturbations and maintain ecosystem functioning. Identifying these vital interactions in real communities typically requires arduous identification of interactions in complex food webs. Here, we overcome this obstacle by developing predator-trait models to predict average body-mass ratios based on a database comprising 290 food webs from freshwater, marine and terrestrial ecosystems across all continents. We analyzed how species traits constrain body-size architecture by changing the slope of the predator-prey body-mass scaling. Across ecosystems, we found high body-mass ratios for predator groups with specific trait combinations including (1) small vertebrates and (2) large swimming or flying predators. Including the metabolic and movement types of predators increased the accuracy of predicting which species are engaged in high body-mass ratio interactions. We demonstrate that species traits explain striking patterns in the body-size architecture of natural food webs that underpin the stability and functioning of ecosystems, paving the way for community-level management of the most complex natural ecosystems.

Carnivorous plants and spiders are known to compete for resources. In North America, spiders of the genus Agelenopsis are known to build funnel-webs, using Sarracenia purpurea pitchers as a base, retreat and storage room. They also very likely profit from the insect attraction of S. purpurea. In a fen in Europe, S. purpurea was introduced ~65 years ago and co-occurs with native insect predators. Despite the absence of common evolutionary history, we observed native funnels-spiders (genus Agelena) building funnel webs on top of S. purpurea in similar ways as Agelelopsis. Furthermore, we observed specimen of the raft-spider (Dolmedes fimbriatus) and the pygmy-shrew (Sorex minutus) stealing prey-items out of the pitchers. We conducted an observational study, comparing plots with and without S. purpurea, to test if Agelena were attracted by S. purpurea, and found that their presence indeed increases Agelena abundance. Additionally, we tested if this facilitation was due to the structure provided for building webs or enhanced prey availability. Since the number of webs matched the temporal pattern of insect attraction by the plant, we conclude that the gain in food is likely the key factor for web installation. Our results provide an interesting case of facilitation by an introduced plant for a local predator, which has developed in a very short time scale.

We studied ballooning spiders captured weekly over an 11-y period using a 12.2-m- high suction trap in an agricultural landscape of western Switzerland. We analyzed population trends, changes in phenology, and species composition. Yearly trends in population size of the most abundant species were studied with nonparametric correlations. We found that these trends were markedly different for ground-living (92% of all decreasing species) and upper-strata species (75% of all increasing species). These contrasting tendencies can be explained by a stronger effect of meteorological conditions on species living at ground level and by an observed decrease in habitat availability for open-habitat species. The phenology of ballooning spiders was analyzed using mixtures of Gaussian curves fitted to the yearly capture data; this was possible for the 7 most abundant species. From these, we estimated the dates of the ballooning peaks for each species and each year (1 to 4 peaks depending on the species). We found that the dates remained mostly constant: the timing of dispersal peaks showed no annual trend for all but 1 species. Using nonparametric correlation, we analyzed the relationships between the dates of the peaks and the meteorological parameters occurring before the peak dates. In line with the absence of trend, the dates of the peaks were only weakly related to meteorological conditions. The extreme climatic event of 2003 had a strong impact by reducing populations of ground-living species. Using Canonical Correspondence Analysis and clustering methods, we identified a strong shift in the phenological structure of the ballooning spider assemblage in that year. In all, despite noticeable trends in population size during the study period, the dispersal phenology did not change, which contrasts with observations from other arthropod groups.

Understanding the effects of biodiversity on community persistence and productivity is key tomanaging both natural and production systems. Because rare species face greater danger of extinction, species evenness, a measure of how similar abundances are across species in a community, is seen as a key component of biodiversity. However, previous studies have failed to find a consistent association of species evenness with species survival and biomass production. Here we provide a theoretical framework for the relationship among these three elements. We demonstrate that the lack of consistent outcomes is not an idiosyncratic artifact of different studies but can be unified under one common framework. Applying a niche theory approach, we confirmthat under demographic stochasticity evenness is a general indicator of the risk of future species extinctions in a community, in accordance with the majority of empirical studies. In contrast, evenness cannot be used as a direct indicator of the level of biomass production in a community. When a single species dominates, as expressed by the constraints imposed by the population dynamics, biomass production depends on the niche position of the dominating species and can increase or decrease with evenness. We demonstrate that high species evenness and an intermediate level of biomass production is the configuration that maximizes the average species survival probability in response to demographic stochasticity

Understanding how trophic levels respond to changes in abiotic and biotic conditions is key for predicting how food webs will react to environmental perturbations. Different trophic levels may respond disproportionately to change, with lower levels more likely to react faster, as they typically consist of smaller-bodied species with higher reproductive rates. This response could cause a mismatch between trophic levels, in which predators and prey will respond differently to changing abiotic or biotic conditions. This mismatch between trophic levels could result in altered top-down and bottom-up control and changes in interaction strength. To determine the possibility of a mismatch, we conducted a reciprocal-transplant experiment involving Sarracenia purpurea food webs consisting of bacterial communities as prey and a subset of six morphologically similar protozoans as predators. We used a factorial design with four temperatures, four bacteria and protozoan biogeographic origins, replicated four times. This design allowed us to determine how predator and prey dynamics were altered by abiotic (temperature) conditions and biotic (predators paired with prey from either their local or non-local biogeographic origin) conditions. We found that prey reached higher densities in warmer temperature regardless of their temperature of origin. Conversely, predators achieved higher densities in the temperature condition and with the prey from their origin. These results confirm that predators perform better in abiotic and biotic conditions of their origin while their prey do not. This mismatch between trophic levels may be especially significant under climate change, potentially disrupting ecosystem functioning by disproportionately affecting top-down and bottom-up control.

Species abundance in local communities is determined by bottom-up and top-down processes, which can act directly and indirectly on the focal species. Studies examining these effects simultaneously are rare. Here we explore the direct top-down and direct and indirect bottom-up forces regulating the abundance and predation success of an intermediate predator, the web-building spider Argiope bruennichi (Araneae: Araneidae). We manipulated plant diversity (2, 6, 12 or 20 sown species) in 9 wildflower strips in a region of intensive farmland. To identify the major factors regulating the distribution and abundance of A. bruennichi, we quantified three characteristics of vegetation (species diversity, composition and vegetation structure) as well as the spider's prey community and natural enemies. The distribution and abundance of A. bruennichi was regulated by combined bottom-up and top-down processes as well as by direct and indirect interactions between trophic levels. Four main factors were identified: (1) the strong direct effect of vegetation structure, (2) the positive effect of plant species diversity, which affected spider abundance directly and indirectly through increased densities and size of flower-visiting prey species, (3) the positive or negative direct effects of different plant species, and (4) the strongly negative direct effect of predacious hornets. The advantage of taking a global approach to understand the regulation of species abundance is highlighted first by the quantification of the relative importance of factors, with a surprisingly strong effect of hornet predators, and second by the discovery of a direct effect of plant diversity, which raises intriguing questions about habitat selection by this spider.

Wildflower strips are used to increase natural enemies of crop pests and to conserve insect diversity on farmland. Mollusks, especially slugs, can affect the vegetation development in these strips considerably. Although recent theoretical work suggests that more diverse plant communities will exhibit greater resistance against herbivore pressure, empirical studies are scarce. We conducted a semi-natural experiment in wildflower strips, manipulating trophic structure (reduction in herbivorous mollusks and reduction in major predators) and plant diversity (2, 6, 12, 20 and 24 sown species). This design allowed us to assess the effect of plant diversity, biomass and composition on mollusks, and vice versa, the effect of mollusc abundance on vegetation. Seven species of mollusks were found in the strips, with the slugs Arion lusitanicus, Deroceras reticulatum and Deroceras panormitanum being most frequent. We found a negative relationship between plant diversity and mollusk abundance, which was due predominantly to a decrease in the agricultural pest species A. lusitanicus. These results are consistent with the hypothesis that plant diversity can reduce the impact of herbivores. However, plant identity also had an effect on mollusks, and accounted for a much larger fraction of the variation in mollusk communities than biodiversity effects. While overall plant diversity decreased during the 3 years of the study, in the final year the highest plant diversity was found in the plots where mollusk populations were experimentally reduced. We conclude that selective feeding by generalist herbivores leads to changes in plant community composition and hence reduced plant diversity. Our results highlight the importance of plant biodiversity as protection against generalist herbivores, which if abundant can in the long term negatively impact plant diversity, driving the system along a “low plant diversity – high mollusk abundance” trajectory.

The objective of this study was to compare butterfly abundances and diversity between wildflower strips and extensively used meadows to identify which butterfly species can be supported by establishing wildflower strips. Butterflies were recorded along transects during one season in twenty-five sown wildflower strips and eleven extensively used meadows in a Swiss lowland agricultural landscape (600 ha). In total 1,669 butterflies of 25 species were observed (25 in the strips, 18 in meadows). This can be related to 38 species recorded in the region (lowland part of Kanton Fribourg) within the Swiss Biodiversity Monitoring Programme. In wildflower strips the number of butterflies per transect meter was significantly higher than in the meadows, but there was no significant difference in species richness. Butterfly communities, though, were quite different between the two habitat types. Habitat type, abundances of flowering plants and presence of forest within 50 m were identified as factors influencing butterfly species richness. Butterfly abundances were affected by habitat type and abundance of flowering plants. In wildflower strips, 65% of all flower visits by butterflies were observed on Origanum. It can be concluded that sown wildflower strips can support a substantial part of a regions species pool. This is mostly true for common species, but can apply to rare species when, for example, larval food plant requirements are met.

Several stochastic models have tried to capture the architecture of food webs. This approach is interesting, but it is limited by the fact that different assumptions can yield similar results. To overcome this limitation, we develop a purely statistical approach. Body size in terms of an optimal ratio between prey and predator is used as explanatory variable. In 12 observed food webs, this model predicts, on average, 20% of interactions. To analyze the unexplained part, we introduce a latent term: each species is described by two latent traits, foraging and vulnerability, that represent nonmeasured characteristics of species once the optimal body size has been accounted for. The model now correctly predicts an average of 73% of links. The key features of our approach are that latent traits quantify the structure that is left unexplained by the explanatory variable and that this quantification allows a test of whether independent biological information, such as microhabitat use, camouflage, or phylogeny, explains this structure. We illustrate this method with phylogeny and find that it is linked to one or both latent traits in nine of 12 food webs. Our approach opens the door to the formulation of more complex models that can be applied to any kind of biological network.

Sown wildflower strips are increasingly being established in European countries within agri-environmental schemes to enhance biodiversity, especially in intensively used agricultural areas. 2. The regulations vary between countries regarding the seed mixture, intensity of management and period of time over which subsidies are given. Insects in particular are intended to benefit from these schemes. 3. This review treats studies of insect diversity and abundance in sown wildflower strips. Schemes on wildflower strips in several countries in Central and Northern Europe are compared. 4. In a significant majority of studies, sown wildflower strips support higher insect abundances and diversity than cropped habitats. In general, numbers and diversity also tend to be higher than in other margin types such as sown grass margins and natural regeneration, but pollen- and nectar-rich flower mixtures may outperform them. 5. Common species are the main beneficiaries of the establishment of wildflower strips, although some studies point out the presence of rare or declining insect spe

The modelling of prey–predator interactions is of major importance for the understanding of population dynamics. Classically, these interactions are modelled using ordinary differential equations, but this approach has the drawbacks of assuming continuous population variables and of being deterministic. We propose a general approach to stochastic modelling based on the concept of functional response for a prey depletion process with a constant number of predators. Our model could involve any kind of functional response, and permits a likelihood-based approach to statistical modelling and stable computation using matrix exponentials. To illustrate the method we use the Holling–Juliano functional response and compare the outcomes of our model with a deterministic counterpart considered by Schenk and Bacher [2002. Functional response of a generalist insect predator to one of its prey species in the field. Journal of Animal Ecology 71 (3), 524–531], who observed the depletion of Cassida rubiginosa due to its exclusive predator, Polistes dominulus. The predation was found to be Holling type III, reflecting the ability of the predator to regulate its prey. Our approach corroborates this result, but suggests that the prey depletion census should have been performed more often, and that predation features were significantly different between the two years for which data are available.

Food webs depict who eats whom in communities. Ecologists have examined statistical metrics and other properties of food webs, but mainly due to the uneven quality of the data, the results have proved controversial. The qualitative data on which those efforts rested treat trophic interactions as present or absent and disregard potentially huge variation in their magnitude, an approach similar to analyzing traffic without differentiating between highways and side roads. More appropriate data are now available and were used here to analyze the relationship between trophic complexity and diversity in 59 quantitative food webs from seven studies (14-202 species) based on recently developed quantitative descriptors. Our results shed new light on food-web structure. First, webs are much simpler when considered quantitatively, and link density exhibits scale invariance or weak dependence on food-web size. Second, the "constant connectance'' hypothesis is not supported: connectance decreases with web size in both qualitative and quantitative data. Complexity has occupied a central role in the discussion of food-web stability, and we explore the implications for this debate. Our findings indicate that larger webs are more richly endowed with the weak trophic interactions that recent theories show to be responsible for food-web stability.

The consensus that complexity begets stability in ecosystems was challenged in the seventies, a result recently extended to ecologically-inspired networks. The approaches assume the existence of a feasible equilibrium, i.e. with positive abundances. However, this key assumption has not been tested. We provide analytical results complemented by simulations which show that equilibrium feasibility vanishes in species rich systems. This result leaves us in the uncomfortable situation in which the existence of a feasible equilibrium assumed in local stability criteria is far from granted. We extend our analyses by changing interaction structure and intensity, and find that feasibility and stability is warranted irrespective of species richness with weak interactions. Interestingly, we find that the dynamical behaviour of ecologically inspired architectures is very different and richer than that of unstructured systems. Our results suggest that a general understanding of ecosystem dynamics requires focusing on the interplay between interaction strength and network architecture., Author summary Finding conditions leading to stable equilibria, i.e. equilibria robust to small perturbations, is a key feature in understanding how persistent are ecological systems. The number of species involved in the ecosystem as well as the number of relations between them has historically been pointed out to be the prominent ingredients determining stability. The latter is however always computed for a particular equilibrium, which needs and is usually assumed to contain only positive abundances. This assumption has until now only barely been tested in relation to system size. We provide here analytical results complemented by numerical simulations which show that it is almost impossible to find equilibria containing only positive abundances in species rich systems or, if so, the parameters are constrained to lead to ecologically-nonsensical abundances in unstructured systems. Interestingly, only ecologically-inspired architectures permit one to obtain feasible equilibria with a rich and sensible dynamical behavior.

Summary 1 There is a current debate about the appropriateness of prey-dependent vs. ratio-dependent functional responses in predator–prey models. This is an important issue as systems governed by these models exhibit quite different dynamical behaviour. However, the issue is not yet resolved on a theoretical basis, and there is a lack of experimental evidence in natural systems. We used a paper wasp–shield beetle system in a natural setting to assess the validity of either approach. 2 We manipulated the abundance of herbivorous insect prey on thistle plants and of predatory paper wasps in the immediate environment of the prey by opening or closing cages containing wasp nests. 3 The number of wasps foraging at the site increased when cages were opened, but rapidly reached an asymptote, indicating predator interference. The predation rate per predator decreased with the number of wasps in the environment. Thus, the functional response depended on both prey and predator density. 4 Neither a pure prey- nor a pure ratio-dependent model fitted perfectly our observations. However, the functional response of the paper wasps towards shield beetle larvae was closer to ratio-dependence. To our knowledge, this is the first experimental evidence discriminating between ratio- and prey-dependence in a natural setting with unconfined predators and prey. 5 Predator interference was most probably responsible for the specific form of the functional response found. We found indications that both direct (e.g. aggression) and indirect interference mechanisms (e.g. depletion of easy-to-find prey) were at work in our system. We conclude that predator density cannot be ignored in models of predator–prey interactions.

1. Parasitism is a non-negligible cost of reproduction in wild organisms, and hosts are selected to partition resources optimally between current and future reproduction. While parents can compensate for the cost of parasitism by increasing their current reproductive investment, such change in resource allocation is expected to carry-over costs on future reproduction. 2. Life history theory predicts that because long-lived organisms have a high residual reproductive value, they should be more reluctant to increase parental effort in response to parasites. Also, when rearing successive infested broods, the cost of parasitism can cumulate over the years and hence be exacerbated by past infestations. 3. We tested these two predictions in the alpine swift Apus melba, a long-lived colonial bird that is infested intensely by the nest-based blood sucking louse-fly Crataerina melbae. For this purpose, we manipulated ectoparasite load over 3 consecutive years and measured reproductive parameters in successive breeding attempts of adults assigned randomly to 'parasitized' and 'deparasitized' treatments. 4. In current reproduction, fathers of experimentally parasitized broods produced a similar number of offspring as fathers from the deparasitized treatment, but the rearing period was prolonged by 4 days. Fathers that were assigned to the parasitized treatment in year x produced significantly fewer fledglings the following year x + 1 than those of the deparasitized treatment. The number of young produced by fathers in year x + 1 was correlated negatively with the number of days they cared for their brood in the previous year x. We also found a significant interaction between treatments performed over 2 successive years, with fathers of parasitized broods suffering a larger fitness loss if in the past they had already cared for a parasitized brood rather than for a deparasitized one. Similar effects of parasitism, although partly non-significant (0.05 < P-values > 0.10), were found in mothers. 5. Altogether, our results show that parasites can modify resource allocation between current and future reproduction in long-lived hosts, and that the cost of parasitism can cumulate over the years. It emphasizes the fact that effects of parasites can depend on past infestations and become apparent in future reproduction only.

If interspecific competition is a strong structuring force of communities, ecologically similar species should tend to have spatial ranges at local scale that do not overlap. Experimental testing of this hypothesis becomes impracticable with large communities. One possibility to tackle this issue is a correlational approach, by comparing the matrix of niche overlap with that of spatial overlap. The use of the standard Mantel test is however impaired by the non-linearity in the relationship of the two descriptors: in a competitively structured assemblage, species with high niche overlap are expected to be segregated spatially, but species with small niche overlap may or may not exhibit high spatial overlap. To overcome this problem, we devised an original randomization test, which was run for three data-sets comprising frogs, lizards, and birds along altitudinal gradients. The test yielded intriguing results: reptiles and birds revealed an adjustment that would reduce the potential for interspecific competition, while amphibians showed the opposite trend, that is, ecologically similar species co-occurred more than expected by chance. Frogs may be more constrained by resource requirements, possibly breeding sites, than by competition. Our test will help to assess the generality of this pattern with other data-sets.

Food-web descriptors serve as a means for among-web comparisons that are necessary for the discovery of regularities in respect to food-web structure. Qualitative descriptors were however found to be highly sensitive to varying levels of sampling effort. To circumvent these shortcomings, quantitative counterparts were proposed which take the magnitude of trophic interaction between species into consideration. For 14 properties we examined the performance with increasing sampling effort of a qualitative, an unweighted quantitative (giving the same weight to each taxon), and a weighted quantitative version (weighing each taxon by the amount of incoming and outgoing flows). The evaluation of 10 extensively documented quantitative webs formed the basis for this analysis. The quantitative versions were found to be much more robust against variable sampling effort. This increase in accuracy is accomplished at the cost of a slight decrease in precision as compared to the qualitative properties. Conversely, the quantitative descriptors also proved less sensitive to differences in evenness in the distribution of link magnitude. By more adequately incorporating the information inherent to quantitative food-web compilations, quantitative descriptors are able to better represent the web, and are thus more suitable for the elucidation of general trends in food-web structure.

The three species of chameleons inhabiting the primary forest on the western slope of Mount Kupe, Cameroon, Chamaeleo montium, Chamaeleo pfefferi, and Chamaeleo quadricornis, are compared with respect to elevational distribution, diet, and body size. Chameleo montium is restricted to lower elevations and markedly separated in elevational distribution from its congeners, which are exclusively found in the submontane forest. The three species prey almost entirely on arthropods, mostly coleopterans, heteropterans, hymenopterans, dipterans, and spiders; niche breadth values that incorpate prey availability are high and suggest opportunistic feeding. Niche overlaps are high with respect to prey type, but the smallest species, C. pfefferi, differs significantly from its congeners in terms of prey volume. Body size difference is largest between the two syntopic species, C. pfefferi and C. quadricornis, and identically low to the allotopic C. montium. The results on dietary segregation and morphological similarity are in accordance with competition limiting elevational distributions of the chameleons on Mount Kupe.

Introduced top predators have the potential to disrupt community dynamics when prey species are naive to predation. The impact of introduced predators may also vary depending on the stage of community development. Early-succession communities are likely to have small-bodied and fast-growing species, but are not necessarily good at defending against predators. In contrast, late-succession communities are typically composed of larger-bodied species that are more predator resistant relative to small-bodied species. Yet, these aspects are greatly neglected in invasion studies. We therefore tested the effect of top predator presence on early- and late-succession communities that were either naive or non-naive to top predators. We used the aquatic community held within the leaves of Sarracenia purpurea. In North America, communities have experienced the S. purpurea top predator and are therefore non-naive. In Europe, this predator is not present and its niche has not been filled, making these communities top-predator naive. We collected early- and late-succession communities from two non-naive and two naive sites, which are climatically similar. We then conducted a common-garden experiment, with and without the presence of the top predator, in which we recorded changes in community composition, body size spectra, bacterial density, and respiration. We found that the top predator had no statistical effect on global measures of community structure and functioning. However, it significantly altered protist composition, but only in naive, early-succession communities, highlighting that the state of community development is important for understanding the impact of invasion.

A food web customarily describes the qualitative feeding relationships in a community. Descriptors have been used to extract ecologically meaningful information from such data, e.g., the proportion of top species (the proportion of taxa without consumers) or vulnerability (the average number of consumers per taxon). Analyses of collections of food webs based on these properties have revealed regularities that fostered the formulation of models of food-web structure. However, it has been shown that most of these qualitative descriptors are highly sensitive to the varying levels of sampling effort used to document a food web. The principal problem is that webs described extensively include trophic links of highly uneven magnitude, with typically few strong/important links and a wealth of weak ones; with qualitative descriptors, the same weight is given to all trophic interactions. To overcome this problem, food webs should be described and analyzed quantitatively. Consequently, we propose here a suite of food-web descriptors, which are built on information- theory indices and take the magnitude of the trophic interactions into account. We define descriptors having a similar meaning as the classical qualitative indices. Two versions of each quantitative descriptor are proposed, one giving the same weight to each taxon, and one weighting each taxon by the total amount of its incoming and outgoing biomass flows. We use a published quantitative food web to exemplify the computation of the new descriptors, and discuss their potential and limitations.

BACKGROUND: Hyperhomocysteinaemia has been identified as an independent cardiovascular risk factor and is found in more than 85% of patients on maintenance haemodialysis. Previous studies have shown that folic acid can lower circulating homocysteine in dialysis patients. We evaluated prospectively the effect of increasing the folic acid dosage from 1 to 6 mg per dialysis on plasma total homocysteine levels of haemodialysis patients with and without a history of occlusive vascular artery disease (OVD). METHODS: Thirty-nine stable patients on high-flux dialysis were studied. Their mean age was 63 +/-11 years and 17 (43%) had a history of OVD, either coronary and/or cerebral and/or peripheral occlusive disease. For several years prior to the study, the patients had received an oral post-dialysis multivitamin supplement including 1 mg of folic acid per dialysis. After baseline determinations, the folic acid dose was increased from 1 to 6 mg/dialysis for 3 months. RESULTS: After 3 months, plasma homocysteine had decreased significantly by approximately 23% from 31.1 +/- 12.7 to 24.5 +/- 9 micromol/l (P = 0.0005), while folic acid concentrations had increased from 6.5 +/- 2.5 to 14.4+/-2.5 microg/l (P < 0.0001). However, the decrease of homocysteine was quite different in patients with and in those without OVD. In patients with OVD, homocysteine decreased only marginally by approximately 2.5% (from 29.0 +/- 10.3 to 28.3 +/- 8.4 micromol/l, P = 0.74), whereas in patients without OVD there was a significant reduction of approximately 34% (from 32.7+/-14.4 to 21.6+/-8.6 micromol/l, P = 0.0008). Plasma homocysteine levels were reduced by > 15% in three patients (18%) in the group with OVD compared with 19 (86%) in the group without OVD (P = 0.001), and by > 30% in none of the patients (0%) in the former group compared with 13 (59%) in the latter (P = 0.001). CONCLUSIONS: These results indicate that the homocysteine-lowering effect of folic acid administration appears to be less effective in haemodialysis patients having occlusive vascular disease than in those without evidence of such disease.

The relative effects of the elevational gradient and of environmental discontinuities (ecotones) on the structure of a herpetofaunal assemblage in a tropical upland forest were contrasted by means of canonical correspondence analysis. Qualitative descriptors were used to define the elevational positions of the ecotones of interest, namely transitions in forest type and presence/absence of water bodies. The elevational gradient was coded in a form that accommodated different types of community response. Analyses were run for four subsets of the entire assemblage: (1) reptiles, (2) amphibians, (3) amphibians dependent on streams for reproduction, and (4) amphibians that do not use streams for reproduction. All subsets showed a significant relationship with the gradient, which suggested that most species respond to the physical continuum associated with the change in elevation. A response to ecotones was revealed for the amphibian subset only and associated with the presence or absence of watercourses. However, this response disappeared within subsets 3 and 4. A variation partitioning analysis was used to assess the individual and common contributions of gradient and ecotone descriptors to the elevational variation in the structure of subsets 1 and 2. The gradient descriptors explained more variation in the reptile subset than did ecotones, while the reverse was found in the amphibian subset. The dependence of most amphibians on aquatic breeding sites that were not available at all elevations reduced the relative importance of the gradient on the species distributions in subset 2 and accounted for the difference to the reptiles. In all, these findings add to the results of previous null model tests on the same four subsets, where competitive interactions were assigned a minor importance in limiting elevational distributions. The response patterns revealed by the present approach, with ecotones and gradient contrasted in a single analysis, emphasised the role of individual responses to the gradient according to the species' physiological tolerance limits.

Studies of initial compilations of food webs resulted in the intriguing generalization that the link density property, L/S (the number of trophic links L divided by the total number of species S in a web), may be scale invariant; that is, this ratio may remain constant across webs of

Declining plant species richness in agro-ecosystems and thus reduced habitat quality can have cascading effects on ecosystem functioning, leading to reduced pollination and biological control. Here we test if plant diversity can affect arthropod diversity and abundance on a very small scale, manipulating plant species richness (2, 6, 12 and 20 sown species) in small adjacent subplots (63 9 m) in 10 wildflower strips in an agricultural landscape. We simultaneously analyzed the effect of plant species richness, vegetation structure, and plant composition on the species richness and abundance of cavity-nesting wild bees, wasps, their prey and natural enemies, and on the structure of their food webs. By separating the trap-nesting species into functional groups according to their prey, we aimed to understand the underlying patterns for the effects of plant diversity. Increasing plant species richness had a significant effect only on spider-predating wasps, the group of wasps trophically most distant from plants. In contrast, bees and food-web structure were unaffected by plant diversity. Spider-predating wasp abundance negatively correlated with the abundance of spiders, suggesting top-down control. Interestingly, the abundance of spiders was the only variable that was strongly affected by plant composition. The hypothesis that the effect of plant diversity decreases with increasing trophic level is not supported by our study, and the mobility of species appears to play a greater role at this small spatial scale

In ecological networks, niche-overlap graphs are considered as complex systems. They represent the competition between two predators that share common resources. The purpose of this paper is to investigate the structural properties of these graphs considered as weighted networks and compare their measures with the ones calculated for the binary networks. To conduct this study, we select four classical network measures : the degree of nodes, the clustering coefficient, the assortativity, and the betweenness centrality. These measures were used to analyse different type of networks such as social networks, biological networks, world wide web, etc. Interestingly, we identify significant differences between the structure of the binary and the weighted niche-overlap graphs. This study indicates that weight information reveals different features that may provide other implications on the dynamics of these networks.

Studies on biodiversity and ecosystem functioning (BEF) have elicited debate over the interpretation of the positive relationship between species richness and plant productivity. Manipulating richness cannot be achieved without affecting composition; it is thus essential to consider the latter in statistical models. We firstly review existing approaches that use species richness as an explanatory variable and propose modifications to improve their performance. We use an original data set to illustrate the analyses. The classical method where composition is coded as a factor with a level for each different species mixture can be improved by defining the levels using clustering. Methods based on ordinations reduce the dimensionality of plant composition and use the new coordinates as fixed effects; they provide a much better fit to our observations. Secondly, we develop a new method where composition is included as a similarity matrix affecting the residual variance-covariance. Similarity in composition between plots is treated in the same way as shared evolutionary history between species in phylogenetic regression. We find that it outperforms the other models. We discuss the different approaches and suggest that our method is particularly suited for observational studies or for manipulative studies where plant diversity is not kept constant by weeding. By treating species composition in an intuitive and sensible way, it offers a valuable and powerful complement to existing models. © 2014 British Ecological Society., BEF; Biodiversity; Composition; Ecosystem function; Ecosystem services;Mixed effects models; Residual correlation structure; Species richness

Historically, ecologists have been more interested in organisms feeding at the tops of food chains than in organisms feeding at or near the bottom. The problem of taxonomic and trophic inconsistency within and among described food webs is central to criticisms of contemporary food web research. To study the effects of taxonomic and trophic aggregation on food web properties, 38 published food webs, each containing a large fraction of investigator-defined biological species, were systematically aggregated by taxonomy and trophic (functional) group similarity. During each step of taxonomic and trophic aggregation, eight food web properties (MIN, MAX, mean chain lengths; the fractions of basal, intermediate and top species; the ratio of all links by the total number of species, L/S; and rigid circuits) were calculated and their departures from the original, unaggregated version were recorded. We found only two properties showing wide systematic departure from initial values after both taxonomic and trophic group aggregation: the fraction of basal species and L/S. One reason for the relative ‘constancy’ of the six other properties was due in part to large numbers of trophically equivalent species (species with identical sets of prey and predators) found in these and other published webs. In the 38 webs, the average number of trophically equivalent species was 45% and ranged from a low of 13% in aquatic webs to a high of 71% in certain terrestrial systems (i.e., carrion webs). Six of the eight properties (MIN, MAX and mean chain lengths, the fractions of top and basal species, and the L/S ratio) were found to be more sensitive to taxonomic than to trophic aggregation. The relatively smaller variations observed in trophically lumped versions suggest that food web properties more aptly reflect functional, rather than taxonomic, attributes of real food webs. These findings parallel earlier trophic-based results, and bolster the conclusion that uneven lumping of taxonomic and trophic groups in published food web reports do not modify markedly the scaling behaviour of most of their descriptive properties.

Networks play a prominent role in the study of complex systems of interacting entities in biology, sociology, and economics. Despite this diversity, we demonstrate here that a statistical model decomposing networks into matching and centrality components provides a comprehensive and unifying quantification of their architecture. The matching term quantifies the assortative structure in which node makes links with which other node, whereas the centrality term quantifies the number of links that nodes make. We show, for a diverse set of networks, that this decomposition can provide a tight fit to observed networks. Then we provide three applications. First, we show that the model allows very accurate prediction of missing links in partially known networks. Second, when node characteristics are known, we show how the matching–centrality decomposition can be related to this external information. Consequently, it offers us a simple and versatile tool to explore how node characteristics explain network architecture. Finally, we demonstrate the efficiency and flexibility of the model to forecast the links that a novel node would create if it were to join an existing network., The work was supported by the National Centre of Competence in Research ‘Plant Survival’, the Swiss National Science Foundation grant 31003A-138489 (both to L.-F. Bersier), by SystemsX.ch, the Swiss Initiative in Systems Biology (to L.-F. Bersier and C. Mazza), and by the FP7-REGPOT-2010-1 programme (project 264125 EcoGenes), and an ERC Advanced Grant (both to J. Bascompte).

The structure of networks has always been interesting for researchers. Investigating their unique architecture allows to capture insights and to understand the function and evolution of these complex systems. Ecological networks such as food-webs and niche-overlap graphs are considered as complex systems. The main purpose of this work is to compare the topology of 15 real niche-overlap graphs with random ones. Five measures are treated in this study: (1) the clustering coefficient, (2) the between ness centrality, (3) the assortativity coefficient, (4) the modularity and (5) the number of chord less cycles. Significant differences between real and random networks are observed. Firstly, we show that niche-overlap graphs display a higher clustering and a higher modularity compared to random networks. Moreover we find that random networks have barely nodes that belong to a unique sub graph (i.e. between ness centrality equal to 0) and highlight the presence of a small number of chord less cycles compared to real networks. These analyses may provide new insights in the structure of these real niche-overlap graphs and may give important implications on the functional organization of species competing for some resources and on the dynamics of these systems.

International audience; The functional response is a key component of trophic interactions since it quantifies the per capita rate of prey consumption. Determining whether this rate depends on the prey density only (which is the standard assumption), on both prey and predator densities, or simply on their ratio is essential to understand interacting populations. Several experiments have convincingly demonstrated ratio dependence but, with very few exceptions, they were conducted in laboratory conditions. The difficulty of collecting the required data (initial prey density, prey consumption, predator density) probably explains the lack of evidence from functional responses observed in natural systems. A field experiment was previously conducted with a paper wasp and its prey, shield beetle larvae. Both densities were manipulated and the prey consumption was measured. A first analysis led to the conclusion that the functional response of the wasps depended on both prey and predator densities but could not be considered being ratio dependent. Here, we perform an improved analysis of these data, making better justified assumptions and using more appropriate statistical methods. We fit several functional response models to the data and select the best one with information-theoretic criteria. We also estimate the model parameters and their confidence intervals. This more reliable analysis significantly modifies the original conclusion. Both model selection and parameter estimation indicate that ratio dependence governs the functional response of paper wasps preying on shield beetles in the field. Therefore, ratio dependence is not a laboratory artefact and should be more systematically considered as a potential model for describing functional responses.

Understanding the environmental factors that structure biodiversity and food webs among communities is central to assess and mitigate the impact of landscape changes. Wildflower strips are ecological compensation areas established in farmland to increase pollination services and biological control of crop pests and to conserve insect diversity. They are arranged in networks in order to favour high species richness and abundance of the fauna. We describe results from experimental wildflower strips in a fragmented agricultural landscape, comparing the importance of landscape, of spatial arrangement and of vegetation on the diversity and abundance of trap-nesting bees, wasps and their enemies, and the structure of their food webs. The proportion of forest cover close to the wildflower strips and the landscape heterogeneity stood out as the most influential landscape elements, resulting in a more complex trap-nest community with higher abundance and richness of hosts, and with more links between species in the food webs and a higher diversity of interactions. We disentangled the underlying mechanisms for variation in these quantitative food web metrics. We conclude that in order to increase the diversity and abundance of pollinators and biological control agents and to favour a potentially stable community of cavity-nesting hymenoptera in wildflower strips, more investment is needed in the conservation and establishment of forest habitats within agro-ecosystems, as a reservoir of beneficial insect populations.

Body mass is a fundamental characteristic that affects metabolism, life history, and population abundance and frequently sets bounds on who eats whom in food webs. Based on a collection of topological food webs, Ulrich Brose and colleagues presented a general relationship between the body mass of predators and their prey and analyzed how mean predator–prey body-mass ratios differed among habitats and predator metabolic categories. Here we show that the general body-mass relationship conceals significant variation associated with both predator and prey phylogeny. Major-axis regressions between the log body mass of predators and prey differed among taxonomic groups. The global pattern for Kingdom Animalia had slope > 1, but phyla and classes varied, and several had slopes significantly

Ecophylogenetics can be viewed as an emerging fusion of ecology, biogeography and macroevolution. This new and fast-growing field is promoting the incorporation of evolution and historical contingencies into the ecological research agenda through the widespread use of phylogenetic data. Including phylogeny into ecological thinking represents an opportunity for biologists from different fields to collaborate and has provided promising avenues of research in both theoretical and empirical ecology, towards a better understanding of the assembly of communities, the functioning of ecosystems and their responses to environmental changes. The time is ripe to assess critically the extent to which the integration of phylogeny into these different fields of ecology has delivered on its promise. Here we review how phylogenetic information has been used to identify better the key components of species interactions with their biotic and abiotic environments, to determine the relationships between diversity and ecosystem functioning and ultimately to establish good management practices to protect overall biodiversity in the face of global change. We evaluate the relevance of information provided by phylogenies to ecologists, highlighting current potential weaknesses and needs for future developments. We suggest that despite the strong progress that has been made, a consistent unified framework is still missing to link local ecological dynamics to macroevolution. This is a necessary step in order to interpret observed phylogenetic patterns in a wider ecological context. Beyond the fundamental question of how evolutionary history contributes to shape communities, ecophylogenetics will help ecology to become a better integrative and predictive science. © 2012 CNRS. Biological Reviews © 2012 Cambridge Philosophical Society.

1. Tokeshi (1990) proposed a goodness-of-fit test to distinguish among a general class of stochastic species abundance models using field data. This test is a good first step, but it is impaired by some shortcomings: there is no adjustment for the number of species (the rejection rate increases with the number of species in a data set); the variance of the generated abundance distributions are not taken into account (data sets with higher variance than a model are not rejected). 2. We propose an alternative Monte Carlo test that overcomes these problems. This is a versatile test that can be used with any stochastic model generating distributions.