Negative selection effects suppress relationships between bacterial diversity and ecosystem functioning

I assembled bacterial communities to explore the effects of bacterial diversity on multiple ecosystem functions, including bacterial community biovolume, decomposition of particulate organic matter, and biomass transfer to the next trophic level. The experiment used a two-way factorial design with four levels of bacterial diversity (one to four species) and the absence/presence of a bacterivorous ciliated protist Tetrahymena pyriformis as two main factors, and all possible combinations of the four bacterial taxa nested within each diversity level. Bacterial community biovolume increased as bacterial diversity increased, a result due largely to positive selection effects. Decomposition and consumer abundance, however, were unaffected by bacterial diversity, though both varied among bacterial composition treatments. Negative selection effects, the dominance of species that do not contribute significantly to ecosystem functioning, accounted for the lack of diversity effects on decomposition and consumer abundance. The presence of Tetrahymena reduced bacterial community biovolume but increased decomposition, without altering the diversity-functioning relationships. Decomposition was strongly linked with consumer abundance such that communities supporting larger consumer biomass exhibited higher decomposition rates. This study suggests that if the negative selection effect is common, as it might be when examining ecosystem variables other than biomass (due to the presence of keystone species that can contribute disproportionably to ecosystem functioning relative to their abundances), basic bacteria-mediated ecosystem processes, such as decomposition and energy transfer to the next trophic level, may not increase with bacterial diversity.     

Consequences of dominance: a review of evenness effects on local and regional ecosystem processes

The composition of communities is strongly altered by anthropogenic manipulations of biogeochemical cycles, abiotic conditions, and trophic structure in all major ecosystems. Whereas the effects of species loss on ecosystem processes have received broad attention, the consequences of altered species dominance for emergent properties of communities and ecosystems are poorly investigated. Here we propose a framework guiding our understanding of how dominance affects species interactions within communities, processes within ecosystems, and dynamics on regional scales. Dominance (or the complementary term, evenness) reflects the distribution of traits in a community, which in turn affects the strength and sign of both intraspecifc and interspecific interactions. Consequently, dominance also mediates the effect of such interactions on species coexistence. We review the evidence for the fact that dominance directly affects ecosystem functions such as process rates via species identity (the dominant trait) and evenness (the frequency distribution of traits), and indirectly alters the relationship between process rates and species richness. Dominance also influences the temporal and spatial variability of aggregate community properties and compositional stability (invasibility). Finally, we propose that dominance affects regional species coexistence by altering metacommunity dynamics. Local dominance leads to high beta diversity, and rare species can persist because of source-sink dynamics, but anthropogenically induced environmental changes result in regional dominance and low beta diversity, reducing regional coexistence. Given the rapid anthropogenic alterations of dominance in many ecosystems and the strong implications of these changes, dominance should be considered explicitly in the analysis of consequences of altered biodiversity.     

A simple plant mutation abets a predator-diversity cascade

Resource consumption often increases with greater consumer biodiversity. This could result either from complementarity among consumers or the inclusion of particular key species, and it is often difficult to differentiate between these two mechanisms. We exploited a simple plant mutation (reduced production of surface waxes) to alter foraging within a community of aphid predators, and thus perhaps shift the nature of resulting predator diversity effects. We found that greater predator species richness dramatically increased prey suppression and plant biomass only on mutant, reduced-wax pea plants (Pisum sativum). On pea plants from a sister line with wild type, waxier plant surfaces, predator species richness did not influence predators' impacts on herbivores or plants. Thus, a change in plant surface structure acted to turn on, or off, the cascading effects of predator diversity. Greater predator richness encouraged higher densities of true predators but did not lead to greater reproduction by a parasitoid, Aphidius ervi; fecundity of each natural enemy species was similar for the two plant types. Behavioral observations indicated that although A. ervi was less likely to forage within species-rich predator communities, low-wax plants mitigated this interference by encouraging generally greater A. ervi foraging and thus high rates of aphid dislodgement (aphids dropped from plants to escape A. ervi, but not the other predators). Thus, only species-rich, low-wax plants simultaneously encouraged strong species-specific effects of A. ervi, and strong complementarity among the other predator species. In summary, our study provides evidence that diversity effects in predator assemblages are sensitive to habitat characteristics. Further, we show that a simple plant morphological trait, controlled by a single gene mutation, can dramatically alter the cascading effects of predator species richness on herbivores and plants.     

Population and community resilience in multitrophic communities

Diversity-stability relationships have long been a topic of controversy in ecology, but one whose importance has been re-highlighted by increasing large-scale threats to global biodiversity. The ability of a community to recover from a perturbation (or resilience) is a common measure of stability that has received a large amount of theoretical attention. Yet, general expectations regarding diversity-resilience relations remain elusive. Moreover, the effects of productivity and its interaction with diversity on resilience are equally unclear. We examined the effects of species diversity, species composition, and productivity on population-and community-level resilience in experimental aquatic food webs composed of bacteria, algae, heterotrophic protozoa, and rotifers. Productivity manipulations were crossed with manipulations of the number of species and species compositions within trophic groups. Resilience was measured by perturbing communities with a nonselective, density-independent, mortality event and comparing responses over time between perturbed communities and controls. We found evidence that species diversity can enhance resilience at the community level (i.e., total community biomass), though this effect was more strongly expressed in low-productivity treatments. Diversity effects on resilience were driven by a sampling/selection effect, with resilient communities showing rapid response and dominance by a minority of species (primarily unicellular algae). In contrast, diversity had no effect on mean population-level resilience. Instead, the ability of a community's populations to recover from perturbations was dependent on species composition. We found no evidence of an effect of productivity, either positive or negative, on community- or population-level resilience. Our results indicate that the role of diversity as an insurer of stability may depend on the level of biological organization at which stability is measured, with effects emerging only when focusing on aggregate community properties.     

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.     

A direct comparison of the consequences of plant genotypic and species diversity on communities and ecosystem function

Biodiversity loss is proceeding at an unprecedented rate, yet we lack a thorough understanding of the consequences of losing diversity at different scales. While species diversity is known to impact community and ecosystem processes, genotypic diversity is assumed to have relatively smaller effects. Nonetheless, a few recent studies suggest that genotypic diversity may have quantitatively similar ecological consequences compared to species diversity. Here we show that increasing either genotypic diversity of common evening primrose (Oenothera biennis) or species diversity of old-field plant species resulted in nearly equivalent increases (approximately 17%) in aboveground primary production. The predominant mechanism explaining this effect, niche complementarity, was similar for each type of diversity. Arthropod species richness also increased with both types of plant diversity, but the mechanisms leading to this effect differed: abundance-driven accumulation of arthropod species was important in plant genotypic polycultures, whereas resource specialization was important in plant species polycultures. Thus, similar increases in primary productivity differentially impacted higher trophic levels in response to each type of plant diversity. These results highlight important ecological similarities and differences between genotypic and species diversity and suggest that genotypic diversity may play a larger role in community and ecosystem processes than previously realized.     

Seaweed diversity enhances nitrogen uptake via complementary use of nitrate and ammonium

The consequences of declining biodiversity remain controversial, in part because many studies focus on a single metric of ecosystem functioning and fail to consider diversity's integrated effects on multiple ecosystem functions. We used tide pool microcosms as a model system to show that different conclusions about the potential effects of producer diversity on ecosystem functioning may result when ecosystem functions are measured separately vs. together. Specifically, we found that in diverse seaweed assemblages, uptake of either nitrate or ammonium alone was equal to the average of the component monocultures. However, when nitrate and ammonium were available simultaneously, uptake by diverse assemblages was 22% greater than the monoculture average because different species were complementary in their use of different nitrogen forms. Our results suggest that when individual species have dominant effects on particular ecosystem processes (i.e., the sampling effect), multivariate complementarity can arise if different species dominate different processes. Further, these results suggest that similar mechanisms (complementary nutrient uptake) may underlie diversity-functioning relationships in both algal and vascular-plant-based systems.     

Physical and biological disturbances interact differently with productivity: effects on floral and faunal richness

Physical and biological disturbances are ecological processes affecting patterns in biodiversity at a range of scales in a variety of terrestrial and aquatic systems. Theoretical and empirical evidence suggest that effects of disturbance on diversity differ qualitatively and quantitatively, depending on levels of productivity (e.g., the dynamic equilibrium model). In this study we contrasted the interactive effects between physical disturbance and productivity to those between biological disturbance and productivity. Furthermore, to evaluate how these effects varied among different components of marine hard-substratum assemblages, analyses were done separately on algal and invertebrate richness, as well as richness of the whole assemblage. Physical disturbance (wave action) was simulated at five distinct frequencies, while biological disturbance (grazing periwinkles) was manipulated as present or absent, and productivity was manipulated as high or ambient. Uni- and multivariate analyses both showed significant effects of physical disturbance and interactive effects between biological disturbance and productivity on the composition of assemblages and total species richness. Algal richness was significantly affected by productivity and biological disturbance, whereas invertebrate richness was affected by physical disturbance only. Thus, we show, for the first time, that biological disturbance and physical disturbance interact differently with productivity, because these two types of disturbances affect different components of assemblages. These patterns might be explained by differences in the distribution (i.e., press vs. pulse) and degree of selectivity between disturbances. Because different types of disturbance can affect different components of assemblages, general ecological models will benefit from using natural diverse communities, and studies concerned with particular subsets of assemblages may be misleading. In conclusion, this study shows that the outcome of experiments on effects of disturbance and productivity on diversity is greatly influenced by the composition of the assemblage under study, as well as on the type of disturbance that is used as an experimental treatment.     

Organisation trophique de quatre peuplements de substrats rocheux selon un gradient de pollution industrielle (Golfe de Fos,France)

This study describes the trophic organization of the fauna of four superficial rock-bottom assemblages localized along a gradient of industrial pollution (Gulf of Fos, Mediterranean Sea). Six stations were studied over a period of 13 or 15 mo between May 1974 and August 1975. The faunal species (188 species) were classified into four major trophic groups: suspension-feeders, detritivores, herbivores and carnivores, which were all present in significant numbers in all four assemblages. A fifth group, the omnivores, was poorly represented. Qualitatively, the herbivores dominated by number of species (mainly Crustacea and Polychaeta). Quantitatively, the trophic groups were distributed in decreasing numbers along an increasing pollution gradient. Statistical analysis revealed a destabilization of the balance existing between the four major trophic groups along an increasing pollution gradient. Abundance of the four trophic groups exhibited spatio-temporal variations that evolved homogeneously during the annual biological cycle. When an environmental disturbance occurs, a particular trophic group will dominate, depending on the type of disturbance. At the reference station, located in relatively unpolluted waters, the three trophic groups suspension-feeders, herbivores and carnivores seemed to be well-balanced, with a slight dominance of the latter; the two other groups were distributed equally. At moderately polluted stations, suspension-feeders dominated. The most polluted stations were dominated by herbivores. Occurrence of detritivores seemed to be negatively related to the degree of thermal pollution, which provoked massive mortality and important detritus production resulting from internal stress caused by rapid temperature variations. At the population-structure level, such temperature variations are accompanied by important recruitment in a limited number of dominant species. Examination of the trophic organization of rock-bottom assemblages for which structural evolution is already well known provided complementary information on the modality of dysfunction in time and space of certain trophic-web levels, as related to progressive alterations in the rock-bottom assemblages. Such dysfunction can appear very early, even before evolution of the assemblage's physiognomical characteristics. Trends in trophic organization reflect the influence of certain edaphic factors such as particle-rich currents, the impact of local thermal pollution, chemical pollution, or low salinity.     

The impact of seabed disturbance on nematode communities: linking field and laboratory observations

Physical disturbance is a key factor in controlling the spatial and temporal composition of shallow-water benthic communities. Like shallow waters, deeper waters are increasingly subject to a range of anthropogenic disturbances, which can lead to significant alterations in sedimentation patterns. These alterations often exceed naturally occurring changes. We used a combined analysis of six independent data sets arising from large-scale field surveys and small-scale laboratory experiments to investigate the effects of seabed disturbance on nematode communities. Disturbance response was documented as a function of disturbance type (coastal development, dredged material disposal, bottom trawling, glacial fjord) and intensity (low, medium, high). Natural and man-induced seabed disturbance exerted differential effects on exposed populations, generating changes in the taxonomic (genus) and functional (feeding type) attributes of their assemblages. The genus composition of nematode assemblages from geographically separate seas converged with increased level of various types of man-made disturbance. Assemblages present along a gradient of natural disturbance in a glacial fjord followed an opposite response vector, suggesting that community changes induced by anthropogenic activities, or experimental treatments simulating the principal impacts of these, inherently differ from disturbance of natural origin. Changes in trophic diversity and structure were primarily driven by factors confounded with physical disturbance, such as metal contamination. Coupling the results of analyses at multiple scales proved a useful means of providing deeper insights into the general response of ecological communities to environmental change.     

The effects of direct and indirect constraints on biological communities

Human activities are expected to result in a diversity of directional or stochastic constraints that affect species either directly or by indirectly impacting their resources. However, there is no theoretical framework to predict the complex and various effects of these constraints on ecological communities. We developed a dynamic model that mimics the use of different resource types by a community of competing species. We investigated the effects of different environmental constraints (affecting either directly the growth rate of species or having indirect effects on their resources) on several biodiversity indicators. Our results indicate that (i) in realistic community models (assuming uneven resource requirements among species) the effects of perturbations are strongly buffered compared to neutral models; (ii) the species richness of communities can be maximized for intermediate levels of direct constraints (unimodal response), even in the absence of trade-off between competitive ability and tolerance to constraints; (iii) no such unimodal response occurs with indirect constraints; (iv) an increase in the environmental (e.g., climatic) variance may have different effects on community biomass and species richness.     

江苏省不同营养状况湖泊底栖动物群落结构与多样性比较

为了解江苏省湖泊底栖动物群落结构和多样性并研究其对水环境质量变化的响应,于2012年春秋两季对江苏省16个湖泊51个采样点湖泊底栖动物群落结构与多样性以及湖泊综合营养状态指数进行调查,分析水质指标与底栖动物指数间Pearson相关关系。结果表明,江苏省16个湖泊营养状态指数范围为35.5~66.4,其中约81.2%的湖泊处于轻度-中度富营养状态,表明水质从好到中度污染;湖泊底栖动物优势种为寡毛类的霍甫水丝蚓(Limnodrilus hoffmeisteri),优势度为13.0%;香浓多样性指数(Shannon-Wiener diversity index)范围为0.00~2.20,级别范围由极贫乏到较丰富状态,表明水质污染程度从重度到轻度污染。从全省尺度看,水质评价结果与生物学(香浓多样性指数、Pielou均匀度指数和Biotic Index(BI)指数)评价结果存在一定差异。与历史数据相比,江苏省湖泊底栖动物清洁敏感物种减少甚至消失,总体群落结构趋于小型化。16个湖泊水体总氮和总磷与底栖动物密度呈显著负相关,而综合营养状态指数与底栖动物指数(香浓多样性指数、Pielou均匀度指数和BI指数)间关系不显著。上述研究结果表明综合营养状态指数无法完全客观反映湖泊水生态健康状况,因此需要综合水质、水文、水生生物、生境状况等因素发展新的评价指标体系。     

Effectiveness of terrestrial protected areas for conservation of lake fish communities

Freshwater protected areas are rare even though freshwater ecosystems are among the most imperiled in the world. Conservation actions within terrestrial protected areas (TPAs) such as development or resource extraction regulations may spill over to benefit freshwater ecosystems within their boundaries. Using data from 175 lakes across Ontario, Canada, we compared common indicators of fish‐assemblage status (i.e., species richness, Shannon diversity index, catch per unit effort, and normalized‐length size spectrum slopes) to evaluate whether TPAs benefit lake fish assemblages. Nearest neighbor cluster analysis was used to generate pairs of lakes: inside versus outside, inside versus bordering, and bordering versus outside TPAs based on lake characteristics. The diversity and abundance indicators did not differ significantly across comparisons, but normalized‐length size spectrum slopes (NLSS) were significantly steeper in lakes outside parks. The latter indicated assemblage differences (greater abundances of small‐bodied species) and less‐efficient energy transfer through the trophic levels of assemblages outside parks. Although not significantly different, pollution‐ and turbidity‐tolerant species were more abundant outside parks, whereas 3 of the 4 pollution‐intolerant species were more abundant within parks. Twenty‐one percent of the difference in slopes was related to higher total dissolved solids concentrations and angling pressure. Our results support the hypothesis that TPAs benefit lake fish assemblages and suggest that NLSS slopes are informative indicators for aquatic protected area evaluations because they represent compositional and functional aspects of communities.     

Predators, prey, and transient states in the assembly of spatially structured communities

Ecological theory suggests that both dispersal limitation and resource limitation can exert strong effects on community assembly. However, empirical studies of community assembly have focused almost exclusively on communities with a single trophic level. Thus, little is known about the combined effects of dispersal and resource limitation on assembly of communities with multiple trophic levels. We performed a landscape-scale experiment using spatially arranged mesocosms to study effects of dispersal and resource limitation on the assembly dynamics of aquatic invertebrate communities with two trophic levels. We found that interplay between dispersal and resource limitation regulated the assembly of predator and prey trophic levels in these pond communities. Early in assembly, predators and prey were strongly dispersal limited, and resource (i.e., prey) availability did not influence predator colonization. Later in assembly, after predators colonized, resource limitation was the strongest driver of predator abundance, and dispersal limitation played a negligible role. Thus, habitat isolation affected predators directly by reducing predator colonization rate, and indirectly through the effect of distance on prey availability. Dispersal and resource limitation of predators resulted in a transient period in which predators were absent or rare in isolated habitats. This period may be important for understanding population dynamics of vulnerable prey species. Our findings demonstrate that dispersal and resource limitation can jointly regulate assembly dynamics in multi-trophic systems. They also highlight the need to develop a temporal picture of the assembly process in multi-trophic communities because the availability and spatial distribution of limiting resources (i.e., prey) and the distribution of predators can shift radically over time.     

Biotic habitat complexity controls species diversity and nutrient effects on net biomass production

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.     

Asymmetry in community regulation: effects of predators and productivity

Eutrophication and predator additions and extinctions are occurring in ecosystems worldwide. Although theory predicts that both will strongly alter the distribution of biomass in whole communities, empirical evidence has not been consolidated to quantitatively determine whether these theoretical predictions are generally borne out in real ecosystems. Here we analyze data from two types of trophic cascade studies, predator removals in factorial combination with fertilization and observed productivity gradients, to assess the role of top-down and bottom-up forces in structuring multi-trophic communities and compare results from these analyses to those from an extensive database of trophic cascade studies. We find that herbivore biomass declines and plant biomass increases in the presence of predators, regardless of system productivity. In contrast, while plants are increased by fertilization, this effect does not significantly increase herbivores in either the presence or absence of predators. These patterns are consistent among marine, freshwater, and terrestrial ecosystems and are largely independent of study size and duration. Thus, top-down effects of predation are transferred through more trophic levels than are bottom-up effects of eutrophication, showing strong asymmetry in the direction of control of biomass distribution in communities.     

Biomass and trophic structure of the macrobenthos in the south-west lagoon of New Caledonia

The biomass of the macrobenthic communities of the southwest lagoon of New Caledonia is analysed with special reference to tropical soft-bottom assemblages. The samples were obtained in 1986 from 350 Smith-McIntyre grabs as well as by diving over an area of 3 500 m², and covered a wide size-range of benthic organisms. The classification of the biomass data into taxonomic groups revealed the following: (1) a zoobenthic biomass (mean=13 g ash-free dry wt m^-2) that is high compared to values previously published for tropical systems, but fairly low compared to those recorded for medium and high latitudes; (2) a large macrophytic biomass (45% of ash-free dry wt); (3) dominance by weight of sponges, molluscs and cnidarians over the other faunal groups. The distribution of the biomass among the trophic groups revealed the following: (1) the dominance by weight of suspension-feeders, which account for 67% of the total zoobenthic ash-free dry wt; (2) the absence of trophic group amensalism between filter-feeders and deposit-feeders whatever the degree of bottom muddiness; (3) the existence of a relationship between the biomass of surface deposit-feeders and ATP at the surface layer of the sediment, suggesting that this group feeds preferentially on living particles. Differences in trophic structure between the muddy-bottom, greysand and white-sand communities of the lagoon reflect the functional characteristics of each bottom type.     

Malthusian overfishing and efforts to overcome it on Kenyan coral reefs

This study examined trends along a gradient of fishing intensity in an artisanal coral reef fishery over a 10-year period along 75 km of Kenya's most populated coastline. As predicted by Malthusian scenarios, catch per unit effort (CPUE), mean trophic level, the functional diversity of fished taxa, and the diversity of gear declined, while total annual catch and catch variability increased along the fishing pressure gradient. The fishery was able to sustain high (approximately 16 Mg x km(-2) x yr(-1)) but variable yields at high fishing pressure due to the dominance of a few productive herbivorous fish species in the catch. The effect of two separate management strategies to overcome this Malthusian pattern was investigated: fisheries area closure and elimination of the dominant and most "competitive" gear. We found that sites within 5 km of the enforced closure showed significantly lower total catch and CPUE, but increased yield stability and trophic level of catch than predicted by regression models normalized for fishing effort. Sites that had excluded illegal beach seine use through active gear management exhibited increased total catch and CPUE. There was a strong interaction between closure and gear management, which indicates that, for closures to be effective at increasing catch, there must be simultaneous efforts at gear management around the periphery of the closures. We propose that Malthusian effects are responsible for the variation in gear and catch and that active management through reduced effort and reductions in the most competitive gear have the greatest potential to increase the functional and trophic diversity and per-person productivity.     

Long-term trends in the trophic structure of the North Sea fish community: evidence from stable-isotope analysis, size-spectra and community metrics

Fishing has wide-ranging impacts on marine ecosystems. One of the most pervasive signs of intensive fishing is "fishing down the food web", with landings increasingly dominated by smaller species from lower trophic levels. Decreases in the trophic level of landings are assumed to reflect those in fish communities, because size-selective mortality causes decreases in the relative abundance of larger species and in mean body size within species. However, existing analyses of fishing impacts on the trophic level of fish communities have focused on the role of changes in species composition rather than size composition. This will provide a biased assessment of the magnitude of fishing impacts, because fishes feed at different trophic levels as they grow. Here, we combine body size versus trophic level relationships for North Sea fishes (trophic level assessed using nitrogen stable-isotope analysis) with species-size-abundance data from two time-series of trawl-survey data (whole North Sea 1982-2000, central and northern North Sea 1925-1996) to predict long-term trends in the trophic structure of the North Sea fish community. Analyses of the 1982-2000 time-series showed that there was a slow but progressive decline in the trophic level of the demersal community, while there was no trend in the trophic level of the combined pelagic and demersal community. Analyses of the longer time-series suggested that there was no trend in the trophic level of the demersal community. We related temporal changes in trophic level to temporal changes in the slopes of normalised biomass size-spectra (which theoretically represent the trophic structure of the community), mean log₂ body mass and mean log₂ maximum body mass. While the size-based metrics of community structure showed long-term trends that were consistent with the effects of increased fishery exploitation, these trends were only correlated with trophic level for the demersal community. Our analysis suggests that the effects of fishing on the trophic structure of fish communities can be much more complex than previously assumed. This is a consequence of sampled communities not reflecting all the pathways of energy transfer in a marine ecosystem and of the absence of historical data on temporal and spatial changes in the trophic level of individuals. For the North Sea fish community, changes in size structure due to the differential effects of fishing on species and populations with different life histories are a stronger and more universal indicator of fishing effects than changes in mean trophic level.     

Predators have large effects on ecosystem properties by changing plant diversity, not plant biomass

Cascading effects of predators on total plant trophic-level biomass tend to be weaker in terrestrial than in aquatic systems. Accordingly, it is hypothesized that top predator effects on terrestrial plant diversity and on ecosystem function should likewise be weak or unimportant. This report presents a test of this hypothesis using data from a long-term field experiment. The five-year experiment manipulated the trophic structure of an old field ecosystem by excluding either predators or predators and herbivores relative to an unmanipulated, natural control. Long-term manipulations led to systematic treatment effects on community properties (plant trophic-level biomass, plant species biomass, plant species evenness) and on ecosystem properties (supply rate of solar radiation, N mineralization rate). The strengths of top predator effects on community properties were modest compared with nonterrestrial systems. But, predator-caused changes in plant community structure via alteration of plant dominance, and hence plant species evenness, strengthened effects on ecosystem properties. Counter to the hypothesis, weak trophic cascades do not necessarily lead to weak indirect effects of predators on ecosystem properties.