Trophic structure of the spider community of a Mediterranean citrus grove: A stable isotope analysis

Spiders are dominant terrestrial predators that consume a large variety of prey and engage in intraguild predation. Although the feeding habits of certain species are well known, the trophic structure of spider assemblages still needs to be investigated. Stable isotope analysis enables characterisation of trophic relationships between organisms because it tracks the energy flow in food webs and indicates the average number of trophic transfers between a given species and the base of the web, thus being a useful tool to estimate the magnitude of intraguild predation in food webs. Using this technique, we studied the trophic groups of spiders and their links within the arthropod food web of a Mediterranean organic citrus grove. We assessed the trophic positions of the 25 most common spider species relative to other arthropod predators and potential prey in the four seasons of the year, both in the canopy and on the ground. The analyses showed great seasonal variation in the isotopic signatures of some arthropod species, as well as the existence of various trophic groups and a wide range of trophic levels among spiders, even in species belonging to the same family. Differences in δ¹⁵N between spiders and the most abundant prey in the grove usually spanned two trophic levels or more. Our findings provide field evidence of widespread intraguild predation in the food web and caution against using spider families or guilds instead of individual species when studying spider trophic interactions.     

High‐resolution food webs based on nitrogen isotopic composition of amino acids

Food webs are known to have myriad trophic links between resource and consumer species. While herbivores have well‐understood trophic tendencies, the difficulties associated with characterizing the trophic positions of higher‐order consumers have remained a major problem in food web ecology. To better understand trophic linkages in food webs, analysis of the stable nitrogen isotopic composition of amino acids has been introduced as a potential means of providing accurate trophic position estimates. In the present study, we employ this method to estimate the trophic positions of 200 free‐roaming organisms, representing 39 species in coastal marine (a stony shore) and 38 species in terrestrial (a fruit farm) environments. Based on the trophic positions from the isotopic composition of amino acids, we are able to resolve the trophic structure of these complex food webs. Our approach reveals a high degree of trophic omnivory (i.e., noninteger trophic positions) among carnivorous species such as marine fish and terrestrial hornets.This information not only clarifies the trophic tendencies of species within their respective communities, but also suggests that trophic omnivory may be common in these webs.     

Measuring the trophic ecology of ants using stable isotopes

Ants are prominent components of most terrestrial arthropod food webs, yet due to their highly variable diet, the role ants play in arthropod communities can be difficult to resolve. Stable isotope analysis is a promising method for determining the dietary history of an organism, and has the potential to advance our understanding of the food web ecology of social insects. However, some unique characteristics of eusocial organisms can complicate the application of this technique to the study of their trophic ecology. Using stable isotopes of N and C, we investigated levels of intraspecific variation both within and among colonies. We also examined the effect of a common preservation technique on δ¹⁵N and δ¹³C values. We discuss the implications of our results on experimental design and sampling methods for studies using stable isotopes to investigate the trophic ecology of social insects. Received 4 February 2005; revised 23 June 2005; accepted 4 July 2005.     

Predator hunting mode influences patterns of prey use from grazing and epigeic food webs

Multichannel omnivory by generalist predators, especially the use of both grazing and epigeic prey, has the potential to increase predator abundance and decrease herbivore populations. However, predator use of the epigeic web (soil surface detritus/microbe/algae consumers) varies considerably for reasons that are poorly understood. We therefore used a stable isotope approach to determine whether prey availability and predator hunting style (active hunting vs. passive web-building) impacted the degree of multichannel omnivory by the two most abundant predators on an intertidal salt marsh, both spiders. We found that carbon isotopic values of herbivores remained constant during the growing season, while values for epigeic feeders became dramatically more enriched such that values for the two webs converged in August. Carbon isotopic values for both spider species remained midway between the two webs as values for epigeic feeders shifted, indicating substantial use of prey from both food webs by both spider species. As the season progressed, prey abundance in the grazing food web increased while prey abundance in the epigeic web remained constant or declined. In response, prey consumption by the web-building spider shifted toward the grazing web to a much greater extent than did consumption by the hunting spider, possibly because passive web-capture is more responsive to changes in prey availability. Although both generalist predator species engaged in multichannel omnivory, hunting mode influenced the extent to which these predators used prey from the grazing and epigeic food webs, and could thereby influence the strength of trophic cascades in both food webs.     

Stable isotope ecology in insects: a review

1. The use of stable isotope analysis (SIA) in ecological research has dramatically increased in recent years largely because it allows researchers to investigate ecological questions that have been previously difficult to address. 2. Ecological applications of SIA include estimating fundamental niche space and overlap, evaluating trophic or species level interactions, and investigating food web structure. Increasingly, researchers have been incorporating SIA in studies of animal migration, disease transmission, diet composition, nutrient assimilation, and body condition among others. 3. Studies using SIA to evaluate the ecology of terrestrial insects have lagged behind other taxonomic groups. This poor representation of stable isotope studies in publications likely stems from a lack of familiarity of entomologists with this technique. 4. An improved understanding of SIA, as well as the advantages and disadvantages specifically related to insect research, will benefit the field of entomology. In addition, insect-model systems provide unique opportunities for entomologists to incorporate SIA in their research to advance our knowledge of insect biology and the stable isotope ecology of insects. 5. We provide background information on stable isotopes, explain sources of isotopic variation, describe the processes of how isotopes are differentially routed and incorporated into an individual's tissues, explain the principles that influence isotopic fractionation and discrimination, highlight different methods and advancements in SIA, review innovative stable isotope studies, and provide an overview of common mistakes, considerations, and future directions entomologists can explore.     

Linking aboveground and belowground food webs through carbon and nitrogen stable isotope analyses

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) have been used for more than two decades in analyses of food web structure. The utility of isotope ratio measurements is based on the observation that consumer δ¹³C values are similar (<1‰ difference) to those of their diet, while consumer δ¹⁵N values are about 3‰ higher than those of their diet. The technique has been applied most often to aquatic and aboveground terrestrial food webs. However, few isotope studies have examined terrestrial food web structure that includes both above- and belowground (detrital) components. Here, we review factors that may influence isotopic signatures of terrestrial consumers in above- and belowground systems. In particular, we emphasize variations in δ¹³C and δ¹⁵N in belowground systems, e.g., enrichment of ¹³C and ¹⁵N in soil organic matter (likely related to soil microbial metabolism). These enrichments should be associated with the high ¹³C (~3‰) enrichment in belowground consumers relative to litter and soil organic matter and with the large variation in δ¹⁵N (~6‰) of the consumers. Because such enrichment and variation are much greater than the trophic enrichment generally used to estimate consumer trophic positions, and because many general predators are considered dependent on energy and material flows from belowground, the isotopic variation in belowground systems should be taken into account in δ¹³C and δ¹⁵N analyses of terrestrial food webs. Meanwhile, by measuring the δ¹³C of key predators, the linkage between above- and belowground systems could be estimated based on observed differences in δ¹³C of primary producers, detritivores and predators. Furthermore, radiocarbon (¹⁴C) measurements will allow the direct estimation of the dependence of predators on the belowground systems.     

Structure of tropical river food webs revealed by stable isotope ratios

Fish assemblages in tropical river food webs are characterized by high taxonomic diversity, diverse foraging modes, omnivory, and an abundance of detritivores. Feeding links are complex and modified by hydrologic seasonality and system productivity. These properties make it difficult to generalize about feeding relationships and to identify dominant linkages of energy flow. We analyzed the stable carbon and nitrogen isotope ratios of 276 fishes and other food web components living in four Venezuelan rivers that differed in basal food resources to determine 1) whether fish trophic guilds integrated food resources in a predictable fashion, thereby providing similar trophic resolution as individual species, 2) whether food chain length differed with system productivity, and 3) how omnivory and detritivory influenced trophic structure within these food webs. Fishes were grouped into four trophic guilds (herbivores, detritivores/algivores, omnivores, piscivores) based on literature reports and external morphological characteristics. Results of discriminant function analyses showed that isotope data were effective at reclassifying individual fish into their pre-identified trophic category. Nutrient-poor, black-water rivers showed greater compartmentalization in isotope values than more productive rivers, leading to greater reclassification success. In three out of four food webs, omnivores were more often misclassified than other trophic groups, reflecting the diverse food sources they assimilated. When fish δ¹⁵N values were used to estimate species position in the trophic hierarchy, top piscivores in nutrient-poor rivers had higher trophic positions than those in more productive rivers. This was in contrast to our expectation that productive systems would promote longer food chains. Although isotope ratios could not resolve species-level feeding pathways, they did reveal how top consumers integrate isotopic variability occurring lower in the food web. Top piscivores, regardless of species, had carbon and nitrogen profiles less variable than other trophic groups.     

Omnivory and stability of food webs

The role and prevalence of omnivory, defined as feeding on more than one trophic level, are critical to understand food web structure and dynamics. Whether omnivory stabilizes or destabilizes food webs depends on the assumptions of theoretical models. Recently, Tanabe and Namba [Tanabe, K., Namba, T., 2005. Omivory creates chaos in simple food web models. Ecology 86, 3411–3414] found that omnivory can create chaos in a simple food web model with linear functional responses and 12 model parameters. In this paper, first we numerically examined bifurcation diagrams with all the parameters as bifurcation parameters, including self-limitation of the intermediate consumer and predator. Chaos spontaneously appears when the intraguild predator’s consumption rates are low for nutrient-rich intraguild prey and high for nutrient-poor basal resource and the intraguild prey reproduces efficiently feeding on the basal resource. Second, we investigated effects of the addition of a species into the basic model food web which exhibits chaos. The additional species is assumed to consume only one of the basal resource, intermediate consumer, or omnivorous predator. Consequences of the addition greatly depend on the trophic level on which the additional species feeds. While the increased diversity of predators feeding on the intermediate consumer stabilizes the web, the increased diversity of prey feeding on the basal resource induces collapse of the food web through exploitative competition for the basal resource. The food chain with the top predator feeding on the omnivorous predator is highly unstable unless the mortality of the top predator is extremely low. We discuss the possibility of real-world chaos and the reason why stability of food webs strongly depends on the topological structure of the webs. Finally, we consider the implications of our results for food web theory and resource management.     

稳定同位素技术在鲨鱼摄食和洄游行为研究中的应用

随着稳定同位素分析技术的不断成熟,其在生态学领域中的应用也增长迅速,并成为动物摄食生态学的重要研究工具.鲨鱼因其在生物系统进化过程中的独特地位和海洋生态系统中的重要作用已成为海洋食物网研究的重点,然而国内针对鲨鱼摄食习性和洄游行为等方面的研究仍处于起步阶段.本文在总结了国内外鲨鱼稳定同位素分析组织样品选取和样品预处理方法的基础上,系统归纳了稳定同位素技术在鲨鱼摄食生态学,尤其在其摄食和洄游行为研究领域中的应用,着重分析稳定同位素技术在鲨鱼稳定同位素判别值和更新速率、食性分析、营养级评估、洄游路径分析和生态位分布等核心问题上的应用现状和发展前景,以期为国内学者开展鲨鱼类基础生物、生态学研究提供有益参考.     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Use of compound-specific nitrogen isotope analysis of amino acids in trophic ecology: assumptions,applications, and implications

Knowledge of the diet source and trophic position of organisms is fundamental in food web science. Since the 1980s, stable isotopes of light elements such as ¹³C and ¹⁵N have provided unique information on the food web structure in a variety of ecosystems. More recently, novel isotope tools such as stable isotopes of heavy elements, radioisotopes, and compound-specific isotope analysis, have been examined by researchers. Here I reviewed the use of compound-specific nitrogen isotope analysis of amino acids (CSIA-AA) as a useful dietary tracer in food web ecology. Its three key features—(1) offsetting against isotopic variation; (2) universality of the trophic discrimination factor; and (3) sensitivity to source mixing—were compared with conventional isotope analysis for the bulk tissue of organisms. These three advantages of CSIA-AA will allow future researchers to (1) estimate the integrated trophic position (iTP) of animal communities; (2) infer trophic transfer efficiency (TTE) in food webs; and (3) quantify contributions from different resources to animals, all of which are crucial for understanding the relationship between biodiversity and multi-trophic ecosystem functioning. Further development of trophic ecology will be facilitated by both methodological refinement of CSIA-AA and its application to a wider range of organisms, food webs, and ecosystems.     

Tracing energy flow in stream food webs using stable isotopes of hydrogen

1. Use of the natural ratios of carbon and nitrogen stable isotopes as tracers of trophic interactions has some clear advantages over alternative methods for food web analyses, yet is limited to situations where organic materials of interest have adequate isotopic separation between potential sources. This constrains the use of natural abundance stable isotope approaches to a subset of ecosystems with biogeochemical conditions favourable to source separation. 2. Recent studies suggest that stable hydrogen isotopes (δD) could provide a robust tracer to distinguish contributions of aquatic and terrestrial production in food webs, but variation in δD of consumers and their organic food sources are poorly known. To explore the utility of the stable hydrogen isotope approach, we examined variation in δD in stream food webs in a forested catchment where variation in δ¹³C has been described previously. 3. Although algal δD varied by taxa and, to a small degree, between sites, we found consistent and clear separation (by an average of 67‰) from terrestrial carbon sources. Environmental conditions known to affect algal δ¹³C, such as water velocity and stream productivity did not greatly influence algal δD, and there was no evidence of seasonal variation. In contrast, algal δ¹³C was strongly affected by environmental factors both within and across sites, was seasonally variable at all sites, and partially overlapped with terrestrial δ¹³C in all streams with catchment areas larger than 10 km². 4. While knowledge of isotopic exchange with water and trophic fractionation of δD for aquatic consumers is limited, consistent source separation in streams suggests that δD may provide a complementary food web tracer to δ¹³C in aquatic food webs. Lack of significant seasonal or spatial variation in δD is a distinct advantage over δ¹³C for applications in many aquatic ecosystems.     

基于稳定同位素的湿地食物源判定和食物网构建研究进展

湿地生物营养动力学是湿地生态系统结构和功能评价研究的基础.碳、氮稳定同位素作为识别营养关系的方法,已在湿地生态系统食物来源、组成和食物链传递研究中得到广泛运用.本文系统综述了稳定同位素食物贡献度计算模型和营养级确定的基本方法和理论;讨论了动物营养分馏值和基线的选择依据;概括了湿地生态系统典型食物源及其稳定同位素变化特征;总结了草食、杂食和肉食等不同营养级动物的食物来源.指出了稳定同位素在湿地食物源溯源和食物网研究中的不足;基于国内外研究现状和发展趋势及需求,展望了未来同位素技术在湿地食物网生态学研究中的运用前景和研究重点,提出需要加强稳定同位素营养分馏和基线的影响因素、样品处理和保存方式研究以及胃含物、分子标记物和多元素同位素结合分析.     

Niche‐dependent trophic position distributions among primary,secondary and tertiary consumers

Many consumers display flexible feeding strategies that vary among individuals or populations, through life‐history, or spatiotemporally. Despite the recognized influence of flexible feeding on the structure and dynamics of food webs, the consequences of these feeding strategies on the actual shape and characteristics of trophic position distributions have received less attention. We proposed and tested several a priori hypotheses to predict the likely effect of niche‐dependent (e.g. herbivore, secondary consumer) foraging on the shape and statistical properties of consumer trophic position distributions using natural abundance stable isotope data from a diverse dataset of consumers. We found evidence that the structural characteristics of consumer trophic position distributions varied as a function of trophic niche. Herbivores and tertiary consumers tended to be ‘packed’ closely near their mean trophic position, with few individuals realizing trophic positions markedly higher or lower than the mean. Conversely, secondary consumers often displayed broad trophic position distributions with many individuals dispersed away from the center of the distribution. We examined the effect of applying constant versus dynamic isotope trophic fractionation models and found that both models yielded similar although not identical results. Our findings suggest that trophic level omnivory supports a larger fraction of consumer diet at intermediate trophic positions than at either the lowest or the highest positions in aquatic food webs. These results suggest that vertical trophic niche declines among higher order consumers despite general evidence that the range of potential foraging options (i.e. horizontal trophic niche) tends to increase at higher trophic positions. Although further work is needed to test the generality of these patterns in other ecosystems, proactively examining trophic position distributions and reporting appropriate measures of central tendency (e.g. arithmetic versus geometric means) will increase the accuracy of individual trophic studies as well as the applicability of results for meta‐analytical food web models.     

稳定性同位素技术在生态学上的应用

稳定性同位素技术早在20世纪70年代末期就被引入到生态学领域。最初是利用植物稳定性碳同位素的差异。开展了许多有关营养流动方面的研究;到90年代,稳定性碳和氮同位素被用来分析动物的食性、营养级位置关系以及食物链结构;本世纪初,由于技术的进步,稳定性同位素(特别是氢同位素)被用来开展动物迁徙习性方面的研究。到目前为止,国内有关这方面的研究还鲜有报道,而且对自然界存在的稳定性同位素的理解还存在一定偏差。本文主要介绍了稳定性同位素效应及其分馏原理、稳定性同位素在示踪动物食性信息、确定营养级位置关系、分析食物网结构以及研究动物迁徙生态学中的作用等方面的内容。     

Combined stable‐isotope and fatty‐acid analyses demonstrate that large wood increases the autochthonous trophic base of a macroinvertebrate assemblage

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Effects of Multi-chain Omnivory on the Strength of Trophic Control in Lakes

Omnivory has been implicated in both diffusing and intensifying the effects of consumer control in food chains. Some have postulated that the strong, community level, top-down control apparent in lakes is not expressed in terrestrial systems because terrestrial food webs are reticulate, with high degrees of omnivory and diverse plant communities. In contrast, lake food webs are depicted as simple linear chains based on phytoplankton-derived energy. Here, we explore the dynamic implications of recent evidence showing that attached algal (periphyton) carbon contributes substantially to lake primary and secondary productivity, including fish production. Periphyton production represents a cryptic energy source in oligotrophic and mesotrophic lakes that is overlooked by previous theoretical treatment of trophic control in lakes. Literature data demonstrate that many fish are multi-chain omnivores, exploiting food chains based on both littoral and pelagic primary producers. Using consumer-resource models, we examine how multiple food chains affect fourth-level trophic control across nutrient gradients in lakes. The models predict that the stabilizing effects of linked food chains are strongest in lakes where both phytoplankton and periphyton contribute substantially to production of higher trophic levels. This stabilization enables a strong and persistent top down control on the pelagic food chain in mesotrophic lakes. The extension of classical trophic cascade theory to incorporate more complex food web structures driven by multi-chain predators provides a conceptual framework for analysis of reticulate food webs in ecosystems.     

Omnivory as a stabilizing feature of natural communities

Abstract Omnivory-defined broadly as feeding on more than one trophic level-occupies a prominent position in discussions of food web architecture and dynamics, due in large part to an enduring conflict regarding omnivory's role in community dynamics. According to classical results from mathematical food web theory, omnivory destabilizes ecological communities, whereas more recent conceptual syntheses suggest that omnivory should be a strongly stabilizing factor in food webs. Working with an arthropod assemblage at Mount Saint Helens, I experimentally addressed this controversy using a two-way factorial design that crossed a manipulation of the degree of omnivory with another "disturbance" manipulation that targeted a specific component of the assemblage. In this statistical design, significant interaction effects (i.e., how the community impacts of the disturbance varied with the degree of omnivory) identified key stabilizing or destabilizing influences of omnivory. Overall, my experimental results indicated that increasing the degree of omnivory stabilized community dynamics, in keeping with recent conceptual syntheses.     

Grassland harvesting alters ant community trophic structure: An isotopic study in tallgrass prairies

Disturbances have long been recognized as important forces for structuring natural communities but their effects on trophic structure are not well understood, particularly in terrestrial systems. This is in part because quantifying trophic linkages is a challenge, especially for small organisms with cryptic feeding behaviors such as insects, and often relies on conducting labor‐intensive feeding trials or extensive observations in the field. In this study, we used stable isotopes of carbon and nitrogen to examine how disturbance (annual biomass harvesting) in tallgrass prairies affected the trophic position, trophic range, and niche space of ants, a widespread grassland consumer. We hypothesized that biomass harvest would remove important food and nesting resources of insects thus affecting ant feeding relationships and trophic structure. We found shifts in the feeding relationships inferred by isotopic signatures with harvest. In particular, these shifts suggest that ants within harvest sites utilized resources at lower trophic levels (possibly plant‐based resources or herbivores), expanded trophic breadth, and occupied different niche spaces. Shifts in resource use following harvest could be due to harvest‐mediated changes in both the plant and arthropod communities that might affect the strength of competition or alter plant nitrogen availability. Because shifts in resource use alter the flow of nutrients across the food web, disturbance effects on ants could have ecosystem‐level consequences through nutrient cycling.