Aboveground mammal and invertebrate exclusions cause consistent changes in soil food webs of two subalpine grassland types,but mechanisms are system‐specific

Ungulates, smaller mammals, and invertebrates can each affect soil biota through their influence on vegetation and soil characteristics. However, direct and indirect effects of the aboveground biota on soil food webs remain to be unraveled. We assessed effects of progressively excluding aboveground large‐, medium‐ and small‐sized mammals as well as invertebrates on soil nematode diversity and feeding type abundances in two subalpine grassland types: short‐ and tall‐grass vegetation. We explored pathways that link exclusions of aboveground biota to nematode feeding type abundances via changes in plants, soil environment, soil microbial biomass, and soil nutrients. In both vegetation types, exclusions caused a similar shift toward higher abundance of all nematode feeding types, except plant feeders, lower Shannon diversity, and lower evenness. These effects were strongest when small mammals, or both small mammals and invertebrates were excluded in addition to excluding larger mammals. Exclusions resulted in a changed abiotic soil environment that only affected nematodes in the short‐grass vegetation. In each vegetation type, exclusion effects on nematode abundances were mediated by different drivers related to plant quantity and quality. In the short‐grass vegetation, not all exclusion effects on omni–carnivorous nematodes were mediated by the abundance of lower trophic level nematodes, suggesting that omni–carnivores also depended on other prey than nematodes. We conclude that small aboveground herbivores have major impacts on the soil food web of subalpine short‐ and tall‐grass ecosystems. Excluding aboveground animals caused similar shifts in soil nematode assemblages in both subalpine vegetation types, however, mechanisms turned out to be system‐specific.     

SSU Ribosomal DNA-Based Monitoring of Nematode Assemblages Reveals Distinct Seasonal Fluctuations within Evolutionary Heterogeneous Feeding Guilds

Soils are among the most complex, diverse and competitive habitats on Earth and soil biota are responsible for ecosystem services such as nutrient cycling, carbon sequestration and remediation of freshwater. The extreme biodiversity prohibits the making of a full inventory of soil life. Hence, an appropriate indicator group should be selected to determine the biological condition of soil systems. Due to their ubiquity and the diverse responses to abiotic and biotic changes, nematodes are suitable indicators for environmental monitoring. However, the time-consuming microscopic analysis of nematode communities has limited the scale at which this indicator group is used. In an attempt to circumvent this problem, a quantitative PCR-based tool for the detection of a consistent part of the soil nematofauna was developed based on a phylum-wide molecular framework consisting of 2,400 full-length SSU rDNA sequences. Taxon-specific primers were designed and tested for specificity. Furthermore, relationships were determined between the quantitative PCR output and numbers of target nematodes. As a first field test for this DNA sequence signature-based approach, seasonal fluctuations of nematode assemblages under open canopy (one field) and closed canopy (one forest) were monitored. Fifteen taxa from four feeding guilds (covering ∼ 65% of the free-living nematode biodiversity at higher taxonomical level) were detected at two trophic levels. These four feeding guilds are composed of taxa that developed independently by parallel evolution and we detected ecologically interpretable patterns for free-living nematodes belonging to the lower trophic level of soil food webs. Our results show temporal fluctuations, which can be even opposite within taxa belonging to the same guild. This research on nematode assemblages revealed ecological information about the soil food web that had been partly overlooked.     

Effects of native and exotic range‐expanding plant species on taxonomic and functional composition of nematodes in the soil food web

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range‐expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range‐expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root‐feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root‐feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root‐feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range‐expanding plant species can have fewer root‐feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web.     

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.     

Testing for top‐down cascading effects in a biomass‐driven ecological network of soil invertebrates

1. To investigate the structural changes of a food‐web architecture, we considered real data coming from a soil food web in one abandoned pasture with former low‐pressure agriculture management and we reproduced the corresponding ecological network within a multi‐agent fully programmable modeling environment in order to simulate dynamically the cascading effects due to the removal of entire functional guilds.
2. We performed several simulations differing from each other for the functional implications. At the first trophic level, we simulated a removal of the prey, that is, herbivores and microbivores, while at the second trophic level, we simulated a removal of the predators, that is, omnivores and carnivores. The five main guilds were removed either separately or in combination.
3. The alteration in the food‐web architecture induced by the removal of entire functional guilds was the highest when the entire second trophic level was removed, while the removal of all microbivores caused an alteration in the food‐web structure of less than 5% of the total changes due to the removal of opportunistic and predatory species.
4. Omnivores alone account for the highest shifts in time of the numerical abundances of the remaining species, providing computational evidence of the importance of the degree of omnivory in the stabilization of soil biota.

     

Plant species identity and diversity effects on different trophic levels of nematodes in the soil food web

Previous studies on biodiversity and soil food web composition have mentioned plant species identity, as well as plant species diversity as the main factors affecting the abundance and diversity of soil organisms. However, most studies have been carried out under limitations of time, space, or appropriate controls. In order to further examine the relation between plant species diversity and the soil food web, we conducted a three-year semi-field experiment in which eight plant species (4 forb and 4 grass species) were grown in monocultures and mixtures of two, four and eight plant species. In addition there were communities with 16 plant species. We analyzed the abundance and identity of the nematodes in soil and roots, including feeding groups from various trophic levels (primary and secondary consumers, carnivores, and omnivores) in the soil food web.
Plant species diversity and plant identity affected the diversity of nematodes. The effect of plant diversity was attributed to the complementarity in resource quality of the component plant species rather than to an increase in total resource quantity. The nematode diversity varied more between the different plant species than between different levels of plant species diversity, so that plant identity is more important than plant diversity. Nevertheless the nematode diversity in plant mixtures was higher than in any of the plant monocultures, due to the reduced dominance of the most abundant nematode taxa in the mixed plant communities. Plant species identity affected the abundances of the lower trophic consumer levels more than the higher trophic levels of nematodes. Plant species diversity and plant biomass did not affect nematode abundance. Our results, therefore, support the hypothesis that resource quality is more important than resource quantity for the diversity of soil food web components and that plant species identity is more important than plant diversity per se.     

Free‐living nematodes as prey for higher trophic levels of forest soil food webs

Nematodes are the most abundant invertebrates in soils and are key prey in soil food webs. Uncovering their contribution to predator nutrition is essential for understanding the structure of soil food webs and the way energy channels through soil systems. Molecular gut content analysis of consumers of nematodes, such as soil microarthropods, using specific DNA markers is a novel approach for studying predator–prey interactions in soil. We designed new specific primer pairs (partial 18S rDNA) for individual soil‐living bacterial‐feeding nematode taxa (Acrobeloides buetschlii, Panagrellus redivivus, Plectus velox and Plectus minimus). Primer specificity was tested against more than 100 non‐target soil organisms. Further, we determined how long nematode DNA can be traced in the gut of predators. Potential predators were identified in laboratory experiments including nine soil mite (Oribatida, Gamasina and Uropodina) and ten springtail species (Collembola). Finally, the approach was tested under field conditions by analyzing five mite and three collembola species for feeding on the three target nematode species. The results proved the three primer sets to specifically amplify DNA of the respective nematode taxa. Detection time of nematode DNA in predators varied with time of prey exposure. Further, consumption of nematodes in the laboratory varied with microarthropod species. Our field study is the first definitive proof that free‐living nematodes are important prey for a wide range of soil microarthropods including those commonly regarded as detritivores. Overall, the results highlight the eminent role of nematodes as prey in soil food webs and for channelling bacterial carbon to higher trophic levels.     

Habitat size influences food web structure in drying streams

Biodiversity in running waters is threatened by an increased severity and incidence of low‐flow extremes resulting from global climate change and a growing human demand for freshwater resources. Although it is unknown how and to what extent riverine communities will change in the face of these threats, considerable insight will be gained from efforts aimed at quantifying habitat size‐related controls on the trophic relationships among taxa in streams experiencing extreme flow loss. Here we report on a detailed space‐for‐time survey of replicate stream food webs sampled along the perennial‐ to‐drying continuum in each of fourteen different intermittent South Island, New Zealand streams. We quantified several structural attributes of food webs at fifty‐eight sites, including two taxonomically‐based metrics (web size, predator:prey ratio) and three stable isotope‐based metrics (food chain length [FCL], trophic area, δ¹³C range); we also quantified habitat size‐, disturbance‐, and resource‐related covariates at each site. Food web structure varied widely across sample sites within and across study streams and much of this variation was explained by habitat size. Consistent with our predictions, we found that food webs became smaller (ca 30 to ca 15 taxa, ca 20‐fold reduction in stable isotope‐based trophic area) and shorter (maximum trophic position [FCL] from 4.1 to 2.0, 25% reduction in predator:prey ratio) as we moved from the largest to smaller habitats. These results, and a comparison of our findings with those from a similar assessment conducted in perennial streams, suggest that there are perturbation thresholds which may trigger food web collapse when exceeded, and further imply that food webs may ultimately be ‘sized’ to minimum flows rather than average flow conditions. Our work provides a basis for making general predictions about how habitat contraction, and flow loss in particular, may affect communities and additionally provides insight on mechanisms warranting further attention.     

Soil Nematode Responses to Increases in Nitrogen Deposition and Precipitation in a Temperate Forest

The environmental changes arising from nitrogen (N) deposition and precipitation influence soil ecological processes in forest ecosystems. However, the corresponding effects of environmental changes on soil biota are poorly known. Soil nematodes are the important bioindicator of soil environmental change, and their responses play a key role in the feedbacks of terrestrial ecosystems to climate change. Therefore, to explore the responsive mechanisms of soil biota to N deposition and precipitation, soil nematode communities were studied after 3 years of environmental changes by water and/or N addition in a temperate forest of Changbai Mountain, Northeast China. The results showed that water combined with N addition treatment decreased the total nematode abundance in the organic horizon (O), while the opposite trend was found in the mineral horizon (A). Significant reductions in the abundances of fungivores, plant-parasites and omnivores-predators were also found in the water combined with N addition treatment. The significant effect of water interacted with N on the total nematode abundance and trophic groups indicated that the impacts of N on soil nematode communities were mediated by water availability. The synergistic effect of precipitation and N deposition on soil nematode communities was stronger than each effect alone. Structural equation modeling suggested water and N additions had direct effects on soil nematode communities. The feedback of soil nematodes to water and nitrogen addition was highly sensitive and our results indicate that minimal variations in soil properties such as those caused by climate changes can lead to severe changes in soil nematode communities.     

Selective alteration of soil food web components by invasive giant goldenrod Solidago gigantea in two distinct habitat types

Apart from relatively well‐studied aboveground effects, invasive plant species will also impact the soil food web. So far, most research has been focusing on primary decomposers, while studies on effects at higher trophic levels are relatively scarce. Giant goldenrod Solidago gigantea, native to North America, is a widespread and common invasive species in most European countries. We investigated its impact on plant communities and on multiple trophic levels of the soil food web in two contrasting habitats: riparian zones and semi‐natural grasslands. In 30 pairs of invaded and uninvaded plots, floristic composition, pH, fungal biomass and the densities of 11 nematode taxa were determined by using a quantitative PCR‐based method. In the two habitats, the invader outcompeted both rare and dominant plant species. Belowground, S. gigantea invasion reduced pH, increased overall fungal biomass as well as the density of a single lineage of fungivorous nematodes, the family Aphelenchoididae. The densities of two other, phylogenetically distinct lineages of fungivorous nematodes, Aphelenchidae and Diphtherophoridae, were unaffected by the local increase in fungal biomass. Apparently this plant species induces a local asymmetric boost of the fungal community, and only Aphelenchoididae were able to benefit from this invader‐induced change. The alternative explanation – the results are explained by a subtle, S. gigantea‐induced 0.1–0.2 units decrease of pH – seems unlikely, as pH optima for nematode taxa are relatively broad. Thus, apart from readily observable aboveground effects, the invasive plant species S. gigantea affects fungal biomass as well as a specific part of the fungivorous nematode community in a soil type‐independent manner.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.     

氮磷输入对过度放牧退化草原土壤线虫群落的影响

线虫是陆地生态系统中数量最多的一类无脊椎后生动物,在土壤碎屑食物网中占据多个营养级,并在能量流动和养分循环中发挥重要的生态功能.土壤线虫的群落结构和多样性水平可以反映生态系统功能的变化.多年来,过度放牧等不合理的开发利用导致我国大面积的草原处在不同程度的退化之中,外源性养分输入是维持其养分平衡进而恢复其生态和生产功能的重要措施.本研究以内蒙古锡林郭勒退化草原为对象,就氮磷输入对土壤线虫丰度和群落结构的单独效应和交互效应进行了研究.本研究获得线虫38属,其中垫咽属、滑刃属、索努斯属和盾垫属是该退化草原线虫群落优势属.氮输入降低了土壤线虫总丰度、捕食杂食线虫丰度以及植物寄生线虫丰度,而磷输入提高了土壤线虫总丰度、食真菌线虫丰度、捕食杂食类线虫丰度以及植物寄生线虫丰度,氮输入抑制了磷输入对线虫总丰度、捕食杂食线虫和植物寄生线虫丰度的促进作用.氮磷输入对线虫多样性无影响,可能与不同养分输入下稳定的植物群落多样性有关.氮输入显著提升了退化草原线虫成熟度指数,降低了植物寄生线虫成熟度指数(PPI),并且能缓解磷输入对PPI和瓦斯乐斯卡指数的负面作用,表明氮输入提高退化草地土壤健康程度,促进线虫群落和食物网的稳定性.上述研究结果有助于从土壤生物学角度理解氮磷输入对退化草地恢复的影响机理.     

Soil nematode abundances drive agroecosystem multifunctionality under short-term elevated CO2 and O3

The response of soil biotas to climate change has the potential to regulate multiple ecosystem functions. However, it is still challenging to accurately predict how multiple climate change factors will affect multiple ecosystem functions. Here, we assessed the short-term responses of agroecosystem multifunctionality to a factorial combination of elevated CO₂ (+200 ppm) and O₃ (+40 ppb) and identified the key soil biotas (i.e., bacteria, fungi, protists, and nematodes) concerning the changes in the multiple ecosystem functions for two rice varieties (Japonica, Nanjing 5055 vs. Wuyujing 3). We provided strong evidence that combined treatment rather than individual treatments of short-term elevated CO₂ and O₃ significantly increased the agroecosystem multifunctionality index by 32.3% in the Wuyujing 3 variety, but not in the Nanjing 5055 variety. Soil biotas exhibited an important role in regulating multifunctionality under short-term elevated CO₂ and O₃, with soil nematode abundances better explaining the changes in ecosystem multifunctionality than soil biota diversity. Furthermore, the higher trophic groups of nematodes, omnivores-predators served as the principal predictor of agroecosystem multifunctionality. These results provide unprecedented new evidence that short-term elevated CO₂ and O₃ can potentially affect agroecosystem multifunctionality through soil nematode abundances, especially omnivores-predators. Our study demonstrates that high trophic groups were specifically beneficial for regulating multiple ecosystem functions and highlights the importance of soil nematode communities for the maintenance of agroecosystem functions and health under climate change in the future.     

农田土壤线虫多样性研究现状及展望

目前土壤生物多样性已成为土壤生态学研究的热点问题之一。土壤生物以不同的方式改变着土壤的物理、化学和生物学特性。在农田生态系统中, 土壤动物是分解作用和养分矿化作用等生态过程的主要调节者。线虫作为土壤中数量最丰富的后生动物, 其生活史和取食类型多样, 在生态系统中发挥着重要作用。本文介绍了农田生态系统中影响线虫多样性的主要因素; 回顾了土壤线虫的物种多样性、营养类群多样性、生活史多样性和功能多样性的研究现状; 并提出了今后农田生态系统线虫多样性研究的重点。建议通过综合土壤线虫的生活史策略和营养类群等信息, 深入了解其生物多样性和土壤生态系统功能, 从而更好地发挥土壤线虫对农田生态系统变化的生物指示作用。     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

Role of Nematodes in Soil Health and Their Use as Indicators

The composition of nematode communities (plant-parasitic and free-living) may be used as bioindicators of soil health or condition because composition correlates well with nitrogen cycling and decomposition, two critical ecological processes in soil. Maturity and trophic diversity indices withstand statistical rigor better than do abundances, proportions, or ratios of trophic groups. Maturity indices respond to a variety of land-management practices, based largely on inferred life history characteristics of families. Similarity indices may be more useful than diversity indices because they reflect taxon composition. Improving existing indices or developing alternative indices refined by a greater understanding of the biology of key taxa may enhance the utility of nematodes as bioindicators.     

Trophic level and basal resource use of soil animals are hardly affected by local plant associations in abandoned arable land

Plants provide resources and shape the habitat of soil organisms thereby affecting the composition and functioning of soil communities. Effects of plants on soil communities are largely taxon‐dependent, but how different functional groups of herbaceous plants affect trophic niches of individual animal species in soil needs further investigation. Here, we studied the use of basal resources and trophic levels of dominating soil meso‐ and macrofauna using stable isotope ratios of carbon and nitrogen in arable fallow systems 3 and 14–16 years after abandonment. Animals were sampled from the rhizosphere of three plant species of different functional groups: a legume (Medicaco sativa), a nonlegume herb (Taraxacum officinale), and a grass (Bromus sterilis). We found virtually no consistent effects of plant identity on stable isotope composition of soil animals and on thirteen isotopic metrics that reflect general food‐web structure. However, in old fallows, the carbon isotope composition of some predatory macrofauna taxa had shifted closer to that of co‐occurring plants, which was particularly evident for Lasius, an aphid‐associated ant genus. Trophic levels and trophic‐chain lengths in food webs were similar across plant species and fallow ages. Overall, the results suggest that variations in local plant diversity of grassland communities may little affect the basal resources and the trophic level of prey consumed by individual species of meso‐ and macrofauna belowground. By contrast, successional changes in grassland communities are associated with shifts in the trophic niches of certain species, reflecting establishment of trophic interactions with time, which shapes the functioning and stability of soil food webs.     

Species‐specific ontogenetic diet shifts attenuate trophic cascades and lengthen food chains in exploited ecosystems

Ontogenetic diet shifts are pervasive in food websbut rules governing their emergence and the implications for trophic cascades are only partly understood. Recent theoretical advances in multispecies size spectrum models (MSSMs) predict that the emergence of ontogenetic diet shifts are driven primarily by size‐selective predation and changes in the relative abundances of suitably sized prey. Howeverthese assumptions have not yet been tested with data. Herewe developed alternative MSSMs based on different assumptions about the nature of species and size‐based preferences and tested them using an extensive dietary database for the Eastern Bering Sea (EBS). MSSMs with both size and species‐specific prey preferences correctly predicted approximately three‐fold more of the diet links than those that assumed fixed species preferences. Importantlythese model assumptions also had a profound effect on the strength of fishing‐induced trophic cascades and the emergent trophic structure of the community with and without fishing. The diet‐informed models exhibited lower predation mortality ratesparticularly for small individuals (less than 1 g) whichin turnreduced the intensity and reach of fishing‐induced trophic cascades up the size spectrum. If the level and size dependency of piscivory observed in EBS predators is typical of other systemsthe potential for fishing‐induced trophic cascades may be over‐stated in MSSMs as they are currently formulated and parameterized. Representation of species‐specific ontogenetic shifts in diet can strongly influence system responses to perturbationsand the extensions we propose should accelerate adoption of MSSMs as frameworks for exploring size‐based food web theory and developing modeling tools to support strategic management decisions.