Nematode Indicators of Organic Enrichment

The organisms of the soil food web, dependent on resources from plants or on amendment from other sources, respond characteristically to enrichment of their environment by organic matter. Primary consumers of the incoming substrate, including bacteria, fungi, plant-feeding nematodes, annelids, and some microarthropods, are entry-level indicators of enrichment. However, the quantification of abundance and biomass of this diverse group, as an indicator of resource status, requires a plethora of extraction and assessment techniques. Soluble organic compounds are absorbed by bacteria and fungi, while fungi also degrade more recalcitrant sources. These organisms are potential indicators of the nature of incoming substrate, but current methods of biomass determination do not reliably indicate their community composition. Guilds of nematodes that feed on bacteria (e.g., Rhabditidae, Panagrolaimidae) and fungi (e.g., Aphelenchidae, Aphelenchoididae) are responsive to changes in abundance of their food. Through direct herbivory, plant-feeding nematodes (e.g., many species of Tylenchina) also contribute to food web resources. Thus, analysis of the nematode community of a single sample provides indication of carbon flow through an important herbivore channel and through channels mediated by bacteria and fungi. Some nematode guilds are more responsive than others to resource enrichment. Generally, those bacterivores with short lifecycles and high reproductive potential (e.g., Rhabditidae) most closely mirror the bloom of bacteria or respond most rapidly to active plant growth. The feeding habits of some groups remain unclear. For example, nematodes of the Tylenchidae may constitute 30% or more of the individuals in a soil sample; further study is necessary to determine which resource channels they portray and the appropriate level of taxonomic resolution for this group. A graphic representation of the relative biomass of bacterivorous, fungivorous, and herbivorous nematodes provides a useful tool for assessing the importance of the bacterial, fungal, and plant resource channels in an extant food web.     

Interactions between a detrital resource pulse and a detritivore community

Detritivore communities influence the decomposition of detrital resources in virtually all natural systems. Conversely, detrital resources can also have considerable bottom-up effects on detritivore communities. While many investigations have examined detritivory and decomposition processes, few have considered interactions between detritivores and detritus as concurrent processes in the same system, or in the context of natural detrital pulses. In many systems, resource pulses contribute substantial detrital inputs to belowground systems. These detrital pulses may influence interactions between the detritivore community and detrital decomposition. I conducted field experiments to investigate interactions between detrital resource pulses of periodical cicada (Magicicada spp.) carcasses and scavenging detritivorous macroarthropods. Cicada litterfall pulses influenced several broad groups in the macroarthropod community, including relatively specialized necrophilous taxa and relatively generalized detritivores, omnivores and predators. Conversely, detritivore activity increased the rate of cicada carcass decomposition by 4,082% compared to caged control carcasses. These results suggest that interactions between pulses of cicada detritus and the detritivore community influence both the persistence of ephemeral detrital resources, and the distribution, abundance and behavior of detritivore populations.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Population- and ecosystem-level effects of predation on microbial-feeding nematodes

We studied the role of nematode predation in the functioning of detrital food webs assembled in microcosms. The microcosms contained defaunated humus and litter materials, a diverse microbial community with bacteria, fungi and protozoa, and a birch (Betula pendula) seedling infected with mycorrhizal fungi. Different levels of top-down control upon microbivorous nematodes were set up by assembling food webs either without predators, or in combinations with a specialist and a non-specialist predatory mite (Mesostigmata). The nematode community was composed of either (1) three species of bacterivorous, or (2) three species of fungivorous nematodes or (3) both groups together. After two growing periods for the birch (38 weeks), the microcosms were destructively sampled for animal and microbial biomasses, concentration of mineral N in the soil, plant biomass and plant N concentration. The specialist predator reduced biomasses of both bacterial- and fungal-feeding nematodes by more than 50%, whereas the non-specialist predator weakly increased the biomass of fungivorous nematodes. Thus, under high predation pressure, the biomass of microbivores changed as predicted by trophic dynamic models assuming strong top-down control and uniformly behaving trophic levels. Despite this, microbial biomass was unaffected by the predators. However, microbial respiration increased slightly in the presence of predators. Assuming that microbial respiration correlates with microbial productivity, the increase in microbial respiration indicates a cascading productivity regulation. The composition of the microbivore community had only a minor effect on the outcome of the top-down control on microbes. The >50% reduction in nematode biomass and respiration coincided with <16% increase in microbial respiration and did not affect microbial biomass. Presence of the specialist predator slightly reduced soil NH⁺ ₄ concentration in communities with fungivore nematodes but plant growth and N uptake remained unchanged. Thus, the structure of the community only weakly controlled nutrient mineralisation. Received: 18 May 1998 / Accepted: 3 May 1999     

转植酸酶基因玉米种植对土壤线虫群落的影响

土壤是生态系统中物质循环和能量转化的重要场所,转基因作物外源基因对土壤非靶标生物的影响已经引起人们的广泛关注.我国转植酸酶基因玉米(即自交系BVLA430101)于2009年9月27日获得了生物安全证书,该转植酸酶基因玉米可提高饲料利用效率,减少动物粪便造成的环境污染.本文以转植酸酶基因玉米(简称转基因玉米)和常规对照亲本玉米(简称对照玉米)为试验材料,通过大田试验研究了转基因玉米和对照玉米种植对土壤线虫群落组成与生态指标的影响.结果表明： 转基因玉米和对照玉米大田土壤分别分离出29个属和26个属的线虫.与对照玉米相比,转基因玉米种植田食细菌线虫相对多度与数量、捕/杂食线虫数量和土壤线虫总数,以及群落多样性指数等都有升高的趋势,而植食线虫相对多度与线虫总成熟度指数呈降低趋势.重复测量方差分析表明,整个生长季节内转基因玉米与对照玉米田间不同营养类群土壤线虫相对多度与数量及生态指标均无显著差异;而T检验分析表明,玉米乳熟期转基因玉米田食细菌线虫、捕/杂食线虫数量和土壤线虫总数显著高于对照玉米田,这可能与乳熟期转基因玉米田土壤总氮含量显著升高有关.     

典型黑土区农田土壤线虫群落的纬度分布格局及其驱动机制

东北黑土区是保障我国粮食安全的重要土壤资源, 了解该区域内农田土壤线虫的群落组成及其对环境驱动因子的响应机制, 对于研究黑土区农田土壤生态系统的生物多样性分布格局具有重要意义。2018年9月, 我们在42°50°‒49°08° N的典型黑土区采集了93个农田土壤样品, 利用形态学鉴定技术分析了土壤线虫群落的组成与结构。共鉴定出47个线虫属(相对丰度 > 1%), 其中食细菌线虫中的拟丽突属(Acrobeloides)是典型黑土区农田土壤中的优势属(相对丰度 > 5%)。土壤线虫总丰富度和总多度均随纬度增加而显著增加, 然而类似的变化趋势只出现在食细菌和杂食/捕食线虫中。土壤有机碳是影响土壤线虫丰富度和多度最为重要的环境因子, 其次是月平均温度。典型黑土区农田土壤线虫群落结构以47° N为分界线分为南部和北部两类, 主要归因于线虫群落中植物寄生和杂食/捕食线虫的相对多度在南、北特征属中存在差异。土壤pH值和容重分别是影响南部与北部黑土区线虫群落最重要的环境因子。本研究明确了典型黑土区农田土壤线虫群落的纬度分布格局及其与环境因素的关系, 可为揭示农业活动干扰下土壤生物对环境因子的响应机制提供基础数据和理论参考。     

Inferring trophic transfers from pulse-dynamics in detrital food webs

In semiarid ecosystems, decomposers are active during numerous short periods following rainfall events, and most inactive in the intervening dry periods. Many studies concern season-long dynamics of decomposer populations, but less is known of the short-term dynamics during wet periods. These short-term dynamics may provide the key to understanding interactions between microbes and fauna.The dynamics of populations in the detrital food web were followed after wetting large intact soil cores that had been removed from native shortgrass steppe, winter wheat, and fallow plots. The cores were sampled over a ten day period for bacteria, fungi, protozoa, and various functional groups of microarthropods and nematodes. The native sod had appreciably greater biomass of fungi, nematodes and microarthropods than did the cultivated plots, but there was no difference in bacteria or protozoans. The observed dynamics after wetting were different in two experiments which differed in temperature, soil water level, and the initial sizes of the populations. These results were interpreted in relation to a model of the structure of the detrital food web, and estimates were made of the rates of trophic transfers in the web. Consumption by protozoa was great enough for them to account for bacterial turnover, but consumption by fungivorous nematodes and microarthropods appeared to be too small to account for fungal turnover.Progress in understanding the dynamics of detrital food webs requires a better definition of the functional groups of soil organisms, their resources, predators and population parameters, and the effects of soil structure and water content on trophic relationships.     

The role of microarthropods and nematodes in decomposition in a semi-arid ecosystem

Summary We sampled the soil microarthropod community monthly in the oak-mesquite sand hill ecosystem. Small fungiphagous prostigmated mites (pyemotids, lordalychids and tarsonemids) that dominated the soil fauna in winter were replaced by large predaceous mites (rhodacarids and laelapids) in summer and autumn.We compared organic matter loss and microarthropod and nematode populations in shinnery oak (Quercus harvardii) using insecticide and untreated litter in fiberglass litterbags.Microarthropods extracted from litterbags showed a seasonal pattern similar to the soil cores except that collembolans and psocopterans were abundant in the litter and not in the soil cores. Numbers of free living nematodes were consistently greater than from untreated litter. The ratio of non-stylet to stylet bearing nematodes extracted from litter decreased from 4:1 in one month bags to 0.8:1.0 in the one year bags. Laboratory experiments showed that rhodacarid mites fed voraciously on nematodes.Untreated litter exhibited higher rates of organic matter loss than the insecticide treated litter; 20% and 35% respectively.We suggest that the abundant mesostigmatid mites prey on free living nematodes and that eliminating the predators allows the nematodes to overgraze the fungi and bacteria. The soil modifies the microclimate in buried litter allowing for higher biological activity, hence higher rates of decomposition.     

紫色土线虫对长期不同施肥措施的响应

土壤线虫在农田生态系统中数量丰富且对土壤环境变化敏感, 可用于评估不同田间管理条件下的土壤健康。本文探究了紫色土区长期不同施肥措施及土壤团聚体粒径对线虫群落的分布及其生态功能多样性的影响。试验设置了5个施肥处理: 不施肥(对照, CK)、单施化肥(NPK)、生物炭 + 化肥(BCNPK)、商品猪粪 + 化肥(OMNPK)和秸秆 + 化肥(RSDNPK)。团聚体粒径分为: 原状土(BS)、大团聚体(> 2 mm; LA)和小团聚体(0.25-2 mm; SA)。结果表明, 与对照相比, 施肥可促进线虫数量增长, 其中单施化肥处理下增幅最小(66%); 有机物料与化肥配施对线虫数量的提升更为显著, 秸秆 + 化肥处理下增幅达206%。不同施肥处理间线虫类群相对丰度差异显著, 大小均表现为: 食细菌线虫(BA) > 杂食/捕食线虫(OP) > 植食线虫(PP) > 食真菌线虫(FU)。小团聚体较其他土壤团聚体的杂食/捕食线虫丰度更低, 食细菌线虫丰度较高。秸秆与化肥配施处理下线虫群落结构指数和富集指数显著增加, 且各施肥处理下线虫功能足迹呈现明显差异。有机肥与化肥配施(尤其是秸秆 + 化肥)可提高土壤养分供应且有利于形成稳定健康的土壤生态系统, 助推区域农业的可持续发展。     

The Impacts of Above- and Belowground Plant Input on Soil Microbiota: Invasive <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Versus Native <Emphasis Type="Italic">Phragmites australis</Emphasis>

Invasive plants affect soil food webs through various resource inputs including shoot litter, root litter and living root input. The net impact of invasive plants on soil biota has been recognized; however, the relative contributions of different resource input pathways have not been quantified. Through a 2 × 2 × 2 factorial field experiment, a pair of invasive and native plant species (Spartina alterniflora vs. Phragmites australis) was compared to determine the relative impacts of their living roots or shoots and root litter on soil microbial and nematode communities. Living root identity affected bacteria-to-fungi PLFA ratios, abundance of total nematodes, plant-feeding nematodes and omnivorous nematodes. Specifically, the plant-feeding nematodes were 627% less abundant when living roots of invasive S. alterniflora were present than those of native P. australis. Likewise, shoot and root biomass (within soil at 0–10 cm depth) of S. alterniflora was, respectively, 300 and 100% greater than those of P. australis. These findings support the enemy release hypothesis of plant invasion. Root litter identity affected other components of soil microbiota (that is, bacterial-feeding nematodes), which were 34% more abundant in the presence of root litter of P. australis than S. alterniflora. Overall, more variation associated with nematode community structure and function was explained by differences in living roots than root or shoot litter for this pair of plant species sharing a common habitat but contrasting invasion degrees. We conclude that belowground resource input is an important mechanism used by invasive plants to affect ecosystem structure and function.     

Indirect effects of predatory trout on organic matter processing in detritus-based stream food webs

Indirect effects of predators on basal resources in allochthonous-based food webs are poorly understood. We investigated indirect effects of predatory brown trout ( Salmo trutta ) on detritus dynamics in southern beech ( Nothofagus spp.) forest streams in New Zealand through predation on the obligate detritivore, Zelandopsyche ingens (Trichoptera, Oeconesidae). Trout presence/absence and Z. ingens density were manipulated in flow-through tanks to investigate the lethal and sub-lethal effects of trout on litter processing by Z. ingens . An experiment that allowed trout access to Z. ingens showed trout predation reduced densities of Z. ingens resulting in slower breakdown of coarse particulate organic matter (CPOM) and reduced production of fine particulate organic matter (FPOM). A second experiment that prevented trout access to Z. ingens , but allowed the transmission of trout cues, resulted in no change in litter processing rates in the presence of trout. Litter processing rates were higher in high Z. ingens density treatments compared to low density treatments. Thus, trout effects on litter processing were due to reduced Z. ingens densities, not trout-induced modifications to Z. ingens feeding behaviour. Field assays of litter processing rates using artificial leaf packs in natural streams showed significant reductions in CPOM loss in trout streams compared to fishless streams. Z. ingens dominated biomass in fishless stream leaf packs, but a facultative shredder, Olinga feredayi , dominated trout stream leaf packs. Thus, the absence of Z. ingens drove differences in processing rates between trout and fishless streams and the indirect effects of trout on litter processing observed in mesocosms were evident in complex, natural food webs. Overall our study provides evidence that predators can influence resource dynamics in donor-controlled food webs through their effects on consumers.     

Top-down and bottom-up influences on populations of a stream detritivore

1. Knowledge of the influence of predatory fish in detritus‐based stream food webs is poor. We tested whether larval abundance of the New Zealand leaf‐shredding caddisfly, Zelandopsyche ingens (family Oeconesidae), was affected by the presence of predatory brown trout, Salmo trutta and the abundance of their primary detrital resource (Nothofagus leaves). 2. The density of Z. ingens and the biomass of leaves were determined in seven fishless streams and four trout streams in the Cass region, central South Island, on four occasions spanning 5 years. 3. Physicochemical conditions were similar in trout and fishless streams, but ancova indicated that Z. ingens numbers were positively related to leaf biomass and that caddisfly numbers were significantly greater in fishless streams than trout streams for any given biomass of leaf. The cases of trout stream larvae were also heavier per unit length than those in fishless streams. 4. Our results provide evidence for both top‐down and bottom‐up influences on a detritus‐based stream food web. Although stream detritivores may benefit from a habitat that provides both food and a degree of protection from predators, top‐down effects of predators on detritivore population abundance were still important. Thus, detrital resource availability may determine maximum attainable population size, whereas predation is likely to reduce the population to a level below that.     

Stoichiometric imbalances between detritus and detritivores are related to shifts in ecosystem functioning

How are resource consumption and growth rates of litter‐consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N:P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two‐species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N:P ratios of the consumers’ body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N‐demand arising from the production of N‐rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N‐demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed‐species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer–resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.     

Influence of an invasive ant on grazing and detrital communities and nutrient fluxes in a tropical forest

Aim  Pathways linking grazing and detrital subsystems of terrestrial ecosystems are important for ecosystem processes and function, but remain poorly understood. The invasion of a generalist predator creates a unique opportunity to study the effects of predation across these subsystems. We examine here, the effects of a non-native generalist predator, the little red fire ant ( Wasmannia auropunctata, Roger) on both grazing and detrital invertebrate communities and ecosystem processes in a rain forest understorey.
Location  Gamba Protected Area Complex, south-western Gabon, Africa.
Methods  We measured abundances and diversities of understorey grazing and detrital invertebrate communities, soil nutrients, herbivory, litter fragmentation rates and leaf chemistry of a dominant understorey shrub inside and outside of 19 separate invasion fronts. We then explored possible trophic cascades and pathways of interaction using path analysis.
Results  Results suggest that invasive ants may alter herbivory regimes, grazing and detrital communities, and may indirectly alter litter decomposition and nutrient cycling in the soil by suppressing important microbivore and detritivore populations with consequences for leaf chemistry.
Main conclusions  These results demonstrate that generalist predators may be major drivers of both grazing and detrital subsystems by inducing strong shifts in adjacent communities that ultimately affect ecosystem processes.     

Invasive plants have different effects on trophic structure of green and brown food webs in terrestrial ecosystems: a meta‐analysis

Although invasive plants are a major source of terrestrial ecosystem degradation worldwide, it remains unclear which trophic levels above the base of the food web are most vulnerable to plant invasions. We performed a meta‐analysis of 38 independent studies from 32 papers to examine how invasive plants alter major groupings of primary and secondary consumers in three globally distributed ecosystems: wetlands, woodlands and grasslands. Within each ecosystem we examined if green (grazing) food webs are more sensitive to plant invasions compared to brown (detrital) food webs. Invasive plants have strong negative effects on primary consumers (detritivores, bacterivores, fungivores, and/or herbivores) in woodlands and wetlands, which become less abundant in both green and brown food webs in woodlands and green webs in wetlands. Plant invasions increased abundances of secondary consumers (predators and/or parasitoids) only in woodland brown food webs and green webs in wetlands. Effects of invasive plants on grazing and detrital food webs clearly differed between ecosystems. Overall, invasive plants had the most pronounced effects on the trophic structure of wetlands and woodlands, but caused no detectable changes to grassland trophic structure.     

Plant diversity and identity effects on predatory nematodes and their prey

There is considerable evidence that both plant diversity and plant identity can influence the level of predation and predator abundance aboveground. However, how the level of predation in the soil and the abundance of predatory soil fauna are related to plant diversity and identity remains largely unknown. In a biodiversity field experiment, we examined the effects of plant diversity and identity on the infectivity of entomopathogenic nematodes (EPNs, Heterorhabditis and Steinernema spp.), which prey on soil arthropods, and abundance of carnivorous non‐EPNs, which are predators of other nematode groups. To obtain a comprehensive view of the potential prey/food availability, we also quantified the abundance of soil insects and nonpredatory nematodes and the root biomass in the experimental plots. We used structural equation modeling (SEM) to investigate possible pathways by which plant diversity and identity may affect EPN infectivity and the abundance of carnivorous non‐EPNs. Heterorhabditis spp. infectivity and the abundance of carnivorous non‐EPNs were not directly related to plant diversity or the proportion of legumes, grasses and forbs in the plant community. However, Steinernema spp. infectivity was higher in monocultures of Festuca rubra and Trifolium pratense than in monocultures of the other six plant species. SEM revealed that legumes positively affected Steinernema infectivity, whereas plant diversity indirectly affected the infectivity of Heterorhabditis EPNs via effects on the abundance of soil insects. The abundance of prey (soil insects and root‐feeding, bacterivorous, and fungivorous nematodes) increased with higher plant diversity. The abundance of prey nematodes was also positively affected by legumes. These plant community effects could not be explained by changes in root biomass. Our results show that plant diversity and identity effects on belowground biota (particularly soil nematode community) can differ between organisms that belong to the same feeding guild and that generalizations about plant diversity effects on soil organisms should be made with great caution.     

宁夏枸杞根际土壤线虫群落特征

宁夏枸杞具有很高的药用价值和营养价值,在宁夏大面积种植,带来了巨大的经济效益.研究不同条件下宁夏枸杞根际土壤线虫群落特征,对防止枸杞园土壤退化具有重要意义.以宁夏枸杞之乡——中宁为研究区,系统分析了季节、树龄和土层对枸杞根际土壤线虫数量和生态指数的影响.结果表明: 夏季宁夏枸杞根际土壤线虫数量最多,小杆属和拟丽突属为优势属;春季线虫数量最少,小杆属、针属和盘旋属为优势属;从春季到秋季,线虫稀有属种类和个体密度逐渐减少.随着树龄的增加,夏季根际土壤线虫总数逐渐减少;春季和秋季枸杞根际线虫总数先增加后减少,其中9年树龄根际土壤线虫总数最多.各树龄根际均为食细菌线虫所占比例最大,食真菌线虫和捕食-杂食类线虫比例很小,树龄<3年和>9年时植物寄生线虫比例相对较大.夏季枸杞根际20～40 cm比0～20 cm土壤线虫数量增加49.4%.随着树龄的增加,枸杞根际土壤线虫多样性指数、均匀度指数和丰富度指数均呈减小-增大-减小的趋势,而优势度指数先减小后增大.土壤pH值与土壤线虫优势度指数呈显著正相关;土壤有机质和速效磷与植物寄生线虫分别呈显著正相关和负相关;土壤碱解氮与线虫总数呈显著正相关;速效钾与线虫多个生态指数均呈显著负相关.整体上,随着树龄的增加,春季和秋季土壤线虫数量先增加后减少,夏季土壤线虫数量呈逐渐减少的趋势;线虫多样性下降,土壤逐渐退化.此外,在幼林和树龄>9年后需要抑制土壤植物寄生线虫的繁殖,以减少后期因线虫引起的病害.     

Top‐down and bottom‐up diversity cascades in detrital vs. living food webs

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrub Piper cenocladum (Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.     

No bottom-up effects of food addition on predators in a tropical forest

Several studies in temperate forests have demonstrated effects of litter addition on decomposers and predators. However, adding litter does not allow separating the effects of food availability and habitat space. We investigated the response of decomposers and predators to increased food resources and space in forests of the southern Mata Atlântica of Brazil. In two forest ecosystems representing an early successional stage of secondary forests and old-growth forest, we added nutrient-rich organic material, artificial litter of no nutritional value, or a combination of both to the soil surface of 120 plots to separate the effects of habitat space and food on soil food webs. We sampled litter- and soil-dwelling arthropods after three months using pitfall traps, soil sample extraction, and sticky traps just above the soil. Adding artificial litter had no positive effect on any of the 17 analyzed arthropod groups. Combining all sampled arthropods the effect was even significantly negative. Adding food had a positive effect on the abundance of decomposers, but not predators. We found no interactions between added artificial litter and added organic material. Our results suggest that the soil fauna in tropical forests is food limited. The lack of a bottom-up effect on predators suggests that they are not predominantly regulated by the abundance of epigeic prey but rather by competition or predation.     

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.     

Unpacking brown food‐webs: Animal trophic identity reflects rampant microbivory

Detritivory is the dominant trophic paradigm in most terrestrial, aquatic, and marine ecosystems, yet accurate measurement of consumer trophic position within detrital (=“brown”) food webs has remained unresolved. Measurement of detritivore trophic position is complicated by the fact that detritus is suffused with microbes, creating a detrital complex of living and nonliving biomass. Given that microbes and metazoans are trophic analogues of each other, animals feeding on detrital complexes are ingesting other detritivores (microbes), which should elevate metazoan trophic position and should be rampant within brown food webs. We tested these hypotheses using isotopic (¹⁵N) analyses of amino acids extracted from wild and laboratory‐cultured consumers. Vertebrate (fish) and invertebrate detritivores (beetles and moths) were reared on detritus, with and without microbial colonization. In the field, detritivorous animal specimens were collected and analyzed to compare trophic identities among laboratory‐reared and free‐roaming detritivores. When colonized by bacteria or fungi, the trophic positions of detrital complexes increased significantly over time. The magnitude of trophic inflation was mediated by the extent of microbial consumption of detrital substrates. When detrital complexes were fed to vertebrate and invertebrate animals, the consumers registered similar degrees of trophic inflation, albeit one trophic level higher than their diets. The wild‐collected detritivore fauna in our study exhibited significantly elevated trophic positions. Our findings suggest that the trophic positions of detrital complexes rise predictably as microbes convert nonliving organic matter into living microbial biomass. Animals consuming such detrital complexes exhibit similar trophic inflation, directly attributable to the assimilation of microbe‐derived amino acids. Our data demonstrate that detritivorous microbes elevate metazoan trophic position, suggesting that detritivory among animals is, functionally, omnivory. By quantifying the impacts of microbivory on the trophic positions of detritivorous animals and then tracking how these effects propagate “up” food chains, we reveal the degree to which microbes influence consumer groups within trophic hierarchies. The trophic inflation observed among our field‐collected fauna further suggests that microbial proteins represent an immense contribution to metazoan biomass. Collectively, these findings provide an empirical basis to interpret detritivore trophic identity, and further illuminate the magnitude of microbial contributions to food webs.