An insight into the feeding ecology of deep-sea canyon nematodes — Results from field observations and the first in-situC feeding experiment in the Nazaré Canyon

Submarine canyon systems provide a heterogeneous habitat for deep-sea benthos in terms of topography, hydrography, and the quality and quantity of organic matter present. Enhanced meiofauna densities as found in organically enriched canyon sediments suggest that nematodes, as the dominant metazoan meiobenthic taxon, may play an important role in the benthic food web of these sediments. Very little is known about the natural diets and trophic biology of deep-sea nematodes, but enrichment experiments can shed light on nematode feeding selectivity and trophic position. An in-situ pulse-chase experiment (Feedex) was performed in the Nazaré Canyon on the Portuguese margin in summer 2007 to study nematode feeding behaviour. ¹³C-labelled diatoms and bacteria were added to sediment cores which were then sampled over a 14-day period. There was differential uptake by the nematode community of the food sources provided, indicating selective feeding processes. ¹³C isotope results revealed that selective feeding was less pronounced at the surface, compared to the sediment subsurface. This was supported by a higher trophic diversity in surface sediments (Θ⁻¹ = 3.50 ± 0.2) compared to the subsurface (2.78 ± 0.6), implying that more food items may be used by the nematode community at the sediment surface. Predatory and scavenging nematodes contributed relatively more to biomass than other feeding types and can be seen as key contributors to the nematode food web at the canyon site. Non-selective deposit feeding nematodes were the dominant trophic group in terms of abundance and contributed substantially to total nematode biomass. The high levels of ‘fresh’ (bioavailable) organic matter input and moderate hydrodynamic disturbance of the canyon environment lead to a more complex trophic structure in canyon nematode communities than that found on the open continental slope, and favours predator/scavengers and non-selective deposit feeders.     

Deep-sea nematodes actively colonise sediments, irrespective of the presence of a pulse of organic matter: results from an in-situ experiment

A colonisation experiment was performed in situ at 2500 m water depth at the Arctic deep-sea long-term observatory HAUSGARTEN to determine the response of deep-sea nematodes to disturbed, newly available patches, enriched with organic matter. Cylindrical tubes,laterally covered with a 500 μm mesh, were filled with azoic deep-sea sediment and (13)C-labelled food sources (diatoms and bacteria). After 10 days of incubation the tubes were analysed for nematode response in terms of colonisation and uptake. Nematodes actively colonised the tubes, however with densities that only accounted for a maximum of 2.13% (51 ind.10 cm(-2)) of the ambient nematode assemblages. Densities did not differ according to the presence or absence of organic matter, nor according to the type of organic matter added. The fact that the organic matter did not function as an attractant to nematodes was confirmed by the absence of notable (13)C assimilation by the colonising nematodes. Overall, colonisation appears to be a process that yields reproducible abundance and diversity patterns, with certain taxa showing more efficiency. Together with the high variability between the colonising nematode assemblages, this lends experimental support to the existence of a spatio-temporal mosaic that emerges from highly localised, partially stochastic community dynamics.     

A New Approach for the Modelling of Sediment Reworking Induced by a Macrobenthic Community

A new model of bioturbation has been developed to describe short term sediment reworking induced by macrobenthic communities. The design of the model had to consider the mixing processes, firstly, at the organism level, and secondly, at community level. This paper describes the mixing mode of the four types of bioturbators defined by the authors: the biodiffusors, the upward-conveyors, the downward-conveyors and the regenerators. The mathematical formulation of these sub-models consists of ordinary differential equations. They take into account the size of the bioturbated zone, the output fluxes to the water column, tracer decay, physical mixing due to local currents and the type and intensity of the bioturbation processes. These sub-models make it possible to describe correctly the mixing events that have occurred in cores with each type of bioturbator. They also provide the basis for general bioturbation model, that will take into account the respective degrees of involvement of (i) the different bioturbation processes and their characteristics, (ii) the interference between the different processes, and (iii) make possible to predict the particle reworking in order to include it in studies of organic matter in early diagenesis. This revised version was published online in June 2006 with corrections to the Cover Date.     

Feeding ecology of shallow water meiofauna: insights from a stable isotope tracer experiment in Potter Cove, King George Island, Antarctica

Antarctic meiofauna is still strongly understudied, and so is its trophic position in the food web. Primary producers, such as phytoplankton, and bacteria may represent important food sources for shallow water metazoans, and the role of meiobenthos in the benthic-pelagic coupling represents an important brick for food web understanding. In a laboratory, feeding experiment ¹³C-labeled freeze-dried diatoms (Thalassiosira weissflogii) and bacteria were added to retrieved cores from Potter Cove (15-m depth, November 2007) in order to investigate the uptake of 3 main meiofauna taxa: nematodes, copepods and cumaceans. In the surface sediment layers, nematodes showed no real difference in uptake of both food sources. This outcome was supported by the natural δ¹³C values and the community genus composition. In the first centimeter layer, the dominant genus was Daptonema which is known to be opportunistic, feeding on both bacteria and diatoms. Copepods and cumaceans on the other hand appeared to feed more on diatoms than on bacteria. This may point at a better adaptation to input of primary production from the water column. On the other hand, the overall carbon uptake of the given food sources was quite low for all taxa, indicating that likely other food sources might be of relevance for these meiobenthic organisms. Further studies are needed in order to better quantify the carbon requirements of these organisms.     

Influence of decomposer food web structure and nitrogen availability on plant growth

We studied the sensitivity of soil microbial communities and ecosystem processes to variation in the vertical and horizontal structure of decomposer food web under nitrogen poor and N-enriched conditions. Microcosms with humus and litter layer of boreal forest floor, birch seedlings infected with mycorrhizal fungi, and decomposer food webs with differing trophic group and species composition of soil fauna were constructed. During the second growing period for the birch, we irrigated half of the microcosms with urea solution, and the other half with de-ionised water to create two levels of N concentration in the substrate. During the experiment night time respirations of the microcosms were measured, and the water leached through the microcosms was analysed for concentration of mineral N, and nematode numbers. The microcosms were destructively sampled after 37 weeks for plant biomass and N uptake, structure of soil animal and microbial community (indicated by PLFA profiles), and physical and chemical properties of the humus and litter materials. Predatory mites and nematodes had a negative influence on the biomass of their microbivorous and microbi-detritivorous prey, and microbi-detritivores affected the biomass and community structure of microbes (indicated by PLFA-analysis). Moreover, predatory mites and nematodes increased microbial biomass and changed the microbial community structure. The decomposer food web structure affected also N uptake and growth of plants. Microbi-detritivorous fauna had a positive effect, whereas predators of microbial and detritus feeding fauna exerted a negative influence on plant N uptake and biomass production. The impact of a trophic group on the microbes and plant was also strongly dependent on species composition within the group. Nitrogen addition magnified the influence of food web structure on microbial biomass and plant N uptake. We suggest that addition of urea-N to the soil modified the animal-microbe interaction by increasing microbial growth and altering community structure of microbes. The presence of microbi-detritivores and predators reduced loss of carbon from the microcosms, and the food web structure influenced also water holding capacity of the materials. The changes in plant growth, nutrient cycling, size of N and C pools, and in the physical properties of the soil emphasize the importance and diversity of indirect consequences of decomposer food web structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

Feeding of biofilm-dwelling nematodes examined using HPLC-analysis of gut pigment contents

The natural feeding behaviour of the nematodes Chromadorina bioculata (Schultze in Carus 1857) and Chromadorina viridis (Linstow 1876) was studied in situ, within epilithic biofilms of the Garonne River (France). Based on their feeding-type characteristics and population dynamics, it was hypothesised that these species feed selectively on microphytobenthos (MPB) within the biofilm, and that among MPB groups, diatoms are preferred. High-performance liquid chromatography (HPLC) was used for separation, identification and quantification of pigments both in nematode guts and in the biofilm. This is the first time that nematode gut pigment contents were examined under natural conditions. Diatoms dominated the MPB which also comprised cyanobacteria and green microalgae. The comparison between chlorophyll a content in nematode guts versus in the biofilm showed that C. bioculata and C. viridis fed opportunistically (non-selectively) on MPB within the biofilm. Only diatom biomarker pigments were found in nematode guts suggesting that they could preferentially fed on diatoms among MPB groups. However, the non-detection of biomarker pigments for other microphyte groups could be also linked to HPLC detection limits. It was estimated that Chromadorina nematodes daily ingested on average 0.03–0.67% of the MPB standing stock. This grazing covered only a small part of their energetic requirements, suggesting that besides MPB they probably also fed on other biofilm food sources. Some considerations on the applicability of the HPLC gut pigment analysis technique for the examination of nematode feeding are also presented.     

Do Meio- and Macrobenthic Nematodes Differ in Community Composition and Body Weight Trends with Depth?

Nematodes occur regularly in macrobenthic samples but are rarely identified from them and are thus considered exclusively a part of the meiobenthos. Our study compares the generic composition of nematode communities and their individual body weight trends with water depth in macrobenthic (>250/300 μm) samples from the deep Arctic (Canada Basin), Gulf of Mexico (GOM) and the Bermuda slope with meiobenthic samples (<45 μm) from GOM. The dry weight per individual (μg) of all macrobenthic nematodes combined showed an increasing trend with increasing water depth, while the dry weight per individual of the meiobenthic GOM nematodes showed a trend to decrease with increasing depth. Multivariate analyses showed that the macrobenthic nematode community in the GOM was more similar to the macrobenthic nematodes of the Canada Basin than to the GOM meiobenthic nematodes. In particular, the genera Enoploides, Crenopharynx, Micoletzkyia, Phanodermella were dominant in the macrobenthos and accounted for most of the difference. Relative abundance of non-selective deposit feeders (1B) significantly decreased with depth in macrobenthos but remained dominant in the meiobenthic community. The occurrence of a distinct assemblage of bigger nematodes of high dry weight per individual in the macrobenthos suggests the need to include nematodes in macrobenthic studies.     

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     

Effect of temperature on growth and nitrogen mineralization of fungi and fungal-feeding nematodes

Dish and column microcosms containing alfalfa-sand medium were established to determine effect of temperature on growth and N mineralization ability of two fungi (Rhizoctonia solaniand Botrytis cinerea) and two nematodes (Aphelenchus avenaeand Aphelenchoides composticola). The microcosms were incubated at 15, 20, 25 and 29 °C for 21 days. In the dish microcosms, hyphal growth rates of both fungal species increased with temperature in the range of 15–25 °C. Above that temperature range, the growth rate of R. solani remained almost constant while that of B. cinereadecrease considerably. The population growth rate of A. avenae increased with temperature between 15 and 29 °C on colonies of R. solani and B. cinerea in dish microcosms. The growth rate of A. composticola also increased in the range of 15–25 °C but decreased greatly beyond that temperature range independent of the fungal species as food source. Inorganic N (NH₄ ⁺ + NO₃ ^–) was collected from each column microcosm by leaching every 3 days. In the columns containing R. solani, there was a significant effect of temperature on the amount of N detected in the fungus+A avenaeor A. composticolabut not in the fungus alone columns. The total amount of N was greatest at 29 °C for A. avenaeand at 20 °C for A. composticola columns, concurrent with the population growth rates of the nematodes. In the columns containing B. cinerea, the effect of temperature on the amount of inorganic N was not significant in either the fungus alone or fungus+nematode columns, although the population growth rates of the both nematode species were highest at 20 °C. For B. cinerea, the N amount across temperatures was the same or larger for the fungus alone as for the fungus+nematode columns. In general, the contribution of fungal-feeding nematodes to N mineralization was small in any combinations of fungus and nematode species at any temperature. Similarity in C/N ratio of the fungal and nematode biomass, organic substrate C/N ratios too low for measurable increase in net mineralization by the nematodes and small reproduction of the nematodes in the column microcosms were probable contributory factors.     

Cryptophytes: An emerging algal group in the rapidly changing Antarctic Peninsula marine environments

The western Antarctic Peninsula (WAP) is a climatically sensitive region where foundational changes at the basis of the food web have been recorded; cryptophytes are gradually outgrowing diatoms together with a decreased size spectrum of the phytoplankton community. Based on a 11-year (2008–2018) in-situ dataset, we demonstrate a strong coupling between biomass accumulation of cryptophytes, summer upper ocean stability, and the mixed layer depth. Our results shed light on the environmental conditions favoring the cryptophyte success in coastal regions of the WAP, especially during situations of shallower mixed layers associated with lower diatom biomass, which evidences a clear competition or niche segregation between diatoms and cryptophytes. We also unravel the cryptophyte photo-physiological niche by exploring its capacity to thrive under high light stress normally found in confined stratified upper layers. Such conditions are becoming more frequent in the Antarctic coastal waters and will likely have significant future implications at various levels of the marine food web. The competitive advantage of cryptophytes in environments with significant light level fluctuations was supported by laboratory experiments that revealed a high flexibility of cryptophytes to grow in different light conditions driven by a fast photo-regulating response. All tested physiological parameters support the hypothesis that cryptophytes are highly flexible regarding their growing light conditions and extremely efficient in rapidly photo-regulating changes to environmental light levels. This plasticity would give them a competitive advantage in exploiting an ecological niche where light levels fluctuate quickly. These findings provide new insights on niche separation between diatoms and cryptophytes, which is vital for a thorough understanding of the WAP marine ecosystem.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

A comparison of the gut contents of two co-existing lake-dwelling Naididae (Oligochaeta), Nais pseudobtusa and Chaetogaster diastrophus

SUMMARY. The gut contents of Nais pseudobtusa and Chaetogaster diastrophus , two co-existing littoral-dwelling Naididae (Oligochaeta), differed over a period of one and a half years due to strong selection by C. diastrophus for specific diatom genera, Cymbella spp. and Navicula spp. In contrast, N. pseudobtusa showed less selection and the gut contents consisted mainly of detritus, diatoms and mineral particles. These differences in the food niche are likely to contribute to the co-existence of the two species in this habitat.
Seasonal changes in the relative abundance of diatom genera in the habitat greatly influenced the gut contents of N. pseudobtusa ; however, these changes had less influence on those of C. diastrophus . This and the highly selective nature of the gut contents of C. diastrophus suggests that the two species have different feeding strategies. It is suggested that C. diastrophus actively seeks out certain diatom genera in preference to other food items, while the unseslective gut contents of N. pseudobtusa are more typical of an opportunist feeder.     

Microbial Food Webs in Marine Sediments. I. Trophic Interactions and Grazing Rates in Two Tidal Flat Communities

Abstract The role of grazing by marine sediment flagellates, ciliates, and meiobenthic animals in controlling production of their bacterial and diatom prey was investigated. Several novel or modified techniques were used to enumerate prey (bacteria and diatoms), measure bacterial production, quantify proto- and micrometazoan predators, and evaluate rates of bacterivory and herbivory. The results indicated that, in a temperate, marine intertidal flat composed of fine sand, colorless nanoflagellates, ciliates, and nematodes were the most important bacterivores. Together, these organisms were responsible for removing up to 53% of bacterial production, by grazing. The observed rates of bacterivory were high enough to hypothesize that periods of grazing control of bacterial production might occur regularly in similar habitats. Colorless microflagellates, ciliates, and nematodes had high rates of diatom consumption. The combined small diatom consumption rate was equivalent to 132% of diatom standing stock per day. Trophic interactions between diatoms and micro- and meiobenthos might be a factor limiting growth of small (around 10 μm) diatoms. In coarse sands of an open beach, all micrograzers except pigmented nanoflagellates were rare, whereas bacterial and diatom assemblages were rather abundant and active. In this type of sediment, the micrograzers were able to consume only a marginal percentage of bacterial production (<1%) and diatom standing stock (3.8%), thus playing a minor role in controlling the dynamics of their prey. Received: 11 June 1996; Accepted: 13 August 1996     

Does bioturbation by a benthic fish modify the effects of sediment contamination on saltmarsh benthic microalgae and meiofauna?

A microcosm experiment was conducted using a replicated factorial design to determine if a benthic fish modifies the effects of sediment-bound contaminants (diesel fuel and two levels of a Cu, Cr, Pb, Hg and Cd mixture) on saltmarsh benthic primary producers and consumers. The naked goby, Gobiosoma bosc, a burrowing fish that preys on small macrofauna, was added to experimental microcosms. Goby burrowing and foraging significantly increased turbidity and disrupted the sediment surface. Results were typified by complex and varied responses with many statistically significant effects and interactions among treatments. Although G. bosc modified the responses to both diesel and metal pollution in invertebrates (but not microalgae), bioturbation did not increase or decrease the toxic effects of metals or diesel, and diesel-metal interactions did not vary in response to G. bosc. Specifically, G. bosc inhibited a trend toward diesel-induced increases in nematode abundance, and diesel toxicity inhibited increases in ostracod abundance stimulated by G. bosc. Diatoms, nematodes and the copepod Pseudostenhelia wellsi decreased in treatments with G. bosc. However, G. bosc lead to increases in cyanobacteria and ostracods and a trend toward increases in the copepod Pseudobradya sp. Our findings suggest that microcosm experiments are potentially poor mimics of natural systems without bioturbation. Conclusions about the direct and indirect effects of contaminants may differ with and without bioturbation. Finally, our work suggests that the direct effects of toxicants may inhibit or mask bioturbation effects that stimulate population growth of some meiofauna.     

Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

黄土高原半干旱区不同苜蓿-作物种植方式下土壤线虫群落组成及代谢足迹

线虫群落在土壤生态系统物质循环和能量流动过程中起着重要的作用。本研究以宁夏南部山区3种苜蓿-作物种植方式[苜蓿连作(A-A)、苜蓿-玉米轮作(A-C)、苜蓿-马铃薯轮作(A-P)]为对象,探讨不同种植方式下土壤理化性质、线虫群落组成结构及代谢足迹特征,评价黄土高原半干旱区苜蓿-作物种植方式对土壤食物网结构和功能的影响。结果表明: 1)与苜蓿连作相比,苜蓿-玉米和苜蓿-马铃薯轮作方式下土壤有机碳含量分别增加了4.6%、7.4%,全氮含量分别增加了4.0%、5.2%,土壤微生物生物量碳和氮也有显著提高;2)在苜蓿连作方式下,线虫总多度为211 条·100 g^-1干土,植食线虫为优势营养类群(35.7%),而在苜蓿-玉米和苜蓿-马铃薯轮作方式下,土壤线虫总多度较苜蓿连作有所增加(分别增加49.5%、93.7%),捕-杂食线虫成为优势营养类群(所占比例分别为45.7%、37.6%);3)相较于苜蓿连作,苜蓿-作物轮作方式下,植物寄生线虫指数(PPI)显著降低,表明植物寄生线虫在土壤食物网中的危害减轻;而线虫通路指数(NCR)有所增加,表明苜蓿-作物轮作方式下,土壤有机质分解过程中细菌分解作用进一步增强;4)苜蓿-作物轮作方式下,土壤线虫成熟度指数(MI)及其复合足迹、富集足迹、结构足迹均显著提高,土壤线虫的生产力和代谢活性显著增强,线虫群落的结构和功能更为成熟稳定。研究表明,相比苜蓿连作,苜蓿-作物轮作改善了土壤养分状况,使得土壤食物网的资源输入和能量利用效率均有所增加,受干扰程度显著降低,土壤生态系统更为稳定健康,因而有利于农业的可持续发展。     

Is there competition among ciliates and nematodes?

1. Biotic interaction between the ciliate Cyclidium glaucoma and the nematode Caenorhabditis elegans was investigated by manipulating the densities of the organisms in microcosms with and without sediment. 2. After 11 days the abundance of ciliates, nematodes and bacteria as well as extracellular enzyme activity were determined. Ciliates had a negative effect on nematode abundance in microcosms without sediment and in microcosms with sandy sediment, whereas in muddy sediment the effect was less distinctive. An effect of nematodes on ciliates was not observed. 3. The common resource bacteria were not affected negatively by the activity of the grazers. Overall grazer biomass increased with the addition of sediment to the microcosms, suggesting a rise of the carrying capacity in the experimental system. Especially in muddy sediment the abundance of bacteria and extracellular enzyme activity was higher compared to the microcosms without sediment. 4. The results of the experiment suggest a strong interspecific competition between nematodes and ciliates, where nematodes are, at least temporary, strongly affected.     

Influence of bacterivorous nematodes on the decomposition of cordgrass

The influence of bacterivorous nematodes (Diplolaimelloides meyli, Diplolaimelloides oschei, Diplolaimella dievengatensis, Panagrolaimus paetzoldi) on the decomposition of a macrophyte (Spartina anglica) in an aquatic environment was investigated by using laboratory microcosm experiments. Several earlier studies have shown enhancement of the decomposition process in the presence of nematodes, but nematode species-specific effects were never tested. In this study four bacterivorous nematode species were applied separately to microcosms to investigate such species-specific influences.No stimulation of the decomposition process nor of the microbial community was observed in the presence of the nematodes, both were highest in the absence of nematodes. However, clear differences were found between nematode treatments. P. paetzoldi reached much higher numbers than the other species, causing a decrease in microbial activity, probably due to (over)grazing. Remarkably this low microbial activity did not result in a slow-down of the decomposition process compared to the other nematode treatments, raising the question whether P. paetzoldi might be able to directly assimilate detrital compounds. Other nematode species reached much lower densities, but nevertheless an influence on the decomposition process was observed. However, this experiment does not support the view that bacterivorous nematodes enhance decomposition rate.The experimental results show that in nematode communities the use of functional groups is inadequate for biodiversity studies. The four nematode species used in this study belong to the same functional group, but are clearly not functionally redundant since they all have a different influence on the cordgrass decomposition. This suggests that the relationship between nematode species diversity and ecosystem functioning may be idiosyncratic.     

Bioturbating space enhances the effects of non‐additive interactions among benthic ecosystem engineers on cross‐habitat nutrient regeneration

Understanding the mechanisms that drive complementary interactions among species is key to the progress of the debate over the role of biodiversity in ecosystem functioning. In addition, interspecific interactions among physical ecosystem engineers have rarely been framed in the context of biodiversity experiments. Here, we provide an empirical test of how the physical niche space of species influences the effects of the biodiversity of bioturbators on cross‐habitat nutrient fluxes in benthic sediments. In the laboratory, we orthogonally manipulated the number and composition of three benthic invertebrate bioturbator species that differ in the dimensions of their bioturbating space niche; i.e. their vertical distribution in the sediment over a gradient of sediment depth and volume. The ammonium (NH₄‐N) flux from the sediment to the water was positively related to bioturbator species richness only in the sediments with the deepest depth and greatest volume. The non‐additive effects of bioturbator species richness on the benthic–pelagic NH₄‐N flux increased linearly with sediment depth and volume, but only in the three‐species mixtures. Furthermore, no individual species dominated the rates of H₄‐N fluxes, indicating that biodiversity effects were mainly driven by complementarity. These results suggest that sediment bioturbating space mediates the magnitude of non‐additive effects among the three invertebrate species and sheds light on the importance of physical niche space in modulating the positive effects of biodiversity on ecosystem functioning.     

Soil community composition drives aboveground plant–herbivore–parasitoid interactions

Soil organisms can influence higher trophic level aboveground organisms, but only very few studies have considered such effects. We manipulated soil community composition of model grassland ecosystems by introducing nematode communities, microorganisms, neither or both groups. Above ground, aphids ( Rhopalosiphum padi ) and parasitoids ( Aphidius colemani ) were introduced, and we measured individual performance and population dynamics of plants, aphids and parasitoids. In microcosms with nematode inoculations either with or without microorganism inoculation, aphids offspring production was significantly reduced by 31%. Aphid populations on both host plants Agrostis capillaris and Anthoxanthum odoratum were lowest in microcosms with combined nematode and microorganism inoculations. Opposite results were found for parasitoids. While the number of emerged parasitoids did not differ between treatments, parasitoid mortality and the proportion of males were significantly lower in microcosms with nematode and microorganism inoculations. Parasitized aphids were significantly larger in microcosms with nematodes inoculated. Plant biomass did not differ, but in the preferred host plant A. odoratum , foliar phenolic content was reduced in the presence of nematodes, and also the concentration of amino acids in the phloem. This study shows that the composition of the soil community matters for aboveground multitrophic interactions.