Linking aboveground and belowground communities: the indirect influence of aphid species identity and diversity on a three trophic level soil food web

There has been a growing recent interest in how foliar herbivory may indirectly affect the belowground sub-system, but little is known about the belowground consequences of the identity, species composition or diversity of foliar herbivores. We performed an experiment, utilising model grassland communities containing three plant species, in which treatments consisted of addition of each of eight aphid species in single and in two- four- and eight-species combinations, as well as an aphid-free treatment. While aphid species treatments did not affect total plant biomass or productivity, aphid species identity had important effects on the relative abundance of the three plant species. This in turn affected the abundances of each of three groups of secondary consumers in the soil food web (bacterial- and fungal-feeding nematodes, and enchytraeids) but not primary consumers (microbes, herbivorous nematodes) or tertiary consumers (predatory nematodes). The fact that some trophic levels responded to treatments while others did not is consistent with trophic dynamic theory. Aphid species treatments also affected the community composition within each of the herbivorous, microbe-feeding and top predatory nematode groups, as well as diversity within the first two of these groups. However, aphid species diversity per se had few effects. There were specific instances in which specific aboveground and belowground response variables in two aphid species combinations differed significantly from those in both of the corresponding single aphid species treatments (apparently as a consequence of resource use complementarity between coexisting aphid species), but no instance in which increasing aphid diversity beyond two species had any effect. Our results provide evidence that the identity of aboveground consumers can have effects that propagate through multiple trophic levels in soil food webs in terms of consumer abundance, and composition and diversity within trophic levels.     

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.     

Co-flowering neighbors influence the diversity and identity of pollinator groups visiting plant species

The generalization–specialization continuum exhibited in pollination interactions currently receives much attention. It is well-known that the pollinator assemblage of particular species varies temporally and spatially, and therefore the ecological generalization on pollinators may be a contextual attribute. However, the factors causing such variation and its ecological and evolutionary consequences are still poorly understood. This variation can be caused by spatial or temporal variation in the pollinator community, but also by variation in the plant community. Here, we examined how the floral neighbourhood influenced the generalization on pollinators and the composition of pollinators of six plant species differing in generalization levels and main pollinators. The diversity, identity and density of floral species affected both the level of generalization on pollinators and the composition of visitors of particular plant species. Although the relationships to floral neighbourhood varied considerably among species, generalization level and visitation by uncommon pollinators generally increased with floral diversity and richness. The generalization level of the neighbourhood was negatively related to the generalization level of the focal species in two species. The number of flowers of the pollinator-sharing species and the number of flowers of the focal species had different effects on the composition of visits in different species; attributable to differences in facilitation/competition for pollinator attraction. We propose that an important ecological implication of our results is that variation in species interactions caused by the pollination context may result in increased community stability. The main evolutionary implication of our results is that selection on flower and pollinator traits may depend, to an unknown extent, on the composition of the co-flowering plant community.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.     

The influence of a parasitoid's response to temperature on the performance of a tri‐trophic food web

1. Food web interactions are complex and can respond to environmental changes in unpredictable ways that do not necessarily equate to the individual responses of each of the components of the food web. 2. Biomass can be used to evaluate the productivity of the three individual trophic levels, in the form of the Net Generational Productivity (NGP) and the performance of the entire food web with the newly developed tri‐trophic food web performance ratio (?_3t). 3. These parameters were used to evaluate the performance of nine plant‐based tri‐trophic food webs composed of: potato, Solanum tuberosum L. and two cultivars of bell pepper Capsicum annuum L; three biotypes of the aphid Macrosiphum euphorbiae Thomas (Hemiptera: Aphididae) and the parasitoid wasp Aphidius ervi (Haliday) (Hymenoptera: Braconidae). 4. The NGP showed that the thermal window for biomass productivity for each trophic level is different and is reduced by approximately 4 °C with respect to the inferior level. Aphidius ervi had the smallest thermal window for biomass productivity and development. 5. The present results showed that the performance (?_3t) of the tested food webs is influenced in a top‐down fashion, where the intra‐specific variation of the food web, namely the host plant, played a major role in the productivity of each of the subsequent trophic levels. 6. The ?_3t suggested that exposure to high and low temperatures might severely affect the effectiveness of A. ervi as a biocontrol agent of the aphid M. euphorbiae in bell pepper and potato crops.     

内蒙草原不同植物功能群及物种对土壤微生物组成的影响

为了分析不同植物群落组成对内蒙古典型草原土壤微生物群落组成的影响,本研究利用植物功能群剔除处理实验平台,采用荧光定量PCR(real-timePCR)和自动核糖体间隔区基因分析(automated ribosomal intergenic spacer analysis,ARISA)技术,对不同植物功能群组成的非根际土壤和常见物种的根际土壤中细菌和真菌的数量及群落结构进行了分析。结果表明,在非根际土壤中,不同植物功能群组成对细菌数量有显著影响,而对真菌数量及细菌和真菌的群落结构影响不明显;在根际土壤中,不同植物物种对细菌、真菌的数量都有显著影响。此外,聚类分析表明,不同物种的根际土中细菌和真菌的群落结构也有所不同,尤其以细菌的群落结构变化较为明显。研究结果表明不同植物物种可以通过根系影响土壤微生物群落组成。     

水盐变化对荒漠植物功能多样性与物种多样性关系的影响

研究植物功能多样性与物种多样性关系对土壤水盐环境的响应规律,有助于掌握胁迫环境下植物多样性的维持机理.本研究分析了艾比湖高、低土壤水盐环境下荒漠植物功能多样性、物种多样性差异及二者间关系在不同水盐环境下的变化规律.结果表明:(1)高水盐环境下植物功能离散度显著低于低水盐环境(P<0.05),物种多样性指数Fish-α显著高于低水盐环境;(2)一元与多元回归分析均表明,高水盐环境下物种多样性对功能多样性RaoQ与FRic指数有显著促进作用,低水盐环境下物种多样性对功能多样性和均匀度有显著促进作用,但对功能离散度具有显著抑制作用(P<0.05);(3)物种多样性对功能多样性的影响在低水盐环境下更显著,预测力更强;(4)功能丰富度、离散度与均匀度能较敏感地反映出荒漠植物功能多样性对水盐变化的响应.本研究有助于理解荒漠植物群落构建机制,并为荒漠植物恢复与管理提供理论参考.     

Differential responses of soil bacteria,fungi, archaea and protists to plant species richness and plant functional group identity

Plants are known to influence belowground microbial community structure along their roots, but the impacts of plant species richness and plant functional group (FG) identity on microbial communities in the bulk soil are still not well understood. Here, we used 454‐pyrosequencing to analyse the soil microbial community composition in a long‐term biodiversity experiment at Jena, Germany. We examined responses of bacteria, fungi, archaea, and protists to plant species richness (communities varying from 1 to 60 sown species) and plant FG identity (grasses, legumes, small herbs, tall herbs) in bulk soil. We hypothesized that plant species richness and FG identity would alter microbial community composition and have a positive impact on microbial species richness. Plant species richness had a marginal positive effect on the richness of fungi, but we observed no such effect on bacteria, archaea and protists. Plant species richness also did not have a large impact on microbial community composition. Rather, abiotic soil properties partially explained the community composition of bacteria, fungi, arbuscular mycorrhizal fungi (AMF), archaea and protists. Plant FG richness did not impact microbial community composition; however, plant FG identity was more effective. Bacterial richness was highest in legume plots and lowest in small herb plots, and AMF and archaeal community composition in legume plant communities was distinct from that in communities composed of other plant FGs. We conclude that soil microbial community composition in bulk soil is influenced more by changes in plant FG composition and abiotic soil properties, than by changes in plant species richness per se.     

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.     

Plant diversity surpasses plant functional groups and plant productivity as driver of soil biota in the long term

Background

One of the most significant consequences of contemporary global change is the rapid decline of biodiversity in many ecosystems. Knowledge of the consequences of biodiversity loss in terrestrial ecosystems is largely restricted to single ecosystem functions. Impacts of key plant functional groups on soil biota are considered to be more important than those of plant diversity; however, current knowledge mainly relies on short-term experiments.

Methodology/Principal Findings

We studied changes in the impacts of plant diversity and presence of key functional groups on soil biota by investigating the performance of soil microorganisms and soil fauna two, four and six years after the establishment of model grasslands. The results indicate that temporal changes of plant community effects depend on the trophic affiliation of soil animals: plant diversity effects on decomposers only occurred after six years, changed little in herbivores, but occurred in predators after two years. The results suggest that plant diversity, in terms of species and functional group richness, is the most important plant community property affecting soil biota, exceeding the relevance of plant above- and belowground productivity and the presence of key plant functional groups, i.e. grasses and legumes, with the relevance of the latter decreasing in time.

Conclusions/Significance

Plant diversity effects on biota are not only due to the presence of key plant functional groups or plant productivity highlighting the importance of diverse and high-quality plant derived resources, and supporting the validity of the singular hypothesis for soil biota. Our results demonstrate that in the long term plant diversity essentially drives the performance of soil biota questioning the paradigm that belowground communities are not affected by plant diversity and reinforcing the importance of biodiversity for ecosystem functioning.     

Sensitivity of ecosystem functioning to changes in trophic structure, functional group composition and species diversity in belowground food webs

The objective of the present paper, using decomposer food webs as a tool, is to explore the levels of the ecological hierarchy (trophic groups, feeding guilds, species populations) at which reduction in complexity brings about significant changes in ecosystem performance. A review is given of various mini-ecosystem studies that have recently been conducted at the University of Jyväskylä. It is hypothesized that the typical features of soils as a habitat, and the peculiarities of belowground food webs, such as the commonness of indirect interactions (mediated through abiotic resources) among the biota, together with the high frequency of polyphagy/omnivory among soil organisms, produce a diversity-ecosystem functioning relationship that is likely to differ from those of aquatic and aboveground food webs. Experiments showed that alterations in trophic levels were reflected in significant changes in decomposition processes, which, in turn, had substantial impact on primary production. Similarly, heterogeneity within trophic levels was shown to be associated with increased growth of birch and pine seedlings. In contrast, species diversity within a feeding guild had little or no influence on ecosystem-level processes. However, the species-specific properties of individual taxa were shown to be more influential in affecting plant growth than species number per se. For example, the presence of an omnivorous enchytraeid species in the mini-ecosystems was observed to consistently be associated with high biomass production of tree seedlings. It is concluded that the so-called trophic dynamic models based on direct feeding interactions are of limited value in predicting the outcomes of interactions taking place belowground.     

Allochtonous input and trophic level heterogeneity: impact on an aquatic food web

The impact of food web complexity in open and closed food webs was explored experimentally under controlled laboratory conditions. We used an aquatic model community consisting of two forms of primary producers, phytoplankton ( Scenedesmus obtusiuscusculus ) and periphyton ( Nitzschia perminuta ) and two types of consumers, Daphnia pulex feeding on phytoplankton and Chydorous sphaericus feeding on both periphyton and phytoplankton. Three different food webs all having the phytoplankton and periphyton, but having either one of the consumers or both were set up. These food webs were studied in an open and closed treatment. In the open treatment, phytoplankton was continuously flowing through the aquaria whereas in the closed system all the phytoplankton was delivered at the start of the experiment.
D. pulex had a positive effect on the density of C. sphaericus in both the open and closed treatments. In the open treatment C. sphaericus increased to very high numbers and was able to depress phytoplankton and thereby negatively affect D. pulex . Our study shows that the explicit handling of the population dynamics of both grazers allowed us to show how a compensatory increase in one primary producer due to increased grazing on another primary producer creates a negative feedback between consumers: C. sphaericus increased and negatively affected D. pulex .     

Cascading predator control interacts with productivity to determine the trophic level of biomass accumulation in a benthic food web

Large-scale exploitation of higher trophic levels by humans, together with global-scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top–down and bottom–up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short-term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer-controlled early life-stages of algae, indicating that both the number of trophic levels and the life-stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food-web structure.     

The response of three chydorid species to temperature, pH and food

The responses of three chydorid species, Chydorus sphaericus(O.F. Müller), Alona affinis (Leydig) and Alonopsis elongata (Sars) to temperature, pH and food type were examined. Egg development time of all species decreased with increasing temperature, although the degree of change was different for each species. C. sphaericus had the fastest development time at all temperatures, and A. elongata the slowest. pH also affected the egg development time of each species differently. A. elongata failed to reproduce at low and medium pH, the egg development time of C. sphaericus was fastest at high pH while that of A. affinis was fastest at low pH. Food type was found to have significant effects on the population growth of individual species. C. sphaericus populations grew equally well in all three food types provided, and grew more than the other two species when fed on an algae culture, and filtered pond water. A. affinisand A. elongata populations grew best when fed on a detritus food source. The responses of each species to the different variables tested are discussed in relation to field observations of their distribution and abundance.     

The differential impact of a native and a non‐native ragwort species (Senecioneae) on the first and second trophic level of the rhizosphere food web

Whereas the impact of exotic plant species on above‐ground biota is relatively well‐documented, far less is known about the effects of non‐indigenous plants on the first and second trophic level of the rhizosphere food web. Here, rhizosphere communities of the invasive narrow‐leaved ragwort Senecio inaequidens and the native tansy ragwort Jacobaea vulgaris, co‐occurring in three semi‐natural habitats are compared. For both species, two life stages were taken into consideration. Quantitative PCR assays for the analyses of bacterial and fungal communities at a high taxonomic level were optimized, and it was investigated whether changes in the primary decomposer community were translated in alterations in bacterivorous and fungivorous nematode communities. In contrast to J. vulgaris, small but significant reductions were observed for Actinobacteria and Bacteroidetes (both p < 0.05) in case of the invasive S. inaequidens. More pronounced changes were detected for the overall nematode community density, and, more specifically, for the bacterivorous genus Anaplectus and the family Monhysteridae (both p < 0.05), as well as the necromenic Pristionchus (p < 0.001). At high taxonomic level, no differences were observed in fungal rhizosphere communities between native and non‐native ragwort species. The impact of plant developmental stages on rhizosphere biota was prominent. The overall bacterial and fungal biomasses, as well as a remarkably consistent set of constituents (Actinobacteria, α‐ and β‐Proteobacteria and Bacteroidetes) were negatively affected by plant stage for both ragwort species. Although later developmental stages of plants generally coincided with lower levels for individual nematode taxa, densities of the fungivorous genera Diphtherophora and Tylolaimophorus remain unaltered. Hence, even at a high taxonomic level, differential effects of native and non‐native ragwort could be pinpointed. However, plant developmental stage has a more prominent impact and this impact was similar in nature for both native and non‐native ragwort species.     

Legacy effects of drought on plant growth and the soil food web

Soils deliver important ecosystem services, such as nutrient provision for plants and the storage of carbon (C) and nitrogen (N), which are greatly impacted by drought. Both plants and soil biota affect soil C and N availability, which might in turn affect their response to drought, offering the potential to feed back on each other’s performance. In a greenhouse experiment, we compared legacy effects of repeated drought on plant growth and the soil food web in two contrasting land-use systems: extensively managed grassland, rich in C and with a fungal-based food web, and intensively managed wheat lower in C and with a bacterial-based food web. Moreover, we assessed the effect of plant presence on the recovery of the soil food web after drought. Drought legacy effects increased plant growth in both systems, and a plant strongly reduced N leaching. Fungi, bacteria, and their predators were more resilient after drought in the grassland soil than in the wheat soil. The presence of a plant strongly affected the composition of the soil food web, and alleviated the effects of drought for most trophic groups, regardless of the system. This effect was stronger for the bottom trophic levels, whose resilience was positively correlated to soil available C. Our results show that plant belowground inputs have the potential to affect the recovery of belowground communities after drought, with implications for the functions they perform, such as C and N cycling.     

Challenging urban species diversity: contrasting phylogenetic patterns across plant functional groups in Germany

Cities are hotspots of plant species richness, harboring more species than their rural surroundings, at least over large enough scales. However, species richness does not necessarily cover all aspects of biodiversity such as phylogenetic relationships. Ignoring these relationships, our understanding of how species assemblages develop and change in a changing environment remains incomplete. Given the high vascular plant species richness of urbanized areas in Germany, we asked whether these also have a higher phylogenetic diversity than rural areas, and whether phylogenetic diversity patterns differ systematically between species groups characterized by specific functional traits. Calculating the average phylogenetic distinctness of the total German flora and accounting for spatial autocorrelation, we show that phylogenetic diversity of urban areas does not reflect their high species richness. Hence, high urban species richness is mainly due to more closely related species that are functionally similar and able to deal with urbanization. This diminished phylogenetic information might decrease the flora's capacity to respond to environmental changes.