Genetic variation in a tropical tree species influences the associated epiphytic plant and invertebrate communities in a complex forest ecosystem

Genetic differences among tree species, their hybrids and within tree species are known to influence associated ecological communities and ecosystem processes in areas of limited species diversity. The extent to which this same phenomenon occurs based on genetic variation within a single tree species, in a diverse complex ecosystem such as a tropical forest, is unknown. The level of biodiversity and complexity of the ecosystem may reduce the impact of a single tree species on associated communities. We assessed the influence of within-species genetic variation in the tree Brosimum alicastrum (Moraceae) on associated epiphytic and invertebrate communities in a neotropical rainforest. We found a significant positive association between genetic distance of trees and community difference of the epiphytic plants growing on the tree, the invertebrates living among the leaf litter around the base of the tree, and the invertebrates found on the tree trunk. This means that the more genetically similar trees are host to more similar epiphyte and invertebrate communities. Our work has implications for whole ecosystem conservation management, since maintaining sufficient genetic diversity at the primary producer level will enhance species diversity of other plants and animals.     

Aquatic invertebrate communities in tank bromeliads: how well do classic ecological patterns apply?

Tank bromeliads (Bromeliaceae) often occur in high densities in the Neotropics and represent a key freshwater habitat in montane forests, housing quite complex invertebrate communities. We tested the extent to which there are species richness–altitude, richness–environment, richness–size, richness–habitat complexity and richness–isolation relationships for the aquatic invertebrate communities from 157 bromeliads in Cusuco National Park, Honduras. We found that invertebrate species richness and abundance correlated most strongly, and positively, with habitat size, which accounted for about a third of the variance in both. Apart from bromeliad size (equivalent of the species–area relationship), we found remarkably little evidence of classic biogeographic and ecological relationships with species richness in this system. Community composition correlated with altitude, bromeliad size and position, though less than 20% of the variation was accounted for by the tested variables. The turnover component of dissimilarity between the communities correlated with altitude, while the nestedness-resultant component was related to bromeliad size. The unexplained variance could reflect a large stochastic component in the system, associated with the ephemerality of the habitat patches (both the plants themselves and the fluctuations in their water content) and stochasticity due to the dispersal dynamics in the system. We conclude that there is a small contribution of classic biogeographic factors to the diversity and community composition of aquatic invertebrates communities in bromeliads. This may be due to the highly dynamic nature of this system, with small patch sizes and high emigration rates. The patterns may mostly be driven by factors affecting colonisation success.     

Environmental determinants of macroinvertebrate diversity in small water bodies: insights from tank-bromeliads

The interlocking leaves of tank-forming bromeliads (Bromeliaceae) collect rainwater and detritus, thus creating a freshwater habitat for specialized organisms. Their abundance and the possibility of quantifying communities with accuracy give us unparalleled insight into how changes in local to regional environments influence community diversity in small water bodies. We sampled 365 bromeliads (365 invertebrate communities) along a southeastern to northwestern range in French Guiana. Geographic locality determined the species pool for bromeliad invertebrates, and local environments determined the abundance patterns through the selection of traits that are best adapted to the bromeliad habitats. Patterns in community structure mostly emerged from patterns of predator species occurrence and abundance across local–regional environments, while the set of detritivores remained constant. Water volume had a strong positive correlation with invertebrate diversity, making it a biologically relevant measure of the pools’ carrying capacity. The significant effects of incoming detritus and incident light show that changes in local environments (e.g., the conversion of forest to cropping systems) strongly influence freshwater communities. Because changes in local environments do not affect detritivores and predators equally, one may expect functional shifts as sets of invertebrates with particular traits are replaced or complemented by other sets with different traits.     

Contrasting covariation of above- and belowground invertebrate species across plant genotypes

1. Invertebrate species generally do not respond independently to genotypic variation in plants, giving rise to clusters of species that naturally associate with or avoid certain genotypes. This covariation causes coevolution to be diffuse rather than pairwise. Studies on this topic, however, have never considered the belowground invertebrate community, leaving a critical gap in our understanding. 2. We investigated the covariation among naturally colonising above- and belowground invertebrate species across six genetically distinct populations of the dune grass Ammophila arenaria. After having grown from seed in a common garden, plants were randomised in a single field site to exclude all but broad-sense genetic variation. 3. Strong positive covariation across genotypes among both above- and belowground invertebrates was detected, while correlations between these two groups were negative. This clustering of above- and belowground species matched well with order level taxonomy. Host range, trophic level and food type on the other hand did not correspond well with the clusters. Within the cluster of aboveground fauna, subsequent groupings were not related to any phylogenetic or ecological characteristic, although correlations within these subgroups were very high. We furthermore demonstrated significant differences in multiple invertebrate species occurrence between plant genotypes, in general as well as at the above- and belowground level. 4. The observed strong covariation suggests diffuse coevolution between A. arenaria and its associated invertebrate species. The trade-off between root and shoot invertebrates could however hamper directional selection on resistance to either group. 5. Our results clearly demonstrate the need for studies of plant-animal interactions to include the belowground fauna, as this might drastically alter our general conception of how plants and their associated animal communities interact and how these interactions shape the process of evolution.     

The influence of range‐wide plant genetic variation on soil invertebrate communities

Plant genetic variation can have far‐reaching effects on associated communities and ecosystems. Heritable variation in ecologically relevant plant traits is often non‐randomly distributed across a species’ range and can exhibit geographic clines. In the event of range expansions and migration, previously unfamiliar genotypes may have large impacts on resident communities and ecosystems due to the introduction of novel and heritable phenotypic variation. Here we test the hypothesis that geographic origin of a focal plant genotype has effects on belowground invertebrate communities using a common garden field experiment. We sampled soil invertebrates from 103 Oenothera biennis genotypes, which were collected from across the species’ range and planted into a common garden field experiment at the northern range limit. We enumerated 24 000 individuals from 190 morphospecies and found that the diversity, abundance, and composition of soil invertebrate communities varied greatly among plant genotypes. Despite strong effects of plant genotype, we found few genetic correlations between plant traits and soil invertebrate community variables. However, herbivore damage was strongly related to variation in the soil invertebrate community. Geographic origin of plant genotypes had at most a weak effect on belowground communities. We speculate that predicting the extended effects of population movement on associated communities will require detailed knowledge of the trait variation occurring within focal species across particular environmental gradients.     

Size distribution of aquatic invertebrates in two prairie wetlands, with and without fish, with implications for community production

1. We compared the size distribution of aquatic invertebrates in two prairie wetlands, one supporting a population of fathead minnows and the other fishless. Both wetlands were sampled in three depth zones on three dates, allowing assessment of temporal and spatial variation.
2. We determined biomass of aquatic invertebrates in 17 log₂ size classes, and used these data to develop normalized size spectra. We also coupled size distributions with an allometric model to estimate relative production at the community level.
3. The composition of the invertebrate communities differed greatly between sites, and invertebrate biomass was higher in nearly all size classes in the fishless wetland. Intercepts of normalized size spectra were significantly different between wetlands, but slopes generally were not, indicating differences in standing-stock biomass but similar size structures between the two invertebrate communities. Higher standing-stock biomass in the fishless wetland resulted in higher relative production per unit area, but similar size distributions resulted in similar mass-specific production (P/B) between wetlands.
4. Our results indicate that invertebrate communities in prairie wetlands may have relatively consistent size structures in spite of large differences in community composition and standing-stock biomass. We hypothesize that the observed differences are because of predation by the minnow population and/or differences in the macrophyte communities between the two sites. However, the relative importance of macrophytes and fish predation in structuring invertebrate communities in prairie wetlands is poorly known.     

空间和环境因子对黄河口自然和淡水恢复湿地底栖动物群落的差异影响

生物多样性的形成和维持机制是生态学研究的核心问题,其中环境和空间因子在群落构建中的相对重要性是生态学家面临的重要挑战。为探究黄河口湿地底栖动物群落的关键影响因子,及环境和空间因子对底栖动物群落结构的相对调控作用。于2017年10月与2018年5月对黄河口湿地32个样点（淡水恢复湿地19个和自然湿地13个）的底栖动物和水体理化指标进行采集分析。非度量多维标度排序（NMDS）结果显示,黄河口淡水恢复湿地和自然湿地的底栖动物群落结构显著不同。典范对应分析（CCA）表明,影响淡水恢复湿地底栖动物群落结构的环境因子主要为电导率、盐度和氧化还原电位;而自然湿地底栖动物群落结构主要受pH和无机碳的影响;盐度是两类湿地底栖动物群落组成差异的关键因子。变差分解（VPA）结果显示,环境过滤对淡水恢复湿地底栖动物群落起主导作用;在自然湿地中,空间因子对底栖动物群落具有主要的调控作用,同时环境和空间因子的相互作用也至关重要。本研究明确了黄河口的自然和恢复湿地中环境和空间因素对底栖动物群落特征的相对作用,对黄河三角洲河口湿地中生物多样性的保护和生态系统管理提供参考。     

Intraspecific chemical diversity among neighbouring plants correlates positively with plant size and herbivore load but negatively with herbivore damage

Intraspecific plant diversity can modify the properties of associated arthropod communities and plant fitness. However, it is not well understood which plant traits determine these ecological effects. We explored the effect of intraspecific chemical diversity among neighbouring plants on the associated invertebrate community and plant traits. In a common garden experiment, intraspecific diversity among neighbouring plants was manipulated using three plant populations of wild cabbage that differ in foliar glucosinolates. Plants were larger, harboured more herbivores, but were less damaged when plant diversity was increased. Glucosinolate concentration differentially correlated with generalist and specialist herbivore abundance. Glucosinolate composition correlated with plant damage, while in polycultures, variation in glucosinolate concentrations among neighbouring plants correlated positively with herbivore diversity and negatively with plant damage levels. The results suggest that intraspecific variation in secondary chemistry among neighbouring plants is important in determining the structure of the associated insect community and positively affects plant performance.     

How effective are non-destructive sampling methods to assess aquatic invertebrate diversity in bromeliads?

With the growing interest in small aquatic water bodies, especially as naturally replicated model systems for ecological research, aquatic invertebrate communities in phytotelmata are increasingly receiving attention these days. The recognition of the substantial contribution to the regional species pool of specialised species draws further attention to these small and often temporary habitats. The methods currently used for studying communities in some types of phytotelma, such as bromeliads, tend to be destructive, typically involving complete dissection of the plant. The expected increase in sampling intensity associated with the increasing interest in phytotelmata may result in a negative impact on plant populations in some areas, decreasing numbers in an unsustainable way, especially in locations with ongoing, intensive research. We therefore aimed to investigate whether less-destructive sampling methods can achieve sufficient data quality to allow their use as alternatives to complete plant dissection. We tested the effectiveness of three such methods in measuring the aquatic invertebrate communities in tank bromeliads (Tillandsia guatemalensis) in Cusuco National Park, Honduras. The three methods were pipetting the water out of the bromeliad, turning the bromeliad upside down and dissecting only the outer part of the plant (the oldest, often deteriorating leaves). Overall, we found that these methods were poor predictors of richness and abundance of the organisms in communities. However, we found big differences between taxonomic groups, depending in part on the ecology of the organisms, and we suggest that some less-destructive alternative methods may be appropriate for studying some specific groups (e.g. Culicidae). Based on these results and a rapid survey of the abundance of bromeliads in the national park, we question whether intensive, ongoing research into aquatic invertebrate communities in similar phytotelma populations is sustainable. From the point of view of conservation, alternative model systems need to be found.     

Effects of Seasonality and Dispersal on the Ciliate Community Inhabiting Bromeliad Phytotelmata in Riparian Vegetation of a Large Tropical River

This study evaluated the influence of rainfall amount on the abundance, species richness, and species occurrence and abundance distribution of the ciliate community associated with the bromeliad Aechmea distichantha. The plants were collected from a rock wall of about 10‐km long at the left bank of Paraná River. We assessed the effects of both spatial and temporal variables on the community attributes, as well as whether plants geographically closer have a similar abundance distribution and species composition. The ciliate community was substantially distinct between both hydrological periods, with greater values of species richness and abundance in the rainy period. No spatial structuring (differences in the species occurrence and abundance distribution among strata) or geographical similarity (similarity in ciliate species composition among the plants) was found. Multiple regression analysis showed a positive relationship only between the ciliate abundances and water volumes for both periods. Although few of the formulated predictions were confirmed, our study provides valuable information on the ecological aspects of the ciliate community inhabiting bromeliad phytotelmata.     

Host‐plant genotypic diversity and community genetic interactions mediate aphid spatial distribution

Genetic variation in plants can influence the community structure of associated species, through both direct and indirect interactions. Herbivorous insects are known to feed on a restricted range of plants, and herbivore preference and performance can vary among host plants within a species due to genetically based traits of the plant (e.g., defensive compounds). In a natural system, we expect to find genetic variation within both plant and herbivore communities and we expect this variation to influence species interactions. Using a three‐species plant‐aphid model system, we investigated the effect of genetic diversity on genetic interactions among the community members. Our system involved a host plant (Hordeum vulgare) that was shared by an aphid (Sitobion avenae) and a hemi‐parasitic plant (Rhinanthus minor). We showed that aphids cluster more tightly in a genetically diverse host‐plant community than in a genetic monoculture, with host‐plant genetic diversity explaining up to 24% of the variation in aphid distribution. This is driven by differing preferences of the aphids to the different plant genotypes and their resulting performance on these plants. Within the two host‐plant diversity levels, aphid spatial distribution was influenced by an interaction among the aphid's own genotype, the genotype of a competing aphid, the origin of the parasitic plant population, and the host‐plant genotype. Thus, the overall outcome involves both direct (i.e., host plant to aphid) and indirect (i.e., parasitic plant to aphid) interactions across all these species. These results show that a complex genetic environment influences the distribution of herbivores among host plants. Thus, in genetically diverse systems, interspecific genetic interactions between the host plant and herbivore can influence the population dynamics of the system and could also structure local communities. We suggest that direct and indirect genotypic interactions among species can influence community structure and processes.     

Does Facilitation of Faunal Recruitment Benefit Ecosystem Restoration? An Experimental Study of Invertebrate Assemblages in Wetland Mesocosms

We used wetland mesocosms (1) to experimentally assess whether inoculating a restored wetland site with vegetation/sediment plugs from a natural wetland would alter the development of invertebrate communities relative to unaided controls and (2) to determine if stocking of a poor invertebrate colonizer could further modify community development beyond that due to simple inoculation. After filling mesocosms with soil from a drained and cultivated former wetland and restoring comparable hydrology, mesocosms were randomly assigned to one of three treatments: control (a reference for unaided community development), inoculated (received three vegetation/sediment cores from a natural wetland), and stocked + inoculated (received three cores and were stocked with a poorly dispersing invertebrate group—gastropods). All mesocosms were placed 100 m from a natural wetland and allowed to colonize for 82 days. Facilitation of invertebrate colonization led to communities in inoculated and stocked + inoculated treatments that contrasted strongly with those in the unaided control treatment. Control mesocosms had the highest taxa richness but the lowest diversity due to high densities and dominance of Tanytarsini (Diptera: Chironomidae). Community structure in inoculated and stocked + inoculated mesocosms was more similar to that of a nearby natural wetland, with abundance more evenly distributed among taxa, leading to diversity that was higher than in the control treatment. Inoculated and stocked + inoculated communities were dominated by non‐aerial invertebrates, whereas control mesocosms were dominated by aerial invertebrates. These results suggest that facilitation of invertebrate recruitment does indeed alter invertebrate community development and that facilitation may lead to a more natural community structure in less time under conditions simulating wetland restoration.     

Complex indirect effects of epiphytic bromeliads on the invertebrate food webs of their support tree

Ecosystem engineers are species that affect others through the provision of habitat rather than consumptive resources. The extent to which ecosystem engineers can indirectly affect entire food webs, however, is poorly understood. Epiphytic tank bromeliads (Bromeliaceae) are ecosystem engineers that are common throughout the Neotropics, and are associated with a variety of predatory arthropods. Here, we examine if bromeliads, by increasing predator densities, indirectly benefit their support tree through reduction in herbivorous insects and leaf damage. We observed and manipulated bromeliad densities in Costa Rican orange orchards, and measured impacts on leaf damage and arboreal and bromeliad invertebrate communities in two different seasons. Our results show that bromeliads are associated with predatory and herbivorous invertebrates but not leaf damage. Bromeliads were correlated with increased densities of their associated predators, especially ants and hunting spiders, but we could not confirm a causal link. Associations with bromeliads changed over time, with seasonal shifts interfering with responses to our manipulations. Bromeliads had a reduced association with predators in the dry season. Moreover, a null association between bromeliads and herbivorous invertebrates in the dry season unexpectedly became positive in the wet season. In summary, we have only limited evidence that bromeliads indirectly promote the top‐down control of arboreal herbivores; instead, our manipulations suggest that bromeliads increase herbivore densities in the wet season. This research suggests that although bromeliads may act as ecosystem engineers, indirectly influencing the invertebrate food web on support trees, their effects are trophically complex and seasonally dependent. Abstract in Spanish is available with online material.     

Influence of microclimate and species interactions on the composition of plant and invertebrate communities in alpine northern Norway

We assessed the effects of both biotic processes and abiotic factors on the community composition of vascular plant species and invertebrates at a site in northern Norway. Plant species were assigned to functional (woody versus herbaceous) and biogeographic (boreal versus alpine) groups. Invertebrate species were classified as either herbivore or predator. When species interactions and effects of the abiotic environment were partitioned, boreal species appeared to influence the distribution of alpine species and woody species the distribution of herbaceous species. Analysis of partial correlations indicated that facilitation was the dominant mode of interaction between the two pairs of plant groups. Among abiotic factors, the thermal environment probably influenced all components of the plant and invertebrate communities, except for predatory invertebrates, and wind appeared important in determining the composition of woody and alpine components of the plant community but not the herbaceous component. The composition of the boreal component of the plant community apparently influenced the composition of all invertebrate communities, except for predatory invertebrates. The composition of the woody component of the plant community influenced the composition of both herbivore and predator communities. The alpine plant-community composition influenced predatory invertebrate community composition. Woody plant community composition influenced the composition of both herbivore and predator communities. Our analytic approach, based on two kinds of structural equation models (d-separation and path analysis), provides a useful method for identifying the biotic as well as abiotic factors that influence community structure.     

Exotic vertebrate and invertebrate herbivores differ in their impacts on native and exotic plants: a meta-analysis

Herbivores modify various ecological processes including interactions between native and exotic plants that may affect invasion success by the exotic plants. It is unknown whether different types of exotic herbivores have similar effects on native and exotic plants. Using two distinct data sets, we ran meta-analyses to compare exotic vertebrate and invertebrate herbivore preferences for, and effects on performance and population sizes of native and exotic plants. We found that exotic vertebrate herbivores have positive effects on exotic plant performance and population sizes, and no significant effects on native plants. Exotic invertebrates have significant negative effects on performance and population sizes of both exotic and native plants. Vertebrates prefer to feed on native plants relative to exotic plants, while invertebrates prefer the exotic plants to native plants. Thus the exotic vertebrate herbivores may aid invasiveness of exotic plants, in accordance with the invasional meltdown hypothesis, while exotic invertebrate herbivores probably have no net effect on invasion process of the exotic plants. Invertebrate herbivore preferences for exotic plants support the biotic resistance hypothesis, as the native plants probably resist the invertebrate herbivory. We also tested an evolutionary logic that posits that herbivores with similar evolutionary history as plants will affect the plants less negatively than plants with which they have not co-evolved. Our results indicate that there is no consistent pattern in effects of exotic vertebrate and invertebrate herbivores on exotic plants with or without which they have co-evolved.     

Understorey environments influence functional diversity in tank‐bromeliad ecosystems

1. A substantial fraction of the freshwater available in neotropical forests is impounded within the rosettes of bromeliads that form aquatic islands in a terrestrial matrix. The ecosystem functioning of bromeliads is known to be influenced by the composition of the contained community but it is not clear whether bromeliad food webs remain functionally similar against a background of variation in the understorey environment. 2. We considered a broad range of environmental conditions, including incident light and incoming litter, and quantified the distribution of a very wide range of freshwater organisms (from viruses to macroinvertebrates) to determine the factors that influence the functional structure of bromeliad food webs in samples taken from 171 tank‐bromeliads. 3. We observed a gradient of detritus‐based to algal‐based food webs from the understorey to the overstorey. Algae, rotifers and collector and predatory invertebrates dominated bromeliad food webs in exposed areas, whereas filter‐feeding insects had their highest densities in shaded forest areas. Viruses, bacteria and fungi showed no clear density patterns. Detritus decomposition is mainly due to microbial activity in understorey bromeliads where filter feeders are the main consumers of microbial and particulate organic matter (POM). Algal biomass may exceed bacterial biomass in sun‐exposed bromeliads where amounts of detritus were lower but functional diversity was highest. 4. Our results provide evidence that tank‐bromeliads, which grow in a broad range of ecological conditions, promote aquatic food web diversity in neotropical forests. Moreover, although bromeliad ecosystems have been categorised as detritus‐based systems in the literature, we show that algal production can support a non‐detrital food web in these systems.     

优势种去除对崇明东滩盐沼湿地生态系统的影响

盐沼生态系统环境梯度明显,物种组成较简单,是研究生物多样性与生态系统功能关系的理想对象。本研究以崇明东滩盐沼湿地为研究区域,研究优势种去除对植物群落结构以及底栖动物群落的影响。结果表明:(1)去除处理仅对植物群落分株密度有极显著效应(P0.01)。去除组和对照组物种组成差异随时间增加而减小,处理效应逐渐减弱。(2)去除组底栖动物密度均低于对照组,但差异不显著。(3)盐沼植物群落特征与底栖动物群落有密切关系,植物密度、冠层高度与底栖动物密度相关性极显著。去除优势种后,植物群落分株密度升高,群落内剩余物种占比有所上升,次优势种对群落的补偿效应具有较大贡献;而底栖动物群落密度下降,其生物量和多样性指数的变化趋势与密度并不一致。上述结果表明生物多样性变化影响了盐沼湿地生态系统植物群落和底栖动物群落结构,进而可能影响物质循环和能量流动过程。     

Seasonal Changes in the Rhizosphere Microbial Communities Associated with Field-Grown Genetically Modified Canola (Brassica napus)

The introduction of transgenic plants into agricultural ecosystems has raised the question of the ecological impact of these plants on nontarget organisms, such as soil bacteria. Although differences in both the genetic structure and the metabolic function of the microbial communities associated with some transgenic plant lines have been established, it remains to be seen whether these differences have an ecological impact on the soil microbial communities. We conducted a 2-year, multiple-site field study in which rhizosphere samples associated with a transgenic canola variety and a conventional canola variety were sampled at six times throughout the growing season. The objectives of this study were to identify differences between the rhizosphere microbial community associated with the transgenic plants and the rhizosphere microbial community associated with the conventional canola plants and to determine whether the differences were permanent or depended on the presence of the plant. Community-level physiological profiles, fatty acid methyl ester profiles, and terminal amplified ribosomal DNA restriction analysis profiles of rhizosphere microbial communities were compared to the profiles of the microbial community associated with an unplanted, fallow field plot. Principal-component analysis showed that there was variation in the microbial community associated with both canola variety and growth season. Importantly, while differences between the microbial communities associated with the transgenic plant variety were observed at several times throughout the growing season, all analyses indicated that when the microbial communities were assessed after winter, there were no differences between microbial communities from field plots that contained harvested transgenic canola plants and microbial communities from field plots that did not contain plants during the field season. Hence, the changes in the microbial community structure associated with genetically modified plants were temporary and did not persist into the next field season.     

Seasonal changes in the rhizosphere microbial communities associated with field-grown genetically modified canola (Brassica napus)

The introduction of transgenic plants into agricultural ecosystems has raised the question of the ecological impact of these plants on nontarget organisms, such as soil bacteria. Although differences in both the genetic structure and the metabolic function of the microbial communities associated with some transgenic plant lines have been established, it remains to be seen whether these differences have an ecological impact on the soil microbial communities. We conducted a 2-year, multiple-site field study in which rhizosphere samples associated with a transgenic canola variety and a conventional canola variety were sampled at six times throughout the growing season. The objectives of this study were to identify differences between the rhizosphere microbial community associated with the transgenic plants and the rhizosphere microbial community associated with the conventional canola plants and to determine whether the differences were permanent or depended on the presence of the plant. Community-level physiological profiles, fatty acid methyl ester profiles, and terminal amplified ribosomal DNA restriction analysis profiles of rhizosphere microbial communities were compared to the profiles of the microbial community associated with an unplanted, fallow field plot. Principal-component analysis showed that there was variation in the microbial community associated with both canola variety and growth season. Importantly, while differences between the microbial communities associated with the transgenic plant variety were observed at several times throughout the growing season, all analyses indicated that when the microbial communities were assessed after winter, there were no differences between microbial communities from field plots that contained harvested transgenic canola plants and microbial communities from field plots that did not contain plants during the field season. Hence, the changes in the microbial community structure associated with genetically modified plants were temporary and did not persist into the next field season.     

After‐life effects: living and dead invertebrates differentially affect plants and their associated above‐ and belowground multitrophic communities

Above–belowground (AG–BG) studies typically focus on plant‐mediated effects inflicted by living organisms. However, animal cadavers may also play an important role in AG–BG interactions. Here, we explore whether living and dead foliar‐feeding and soil‐dwelling invertebrates differentially affect plants and their associated AG and BG multitrophic communities. In a mesocosm study we separated effects of living and dead locusts (AG herbivores) and earthworms (BG detritivores) on experimental multitrophic communities consisting of eight plant species, an AG aphid and parasitoid community and a BG nematode community. We measured root and shoot biomass and determined plant community composition and densities of aphids, parasitoids and nematodes. Living locusts decreased total shoot and root biomass in the mesocosms, whereas living earthworms enhanced total root biomass. Cadavers of both invertebrates strongly increased total root and shoot biomass, and changed the plant community composition mainly via enhanced growth of grasses. Earthworm cadavers affected plant biomass and community composition more strongly than their living counterparts, while this was reversed for locusts. Structural equation models showed that aphids and parasitoids were influenced via changes in plant community composition. Nematode densities in the soil, especially those of bacterivorous and entomopathogenic nematodes, were strongly increased by dead invertebrates, but unaffected by living ones. We conclude that effects of invertebrates on plant growth and densities of AG and BG organisms strongly depend on whether the invertebrates are dead or alive. Remarkably, invertebrate cadavers may inflict even stronger effects than their living counterparts. Hence, our study reveals an important, but often neglected, role of animal cadavers in AG–BG studies.