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Across the globe, ecological communities are confronted with multiple global environmental change drivers, and they are responding in complex ways ranging from behavioral, physiological, and morphological changes within populations to changes in community composition and food web structure with consequences for ecosystem functioning. A better understanding of global change‐induced alterations of multitrophic biodiversity and the ecosystem‐level responses in terrestrial ecosystems requires holistic and integrative experimental approaches to manipulate and study complex communities and processes above and below the ground. We argue that mesocosm experiments fill a critical gap in this context, especially when based on ecological theory and coupled with microcosm experiments, field experiments, and observational studies of macroecological patterns. We describe the design and specifications of a novel terrestrial mesocosm facility, the iDiv Ecotron. It was developed to allow the setup and maintenance of complex communities and the manipulation of several abiotic factors in a near‐natural way, while simultaneously measuring multiple ecosystem functions. To demonstrate the capabilities of the facility, we provide a case study. This study shows that changes in aboveground multitrophic interactions caused by decreased predator densities can have cascading effects on the composition of belowground communities. The iDiv Ecotrons technical features, which allow for the assembly of an endless spectrum of ecosystem components, create the opportunity for collaboration among researchers with an equally broad spectrum of expertise. In the last part, we outline some of such components that will be implemented in future ecological experiments to be realized in the iDiv Ecotron., The iDiv Ecotron is a novel and unique research platform that was explicitly designed to address the perpetual claim that biodiversity–ecosystem function (BEF) research in terrestrial ecosystems needs to move beyond the manipulation of diversity at single trophic levels to embrace the multitrophic complexity of ecological communities.

Animals, such as termites, have largely been overlooked as global-scale drivers of biogeochemical cycles1,2, despite site-specific findings3,4. Deadwood turnover, an important component of the carbon cycle, is driven by multiple decay agents. Studies have focused on temperate systems5,6, where microbes dominate decay7. Microbial decay is sensitive to temperature, typically doubling per 10°C increase (decay effective Q10 = ~2)8–10. Termites are important decayers in tropical systems3,11–13 and differ from microbes in their population dynamics, dispersal, and substrate discovery14–16, meaning their climate sensitivities also differ. Using a network of 133 sites spanning 6 continents, we report the first global field-based quantification of temperature and precipitation sensitivities for termites and microbes, providing novel understandings of their response to changing climates. Temperature sensitivity of microbial decay was within previous estimates. Termite discovery and consumption were both much more sensitive to temperature (decay effective Q10 = 6.53), leading to striking differences in deadwood turnover in areas with and without termites. Termite impacts were greatest in tropical seasonal forests and savannas and subtropical deserts. With tropicalization17 (i.e., warming shifts to a tropical climate), the termite contribution to global wood decay will increase as more of the earth’s surface becomes accessible to termites. Acknowledgements. For materials, field, and lab assistance, we thank the Australian Landscape Trust Team, Juan Martínez de Aragón, Eric Roy, Hugh Wilson, Kylie Brice, Bethanie Coleman, Brendan Delroy, Coline Deveautour, Suzanne Donn, Generci Assis Neves, Ricardo Marques Barreiros, Ricardo Marques Barreiros, Generci Assis Neves from Resineves Agroflorestal company, Alfred Lochner, Pousada Vale do Céu, Humberto Robles, María M. Rivera, and Grant Kirker. Steve Allison and Tom Crowther provided comments on the manuscript and Abbey Yatsko helped with figure and manuscript formatting. Funding for this work was provided by US National Science Foundation (DEB-1655759, 2149151, 1713502, 1713435, 1647502, 1546686, 1831952), George Washington University, USDA Forest Service, Centre College Faculty Development Funds, Australia Terrestrial Ecosystem Research Network (TERN) National Collaborative Research Infrastructure Strategy (NCRIS), Royal Society-FCDO Africa Capacity Building Initiative, New Phytologist Foundation, Fondecyt grant:1160329, Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Brasil (CAPES), Department of Ecology and Conservation of the Federal University of Lavras, CNPq, FAPEMIG, Australian Academy of Science 2017 Thomas Davies Research Grant, Australian Research Council (DP160103765), UK National Environment Research Council (NE/L000016/1), Fundação de Amparo à Pesquisa do Estado de São Paulo, Brazil (NERC - FAPESP)19/07773-1, Environment Research and Technology Development Fund (ERTDF; JPMEERF15S11420) of the Environmental Restoration and Conservation Agency (ERCA) of Japan, COLCIENCIAS (No. FP44842-046-2017), Spanish Government (grant PID2019-110521GB-I00), Catalan Government (grant SGR 2017-1005), Fundación Ramón Areces (grant ELEMENTAL-CLIMATE), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT) PICT 2019- 2472, National Agency for the Promotion of Research, Technological Development and Innovation, Scientific and Technological Research Project 2018-01561 (PICT 2018- 01561), iDiv German Research Foundation (DFG– FZT 118, 202548816), European Research Council (ERC), European Union's Horizon 2020 research and innovation program (grant agreement no. 677232).

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity., Nature Communications, 12 (1), ISSN:2041-1723

Recent research shows that earthworms can alter defense traits of plants against herbivores and pathogens by affecting soil biochemistry. Yet, the effects of invasive earthworms on defense traits of native plants from previously earthworm‐free ecosystems as well as the consequences for multitrophic interactions are virtually unknown.Here we use a combination of an observational study and a complementary experimental study to investigate the effects of invasive earthworms on leaf defense traits, herbivore damage and pathogen infection in two poplar tree species (Populus balsamifera and Populus tremuloides) native to North American boreal forests.Our observational study showed that earthworm invasion was associated with enhanced leaf herbivory (by leaf‐chewing insects) in saplings of both tree species. However, we only detected significant shifts in the concentration of chemical defense compounds in response to earthworm invasion for P. balsamifera. Specifically, leaf phenolic concentrations, including salicinoids and catechin, were lower in P. balsamifera from earthworm‐invaded sites.Our experimental study confirmed an earthworm‐induced reduction in leaf defense levels in P. balsamifera for one of the defense compounds, tremulacin. The experimental study additionally showed that invasive earthworms reduced leaf dry matter content, potentially increasing leaf palatability, and enhanced susceptibility of trees to infection by a fungal pathogen, but not to aphid infestation, in the same tree species. Synthesis. Our results show that invasive earthworms can decrease the concentrations of some chemical defense compounds in P. balsamifera, which could make them susceptible to leaf‐chewing insects. Such potential impacts of invasive earthworms are likely to have implications for tree survival and competition, native tree biodiversity and ecosystem functioning., Invasive earthworms can dramatically alter native biodiversity and ecosystem functioning. Yet, we know little how invasive earthworms affect the chemical defense of native trees. Here using a combination of observational and experimental studies, we show that invasive earthworms from Europe can decrease the concentrations of some chemical defense compounds in a native North American deciduous tree, which could make them susceptible to leaf‐chewing insects.

Two major human-made problems in rice production systems in the north of Vietnam concern the low plant-available silicon content of soils and the low biodiversity. The results of the LEGATO project suggest a change to an environmentally friendly rice production system that will help to recover biodiversity. We propose here a framework for a demonstration and dissemination model that will be exemplary for the farmers once it has been successfully realized. We advocate local-option models in different districts to demonstrate to farmers. The methods should be adapted to local and ecoregional differences in climate and land-use tradition, and they explicitly take into account soil care, organic fertilizer, manual weeding, native nectar-rich plant bunds, manual pest snail collection, hymenopteran nesting aids, and biodiversity and yield monitoring.

The transdisciplinary research project LEGATO analysed the combined generation of provisioning, regulating and cultural ecosystem services in wet rice agriculture in South-East Asia and applied ecological engineering to future-proof it against global change and environmental pollution challenges. Due to its transdisciplinary character and the sheer size, a systematic stakeholder involvement was inevitable. Starting with stakeholder identification by snowballing from a limited number of contacts, we derived a multi-level stakeholder analysis and tried to involve those identified as relevant. Applying different means and aiming at different depth of involvement, the effort can be judged successful. The paper describes the methods used to identify and classify stakeholders, and key elements of the stakeholder management, guided by the BiodivERsA Stakeholder Activation Handbook. While on the local level, farmers and extension workers turned out to be influential and interested stakeholders in both countries, as well as local (PH) or provincial authorities (VN), differences were manifest on higher levels due to the divergent institutional setting. National-level agents were hard to get interested in both countries, and influential agents along the production chain, including middle men, were not really interested in collaboration.

1. For managed temperate forests, conservationists and policymakers favour fine-grained uneven-aged management over more traditional coarse-grained even-aged management, based on the assumption that within-stand habitat heterogeneity enhances biodiversity. There is, however, little empirical evidence to support this assumption. We investigated for the first time how differently grained forest management systems affect the biodiversity of multiple above- and below-ground taxa across spatial scales. 2. We sampled 15 taxa of animals, plants, fungi and bacteria within the largest contiguous beech forest landscape of Germany and classified them into functional groups. Selected forest stands have been managed for more than a century at different spatial grains. The even-aged (coarse-grained management) and uneven-aged (fine-grained) forests are comparable in spatial arrangement, climate and soil conditions. These were compared to forests of a nearby national park that have been unmanaged for at least 20 years. We used diversity accumulation curves to compare γ-diversity for Hill-numbers 0D (species richness), 1D (Shannon diversity) and 2D (Simpson diversity) between the management systems. Beta diversity was quantified as multiple-site dissimilarity. 3. Gamma diversity was higher in even-aged than in uneven-aged forests for at least one of the three Hillnumbers for six taxa (up to 77%), while eight showed no difference. Only bacteria showed the opposite pattern. Higher γ-diversity in even-aged forests was also found for forest specialists and saproxylic beetles. 4. Between-stand β-diversity was higher in even-aged than in uneven-aged forests for one third (all species) and half (forest specialists) of all taxa, driven by environmental heterogeneity between age-classes, while α-diversity showed no directional response across taxa or for forest specialists. 5. Synthesis and applications. Comparing even-aged and uneven-aged forest management in Central European beech forests, our results show that a mosaic of different age-classes is more important for regional biodiversity than high within-stand heterogeneity. We suggest reconsidering the current trend of replacing even-aged management in temperate forests. Instead, the variability of stages and stand structures should be increased to promote landscape scale biodiversity.

Background and aimsMinirhizotrons are commonly used to study root turnover which is essential for understanding ecosystem carbon and nutrient cycling. Yet, extracting data from minirhizotron images requires intensive annotation effort. Existing annotation tools often lack flexibility and provide only a subset of the required functionality. To facilitate efficient root annotation in minirhizotrons, we present the user-friendly open source tool rhizoTrak.Methods and resultsrhizoTrak builds on TrakEM2 and is publically available as Fiji plugin. It uses treelines to represent branching structures in roots and assigns customizable status labels per root segment. rhizoTrak offers configuration options for visualization and various functions for root annotation mostly accessible via keyboard shortcuts. rhizoTrak allows time-series data import and particularly supports easy handling and annotation of time series images. This is facilitated via explicit temporal links (connectors) between roots which are automatically generated when copying annotations from one image to the next. rhizoTrak includes automatic consistency checks and guided procedures for resolving conflicts. It facilitates easy data exchange with other software by supporting open data formats.ConclusionsrhizoTrak covers the full range of functions required for user-friendly and efficient annotation of time-series images. Its flexibility and open source nature will foster efficient data acquisition procedures in root studies using minirhizotrons.

The golden apple snail or GAS (Pomacea canaliculata) is an important invasive pest in irrigated rice that feeds on young rice plants. In many countries in SE-Asia, governments have recently decreased their support of training courses for snail management, because farmers are now considered to know how to effectively manage this pest. Although a great number of sustainable control methods is recommended which do not involve the use of pesticides, it is uncertain whether these are taken up by farmers. Probably, the easiest way to control GAS, is the application of synthetic ‘instant kill’ molluscicides, which can have detrimental effects on the environment, non-target species, and health. The aim of this study was to develop ideas for solutions on how to achieve a sustainable management of GAS without or at least a decreased use of molluscicides. In a large-scale approach, we conducted interviews with rice farmers in seven regions across Vietnam and the Philippines, assessing the participation in training courses, knowledge on snail ecology, the methods of controlling and the utilization of the snail, and the farmers’ suggestions on how to improve pest snail control. Only 23% of the farmers had previously received training in GAS management. We found that training neither had positive nor negative effects on the number of sustainable methods applied, molluscicide avoidance, concern about using molluscicides, or on the farmers’ knowledge about GAS. As much as 74% of the respondents applied molluscicides. Contrary to recommendations, farmers applied only few sustainable control methods. All farmers had clear knowledge gaps about GAS, especially in species identification, which can even further the ongoing decline of native mollusks in rice landscapes. We conclude that the decision to phase out information campaigns has been taken to rash, and that trainings in our study regions carried through previously had limited success, and thus need revision. To decrease molluscicide use, and to promote sustainable management on the large scale, we synthesized our results, and we suggest that information for farmers might be provided through media often accessed, such as TV, radio or the internet (e.g. by entertainment − education). We further discuss the potential of community cooperation to achieve sustainability. As there are distinct limitations to these approaches, we have developed the concept of local GAS management and utilization companies (GASMUC) which could take over sustainable control and utilization of GAS, and native mollusk conservation for an entire community.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

In a recent Forum paper, Wardle (Journal of Vegetation Science, 2016) questions the value of biodiversity–ecosystem function (BEF) experiments with respect to their implications for biodiversity changes in real world communities. The main criticism is that the previous focus of BEF experiments on random species assemblages within each level of diversity has ‘limited the understanding of how natural communities respond to biodiversity loss.’ He concludes that a broader spectrum of approaches considering both non-random gains and losses of diversity is essential to advance this field of research. Wardle's paper is timely because of recent observations of frequent local and regional biodiversity changes across ecosystems. While we appreciate that new and complementary experimental approaches are required for advancing the field, we question criticisms regarding the validity of BEF experiments. Therefore, we respond by briefly reiterating previous arguments emphasizing the reasoning behind random species composition in BEF experiments. We describe how BEF experiments have identified important mechanisms that play a role in real world ecosystems, advancing our understanding of ecosystem responses to species gains and losses. We discuss recent examples where theory derived from BEF experiments enriched our understanding of the consequences of biodiversity changes in real world ecosystems and where comprehensive analyses and integrative modelling approaches confirmed patterns found in BEF experiments. Finally, we provide some promising directions in BEF research.

International audience; Concern about the functional consequences of unprecedented loss in biodiversity has prompted biodiversity-ecosystem functioning (BEF) research to become one of the most active fields of ecological research in the past 25 years. Hundreds of experiments have manipulated biodiversity as an independent variable and found compelling support that the functioning of ecosystems increases with the diversity of their ecological communities. This research has also identified some of the mechanisms underlying BEF relationships, some context-dependencies of the strength of relationships, as well as implications for various ecosystem services that mankind depends upon. In this paper, we argue that a multitrophic perspective of biotic interactions in random and non-random biodiversity change scenarios is key to advance future BEF research and to address some of its most important remaining challenges. We discuss that the study and the quantification of multitrophic interactions in space and time facilitates scaling up from small-scale biodiversity manipulations and ecosystem function assessments to management-relevant spatial scales across ecosystem boundaries. We specifically consider multitrophic conceptual frameworks to understand and predict the context-dependency of BEF relationships. Moreover, we highlight the importance of the eco-evolutionary underpinnings of multitrophic BEF relationships. We outline that FAIR data (meeting the standards of findability, accessibility, interoperability, and reusability) and reproducible processing will be key to advance this field of research by making it more integrative. Finally, we show how these BEF insights may be implemented for ecosystem management, society, and policy. Given that human well-being critically depends on the multiple services provided by diverse, multitrophic communities, integrating the approaches of evolutionary ecology, community ecology, and ecosystem ecology in future BEF research will be key to refine conservation targets and develop sustainable management strategies.

LEGATO stands for “Land-use intensity and Ecological EnGineering—Assessment Tools for risks and Opportunities in irrigated rice based production systems.”

Rice is among the most important crops for human nutrition. The need to increase yields is commonly achieved by agricultural intensification that often comes along with negative impacts on the environment. In 2011, the interdisciplinary LEGATO project was launched with the aim to advance sustainable rice production. A key feature for sustainable rice production is silicon (Si) availability. Si can e.g. improve rice crop resistance against pathogens and prevent the uptake of toxic metals. Herein, we introduce the characteristics of the seven LEGATO study regions in Vietnam and the Philippines (i.e., climate, geology, soils, agricultural practices) ranging from lowlands with intensive production to mountain regions with extensive and traditional production systems. Secondly, we show data on concentrations of plant-available Si (Si pa ) in topsoils of paddy fields (Ap + Arp horizons) and of sites with an alternative land-use (forest, upland crops), and discuss factors determining spatial differences in Si pa concentrations. Results show that Si pa concentrations in topsoils of paddies (assessed by acetate extraction) greatly differ between Philippine (141–322 mg Si kg −1 ) and Vietnamese (20–51 mg Si kg −1 ) regions. This can be explained by differences in geo-/pedologic conditions between the countries. Large Si pa concentrations in the Philippines are due to recent rock formation by active volcanism, hence, by a large Si pa input due to mineral weathering in recent geologic history. Land-use can also affect Si pa in topsoils: in Philippine regions, Si pa concentrations were significantly larger for paddies than for other land-use types. In some of the Vietnamese paddies, Si pa concentrations are below critical values, thus might limit plant growth. Many Vietnamese farmers export Si from fields by removing straw residues with the harvest. Our subsequent research thus aims to test whether changes in harvest residue management can improve the Si supply to plants in Vietnamese regions.

Land-use intensification is a key driver of biodiversity change. However, little is known about how it alters relationships between the diversities of different taxonomic groups, which are often correlated due to shared environmental drivers and trophic interactions. Using data from 150 grassland sites, we examined how land-use intensification (increased fertilization, higher livestock densities, and increased mowing frequency) altered correlations between the species richness of 15 plant, invertebrate, and vertebrate taxa. We found that 54% of pairwise correlations between taxonomic groups were significant and positive among all grasslands, while only one was negative. Higher land-use intensity substantially weakened these correlations (35% decrease in r and 43% fewer significant pairwise correlations at high intensity), a pattern which may emerge as a result of biodiversity declines and the breakdown of specialized relationships in these conditions. Nevertheless, some groups (Coleoptera, Heteroptera, Hymenoptera and Orthoptera) were consistently correlated with multidiversity, an aggregate measure of total biodiversity comprised of the standardized diversities of multiple taxa, at both high and low land-use intensity. The form of intensification was also important; increased fertilization and mowing frequency typically weakened plant–plant and plant–primary consumer correlations, whereas grazing intensification did not. This may reflect decreased habitat heterogeneity under mowing and fertilization and increased habitat heterogeneity under grazing. While these results urge caution in using certain taxonomic groups to monitor impacts of agricultural management on biodiversity, they also suggest that the diversities of some groups are reasonably robust indicators of total biodiversity across a range of conditions. Read More: http://www.esajournals.org/doi/10.1890/14-1307.1

Numerous important ecosystem functions and services depend on soil biodiversity. However, little is known about the mechanisms which maintain the vast belowground biodiversity and about the filters shaping soil community composition. Yet, biotic interactions like facilitation and dispersal by animals are assumed to play a crucial role, particularly as most soil animal taxa are strongly limited in their active dispersal abilities. Here, we report on a newfound interaction of potentially high ubiquity and importance in soil communities: the endozoochorous dispersal of soil fauna by gastropods. We focus on the dispersal-limited group of oribatid mites, one of the most diverse and abundant soil animal groups. In a field survey in a German riparian forest, 73% of 40 collected slugs (Arion vulgaris) egested a total of 135 oribatid mites, belonging to 35 species. Notably, 70% of the egested mites were alive and survived the gut passage through slugs. Similar results were found for Roman snails (Helix pomatia), indicating the generality of our findings across different gastropod taxa. Complementary laboratory experiments confirmed our field observations, revealing that oribatid mites are, indeed, ingested and egested alive by slugs, and that they are able to independently escape the faeces and colonise new habitats. Our results strongly indicate that gastropods may help soil organisms to disperse within habitats, to overcome dispersal barriers, and to reach short-lived resource patches. Gastropods might even disperse whole multi-trophic micro-ecosystems, a discovery that could have profound implications for our understanding of dispersal mechanisms and the distribution of soil biodiversity.

Plant–animal interactions are not isolated pairwise relationships but are always accompanied by diverse assemblages of microbes. Additional to direct effects of microorganisms on their hosts, recent investigations demonstrated that bacteria associated with plants can modify the behavior of organisms of higher trophic levels. However, in the context of herbivory, functions of non-phytopathogenic bacteria colonizing leaf surfaces remain understudied. This study showed that naturally occurring epiphytic bacteria affect the feeding behavior of a generalist herbivore. Epiphytic bacteria isolated from leaves of Lactuca sativa var. capitata were screened for their potential to influence feeding choices of the slug Arion vulgaris. Cultivated bacteria were inoculated in artificial food substrates or on sterile leaves of gnotobiotic lettuce plants and were offered to slugs in different behavioral bioassays. A large proportion of bacterial strains tested induced behavioral alterations in the feeding choices of slugs. Behavioral responses of slugs were further modified by antibiotic treatment of slugs prior to choice tests indicating that both bacteria associated with plants and animals affect plant–animal interactions. Our results emphasize the important role of bacteria in plant–animal interactions and suggest a prominent role of bacteria in herbivory in natural, horticultural, and agricultural systems.

Summary 1. As a rule, communities consist of few abundant and many rare species, which is reflected in the characteristic shape of species abundance distributions (SADs). The processes that shape these SADs have been a longstanding problem for ecological research. Although many studies found strong negative effects of increasing land-use intensity on diversity, few reports consider land-use effects on SADs. 2. Arthropods (insects and spiders) were sampled on 142 grassland plots in three regions in Germany, which were managed with different modes (mowing, fertilization and/or grazing) and intensities of land use. We analysed the effect of land use on three parameters characterizing the shape of SADs: abundance decay rate (the steepness of the rank abundance curve, represented by the niche-preemption model parameter), dominance (Berger-Parker dominance) and rarity (Fisher’s alpha). Furthermore, we tested the core-satellite hypothesis by comparing the species’ rank within the SAD to their distribution over the land-use gradient. 3. When data on Araneae, Cicadina, Coleoptera, Heteroptera and Orthoptera were combined, abundance decay rate increased with combined land-use intensity (including all modes). Among the single land-use modes, increasing fertilization and grazing intensity increased the decay rate of all taxa, while increasing mowing frequency significantly affected the decay rate only in interaction with fertilization. Results of single taxa differed in their details, but all significant interaction effects included fertilization intensity. Dominance generally increased with increasing fertilization and rarity decreased with increasing grazing or mowing intensity, despite small differences among taxa and regions. The majority of species found on

Because arthropods dominate terrestrial ecosystems in species number and biomass, they can potentially play a major role as environmental indicators in applied ecology and nature conservation. We tested the suitability of arthropods as indicators for particular forest types based on a comprehensive standardized sampling of various taxa by different trap types in 93 mature temperate forest sites in three regions of Germany. We tested whether indicator species (1) can be derived for different forest types across regions, (2) are more widespread and more abundant than non-indicator ones, (3) belong to a particular taxon or trophic guild, and (4) are consistent between regions and years. Among 2041 sampled arthropod species, only four were significant indicator species for the same forest type in all region, and no single taxon or guild performed better than other groups. Indicators were generally more abundant and more widespread than non-indicators, but both abundance and distribution varied widely between species. When the analysis was repeated using data from the next year, indicator values of species significantly correlated between years, but the identity of more than 50 % of significant indicators changed, suggesting high among-year variability. We conclude that overall, arthropods did not turn out to be reliable indicators, at least at the scale of Germany. If anything, arthropod indicator species should be defined at the regional scale. Furthermore, indicators should be selected across taxa and trophic levels. Future evaluation of indicator species among arthropods should be conducted over several years based on standardized sampling protocols to develop a reliable definition of indicator species despite the high fluctuations in abundance of species among arthropods, which might either mask or overestimate the indicator value of particular species.

The importance of spatial scale for β-diversity has been shown in several studies, but it is unclear how spatial diversity patterns correlate among different organismic groups. We studied spatial diversity organization of plants and several trophic guilds of beetles in beech-dominated forests in two regions of Germany to test whether different trophic guilds are organized independently in space. We applied multiplicative diversity partitioning using a nested hierarchical design of four increasingly broader spatial levels (subplot, plot, forest class, region) and tested for correlations among trophic guilds by using Pearson product moment correlations and Mantel-tests. We observed similar general diversity patterns at different trophic guilds showing a high contribution of β-diversity to total γ-diversity and found β-diversity to be higher at different spatial scales and α-diversity to be lower than expected by random distributions of individuals. Results, however, partly depended on the weighting of rare and abundant species. Beta-diversity in our study was caused mainly by species spatial turnover rather than by nestedness. Correlations of α-diversity between trophic guilds were low whereas correlations of β-diversity above subplot level were high. Importantly, more strongly connected trophic guilds revealed not generally stronger relationships than less strongly connected guilds. Three important implications for conservation can be deduced from our results: (1) heterogeneity of beech forests at different spatial scales should be supported in conservation strategies to enhance biodiversity and related functions; (2) the observed high importance of spatial turnover in relation to nestedness implies a concentration of conservation efforts to a large number of not necessarily the richest sites, and (3) recommendation for particular conservation strategies (e.g. selection of priority sites for conservation at regional scale) based on single indicator taxa or functional guild is difficult because of the varied response of the species in our study.

Organic management is one of the most popular strategies to reduce negative environmental impacts of intensive agriculture. However, little is known about benefits for biodiversity and potential worsening of yield under organic grasslands management across different grassland types, i.e. meadow, pasture and mown pasture. Therefore, we studied the diversity of vascular plants and foliage-living arthropods (Coleoptera, Araneae, Heteroptera, Auchenorrhyncha), yield, fodder quality, soil phosphorus concentrations and land-use intensity of organic and conventional grasslands across three study regions in Germany. Furthermore, all variables were related to the time since conversion to organic management in order to assess temporal developments reaching up to 18 years. Arthropod diversity was significantly higher under organic than conventional management, although this was not the case for Araneae, Heteroptera and Auchenorrhyncha when analyzed separately. On the contrary, arthropod abundance, vascular plant diversity and also yield and fodder quality did not considerably differ between organic and conventional grasslands. Analyses did not reveal differences in the effect of organic management among grassland types. None of the recorded abiotic and biotic parameters showed a significant trend with time since transition to organic management, except soil organic phosphorus concentrations which decreased with time. This implies that permanent grasslands respond slower and probably weaker to organic management than crop fields do. However, as land-use intensity and inorganic soil phosphorus concentrations were significantly lower in organic grasslands, overcoming seed and dispersal limitation by re-introducing plant species might be needed to exploit the full ecological potential of organic grassland management. We conclude that although organic management did not automatically increase the diversity of all studied taxa, it is a reasonable and useful way to support agro-biodiversity.

Earthworms dominate invertebrate biomass in many non-acidic terrestrial ecosystems and represent key detritivores. Anecic earthworms live in the soil but primarily feed on soil surface litter. Recent studies indicate that predation of seeds and seedlings may also contribute to earthworm nutrition. However, the biological significance of this behavior is unknown. Here, we present to our knowledge the first photographic and video evidence that the anecic earthworm species Lumbricus terrestris L. attacks living plants and damages leaves still attached to plants. This behavior could be observed repeatedly and in different locations, indicating that it may be a common process. Though based on uncontrolled observations and in situations of low litter availability, this herbivore behavior suggests that the role of earthworms within ecosystems should not be restricted to litter burial and decomposition, but should include aboveground herbivory. Our observations do not allow us to determine with certainty if the earthworms directly fed on fresh plant material or – more likely – transported it to its permanent burrows to facilitate microbial decay. We propose that this behavior and its implications for the composition and functioning of plant communities merits further scientific attention. For instance, given the detrimental effects of some common molluscicides on earthworm performance, assumed mollusk effects on plant growth and seedling survival reported in previous studies could be – at least in part – due to the action of anecic earthworms. Future studies should investigate the biological significance and the context-dependency of this behavior.

Agri-environmental schemes involving organic farming or set-aside management aim at promoting biodiversity and restoring ecosystem functioning in agrarian landscapes. Application of pesticides in these crop fields is strongly regulated facilitating the spread of weeds but also allowing for the establishment of endangered herbs and a variety of animals. Recent studies found gastropods and earthworms to be legitimate dispersers of seeds of wild plants. We assumed that both groups also play a significant role in the spread and establishment of wild plants within crop fields. Therefore, we are conducting a series of experiments in three different study systems on (1) the role of earthworms and gastropods as dispersers of rare herbs and weeds in an organic rye field in Germany, (2) the seed feeding behavior of gastropods of plants sown in fallow ground in Switzerland, and (3) weed dispersal in irrigated rice fields by golden apple snails in the Philippines.

Land-use intensification is a major driver of biodiversity loss(1,2). Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in beta-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (alpha)-diversity(1,3) and neglected biodiversity loss at larger spatial scales. Studies addressing beta-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above-and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in alpha-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on beta-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in beta-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local alpha-diversity in aboveground groups, whereas the alpha-diversity increased in belowground groups. Correlations between the alpha-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Forest canopies are an important part of forest ecosystems and comprise the majority of arthropod diversity. As tree crowns provide various microhabitats, a suitable trapping standard for canopy-dwelling arthropods is required. However, vertical trap position is often not standardized. We developed six vertical placement strategies for flight-interception traps based on information on tree height and crown base obtained from forest inventory data. Strategies differed in how crown base and height of trees were weighted in the calculations of trap height. Forest inventory data from almost 1600 plots in three regions of Germany. Were used to calculate the theoretical suitability of the strategies. The strategy to place traps at the canopy centre was predicted to result in very few trees for which placement would not be possible because trees are too short or crown base is too high, and this strategy by definition resulted in the lowest deviation of trap position from the canopy centre. The strategy Centre was tested in the field by installing three flight-interception traps each in 150 deciduous or coniferous stands of various age structures. On average, 53.8% of the traps were installed at exactly the desired height and for 86.9% of the traps the difference was

Land use intensification in forests is a main driver of global biodiversity loss. Although historical state of land use differs between subtropical and temperate zones, gradients of land-use intensities similarly range from unmanaged to very intensively managed forests. Irrespective of similar land use forces in both climate zones, comparative studies on land use effects are still rare. Such studies are, however, promising in discovering more general impacts and geographical specifics of land use intensification. We studied litter-dwelling invertebrates along a gradient of increasing land use intensity in subtropical forests in Southern Brazil and temperate forests in Central Europe using similar sampling designs. Effects of land use intensity on the entire community were analyzed on the level of orders and feeding guilds. In both climate zones a similar number of individuals were caught when standardizes to 100 pitfall trap days, but taxa richness was higher in the subtropics. Moreover, community composition differed between both climate zones. In both regions, land use intensity did not affect taxa richness, but invertebrate abundance was affected in opposite ways; while increasing land use intensity resulted in a decrease of invertebrate abundance in the subtropics, an increase was observed in the temperate zone and this was mostly consistent regarding different feeding guilds. Management practices should take into account that the effect of land use intensity on biodiversity can differ drastically among climatic regions.

1. In obligate symbioses with horizontal transmission, the population dynamics of the partner organisms are highly interdependent. Host population size limits symbiont number, and distribution of partners is restricted by the presence and thus dispersal abilities of their respective partner. The Crematogaster decamera–Macaranga hypoleuca ant–plant symbiosis is obligate for both partners. Host survival depends on colonisation by its ant partner while foundress queens require hosts for colony establishment. 2. An experimental approach and population genetic analyses were combined to estimate dispersal distances of foundresses in their natural habitat in a Bornean primary rainforest. 3. Colonisation frequency was significantly negatively correlated with distance to potential reproductive colonies. Results were similar for seedlings at natural densities as well as for seedlings brought out in the area experimentally. Population genetic analysis revealed significant population differentiation with an FST of 0.041 among foundresses (n = 157) located at maximum 2280 m apart. In genetic spatial autocorrelation, genotypes of foundresses were significantly more similar than expected at random below 550 m and less similar above 620 m. Direct estimation of dispersal distances by pedigree analysis yielded an average dispersal distance of 468 m (maximum 1103 m). 4. For ants that disperse on the wing, genetic differentiation at such small spatial scales is unusual. The specific nesting requirements of the queens and the necessity for queens to find a host quickly could lead to colonisation of the first suitable seedling encountered, promoting short dispersal distances. Nonetheless, dispersal distances of C. decamera queens may vary with habitat or host spatial distribution.

In beech-dominated forests in Central Europe, many spring geophytes show adaptations to seed dispersal by ants (myrmecochory). Ants, however, can be rare in such moist forests. Motivated by observations of slug feeding on seeds we investigated the seed consumption of two plant species, Anemone nemorosa and Asarum europaeum, by slugs, in a series of experiments. In a seed predation experiment in a beech forest, we found that seed removal was strongly reduced when gastropods were excluded from the seed depots. The contribution of insects, including ants, and rodents to seed removal was relatively less but differed between May and July. In the laboratory, slug species, in particular Arion sp., consumed seeds of both plant species. Slugs either consumed the elaiosomes of seeds or swallowed seeds intact. Swallowed seeds were defecated undamaged and germinated as well as control seeds when buried overwinter, indicating the potential for seed dispersal by slugs. We also recovered seeds of myrmecochores in the faeces of several slugs caught in forests. In a slug release experiment in the forest, slugs moved up to 14.6 m (mean 4.4 m) in 15 h, which is the median gut passage time of seeds based on measurements made in the laboratory. We also found that when slug-defecated seeds were offered to rodents, these were less attractive than control seeds, suggesting that passage through the slug gut reduces seed predation risk. Our results demonstrate that slugs are significant consumers of elaiosomes or entire seeds of ant-dispersed plants and that they can function as seed dispersers of these plants.

Background Recent studies suggested that the earthworm Lumbricus terrestris might act as a seedling predator by ingesting emerging seedlings, and individuals were observed damaging fresh leaves of various plant species in the field. To evaluate the significance of herbivore behavior of L. terrestris for plant and earthworm performance we exposed 23- to 33-days-old seedlings of six plant species to earthworms in two microcosm experiments. Plants belonged to the three functional groups grasses, non-leguminous herbs, and legumes. Leaf damage, leaf mortality, the number of leaves as well as mortality and growth of seedlings were followed over a period of up to 26 days. In a subset of replicates 0.1 g of soil surface litter of each of the six plant species was provided and consumption was estimated regularly to determine potential feeding preferences of earthworms. Results There was no difference in seedling growth, the number of live seedlings and dead leaves between treatments with or without worms. Fresh leaves were damaged eight times during the experiment, most likely by L. terrestris, with two direct observations of earthworms tearing off leaf parts. Another nine leaves were partly pulled into earthworm burrows. Lumbricus terrestris preferred to consume legume litter over litter of the other plant functional groups. Earthworms that consumed litter lost less weight than individuals that were provided with soil and live plants only, indicating that live plants are not a suitable substitute for litter in earthworm nutrition. Conclusion Our results demonstrate that L. terrestris damages live plants; however, this behavior occurs only rarely. Pulling live plants into earthworm burrows might induce microbial decomposition of leaves to make them suitable for later consumption. Herbivory on plants beyond the initial seedling stage may only play a minor role in earthworm nutrition and has limited potential to influence plant growth.

Forest management is known to influence species diversity of various taxa but inconsistent or even contrasting effects are reported for arthropods. Regional differences in management as well as differences in regional species pools might be responsible for these inconsistencies, but, inter-regional replicated studies that account for regional variability are rare. We investigated the effect of forest type on the abundance, diversity, community structure and composition of two important ground-dwelling beetle families, Carabidae and Staphylinidae, in 149 forest stands distributed over three regions in Germany. In particular we focused on recent forestry history, stand age and dominant tree species, in addition to a number of environmental descriptors. Overall management effects on beetle communities were small and mainly mediated by structural habitat parameters such as the cover of forest canopy or the plant diversity on forest stands. The general response of both beetle taxa to forest management was similar in all regions: abundance and species richness of beetles was higher in older than in younger stands and species richness was lower in unmanaged than in managed stands. The abundance ratio of forest species-to-open habitat species differed between regions, but generally increased from young to old stands, from coniferous to deciduous stands and from managed to unmanaged stands. The response of both beetle families to dominant tree species was variable among regions and staphylinid richness varied in the response to recent forestry history. Our results suggest that current forest management practices change the composition of ground-dwelling beetle communities mainly by favoring generalists and open habitat species. To protect important forest beetle communities and thus the ecosystem functions and services provided by them, we suggest to shelter remaining ancient forests and to develop near-to-nature management strategies by prolonging rotation periods and increasing structural diversity of managed forests. Possible geographic variations in the response of beetle communities need to be considered in conservation-orientated forest management strategies. (C) 2014 Elsevier B.V. All rights reserved.

Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.

Endozoochory plays a prominent role for the dispersal of seed plants, and dispersal vectors are well known. However, for taxa such as ferns and bryophytes, endozoochory has only been suggested anecdotally but never tested in controlled experiments. We fed fertile leaflets of three ferns and capsules of four bryophyte species to three slug species. We found that, overall, spores germinated from slug feces in 57.3 % of all 89 fern and in 51.3 % of all 117 bryophyte samples, showing that the spores survived gut passage of slugs. Moreover, the number of samples within which spores successfully germinated did not differ among plant species but varied strongly among slug species. This opens new ecological perspectives suggesting that fern and bryophyte endozoochory by gastropods is a so-far-overlooked mode of dispersal, which might increase local population sizes of these taxa by spore deposition on suitable substrates.

Seed dispersal by ants (myrmecochory) is widespread, and seed adaptations to myrmecochory are common, especially in the form of fatty appendices (elaiosomes). In a recent study, slugs were identified as seed dispersers of myrmecochores in a central European beech forest. Here we used 105 beech forest sites to test whether myrmecochore presence and abundance is related to ant or gastropod abundance and whether experimentally exposed seeds are removed by gastropods. Myrmecochorous plant cover was positively related to gastropod abundance but was negatively related to ant abundance. Gastropods were responsible for most seed removal and elaiosome damage, whereas insects (and rodents) played minor roles. These gastropod effects on seeds were independent of region or forest management. We suggest that terrestrial gastropods can generally act as seed dispersers of myrmecochorous plants and even substitute myrmecochory, especially where ants are absent or uncommon.

Seed dispersal is one of the most important mechanisms shaping biodiversity, and animals are one of the key dispersal vectors. Animal seed dispersal can directly or indirectly be altered by invasive organisms through the establishment of new or the disruption of existing seed dispersal interactions. So far it is known for a few gastropod species that they ingest and defecate viable plant seeds and consequently act as seed dispersers, referred to as gastropodochory. In a multi-species experiment, consisting of five different plant species and four different gastropod species, we tested with a fully crossed design whether gastropodochory is a general mechanism across native gastropod species, and whether it is altered by the invasive alien slug species Arion lusitanicus. Specifically, we hypothesized that a) native gastropod species consume the seeds from all tested plant species in equal numbers (have no preference), b) the voracious invasive alien slug A. lusitanicus – similarly to its herbivore behaviour – consumes a higher amount of seeds than native gastropods, and that c) seed viability is equal among different gastropod species after gut passage. As expected all tested gastropod species consumed all tested plant species. Against our expectation there was a difference in the amount of consumed seeds, with the largest and native mollusk Helix pomatia consuming most seeds, followed by the invasive slug and the other gastropods. Seed damage and germination rates did not differ after gut passage through different native species, but seed damage was significantly higher after gut passage through the invasive slug A. lusitanicus, and their germination rates were significantly reduced.

Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity–multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a land-use intensity (LUI) gradient. The diversity of above- and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the community-level effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Species-specific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities.