Volkmar Wolters

HABITAT LOSS, TROPHIC COLLAPSE, AND THE DECLINE OF ECOSYSTEM SERVICES

The provisioning of sustaining goods and services that we obtain from natural ecosystems is a strong economic justification for the conservation of biological diversity. Understanding the relationship between these goods and services and changes in the size, arrangement, and quality of natural habitats is a fundamental challenge of natural resource management. In this paper, we describe a new approach to assessing the implications of habitat loss for loss of ecosystem services by examining how the provision of different ecosystem services is dominated by species from different trophic levels. We then develop a mathematical model that illustrates how declines in habitat quality and quantity lead to sequential losses of trophic diversity. The model suggests that declines in the provisioning of services will initially be slow but will then accelerate as species from higher trophic levels are lost at faster rates. Comparison of these patterns with empirical examples of ecosystem collapse (and assembly) suggest similar patterns occur in natural systems impacted by anthropogenic change. In general, ecosystem goods and services provided by species in the upper trophic levels will be lost before those provided by species lower in the food chain. The decrease in terrestrial food chain length predicted by the model parallels that observed in the oceans following overexploitation. The large area requirements of higher trophic levels make them as susceptible to extinction as they are in marine systems where they are systematically exploited. Whereas the traditional species-area curve suggests that 50% of species are driven extinct by an order-of-magnitude decline in habitat abundance, this magnitude of loss may represent the loss of an entire trophic level and all the ecosystem services performed by the species on this trophic level.

Global decomposition experiment shows soil animal impacts on decomposition are climate-dependent

Climate and litter quality are primary drivers of terrestrial decomposition and, based on evidence from multisite experiments at regional and global scales, are universally factored into global decomposition models. In contrast, soil animals are considered key regulators of decomposition at local scales but their role at larger scales is unresolved. Soil animals are consequently excluded from global models of organic mineralization processes. Incomplete assessment of the roles of soil animals stems from the difficulties of manipulating invertebrate animals experimentally across large geographic gradients. This is compounded by deficient or inconsistent taxonomy. We report a global decomposition experiment to assess the importance of soil animals in C mineralization, in which a common grass litter substrate was exposed to natural decomposition in either control or reduced animal treatments across 30 sites distributed from 43°S to 68°N on six continents. Animals in the mesofaunal size range were recovered from the litter by Tullgren extraction and identified to common specifications, mostly at the ordinal level. The design of the trials enabled faunal contribution to be evaluated against abiotic parameters between sites. Soil animals increase decomposition rates in temperate and wet tropical climates, but have neutral effects where temperature or moisture constrain biological activity. Our findings highlight that faunal influences on decomposition are dependent on prevailing climatic conditions. We conclude that (1) inclusion of soil animals will improve the predictive capabilities of region- or biome-scale decomposition models, (2) soil animal influences on decomposition are important at the regional scale when attempting to predict global change scenarios, and (3) the statistical relationship between decomposition rates and climate, at the global scale, is robust against changes in soil faunal abundance and diversity.

Soil food web properties explain ecosystem services across European land use systems

Intensive land use reduces the diversity and abundance of many soil biota, with consequences for the processes that they govern and the ecosystem services that these processes underpin. Relationships between soil biota and ecosystem processes have mostly been found in laboratory experiments and rarely are found in the field. Here, we quantified, across four countries of contrasting climatic and soil conditions in Europe, how differences in soil food web composition resulting from land use systems (intensive wheat rotation, extensive rotation, and permanent grassland) influence the functioning of soils and the ecosystem services that they deliver. Intensive wheat rotation consistently reduced the biomass of all components of the soil food web across all countries. Soil food web properties strongly and consistently predicted processes of C and N cycling across land use systems and geographic locations, and they were a better predictor of these processes than land use. Processes of carbon loss increased with soil food web properties that correlated with soil C content, such as earthworm biomass and fungal/bacterial energy channel ratio, and were greatest in permanent grassland. In contrast, processes of N cycling were explained by soil food web properties independent of land use, such as arbuscular mycorrhizal fungi and bacterial channel biomass. Our quantification of the contribution of soil organisms to processes of C and N cycling across land use systems and geographic locations shows that soil biota need to be included in C and N cycling models and highlights the need to map and conserve soil biodiversity across the world.

Invertebrate control of soil organic matter stability

Abstract The control of soil organic matter (SOM) stability by soil invertebrates is evaluated in terms of their impact on the inherent recalcitrance, accessibility to microorganisms, and interaction with stabilizing substances of organic compounds. Present knowledge on internal (ingestion and associated transformations) and external (defecation, constructions) control mechanisms of soil invertebrates is also reviewed. Soil animals contribute to the stabilization and destabilization of SOM by simultaneously affecting chemical, physical, and microbial processes over several orders of magnitude. A very important aspect of this is that invertebrates at higher trophic levels create feedback mechanisms that modify the spatio-temporal framework in which the micro-food web affects SOM stability. Quantification of non-trophic and indirect effects is thus essential in order to understand the long-term effects of soil biota on SOM turnover. It is hypothesized that the activities of invertebrates which lead to an increase in SOM stability partly evolved as an adaptation to the need for increasing the suitability of their soil habitat. Several gaps in knowledge are identified: food selection and associated changes in C pools, differential effects on SOM turnover, specific associations with microorganisms, effects on dissolution and desorption reactions, humus-forming and humus-degrading processes in gut and faeces, and the modification of invertebrate effects by environmental variables. Future studies must not be confined merely to a mechanistic analysis of invertebrate control of SOM stability, but also pay considerable attention to the functional and evolutionary aspects of animal diversity in soil. This alone will allow an integration of biological expertise in order to develop new strategies of soil management which can be applied under a variety of environmental conditions.

RELATIONSHIP AMONG THE SPECIES RICHNESS OF DIFFERENT TAXA

Spatially explicit forecasting of changes in species richness is key to designing informative scenarios on the development of diversity on our planet. It might be possible to predict changes in the richness of inadequately investigated groups from that of groups for which enough information is available. Here we evaluate the reliability of this approach by reviewing 237 richness correlations extracted from the recent literature. Of the 43 taxa covered, beetles, vascular plants, butterflies, birds, ants, and mammals (in that order) were the most common ones examined. Forests and grasslands strongly dominated the ecosystem types studied. The variance explanation (R2) could be calculated for 152 cases, but only 53 of these were significant. An average correlation effect size of 0.374 (95% CI = +/- 0.0678) indicates positive but weak correlations between taxa within the very heterogeneous data set; None of the examined explanatory variables (spatial scale, taxonomic distance, trophic position, biome) could account for this heterogeneity. However, studies focusing on 10-km2 grid cells had the highest variance explanation. Moreover, within-phylum between-class comparisons had marginally significantly lower correlations than between-phylum comparisons. And finally, the explanatory power of studies conducted in the tropics was significantly higher than that of studies conducted in temperate regions. It is concluded that the potential of a correlative approach to species richness is strongly diminished by the overall low level of variance explanation. So far, no taxon has proved to be a universal or even particularly good predictor for the richness of other taxa. Some suggestions for future research are inclusion of several taxa in models aiming at regional richness predictions, improvement of knowledge on species correlations in human dominated systems, and a better understanding of mechanisms underlying richness correlations.

Pollinator dispersal in an agricultural matrix: opposing responses of wild bees and hoverflies to landscape structure and distance from main habitat

Semi-natural habitats provide essential resources for pollinators within agricultural landscapes and may help maintain pollination services in agroecosystems. Yet, whether or not pollinators disperse from semi-natural habitat elements into the adjacent agricultural matrix may to a large extent depend on the quality of this matrix and the corresponding pollinator-specific life history traits. To investigate the effects of matrix quality on the distance decay of wild bees and hoverflies, six transects along vegetated field tracks originating at a large semi-natural main habitat and leading into the adjacent agricultural matrix were established in the Wetterau Region, central Hesse, Germany. Species richness of wild bees did not change with distance from the main habitat in landscapes with sufficient grassland cover in the surrounding landscape, but significantly declined when semi-natural grasslands where scarce and isolated in the adjacent agricultural matrix. Abundance of wild bees declined with distance regardless of matrix quality. Species richness of hoverflies did not decline with increasing distance in any landscape. Abundance even increased with distance to the main habitat independently of matrix quality. Thus, our data show that taxa of the pollinator guild may perceive landscapes quite differently. Because of their differing dispersal modes and resource requirements as compared to wild bees, hoverflies may play an important role in maintaining pollination services in agricultural landscapes unsuitable for bee species. Our results highlight the need for considering these taxon-specific differences when predicting the effect of landscape structure on pollinators.

Effects of global changes on above- and belowground biodiversity in terrestrial ecosystems: implications for ecosystem functioning.

bove- and belowground organisms are criticalforthe biogeochemical cycles that sustain the Earth,butthere is limited knowledge on the extent to which the biotabelow ground and the functions they perform are dependenton the biota above ground,and vice versa.Hooper et al.(2000) provide a synthesis ofthe patterns and mechanismslinking above- and belowground biodiversity.The close re-lationship between vegetation change and soil carbon (C)dynamics (Jobbagy and Jackson 2000) suggests that anydisruption ofthe coupling between plants and soil organ-isms as a result ofglobal change may have deleterious con-sequences for functioning ofterrestrial ecosystems.However,most ofthe scientific evidence supporting this hypothesiscomes from correlative approaches.The complexity ofthenumerous interactions between various environmental

PLFA profiles of microbial communities in decomposing conifer litters subject to moisture stress

Abstract The influence of moisture stress on microbial communities in decomposing coniferous litters was investigated using phospholipid fatty acid (PLFA) profiling. Studies were carried out in German and Greek forest plots under contrasting climatic conditions from the late summer to the early winter periods. Litterbags containing spruce (Germany) or pine (Greece) needles were subjected to different irrigation treatments over 4 months. The influences of climate and litter type on microbial community structure were larger than those imposed by irrigation or moisture stress treatments. In the German spruce litter, the PLFA signatures indicated that there was initially a larger bacterial than fungal biomass and both components decreased with time. Concentrations of individual PLFA, proportions of PLFA subgroups and principal component (PC) scores showed that, apart from sample date, mesh size was more important than irrigation treatment in determining microbial community structure; though treatment effects were less apparent in the third (winter) sample. Pine litter in the Greek site, with a Mediterranean climate, had a larger fungal than bacterial biomass. Little effect of treatment on individual PLFA concentrations or PC scores was measured, though both fungal and bacterial communities increased significantly with regular irrigation in the third (winter) sample. Effects of mesh size in the German spruce litter were related to differences in the abundance of microarthropods. This effect was absent from the Greek pine litter where there was a relatively low abundance of fauna. The final spruce litter sample, taken in winter, exhibited very different PC scores from other samples, suggesting marked changes in the microbial community in response to snow melt. Certain long chain fatty acids associated with eukaryotes were only found on this occasion. This study has shown that structure of bacterial communities associated with decomposing conifer litters is highly sensitive to changes in environmental conditions. There was, however, little indication that these differences in biota were functionally important for the initial phases of plant litter decomposition.

Biodiversity of soil animals and its function

Abstract Current knowledge suggests a high redundancy of soil organism communities, i.e. saturation of function at low levels of species richness. This does not imply, however, that research on soil organism biodiversity and its function is irrelevant. First, it is well established that several species of the decomposer community are functionally more important than others. The ‘step’ hypothesis developed in this paper shows that under these conditions random loss of species is much more likely to impact ecosystem processes than expected from the redundancy hypothesis. Second, redundant species may gain functional significance by interacting with functionally important species. Third, the number of ‘important species’ is increased by the multiplicity of functions carried out by soil biota. And finally, alteration in species composition will not be a random process. In fact, functionally important soil biota might be among the first to be affected by large-scale changes in land use. Even if we were to accept that conservation efforts should be confined to the functionally most important species, we would have no idea how to do so. Crossing the ‘spatial barrier’ seems to be the biggest challenge for future investigations on soil biodiversity, because traditional approaches of community ecology will not be sufficient to answer the questions originating from large-scale impoverishment of the soil fauna. Some examples of promising macroecological topics are discussed: (i) the impact of the regional species pool on local species richness, (ii) the relationship between α- and γ-diversity, and (iii) abundance-occupancy relationships. It is argued that research in this direction will be essential for answering the question of how populations and communities must be organised to resist alterations of the soil habitat at the landscape-level.

Intensive agriculture reduces soil biodiversity across Europe

Soil biodiversity plays a key role in regulating the processes that underpin the delivery of ecosystem goods and services in terrestrial ecosystems. Agricultural intensification is known to change the diversity of individual groups of soil biota, but less is known about how intensification affects biodiversity of the soil food web as a whole, and whether or not these effects may be generalized across regions. We examined biodiversity in soil food webs from grasslands, extensive, and intensive rotations in four agricultural regions across Europe: in Sweden, the UK, the Czech Republic and Greece. Effects of land-use intensity were quantified based on structure and diversity among functional groups in the soil food web, as well as on community-weighted mean body mass of soil fauna. We also elucidate land-use intensity effects on diversity of taxonomic units within taxonomic groups of soil fauna. We found that between regions soil food web diversity measures were variable, but that increasing land-use intensity caused highly consistent responses. In particular, land-use intensification reduced the complexity in the soil food webs, as well as the community-weighted mean body mass of soil fauna. In all regions across Europe, species richness of earthworms, Collembolans, and oribatid mites was negatively affected by increased land-use intensity. The taxonomic distinctness, which is a measure of taxonomic relatedness of species in a community that is independent of species richness, was also reduced by land-use intensification. We conclude that intensive agriculture reduces soil biodiversity, making soil food webs less diverse and composed of smaller bodied organisms. Land-use intensification results in fewer functional groups of soil biota with fewer and taxonomically more closely related species. We discuss how these changes in soil biodiversity due to land-use intensification may threaten the functioning of soil in agricultural production systems.