▪ Abstract We explore empirical and theoretical evidence for the functional significance of plant-litter diversity and the extraordinary high diversity of decomposer organisms in the process of litter decomposition and the consequences for biogeochemical cycles. Potential mechanisms for the frequently observed litter-diversity effects on mass loss and nitrogen dynamics include fungi-driven nutrient transfer among litter species, inhibition or stimulation of microorganisms by specific litter compounds, and positive feedback of soil fauna due to greater habitat and food diversity. Theory predicts positive effects of microbial diversity that result from functional niche complementarity, but the few existing experiments provide conflicting results. Microbial succession with shifting enzymatic capabilities enhances decomposition, whereas antagonistic interactions among fungi that compete for similar resources slow litter decay. Soil-fauna diversity manipulations indicate that the number of trophic levels, spec...

Biodiversity and Litter Decomposition in Terrestrial Ecosystems

Chemical taxonomy based upon the composition of lipids is widely applied to investigate microbial communities and fatty acids have recently been employed to connect soil microbial and faunal food webs as well as to elucidate functional groups at higher trophic levels. The additional use of compound-specific isotopic analysis of 13C/12C ratios in fatty acids allows assessing specific trophic links and belowground carbon fluxes. In this review systematic patterns and processes underlying variations in the composition of fatty acids and their 13C/12C ratio are described. The emphasis is on biomarker fatty acids, their incorporation and modification, effects of pool size, and analytical methods. Further development of the application of fatty acid profiling to soil ecology should include both advances in experimental research and growth of theory. Accordingly, areas in which future experimentation can lead to progress in soil food web analysis are identified. Overall, combining fatty acid biomarker and their isotopic ratios will allow detailed insight into belowground trophic interactions.

The fat that matters: Soil food web analysis using fatty acids and their carbon stable isotope signature

Global environmental changes are expected to impact the abundance of plants and animals aboveground, but comparably little is known about the responses of belowground organisms. Using meta-analysis, we synthe- sized results from over 75 manipulative experiments in order to test for patterns in the effects of elevated CO2, warming, and altered precipitation on the abundance of soil biota related to taxonomy, body size, feeding habits, eco- system type, local climate, treatment magnitude and duration, and greenhouse CO2 enrichment. We found that the positive effect size of elevated CO2 on the abundance of soil biota diminished with time, whereas the negative effect size of warming and positive effect size of precipi- tation intensified with time. Trophic group, body size, and experimental approaches best explained the responses of soil biota to elevated CO2, whereas local climate and ecosystem type best explained responses to warming and altered precipitation. The abundance of microflora and microfauna, and particularly detritivores, increased with elevated CO2, indicative of microbial C limitation under ambient CO2. However, the effects of CO2 were smaller in field studies than in greenhouse studies and were not sig- nificant for higher trophic levels. Effects of warming did not depend on taxon or body size, but reduced abundances were more likely to occur at the colder and drier sites. Precipitation limited all taxa and trophic groups, particu- larly in forest ecosystems. Our meta-analysis suggests that the responses of soil biota to global change are predictable and unique for each global change factor.

A meta-analysis of responses of soil biota to global change

The large number of animals that coexist in soil without any clear niche differentiation has puzzled biologists for a long time. We investigated stable isotope ratios ( 15 N/ 14 N) in a diverse group of soil microarthropods, oribatid mites, to evaluate trophic niche differentiation. The natural variation of the stable isotopes 15 N/ 14 N was measured in 36 species/taxa from four beech and beech-oak forests. Signatures of d 15 N formed a gradient spanning over 12 d units suggesting that (a) different species occupy different trophic niches and (b) oribatid mites span three to four trophic levels. This study for the first time documented strong trophic niche differentiation in decomposer microarthropods. The results suggest that trophic niche differentiation within taxonomic groups significantly contributes to the high diversity of soil animal taxa. q 2004 Elsevier Ltd. All rights reserved.

Trophic niche differentiation in soil microarthropods (Oribatida, Acari): evidence from stable isotope ratios ( 15 N/ 14 N)

In soil a high number of species co-exist without extensive niche differentiation, which was assigned as ‘the enigma of soil animal species diversity’. In particular, the detritivores are regarded as food generalists. We have investigated nitrogen stable isotope ratios (15N/14N) of a major decomposer group, the Collembola, to evaluate trophic relationship and determine feeding guilds. Additionally, the δ15N values of potential food sources such as mosses, lichens and other plant derived material (bark, nuts, leaves) were analysed. The natural variation in nitrogen isotopes was assessed in 20 Collembola taxa from three deciduous forest stands. The δ15N signature formed a continuum from phycophages/herbivores to primary and secondary decomposers, reflecting a gradual shift from more detrital to more microbial diets. The δ15N gradient spanned over 9 δ units, which implies a wide range in food sources used. Assuming a shift in 15N of about 3 ‰ per trophic level, the results indicate a range of three trophic levels. These variations in 15N/14N ratios suggest that trophic niches of Collembola species differ and this likely contributes to Collembola species diversity.

Feeding guilds in Collembola based on nitrogen stable isotope ratios

The structure of the soil food web in two beech (Fagus sylvatica) forests, the Gottinger Wald and the Solling forest (Northern Germany), was investigated using variations in tissue 15N concentrations of animal species or taxa. The Gottinger Wald is located on a limestone plateau and characterized by mull humus with high macrofauna activity, particularly of Lumbricidae, Diplopoda and Isopoda. In contrast, the Solling forest is located on a sandstone mountain range and characterized by moder humus. The soil fauna of this forest is dominated by mesofauna, particularly by Collembola, Enchytraeidae and Oribatida. In June 1995 soil fauna was sampled using heat extraction. Three soil layers were analysed at each of the sites. 15N/14N ratios of bulk material increased strongly with soil depth in both forests. This also applied to the water-soluble fraction at the Gottinger Wald, but not at the Solling. Generally, the water-soluble fraction was more enriched in 15N than the bulk materials. For most animals studied 15N/14N ratios varied little with soil depth. In both forests soil animals could be classified either as saprophages, including microphytophages, or predators. On average, the δ15N of predatory taxa (Chilopoda, Araneida, Gamasina, Staphylinidae) exceeded that of saprophagous or microphytophagous taxa (Lumbricidae, Isopoda, Diplopoda, Collembola, Oribatida, Enchytraeidae) by 4.4 and 3.9‰ for the Gottinger Wald and the Solling, respectively. We assume that most of the saprophagous or microphytophagous taxa studied consist of primary and secondary decomposers and hypothesize that predators prey more on secondary than primary decomposers. Generally, average δ15N values differed little between saprophagous (Lumbricidae, Diplopoda, Isopoda) and microphytophagous taxa (Collembola, Oribatida). The variations in δ15N values of species within these taxa consistently exceeded the variation between them, indicating that the species of each of these taxa form a continuum from primary to secondary decomposers. Also, variations in δ15N values within predatory taxa in most cases exceeded that between taxa excluding top predators like Sorex. We conclude that using higher taxonomic units in soil food web analysis is problematic and in general not consistent with nature. Higher taxonomic units may only be useful for depicting very general trophic groupings such as predators or microbi-detritivores.

The soil food web of two beech forests (Fagus sylvatica) of contrasting humus type: stable isotope analysis of a macro- and a mesofauna-dominated community.

Marin Falca

Papers

The soil food web of two beech forests (Fagus sylvatica) of contrasting humus type: stable isotope analysis of a macro- and a mesofauna-dominated community.