Graham H. R. Osler

TOWARD A COMPLETE SOIL C AND N CYCLE: INCORPORATING THE SOIL FAUNA

Increasing pressures on ecosystems through global climate and other land-use changes require predictive models of their consequences for vital processes such as soil carbon and nitrogen cycling. These environmental changes will undoubtedly affect soil fauna. There is sufficient evidence that soil fauna have significant effects on all of the pools and fluxes in these cycles, and soil fauna mineralize more N than microbes in some habitats. It is therefore essential that their role in the C and N cycle be understood. Here we introduce a new framework that attempts to reconcile our current understanding of the role of soil fauna within the C and N cycle with biogeochemical models and soil food web models. Using a simple stoichiometric approach to integrate our understanding of N mineralization and immobilization with the C:N ratio of substrates and faunal life history characteristics, as used in food web studies, we consider two mechanisms through which soil fauna can directly affect N cycling. First, fauna that are efficient assimilators of C and that have prey with similar C:N ratios as themselves, are likely to contribute directly to the mineral N pool. Second, fauna that are inefficient assimilators of C and that have prey with higher C:N ratios than themselves are likely to contribute most to the dissolved organic matter (DOM) pool. Different groups of fauna are likely to contribute to these two pathways. Protists and bacteria-feeding nematodes are more likely to be important for N mineralization through grazing on microbial biomass, while the effects of enchytraeids and fungal-feeding microarthropods are most likely to be important for DOM production. The model is consistent with experimental evidence and, despite its simplicity, provides a new framework in which the effects of soil fauna on pools and fluxes can be understood. Further, the model highlights our gaps in knowledge, not only for effects of soil fauna on processes, but also for understanding of the soil C and N cycle in general.

The Enigma of Soil Animal Species Diversity Revisited: The Role of Small-Scale Heterogeneity

Background “The enigma of soil animal species diversity” was the title of a popular article by J. M. Anderson published in 1975. In that paper, Anderson provided insights on the great richness of species found in soils, but emphasized that the mechanisms contributing to the high species richness belowground were largely unknown. Yet, exploration of the mechanisms driving species richness has focused, almost exclusively, on above-ground plant and animal communities, and nearly 35 years later we have several new hypotheses but are not much closer to revealing why soils are so rich in species. One persistent but untested hypothesis is that species richness is promoted by small-scale environmental heterogeneity. Methodology/Principal Findings To test this hypothesis we manipulated small-scale heterogeneity in soil properties in a one-year field experiment and investigated the impacts on the richness of soil fauna and evenness of the microbial communities. We found that heterogeneity substantially increased the species richness of oribatid mites, collembolans and nematodes, whereas heterogeneity had no direct influence on the evenness of either the fungal, bacterial or archaeal communities or on species richness of the large and mobile mesostigmatid mites. These results suggest that the heterogeneity-species richness relationship is scale dependent. Conclusions Our results provide direct evidence for the hypothesis that small-scale heterogeneity in soils increase species richness of intermediate-sized soil fauna. The concordance of mechanisms between above and belowground communities suggests that the relationship between environmental heterogeneity and species richness may be a general property of ecological communities.

Detritivores as indicators of landscape stress and soil degradation

Detritivores are small- to medium-sized invertebrates that comminute and break down organic materials such as leaves, twigs and roots, especially within or upon the soil surface, or nearby. Detritivores constitute the majority of the invertebrate biomass pyramid in most environments and provide a key role in organic matter turnover; they also provide alternative food for polyphagous predators that can be active in pest control on crops. Many arthropod taxa are detritivores in soil and litter layers. Here, we focus on the bioindicator potential of three key detritivore groups: slaters, millipedes and oribatid mites. There are possibly 300 species of slaters (terrestrial isopods or Oniscidea) in Australia with 13 of these being introduced, mostly from north-western Europe. These non-native species are the dominant species in disturbed environments such as intensively managed forests and agricultural fields. Slaters are promising indicators of landscape disturbance, soil contamination and tillage. Millipedes are potentially important indicators of stress in agricultural landscapes, given their sensitivity to litter and soil moisture gradients and to physical and chemical perturbations. However, because there is a close association between the millipede fauna and moist plant communities in Australia, they are generally absent from drier landscapes and, therefore, their use as bioindicators in agricultural environments here is problematic. An exception to this association is the increasingly ubiquitous introduced Black Portuguese millipede. This species is tolerant of much drier conditions than most natives, and is likely to change the nature of nutrient cycling processes in pastures and native grasslands in much of southern Australia. Oribatid mites are present in all Australian terrestrial ecosystems. The few studies that have examined their response to disturbance and land use in Australia are consistent with the body of work conducted outside Australia. This consistent response means that the oribatids may be developed as indicators in agricultural, pasture and forested environments. However, the paucity of information on oribatids over appropriate spatial scales in Australia makes the use of this group extremely difficult at this time.

Association of vegetation and soil mite assemblages with isolated Scots pine trees on a Scottish wet heath

Isolated trees may significantly enhance biodiversity at the landscape level. However, our understanding of their impacts is still poor, particularly in environments with high soil moisture where research on this topic has been comparatively limited. We examined understorey vegetation and soil oribatid mite assemblages under live and dead Scots pine trees and in open treeless areas, all within the same Scottish upland wet heath system, to determine whether isolated live trees affected the understorey and mite components of the ecosystem, and whether these effects occurred in parallel. We also explored whether these responses might result from tree-driven reductions in soil moisture content. Live trees reduced soil moisture (relative to wet heath and beneath dead trees) and appeared to change vegetation from wet heath to dry heath type communities. These effects were strongly related to tree trunk diameter (tree size). No major effects of dead trees on understorey vegetation or soil moisture were apparent. Higher mite species abundance and richness were found under live trees than in treeless open heath. Although mite abundances were lower under dead trees than live trees, richness remained similar, thus different factors seem to be regulating mite abundance and community composition. These findings indicate that landscape-level biodiversity responses to environmental change such as habitat fragmentation cannot be predicted from vegetation patterns alone, and that even in heavily fragmented landscapes comparatively small patches such as isolated individual trees can enhance biodiversity.