Matty P. Berg

Soil invertebrate fauna enhances grassland succession and diversity

One of the most important areas in ecology is to elucidate the factors that drive succession in ecosystems and thus influence the diversity of species in natural vegetation. Significant mechanisms in this process are known to be resource limitation and the effects of aboveground vertebrate herbivores. More recently, symbiotic and pathogenic soil microbes have been shown to exert a profound effect on the composition of vegetation and changes therein. However, the influence of invertebrate soil fauna on succession has so far received little attention. Here we report that invertebrate soil fauna might enhance both secondary succession and local plant species diversity. Soil fauna from a series of secondary grassland succession stages selectively suppress early successional dominant plant species, thereby enhancing the relative abundance of subordinate species and also that of species from later succession stages. Soil fauna from the mid-succession stage had the strongest effect. Our results clearly show that soil fauna strongly affects the composition of natural vegetation and we suggest that this knowledge might improve the restoration and conservation of plant species diversity.

Towards an assessment of multiple ecosystem processes and services via functional traits

Managing ecosystems to ensure the provision of multiple ecosystem services is a key challenge for applied ecology. Functional traits are receiving increasing attention as the main ecological attributes by which different organisms and biological communities influence ecosystem services through their effects on underlying ecosystem processes. Here we synthesize concepts and empirical evidence on linkages between functional traits and ecosystem services across different trophic levels. Most of the 247 studies reviewed considered plants and soil invertebrates, but quantitative trait–service associations have been documented for a range of organisms and ecosystems, illustrating the wide applicability of the trait approach. Within each trophic level, specific processes are affected by a combination of traits while particular key traits are simultaneously involved in the control of multiple processes. These multiple associations between traits and ecosystem processes can help to identify predictable trait–service clusters that depend on several trophic levels, such as clusters of traits of plants and soil organisms that underlie nutrient cycling, herbivory, and fodder and fibre production. We propose that the assessment of trait–service clusters will represent a crucial step in ecosystem service monitoring and in balancing the delivery of multiple, and sometimes conflicting, services in ecosystem management.

Functional traits as indicators of biodiversity response to land use changes across ecosystems and organisms.

Rigorous and widely applicable indicators of biodiversity are needed to monitor the responses of ecosystems to global change and design effective conservation schemes. Among the potential indicators of biodiversity, those based on the functional traits of species and communities are interesting because they can be generalized to similar habitats and can be assessed by relatively rapid field assessment across eco-regions. Functional traits, however, have as yet been rarely considered in current common monitoring schemes. Moreover, standardized procedures of trait measurement and analyses have almost exclusively been developed for plants but different approaches have been used for different groups of organisms. Here we review approaches using functional traits as biodiversity indicators focussing not on plants as usual but particularly on animal groups that are commonly considered in different biodiversity monitoring schemes (benthic invertebrates, collembolans, above ground insects and birds). Further, we introduce a new framework based on functional traits indices and illustrate it using case studies where the traits of these organisms can help monitoring the response of biodiversity to different land use change drivers. We propose and test standard procedures to integrate different components of functional traits into biodiversity monitoring schemes across trophic levels and disciplines. We suggest that the development of indicators using functional traits could complement, rather than replace, the existent biodiversity monitoring. In this way, the comparison of the effect of land use changes on biodiversity is facilitated and is expected to positively influence conservation management practices.

Highly consistent effects of plant litter identity and functional traits on decomposition across a latitudinal gradient.

Plant litter decomposition is a key process in terrestrial carbon cycling, yet the relative importance of various control factors remains ambiguous at a global scale. A full reciprocal litter transplant study with 16 litter species that varied widely in traits and originated from four forest sites covering a large latitudinal gradient (subarctic to tropics) showed a consistent interspecific ranking of decomposition rates. At a global scale, variation in decomposition was driven by a small subset of litter traits (water saturation capacity and concentrations of magnesium and condensed tannins). These consistent findings, that were largely independent of the varying local decomposer communities, suggest that decomposer communities show little specialisation and high metabolic flexibility in processing plant litter, irrespective of litter origin. Our results provide strong support for using trait-based approaches in modelling the global decomposition component of biosphere-atmosphere carbon fluxes.

Parallel ecological networks in ecosystems

In ecosystems, species interact with other species directly and through abiotic factors in multiple ways, often forming complex networks of various types of ecological interaction. Out of this suite of interactions, predator–prey interactions have received most attention. The resulting food webs, however, will always operate simultaneously with networks based on other types of ecological interaction, such as through the activities of ecosystem engineers or mutualistic interactions. Little is known about how to classify, organize and quantify these other ecological networks and their mutual interplay. The aim of this paper is to provide new and testable ideas on how to understand and model ecosystems in which many different types of ecological interaction operate simultaneously. We approach this problem by first identifying six main types of interaction that operate within ecosystems, of which food web interactions are one. Then, we propose that food webs are structured among two main axes of organization: a vertical (classic) axis representing trophic position and a new horizontal ‘ecological stoichiometry’ axis representing decreasing palatability of plant parts and detritus for herbivores and detrivores and slower turnover times. The usefulness of these new ideas is then explored with three very different ecosystems as test cases: temperate intertidal mudflats; temperate short grass prairie; and tropical savannah.

Dynamics and stratification of bacteria and fungi in the organic layers of a scots pine forest soil

Abstract The abundance and micro-stratification of bacteria and fungi inhabiting the organic layers of a Scots pine forest (Pinus sylvestris L.) were investigated. An experiment using stratified litterbags, containing organic material of four degradation stages (fresh litter, litter, fragmented litter and humus) was performed over a period of 2.5 years. Dynamics and stratification of fluorescent stained bacteria and fungi, ratios between bacterial and fungal biomass, and relationships with moisture and temperature are described. Average bacterial counts in litter and fragmented litter were similar, i.e., approximately 5×109 bacteriag–1 (dry weight) organic matter, and significantly exceeded those in humus. The mean bacterial biomass ranged from 0.338 to 0.252mg carbon (C) g–1 (dry weight) organic matter. Lengths of mycelia were significantly below the usually recorded amounts for comparable temperate coniferous forests. The highest average hyphal length, 53mg–1 (dry weight) organic matter, was recorded in litter and decreased significantly with depth. The corresponding mean fungal biomass ranged from 0.050 to 0.009mg Cg–1 (dry weight). The abundance of bacteria and fungi was influenced by water content, that of fungi also by temperature. A litterbag series with freshly fallen litter of standard quality, renewed bimonthly, revealed a clear seasonal pattern with microbial biomass peaks in winter. The mean hyphal length was 104mg–1 (dry weight) and mean number of bacteria, 2.40×109 bacteria g–1 (dry weight). Comparable bacterial and fungal biomass C were found in the freshly fallen litter [0.154 and 0.132mgCg–1 (dry weight) organic material, respectively]. The ratio of bacterial-to-fungal biomass C increased from 1.2 in fresh litter to 28.0 in humus. The results indicate the existence of an environmental stress factor affecting the abundance of fungi in the second phase of decomposition. High atmospheric nitrogen deposition is discussed as a prime factor to explain low fungal biomass and the relatively short lengths of fungal hyphae in some of the forest soil layers under study.

Dynamics and stratification of functional groups of micro- and mesoarthropods in the organic layer of a Scots pine forest

Abstract This paper addresses the abundance, biomass and microstratification of functional groups of micro- and mesoarthropods inhabiting the organic layers of a Scots pine forest (Pinus sylvestris L.). An experiment using stratified litterbags, containing organic material of four degradation stages, i.e., freshly fallen litter, litter, fragmented litter and humus, was performed over a period of 2.5 years. Statistical data analysis revealed that each organic layer had a different, characteristic species composition that changed with time following successive degradation stages. Species of Acari, Araneae and Collembola were assigned to different functional groups based on taxonomy, microstratification, food type or feeding mode. The abundance and biomass carbon of functional groups were dependent on the organic layer and most functional groups showed a particular preference for one of the upper organic layers. Temporal and spatial differences in density and biomass carbon of functional groups could partly be related to fluctuations in the soil climate, although effects of trophic interactions could not be ruled out. A general decline in abundance and biomass, especially in populations of fungal feeders, during the last year of the study could not be explained by a reduction in litterbag volume, changed litter chemistry or soil climate, but was attributed to an indirect effect of a remarkable increase in soil coverage by wavy hair grass, Deschampsia flexuosa (L.). The analysis demonstrated that species diversity, microhabitat specification, soil fauna succession, and degradation stages of organic material are interrelated. The results obtained indicate that both the chemistry of organic matter and decomposition rates have an important effect on trophic relationships and community structure.

Long‐term effects of plant diversity and composition on soil nematode communities in model grasslands

An important component of plant-soil feedbacks is how plant species identity anddiversity influence soil organism communities. We examine the effects of grassland plant species growing alone and together up to a richness of 12 species on nematode diversity and feeding group composition, eight years after the establishment of experimental grassland plots at the BIODEPTH site in northern Sweden. This is a substantially longer time than most other experimental studies of plant effects on soil fauna. We address the hypotheses that (la) higher species or functional diversity of plants increases nematode diversity, as well as influences nematode community composition. Alternatively, (1b) individual plant species traits are most important for nematode diversity and community composition. (2) Plant effects on soil organisms will decrease with increasing number of trophic links between plants and soil fauna. Plant species identity was often more important than plant diversity for nematode community composition, supporting hypothesis 1b. There was a weak positive relation between plant and nematode richness;which could be attributed to the presence of the legume Trifolium pratense, but also to some other plant species, suggesting a selection or sampling effect. Several plant species in different functional groups affected nematode community composition. For example, we found that legumes increased bacterial-feeding nematodes, most notably r-selected Rhabditida, while fungal-feeding nematodes were enhanced by forbs. Other bacterial feeders and obligate root feeders were positively related to grasses. Plant effects were usually stronger on plant-, bacterial- and fungal-feeding nematodes than on omnivores/predators, which supports hypothesis 2. Our study suggests that plant identity has stronger effects than plant diversity on nematode community composition, but when comparing our results with similar previous studies the effects of particular plant species appear to vary. We also found that more productive plant species affected bacterial-feeding nematodes more than fungal feeders. Moreover, we observed stronger effects the fewer the number of trophic links there were between a nematode feeding group and plants. Although we found clear effects of plants on soil nematodes, these were probably not large enough to result in strong and persistent plant-soil-organism-plant feedback loops.

Trait plasticity in species interactions: a driving force of community dynamics

Evolutionary community ecology is an emerging field of study that includes evolutionary principles such as individual trait variation and plasticity of traits to provide a more mechanistic insight as to how species diversity is maintained and community processes are shaped across time and space. In this review we explore phenotypic plasticity in functional traits and its consequences at the community level. We argue that resource requirement and resource uptake are plastic traits that can alter fundamental and realised niches of species in the community if environmental conditions change. We conceptually add to niche models by including phenotypic plasticity in traits involved in resource allocation under stress. Two qualitative predictions that we derive are: (1) plasticity in resource requirement induced by availability of resources enlarges the fundamental niche of species and causes a reduction of vacant niches for other species and (2) plasticity in the proportional resource uptake results in expansion of the realized niche, causing a reduction in the possibility for coexistence with other species. We illustrate these predictions with data on the competitive impact of invasive species. Furthermore, we review the quickly increasing number of empirical studies on evolutionary community ecology and demonstrate the impact of phenotypic plasticity on community composition. Among others, we give examples that show that differences in the level of phenotypic plasticity can disrupt species interactions when environmental conditions change, due to effects on realized niches. Finally, we indicate several promising directions for future phenotypic plasticity research in a community context. We need an integrative, trait-based approach that has its roots in community and evolutionary ecology in order to face fast changing environmental conditions such as global warming and urbanization that pose ecological as well as evolutionary challenges.

Changes in chemical composition of Pinus sylvestris needle litter during decomposition along a European coniferous forest climatic transect.

The objective of this investigation was to assess the changes in chemical composition (lignin, cellulose, hemicelluloses, non-structural compounds, N, and ash) of decomposing litter. Standard Pinus sylvestris needle litter, originating from southern Sweden, was incubated in litterbags at 15 sites selected from the Netherlands to south Spain. The changes in chemical composition of this litter were determined using near infrared reflectance spectroscopy. The hypothesis was that standard (chemically uniform) litter decomposing under a range of climates would show different dynamics of accumulation and loss of C-fractions, N, and ash, relative to mass loss. It was shown that, for a given mass-loss value (10, 20, 30, 40, or 50%), the proportion of lignin, cellulose, hemicelluloses, non-structural compounds, N, and ash in the decomposing pine needles differed between sites. Lignin concentration in the litter residue at 50% mass loss ranged from approximately 26 to 43%, cellulose from 19 to 27%, hemicelluloses from 7 to 11%, non-structural compounds from 19 to 25%, N from 0.7 to 1.3%, and ash content from 1.4 to 10.1%. Lignin concentrations showed the highest range of variation. Lignin concentrations during decomposition were positively related to moisture factors as significant correlations were found with actual evapotranspiration and were improved in multiple regressions by the mean annual precipitation or the water surplus. Cellulose was degraded further at sites with high precipitation whereas hemicellulose degradation was related to temperature. This leads to the conclusion that the remaining organic matter produced by standard litter decomposition within the studied climatic range of variations tended to be more recalcitrant under wet and warm climatic conditions than under cold or dry climate.