Björn Sohlenius

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.

Nematode community structure as indicator of soil functioning in European grassland soils

This investigation analyses whether soil nematode diversity is correlated with soil functional parameters to serve as bioindicator of soil functioning. The analysis focuses on the interrelations of nematofauna, microflora, and soil nitrogen pools. The sites studied represent six major European grassland types: Northern tundra, atlantic heath, wet grassland, seminatural temperate grassland, East European steppe, and mediterranean garigue. Continental and local climate gradients were combined to a wide and continuous range of microclimate conditions. Nematode richness, as indicated by the number of genera, was highest under temperate conditions and declined towards the climatic extremes. Differences in richness affected all nematode feeding types proportionally. Nematode richness was the only parameter among a range of 15 alternatives tested that exhibited consistent correlations with mass and activity parameters of both nematofauna and microflora in the mineral grassland soils (garigue, wet grassland, seminatural grassland, steppe). In the same soils, the nematode Maturity Index was the best indicator of nitrogen status. We conclude that a high nematode richness can generally be seen as a good indicator of an active nematofauna and microflora in mineral grassland soils, and hence as an indicator of the decomposition function. The prospects of exploiting nematode diversity as an indicator of soil functioning are critically discussed.

Effects of organic matter removal on the soil food web: Forestry practices meet ecological theory

Abstract We examine the long-term effects of removing forest harvesting residues (above-ground whole-tree harvesting; WTH) in the soil food web, using data from two different experiments on slash removal and addition in Sweden. Removal of harvesting residues had negative effects on the abundances of animals at higher trophic positions and on more mobile animals. Predators such as gamasid mites, spiders and staphyliniid and cantharid beetles decreased by 30–50% after WTH at both sites. Microbidetritivorous dipterans and fungivorous Collembola decreased at the pine site, but not at the more productive spruce site. Enchytraeids and nematodes were usually not affected by WTH. No differences in fungal and bacterial biomass were found. The removal of organic matter had fairly long-term, of the order of decades, effects on the soil food web. The effects were quantitative rather than qualitative. No functional groups disappeared after WTH. Our findings do not support the hypothesis that soil food webs are entirely donor-controlled. The more pronounced effects of WTH at higher trophic positions indicates that the major interactions in the food webs we have studied are reciprocal, i.e., that consumers to some degree control the density of their resources.

Effects of the nematofauna on microbial energy and matter transformation rates in European grassland soils

The effect of the nematofauna on the microbiology and soil nitrogen status was studied in 6 major European grassland types (Northern tundra (Abisko, Sweden), Atlantic heath (Otterburn, UK), wet grassland (Wageningen, Netherlands), semi-natural temperate grassland (Linden, Germany), East European steppe (Pusztaszer, Hungary) and Mediterranean garigue (Mt. Vermion, Greece). To extend the range of temperature and humidity experienced locally during the investigation period, soil microclimates were manipulated, and at each site 14 plots were established representing selected combinations of 6 temperature and 6 moisture levels. The investigated soils divided into two groups: mineral grassland soils that were precipitation fed (garigue, wet grassland, seminatural grassland, steppe), and wet organic soils that were groundwater fed (heath, tundra). Effects of the nematofauna on the microflora were found in the mineral soils, where correlations among nematode metabolic activity as calculated from a metabolic model, and microbial activity parameters as indicated by Biolog and ergosterol measurements, were significantly positive. Correlations with bacterial activity were stronger and more consistent. Microbial parameters, in turn, were significantly correlated with the size of the soil nitrogen pools NH4, NO3, and Norganic. Furthermore, model results suggested that there were remarkable direct effects of nematodes on soil nitrogen status. Calculated monthly nematode excretion contributed temporarily up to 27% of soluble soil nitrogen, depending on the site and the microclimate. No significant correlation among nematodes and microbial parameters, or nitrogen pools, were found in the wet organic soils. The data show that the nematofauna can under favourable conditions affect soil nitrogen status in mineral grassland soils both directly by excretion of N, and indirectly by regulating microbial activity. This suggests that the differences in nitrogen availability observed in such natural grasslands partly reflect differences in the activity of their indigenous nematofauna.

Influence of clear-cutting and forest age on the nematode fauna in a Swedish pine forest soil

Abstract The abundance and composition of the soil nematode fauna in three seral stages of a pine forest chronosequence located in central Sweden was investigated. The aim of the study was to investigate faunal development during the forest cycle (120–160 years). Three sites, clear-cut 3–12 years before sampling but still without pine trees were compared to three sites with 10–130 year-old trees. Together, these provided sites that could be classified as clear-cut, young stand and mature forest. In most sites without trees and with a poor field layer vegetation, nematode abundance was much lower than in the forested plots. Similar major species/genera of nematodes were found in all age stages of the forest development. The development of the fauna could be related both to climatic shifts and to changes in production of food sources. The ratio of fungal to bacterial feeders and the proportions of two groups of bacterial feeders (Rhabditida BF and Adenophorea BF) varied in distinct ways related to soil layer and seral stage. Thus, in the clear-cut sites the proportion of fungal feeders was quite low. This was also the case with Rhabditida BF. These two groups also contributed a rather low proportion in the litter layer, whereas, their importance successively increased with increasing depth of the soil layers. Proportions of fungal feeding members of the Dorylaimida were highest in the 10–20 year-old stands. The effect of forest trees on nematodes was quite large and the faunal structure in sites with pine trees differed quite markedly from the sites without trees. Apparently, there are certain nematode species whose contribution to the fauna is greatest in the early tree-free stages, others are more abundant in the young forest and still others have their greatest abundance in the mature forest. There are also species whose abundance seems to be unaffected by forest age. Nematode abundance was correlated with the abundance of some other groups of soil organisms.

The geographic distribution of metazoan microfauna on East Antarctic nunataks

The abundance and distribution of nematodes, rotifers and tardigrades in samples from nunataks in continental Antarctica were investigated during four Antarctic expeditions in the austral summers of 1991/92, 1993/94, 1996/97 and 2001/02. Altogether 368 samples were collected from 14 nunataks and one oasis in East Antarctica. Nematodes were found in 35%, rotifers in 67% and tardigrades in 40% of all samples. Fifty-four microfaunal taxa were identified. Of these, 27 were nematodes, 8 tardigrades and 19 rotifers. The size and geographic location of the nunatak and oasis influenced the abundance and taxonomic composition of the microfauna. The highest abundance and diversity of nematodes were found on large nunataks close to the coast. Nematodes were not found on small inland nunataks. Very high population densities of tardigrades and rotifers were found on two small and isolated nunataks. No microfauna was found on the two southernmost nunataks (Okkenhaugrusta and Vardeklettane), or on the smallest one (Utsikta). The Sørensen’s Quotient of Similarity was generally low, especially between faunas on nunataks in different mountain ridges. The results indicate rather limited rates of dispersal and colonization between nunataks.

Distribution patterns of microfauna (nematodes, rotifers and tardigrades) on nunataks in Dronning Maud Land, East Antarctica

In a previous paper the occurrence of nematodes, rotifers and tardigrades in 29 samples from nunataks in Dronning Maud Land, East Antarctica taken 1991/1992 was recorded. In this study, a more thorough description of animal distribution is possible by adding 64 new samples (1993/1994), thus increasing the total number from these studies to 93 soil samples from 11 nunataks. The number of species recorded from the area has now increased from 27 to 34. The populations were patchily distributed and the species composition differed between nunatak areas in Vestfjella and Heimefrontfjella. In most samples only one or a few species of nematodes or tardigrades were found. The number of rotifer species in a particular sample varied from nought to seven. Rotifers occurred in most of the samples (76%) with a maximum density of 750 specimens/gram dry weight. The nematode distribution was more restricted. Nematodes were found in high abundance on only four of the nunataks and occurred in 35% of the samples with a maximum density of 210 specimens/gram dry weight. Tardigrades were found in 39% of the samples with a maximum density of 580 specimens/gram dry weight. Some of the tardigrade species were widely distributed, whereas others were found only on specific nunataks. The distribution pattern of the fauna on two of the nunataks, Basen and Fossilryggen, was studied in greater detail and the results showed little difference in peak abundance and species composition between samplings carried out during two southern summer seasons.

Nematodes, rotifers and tardigrades from nunataks in Dronning Maud Land, East Antarctica

Four species of nematodes, sixteen species of rotifers and seven species of tardigrades were found in samples from nunataks in Vestfjella and Heimefrontfjella in Dronning Maud Land, East Antarctica. The distribution of the fauna was very aggregated and some samples contained no animals. The degree of similarity in species composition between various nunataks was low. The highest abundances and number of species were found on the largest and highest nunataks in the most southern position.

Species diversity and random distribution of microfauna in extremely isolated habitable patches on Antarctic nunataks

Populations of metazoan microfauna (nematodes, rotifers and tardigrades) are patchily distributed on mountain outcrops penetrating the ice sheet (nunataks) in continental Antarctica. The abundance and fauna composition of microscopic animals vary greatly also among samples from similar types of habitats. Occurrence of similar seemingly habitable sites without microfauna and sites with various combinations of animal taxa indicates that stochastic colonization processes as well as local environmental factors and historical factors influence faunal composition in a specific habitable patch. The abundance of nematodes, rotifers and tardigrades in various combinations of co-occurrence was analyzed. One objective was to investigate if biotic interactions structuring these simple communities could be observed. The 368 samples analysed originate from three kinds of habitats, viz. mosses, ornithogenic soils and fellfield soils, obtained from 14 nunataks in Dronning Maud Land, East Antarctica. It is suggested that high population densities of any of the three animal groups, when they were found alone and lower densities, when they coexisted with other taxa could indicate the presence of competition or predation. However, the great variability in microfauna densities for similar habitable patches made it difficult to find significant differences among population densities in samples with varying complexity.

Annual and long-term fluctuations of the nematode fauna in a Swedish Scots pine forest soil

Summary Samples from an old Scots pine forest at Ivantjarnsheden in the middle of Sweden were used to study predictability and patterns of variation of soil nematode communities. There were two annual sampling series (1974–75 and 1977–78) and one long-term series sampled in September ten times over a period of 25 years. The abundance and the composition of the fauna fluctuated rather considerably in both the annual and long-term series. In the annual series abundance and species composition varied in a way which can partly be explained by changes in temperature and moisture. Total nematode abundance was influenced by soil water contents as indicated by co-variations with precipitation. Although the variations in abundance and fauna composition were large no systematic changes could be detected during 25 years. The differences in faunal structure between the two annual series were greater than between the annual and the long-term series. In all series there was a distinct vertical stratification of the fauna. In the superficial moss and litter layers species belonging to Adenophorea ( Plectus ) dominated. In deeper layers members of Rhabditida ( Acrobeloides ) contributed a greater proportion of the fauna. Variations of the annual series indicate that coexistence of different nematode species is facilitated by differences in response to temperature and moisture. The abundance of fungal and bacterial feeders changed in a regular way. During the summer the proportions of fungal and bacterial feeders were almost equal, but during the wet and cold winter the proportion of bacterial feeders increased. Rapidly growing bacterial feeding species belonging to Rhabditida were common in late summer and early autumn, whereas the more slowly growing bacterial feeders belonging to Adenophorea were most abundant during the winter. Although the community fluctuated rather much the average values indicated a rather high degree of predictability and also a high similarity with nematode faunas of other pine forest soils.