Biol Fertil Soils (1998) 26:5057

Springer-Verlag 1998

O R I G I N A L PA P E R

J. K. Nieminen H. Seta la

Enclosing decomposer food web: implications for community structure and function

Received: 5 February 1997

Abstract We designed a field experiment to evaluate how restriction of soil faunal movements affects decomposer community structure, food web architecture, and decomposition of organic matter. Intact soil cores (3 cm thick, diameter 16 cm) were placed either in open (mesh size 1 mm, allowing all meso- and microfauna to move through) or closed (27 lm, animal movement prevented except for the smallest microfauna) mesh bags in early May. Before being buried in the forest floor of a mixed spruce stand, hay litter was placed in the mesh bags in separate litter bags. The samplings took place 2 and 6 months after establishing the experiment. Additional field samples were taken from the adjacent soil to determine possible side effects of the mesh-bags. Physicochemical conditions, decomposition rate of hay litter, and total respiration of soil cores were identical in the two bag treatments. Enchytraeids increased significantly in the closed treatment, while macrofauna, such Coleoptera larvae and dipteran larvae, went close to extinction in the closed bags. The elevated enchytraeid number is in accordance with the findings of closed microcosm studies, and is best explained by reduced predation by macrofauna. Although a set of 14 mite taxa was found to distinctively reflect the degree of isolation, neither the total number of individuals nor the number of microarthropod taxa differed between the bag treatments, or between the bags and the field samples. It is concluded that in the time-span of one growing season, reduction in the spatial scale does not necessarily reduce the diversity of fauna but can significantly change the decomposer food-web architecture. Key words Decomposer food web Dispersion Isolation Litter-bags Spatial scale

Introduction
The concepts of scale and isolation broadly reviewed by Levin (1992) have recently become a general topic of contemporary ecology. In soil ecology constriction of spatiotemporal dimensions has been a widely applied method to investigate, for example, the relationship between decomposer community structure and its function (Beyers and Odum 1993; Verhoef 1996). The litter bag method representing a field exclosure/enclosure technique (Swift et al. 1979), and laboratory microcosm techniques (Seta la et al. 1988; Taylor et al. 1989; Seta la and Huhta 1991; Tanaka et al. 1994) serve as examples of such an approach. While the litter bag and microcosm experiments are easily performed, extrapolation of results to natural ecosystems may not be straightforward; apart from the reduced spatial scale, the systems under study become more or less isolated from their environment. Consequently, the ability of organisms to freely exploit their natural habitat is limited to the scale of the experimental unit, thus potentially affecting the extrapolation of the results to the field (Teuben and Verhoef 1992). At least some soil fauna evidently move on a scale that is larger than the size of an enclosure. This was demon gvar and Kjndal (1981) in a succession strated by Ha study in which the rapid colonisation of litter bags by mites and collembolans reflected the motility of the microarthropod community. Under controlled laboratory conditions a fungivorous collembolan was found to disperse up to 10 cm per day (Bengtsson et al. 1994). Thus, preventing soil fauna from dispersing in their habitat may cause unnatural consequences in spatially restricted systems, thus reducing the reliability of microcosm and litter bag experiments in simulating natural phenomena in soils (Beyers and Odum 1993). Although the importance of scale and isolation in field and laboratory studies needs direct testing (Ruth et al. 1994), few attempts have been made to cope with the problems involved in laboratory techniques (but see Elliott et al. 1987; Huhta and Seta la 1990; Seta la et al. 1997). Here we report the results of a field experiment designed

J. K. Nieminen H. Seta la (u) University of Jyva skyla , Department of Biological and Environmental Science, P. O. Box 35, SF-40351 Jyva skyla , Finland Fax: 358 14 60 23 21; e-mail: hsetala@jyu.fi

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to evaluate whether restriction of faunal dispersing affects (1) the structure of the soil animal community, and (2) the decomposition rate of organic material. The study was designed to take place during the biologically most active period of the year, from early May to late October, to allow several generations of micro- and mesofauna to turn over. A scale consistent with the latest laboratory studies (Seta la and Huhta 1991; Laakso et al. 1995; Seta la 1995) was chosen; intact soil cores with a diameter of 16 cm were placed in either closed (mimicking closed laboratory enclosures, allowing only some microfauna to move through) or open mesh bags (practically all animal movement possible; control for the closed treatment). Non-manipulated field data were used to control the effects brought about by the mesh bags themselves. We hypothesised that (1) isolation would reduce the diversity of soil fauna and simplify the food web configuration of the detrital community in the closed mesh bags, while communities in the open bags would resemble natural ones, and (2) as an outcome of biological processes decomposition rate of organic material would consequently be different.

formed as above, with the exception that five field samples were taken and included in the respiration measurement. SPSS for Windows (version 6.1.) was used for the data analysis, including analyses of variance (Anova), t-tests, and discriminant analysis. Canonical correspondence analysis was performed using the CANOCO program (Braak 1988). Square-root transformed variables were used in the analyses; original data are presented in the Tables and Appendix.

Results
Physicochemical parameters The physicochemical conditions in the soil cores did not differ between the two types of mesh bags. The moisture content of the litter and the soil varied between 74.3 and 78.7% (of fresh mass) throughout the study, except for July when the moisture content in the surrounding soil (55.8%) was considerably drier than the soil material in the mesh bags. The pH of the soil varied between 4.5 and 4.9 during the study. Decomposition of hay litter

Materials and methods

For the experiment, 40 soil cores (3 cm thick, 16 cm, F and H layer material) were collected from a 2 m2 area in a mixed spruce stand in Konnevesi, central Finland. The uppermost loose litter and living plants were removed, and replaced with a litter bag (mesh size of 1 mm) with 2 g (dry mass) of hay (Festuca ) litter. The soil cores with the hay bags were then packed in either open (mesh size 1 mm, animal movement allowed, n = 20) or closed mesh bags (mesh size 27 lm, dispersing of fauna other than microfauna prevented, n = 20). The experiment was established on 4 May 1995. Forty new holes with the same measurements as the bag size were dug in ten adjacent rows with a minimum distance of 30 cm between the holes. The experimental units were placed into the holes so that in each row every second unit had an open bag. Finally, the meshbags were covered with a blanket of litter. Field samples (from now on referred to as initial samples) were taken for determining the initial decomposer community structure. After 2 months (3 July 1995) one unit of each treatment was sampled at random in each row to produce a set of ten bags per treatment. Additionally, four 25-cm2 soil samples (hence forth referred to as field samples) were taken from the soil between the mesh bags for detecting the possible side effects of the mesh-bag method. Having been weighed for fresh mass the soil cores, together with hay bags, were closed in airtight Plexi-glass chambers connected to an infrared carbon-analyser. After a settling period of about 24 hours the evolution of carbon dioxide was measured at 3 min intervals for another 24 hours. After the CO2 analysis the soil cores were analysed for pH(H2O), water content (48 h at 70 C), mass loss of the hay litter, and animal populations. To quantify the fauna in the mesh-bags the samples were divided into four parts as follows. Enchytraeids were extracted from 25-cm2 soil-samples using Baerman funnels. The same funnels were used to extract nematodes, tardigrades and rotifers from 5 g (fresh mass) of 10-cm2 samples for 24 hours. An automatic high gradient extractor (Macfadyen 1961) was used to extract microarthropods from 25-cm2 samples. Mites and springtails were identified and counted. The remaining 141-cm2 samples were used to extract larger animals using a modified Tullgren apparatus (Huhta 1972). The second sampling took place on 23 October 1995, 25 weeks after the start of the experiment. Sampling and analyses were per-

There was a rapid loss in hay mass during the experiment with no differences between the open and closed treatment. After 2 months only 0.670.02 g (dry mass; meanSEM and 0.690.01 g) of the initial 2 g were left in open and closed bags, respectively. The average mass remaining at the end of the experiment was 0.440.03 g in the open treatment, and 0.390.03 in the closed treatment. Carbon dioxide evolution was slightly but not significantly (Anova, P<0.17) higher in the open than in the closed treatment. In October, the field samples respired significantly more (104.516.3 lg CO2 h1 g1 fresh mass, meanSE) than the soil cores in the closed bags (74.111.3). Decomposer community structure Microfauna Mean population numbers of nematodes, rotifers and tardigrades are summarised in Table 1. Altogether, 15 genera of Nematoda were detected, 3 of which were fungal feeders (Aphelenchoides spp., Tylenchus spp., Tylencholaimus spp.), 10 bacterial feeders (Acrobeloides spp. and Rhabditis spp. comprising more than 90% of this group), one predator (Prionchulus sp.), and one omnivore (Dorylaimus sp.). The closed bags were evidently intolerable for rotifers, whereas tardigrades were equally abundant in every set of samples. Total numbers of nematodes did not differ between the bag treatments, but in October there were significantly more nematodes in the mesh bags than in the surrounding soil (Students t-test, t = 3.38, P & 0.003). The genus level nematode data was subjected to the canonical correspondence analysis. The first two eigen-

52 Table 1 Total numbers of Nematoda, Tardigrada and Rotifera (103 individuals/m2), and mean incidences (SEM) of Acari taxa in open May Field, n = 5 Nematoda Tardigrada Rotifera Acari 1717423 30.06.8 11.32.5 0.750.01 July Field, n = 5 0.660.02 Open, n = 10 3249793 29.06.4 3.91.6 0.290.01 * Closed, n = 10 3255898 28.87.8 1.91.4 0.280.01 and closed mesh bags and in field samples (* significant difference from the field mean within a sampling at a = 0.05 risk level) October Field, n = 5 615150 20.419.4 5.87.2 0.510.01 Open, n = 10 1519206 * 23.822.1 6.707.0 0.400.01 Closed, n = 10 2411557 * 31.545.4 0.983.1 0.430.011 *

values, k1 = 0.10 and k2 = 0.06 (P<0.01), indicate that the nematodes were not randomly distributed among the samples. Most of the variation existed between the field and bag samples (the ordination diagram is not presented). Enchytraeidae Enchytraeid populations almost exclusively composed of Cognettia sphagnetorum Vejd grew at greatest density in closed mesh-bags. In July the difference between the treatments was small (about 15 000 individuals/m2 in each treatment), but at the end of the experiment the closed bags harboured about 4 times more enchytraeids (35 6006 425 individuals m2; meanSE) than the open bags (8 2001 424), or the field samples (9 2001 431) (Anova, P<0.001). Enchytraeid densities tended to be negatively correlated with the abundances of the predatory mite Pergamasus (r = 0.257, P = 0.072), other Gamasina (r = 0.252, P = 0.077), Coleoptera larvae (r = 0.261, P = 0.084) and Diptera (r = 0.255, P = 0.091). When enchytraeid numbers are plotted against the sum of Coleoptera larvae and Araneae (later on referred to as macropredators), it can be seen that the more macropredators were involved the fewer enchytraeids were present (Fig. 1). Microarthropoda A total of 40 microarthropod taxa were determined (see Appendix). In July a pronounced bag effect occurred: more taxa were found in the field than in the mesh bags. In October, however, this effect was insignificant, but the number of taxa among the mesh bags varied substantially more than the field samples from the adjacent soil. Incidences of all microarthropods at the final sampling are given in Appendix, and mean incidences of Acari in Table 1. Incidence (I ) of a taxon j is defined as a proportion of cases with j within a sample: Ij nj =N where nj is the number of cases with taxon j, and N is the sample size. Mite taxa were present in significantly fewer cases in the bags than in the field samples, whereas the mean incidences of Collembola did not differ significantly between the treatments (Anova, df = 6, F = 0.349, P>0.9).

Fig. 1 Regression between Macropredators (Coleoptera, Arancae, p 2 ) when present Chilopoda) and Enchytraeids (both individuals = m p in the samples. The model is Enchytraeidae = (168.7522.51) p (2.991.43) Macropredators, and the significance of the constant and regression coefficient is P<0.0005 and P<0.045, respectively

Total numbers of microarthropods did not differ between the treatments, but as with the number of the taxa, the variability of the population densities in mesh bags was conspicuous. Collembolans seemed to have been rather insensitive to the treatments, although Folsomia spp., Isotomiella minor and Onychiuridae were generally more abundant in the bags than outside in the field in October. Oribatida reflected exactly the same pattern as total numbers of mites, while Prostigmata and Astigmata did not vary after a drop in population numbers between May and July. The predatory Uropodina increased in numbers in the closed bags from July to October. At the same time Gamasina grew more abundant in the open bags, but the treatments did not differ significantly due to one outlier in October. To evaluate the ability of a systems openness to classify the mite data, discriminant analysis was applied. After discarding the variables with high Wilks lambda (i. e. those with a poor effect on classification), ten oribatid

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closed bags. Paleacaroidea was scarce in all treatments. Of the predators Pergamasus was observed in May and October only, while Eviphis was almost absent in October. The open mesh-bags and the field samples had significantly higher mean numbers of predators summarised as unidentified Gamasina than the closed bags had. Macrofauna Larvae of Diptera and Coleoptera were the two dominant groups of larger animals. Others [Araneae, Coleoptera adults, Chilopoda, earthworms (Dendrobaena octaedra ) and Hymenoptera] were only occasionally encountered. The effect of mesh size on the numbers of macrofauna was significant, whereas the interaction between mesh size and sampling time (JulyOctober) was not. Macrofauna were nearly absent in closed bags (Fig. 3). The drop from initial numbers was marked even at the first sampling in July.

Fig. 2 Distribution of mites (discriminant scores of 14 taxa) in a biplot with respect to three habitats of varying openness (Group 0 field samples, Group 1 mesh bags with 1-mm mesh size, Group 2 mesh bags with 27-lm openings)

groups, three predatory groups, and Prostigmata + Astigmata remained in the final analysis. Eigenvalues and significances of the two discriminant functions formed were k1 = 2,86, P<0.0005, and k2 = 0.74, P = 0.025, respectively. The discriminant scores for the 54 cases are plotted in Fig. 2. Over 90% of the cases became correctly classified in this analysis. The two bag treatments and the field values appeared to form three distinguishable groups; the field samples differed more from the two bag treatments than the latter did from each other. Of the 14 taxa that best characterised the mite data with respect to the degree of isolation, Oppioidea and Tectocepheus, the juveniles in particular, and Prostigmata + Astigmata were clearly the most common and abundant groups. In contrast to their numbers in the field samples, their numbers increased in the bags from July to October. The majority of the relatively rare oribatid genera Camisia, Carabodes, Eupelops and Scheloribates were found in field samples and in open bags. Steganacarus differed from these in being absent in open but present in

Discussion
We shall examine the results of this study from three standpoints, starting with an analysis of the impact of enclosement on the faunal community at species/genus level. Since microarthropods comprised the highest diversity of the encountered taxa, the community analysis is largely based on this group of fauna. The impact of enclosures on the decomposer food web configuration is investigated next, and finally the decomposition rate of organic material is examined in relation to the mesh size of the bag. Because the physicochemical conditions in the bags did not differ between the open and closed treatment, we conclude that any effects concerning the decomposer community structure or function relating to the mesh size of the bag are to be taken for direct effects or restricting animal movements.

Fig. 3 Total numbers (individuals g1 dry mass) of macrofauna in the open and closed mesh bags and in the field, presented as boxplots [minimum, Q 1, median, Q 3, maximum; o and *, followed by a case number, stand for outliers and extremes, respectively; P <0.015 for the effect of the size of the mesh (t -test, n = 20) (Closed mesh bags with 27-lm opening, Open 1-mm opening, VII July, X October, N number of replicates)

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Community composition of microarthropods The initial numbers of microarthropods in the field samples were comparable with the values given by Huhta et al. (1986) for similar soils, whereas the summer densities in the mesh bags and in the field samples were much lower, most likely due to the exceptionally long dry periods during the summer 1995. The initially reduced species richness of microarthropods, the greater variation of the quality and quantity of this fauna, and the significantly lower incidence of species in the bags than in the field imply that chance played a bigger role in the former habitat. In other words, despite the similarity in the mean number of species, the probability of encountering every one species in a sample was lower in the bag-treatments than in the field. This particularly holds for the closed mesh bags, in which the species assemblage must depend on the initial species pool because the entry of animals was prevented during the experiment. Since the development of a community can be remarkably sensitive to the initial conditions of the system (Drake 1991; Wilson 1992), the great variance in the community structure in the closed bags is not surprising. This cannot, however, explain the reduced incidences of taxa in the open bags. It seems that even the open bags restricted free dispersal of some mesofauna, or that the material in the bags, regardless of the size of the mesh, lost its attractiveness to the fauna. Whether this possible restriction is physical, biological, or related to possible harmful effects for the fauna while establishing the experiment, remains open. Beyers and Odum (1993) hypothesised the commonly occurring diversity reduction in closed experimental systems to be due to loss of space, loss of species with a large area demand, and loss of available niches. Wright and Coleman (1993) showed that the diversity of nematode communities tended to decline in isolation, while the size (volume) of the isolate had little influence on the nematode community composition. Our results partly support these hypotheses and experimental evidence that community composition of soil fauna some micorarthropods and nematodes in particular can be sensitive to isolation. However, that only 14 microarthropod taxa out of 40 proved sensitive to the treatment setting indicates that a relatively small area (200 cm2) is large enough to maintain a diverse community of microarthropods, at least for a period of one growing season. Therefore we conclude that the worry of losing species diversity in experimental soil ecology is a smaller concern than that of influencing other abio-biotic properties of the manipulated system. Effects of enclosure on trophic structure of the decomposer food web Although categorising decomposer food webs into clearly distinguishable trophic components is difficult (Moore et al. 1988; Beare et al. 1995), some generalisations concerning the enclosure effects on trophic organisation of the

fauna can nevertheless be made. The strong reduction in large-sized taxa, such as detritivorous dipterans, and particularly predaceous spiders and Coleoptera, in the closed mesh bags is clearly the most prominent effect brought about by the experimentation. As implied by the regression analysis, the reduction of these macropredators and detritivores was an important factor, although not the only one, in explaining the increased numbers of enchytraeids in the closed bags. Indeed, besides macropredators, larger Gamasina, which are known to prey upon enchytraeids (Walter et al. 1988), were also more abundant in the open bags than in the closed bags. Based on the abundance of these large and medium-sized carnivores, it is evident that the intensity of predation upon enchytraeid worms was less pronounced in the closed bags than in the open ones, or in the field. Since the abiotic conditions between the open and the closed bags were identical, it is reasonable to conclude that the massive increase of enchytraeid populations in the closed bags must, at least partly, result from biotic interactions. On the other hand, the reduction in numbers of highly mobile predators in the closed systems evidently resulted from the radically restricted spatial scale in the bags. Moreover, removal of most of the original litter layer from the soil core while establishing the experiment may also have reduced the numbers of large detritivores and predators typically existing in the uppermost layers of the soil (Wallwork 1970). It has been argued that some interspecific interactions become less important in closed systems than in open systems, whereas other species interactions may become overly emphasised in enclosures (Elliott et al. 1987; Beyers and Odum 1993; Lawton 1995). Results of the present study and those of Kajak and Jakubczyk (1977) strengthen the concern that unlimited growth of some species, such as Cognettia sphagnetorum, may have an unproportionally important role in directing the community and ecosystem dynamics in closed systems, and thus largely account for results obtained, for example, in laboratory experiments. It is known that enchytraeids can ecologically engineer their milieu by processing soil organic matter and changing the pore size distribution in the soil (Didden 1990). Therefore the capability of enchytraeids to indirectly influence the development of other members of the decomposer community should not be underestimated. Food web architecture and decomposition of organic matter Invariability of the process variables (community respiration, and decomposition rate of hay litter) in terms of isolation indicates that the decomposition processes, though by ecologically different paths, resulted in the same outcome in both treatments. Because of the large contribution of microbes to total respiratory metabolism in forest soils (Persson et al. 1980), the evolution of carbon dioxide in our soil samples largely reflects microbial activity. Therefore, the similar decomposer activity between the treatments is not surprising; obviously organisms low in

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the detrital food web are not sensitive to manipulation of this kind. In other words, in parallel to the results by Seta la et al. (1997), predatory induced changes at the top of the food web did not cascade down to the basal resources as is speculated to be possible in food webs rich in species (Pimm et al. 1991).

References
Anderson JM (1977) The organization of soil animal communities. In: Lohm U, Persson T (eds) Soil organisms as components of ecosystems. Proceedings of the VI international soil zoology colloquium (Ecological bulleting, vol 25) Stockholm pp 1523 Anderson JM (1988) Spatiotemporal effects of invertebrates on soil processes. Biol Fertil Soils 6:216227 Beare MH, Coleman DC, Crossley DA, Hendrix PF, Odum EP (1995) A hierarchical approach to evaluating the significance of soil biodiversity to biogeochemical cycling. Plant Soil 170:522 Bengtsson G, Hedlund K, Rundgren S (1994) Food- and density-dependent dispersal: evidence from a soil collembolan. J Anim Ecol 63:513520 Bengtsson J, Seta la H, Zheng D (1995) Food webs and nutrient cycling in soils: interactions and positive feedbacks. In: Polis GA, Winemiller KO (eds) Food webs: integration of patterns and dynamics. Chapman and Hall, New York, pp 3038 Beyers RJ, Odum HT (1993) Ecological microcosms. Springer, Berlin Heidelberg New York Braak CJF ter (1988) CANOCO an extension of DECORANA to analyse species-environment relationships. Vegetatio 75:159160 Didden WAM (1990) Involvement of Enchytraeidae (Oligochaeta) in soil structure evolution in agricultural fields. Biol Fertil Soils 9:152158 Drake JA (1991) Community-assembly mechanisms and the structure of an experimental species enemble. Am Nat 137:126 Elliott ET, Hunt HW, Walter DE, Moore JC (1987) Microcosms, mesocosms and ecosystems: linking the laboratory to the field. In: Megusar F, Gantar M (eds) Perspectives in microbial ecology. Proceedings of the IV international symposium on microbial ecology. Ljubljana, Yugoslavia, pp 472480 Gascon C, Travis J (1992) Does the spatial scale of experimentation matter? A test with tadpoles and dragonflies. Ecology 73:2237 2243 gvar S, Kjndal BR (1981) Succession, diversity and feeding habHa its of microarthropods in decomposing birch leaves, Pedobiologia 22:385408 Huhta V (1972) Efficiency of different dry funnel techniques in extracting Arthropoda from raw humus forest soil. Ann Zool Fenn 9:4248 Huhta V, Seta la H (1990) Laboratory design to simulate complexity of forest floor for studying the role of fauna in the soil processes. Biol Fertil Soils 10:155162 Huhta V, Hyvo nen R, Kaasalainen P, Koskenniemi A, Muona J, Ma kela I, Sulander M, Vilkamaa P (1986) Soil fauna of Finnish coniferous forests. Ann Zool Fenn 23:345360 Hunt HW, Coleman DC, Ingham RE, Ingham ER, Elliott ET, Moore JC, Rose SL, Reid CPP, Morley CR (1987) The detrital food web in a shortgrass prairie. Biol Fertil Soils 3:5768 Kajak A, Jakubczyk H (1977) Experimental studies on predation in the soil-litter interface. In: Lohm U, Persson T (eds) Soil organisms as components of Ecosystems. Proceedings of the VI international colloquium of soil zoology. (Ecological Bulletin, vol 25) Stockholm, pp 493496 Laakso J, Salminen J, Seta la H (1995) Effects of abiotic conditions and microarthropod predation on the structure and function of soil animal communities. Acta Zool Fenn 196:162167 Lawton JH (1995) Ecological experiments with model systems. Science 169:328331 Levin SA (1992) The problem of pattern and scale in ecology. Ecology 73:19431967 Macfadyen A (1961) Improved funnel-type extractors for soil arthropods. J Anim Ecol 30:171184 Moore JC, Walter DE, Hunt WH (1988) Arthropod regulation of micro- and mesobiota in belowground detrital food webs. Annu Rev Entomol 33:419439 Persson T, Clarholm M, Lundkvist H, So derstro m B, Sohlenius B (1980) Trophic structure, biomass dynamics and carbon metabolism in a Scots pine forest. In: Persson T (ed) Structure and function of northern coniferous forest an ecosystem study. Ecol Bull 23:419459

Conclusions Preventing a community of soil meso- and macrofauna from freely exploiting their habitat brought about surprisingly small effects on the total number of taxa and abundance of individuals in the microarthropod community. However, when the community development is analysed for each of the samples (mesh bags and field samples) separately, the development of the community assemblage showed much greater variation in closed versus open systems. Since forest soil is a patchy habitat for soil organisms (Anderson 1977; Swift et al. 1979), this phenomenon is most likely attributable to differences in the faunal composition in the soil cores when establishing the experiment. That the closed systems lost remarkably few species during the 6 month study period supports the findings by Gascon and Travis (1992) that experiments on small spatial scales do not necessarily distort the numerical dynamics of organisms in closed systems. The most conspicuous of the results was the reduction of large invertebrates, particularly top predators, and the simultaneous increase of enchytraeids in the closed systems. That large and mobile predators are sensitive to restriction in their spatial dimensions is commonly observed in laboratory (Beyers and Odum 1993; Laakso et al. 1995) and field experiments (Sih et al. 1985). The decline in large predators and the subsequent uncontrollable increase of their prey in spatially restricted systems may thus severely distort the natural food web dynamics, and has the potential to induce anomalies in the energy and material transfers in enclosure experiments. However, as far as short-term experiments with soil food webs are concerned, reduction of large predators does not necessarily seem to affect the organisms at the bottom of the food web. Since the great majority of energy bound in dead organic material flows through microbes and their microfaunal grazers, including Protozoa and Nematoda (Persson et al. 1980; Hunt et al. 1987), predatory induced changes at the very top of the food web are unlikely to manifest themselves further down the food web. In the long run, however, the indirect effects brought about by predation may well reach down to the microbial component, thus potentially affecting the decomposition and nutrient turnover in a system (Bengtsson et al. 1995).

Acknowledgements We thank Jari Haimi, Veikko Huhta, Jouni Laakso and Juha Mikola for their valuable comments on the early drafts of the manuscript. The study was supported by the grant provided by the Finnish Biological Association Vanamo.

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57 Appendix Summary of the microarthropod species encountered in the field samples (Field), and in mehs bags with either 1 mm opening (Open) or 27 lm opening (Closed) at the end of the study (23 OcField, n = 5 x Acari Brachychtonidae spp. Camisia spinifer C. L. Koch Carabodes forslundii Chamobates Eupelops sp. Eviphis sp. Heminothrus sp. Nanhermannia sellnicki Forssl. Nothrus silvestris Nicolet Oppioidea Paleacaroidea Parazercon radiatus Berl. Pergamasus sp. Phthiracarus sp. Prostigmata+Astigmata Prozercon kochi Sellnick Scheloribates confundatus Selln. Steganacarus carinatus C. L. Koch Tectocepheus velatus Michael Trachytes sp. Uropodina sp. Veigaia nemorensis C. L. Koch Vulgarogamasus kraepelini Berl. Unidentified juv. Unidentified Gamasina R Collembola Anurophorus septentrionalis Palissa Folsomia spp. Isotoma hiemalis Scho tt I. notabilis Scha ffer Isotomiella minor Scha ffer Lepidocyrtus sp. Megalothorax minimus Willem Neanura muscorum Templeton Onychiuridae sp. 1
sp. 2

tober 1995). Mean number per square metre (x), standard error for mean (SE), and incidence (I) are given for untransformed data Open, n = 10 Closed, n = 10 I 0.6 0.2 0 0.4 0 0.1 0.8 0.4 0.9 0.9 0.1 0.6 0.5 0.4 0.5 0.4 0.3 0 0.6 0.6 0.8 0.4 0.1 0.8 1.0 x 4 720 120 40 1 600 80 80 16 960 760 8 080 15 720 120 3 640 80 720 2 800 1 520 40 560 680 1 520 2 200 240 40 3 280 4 560 70 160 360 10 480 280 280 3 400 0 1 240 680 30 880 2 840 520 200 0 51160 121 320 SE 3 438 85 40 1104 53 80 8 871 464 3 618 11 080 61 1 960 53 291 2 004 1197 40 268 332 1 044 1 480 122 40 1 686 1191 34 293 183 4 653 120 85 1 410 0 276 473 8 580 915 231 200 0 11132 39 564 I 0.6 0.2 0.1 0.4 0.2 0.1 0.9 0.5 0.9 0.9 0.3 0.7 0.2 0.6 0.5 0.3 0.1 0.6 0.5 0.7 0.5 0.4 0.1 0.7 1.0

SE 480 233 253 412 160 0 3 124 506 2 787 5 989 80 1103 150 388 681 98 412 271 1 243 240 852 80 0 686 3 412 16 051 320 1 403 392 98 219 0 233 2 011 371 299 310 0 2 759 17 191

I 0.6 0.4 0.4 1.0 0.2 0 0.8 0.8 1.0 1.0 0.2 0.4 0.6 0.4 0.6 0.4 0.8 0.8 0.6 0.2 0.8 0.2 0 0.6 1.0

x 3 400 120 0 680 0 160 25 140 480 10 080 12 400 80 5 920 520 360 680 1 560 120 0 440 840 1 000 400 120 2 440 79 380

SE 1762 85 0 332 0 160 12 499 222 4 208 5 027 80 2 862 231 183 316 821 61 0 173 379 359 215 120 865 27 273 40 2 151 0 183 409 0 858 430 10 954 723 210 0 0 14 107 40 255

880 320 400 1 440 160 0 4 400 800 3 680 10 320 80 1 200 320 480 1 600 160 960 640 2 160 240 1 680 80 0 1120 14 320 47 440 320 6 400 560 160 1 200 0 720 8 800 1 040 560 400 0 20 400 67 840

0.2 1.0 0.4 0.4 1.0 0 0.8 0.2 1.0 1.0 0.6 0.4 0

40 9 600 0 440 1 880 0 2 400 240 31 560 1 840 440 0 0 48 440 127 820

0.1 1.0 0 0.5 0.9 0 0.9 0.3 1.0 0.8 0.5 0 0

0.4 1.0 0.4 0.6 1.0 0 0.9 0.4 1.0 0.8 0.4 0.1 0

Hypogastruridae sp. 1 sp. 2 Sminthuridae sp. R Total