Effects of oil palm plantations on habitat structure and fish assemblages in Amazon streams

The aim of this research is to assess the effects of oil palm plantations on stream habitat and their fish assemblage diversity. We hypothesize that streams which drain through oil palm plantations tend to be less heterogeneous, limiting the occurrence of many species, than streams that drain through forest fragments, which support higher fish diversity. A total of 17 streams were sampled; eight in forest fragments and nine in oil palm plantations. Environmental and biological variables were sampled along 150 m stretch in each stream. Of the 242 environmental variables measured, ten were considered important to assess the condition of structural habitat, and out of these variables, four were considered relevant in the distinction between streams in oil palm plantations and forest fragments. A total of 7245 fishes were collected, belonging to 63 species. Unlike our original hypothesis, the species richness did not differ between forest fragment and oil palm plantations streams, showing that it is not a good divert measure in streams disturbance assessment. However, fish assemblages differed in species composition, and 56 species were recorded in oil palm plantation streams, while 44 species were recorded in forest fragments streams. Some species were identified as indicators of either altered (Aequidens tetramerus and Apistogramma agassizii) or undisturbed areas (Helogenes marmoratus). Overall, oil palm plantations were proven to change stream habitat structure and fish species distribution, corroborating other studies that have evidenced changes in patterns of biological community structure due to impacts by different land uses.     

Ecological structure of tropical fish assemblages in wet-zone streams of Sri Lanka

The introduction of four fish species into a depauperate stream system in central Sri Lanka provided a 'natural experiment' that enabled us to determine the ecological structure of wet-zone stream fish assemblages. All the species indigenous to this 'introduction' stream system also co-occur naturally with the introduced species in nearby streams. Analyses of habitat use and dietary requirements revealed that most species in the assemblages were segregated on the basis of macrohabitat, microhabitat and food, regardless of origin of the fishes. Macrohabitat and microhabitat utilizations by species were similar in each stream of the 'introduction' system despite differences in stream conditions. Thus species distributions, relative to each other in ecological space, were consistent among streams, and with the combined data. High overlaps along the resource axes of velocity, depth, substratum and food were few. A major proportion of the high overlaps was due to associations with introduced species; especially Barbus nigrofasciatus and Barbus cumingii. The other two introduced species, Barbus titteya and Rasbora vaterifloris , were more specialized, and interacted relatively less with indigenous species. The indigenous species, however, exhibited pronounced complementarity along three resource axes representing depth, velocity and food. Rasbora vaterifloris and B. nigrofasciatus grew to larger sizes in one stream which had fewer indigenous species, suggesting competitive release. In this stream, the most common indigenous species was a dietary specialist that fed on diatoms, and B. nigrofasciatus , which fed heavily on diatoms in other streams, switched to feeding more on macrophytes. Overall, the data suggest that these assemblages are predictable, co-evolved systems with competition serving as an important structuring force that reinforces species segregation.     

Studies on the bottom fauna in a tropical freshwater fish pond

Summary Quantitative samples of the bottom fauna obtained from a tropical freshwater fish pond during a period of two years were studied with regard to species composition, zonal variation and seasonal fluctuation. Two different zones with differences in faunal composition could be distinguished. The littoral zone was dominated by Mollusca, whereas Oligochaetes were predominant in the profundal zone. Most of the organisms showed their peaks in the months January to April. Breeding activities ofVivipara bengalensis, Digoniostoma cerameopoma, Melanoides granifera andPisidium clarkeanum as evidenced by presence of younger length groups in the populations were also observed during the same period (January to April). The influence of environmental factors as temperature, alkalinity and food were discussed as probable causes for the peaks in abundance. The average number of total organisms per square metre was found to be comparatively high indicating a high standing crop. Differences observed in the total number of organisms present have been attributed to inflow of water resulting in sudden alteration of the nutrient condition of the bottom mud. The seasonal variations of the standing crop of bottom fauna during both the years have been found to be influenced by fluctuations of the bivalve mollusc,Lamellidens corrianus. The availability of sufficient amount of benthic fauna for consumption by bottom feeding fishes has been indicated.This work formed part of a thesis submitted for the Degree of Doctor of Philosophy to the University of Calcutta, India in 1964.     

The influence of trophic status and large-scale climatic change on the structure of fish communities in Perialpine lakes

1. A recurrent question in ecology is the influence of environmental factors, particularly nutrients and climatic variables, on community structure and functioning, and their interaction with internal community processes (e.g. competition). 2. Perialpine lakes have been subject to two main kinds of human-induced changes over the last 50 years: eutrophication-reoligotrophication, represented by lake-specific changes in total phosphorus concentration (TP), and long-term global climatic change, captured by average winter temperature (AWT). 3. Changes in fish communities (abundance of seven species from fishery data) in 11 Perialpine lakes during 31 years (1970-2000) were investigated in relation to variation in TP and AWT using models incorporating the effects of fish maturation age, and potentially discriminating effects on adult survival and recruitment. 4. We show that phosphorus concentration affects fish abundance in species-specific ways. These effects are mediated by recruitment rather than by adult survival. Phosphorus effects are probably modulated by interspecific interactions, as increasing TP enhances total community biomass, which in turn is either positively or negatively associated with species abundance depending on species position in trophic chains. 5. Climatic change has very little effect on fish abundances, which is not consistent with the prediction of larger changes in species near their southern distribution boundary. 6. We propose several hypotheses to account for those findings, and place our study in the wider framework of community ecology.     

Taxonomic and functional homogenization of an endemic desert fish fauna

Aim The highly endemic fishes of the arid Southwest USA have been heavily impacted by human activities resulting in one of the most threatened fish faunas in the world. The aim of this study was to examine the patterns and drivers of taxonomic and functional beta diversity of freshwater fish in the Lower Colorado River Basin across the 20th century. Location Lower Colorado River Basin (LCRB). Methods The taxonomic and functional similarities of watersheds were quantified to identify patterns of biotic homogenization or differentiation over the period 1900–1999. Path analysis was used to identify the relative influence of dam density, urban land use, precipitation regimes and non‐native species richness on observed changes in fish faunal composition. Results The fish fauna of the LCRB has become increasingly homogenized, both taxonomically (1.1% based on β_sim index) and functionally (6.2% based on Bray–Curtis index), over the 20th century. The rate of homogenization varied substantially; range declines of native species initially caused taxonomic differentiation (?7.9% in the 1960s), followed by marginal homogenization (observed in the 1990s) in response to an influx of non‐native species introductions. By contrast, functional homogenization of the basin was evident considerably earlier (in the 1950s) because of the widespread introduction of non‐native species sharing similar suites of biological traits. Path analysis revealed that both taxonomic and functional homogenization were positively related to the direct and indirect (facilitation by dams and urbanization) effects of non‐native species richness. Main conclusions Our study simultaneously examines rates of change in multiple dimensions of the homogenization process. For the endemic fish fauna of the LCRB, we found that the processes of taxonomic and functional homogenization are highly dynamic over time, varying both in terms of the magnitude and rate of change over the 20th century.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

The influence of habitat on the spatial variation in fish assemblage composition in an unimpacted tropical River of Ganga basin, India

Effects of local habitat variables on the structure of fish assemblage were evaluated from 50 sampling sites in a tropical River of Central India of the Ganges basin with limited anthropogenic disturbance covering premonsoon, monsoon and postmonsoon periods. Data were analyzed for 5,186 fish individuals of 24 freshwater fish species of conservation and fishery management interest. Out of the total fish species, seven belong to the ??endangered?? and 8 belong to the vulnerable category. A Cyprinid, Puntius sarana, was the most widely distributed species (frequency of occurrence 76%) out of the total species in this study. We used canonical correspondence analyses to determine the influence of environmental conditions on species occurrences and assemblage characteristics. Regarding the microhabitat, hydromorphological parameters (depth and current velocity) followed by temperature, turbidity and total dissolve solid were of significant for the structure of the fish community. Conductivity was another important factor that explained the major proportion of the variability affecting fish in their habitat choice. The other local habitat variables like overhanging vegetation and land use were of secondary but significantly important for the assemblage of the fishes. Our results suggests the importance of local environment influences on the fishes of conservation importance and their assemblage characteristics in an unimpacted river and provide a framework and reference conditions to support restoration efforts of relatively altered fish habitats in tropical rivers of India.     

The fish fauna of mountain streams in the Guanshan National Nature Reserve, Jiangxi, China

Guanshan National Nature Reserve (GNNR) is located in the middle reach of the Jiuling Mountain Range in the southern China and the first national nature reserve within Jiangxi Province. The fish fauna of mountain streams in the GNNR was investigated from September 2006 to August 2007. A total of 379 fish were collected and classified into two families and seven species. Acrossocheilus parallens [relative abundance (RA) 44.3%] was the most abundant fish species. Subdominant species were Onychostoma barbata (RA, 20.3%), Zacco platypus (RA, 14.2%) and Opsariichthys bidens (RA, 12.4%).A greater number of species (7 vs. 2) and individuals (295 vs. 84) were collected from the West Stream compared to the East Stream. Among the seven fish species collected in the GNNR, Acrossocheilus parallens, Onychostoma barbata and Vanmanenia stenosoma are endemic to the Chinese mountain streams. This study suggests that mountain streams in the nation reserves including GNNR are very important for freshwater fish diversity and conservation in Jiangxi Province.     

Simple predictions of instream habitat model outputs for fish habitat guilds in large streams

1. In the context of a generalised modification of hydraulic conditions in medium to large streams, modelling the impacts of stream regulation on fish communities in multiple streams is an important challenge for basic and applied freshwater ecology. Conventional instream habitat models such as PHABSIM link a hydraulic model with preference curves for various species to estimate habitat value changes with discharge in stream reaches. Despite world‐wide applications, they have been scarcely used in multiple sites with multiple species. 2. We assigned 21 size classes of European fish species to four habitat guilds (cluster analysis grouping size classes with comparable microhabitat preference curves). Then, we ran a conventional instream habitat model on 28 French stream reaches belonging to the `barbel zone', to estimate habitat values versus discharge curves for the 21 size classes. We summarised the outputs as mean habitat values for guilds, and tested if they were predictable from average characteristics of reaches (discharge, depth, width, particle size). 3. As was obtained elsewhere for populations, habitat values for guilds were strongly related to average, dimensionless characteristics of reaches. The Reynolds number of reaches, equivalent to a discharge per width unit, reflected most of the discharge‐dependent changes in habitat values (within reaches). In particular, habitat values of species preferring bank (respectively midstream) microhabitats decreased (respectively increased) with increasing Reynolds number. The Froude number at median discharge was the major predictor of reach‐dependent but discharge‐independent variations in habitat values. Habitat values of species preferring riffle versus pool or bank microhabitats were higher in reaches with high Froude numbers. These relationships were consistent with existing knowledge on the different species. 4. Such results suggest that the input variables required to estimate habitat values for fish communities can be greatly simplified, as illustrated by a general estimation of the sensitivity of species preferring midstream habitats to discharge changes in any reach. Cost‐efficient alternatives to conventional instream habitat models should facilitate their validation in multiple sites, a point that remains critical in instream habitat modelling of fish communities.     

The structure and organization of an eelgrass fish fauna

Summary Ten species of fish made up more than 95 percent of the standing crop of the total fish assemblage of a shallow-water eelgrass meadow in southeastern Australia. The relative abundance of each species remained fairly constant from year to year during a four and a half year period. Four species were permanent residents, three moved over the meadow at high tide, and the juveniles of three species changed residence status as they grew. Most species exhibited distinct diel patterns of activity, four species were diurnal and five were nocturnal. Species foraged in different microhabitats and dietary overlap was very low amongst all but two species which had overlapping habitat, prey and prey size preferences, but had different diel activity periods. Data on production by prey and consumption of prey by these two fish species indicated that competition for food was probably responsible for the temporal separation of feeding niches. Predation by birds and fish appeared to be the major influence on the diel activity period of one fish species. The maintenance of the patterns of resource allocation among the remaining species is discussed and the organization of different seagrass fish faunas is compared.This is publication number 265 in the Ministry for Conservation, Victoria publication series. — Present address: Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1, Canada     

Effects of habitat complexity on trophic interactions of three congeneric fish species

Aquatic Ecology - Habitat complexity can substantially alter trophic relationships, such as competitive and predatory interactions, between fish species. This study aimed to evaluate how trophic...     

The influence of chemistry and habitat features on the microcrustacea of some upland Welsh streams

SUMMARY. 1. Microcrustaceans were sampled during September 1990 from three microhabitats (margins, stony riffles and gravel) at thirteen stream sites in mid-Wales.
2. Patterns in the distribution, abundance and community structure of microcrustacean taxa were related to stream chemistry, physiography, substratum composition and marginal habitat structure. Patterns of distribution between microhabitats were also assessed.
3. Chemical variables were the strongest correlates with microcrustacean parameters. Total numbers of microcrustaceans and harpacticoid copepods were reduced at sites with high aluminium concentrations, and ostracods were scarce or absent at the most acidic sites. In contrast, cyclopoid copepods were more abundant and species rich at low pH.
4. Microcrustacean communities were most species rich at sites with a high percentage cover of macrophytes on the substratum. Cladoceran species richness and abundance, and the abundance of several microcrustacean species, were also positively correlated with percentage macrophyte cover.
5. Three site groupings produced by TWINSPAN classification were primarily related to percentage macrophyte cover and marginal habitat structure, with chemistry and land use also of importance.
6. Total microcrustacean abundance was highest in margins, but of all the species, only the harpacticoid Bryocamptus cuspidatus showed a strong microhabitat preference.     

Morphological traits mediate fish occurrences in oil palm-impacted tropical streams

1. Land-use change is a leading driver of biodiversity loss, especially in tropical fresh waters where the conversion of natural forest to monoculture plantations impacts freshwater fish assemblages. The environmental pathways underpinning shifts in fish assemblages, however, are poorly understood, but could potentially be inferred from trait–environment relationships.
2. We addressed this knowledge gap using eco-morphological traits to explain fish occurrences in oil palm-impacted streams of the Endau drainage in Peninsular Malaysia. We also investigated how traits relate to differences in environmental conditions associated with land-use change. We then integrate findings from the above to test how potential pathways of land-use driven environmental changes can impact species occurrences through effects on life history, feeding habits, and mobility.
3. Mixed-effects models show that fishes with superior (upward-facing) mouths and low body mass were more likely to occur in oil palm streams than forest streams, and these traits were associated with grass-dominated riparian zones and reduced woody debris in oil palm streams, respectively. Structural equation models show that mouth positions statistically mediated the effect of riparian vegetation on fish species occurrences in oil palm streams. Specifically, fishes with superior mouths were more likely to feed on terrestrial invertebrates.
4. Our analysis of easily measurable traits revealed pathways of land-use impact that are potentially more widely applicable than conventional taxa-based approaches. Fishes with superior mouths tended to occur in oil palm streams as they were able to more effectively exploit inputs of terrestrial invertebrates that are potentially associated with grass-dominated riparian vegetation.
5. Moreover, shifts in traits may suggest land-use driven changes in stream ecosystem functioning (e.g. in terms of the role of terrestrial subsidies), thereby informing targeted management actions in land-use impacted habitats (e.g. retention/restoration of riparian trees).

     

Food web pathways for fish communities in small tropical streams

1. It is often assumed that invertebrate consumers in small tropical streams are dependent on allochthonous sources, although recent studies indicate that algae can form the base of food webs in tropical streams. Fish in tropical streams can feed across several trophic levels and the origin and path of energy and nutrient flow is uncertain for many species.
2. We collected fish, insects, periphyton, and leaf litter from 20 streams across four Atlantic Forest catchments. We analysed stomach contents of fish to define trophic guild and fish dietary trophic position. We also analysed stable isotopes of carbon and nitrogen of fish and their resources to identify the main basal resources of the food web and to estimate trophic positions and identify the path of energy flow.
3. We found that autochthonous sources were the primary resource base for fish communities. Trophic positions estimated from diet and isotopes were similar and correlated for insectivore and algivore–insectivore fish, but not for algivore–detritivore or omnivore fish. Using path analysis, fish classified as algivore–detritivores appear to have derived their biomass through a diet of primary consumer insects and periphytic algae and thus, are more likely to play a trophic role as algivore–insectivores in these streams. However, omnivores probably derived much of their biomass from aquatic insects.
4. Our findings support other studies of tropical systems in which the main basal resource is autochthonous, even in small streams. We also show that the assignment to a specific trophic guild for some fish species, based on gut contents, does not reflect what they assimilate into their bodies. In some species, food sources that are uncommon can make a disproportionately important contribution to their biomass.
5. This study affirms the important role of inconspicuous algal resources in aquatic food webs, even in small forested streams, and demonstrates the effectiveness of taking a combined approach of diet analysis, isotopic tracing, and modelling to resolve food web pathways where the level of omnivory is high.

     

Null model of biotic homogenization: a test with the European freshwater fish fauna

In recent years, there has been growing concern about how species invasions and extinctions could change the distinctiveness of formerly disparate fauna and flora, a process called biotic homogenization. In the present study, a null model of biotic of homogenization was developed and applied to the European freshwater fish fauna. We found that non-native fish species led to the greatest homogenization in south-western Europe and greatest differentiation in north-eastern Europe. Comparing these observed patterns to those expected by our null model empirically demonstrated that biotic homogenization is a non-random ecological pattern, providing evidence for previous assumptions. The place of origin of non-native species was also considered by distinguishing between exotic (originating from outside Europe) and translocated species (originating from within Europe). We showed that exotic and translocated species generated distinct geographical patterns of biotic homogenization across Europe because of their contrasting effects on the changes in community similarity among river basins. Translocated species promoted homogenization among basins, whereas exotic species tended to decrease their compositional similarity. Quantifying the individual effect of exotic and translocated species is therefore an absolute prerequisite to accurately assess the spatial dynamics of biotic homogenization.     

Geographic shifts in the effects of habitat size on trophic structure and decomposition

Habitat size is known to affect community structure and ecosystem function, but few studies have examined the underlying mechanisms over sufficient size gradients or in enough geographic contexts to determine their generality. Our goal in this study was to determine if the relationship between habitat size and leaf decomposition varied across geographic sites, and which factors may be driving the differences. We conducted replicated observations in a coastal forest in Brazil, and in rainforests in Costa Rica and Puerto Rico. We used leaf litter decomposition and macroinvertebrate composition in bromeliad phytotelmata of varying sizes to determine the relationships between habitat size, trophic structure and decomposition over a wide geographical range. We experimentally disentangled the effects of site and litter quality by quantifying invertebrate control of decomposition of a native and a transplanted litter type within one site. We found that the relationship between bromeliad size and decomposition rates differed among study sites. In rainforests in Costa Rica and Puerto Rico, decomposition was strongly linked to macroinvertebrate trophic structure, which varies with bromeliad size, driving strong bromeliad size‐decomposition relationships. However, in Brazil there was no relationship between bromeliad size and decomposition. Our manipulative experiment suggests that within coastal forest in Brazil, the poor quality of native litter resulted in little invertebrate control of decomposition. Furthermore, the key detritivore in this site builds a predator‐resistant case, which likely prevented effects of bromeliad size on trophic structure from being transmitted to decomposition even when litter quality was increased. We conclude that differences in both leaf litter quality and macroinvertebrate traits among sites determine the link between decomposition and macroinvertebrates, and consequently the decomposition‐bromeliad size relationship. These results show that the response of decomposition to habitat size is context‐dependent, and depends on which component of the food web is the main driver of the function.     

Modifications in habitat use patterns and trophic interrelationships in the fish fauna of an oligotrophic artificial lake: Sainte Croix (Provence,France)

Lake Sainte-Croix is a hydro-electric reservoir on the river Verdon that came into operation about 16 years ago. The area of the lake is about 22 km² and the volume about 767 Mm³. The water level drops by as much as 16 m in winter. Sainte-Croix is a warm monomictic oligotrophic lake. The indigenous fish fauna of the Verdon originally included 8 species. Successive stocking has raised this number to 17. In catches made in 1976–1977 with 27 mm mesh nets, 80 to 90% of the fish with a body size of between 15 and 25 cm were of the four main species indigenous to the river before the dam was built: Leuciscus cephalus, Barbus fluviatilis, Chondrostoma toxostoma, Salmo trutta. Catches made at same period with 14 mm mesh nets gave only Chondrostoma toxostoma and Leuciscus cephalus.Changes in the fish fauna were apparent from catches made in 1984 and 1987 with nets of the same mesh size. There was a marked demographic increase of roach, and more recently of bleak and gudgeon, all three species introduced after the construction of the dam. At the same time, there was a decrease in catch frequency of Ch. toxostoma, L. cephalus and B. fluviatilis in the downstream area of the lake. Stomach content analysis carried out in different seasons on the six most abundant species of the lake (roach, toxostome, chub, bleak, perch and gudgeon) provided evidence on the trophic organisation of the fish fauna. We have measured niche breadth for each species by the Simpson diversity index and niche overlap by the Pianka index. Bleak is the only species to feed on zooplankton in open water. Roach is a particularly opportunistic feeder, sometimes taking plankton, but at Sainte-Croix it is exclusively herbivorous and benthophagous. Niche overlap between these two species is slight or null, depending on the season. On the other hand, there is a high degree of overlap between roach and chub and between roach and toxostome. The decrease in toxostome and chub frequency in the downstream area of the lake, where roach is most abundant, would appear to be a competitive exclusion phenomenon. There is high niche overlap between bleak and young perch, which partly explains the low abundance of the latter species; other contributory factors are the scarcity of zooplankton inshore, and the scarcity of zoobenthos resulting from the drop in the water level in winter.     

The soil fauna of a beech forest on limestone: trophic structure and energy budget

Summary The soil fauna of a mull beech forest on lime-stone in southern Lower Saxony (West Germany) was sampled quantitatively. Biomass estimates, trophic characteristics, and measurement and calculation of the energetic parameters of the constituent animal populations were used to construct an energy budget of the total heterotrophic subsystem of the forest. Mean annual zoomass amounted to about 15 g d wt m^–2; earthworms (about 10 g d wt m^–2) and other groups of the macrofauna were dominant. Protozoa constituted about 1.5 g d wt m^–2. Relative distribution of zoomass among the trophic categories was 50% macrosaprophages, 30% microsaprophages, 12% microphytophages, and 4% zoophages. Total annual consumption rate of the saprophagous and microphytophagous soil fauna (6328 and 4096 kJ m^–2 yr^–1, respectively) was of the same order of magnitude as annual litter fall (canopy leaves 6124 kJ m^–2 yr^–1, flowers and fruits 944 kJ m^–2 yr^–1, herbs 1839 kJ m^–2 yr^–1, fine woody material 870 kJ m^–2 yr^–1, tree roots 3404 kJ m^–2 yr^–1, without coarse woody litter). Primary decomposers (macrosaprophages) were the key group for litter comminution and translocation onto and into the soil, thus contributing to the high decomposition rate (k=0.8) for leaf litter. Consumption rates of the other trophic groups were (values as kJ m^–2 yr^–1): bacteriophages 2954, micromycophages 416, zoophages 153. Grazing pressure of macrophytophages (including rhizophages) was low. Faeces input from the canopy layer was not significant. Grazing pressure on soil microflora almost equalled microbial biomass; hence, a large fraction of microbial production is channelled into the animal component. Predator pressure on soil animals is high, as a comparison between consumption rates by zoophages and production by potential prey — mainly microsaprophages, microphytophages and zoophages — demonstrated. Soil animals contributed only about 11% to heterotrophic respiration. However, there is evidence that animals are important driving variables for matter and energy transfer: key processes are the transformation of dead organic material and grazing on the microflora. It is hypothesized that the soil macrosaprophages are donor-limited.