Differences in burrowing behaviour and spatial interaction between the two earthworm species Aporrectodea nocturna and Allolobophora chlorotica

 To study intraspecific and interspecific interactions between different ecological types of earthworm, the burrowing behaviour of two earthworm species (the anecic earthworm Aporrectodea nocturna and the endogeic earthworm Allolobophora chlorotica) was observed in a microcosm. Earthworms were either alone in the microcosm, together with a conspecific earthworm, or with an earthworm of the other species. Observations under red light, including those of the position of the animals and the burrow dug, were recorded 4 times a day for 8 days and provided the data needed to reconstruct the probable trajectories of each earthworm. Differences in movement and burrowing behaviour were observed. Comparisons between the two species confirmed the expected behaviour of each ecological type: A. nocturna reused its burrow system regularly, whereas A. chlorotica rarely did. Moreover, it was shown that A. chlorotica burrowed less and explored a smaller surface when in the presence of A. nocturna. Besides, A. nocturna burrowed less and explored a smaller surface when in the presence of another A. nocturna. If these interactions occur under natural conditions, they could affect the structure of the burrow systems of the earthworm species examined. Received: 15 January 1999     

Burrow systems made by Aporrectodea nocturna and Allolobophora chlorotica in artificial cores: morphological differences and effects of interspecific interactions

Burrow systems of earthworms contained in artificial cores were analysed through X-ray computed tomography and 3D skeleton reconstructions. Gas diffusion experiments were carried out on these cores to characterize soil transfer properties associated with the different burrow systems. Three types of cores were studied: cores in which Aporrectodea nocturna, an anecic earthworm was introduced (treatment 1), cores in which Allolobophora chlorotica, an endogeic earthworm was introduced (treatment 2) and cores that contained both species (treatment 3). Comparisons of the characteristics of the burrow systems of treatment 1 and 2 show important differences: the burrow system of A. nocturna comprises fewer burrows, which are longer, less branched, more vertical and have a lower sinuosity. The burrow system of A. chlorotica is characterized by lower continuity, which however did not result in a lower soil diffusivity. To study the burrow systems made by the two species in the same core (treatment 3), a separation that was based on differences in pore diameter between the two species and that takes into account the burrow orientation was designed. This separation was proven to be efficient since it resulted in low percentages of errors (around 10%) when applied to the burrow systems of treatments 1 and 2. Comparison of the burrow systems from treatments 1 and 3 demonstrated that the burrow system of A. nocturna was influenced by the presence of A. chlorotica: in treatment 3, A. nocturna made burrows that were smaller, more vertical and less branched. However, these interactions have to be confirmed under natural conditions.     

Sublethal effects of imidacloprid on the burrowing behaviour of two earthworm species: Modifications of the 3D burrow systems in artificial cores and consequences on gas diffusion in soil

Earthworms have been termed ‘ecosystem engineers’ (sensu [Jones, C.G., Lawton, J.H., Shachak, M., 1994. Organisms as ecosysem engineers. Oikos 69, 373-386.]) because of the important roles they play in the soil. As a consequence, it is assumed that if earthworms change their behaviour following exposure to pesticides or pollutants this could have a drastic impact on soil functioning. To test this assumption under laboratory conditions, we studied the burrow systems made by two earthworm species (the anecic Aporrectodea nocturna and the endogeic Allolobophora icterica) in artificial soil cores containing imidacloprid, a widely used neonicotinoid insecticide. After 1-month incubation period, the macropores created in the soil core were analyzed by tomography. In order to further characterize transfer properties associated with burrow systems gas diffusion measurements were also carried out. The burrow systems made by the two earthworm species were very different: A. nocturna made more continuous, less branched, more vertical and wider burrows than A. icterica. Some changes to A. nocturna burrow systems were observed after exposure to imidacloprid (they made a smaller burrow system and burrows were more narrow), but only at the highest concentration of imidacloprid used (0.5 mg kg⁻¹). A. icterica worms were more sensitive to imidacloprid and many differences in their burrow systems (length, sinuosity, branching rate and number of burrows) were observed at both concentrations tested (0.1 and 0. 5 mg kg⁻¹). As a consequence, the continuity of the burrow systems made by both species was altered following imidacloprid treatment. Gas diffusion through the A. nocturna soil cores was reduced but no difference in gas diffusion was observed in the A. icterica soil cores.     

Quantitative estimates of burrow construction and destruction,by anecic and endogeic earthworms in repacked soil cores

Although the role of earthworms in soil functioning is often emphasised, many important aspects of earthworm behaviour are still poorly understood. In this study we propose a simple and cost-effective method for estimating burrow system area and continuity, as well as a new and often neglected parameter, the percentage of burrow refilling by the earthworms own casts. This novel parameter is likely to have a huge influence on the transfer properties of the burrow system. The method uses standard repacked soil cores in PVC cylinders and takes advantages of clay shrinkage and the fact that earthworms were previously shown to prefer to burrow at the PVC/soil interface. In this way, after removing the PVC cylinders off dry cores, the external section of the burrow system made by earthworms along the soil walls could be easily described. We applied this method to characterise the burrow systems of four earthworms species: two anecics (Aporrectodea caliginosa nocturna and Aporrectodea caliginosa meridionalis) and two endogeics (Aporrectodea caliginosa icaliginosa and Allolobophora chlorotica). After one month the burrow's area generated by both anecic species were much larger (about 40 cm²) than the endogeic burrow's area (about 15 cm²). A. nocturna burrow system continuity was higher than that of A. meridionalis and both anecic burrow systems were more continuous than those made by the endogeic earthworms. This was partly explained by the far larger proportion of the burrow area that was refilled with casts: approximately 40% and 50% for Al. chlorotica and A. caliginosa, respectively compared with approximately 20% for the anecic burrows. We discuss whether these estimates could be used in future models simulating the dynamics of earthworm burrow systems by taking into account both burrow creation and destruction by earthworms.     

3D skeleton reconstructions of natural earthworm burrow systems using CAT scan images of soil cores

 Four soil cores (length, 20 cm; diameter, 16 cm) were sampled in a Swiss pre-Alpine meadow with high earthworm abundance (>400 individuals/m²); two cores were taken in October 1993 and the other two cores in April 1994. The cores were described using computer assisted tomography which gives a series of section images every 3 mm. A method for reconstructing the three-dimensional (3D) skeleton of the earthworm burrow system is presented and discussed. This method provides an image of the structural organisation of the burrow system and was found to be adequately sensitive for use in ecological and functional studies. The seasonal variation of these 3D skeletons was investigated using two approaches, i.e. the analysis of: (1) global burrow system characteristics, and (2) individual burrow characteristics. At the scale of the global burrow system no difference was found between seasons (same number of burrows and same total burrow length) except for the vertical segment distribution, which was homogeneous in spring and decreased with depth in the fall. The study of individual burrow characteristics revealed that burrows tended to be more vertical in spring and that their branching intensity was higher in this season. Received: 10 June 1997     

Characterisation of the three-dimensional structure of earthworm burrow systems using image analysis and mathematical morphology

The aim of this work was to exemplify the specific contribution of both two- and three-dimensional (3D) X-ray computed tomography to characterise earthworm burrow systems. To achieve this purpose we used 3D mathematical morphology operators to characterise burrow systems resulting from the activity of an anecic ( Aporrectodea nocturna), and an endogeic species ( Allolobophora chlorotica), when both species were introduced either separately or together into artificial soil cores. Images of these soil cores were obtained using a medical X-ray tomography scanner. Three-dimensional reconstructions of burrow systems were obtained using a specifically developed segmentation algorithm. To study the differences between burrow systems, a set of classical tools of mathematical morphology (granulometries) were used. So-called granulometries based on different structuring elements clearly separated the different burrow systems. They enabled us to show that burrows made by the anecic species were fatter, longer, more vertical, more continuous but less sinuous than burrows of the endogeic species. The granulometry transform of the soil matrix showed that burrows made by A. nocturna were more evenly distributed than those of A. chlorotica. Although a good discrimination was possible when only one species was introduced into the soil cores, it was not possible to separate burrows of the two species from each other in cases where species were introduced into the same soil core. This limitation, partly due to the insufficient spatial resolution of the medical scanner, precluded the use of the morphological operators to study putative interactions between the two species     

Dynamic study of the burrowing behaviour of Aporrectodea nocturna and Allolobophora chlorotica: interactions between earthworms and spatial avoidance of burrows

The behaviour of earthworms belonging to two different species and ecological types (Aporrectodea nocturna and Allolobophora chlorotica) was studied using two-dimensional (2D) terraria. Two experiments were set up to gain insight into the nature of interactions between these earthworms. Firstly, the evolution of the burrow systems was analysed with the density of the earthworms varying from one to five individuals. Secondly, a burrow system was first established by using one earthworm which was then removed before the introduction of a second earthworm. This second earthworm therefore encountered a burrow system created either by a conspecific earthworm or by an earthworm of the other species. These experiments showed that: (1) intra- and interspecific interactions occur between earthworms, (2) these interactions are dependent on the physical presence of the earthworms, and (3) spatial avoidance can occur (A. chlorotica avoiding burrows created by A. nocturna). The results suggest that earthworm burrow systems are "individual structures", rarely used by other earthworms when inhabited. When abandoned, the burrows may be recolonised depending on the ecological type of the earthworm under consideration.     

Quantitative analysis of earthworm burrow systems with respect to biological soil-structure regeneration after soil compaction

 On arable land, tilled with conventional tillage (CT) and conservation tillage (CS) respectively, plots were compacted by wheeling them 6 times with a 5 Mg wheel load in spring 1995. Immediately after compaction, undisturbed soil monoliths were excavated from the compacted and uncompacted plots. The monoliths were defaunated and inoculated with either Lumbricus terrestris or Aporrectodea caliginosa. One monolith from each plot remained uninoculated as a control. After 6 months the monoliths were defaunated again and then scanned with X-ray helical computed tomography. The data were transformed, the void systems inside the monoliths were reconstructed and visualised, and the parameters total void length, total void volume, tortuosity and continuity were quantified. The parameters' values were generally lower in the controls than in the inoculated monoliths. Differences in burrow construction could be explained by the different life strategies of the two earthworm species. Changes in burrow morphology due to tillage system and soil compaction were minor. Only the continuity of the burrow systems clearly changed: decreasing for L. terrestris and increasing for A. caliginosa. This can be explained by a change in the earthworms' burrowing activity to minimise energy expenditure in compacted soil. By extrapolating field data, we concluded that earthworms have great potential for biologically regenerating the soil structure after a single compaction event. Due to higher earthworm abundances in soil managed by CS the regeneration of the soil structure is assumed to be better in these plots than those tilled by CT. Received: 17 December 1997     

Earthworm communities in native savannas and man-made pastures of the Eastern Plains of Colombia

 The structure and seasonal changes of earthworm communities were evaluated in a natural savanna and in a improved grass-legume pasture in a Colombian oxisol over a period of 18 months. One plot of 90×90 m was isolated in each of the systems and each month five samples of 1 m²×0.5 m and ten of 20×20×20 cm were randomly selected from a stratified block design. Species richness was similar in the two evaluated plots (seven species), whereas diversity measured by the index, H (Shannon and Weaver 1949) was clearly different, i.e. H=2.89 in natural savanna and H=1.29 in pasture. This is explained by differences in earthworm community structure. The average annual density in the savanna was 49.8, ranging from 10.8 to 135.8 individuals (ind) m^–2, and biomass was 3.3 g m^–2 (hand-sorting method), ranging from 0.9 to 11.5 g m^–2. In the man-made pasture, density was 80.1 ind m^–2 on average, ranging from 24 to 215.8 ind m^–2 and biomass was more than tenfold higher, ranging from 29.2 to 110.4 g m^–2. This was especially due to the presence of a large glossoscolecid anecic species, Martiodrilus carimaguensis Jiménez and Moreno, which has been greatly favoured by conversion of savanna to pasture. Endogeic species were dominant in the natural savanna whereas the anecic species accounted for 88% of total earthworm biomass in the pasture. Total earthworm density and biomass were significantly different in the two systems studied (t-test). The results indicate a clearly positive response of earthworm communities to improved pastures, a type of land use that is being increasingly adopted in moist neotropical savannas. Received: 20 October 1997     

Surface cast production by the earthworm Aporrectodea nocturna in a pre-alpine meadow in Switzerland

Field and laboratory experiments were carried out to describe the effects of Aporrectodea nocturna on soil characteristics in a pre-alpine meadow and to support the development of a model of cast production. In the prealpine meadow, increased cast production, first observed about 20 years ago around a newly planted hedge, was recorded to a distance of maximal 170 m from the hedge. Numbers of A. nocturna between 130 and 165 m from the hedge decreased from 164 to 16 individuals m^-2. In the same area cast production steadily decreased from about 1.5 kg m^-2 week^-1 to nil, the plant community structure changed and the microbial biomass decreased, but the root biomass and the moisture content did not change. Laboratory experiments demonstrated that high cast production was not a specific feature of the A. nocturna population nor of the soil in the meadow. Diapause of A. nocturna was terminated in the laboratory during September. A model of cast production potential by the earthworm A. nocturna was established using laboratory determinations of the relationships with body weight, temperature, and water potential. The model was shown to predict cast production in the field given the assumption that the water potential was 0 MPa. According to the model, 81% of surface cast production was by juveniles, and 19% by adults of A. nocturna.     

Characterization of the burrow system of the earthworms Lumbricus terrestris and Aporrectodea giardi using X-ray computed tomography and image analysis

 The burrow systems of two earthworm species (Lumbricus terrestris and Aporrectodea giardi) were studied in artificially packed soil columns placed in controlled conditions in the laboratory. At the end of the incubation, which lasted 246 days, the burrow systems were characterized on the undisturbed columns using X-ray computed tomography. This method provided a set of digitized images corresponding to horizontal 3-mm-thick sections. The following parameters were measured using image analysis on each section: number of biopores, their volume, and their individual orientation calculated using an elliptical model of the earthworm channel. The profiles of these parameters through the columns showed that the burrow systems of L. terrestris and A. giardi, which are both anecic species, were very different in terms of total volume, number of burrows, burrow orientation and extension with soil depth. These results led us to conclude that the burrow system of L. terrestris can be considered as a permanent structure whereas that of A. giardi is closer to the burrow system of endogeic species. Received: 12 June 1998     

Burrow systems of endogeic earthworms: Effects of earthworm abundance and consequences for soil water infiltration

By creating burrows, earthworms influence the transfer properties of soils. The effects of endogeic species on soil transfer properties, however, are not yet well understood because these earthworms generally create burrows that are refilled by casts and have no preferential vertical orientation. Thirty soil cores were incubated for various periods (1–3 or 4 weeks) at different earthworm densities (70, 210, 345 or 480 individuals m⁻²). The cores were then scanned using X-ray tomography and the burrow systems were characterised by measuring the total burrow volume, bioturbation volume (refilled burrows and lateral compaction around the burrows), the number of branches, tortuosity and continuity (assessed by computing the number of burrows with a vertical extension greater than 15, 20 and 25% of the core). We also computed the mean geodesic distance, i.e. the mean distance from the bottom to the top of the core assuming that distances inside burrows are null. Rainfall simulations were carried out on 17 cores chosen to encompass the variations observed in the burrow systems. The water transfer efficiency of each core was estimated by measuring two parameters: breakthrough volume and the percentage of water transmitted after 1 h of rain. Burrow and bioturbation volume increased significantly and steadily with time and earthworm density. We estimated that on average Allolobophora chlorotica burrowed 22 cm per week. All other burrow system characteristics also increased with time and earthworm density except the mean geodesic distance, which decreased significantly. This suggests that intraspecific interactions had no significant effect on burrow system geometry. Univariate PLS regressions were used to understand which burrow system characteristics had the strongest influence on water transfer. These regressions showed that the mean geodesic distance was the most important parameter. This means that in addition to individual burrow characteristics, the spatial arrangement of the whole burrow system also had a major effect on transfer properties.     

Urease activity and its relation to soil organic matter, microbial biomass nitrogen and urea-nitrogen assimilation by maize in a Brazilian Oxisol under no-tillage and tillage systems

 We studied the relationship between urease activity (UA) and soil organic matter (SOM), microbial biomass N (N_biom) content, and urea-N fertilizer assimilation by maize in a Dark Red Latosol (Typic Haplustox) cultivated for 9 years under no-tillage (NT), tillage with a disc plough (DP), and tillage with a moldboard plough (MP). Two soil depths were sampled (0–7.5 cm and 7.5–15 cm) at 4 different times during the crop cycle. Urea was applied at four different rates, ranging from 0 to 240 kg N ha^–1. The levels of fertilizer N did not affect the UA, SOM content, and N_biom content. No significant difference between the treatments (NT, DP, and MP) was observed for SOM during the experiment, probably because the major part of the SOM was in recalcitrant pools, since the area was previously cultivated (conventional tillage) for 20 years. The N_biom content explained 97% and 69% of the variation in UA in the upper and deeper soil layer, respectively. UA and biomass N were significantly higher in the NT system compared to the DP and MP systems. The highest maize productivity and urea-N recovery was also observed for the NT system. We observed that the increase in urea-N losses under NT, possibly as a consequence of a higher UA, was compensated for by the increase in N immobilized in the biomass. Received: 2 July 1999     

Assessment of recovery of population, biomass and reproduction of the earthworm Drawida willsi following the application of malathion under field conditions

 The recovery of population, biomass and reproduction of a dominant crop field earthworm (Drawida willsi, Michaelsen) was assessed following the application of two recommended agricultural doses of malathion (2.2 mg kg^–1– single dose and 4.4 mg kg^–1– double dose) in a rice field agroecosystem for a period of 105 days. The average worm population and biomass showed 12% and 21% decline, respectively, in plots treated with single and double doses of malathion, compared with control. The peak of worm population was observed after 60 days in control (plot I) and this peak value was shifted by 15 days (i.e. observed after 75 days) in malathion-treated plots (plots II and III) indicating delay in growth and maturation of the worms following the application of malathion. Inhibition in the total number of cocoons produced, and thus in the rate of reproduction, was also observed in malathion-treated plots. The peak rate of reproduction (1.71 cocoons adult^–1) was observed after 60 days in control plots and after 75 days (1.57 and 1.40 cocoons adult^–1) in single and double dose malathion-treated plots, respectively. One-way analysis of variance showed significant differences in the population size and biomass of D. willsi up to 75 days, in rate of reproduction up to 90 days, and no difference thereafter. This indicates that, under field conditions, D. willsi worms took about 75–90 days after application of malathion to resume normal population, biomass and reproduction, and hence we suggest that a second application of malathion in single and double agricultural doses should be avoided before this time. Received: 7 September 1999     

Accumulation of zinc and its effects on the growth, reproduction and life cycle of Drawida willsi (Oligochaeta), a dominant earthworm in Indian crop fields

 Accumulation of Zn and its effects on the growth, reproduction and life cycle of the earthworm Drawida willsi were determined. D. willsi did not reveal any significant changes in their mass at any of the concentrations of Zn (50, 200 and 400 mg kg^–1) compared to in untreated soils. The Zn concentrations in the exposed earthworms were significantly increased, but they were able to regulate their body content of Zn within a range of 116–125 mg kg^–1 (dry wt) in 200–400 mg kg^–1 Zn-treated soil. Reproduction was significantly reduced when the Zn concentration in soil exceeded 200 mg kg^–1. The drop in reproduction at elevated concentrations of Zn apparently resulted in a delay in completion of the life cycle and a decline in the total population. Received: 9 September 1998     

Computer-assisted tomography of macroporosity and its application to study the activity of the earthworm Aporrectodea nocturna

In a pre-alpine meadow in the Toggenburg (Switzerland), casts of the earthworm Aporrectodea nocturna, initially detected 20 years ago around a newly planted hedge, now extend 170 m from the hedge. The abundance of A. nocturma decreased between 140 and 170 m from the hedge from 237 (site: An+) to 0 (site: An?) individuals m^?2. The worm's activity and the macroporosity it has created have been studied using CAT scanning followed by automated image analysis of pores from 1·5 to 9 mm diameter. Brightness values in images from CAT scanning showed distinct local frequency maxima for stones, soil matrix and macropores. Measured diameters of pores were strongly linearly correlated with diameters of needles used to produce artificial pores. Pores from 1·5 to 3 mm were most abundant. The maximum number of pores from 1·5 to 9 mm (about 1600 m^?2) was larger than the maximum number of all earthworms (about 480 m^?2). Modelling the cast production of A. nocturna indicated that 23·8 kg m^?2 were deposited on the soil's surface within 3 months. The measured macroporosity (1·5–6 mm) was one-quarter to one-sixth of the pore volume corresponding to the removed casts. Thus, net production of pores by earthworms was a result both of the burrowing activity and of the refilling with eroded cast material. Depth distributions of bulk density, total porosity and pores from 1·5 to 9 mm were different at An+ from those at An?. However, diffusion of butane was equal at An+ and An?. Three-dimensional reconstruction suggested that the disposition (continuity, interconnectivity) of pores was more important for gas diffusion than the structure (size distribution, frequency) and bulk soil parameters.     

Simultaneous effects of plants and earthworms on mineralisation of 15N-labelled organic compounds adsorbed onto soil size fractions

 The simultaneous impact of three successive crops of wheat (Triticum aestivum L.) and of the earthworm (Lumbricus terrestris L.) on the mineralisation of ¹⁵N-labelled organic compounds adsorbed to different soil size fractions (sand and organic residues >50 μm; silt 50–2 μm; coarse clay 2–0.2 μm and fine clay <0.2 μm) was studied under controlled conditions in the greenhouse. Unplanted soils (UPS) were used as controls. In planted soils without earthworm (PS) total plant biomass decreased with each cropping by up to 50%. However, in planted soils with earthworms (PES) the total plant biomass loss was only 17%. This pattern was explained by the earthworm effect. Compared to the unplanted soils, the planted soils had an increased (mean +37%) mineralisation of ¹⁵N adsorbed onto fine clays and a partial transfer of ¹⁵N to silt and coarse clay. The quantities of ¹⁵N mineralised and transferred were higher in the planted soils with earthworms, indicating an amplification of the phenomenon in the presence of earthworms. The simultaneous effect of the rhizosphere and the drilosphere did not lead to increased mineralisation of N adsorbed onto coarse clays and silts but instead a greater transfer of N associated with the fine fractions towards the coarser fractions. Received: 25 April 2000     

Impact of earthworms on the diversity of microarthropods in a vertisol (Martinique)

In a study of a 15-year-old pasture in Martinique (French West Indies), abundance and organization of microarthropod communities were correlated with the spatial distribution of the earthworm Polypheretima elongata (Megascolecidae). In patches of high earthworm density (133 individuals m^–2), microarthropod density was significantly higher (80000 individuals m^–2) than in patches with few earthworms (31 worms m^–2 and 49000 microarthropods m^{– 2}). The diversity of microarthropod communities followed a similar pattern, the Shannon index for Collembola communities being, respectively, 3.12 and 1.82 in and outside earthworm patches. These results suggest that mesofauna abundance and diversity might be at least partly determined by the activity of larger invertebrates, as a result of the dramatic effects that the latter group exerts upon soil structure, pore distribution and food resources. Received: 7 February 1997     

Burrow constructions during the development of Lumbricus badensis individuals

Summary Juveniles of the anecic earthworm species Lumbricus badensis hatch from cocoon chambers, which the adult has excavated at depths between 0.4 and 1.5 m. After hatching the young animals migrate to the soil surface through the burrow of the adult. Juveniles live epigeically in self-constructed horizontal tubes during the first growing season. After hibernating in the soil at depths between 30 and 50 cm, juveniles make U-shaped burrows in the mineral soil. Later a main tube is constructed penetrating into the soil. This tube is split into two branches forming a V near the soil surface, and resembling the burrow of the adult. The burrow of an adult earthworm may reach a depth of 2.5 m and ramifies into 5–7 outlet tubes near the soil surface. The construction of the burrows during an individual's development, together with the transition from the epigeic to the anecic way of life, shows the close relationship of the anecic Lumbricus species to epigeic types. The feeding behaviour of the adult and the construction of the humic wall confirm this close relationship.Dedicated to the late Prof. Dr. M.S. Ghilarov     

Spatial changes of soil fungal and bacterial biomass from a sub-alpine coniferous forest to grassland in a humid, sub-tropical region

 Fungal and bacterial biomass were determined across a gradient from a forest to grassland in a sub-alpine region in central Taiwan. The respiration-inhibition and ergosterol methods for the evaluation of the microbial biomass were compared. Soil fungal and bacterial biomass both significantly decreased (P<0.05) with the shift of vegetation from forest to grassland. Fungal and bacterial respiration rates (evolved CO₂) were, respectively, 89.1 μl CO₂ g^–1 soil h^–1 and 55.1 μl CO₂ g^–1 soil h^–1 in the forest and 36.7 μl CO₂ g^–1 soil h^–1 and 35.7 μl CO₂ g^–1 soil h^–1 in the grassland surface soils (0–10 cm). The fungal ergosterol content in the surface soil decreased from the forest zone (108 μg g^–1) to the grassland zone (15.9 μg g^–1). A good correlation (R ²=0.90) was exhibited between the soil fungal ergosterol content and soil fungal CO₂ production (respiration) for all sampling sites. For the forest and grassland soil profiles, microbial biomass (respiration and ergosterol) declined dramatically with depth, ten- to 100-fold from the surface organic horizon to the deepest mineral horizon. With respect to fungal to bacterial ratios for the surface soil (0–10 cm), the forest zone had a significantly (P<0.05) higher ratio (1.65) than the grassland zone (1.05). However, there was no fungal to bacterial ratio trend from the surface horizon to the deeper mineral horizons of the soil profiles. Received: 30 March 2000