Jean-François Ponge

Humus forms in terrestrial ecosystems : a framework to biodiversity

Humus forms are the seat of most biological transformations taking place in terrestrial ecosystems, being at the interface between plants, animals and microbes. The diversity of terrestrial humus forms (mor, moder and mull) can be attributed to the existence of different patterns (strategies) for the capture and use of resources by ecosystems, in ascending order of biodiversity and bioavailability. Arguments are found in the parallel development of humus forms and terrestrial ecosystems, in exclusion mechanisms between organisms living in different humus forms, and in palaeontological studies. The diversification of terrestrial life forms in the course of Earth history, concomitant with an improvement in resource availability due to the development of sedimentary layers at the surface of continents, may explain the successive appearance of more active humus forms enabling the co-existence of an increasing number of organisms. Contradictory reports about the relationships between biodiversity and stability of ecosystems can be explained by the existence of different belowground pathways making ecosystems more stable.

Humus form dynamics during the sylvogenetic cycle in a mountain spruce forest

Abstract The humus forms during the life cycle of a spruce forest are described. A significant change in humus form may be attributed to plant and soil fauna changes. This phenomenon is considered to be fundamental for the renewal of the forest ecosystem. Forest dynamics is perceived as a biphasic cycle, (i) the tree growth phase with a shift from mull towards moder humus form, as a consequence of a decline in earthworm populations and (ii) a humus form improvement from moder towards earthworm mull humus, during the second half of the life of spruce trees. This results from a succession of earthworm species. The particular role of anecic species during the second phase has been highlighted, where they allow endogeic earthworm species and young spruce seedlings to install themselves in the regeneration site, the fall of parent trees not being considered as the chief factor governing humus changes. The life cycle of the spruce ecosystem can nevertheless be impaired by the development of a bilberry heath, with a mor humus form which is detrimental to the germination and growth of spruce seedlings. Earthworm populations of anecic and endogeic species are present in this case but without any burrowing activity.

Succession of fungi and fauna during decomposition of needles in a small area of Scots pine litter

During micromorphological investigations on Scots pine litter, several decomposition stages have been recognized on fallen pine needles, each being associated with the activity of animal and microbial organisms, both. To well-known fungal successions that have been so far described by mycologists we must add succession of animal groups such as nematodes, amoebae, enchytraeids, sciarid larvae, oribatid mites and earthworms. A bacterial development was observed in the L2 layer, following penetration by microfauna (nematodes, amoebae). After that stage pine needles were actively tunnelled by enchytraeids, sciarid larvae and oribatid mites and at the same time were nibbled on by epigeic earthworms (L2 and F1 layers). When the fine root system of pine developed through accumulated old needles (F1 layer), mycorrhizal fungi penetrated the needles and seemed to impede any further bacterial development. Pine foliar tissues were progressively incorporated into the fecal material of earthworms and other members of the soil fauna. A more realistic scheme was suggested for plant litter decomposition in moder humus.

Food resources and diets of soil animals in a small area of Scots pine litter

Abstract The fauna inhabiting a small area (ca. 5 cm × 5 cm) were investigated in a Scots pine stand. After microstratification of the litter layers in the field and fixation in 95% ethyl alcohol, invertebrates, mainly mesofauna, were sorted under a dissecting microscope and mounted or dissected in order to study their intestinal guts. Faeces were mounted or sectioned to obtain information about the activity of other invertebrate groups not represented in the sample and to follow the fate of plant and microbial material after defaecation occurred. Plant material, mainly from moss, bracken, pine needles and bark, was extensively consumed by enchytraeid and lumbricid worms, sciarid larvae and phthiracarid mites. Fungal material was ingested by all groups, either in combination with plant material or alone (camisiid and oppiid mites, some species of Collembola, sciarid and chironomid larvae). Isotomid springtails and chironomid larvae appeared to consume faecal material. The choice and the degree of comminution and digestion of the material differed greatly from one group to another, but without any indication of resource sharing.

Collembolan communities as bioindicators of land use intensification

Springtail communities (Hexapoda: Collembola) were sampled in the Morvan Nature Regional Park (Burgundy, France) in six land use units (LUUs) 1 km2 each, which had been selected in order to cover a range of increasing intensity of land use. Human influence increased from LUU 1 (old deciduous forest) to LUU 6 (agricultural land mainly devoted to cereal crops), passing by planted coniferous forests (LUU 2) and variegated landscapes made of cereal crops, pastures, hay meadows, conifer plantations and small relict deciduous groves in varying proportion (LUUs 3–5). Sixteen core samples were taken inside each LUU, at intersections of a regular grid. Species composition, species richness and total abundance of collembolan communities varied according to land use and landscape properties. Land use types affected these communities through changes in the degree of opening of woody landscape (woodland opposed to grassland) and changes in humus forms (measured by the Humus Index). A decrease in species richness and total abundance was observed from old deciduous forests to cereal crops. Although the regional species richness was not affected by the intensification gradient (40–50 species were recorded in every LUU), a marked decrease in local biodiversity was observed when the variety of land use types increased. In variegated landscapes the observed collapse in local species richness was not due to a different distribution of land use types, since it affected mainly woodland areas. Results indicated the detrimental influence of the rapid afforestation of previous agricultural land, which did not afford time for the development of better adapted soil animal communities.

The Forest Regeneration Puzzle Biological mechanisms in humus layer and forest vegetation dynamics

Forest ecosystem dynamics have long been explained by changes in floristic composition (Foster 1988) and nutrient availability (Vitousek 1984). The passage from pioneer to late-successional plant communities has frequently been modeled (McCook 1994) and, in a few instances, has been studied experimentally (Facelli and Facelli 1993). Interactions between plants (including their immediate environment as a passive partner) have generally been considered to be responsible for forest succession. However, the belowground components of any given ecosystem must also be taken into account before successional and steady-state patterns in plant communities can be fully understood (Cromack 1981, Miles 1985, Nilsson 1994). For example, managed coniferous forests

Vertical distribution of Collembola (Hexapoda) and their food resources in organic horizons of beech forests

Abstract Micro-samples of the surface organic horizons of 13 beech forests in Belgium were fixed immediately after collection in ethanol. Collembola (6255 animals) were sorted directly from micro-samples in the laboratory using a dissecting microscope, while the litter/soil matrix was analysed semi-quantitatively. The vertical distribution of Collembolan species was studied by correspondence analysis. Gut contents of animals were examined under a light microscope and their composition was compared with that of the matrix. A consistent association was found between the vertical distribution of gut contents and that of food resources in the immediate proximity of animals. Species differed in their feeding habits but most of them ingested a wide spectrum of food items. Plasticity in the food regime according to depth could be demonstrated in members of the Onychiuridae family.

The heterogeneity of humus profiles and earthworm communities in a virgin beech forest

Thirty sites, encompassing a range of soil and vegetation conditions in the biological reserve of La Tillaie (Fontainebleau Forest, France) were investigated in April 1992. Beech (Fagus sylvatica L.) was the dominant tree species, with several developmental phases forming the forest patchwork. Sessile oak [Quercus petraea (Mattus.) Liebl.] was present but only as old relictual individuals. Gaps in the canopy cover were abundant, mostly caused by wind storms 2 years previously. The next most recent storm was 25 years before, resulting in distinct patches of full-grown trees. Humus profiles were classified and compared with the distribution of earthworm communities, canopy cover, and soil types. Geomorphology was responsible for the main part of the observed variation. Absence of lime in the substrate and direct contact with a sandstone stratum near the ground surface was associated with the absence of earthworms and the appearance of an OH horizon (moder humus). Elsewhere, earthworms were present and humus profiles did not display any OH horizon (mull or mull-like moder humus), but species composition was variable and strongly influenced by the thickness of the superficial sand deposit overlying limestone. On a thick (1 m or more) sandy substrate earthworm communities were dominated by epigeic species together with the anecic Lumbricus terrestris L. The species richness was higher on a shallower sandy substrate (50 cm) where lime was more accessible to tree roots and burrowing animals. The influence of the forest cycle of beech was visible in the latter case (covering most of the area), with an increase in the thickness of the OL and OF horizons and a decrease in endogeic earthworm populations during the phase of intense growth of beech. This fall in burrowing activity was apparent in gaps created by wind storms and fungal diseases within mature stands as early as 2 years after the fall of the trees.

Manioc peel and charcoal: a potential organic amendment for sustainable soil fertility in the tropics

In tropical areas, where crop production is limited by low soil quality, the development of techniques improving soil fertility without damage to the environment is a priority. In French Guiana, we used subsistence farmer plots on poor acidic soils to test the effect of different organic amendments, bitter manioc peel (M), sawdust (Sw) and charcoal (Ch), on soil nutrient content, earthworm abundance and yard-long bean (Vigna unguiculata sesquipedalis) production. The peregrine Pontoscolex corethrurus was the only earthworm species found. Pod production and plant growth were lowest in unamended soil. The application of a mixture of manioc peel and charcoal (M + Ch) improved legume production compared with other organic mixtures. It combined the favourable effects of manioc peel and charcoal. Manioc peel improved soil fertility through its low C:N ratio and its high P content, while charcoal decreased soil acidity and exchangeable Al and increased Ca and Mg availability, thus alleviating the possible toxic effects of Al on plant growth. The M + Ch treatment was favourable to P. corethrurus, the juvenile population of which reached a size comparable to that of the nearby uncultivated soil. The application of a mixture of manioc peel and charcoal, by improving crop production and soil fertility and enhancing earthworm activity, could be a potentially efficient organic manure for legume production in tropical areas where manioc is cultivated under slash-and-burn shifting agriculture.

Earthworm and enchytraeid activity under different arable farming systems, as exemplified by biogenic structures.

A study was conducted in order to compare soil faunal activity in four experimental farming systems using different tillage, chemical input and crop rotation practices: A conventional system with deep-ploughing (CT), an integrated system with reduced tillage and minimum chemical input (IN), a system with reduced tillage and high chemical input (RT) and a system with minimum tillage and high chemical input (MT). In nine experimental fields with two sampling points each, earthworms were sampled and biogenic structures were identified and counted in topsoil profiles (0–14 cm depth). Components of these profiles were identified by morphological features. Quantitative analyses of these morphological features provided information about soil compaction, earthworm and enchytraeid activity and distribution of roots and crop residues in the soil matrix. The dominant species in the earthworm community was the endogeic Aporrectodea rosea. Earthworm densities were unexpectedly lowest under reduced tillage (6 specimens per m2), and highest under deep-ploughing (67 specimen per m2), the reverse effect being observed with enchytraeid worms, as ascertained by deposition of their faecal pellets in topsoil profiles. Strong very fine granular structure (STVFGR) was most frequent in the integrated farming system (IN). We concluded that in the studied site embracing four farming systems, enchytraeids play an important role in creating a stable soil structure and porosity at the low level of earthworm densities found in the integrated system (IN).