Free-living nematodes present several characteristics that have led to their use as bioindicators of soil quality. Analyzing the structure of nematofauna is a pertinent way to understand soil biological processes. Earthworms play an important role in soil biological functioning and organic matter dynamics. Their effects on soil nematofauna have seldom been studied. We studied the effect of the tropical endogeic earthworm, Pontoscolex corethrurus, on nematode community structure in a 5-month field mesocosm experiment conducted in Madagascar. Ten different treatments with or without earthworms and with or without organic residues (rice, soybean) were compared. Organic residues were applied on the soil surface or mixed with the soil. The abundance of nematodes (bacterial and fungal feeders) was higher in presence of P. corethrurus than in their absence. The type of plant residues as well as their localisation had significant effects on the abundance and composition of soil nematodes. The analysis of nematode community structure showed that earthworm activity led to an overall activation of the microbial compartment without specific stimulation of the bacterial or fungal compartment. 1. Introduction Soil organisms play a leading role in decomposition and mineralization of organic matter (OM) [1]. They are involved in processes that affect carbon (C) sequestration as well as in the modification of soil physical structure and chemical properties. They also interact with other soil fauna and these interactions result in complex food webs [2]. Nematodes are small organisms (ca. 1?mm long at the adult stage) abundant in soil (several million m ? 2 soil), they present a high species diversity (about 11,000 species have already been described). Nematodes live in the film of water between soil particles and present various feeding behaviours. During the last twenty years, many studies have been conducted on these microfaunal organisms because they can be an efficient tool to assess soil quality and soil biological functioning [2–4]. Because they are present at different levels of the soil food web and present variable tolerance toward stress, nematofauna provide information about OM decomposition pathways and soil pollution status [3, 5–7]. Nematodes interact with other soil organisms including earthworms, which also play an important role in soil biological functioning and OM dynamics [8, 9]. Until now, studies on interactions between nematodes and earthworms have focused on the contribution of earthworm burrowing and casting activity to nematofauna abundance,
References
[1]
P. Lavelle and A. Spain, Soil Ecology, Kluwer Academic Publishers, Dordrecht, The Netherlands, 2001.
[2]
H. Ferris, T. Bongers, and R. G. M. de Goede, “A framework for soil food web diagnostics: extension of the nematode faunal analysis concept,” Applied Soil Ecology, vol. 18, no. 1, pp. 13–29, 2001.
[3]
T. Bongers and H. Ferris, “Nematode community structure as a bioindicator in environmental monitoring,” Trends in Ecology and Evolution, vol. 14, no. 6, pp. 224–228, 1999.
[4]
K. Ritz and D. L. Trudgill, “Utility of nematode community analysis as an integrated measure of the functional state of soils: perspectives and challenges: discussion paper,” Plant and Soil, vol. 212, no. 1, pp. 1–11, 1999.
[5]
T. Bongers and M. Bongers, “Functional diversity of nematodes,” Applied Soil Ecology, vol. 10, no. 3, pp. 239–251, 1998.
[6]
D. W. Freckman and E. P. Caswell, “The ecology of nematodes in agroecosystems,” Annual Review of Phytopathology, vol. 23, pp. 275–296, 1985.
[7]
G. W. Yeates and T. Bongers, “Nematode diversity in agroecosystems,” Agriculture, Ecosystems and Environment, vol. 74, no. 1–3, pp. 113–135, 1999.
[8]
T. Desjardins, F. Charpentier, B. Pashanasi, A. Pando-Bahuon, P. Lavelle, and A. Mariotti, “Effects of earthworm inoculation on soil organic matter dynamics of a cultivated ultisol,” Pedobiologia, vol. 47, no. 5-6, pp. 835–841, 2003.
[9]
C. Villenave, F. Charpentier, P. Lavelle, et al., “Effects of earthworms on soil organic matter and nutrient dynamics following earthworm inoculation in field experimental situations,” in Earthworm Management in Tropical Agroecosystems, P. Lavelle, L. Brussaard, and P. Hendrix, Eds., pp. 173–197, Cabi, New York, NY, USA, 1999.
[10]
M. Aira, F. Monroy, and J. Domínguez, “Effects of two species of earthworms (Allolobophora spp.) on soil systems: a microfaunal and biochemical analysis,” Pedobiologia, vol. 47, no. 5-6, pp. 877–881, 2003.
[11]
M. Aira, F. Monroy, J. Domínguez, and S. Mato, “How earthworm density affects microbial biomas and activity in pig manure,” European Journal of Soil Biology, vol. 38, no. 1, pp. 7–10, 2002.
[12]
R. Hyv?nen, S. Andersson, M. Clarholm, and T. Persson, “Effects of lumbricids and enchytraeids on nematodes in limed and unlimed coniferous mor humus,” Biology and Fertility of Soils, vol. 17, no. 3, pp. 201–205, 1994.
[13]
K. Ilieva-Makulec and G. Makulec, “Effect of the earthworm Lumbricus rubellus on the nematode community in a peat meadow soil,” European Journal of Soil Biology, vol. 38, no. 1, pp. 59–62, 2002.
[14]
A. V. Tiunov, M. Bonkowski, M. Bonkowski, J. A. Tiunov, and S. Scheu, “Microflora, protozoa and nematoda in Lumbricus terrestris burrow walls: a laboratory experiment,” Pedobiologia, vol. 45, no. 1, pp. 46–60, 2001.
[15]
T. M. Razafimbelo, “Stockage et protection du carbone dans un sol ferrallitique sous systèmes en semis direct avec couverture végétale des hautes terres malgaches,” Montpellier, 2005.
[16]
S. Coq, B. G. Barthès, R. Oliver, B. Rabary, and E. Blanchart, “Earthworm activity affects soil aggregation and organic matter dynamics according to the quality and localization of crop residues—an experimental study (Madagascar),” Soil Biology and Biochemistry, vol. 39, no. 8, pp. 2119–2128, 2007.
[17]
G. D. Bending, M. K. Turner, and J. E. Jones, “Interactions between crop residue and soil organic matter quality and the functional diversity of soil microbial communities,” Soil Biology and Biochemistry, vol. 34, no. 8, pp. 1073–1082, 2002.
[18]
C. Villenave, T. Bongers, K. Ekschmitt, P. Fernandes, and R. Oliver, “Changes in nematode communities after manuring in millet fields in Senegal,” Nematology, vol. 5, no. 3, pp. 351–358, 2003.
[19]
C. Villenave, K. Ekschmitt, S. Nazaret, and T. Bongers, “Interactions between nematodes and microbial communities in a tropical soil following manipulation of the soil food web,” Soil Biology & Biochemistry, vol. 36, no. 12, pp. 2033–2043, 2004.
[20]
D. W. Freckman, “Bacterivorous nematodes and organic-matter decomposition,” Agriculture, Ecosystems and Environment, vol. 24, no. 1–3, pp. 195–217, 1988.
[21]
R. Lenz and G. Eisenbeis, “The vertical distribution of decomposition activity and of litter-colonizing nematodes in soils under different tillage,” Pedobiologia, vol. 42, no. 3, pp. 193–204, 1998.
[22]
C. Fragoso, G. G. Brown, J. C. Patron, et al., “Agricultural intensification, soil biodiversity and agroecosystem function in the tropics: the role of earthworms,” Applied Soil Ecology, vol. 6, no. 1, pp. 17–35, 1997.
[23]
S. Abiven, S. Recous, V. Reyes, and R. Oliver, “Mineralisation of C and N from root, stem and leaf residues in soil and role of their biochemical quality,” Biology and Fertility of Soils, vol. 42, no. 2, pp. 119–128, 2005.
[24]
J. T. Gilmour, A. Mauromoustakos, P. M. Gale, and R. J. Norman, “Kinetics of crop residue decomposition: variability among crops and years,” Soil Science Society of America Journal, vol. 62, no. 3, pp. 750–755, 1998.
[25]
J. J. s'Jacobb and J. van Bezooijen, A Manual for Practical Work in Nematology, Department of Nematology, Agricultural University, Wageningen, The Netherlands, 1986.
[26]
G. W. Yeates, T. Bongers, R. G. M. de Goede, D. W. Freckman, and S. S. Georgieva, “Feeding habits in soil nematode families and genera—an outline for soil ecologists,” Journal of Nematology, vol. 25, no. 3, pp. 315–331, 1993.
[27]
J. Domínguez, R. W. Parmelee, and C. A. Edwards, “Interactions between Eisenia andrei (Oligochaeta) and nematode populations during vermicomposting,” Pedobiologia, vol. 47, no. 1, pp. 53–60, 2003.
[28]
E. Blanchart, A. Albrecht, J. Alegre, et al., “Effects of earthworms on soil structure and physical properties,” in Earthworm Management in Tropical Agroecosystems, P. Lavelle, L. Brussaard, and P. Hendrix, Eds., pp. 149–172, Cabi, New York, NY, USA, 1999.
[29]
I. Barois and P. Lavelle, “Changes in respiration rate and some physicochemical properties of a tropical soil during transit through Pontoscolex corethrurus (glossoscolecidae, oligochaeta),” Soil Biology and Biochemistry, vol. 18, no. 5, pp. 539–541, 1986.
[30]
Y. Araujo, F. J. Luiz?o, and E. Barros, “Effect of earthworm addition on soil nitrogen availability, microbial biomass and litter decomposition in mesocosms,” Biology and Fertility of Soils, vol. 39, no. 3, pp. 146–152, 2004.
[31]
H. Ferris and M. M. Matute, “Structural and functional succession in the nematode fauna of a soil food web,” Applied Soil Ecology, vol. 23, no. 2, pp. 93–110, 2003.
[32]
D. Djigal, A. Brauman, T. A. Diop, J. L. Chotte, and C. Villenave, “Influence of bacterial-feeding nematodes (Cephalobidae) on soil microbial communities during maize growth,” Soil Biology and Biochemistry, vol. 36, no. 2, pp. 323–331, 2004.
[33]
G. W. Yeates, “Nematodes as soil indicators: functional and biodiversity aspects,” Biology and Fertility of Soils, vol. 37, no. 4, pp. 199–210, 2003.
[34]
L. Wasilewska, “Participation of soil nematodes in grass litter decomposition under diverse biocenotic conditions of meadows,” Ekologia Polska, vol. 40, pp. 75–100, 1992.
[35]
C. J. Wright and D. C. Coleman, “Cross-site comparison of soil microbial biomass, soil nutrient status, and nematode trophic groups,” Pedobiologia, vol. 44, no. 1, pp. 2–23, 2000.
[36]
L. Bj?rnlund and S. Christensen, “How does litter quality and site heterogeneity interact on decomposer food webs of a semi-natural forest?” Soil Biology and Biochemistry, vol. 37, no. 2, pp. 203–213, 2005.
[37]
M. J. Vreeken-Buijs and L. Brussaard, “Soil mesofauna dynamics, wheat residue decomposition and nitrogen mineralization in buried litterbags,” Biology and Fertility of Soils, vol. 23, no. 4, pp. 374–381, 1996.
[38]
D. L. Porazinska, L. W. Duncan, R. McSorley, and J. H. Graham, “Nematode communities as indicators of status and processes of a soil ecosystem influenced by agricultural management practices,” Applied Soil Ecology, vol. 13, no. 1, pp. 69–86, 1999.