Laode Muhammad Harjoni Kilowasid, Tati Suryati Syamsudin, Endah Sulystiawati, Fransiscus-Xaverius Susilo


An understanding of the structure of the soil food web is critical in determining the practices of soil fertility management based on the biological processes in tropical agricultural regions. The objectives of the study were to assess the variation in trophic level biomass and to analyze the dynamics of the energy channels on the increasing age of cocoa plantation. The characteristics of soil food web structure in smallholder cocoa plantation aged 4, 5, 7, 10, and 16 years were analyzed. The results showed that only biomass at the third trophic level increased with plantation age, but not for the biomass at the lower trophic levels. Biomass in all energy channels did not increased as well along with plantation age. We concluded that variation in the soil food web structure was more influenced by biotic factors of macro-arthropods group, such as facilitation, recolonization capabilities and accessibility in the soil habitat of smallholder cocoa plantation.


Keyword: biomass, biotic, energy channel, trophic level.     

Full Text:



Adl, S.M., D. Acosta-Mercado and D.H. Lynn. 2008. Protozoa. In: Carter, M.R. and E.G. Gregorich. 2nd edition. (Ed.), Soil sampling and methods of analysis, Canadian Society of Soil Science, Inc. p. 455 – 469.

Aira, M., L.Sampedro, F. Monroy and J. Dominguez. 2008. Detrivorous earthworms directly modify the structure, thus altering the functioning of a microdecomposer food web. Soil Biology & Biochemistry 40: 2511 – 2516.

Bakken, L. R. and R.A. Olsen. 1985. Buoyant densities and dry-matter content of microorganisma: conversion of a measured biovolume into biomass. Applied and Environmental Microbiology 45 (4): 1188 – 1195.

Bardgett, R.D. and D.A. Wardle. 2010. Aboveground-belowground linkage: biotic interactions, ecosystem processes, and global change. Oxford University Press. New York, p.297.

Beare, M. H., R.W. Parmelee, P.F. Hendrix, W. Cheng, D.C. Coleman and D.A. Jr. Crossley. 1992. Microbial and faunal interactions and effects on litter nitrogen and decomposition in agroecosystems. Ecological Monographs 62 (4): 569 – 591.

Belay-Tedla, A., X. Zhou, B. Su, S. Wan and Y. Lou. 2009. Labile, recalcitrant, and microbial carbon and nitrogen pools of tallgrass prairie soils in the US great plains subjected to experimental warming and clipping. Soil Biology & Biochemistry 41: 110 – 116.

Berg, M., J.P. Kniese, J.J.M. Bedaux and H.A. Verhoef .1998. Dynamics and stratification of functional groups of micro- and mesoarthropods in the organic layer of a scots pine forest. Biol Fertil Soils 26: 268 – 284.

Berg, M., P.C. de Ruiter, W. Didden, M. Jansen, T. Schouten and H. Verhoef. 2001. Community food web, decomposition and nitrogen mineralisation in a stratified scots pine forest soil. Oikos 94: 130 – 142.

Bezemer, T.M, M.T. Fountain, J.M. Barea, S. Christensen, S.C. Dekker, H. Duyts, R. van Hal, J.A. Harvey, K. Hedlund, M. Maraun, J. Mikola, A.G. Mladenov, C. Robin, P.C. de Ruiter, S. Scheu, H. Setälä, P. Šmilauer and W.H. van der Putten. 2010. Divergent composition but similar function of soil food webs of individual plants: plant species and community effects. Ecology 91: 3027 – 3036.

Bonkowsky, M., C. Villenave and B. Griffiths. 2009. Rhizosphere fauna: the functional and structural diversity of intimate interactions of soil fauna with plant roots. Plant Soil 321: 213 – 233.

Brady, C.J and R.A. Noske. 2006. Generalised regressions provide good estimates of insect and spider biomass in the monsoonal tropics of Australia. Australian Journal of Entomology 45: 187 – 191.

Coleman, D.C. 2008. From peds to paradoxes: linkages between soil biota and their influences on ecological processes. Soil Biology & Biochemistry 40: 271 – 289.

Collins, P.T. 1991. Length-biomass relationships for terrestrial Gastropoda and Oligochaeta, Am.Midl. Nat. 128: 404 – 406.

Delabie, J.H.C., B. Jahyny, I.C. do Nascimento, C.S.F. Mariano, S. Lacau, S. Campiolo, S.M. Philpott, and M. Leponce. 2007. Contribution of cocoa plantations to the conservation of native ants (insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic forest fauna of southern Bahia, Brazil. Biodivers Conserv 16: 2359 – 2384.

de Ruiter, P.C., J. Bloem, L.A. Bouwman, W.A.M. Didden, G.H.J. Hoenderboom, G. Lebbink, J.C.Y. Marinissen, J.A. de Vos, M.J. Vreeken-Buijs and K.B. Zwart. 1994. Simulation of dynamics in nitrogen mineralization in the below-ground food webs of two arable farming system. Agric. Ecosyst. Environ. 51: 199 – 208.

Didden, W.A.M., J.C.Y. Marinessen, M.J. Vreeken-Buijs, S.L.G.E. Burgers, R. de Fluiter, M. Geurs and L. Brussaard. 1994. Soil meso-and macrofauna in two agricultural systems: factors affecting population dynamics and evaluation of their role in carbon and nitrogen dynamics. Agriculture, Ecosystems, and Environment 51: 171 – 186.

Directorate General of Esatate, Agricultural Ministry, RI. 2011. Area and production by category of producers. download: November 25. 2011.

Doblas-Miranda, E., F. Sánches-Piñero and A. Gonzalez-Megias. 2009. Different structuring factors but connected dynamics shape litter and below-ground soil macrofaunal food webs. Soil Biology & Biochemistry 41: 2543 – 2550.

Ekelund, F., R. Rønn and B. Griffths. 2001. Quantitative estimation of flagellate community structure and diversity in soil samples. Protist 152: 301 – 314.

Ferris, H. 2010. Form and function: metabolic footprints of nematodes in the soil food web. European Journal of Soil Biology 46: 97 – 104.

Ganihar, S.R. 1997. Biomass estimates of terrestrial arthropods based on body length. J. Biosci. 22(2): 219 – 224.

Gaspar, M.L., M.N. Cabello, R. Pollero and M.A. Aon. 2001. Fluorescein diacetate hydrolysis as a measure of fungal biomass in soil. Current Microbiology 42: 339 – 344.

Green, V.S., D.E. Stott, and M. Diack. 2006. Assay for fluorescein diacetate hydrolytic activity: Optimization for Soil Samples. Soil Biology & Biochemistry 38: 693 – 701.

Gruner, D.S. 2003. Regressions of length and width to predict arthropod biomass in the Hawaiian Islands. Pacific Science. 57(3): 325 – 336.

Hertel, D., M.A. Harteveld and C. Leuschner. 2009. Conversion of a tropical forest into agroforest alters the fine root-related carbon flux to the soil. Soil Biology and Biochemistry 41: 481 – 490.

Holtkamp. R., P. Kardol, A. van der Wal, S.C. Dekker, W.H. van der Putten, and P.C. de Ruiter. 2008. Soil food web during ecosystem development after land abandonment. Applied Soil Ecology. 39: 23 – 34.

Holtkamp, R., A. van der Wal, P. Kardol, W.H. van der Putten, P.C. de Ruiter, and S.C. Dekker. 2011. Modelling C and N mineralisation in soil food web during secondary succession on ex-arable land. Soil Biology & Biochemistry 43: 251 – 260.

Höfer, H., and R. Ott. 2009. Estimating biomass of Neotropical spiders and other arachnids (Araneae, Opiliones, Pseudoscorpiones, Ricinulei) by mass-length regression. The Journal of Arachnology 37: 160 – 169.

Hubbel, S.P. 2001. The unified neutral theory of biodiversity and biogeography. Princeton University Press. New Jersey.

Hunt, H.W., D.C. Coleman, E.R. Ingham, R.E. Ingham, E.T. Elliot, J.C. Moore, S.L. Rose, C.P.P, Reid and C.R. Morley. 1987. The detrital food web in a shortgrass prairie. Biol. Fertil. Soils. 3:57 – 68.

ICCO. 2010. International Cocoa Organization. London.

Kilowasid, L.M.H. 2012. Control of soil fauna community on nitrogen status in cocoa plantation. ITB Bandung. Dissertation. pp.92 - 93. (in Indonesia).

Kilowasid, L.M.H, T.S. Syamsudin, F.X. Susilo and E. Sulistyawati. 2012. Ecological Diversity of soil fauna as ecosystem engineers in small-holder cocoa plantation in South Konawe. J Trop Soils 17: 173 – 180.

Kilowasid, L.M.H, T.S. Syamsudin, F.X. Susilo, E. Sulistyawati and H. Syaf. 2013. Characteristics of soil fauna communities and habitat in small-holder cocoa plantation in South Konawe. J Trop Soils 18 (2): 149 – 159.

Klein, D.A, and M.W. Paschke. 2000. A soil microbial community structural-functional index: the microscopy-based total/active/active fungal/bacterial (TA/AFB) biovolumes ratio. Applied Soil Ecology 14: 257 – 268.

Laakso, J. 1999. Short-term effects of wood ants (Formica aquilonia Yarr.) on soil animal community structure. Soil Biology and Biochemistry 31: 337 – 343.

Li, Y., and G. González. 2008. Soil fungi and macrofauna in the neotropics. In Myster, R. W. (eds.). Post-Agricultural Succession in the Neotropics. Springer. p.93 – 114.

Lorange-Merciris, G., D. Imbert, F. Bernhard-Reversat, P. Lavelle and J.F. Ponge. 2008. Litter N-content influences soil milipede abundance, species richness and feeding preferences in a semi-evergreen dry forest of Guadeloupe (Lesser Antilles). Biol Fertil Soils 45:93 – 98.

Marhaning, A.R., A.A.S. Mills, and S.M. Adl. 2009. Soil community changes during secondary succession to naturalizaed grasslands. Applied Soil Ecology 41: 137 – 147.

Mikola, J and H. Setala. 1999. Interplay of omnivory, energy channels and C availability in a microbial-based soil food web. Biol Fertil Soils 28: 212 – 218.

Moco, M.K.S., E.F. Gama-Rodrigues, A.C. Gama-Rodrigues, R.C.R. Machado and V.C. Baligar. 2009. Soil and litter fauna of cocoa agroforestry systems in Bahia, Brazil. Agroforest Syst 76: 127 – 138.

Moore, J.C. and H.W. Hunt. 1988. Resource compartmentation and the stability of real ecosystem. Nature 333: 261 – 263.

Moore, J.C., D.E. Walter and W.H. Hunt. 1988. Arthtropod regulation of micro- and mesobiota in below-ground detrital food webs. Ann. Rev. Entomol. 33: 419 – 439.

Moore, J.C 1994. Impact of agricultural practices on soil food web structure: theory and application. Agriculture, Ecosystem and Environment 51:239 – 247.

Moore, J. C., E.L. Berlow, D.C. Coleman, P.C. de Ruiter, Q. Dong, A. Hastings, N.C. Johnson, K.S. McCann, K. Melville, P.J. Morin, K. Nadelhoffer, A.D. Rosemond,D.M. Post, J.L. Sabo, K.M. Scow, M.J. Vanni and D.H. Wall. 2004. Detritus, trophic dynamics and biodiversity. Ecology Letters 7: 584 – 600.

Moore, J.C., K. McCann and P.C. de Ruiter. 2005. Modeling trophic patways, nutrient cycling, and dynamic stability in soils. Pedobiologia 49: 499 – 510.

Moya-Larano, J and D.H. Wise. 2007. Direct and indirect effects of ants on a forest-floor food web. Ecology 88(6):1454 – 1465.

Muñoz, F., and J. Beer. 2001. Fine root dynamics of shaded cacao plantations in Costa Rica. Agroforestry Systems 51:119 –130.

Osler, G.H.R. and M. Sommerkorn. 2007. Toward a complete soil C and N cycle: incorporating the soil fauna. Ecology 88: 1611 – 1621.

Petchey, O. L., P.J. Morin, and H. Olff. 2010. The Topology of ecological interaction networks: the state of the art. In: Verhoef H.A. and P.J. Morin (eds.): Community ecology: processes, models, and applications. Oxford University Press. New York. p. 7 – 22.

Persson, T. and U. Lohm. 1977. Energetical significance of the annelids and arthropods in a Swedish grassland soil. Ecological Bulletins No. 23. Swedish Natural Science Research Council. p.210.

Rooney, N., K. McCann, G. Gellner and J.C. Moore. 2006. Structural asymmetry and the stability of diverse food webs. Nature 442: 265 – 269.

Rovira, P. and V.R. Vallejo. 2002. Labile and recalcitrant pool of carbon and nitrogen in organic matter decomposing at different depth in soil: an acid hydrolysis approach. Geoderma 107:109 – 141.

Rovira, P. and V.R. Vallejo. 2007. Labile, recalcitrant, and inert organic matter in mediterranean forest soils. Soil Biology & Biochemistry 39: 202 – 215.

Sabo, J.L., J.L. Bastow, and M.E. Power. 2002. Length-mass relationships for adult aquatic and terrestrial invertebrates in a California watershed. J. N. Am. Benthol. Soc. 21(2): 336 – 343.

Schimel, J.P. and J. Bennet. 2004. Nitrogen mineralization: challenges of a changing paradigm. Ecology 85: 591 – 602.

Schroter, D. V. Wolters, and P.C. de Ruiter. 2003. C and N mineralization in the decomposer food webs of a European forest transect. Oikos 102: 294 – 308.

Smiley, G. L. and J. Kroschel. 2008. Temporal change in carbon stock of cocoa-gliricidia agroforests of Central Sulawesi. Indonesia. Agroforest Syst. 73: 219 – 231.

Smiley, G.L. and J. Kroschel. 2009. Yield development and nutrient dynamics in cocoa-gliricidia agroforests of central Sulawesi. Indonesia. Agroforest Syst.

Stout, J.D. and O.W. Heal. 1967. Protozoa. In: Burges, A. and F. Raw (Eds.): Soil biology. Academic Press. New York. p. 149 – 195.

Sugihara, G., L.F. Bersier, and K. Schoenly. 1997. Effects of taxonomic and trophic aggregation on food web properties. Oecologia 112: 272 – 284.

Susilo, F.X., A.M. Neutal, van Noordwijk, K. Hairiah, G. Brown and M.J. Swift. 2004. Soil biodiversity and food webs. In: van Noordwijk, G. Cadisch, and C.K. Ong: Below-ground interactions in tropical agroecosystems: concept and models with multiple plant components. CAB International Publushing. p. 285 – 308

Swift, M.J. 1997. Biological management of soil fertility as a component of sustainable agriculture: perspectives and prospects with particular reference to tropical regions. In: Brussard, L. and R. Ferrera-Cerrato (ed.): Soil ecology in sustainable agricultural systems. CRC Press LLC. Boca Raton. Chapter 7.

Trolldenier, G. 1996. Bacterial biomass. In: Schinner, F., R. Öhlinger, E. Kandeler and R. Maegisen (Eds.) Methods in soil biology. Springer, Hong Kong. p.135 – 137

van Veen, J.A. and E.A. Paul. 1979. Conversion of biovolume measurements of soil organisms, grown under various tension, to biomass and their nutrient content. Applied and Environmental Microbiology 37: 686 – 692.

Viketoft, M., J. Bengtsson, B. Sohlenius, M.P. Berg, O. Petchey, C. Palmborg and K. Hus-Danell. 2009. Long-term effects of plant diversity and composition on soil nematode communities in model grasslands. Ecology 90(1): 90 – 99.

Wardle, D.A. 2006. The influence of biotic interactions on soil biodiversity. Ecology Letters 9: 870 – 886.

Wallwork, J.A. 1970. Ecology of soil animals. McGraraw-Hill. London.

Williams, R.J and N.D. Martinez. 2004. Limit to trophic levels and omnivory in complex food webs: Theory and data. The American Naturalist 163(3): 458 – 468.

Wise, D.H. 2004. Wandering spiders limit densities of a major microbial-detritivore in the forest-floor food web. Pedobiologia 48: 181 – 188.

Yeates, G.W. 1979. Soil nematodes in terrestrial ecosystems. J. Nematol. 11: 213-229.

Yeates, G.W., T. Bongers, R.G.M. De Goede, D.W. Freckman and S.S. Georgieva.1993. Feeding habits in soil nematode families and genera, An outline for soil ecologists. J. Nematol. 25: 315 – 331.