Abstract
The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.
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References
Achard F, Eva HD, Stibig H-J, Mayaux P, Callego J, Richards T, Malingreau J-P (2002) Determination of deforestation rates of the world’s humid tropical forests. Science 297:999–1002
Armbrecht I, Perfecto I, Vandermeer J (2004) Enigmatic biodiversity correlations: ant diversity responds to diverse resources. Science 304:284–286
Armbrecht I, Rivera L, Perfecto I (2005) Reduced diversity and complexity in the leaf-litter ant assemblage of Colombian coffee plantations. Conserv Biol 19:897–907
Basset Y (2001) Invertebrates in the canopy of tropical rain forest. How much do we really know? Plant Ecol 153:87–107
Belsky JM, Siebert SF (2003) Cultivating cacao: implications of sun-grown cacao on local food security and environmental stability. Agric Human Values 20:270–285
Bolton B (1994) Identification guide to the ant genera of the world. Harvard University Press, Cambridge, Massachusetts, USA
Colwell RK (2004) EstimateS: statistical estimation of species richness and shared species from samples. Version 7. User’s Guide and application. http://purl.oclc.org/estimates
Colwell RK, Coddington JA (1994) Estimating terrestrial biodiversity through extrapolation. Philos Trans R Soc Lond B 345:101–118
Donald PF (2004) Biodiversity impacts of some agricultural commodity production systems. Conserv Biol 18:17–37
Erwin TL (1982) Tropical forests: their richness in Coleoptera and other arthropod species. Coleopt Bull 36:74–75
Gibb H, Hochuli DF (2003) Colonisation by a dominant ant facilitated by anthropogenic disturbance: effects on ant assemblage composition, biomass and resource use. Oikos 103:469
Gradstein SR, Kessler M, Pitopang R (2007) Tree species diversity relative to human land uses in tropical rain forest margins in Central Sulawesi. In: Tscharntke T, Leuschner C, Guhardja E, Zeller M (eds) The stability of tropical rainforest margins: linking ecological, economic and social constraints of land use and conservation. Springer, Berlin, Heidelberg, New York, pp 321–334
Johns ND (1999) Conservation in Brazil’s chocolate forest: the unlikely persistence of the traditional cocoa agroecosystem. Environ Manage 23:31–47
Klein A-M, Steffan-Dewenter I, Buchori D, Tscharntke T (2002) Effects of land-use intensity in tropical agroforestry systems on coffee flower-visiting and trap-nesting bees and wasps. Conserv Biol 16:1003–1014
Lawton JH, Bignell DE, Bolton B, Bloemers GF, Eggleton P, Hammond PM, Hodda M, Holt RD, Larsen TB, Mawsley NA, Stork NE, Srivastava DS, Watt AD (1998) Biodiversity inventories, indicator taxa and effects of habitat modification in tropical forest. Nature 391:72–76
Majer JD (1972) The ant mosaic in Ghana cocoa farms. Bull Entomol Res 62:151–160
Majer JD (1976) The influence of ants and ant manipulation on the cocoa farm fauna. J Appl Ecol 13:157–175
Majer JD, Delabie JHC, Smith MRB (1994) Arboreal ant community patterns in Brazilian cocoa farms. Biotropica 26:73–83
McKinney ML, Lockwood JL (1999) Biotic homogenization: a few winners replacing many losers in the next mass extinction. Tree 14:450–453
Myers N, Mittelmeier RA, Mittelmeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 493:853–858
O’Dowd DJ, Green PT, Lake PS (2003) Invasional ‘meltdown’ on an oceanic island. Ecol Lett 6:812–817
Perfecto I, Mas A, Dietsch T, Vandermeer J (2003) Conservation of biodiversity in coffee agroecosystems: a tri-taxa comparison in southern Mexico. Biodivers Conserv 12:1239–1252
Perfecto I, Vandermeer J (1996) Microclimatic changes and the indirect loss of ant diversity in a tropical agroecosystem. Oecologia 108:577–582
Perfecto I, Vandermeer J, Hanson P, Cartín V (1997) Arthropod biodiversity loss and the transformation of a tropical agro-ecosystem. Biodivers Conserv 6:935–945
Philpott SM, Foster PF (2005) Nest-site limitation in coffee agroecosystems: artificial nests maintain diversity of arboreal ants. Ecol Appl 15:1478–1485
Pineda E, Moreno C, Escobar F, Halffter G (2005) Frog, bat, and dung beetle diversity in the cloud forest and coffee agroecosystems of Vercruz, Mexico. Conserv Biol 19:400–410
Potter L (2001) Agricultural intensification in Indonesia: outside pressures and indigenous strategies. Asia Pac Viewp 42:305–324
Rice RA, Greenberg R (2000) Cacao cultivation and the conservation of biological diversity. Ambio 29:167–172
Room PM (1971) The relative distribution of ant species in Ghana’s cocoa farms. J Anim Ecol 40:735–751
Schroth G, da Fonseca GAB, Harvey CA, Gascon C, Vasconcelos HL, Izac A-MN (2004) Agroforestry and biodiversity conservation in tropical landscapes. Island Press, Washington, DC
Schulze CH, Waltert M, Kessler PJA, Pitopang R, Shahabuddin, Veddeler D, Muhlenberg M, Gradstein SR, Leuschner C, Steffan-Dewenter I, Tscharntke T (2004) Biodiversity indicator groups of tropical land-use systems: comparing plants, birds and insects. Ecol Appl 14:1321–1333
Shahabuddin, Schulze CH, Tscharntke T (2005) Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia). Biodivers Conserv 14:863–877
StatSoft Inc. 1984–2004. STATISTICA for Windows [Software-System for Data-analyses] Version 7.0. Tulsa, USA
Waltert M, Mardiastuti A, Muehlenberg M (2004) Effects of land use on bird species richness in Sulawesi, Indonesia. Conserv Biol 18:1339–1346
Zuidema PA, Leffelaar PA, Gerritsma W, Mommer L, Anten NPR (2005) A physiological production model for cocoa (Theobroma cacao): model presentation, validation and application. Agric Syst 84:195–225
Acknowledgments
This study was funded by the German Research Foundation (DFG), grant SFB-552 “STORMA”. We thank Christian H. Schulze and Simone G. Sporn for valuable suggestions during the fieldwork. Further, we thank Goetz Schroth, Celia Harvey, Jason M. Tylianakis, Tatyana Rand and 3 anonymous referees for their valuable suggestions that improved previous versions of this manuscript. We are very grateful to Boris Büche, Christoph Bayer, Rudy Kohout and Akhmad Rizali for their detailed and careful identification and sorting of ant and beetle species. We thank Ramadanil Pitopang (Herbarium Celebense, Indonesia) for identifying forest trees, Damayanti Buchori (IPB Bogor, Indonesia), Pak Mann, Arifin, Daniel Stietenroth, Adam Malik, Wolfram Lorenz and Surya Tarigan for their help in this work, and the following coleopterists for their identifications after visiting www.beetle-diversity.com: Alberto Ballerio, Roland Gerstmeier, Denis Keith, Otto Merkl, Chris Reid, Wolfgang Schawaller, Dmitry Telnov and Jürgen Wiesner.
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Bos, M.M., Steffan-Dewenter, I. & Tscharntke, T. The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia. Biodivers Conserv 16, 2429–2444 (2007). https://doi.org/10.1007/s10531-007-9196-0
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DOI: https://doi.org/10.1007/s10531-007-9196-0
Keywords
- Arthropods
- Biodiversity
- Cultivated land
- Deforestation
- Habitat preference
- Knockdown fogging
- Lower canopy