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Effects of environmental variables and foliar traits on the transpiration rate of cocoa (Theobroma cacao L.) under different cultivation systems


The response of plant species to environmental conditions influences changes in functional traits associated with the process that determines biological fitness and ecosystem processes. However, documenting these responses remain largely elusive in cultivation systems. We analyzed how environmental variables and leaf traits have effects on the transpiration rate of cocoa (Theobroma cacao L.) trees compared among different cultivation systems. Fieldwork was carried out at the Sara Ana experimental station in Alto Beni, La Paz, Bolivia. We sampled four trees in each of eight plots; four plots for each cultivation system (organic monoculture vs. organic agroforestry). From each tree, two mature, sunlit and healthy leaves were collected to make measurements of foliar traits and environmental variables. We found that canopy cover was higher in the agroforestry system. The specific leaf area was greater in agroforestry but the stomata size was significantly higher in the monoculture system. Temperature had a positive relationship with transpiration, whereas canopy cover and specific leaf area had a negative relationship in the agroforestry system. Cultivation system caused changes in microenvironmental conditions and on the expression of leaf traits that regulate water flow through the plant. Thus, the mutual effects of canopy cover, larger leaves and smaller stomatal size may drive a more efficient water use by reducing the transpiration rate of plants growing in agroforestry systems.

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The datasets generated and/or analyzed during the current study are available in the supplementary material or from the corresponding author on reasonable request.


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We are thankful to the staff of the Sara Ana experimental station for allowing us to conduct the study and support us with guidance and advise during fieldwork. To the staff of the Herbario Nacional de Bolivia (LPB) and the Biology department for their help with leaf data processing. To Wiebke Niether for helpful advice during field work. The study was funded by the Swiss Agency for Development and Cooperation (SDC), the Liechtenstein Development Service (LED), the Biovision Foundation for Ecological Development and the Coop Sustainability Fund and implemented in cooperation with the Research Institute of Organic Agriculture (FiBL) and Ecotop Foundation. The study was carried out under permission and current laws of the Government of Bolivia.

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The study was designed by Francisco Saavedra, Ernesto Jordan and Kasuya Naoki. Ernesto Jordan collected data in the field and laboratory. Francisco Saavedra and Kasuya Naoki conducted data analysis. Ernesto Jordan wrote the first draft of the manuscript. Early versions of the mansucript was wrote by Francisco Saavedra. Monika Schneider revised it critically early versions of the manuscript. All the authors discussed the results and commented on the manuscript.

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Correspondence to Francisco Saavedra.

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Saavedra, F., Jordan Peña, E., Schneider, M. et al. Effects of environmental variables and foliar traits on the transpiration rate of cocoa (Theobroma cacao L.) under different cultivation systems. Agroforest Syst 94, 2021–2031 (2020).

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  • Agroforestry
  • Canopy cover
  • Specific leaf area
  • Stomata size
  • Temperature