Carbon cycle in tropical upland ecosystems: a global review

Abstract. Along with habitat transformation, climate change has profound impacts on biodiversity and may alter ecosystem services on which human welfare depends. Many studies of the carbon cycle have focused on lowland tropical forests; however, upland forests have been less explored despite their pivotal role in carbon sequestration. Here, I synthesized the state of knowledge on the allocation of carbon in its different stocks (aboveground, belowground, and soil) as well as in its main fluxes (plant decomposition, respiration, and litterfall) in tropical upland ecosystems of the planet. In November 2020, a systematic review was carried out to identify references published from 2000 to 2020 through a combination of key terms in Google Scholar and Scopus databases, thus analysing bibliographic, geographical, methodological, and carbon cycling information of the global upland tropics (between 23.5∘ N–23.5∘ S). After analysing a total of 1967 references according to inclusion–exclusion criteria, 135 references published in the last 20 years were selected. Most of the studies were conducted in the tropical and subtropical moist broadleaf forest of South America. The main factors studied were elevation and forest type. Forest structure and soil variables were largely associated when studying carbon cycling in these ecosystems. Estimations of carbon stocks comprised three-fourths of the total studies, while the remaining fraction focused on carbon fluxes. Aboveground biomass and carbon in soils were highly investigated, while plant decomposition and respiration were the components that received the least attention. Even though in the last 20 years there was a slight increase in the number of studies on carbon cycle in tropical upland forests, I found bias associated with the biomes and ecoregions studied (especially in the Andes). Elevation was the main factor examined but other essential aspects such as the successional gradient, landscape management, diversity–productivity relationship, faunal and microbial effect, trophic cascades, and Gadgil effect require more attention. The inclusion of different litter species and origins (i.e. roots and stems) and theoretical frameworks including home-field advantage, substrate–matrix interaction, and phenology–substrate match may provide explanatory mechanisms to better understand litter decomposition over these forests. Despite respiration being a paramount link that is closely tied to above- and belowground compartment, this flux constitutes one of the important gaps to fulfil in future research. For a comprehensive understanding of the carbon cycle in upland forests, it is necessary to obtain information on its main fluxes and integrate them into climate change mitigation plans.