Climate-land-use interactions shape tropical mountain biodiversity and ecosystem functions.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Impacts of climate and land use on N2 O and CH4 fluxes from tropical ecosystems in the Mt. Kilimanjaro region, Tanzania.

In this study, we quantify the impacts of climate and land use on soil N2 O and CH4 fluxes from tropical forest, agroforest, arable and savanna ecosystems in Africa. To do so, we measured greenhouse gases (GHG) fluxes from 12 different ecosystems along climate and land-use gradients at Mt. Kilimanjaro, combining long-term in situ chamber and laboratory soil core incubation techniques. Both methods showed similar patterns of GHG exchange. Although there were distinct differences from ecosystem to ecosystem, soils generally functioned as net sources and sinks for N2 O and CH4 respectively. N2 O emissions correlated positively with soil moisture and total soil nitrogen content. CH4 uptake rates correlated negatively with soil moisture and clay content and positively with SOC. Due to moderate soil moisture contents and the dominance of nitrification in soil N turnover, N2 O emissions of tropical montane forests were generally low (<1.2 kg N ha-1  year-1 ), and it is likely that ecosystem N losses are driven instead by nitrate leaching (~10 kg N ha-1  year-1 ). Forest soils with well-aerated litter layers were a significant sink for atmospheric CH4 (up to 4 kg C ha-1  year-1 ) regardless of low mean annual temperatures at higher elevations. Land-use intensification significantly increased the soil N2 O source strength and significantly decreased the soil CH4 sink. Compared to decreases in aboveground and belowground carbon stocks enhanced soil non-CO2 GHG emissions following land-use conversion from tropical forests to homegardens and coffee plantations were only a small factor in the total GHG budget. However, due to lower ecosystem carbon stock changes, enhanced N2 O emissions significantly contributed to total GHG emissions following conversion of savanna into grassland and particularly maize. Overall, we found that the protection and sustainable management of aboveground and belowground carbon and nitrogen stocks of agroforestry and arable systems is most crucial for mitigating GHG emissions from land-use change.

Species richness is more important for ecosystem functioning than species turnover along an elevational gradient.

Many experiments have shown that biodiversity enhances ecosystem functioning. However, we have little understanding of how environmental heterogeneity shapes the effect of diversity on ecosystem functioning and to what extent this diversity effect is mediated by variation in species richness or species turnover. This knowledge is crucial to scaling up the results of experiments from local to regional scales. Here we quantify the diversity effect and its components-that is, the contributions of variation in species richness and species turnover-for 22 ecosystem functions of microorganisms, plants and animals across 13 major ecosystem types on Mt Kilimanjaro, Tanzania. Environmental heterogeneity across ecosystem types on average increased the diversity effect from explaining 49% to 72% of the variation in ecosystem functions. In contrast to our expectation, the diversity effect was more strongly mediated by variation in species richness than by species turnover. Our findings reveal that environmental heterogeneity strengthens the relationship between biodiversity and ecosystem functioning and that species richness is a stronger driver of ecosystem functioning than species turnover. Based on a broad range of taxa and ecosystem functions in a non-experimental system, these results are in line with predictions from biodiversity experiments and emphasize that conserving biodiversity is essential for maintaining ecosystem functioning.