African rainforests: past, present and future

In recent decades, there has been a surge of interest in tropical forests, as there is increased appreciation of the rich biodiversity they host and the many roles they play in the functioning of the Earth system at local, regional and global scales. Of the world’s major tropical forest regions, most research and policy attention has focused on the Amazon region, the world’s largest tropical forest bloc, and to a lesser extent on South East Asia, the third largest tropical forest region. By contrast, the world’s second largest tropical forest region, the tropical forests of Central and West Africa (termed the Guineo-Congolian region) have been relatively neglected. This has been for a number of reasons, including challenging and fragmented politics, civil conflicts and logistical as well as infrastructure challenges. Nevertheless, there is an extensive amount of research activity in the African rainforest zone that has rarely been compiled in a single interdisciplinary volume. This review paper synthesizes the insights emerging from the theme issue on ‘African rainforests: past, present and future’ of Philosophical Transactions of the Royal Society.JRC.H.5-Land Resources Managemen

The past, present and future of Africa's rainforests

The Africanwet tropics contain the second largest area of tropical rainforest in the world (second only toAmazonia), accounting for roughly 30% of global rainforest cover, the lush green heart of an otherwise generally dry continent. These rainforests have global significance and value as reservoirs of biodiversity, as stores and sinks of atmospheric carbon, as regulators of flow of mighty rivers, as sources of moisture to the atmosphere and engines of the global atmospheric circulation, as a key component of the Earth system and its biogeochemical cycles, and as providers of resources and ecosystem services to local communities and the region’s nations. They also have a unique and particular history of changes in climate and human pressure, and face a range of contemporary pressures. Over the twenty-first century, the African rainforest realmhas the potential to witness massive change, both through an expansion of deforestation, hunting and logging, and through the effects of global climate change. This Theme Issue presents a multidisciplinary perspective on the nature and ecology of the African rainforest biome, and examines the current pressures and future threats to this biome. Compared with the other major rainforest regions, Amazonia and Southeast Asia, the African rainforest realm remains understudied, and in particular there have been very few attempts at interdisciplinary synthesis. This Theme Issue is an attempt to address this deficit, and explores what we know about the African rainforests and the threats they face, and what we need to know is this century of rapid change. In some ways, this can be viewed as a complement to similar Theme Issues of this journal focused on the rainforests of Amazonia [1,2] and Southeast Asia [3].JRC.H.5-Land Resources Managemen

Termites have wider thermal limits to cope with environmental conditions in savannas

1. The most diverse and abundant family of termites, the Termitidae, evolved in African tropical forests. They have since colonised grassy biomes such as savannas. These open environments have more extreme conditions than tropical forests, notably wider extremes of temperature and lower precipitation levels and greater temporal fluctuations (of both annual and diurnal variation). These conditions are challenging for soft-bodied ectotherms, such as termites, to survive in, let alone become as ecologically dominant as termites have. 2. Here, we quantified termite thermal limits to test the hypothesis that these physiological limits are wider in savanna termite species to facilitate their existence in savanna environments. 3. We sampled termites directly from mound structures, across an environmental gradient in Ghana, ranging from wet tropical forest through to savanna. At each location, we quantified both the Critical Thermal Maxima (CTmax) and the Critical Thermal Minima (CTmin) of all the most abundant mound-building Termitidae species in the study areas. We modelled the thermal limits in two separate mixed-effects models against canopy cover at the mound, temperature and rainfall, as fixed effects, with sampling location as a random intercept. 4. For both CTmax and CTmin, savanna species had significantly more extreme thermal limits than forest species. Between and within environments, areas with higher amounts of canopy cover were significantly associated with lower CTmax values of the termite colonies. CTmin was significantly positively correlated with rainfall. Temperature was retained in both models; however, it did not have a significant relationship in either. Sampling location explained a large proportion of the residual variation, suggesting there are other environmental factors that could influence termite thermal limits. 5. Our results suggest that savanna termite species have wider thermal limits than forest species. These physiological differences, in conjunction with other behavioural adaptations, are likely to have enabled termites to cope with the more extreme environmental conditions found in savanna environments and facilitated their expansion into open tropical environments.Royal Society-DFID/FCDO Capacity Building Initiative; NERC ACCE DTP.http://wileyonlinelibrary.com/journal/janeZoology and Entomolog

Carbon dynamics, net primary productivity (NPP) and human appropriated NPP (HANPP) across a forest‐cocoa farm landscape in West Africa

Terrestrial net primary productivity (NPP) is an important metric of ecosystem functioning; however, there is little empirical data on the NPP of human-modified ecosystems, particularly smallholder,perennial crops like cocoa (Theobroma cacao), which are extensive across the tropics. Human appropriated NPP (HANPP) is a measure of the proportion of a natural system’s NPP that has either been reduced through land-use change or harvested directly and, previously, has been calculated to estimate the scale of the human impact on the biosphere. Additionally, human-modification can create shifts in NPP allocation and decomposition, with concomitant impacts on the carbon cycle. This study presents the results of three years of intensive monitoring of forest and smallholder cocoa farms across disturbance, management intensity, distance from forest and farm age gradients. We measured among the highest reported NPP values in tropical forest, 17.57 ± 2.1 and 17.7 ± 1.6 Mg C ha-1 yr-1 for intact and logged forest respectively; however, the average NPP of cocoa farms was still higher, 18.8 ± 2.5 Mg C ha-1 yr-1, which we found was driven by cocoa pod production. We found a dramatic shift in litterfall residence times, where cocoa leaves decomposed more slowly than forest leaves and shade tree litterfall decomposed considerably faster, indicating significant changes in rates of nutrient cycling. The average HANPP value for all cocoa farms was 2.1 ± 1.1 Mg C ha-1 yr-1; however, depending on the density of shade trees it ranged from -4.6 to 5.2 Mg C ha-1 yr-1. Therefore, rather than being related to cocoa yield, HANPP was reduced by maintaining higher shade levels. Across our monitored farms 18.9% of farm NPP was harvested (i.e. whole cocoa pods) and only 1.1% (i.e.cocoa beans) was removed from the system; suggesting that the scale of HANPP in smallholder cocoa agroforestry systems is relatively small

The modern pollen-vegetation relationship of a tropical forest-savannah mosaic landscape, Ghana, West Africa

Transitions between forest and savannah vegetation types in fossil pollen records are often poorly understood due to over-production by taxa such as Poaceae and a lack of modern pollen-vegetation studies. Here, modern pollen assemblages from within a forest-savannah transition in West Africa are presented and compared, their characteristic taxa discussed, and implications for the fossil record considered. Fifteen artificial pollen traps were deployed for 1 year, to collect pollen rain from three vegetation plots within the forest-savannah transition in Ghana. High percentages of Poaceae and Melastomataceae/Combretaceae were recorded in all three plots. Erythrophleum suaveolens characterised the forest plot, Manilkara obovata the transition plot and Terminalia the savannah plot. The results indicate that Poaceae pollen influx rates provide the best representation of the forest-savannah gradient, and that a Poaceae abundance of >40% should be considered as indicative of savannah-type vegetation in the fossil record

Pantropical modelling of canopy functional traits using Sentinel-2 remote sensing data

Tropical forest ecosystems are undergoing rapid transformation as a result of changing environmental conditions and direct human impacts. However, we cannot adequately understand, monitor or simulate tropical ecosystem responses to environmental changes without capturing the high diversity of plant functional characteristics in the species-rich tropics. Failure to do so can oversimplify our understanding of ecosystems responses to environmental disturbances. Innovative methods and data products are needed to track changes in functional trait composition in tropical forest ecosystems through time and space. This study aimed to track key functional traits by coupling Sentinel-2 derived variables with a unique data set of precisely located in-situ measurements of canopy functional traits collected from 2434 individual trees across the tropics using a standardised methodology. The functional traits and vegetation censuses were collected from 47 field plots in the countries of Australia, Brazil, Peru, Gabon, Ghana, and Malaysia, which span the four tropical continents. The spatial positions of individual trees above 10 cm diameter at breast height (DBH) were mapped and their canopy size and shape recorded. Using geo-located tree canopy size and shape data, community-level trait values were estimated at the same spatial resolution as Sentinel-2 imagery (i.e. 10 m pixels). We then used the Geographic Random Forest (GRF) to model and predict functional traits across our plots. We demonstrate that key plant functional traits can be accurately predicted across the tropicsusing the high spatial and spectral resolution of Sentinel-2 imagery in conjunction with climatic and soil information. Image textural parameters were found to be key components of remote sensing information for predicting functional traits across tropical forests and woody savannas. Leaf thickness (R2 = 0.52) obtained the highest prediction accuracy among the morphological and structural traits and leaf carbon content (R2 = 0.70) and maximum rates of photosynthesis (R2 = 0.67) obtained the highest prediction accuracy for leaf chemistry and photosynthesis related traits, respectively. Overall, the highest prediction accuracy was obtained for leaf chemistry and photosynthetic traits in comparison to morphological and structural traits. Our approach offers new opportunities for mapping, monitoring and understanding biodiversity and ecosystem change in the most species-rich ecosystems on Earth

Resistance of African tropical forests to an extreme climate anomaly.

The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015-2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015-2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha-1 y-1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests

Pantropical modelling of canopy functional traits using Sentinel-2 remote sensing data

Funding Information: This work is a product of the Global Ecosystems Monitoring (GEM) network (gem.tropicalforests.ox.ac.uk). J.A.G. was funded by the Natural Environment Research Council (NERC; NE/T011084/1 and NE/S011811/1) and the Netherlands Organisation for Scientific Research (NWO) under the Rubicon programme with project number 019.162LW.010. The traits field campaign was funded by a grant to Y.M. from the European Research Council (Advanced Grant GEM-TRAIT: 321131) under the European Union‘s Seventh Framework Programme (FP7/2007-2013), with additional support from NERC Grant NE/D014174/1 and NE/J022616/1 for traits work in Peru, NERC Grant ECOFOR (NE/K016385/1) for traits work in Santarem, NERC Grant BALI (NE/K016369/1) for plot and traits work in Malaysia and ERC Advanced Grant T-FORCES (291585) to Phillips for traits work in Australia. Plot setup in Ghana and Gabon were funded by a NERC Grant NE/I014705/1 and by the Royal Society-Leverhulme Africa Capacity Building Programme. The Malaysia campaign was also funded by NERC GrantNE/K016253/1. Plot inventories in Peru were supported by funding from the US National Science Foundation Long-Term Research in Environmental Biology program (LTREB; DEB 1754647) and the Gordon and Betty Moore Foundation Andes-Amazon Program. Plots inventories in Nova Xavantina (Brazil) were supported by the National Council for Scientific and Technological Development (CNPq), Long Term Ecological Research Program (PELD), Proc. 441244/2016-5, and the Foundation of Research Support of Mato Grosso (FAPEMAT), Project ReFlor, Proc. 589267/2016. During data collection, I.O. was supported by a Marie Curie Fellowship (FP7-PEOPLE-2012-IEF-327990). GEM trait data in Gabon was collected under authorisation to Y.M. and supported by the Gabon National Parks Agency. D.B. was funded by the Fondation Wiener-Anspach. W.D.K. acknowledges support from the Faculty Research Cluster ‘Global Ecology’ of the University of Amsterdam. M.S. was funded by a grant from the Ministry of Education, Youth and Sports of the Czech Republic (INTER-TRANSFER LTT19018). Y.M. is supported by the Jackson Foundation. We thank the two anonymous reviewers and Associate Editor G. Henebry for their insightful comments that helped improved this manuscript.Peer reviewedPostprin

ENSO Drives interannual variation of forest woody growth across the tropics

Meteorological extreme events such as El Niño events are expected to affect tropical forest net primary production (NPP) and woody growth, but there has been no large-scale empirical validation of this expectation. We collected a large high–temporal resolution dataset (for 1–13 years depending upon location) of more than 172 000 stem growth measurements using dendrometer bands from across 14 regions spanning Amazonia, Africa and Borneo in order to test how much month-to-month variation in stand-level woody growth of adult tree stems (NPPstem) can be explained by seasonal variation and interannual meteorological anomalies. A key finding is that woody growth responds differently to meteorological variation between tropical forests with a dry season (where monthly rainfall is less than 100 mm), and aseasonal wet forests lacking a consistent dry season. In seasonal tropical forests, a high degree of variation in woody growth can be predicted from seasonal variation in temperature, vapour pressure deficit, in addition to anomalies of soil water deficit and shortwave radiation. The variation of aseasonal wet forest woody growth is best predicted by the anomalies of vapour pressure deficit, water deficit and shortwave radiation. In total, we predict the total live woody production of the global tropical forest biome to be 2.16 Pg C yr−1, with an interannual range 1.96–2.26 Pg C yr−1 between 1996–2016, and with the sharpest declines during the strong El Niño events of 1997/8 and 2015/6. There is high geographical variation in hotspots of El Niño–associated impacts, with weak impacts in Africa, and strongly negative impacts in parts of Southeast Asia and extensive regions across central and eastern Amazonia. Overall, there is high correlation (r = −0.75) between the annual anomaly of tropical forest woody growth and the annual mean of the El Niño 3.4 index, driven mainly by strong correlations with anomalies of soil water deficit, vapour pressure deficit and shortwave radiation

Resistance of African tropical forests to an extreme climate anomaly

The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015–2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015–2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha−1 y−1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests