Stephen Adu-Bredu

African rainforests: past, present and future.

The rainforests are the great green heart of Africa, and present a unique combination of ecological, climatic and human interactions. In this synthesis paper, we review the past and present state processes of change in African rainforests, and explore the challenges and opportunities for maintaining a viable future for these biomes. We draw in particular on the insights and new analyses emerging from the Theme Issue on ‘African rainforests: past, present and future’ of Philosophical Transactions of the Royal Society B. A combination of features characterize the African rainforest biome, including a history of climate variation; forest expansion and retreat; a long history of human interaction with the biome; a relatively low plant species diversity but large tree biomass; a historically exceptionally high animal biomass that is now being severely hunted down; the dominance of selective logging; small-scale farming and bushmeat hunting as the major forms of direct human pressure; and, in Central Africa, the particular context of mineral- and oil-driven economies that have resulted in unusually low rates of deforestation and agricultural activity. We conclude by discussing how this combination of factors influences the prospects for African forests in the twenty-first century.

The past, present and future of Africa's rainforests.

The African wet tropics contain the second largest area of tropical rainforest in the world (second only to Amazonia), 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

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Abstract Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.

Forest biomass, productivity and carbon cycling along a rainfall gradient in West Africa

Net Primary Productivity (NPP) is one of the most important parameters in describing the functioning of any ecosystem and yet it arguably remains a poorly quantified and understood component of carbon cycling in tropical forests, especially outside of the Americas. We provide the first comprehensive analysis of NPP and its carbon allocation to woody, canopy and root growth components at contrasting lowland West African forests spanning a rainfall gradient. Using a standardized methodology to study evergreen (EF), semi-deciduous (SDF), dry forests (DF) and woody savanna (WS), we find that (i) climate is more closely related with above and belowground C stocks than with NPP (ii) total NPP is highest in the SDF site, then the EF followed by the DF and WS and that (iii) different forest types have distinct carbon allocation patterns whereby SDF allocate in excess of 50% to canopy production and the DF and WS sites allocate 40%-50% to woody production. Furthermore, we find that (iv) compared with canopy and root growth rates the woody growth rate of these forests is a poor proxy for their overall productivity and that (v) residence time is the primary driver in the productivity-allocation-turnover chain for the observed spatial differences in woody, leaf and root biomass across the rainfall gradient. Through a systematic assessment of forest productivity we demonstrate the importance of directly measuring the main components of above and belowground NPP and encourage the establishment of more permanent carbon intensive monitoring plots across the tropics.

The modern pollen–vegetation relationships of a tropical forest–savannah mosaic landscape, Ghana, West Africa

ABSTRACT 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.

Pollen-vegetation richness and diversity relationships in the tropics

Tracking changes in biodiversity through time requires an understanding of the relationship between modern diversity and how this diversity is preserved in the fossil record. Fossil pollen is one way in which past vegetation diversity can be reconstructed. However, there is limited understanding of modern pollen-vegetation diversity relationships from biodiverse tropical ecosystems. Here, pollen (palynological) richness and diversity (Hill N1) are compared with vegetation richness and diversity from forest and savannah ecosystems in the New World and Old World tropics (Neotropics and Palaeotropics). Modern pollen data were obtained from artificial pollen traps deployed in 1-ha vegetation study plots from which vegetation inventories had been completed in Bolivia and Ghana. Pollen counts were obtained from 15 to 22 traps per plot, and aggregated pollen sums for each plot were > 2,500. The palynological richness/diversity values from the Neotropics were moist evergreen forest = 86/6.8, semi-deciduous dry forest = 111/21.9, wooded savannah = 138/31.5, and from the Palaeotropics wet evergreen forest = 144/28.3, semi-deciduous moist forest = 104/4.4, forest-savannah transition = 121/14.1; the corresponding vegetation richness/diversity was 100/36.7, 80/38.7 and 71/39.4 (Neotropics), and 101/54.8, 87/45.5 and 71/34.5 (Palaeotropics). No consistent relationship was found between palynological richness/diversity, and plot vegetation richness/diversity, due to the differential influence of other factors such as landscape diversity, pollination strategy, and pollen source area. Palynological richness exceeded vegetation richness, while pollen diversity was lower than vegetation diversity. The relatively high global diversity of tropical vegetation was found to be reflected in the pollen rain.

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. This article is part of the discussion meeting issue ‘The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications’.

Trait variations in 28-year-old teak (Tectona grandis) provenance field trials in Ghana, West Africa

Stem straightness, axis persistence, presence/absence of protuberant buds and epicormics, diameter and height have profound influence on timber quality and volume of teak (Tectona grandis). Provenance trials of teak were established in the 1970s in Ghana, as part of the Food and Agriculture Organization (FAO) International Series of Provenance Trials, with the view of selecting teak germplasm for sites with specific environmental conditions. Two field trials were located in dry semi-deciduous (dry) and moist semi-deciduous (moist) ecological zones of Ghana. They consisted of 13 provenances, including four landraces each from Ghana and Indonesia, and two and three provenances from India and Laos, respectively. Trees were assessed at 9, 17 and 28 years to (1) quantify the potential variation in quality and timber volume production, (2) examine possibilities for determining early selection of parameters of superior provenances and (3) select provenances for sites with particular environmental conditions. Production traits were higher on the moist site than the dry site. Mean height was 23.2 and 20.2 m tree−1, stem cross-sectional area at breast height was 0.0896 and 0.0474 m2 tree−1, and stem volume was 0.75 and 0.34 m3 tree−1 for the moist and dry sites, respectively. Nilambur provenance from moist India had the highest mean stem straightness score of 19% above average, whereas Savannakhet from Laos had the best protuberant bud score with 18% of the trees above average. Indonesian landraces performed better in the dry zone, whereas provenances from India and Laos performed better in the moist zone. Phenotypic correlations between age 9 and 28 years were moderate (r 0.54–0.90) to high (r > 0.90) for production and qualitative characteristics, indicating feasibility of early assessment for identification of superior provenances. Nilambur and Savannakhet II provenances proved to be favourable choices for the moist zone, whereas the Temandsang provenance from Indonesia was found to be the best choice for the dry site.

Mapping the Leaf Economic Spectrum across West African Tropical Forests Using UAV-Acquired Hyperspectral Imagery

The leaf economic spectrum (LES) describes a set of universal trade-offs between leaf mass per area (LMA), leaf nitrogen (N), leaf phosphorus (P) and leaf photosynthesis that influence patterns of primary productivity and nutrient cycling. Many questions regarding vegetation-climate feedbacks can be addressed with a better understanding of LES traits and their controls. Remote sensing offers enormous potential for generating large-scale LES trait data. Yet so far, canopy studies have been limited to imaging spectrometers onboard aircraft, which are rare, expensive to deploy and lack fine-scale resolution. In this study, we measured VNIR (visible-near infrared (400–1050 nm)) reflectance of individual sun and shade leaves in 7 one-ha tropical forest plots located along a 1200–2000 mm precipitation gradient in West Africa. We collected hyperspectral imaging data from 3 of the 7 plots, using an octocopter-based unmanned aerial vehicle (UAV), mounted with a hyperspectral mapping system (450–950 nm, 9 nm FWHM). Using partial least squares regression (PLSR), we found that the spectra of individual sun leaves demonstrated significant (p < 0.01) correlations with LMA and leaf chemical traits: r2 = 0.42 (LMA), r2 = 0.43 (N), r2 = 0.21 (P), r2 = 0.20 (leaf potassium (K)), r2 = 0.23 (leaf calcium (Ca)) and r2 = 0.14 (leaf magnesium (Mg)). Shade leaf spectra displayed stronger relationships with all leaf traits. At the airborne level, four of the six leaf traits demonstrated weak (p < 0.10) correlations with the UAV-collected spectra of 58 tree crowns: r2 = 0.25 (LMA), r2 = 0.22 (N), r2 = 0.22 (P), and r2 = 0.25 (Ca). From the airborne imaging data, we used LMA, N and P values to map the LES across the three plots, revealing precipitation and substrate as co-dominant drivers of trait distributions and relationships. Positive N-P correlations and LMA-P anticorrelations followed typical LES theory, but we found no classic trade-offs between LMA and N. Overall, this study demonstrates the application of UAVs to generating LES information and advancing the study and monitoring tropical forest functional diversity.

Compositional patterns of overstorey and understorey woody communities in a forest-savanna boundary in Ghana

ABSTRACT Background: Forest and savanna vegetation in the zone of transition (ZOT) contain distinct woody species due to fire, drought and herbivory barriers that constrain forest species from invading adjacent savannas and vice-versa. Little is known if these barriers cause divergence in species composition between the overstorey and understorey strata in these vegetation types. Aim: We investigated woody species composition across overstorey and understorey strata in the ZOT and explored the relationship between soil fertility and species composition patterns. Methods: We sampled overstorey and understorey woody species and determined soil nutrient concentrations in twenty-five 20 m × 20 m plots in a ZOT in Ghana. Results: Forest and savanna species dominated the overstorey and understorey of their respective environments. However, species composition was decoupled between the overstorey and understorey strata in both forest and savanna vegetations. Few savanna and forest species had individuals co-occurring in both overstorey and understorey such that ~65% of the dominant species was limited to only one stratum. Soil fertility had little effect on these patterns. Conclusion: These patterns indicate that, forest and savanna species face significant recruitment barriers in their respective environments, suggesting that requirements for juvenile establishment may differ from recruitments to the canopy layer.