C Wade Ross1,2, Niall P Hanan1, Lara Prihodko3, Julius Anchang1, Wenjie Ji1, Qiuyan Yu1. 1. Department of Plant and Environmental Sciences New Mexico State University, Las Cruces, NM, USA. 2. Tall Timbers Research Station, Tallahassee, Florida 32312, USA. 3. Animal and Range Sciences, New Mexico State University, Las Cruces, NM, USA.
Abstract
Africa's ecosystems have an important role in global carbon dynamics, yet consensus is lacking regarding the amount of carbon stored in woody vegetation and the potential impacts to carbon storage in response to changes in climate, land use, and other Anthropocene risks. Here, we explore the socio-environmental conditions that shaped the contemporary distribution of woody vegetation across sub-Saharan Africa and evaluate ecosystem response to multiple scenarios of climate change, anthropogenic pressures, and fire disturbance. Our projections suggest climate change will have a small but negative effect on above ground woody biomass at the continental scale, and the compounding effects of population growth, increasing human pressures, and socio-climatic driven changes in fire behavior further exacerbate climate-driven trends. Relatively modest continental-scale trends obscure much larger regional perturbations, with climatic and anthropogenic factors leading to increased carbon storage potential in East Africa, offset by large deficits in West, Central, and Southern Africa.
Africa's ecosystems have an important role in global carbon dynamics, yet consensus is lacking regarding the amount of carbon stored in woody vegetation and the potential impacts to carbon storage in response to changes in climate, land use, and other Anthropocene risks. Here, we explore the socio-environmental conditions that shaped the contemporary distribution of woody vegetation across sub-Saharan Africa and evaluate ecosystem response to multiple scenarios of climate change, anthropogenic pressures, and fire disturbance. Our projections suggest climate change will have a small but negative effect on above ground woody biomass at the continental scale, and the compounding effects of population growth, increasing human pressures, and socio-climatic driven changes in fire behavior further exacerbate climate-driven trends. Relatively modest continental-scale trends obscure much larger regional perturbations, with climatic and anthropogenic factors leading to increased carbon storage potential in East Africa, offset by large deficits in West, Central, and Southern Africa.
Planetary-scale consumption of resources, driven by rapid population and economic
growth, has substantially increased emissions of CO2 and other greenhouse
gases[1,2]. Building on previous treaties, the Paris Agreement[3] developed international policy in an
effort to limit global warming to less than 2°C above pre-industrial levels via
decarbonization of energy systems and increased mitigation efforts, including improved
land management and ecosystem restoration to increase the carbon sink and storage
capacity of terrestrial ecosystems. Achieving ambitious climate mitigation goals
requires credible, accurate, and reliable mapping and monitoring of terrestrial carbon
stock. As the second largest landmass, Africa’s forests, woodlands, and savannas
have a substantial impact on the global carbon budget by actively cycling carbon between
the atmosphere, vegetation, and soil. However, the continent remains one of the largest
sources of uncertainty in the global carbon cycle, functioning as both a sink and a
source of CO2 in response to natural and anthropogenic
perturbations[4-9]. In recent decades, for example, severe droughts
have impacted many humid and sub-humid regions[10,11], leading to tree
mortality and loss of biomass[12].
Conversely, several studies have reported the recovery of African drylands following
severe droughts of the 1970s and 1980s, attributed to post-drought vegetation recovery.
Improvements in land management and agroforestry have also contributed to improved
sustainability, while meeting increased demands for food, fiber, and livestock
production[9,13-15].While Earth’s climate system is unequivocally changing[16,17],
predicting the rate and magnitude of associated changes in terrestrial systems is a
major unresolved challenge for evaluating coupled human-environmental impacts[18,19]. Many African communities, for example, rely on forests, woodlands
and savannas for timber production and energy (fuelwood and charcoal), food (fruits,
nuts, and animal rangeland), traditional medicines, and other crucial resources.
Depending on the context, these land uses can enhance or reduce woody biomass[9,20]. Emissions from land-use (0.32 ± 0.05 Pg C
yr−1) and fire (1.03 ± 0.22 Pg C yr−1),
while highly variable through both space and time, are significant carbon
sources[7,8]. Land use, land-use change, and fire behavior are
therefore central components of global change and carbon dynamics, as the risk of land
degradation from climate change and anthropogenic pressures has never been
greater[21-23].Here, we present a novel approach to deriving continental-scale estimates of
above ground woody biomass by characterizing the climatic, topo-edaphic, and
socioeconomic conditions that have shaped the distribution of tree and shrub biomass
across the African landscape and evaluate ecosystem response to multiple scenarios of
disturbance. We explore the spatial patterns of contemporary woody vegetation using a
new Earth observation (EO) dataset[24]
that integrates optical phenological metrics, microwave and LiDAR, providing an improved
characterization of woody cover and biomass, particularly in the open and low-stature
savannas of sub-Saharan Africa (Methods). We use a
data-driven, machine-learning-based approach to examine how climate, human pressures,
fire, and topo-edaphic conditions impact biomass variability across the continent (Extended Data Fig. 1). Ecosystem response to climate
change is evaluated using the fitted relationships to empirically project and quantify
the potential distribution of biomass for the end of the 21st century.
Contemporary and future climate conditions are represented with long-term mean-annual
aridity (calculated as 1 – [precipitation/potential evapotranspiration]), derived
from a 27-model ensemble (Supplementary Table 1) participating in the fifth phase of the Coupled Model
Inter-comparison Project (CMIP5). Two climate-forcings (representative concentration
pathways, RCP 4.5 and RCP 8.5) are used to investigate ecosystem response under a broad
range of potential climate-change scenarios for the end of this century. We also
evaluate ecosystem response to assumptions regarding future population growth and
socio-climatic driven changes in fire behavior.
Extended Data Fig. 1
Socio-environmental data.
a) Long-term mean-annual aridity based on the
historical data[12] (1981
– 2010) and b) climate zones, with H.A., hyper-arid; A.,
arid; S.A., semi-arid; S.H., Sub-humid; H., humid. c)
Satellite-derived estimates of mean annual (2003 – 2015) burned-area
(%)[31].
d) Digital elevation model[52]. e) The contemporary
human footprint index[32].
f) Hydrologic soil groups[33], with A corresponds to low
runoff-potential soils (e.g., sands); B, moderately low runoff-potential; C,
moderately high runoff-potential; D, high runoff-potential (e.g.,
clays).
Biomass prediction and evaluation
Comparing our biomass predictions with the satellite-derived estimates
yields excellent agreement (Supplementary Table 2) and confirms broad geographic expectations, with
the density of woody biomass greatest near the equator and generally declining with
latitude, but with considerable spatial variability associated with regional-scale
topo-edaphic heterogeneity and climate (Fig.
1). Validation against an independent test set (N = 18,320) indicates that
our model explains ca. 89% of the variation in tree and shrub
biomass with low bias and low variance (Supplementary Table 3). Further
comparison reveals that our biomass predictions are consistently at the low end of
the range of previously reported estimates (Supplementary Table 4). However, it is
important to note that models from previous studies[25-28] were trained exclusively with data sampled from forested
regions, whereas the satellite-derived estimates[24] used to train our model better characterize the open and
low-stature canopies of the savannas and drylands of Africa, which are
underrepresented in most woody-cover datasets despite covering more than half of the
African continent[29].
Figure 1.
Satellite-derived estimates of contemporary (2005) above-ground woody biomass
(Mg ha-1).
By integrating Earth-observation data from optical, microwave, and LiDAR
sensors, Hanan and colleagues24 improved biomass estimates for trees and shrubs
in the open and low-stature dryland and drought-seasonal savannas of sub-Saharan
Africa. Marginal plots on vertical and horizontal axes correspond to woody
biomass density averaged by latitude and longitude, respectively. The map
overlay corresponds to ecological regions, which were derived by aggregating
Terrestrial Ecosystems30 into broader classes (Extended Data Fig. 2). CA is the Cape of Africa; CM is Central
Mesic; EAH is East African Highlands; EM is Eastern Madagascar; ES is Eastern
Sahel; ESS is East Sudanian Savannas; HA is the Horn of Africa; MT is Moist
Tropical; ND is Namib Desert; RV is Rift Valley; SD is Southern Dry; WM is
Western Madagascar; WS is Western Sahel; WSS is West Sudanian Savanna.
Evaluation of the conditional distributions from our covariate set indicates
that climate has the largest effect on the contemporary distribution of woody
biomass, which decreases as aridity[12], fire[31],
human pressures[32], and sand
content[33] increase (Fig. 2). Aridity alone explains 75% of the
variance across sub-Saharan Africa, where mean biomass density is greatest in humid
(103.9 ± 87.6 Mg ha−1, mean ± 1 standard deviation)
and dry sub-humid (27.1 ± 21.2 Mg ha−1) regions, including
the moist tropics (185.9 ± 69 Mg ha−1), eastern Madagascar
(94.9 ± 63 Mg ha−1), and East African Highlands (44.2
± 39 Mg ha−1). Conversely, biomass density is lowest in
Africa’s hyper-arid (9e−3 ± 0.1 Mg
ha−1), arid (1.3 ± 2.3 Mg ha−1), and
semi-arid regions (8.8 ± 9.9 Mg ha−1). In the
water-limited, drought-seasonal savannas of the Sahel, mean biomass density ranges
from 1.0 ± 2 Mg ha−1 in the west to 2.4 ± 3 Mg
ha−1 in the east.
Figure 2.
Sensitivity of above ground woody biomass to socio-environmental
conditions.
Accumulated local effects from the fitted random forest model (n
=79,576) illustrate the relationship between sub-Saharan woody biomass and
socio-environmental covariates, which are ranked in order of largest (top) to
smallest effect (bottom). The x-axis represents the units of the independent
covariate, the y-axis represents the size of the mean effect each covariate has
on woody biomass predictions, and grey shading indicates the 95% confidence
interval. Aridity [calculated as 1 – (precipitation/potential
evapotranspiration)] values ranging from 1.0 – 0.9, 0.9 – 0.8, 0.8
– 0.5, 0.5 – 0.35, and less than 0.35, respectively correspond to
hyper-arid, arid, semi-arid, dry sub-humid, and humid regions. Hydrologic soil
groups A, B, C, and D respectively correspond to soils with low run off
potential (e.g., sandy soils), moderately low, moderately high, and high runoff
potential (e.g., clay soils).
Fire is most extensive in semi-arid and dry sub-humid climate zones (Methods 2.4). Indeed, nearly half (45%) of the
land area impacted by fire each year occurs in the Sudanian savannas—where
mean biomass density ranges from 21.5 ± 24 Mg ha−1 in the
west to 44.6 ± 42.8 Mg ha−1 in the east. Human pressures
also have an overall negative effect on biomass density in our model (Fig. 2). However, the relationship between humans and
woody biomass is nuanced. In remote areas where anthropogenic disturbance is low
(e.g., index scores of ca. 10 or less), biomass declines nearly
linearly in response to increasing anthropogenic pressures. The strength of the
effect weakens and even begins to increase as human pressures increase further,
perhaps reflecting human promotion of forest cover in and around
settlements[20]. For
example, conservation and restoration efforts such as the
‘shelterbelts’ in Nigeria, Mali and Burkina Faso, ‘green
belts’ spanning the Sahel, and agroforestry have had a profound impact on
woody-vegetation recovery in recent decades[9,34,35]. However, the human footprint relationship
is characterized by a large degree of uncertainty, especially in areas with
relatively moderate index scores (e.g., between 20 and 35) and further analysis is
required to fully tease apart the nuanced relationship between woody biomass and
anthropogenic disturbance.
Aridification yields a biomass deficit
Our end-of-the-century projections suggest that woody biomass respectively
declines by 0.5 – 2.5% in response to climate changes under RCP 4.5 and RCP
8.5, representing a biomass deficit of ca. 0.4 – 2.1 Pg
relative to our predictions for the contemporary (i.e., baseline) period. While the
overall spatial patterns of change are similar under both climate scenarios, the
direction and magnitude of the response are amplified under RCP 8.5 (Fig. 3). Broadly speaking, woody vegetation tends to
decline across most of West, Central, and Southern Africa in response to hotter and
drier climate conditions. The already dry landscape of southern Africa is expected
to become more arid, particularly in South Africa, where aridity under RCP 8.5 is
expected increase by 6 – 10% relative to the baseline period (See Methods 2.2).
Figure 3.
Projected change (%) in above ground woody biomass relative to contemporary
estimates.
Each map represents the projected change (%) in the mean density of
woody-biomass (1 km−2) relative to the baseline period in
response to projections in aridity (calculated as 1 –
(precipitation/potential evapotranspiration)] under a) RCP 4.5) and
b) RCP 8.5. The map overlay corresponds to biophysical regions
illustrated in Fig. 1.
Per unit area, the largest responses occur in the Cape and Namib desert
regions, with the density of woody vegetation declining by ca. 15
– 33% depending on the scenario. However, these regions contribute a small
fraction to the bottom line of Africa’s biomass budget, and therefore
represent a relatively small fraction of the future deficit. In regard to biomass
stock (and thus carbon storage), the largest projected deficits in response to
climate changes occur in the central-mesic region and the West Sudanian Savannas,
where woody biomass declines by ca. 0.3 – 2.1 Pg (Fig. 3). Under RCP 4.5, however, the magnitude of
loss is much lower (−0.01 Pg), and the deficit is largely offset by growth in
the Eastern Sahel (+0.01 Pg). In West Africa, biomass deficits under RCP 4.5 emerge
primarily in the western-most extent of the mesic Sudanian and Guinean savanna
regions (e.g., southern Senegal, Guinea, Guinea-Bissau, and southern Mali). However,
a much larger portion of West Africa is expected to incur deficits under RCP 8.5,
with projected losses expanding further east and south across much of coastal West
Africa (Fig. 2b).Most of the projected biomass growth occurs throughout East Africa, where
mean biomass density (Mg ha−1) increases by ca.
21 – 78% in response to warmer and wetter climate conditions. The largest
potential increase for carbon storage occurs in the Horn of Africa (0.5 – 1.2
Pg), followed by the Rift Valley region (0.4 – 0.5 Pg), and the Highlands of
East Africa (0.2 – 0.3 Pg). Although the density of woody vegetation has a
relatively strong response to increasingly humid conditions across much of the
semi-arid landscape of the Central and Eastern Sahel (e.g., northeastern Mali, Niger
and Chad; Extended Data Fig. 2), productivity
will likely remain constrained by mean-annual rainfall. The carbon-storage potential
of the Sahel is therefore expected to remain relatively low, with biomass increasing
by ca. 0.01 – 0.04 Pg.
Extended Data Fig. 2
Biophysical regions and countries of sub-Saharan Africa.
a) Biophysical regions were derived by aggregating The
Nature Conservancy Terrestrial Ecosystems[30] into broader classes.
b) Country borders were mapped using the R sf[64] package.
Fire dynamics have mixed effects
While Africa accounts for the majority of Earth’s fire-derived carbon
emissions (52%)[7], recent research
indicates that burned-area extent has declined across sub-Saharan Africa[36,37]. Evaluation of our burned-area projections, driven by
changes in climate and human pressures, suggest that this trend may continue through
the end of the century, particularly in the semi-arid and dry sub-humid regions of
Africa, where projections of burned-area extent decline by 2 – 15% in
response to the considered forcing scenarios. However, our results for the larger
Sub-Saharan Africa region suggest that Africa’s already large contribution to
global fire-derived carbon emissions may actually increase in the coming decades in
response to aridification and increased fuel and flammability in humid regions,
where the potential loss in woody biomass is relatively large.At the continental scale, the impacts of fire on above ground woody biomass
have virtually no effect under RCP 4.5, as regional gains and losses offset each
other (Supplementary Table
5). Under RCP 8.5, however, changing fire regimes appear to reduce
Africa’s total woody biomass by an additional 1.7 Pg. The major fire-induced
losses in carbon are attributed to reduced precipitation and higher evaporative
demand in the moist tropics, where burned area is projected to increase and biomass
density is one to two orders of magnitude larger than biomass density in other
biophysical regions. Relatively large deficits in response to burned-area
projections under RCP 8.5 are also expected to occur in eastern Madagascar
(−0.5 Pg), as well as in East (−0.2 Pg) and West Sudanaian savannas
(−0.2 Pg). Conversely, burned area is expected to decrease across large
portions of East Africa, which has a small but positive effect on total biomass in
the East African Highlands, the Rift Valley, and the Horn of Africa, presumably due
to reduced fuels flammability in response to an increasingly humid climate. A
reduction in burned area in the East Sudanian Savannas offsets the negative effects
of climate changes under RCP 4.5, resulting in a small increase relative to the
baseline period.
Our findings suggest that human pressures not only exacerbate the
climate-driven deficits that are projected for much of West, Central, and Southern
Africa, but also abate much of the growth response expected to occur across large
portions of East Africa. Depending on the forcing scenario, above ground woody
biomass is expected to decline by 4 – 8% in response to the compounding
effects of changes in climate, fire disturbance, and human
pressures—representing a deficit of ca. 3.4 – 6.7 Pg
relative to our predictions for the baseline period. The largest deficits under
these assumptions occur in the moist tropical and central mesic regions, where
biomass declines by ca. 1.2 – 2.8 Pg. In the East African
Highlands, where biomass increases in response climate changes under both forcing
scenarios, human pressures offset the growth response under RCP 4.5, resulting in a
small deficit (Figure 3). This trend also
occurs under RCP 8.5; however, the effects of climate changes outweigh those from
human pressures because the growth response is much larger under the high emissions
scenario.Regional differences regarding the direction and magnitude of change in
response to human pressures largely depend on projections regarding local population
and economic development assumptions under SSP2[38]—the “middle of the road” Shared
Socioeconomic Pathways scenario (See Methods
2.3 and Extended Data Fig. 3). The
largest relative change occurs in Niger (Fig.
5), where human pressures increase by 50%, followed by Liberia (31%), Chad
(28%), and Egypt (28%). Conversely, index scores increase by just 1% in Lesotho, and
are expected to decrease in South Africa (−0.6%) and Zimbabwe (−5%),
presumably due to regional differences in birth rates and/or emigration. While woody
biomass has a small, positive response to the aforementioned demographic trends in
South Africa and Zimbabwe, the effects of aridification are much stronger, resulting
in an overall deficit under these assumptions.
Extended Data Fig. 3
Human population density.
a), Population density (people per km−2) for 2010
and b) projected population density for 2100 under the “middle of the
road” Shared Socioeconomic Pathways (SSP2)[38].
Figure 5.
Sensitivity of burned area (%) to socio-environmental conditions.
Accumulated local effects from the fitted random forest model illustrate
the relationship between burned-area and socio-environmental covariates in
sub-Saharan Africa, which are ranked in order of highest feature importance
(top) to lowest feature importance (bottom). The x-axis represents the units of
the independent covariate, the y-axis represents the size of the mean effect
each covariate has on woody biomass predictions, and grey shading indicates the
95% confidence interval. Note that larger human footprint values correspond to
regions with higher human pressures due to anthropogenic activity.
Discussion
The compounding effects of climate change, regional increases in human
pressures due to population and socioeconomic growth, and increased fire disturbance
in heavily vegetated regions are expected reduce woody biomass by 4 – 8%
across sub-Saharan Africa, representing a deficit of ca. 3.4
– 6.7 Pg relative to our contemporary estimates. Distinct spatial patterns of
change are similar between the considered climate-forcing scenarios, but the trends
are amplified under RCP 8.5. Our projections suggest that above-ground woody carbon
storage declines across most of West, Central, and Southern Africa in response to
increasingly arid conditions associated with climate changes. A small portion of the
projected deficit is expected to be offset by growth in East Africa and other
regions in response to warmer and wetter climate conditions.Our evaluation of Anthropocene risks is enabled by integrating
Representative Concentration Pathways with Shared-Socioeconomic
Pathways—which were developed to provide a framework for a new generation of
climate change research[39].
Anthropogenic pressures generally exacerbate the climate-driven deficit; however,
considerable scope exists to determine how natural systems respond to socioeconomic
transformation—such as land-use change and/or community-based restoration
efforts—and if non-linearity or tipping points exist where anthropogenic
pressures lead to accelerated impacts. Our assessment of anthropogenic pressures via
the human-footprint index circumvents the limitations of relying solely on remote
sensing, which has difficulty in detecting low intensity pressures[40] and often confounds natural and
anthropogenic land-cover patterns in arid and patchy environments[41]. This approach is, however,
subject to three primary constraints. First, these data do not fully account for all
human activities, such as invasive species and pollution[32] or conservation and restoration efforts.
Second, we model future human pressures from socio-economic narratives of population
dynamics only, leaving infrastructure, energy networks, and other data static.
Third, we use the Shared Socioeconomic Pathways ‘middle of the road’
assumptions (SSP2) regarding population dynamics[38,39]. Therefore, we
consider our projection of anthropogenic pressures to be conservative, as
transportation networks, built environments, and agriculture will presumably expand
with socioeconomic growth.While our models do not incorporate mechanistic relationships or
biogeochemical feedbacks that could alter the climate-driven trends, our results are
generally consistent with those reported by Martens and colleagues[42], who used an adaptive Dynamic
Global Vegetation Model (aDGVM) to quantify ecosystem response to climate forcings
under RCP 4.5 and RCP 8.5. The authors reported that woody vegetation changed
between −8 to 11% under RCP 4.5, and by −22 to −6% under RCP
8.5 when CO2 enrichment was omitted. When the CO2 effect was
included, the authors reported that aboveground vegetation changed between 18% to
43% (RCP4.5) and 37% to 61% (RCP8.5), and that this change was primarily associated
with woody encroachment into grasslands and increased woody cover in savannas.
However, some research suggests that general circulation and Earth system models may
be overly sensitive to CO2 enrichment[43] and that the CO2 effect is weakened when
moisture constraints are strong[44].
There is also evidence to suggest that CO2 enrichment may cause woody
plants to complete their lifecycles faster[45], therefore increasing biomass turnover and offsetting
carbon sequestered in response to CO2 enrichment[46]. Our approach is deliberately data-driven
and empirical, acknowledging that although our models capture the
socio-environmental components that explain much of the variability in the
contemporary distribution of woody biomass, empirical projections may not fully
capture the mechanistic relationships leading to the observed patterns of change.
Indeed, the scenarios presented here are intended to provide insight into the
multidimensional aspects of global change.Climate change—coupled with population growth, economic development
and land-use change—will inevitably lead to long-term and widespread changes
in vegetation structure and carbon storage in Africa and other continents. Our
understanding of these trends—and the appropriate policy and land-management
decisions needed to promote economic wellbeing and carbon
sequestration—should be guided by research that not only examines the
implications of climate-changes, but how economic development, demographic trends,
and land management is likely to change over the next century. This information is
essential to refine our understanding of how Anthropocene risks might impact coupled
biophysical and social systems in the coming years. This is especially important for
promoting and ensuring sustainable land use for the African communities that rely on
local forests, woodlands, and savannas for energy, food, livestock grazing,
traditional medicines, and other essential resources.
Data availability
The datasets used for this analysis can be accessed as described below.Woody cover and biomass data[24] are available GeoTiff files from the Oak Ridge
National Laboratory (ORNL) Distributed Active Archive Center (DAAC;
https://doi.org/10.3334/ORNLDAAC/1777).Aridity data were provided by Feng and Fu[12].The Human Footprint map[32] is available as a GeoTiff file from Dryad
(https://doi.org/10.5061/dryad.052q5).Future projections of human population density based on Shared
Socioeconomic Pathways[38]
are available as GeoTiff files from the Socioeconomic Data and Application
Center (SEDAC; https://doi.org/10.7927/m30p-j498).Contemporary estimates of burned area are available as GeoTiff and
were acquired from Kahiu and Hanan[31].HYSOGs[47] data are
available as GeoTiff from the ORNL DAAC (https://doi.org/10.3334/ORNLDAAC/1566).Shuttle Radar Topography Mission (SRTM) elevation data were acquired
from the United States Geological Survey (USGS) Earth Explorer (https://earthexplorer.usgs.gov/).Biophysical regions were derived from The Nature Conservancy
Terrestrial Ecoregions and are provided as GIS shapefiles (http://maps.tnc.org/gis_data.html).Biomass prediction maps[48] and R code[49] are available from Figshare.
Methods
Overview of methods
We developed predictive relationships between satellite-derived
estimates of woody biomass[24]
and socio-environmental covariates in R 3.4.4[50] with the random forest package[51]. The random forest
algorithm[52] was chosen
as it allows for non-linear, non-monotonic relationships between the target
property and multiple covariates. Predictive relationships were modeled from a
sample of the satellite-derived woody biomass estimates, which was achieved by
extracting grid-cell attributes from our covariate set and satellite-derived
woody biomass estimates at 100,000 randomly generated point locations. Missing
values were removed, leaving 99,471 rows in our modeling matrix. All random
forest (RF) models were developed on a training set (80%) and model performance
was assessed with a held-out validation set (20%). Training (n =79,576) and
validation (n =19,894) sets were determined by splitting the data frame at
random, an optimal mtry value of 4 was identified with the
tuneRF function, and 1,000 individual trees comprised the
forest. The two sample Kolmogorov-Smirnov test was conducted to verify that the
distribution of woody biomass was similar between training and validation
sets[53].
Socio-environmental data
The spatial distribution of woody biomass was modeled with key
socio-environmental drivers, including aridity[12], anthropogenic disturbance[32], burned area
estimates[31],
elevation[54], and
hydrologic soil groups[33,47] (Extended Data Fig. 1). All data were projected to a common
coordinate system (sinusoidal equal-area) and, if necessary, re-sampled to match
the spatial resolution of our woody biomass product (1 km−2).
Bilinear interpolation and nearest neighbor were used to resample continuous and
categorical data, respectively.
Woody biomass
Satellite-derived estimates of woody biomass were provided by Hanan
et al.[24] at 1
km2 resolution for sub-Saharan Africa (Extended Data Fig. 4). These estimates are a
product of data integration, which was achieved by applying allometric
equations to relate biomass with canopy cover and canopy height. Canopy
cover estimates were produced at 1 km2 resolution by combining
phenological metrics from MODIS with Quick-Scatterometer ku-band microwave
retrievals. This product was chosen to represent canopy cover as it provides
an improved assessment of low stature systems by accounting for vegetation
less than 5 m in height, which occupy a considerable fraction of the African
landscape but remain under-represented in widely-used tree-cover datasets
due to mapping challenges presented by their complex landscapes, and the
underestimation of woody plants by methods that exclude short stature trees
and shrubs[29]. Canopy
height was estimated by Simard and colleagues[55] from light detection and ranging
(LiDAR) at 1 km2 resolution. Biomass estimates were derived by
relating canopy cover to canopy height with an allometric equation derived
from the globallometree.org database[56].
Extended Data Fig. 4
Above-ground woody biomass (Mg ha−1)
a), Satellite-derived estimates of above-ground woody
biomass[24]. b),
Predicted above-ground biomass representing the baseline (i.e.,
contemporary) estimates. c), End of century empirical projection of woody
biomass in response to RCP 4.5 and assumptions regarding population growth
and fire regime changes. d), End of century empirical projection of woody
biomass in response to RCP 8.5 and assumptions regarding population growth
and fire regime changes. Data are available for download[48] as GeoTiffs.
Aridity
Global coverage of the aridity-index was provided as a 27-model
ensemble mean by Feng and Fu[12] (Supplementary Table 1). The aridity index (AI = precipitation /
potential evapotranspiration) was calculated using the Penman-Monteith
method, which accounts for the effects of surface-air temperature, humidity,
solar radiation, and wind speed. These data were temporally averaged over
the CMIP5 historical forcing’s (1980 to 2005) to model
satellite-derived estimates of existing woody biomass. Aridity was then
calculated as 1 – AI so that increasingly larger values correspond to
increasingly drier conditions. Ecosystem response to climate change (CMIP5
aridity projections) was modeled under representative concentration pathways
(RCP) 4.5 and RCP 8.5. These RCPs were chosen to represent a broad range of
potential climate change scenarios.
Human footprint
Following Venter and colleagues[32], we adopted the human-footprint
methodology[57] to
project human pressures in response to population growth
assumptions[38] for
end of the 21st century. Contemporary human pressures were
related to satellite-derived estimates of woody biomass via the 2009
‘Human Footprint’ map[32], which was developed through cumulative pressure
mapping by integrating datasets of built environments, population density,
electric infrastructure, croplands, pasture lands, roads, railways, and
navigable waterways. Venter and colleagues[32] developed the Human Footprint index
by first standardizing each dataset on a scale of 0 to 10 to obtain
individual pressure scores for each dataset. Anthropogenic ‘pressure
scores’ are then summarized into a single dataset to produce the
Human Footprint index (Figure 3a). Low
scores (e.g., 0 to 5) correspond to areas that that receive little or no
pressure from human activities. Conversely, highly impacted areas (e.g.,
large cities) are assigned larger scores, with a maximum score of 50.To assess biomass dynamics in response to end-of-century human
pressures, we produced a future Human Footprint map (Figure 3b) by calculating the population pressure
score for 2100. Projected population density was derived from the Global
Population Projection Grids Based on Shared Socioeconomic Pathways
(SSPs)[38]. We
choose SSP2, which represents a conservative (‘middle of the
road’) scenario regarding spatiotemporal patterns of population,
urbanization, and development demographics. Following the human footprint
methodology, a pressure score of 10 was assigned to all grid cells with
1,000 or more people km−2. Population pressure scores for
the remaining grid cells were logarithmically scaled (Eq. 1).
To obtain an index score for the future human footprint, we
summarized the pressures scores provided by Venter and colleagues[32], substituting the 2009
population pressure score with the SSP2 population pressure score. While
future projections of other anthropogenic activities (e.g., agricultural
expansion, transportation, built environments, etc.) have not yet been
developed, they are correlated with the increase in human population
density. Thus, our future human footprint map is considered a conservative
estimate of human pressures for 2100.
Burned area
Long-term (2003 to 2015) mean-annual burned area was provided from
Kahui and Hanan[31]. These
data were produced from Earth observations of monthly burned area estimates
obtained from the Global Fire Emissions Database (GFED) with a spatial
resolution of 0.25° (Extended Data Fig.
1). While most African fires are attributed to land use and
management, climate strongly influences fire intensity, severity, and
wildfire spread. We therefore model contemporary burned area as a function
of climate (i.e. aridity), dry season precipitation, anthropogenic pressures
as indexed by the human footprint, and elevation (Extended Data Fig. 1). Specifically, we used
long-term mean-annual aridity and dry-season precipitation, elevation, and
the Human Footprint map. The same set of randomly generated point locations
described in the methods overview were used to extract the grid-cell
attribute information from the aforementioned covariates. Model evaluation
with the independent validation set (N = 20%) indicates that our random
forest model was able to explain 72% of burned-area variation, corresponding
to a RMSE of 8.8%. Evaluation of accumulated local effects, obtained from
the Interpretable Machine Learning package[58], indicates that aridity is the most
important predictor of burned area, followed by dry-season precipitation,
elevation, and human pressures. Burned area projections for the end of the
21st century (2071 to 2100) were projected in response to
changes in human pressures, mean-annual precipitation and dry-season
precipitation.
Topo-edaphic properties
Hydrologic soil groups (0 – 100 cm) were obtained from the
HYSOGs250m dataset[47],
which integrates data pertaining to soil texture classes, depth to bedrock,
and depth to water table. Three arc-second, gap-filled Shuttle Radar
Topography Mission (SRTM) elevation data was acquired from earthexplorer.usgs.gov. We assume that
the aforementioned topo-edaphic properties (Extended Data Fig. 1) will not change substantially by 2100, and
were therefore treated as constants in our model.
Model evaluation
Model evaluation and data analysis was performed with the tidyverse
package[59], and figures
were produced using raster[60],
rasterVis[61],
ggplot2[62],
colorspace[63],
RColorBrewer[64],
gridExtra[65],
iml[58], and
sf[66] packages.
Data-model fit was evaluated with the coefficient of determination
(R2, equation 2),
root mean squared error (RMSE, equation
3), residual prediction deviation (RPD, equation 4), and the ratio of performance to
interquartile distance (RPIQ, equation
5).
where, are the model predicted values,
y are the observed values,
n is the number of predicted or observed values in the
held-out dataset (testing) with i = 1, 2,…, n,
SD is the standard deviation of the testing set,
RMSE is the root mean square error, and IQ
is the interquartile range.
Socio-environmental data.
a) Long-term mean-annual aridity based on the
historical data[12] (1981
– 2010) and b) climate zones, with H.A., hyper-arid; A.,
arid; S.A., semi-arid; S.H., Sub-humid; H., humid. c)
Satellite-derived estimates of mean annual (2003 – 2015) burned-area
(%)[31].
d) Digital elevation model[52]. e) The contemporary
human footprint index[32].
f) Hydrologic soil groups[33], with A corresponds to low
runoff-potential soils (e.g., sands); B, moderately low runoff-potential; C,
moderately high runoff-potential; D, high runoff-potential (e.g.,
clays).
Biophysical regions and countries of sub-Saharan Africa.
a) Biophysical regions were derived by aggregating The
Nature Conservancy Terrestrial Ecosystems[30] into broader classes.
b) Country borders were mapped using the R sf[64] package.
Human population density.
a), Population density (people per km−2) for 2010
and b) projected population density for 2100 under the “middle of the
road” Shared Socioeconomic Pathways (SSP2)[38].
Above-ground woody biomass (Mg ha−1)
a), Satellite-derived estimates of above-ground woody
biomass[24]. b),
Predicted above-ground biomass representing the baseline (i.e.,
contemporary) estimates. c), End of century empirical projection of woody
biomass in response to RCP 4.5 and assumptions regarding population growth
and fire regime changes. d), End of century empirical projection of woody
biomass in response to RCP 8.5 and assumptions regarding population growth
and fire regime changes. Data are available for download[48] as GeoTiffs.
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