Research Article |
Corresponding author: Javier Bravo-García ( j.bravo@evenor-tech.com ) Academic editor: Janet Franklin
© 2024 Javier Bravo-García, Juan Camarillo-Naranjo, Francisco J. Blanco-Velázquez, Félix González-Peñaloza, María Anaya-Romero.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bravo-García J, Camarillo-Naranjo J, Blanco-Velázquez FJ, González-Peñaloza F, Anaya-Romero M (2024) Mapping the potential habitat suitability and opportunities of bush encroacher species in Southern Africa: a case study of the SteamBioAfrica project. Frontiers of Biogeography 17: e136222. https://doi.org/10.21425/fob.17.136222
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Senegalia mellifera (Benth) Seigler & Ebinger., Dichrostachys cinerea (L.) Wight & Arn. and Terminalia sericea Burch. Ex DC., are three important bush encroacher species that contribute to the well-known ecological process named “thicketization” in Southern Africa. This issue has persisted for many years, impacting species distribution, plant communities, soil, and fauna dynamics. According to climate change projections, Southern Africa is expected to become drier and warmer in future scenarios, creating favourable conditions for proliferation of bush encroacher species. MaxEnt is a general-purpose machine learning method widely utilized in various ecological and biological scenarios to predict the potential suitable habitat of species. This is achieved by incorporating presence-only occurrence records and bioclimatic, and topographic variables. The analysis was performed in a Geographic Information System based on the current and future suitable areas for the respective Shared Socioeconomic Pathways (SSP) scenarios according to INM-CM5-0, UK-ESM1-0-LL and MPIES-M1-2-HR climate models. This was done to assess the potential effects of climate change on the distribution patterns of bush encroacher species. Model performance was assessed by the area under the curve (AUC) of the receiver operator characteristic (ROC), with 0.836, 0.822 and 0.738 as results to Terminalia sericea, Dichrostachys cinerea and Senegalia mellifera respectively. The current results show that Senegalia mellifera presents a habitat suitability of 56% (1,460,353 km²) of the total area, while Terminalia sericea and Dichrostachys cinerea have suitability over 37.9% (996,168 km²) and 43.9% (1,154,645 km²) of the area, respectively. These findings indicate that precipitation and temperature variables are the most important factors in explaining the spatial distribution of the bush encroacher species, predicting a future increase between 8–29.4%, 2.8–24%, and 3–24.2% for Senegalia mellifera, Terminalia sericea, and Dichrostachys cinerea respectively. Furthermore, each species has its own set of important variables and different ecological behaviour patterns. These results imply that an improved understanding of the response of woody species to a changing climate is important for managing bush encroachment in savanna ecosystems.
Based on our analysis, Senegalia mellifera is currently suitable in 56% of the total area; while 43.9% of the total area is suitable for Dichrostachys cinerea, and 37.87% of the total area is currently suitable for Terminalia sericea. In the future scenarios, the habitat suitability increases for all three species compared to the current state.
The unsuitable areas decrease in all the proposed scenarios with respect to the present, predicting a shrub expansion in the future throughout southern Africa. Furthermore, our results imply that Senegalia mellifera is the most potentially affected by bush encroachment in future changes, with a larger distribution area in future scenarios. These findings are supported by many studies, which indicate the probable increase of woody cover and loss of grasslands.
Temperature and precipitation patterns are the main drivers behind the distribution of these bush encroachers, increasing or decreasing the competitiveness of these species according to these variables and their phenotypic plasticity.
The increase in habitat suitability occurs throughout the case study, but there is a clear trend of shrub expansion towards the south of the case study. The change maps show a pattern of shrub movement towards South Africa and Botswana. This would fit with climate predictions for South Africa, which predict a decrease in rainfall in the east and north of the country, being more suitable for shrub competitive species.
Bush encroaching, climate change, Dichrostachys cinerea, habitat suitability, MaxEnt, Senegalia mellifera, Terminalia sericea, thicketization
“Thicketization’’, “bush encroachment” or “woody plant encroachment” are different concepts used to explain the natural phenomenon characterized by an increase in density of woody plant bushes in the different savanna and grassland land uses (
Referring to the impact on rangelands and its effects on species distribution and plant-soil-fauna interactions, it aligns with the modern two-layer theory of savanna ecology (Kulmatiski and Beard 2013). According to this theory, savannas are characterized by a distinct vertical structure consisting of an upper layer dominated by trees and a lower layer dominated by grasses and herbaceous plants. If the herbaceous layer is over exploited, it loses its competitive edge and becomes unable to efficiently utilize water and nutrients, increasing the infiltration rate into the subsoil, favouring the growth of shrubs and trees, enabling them to assume the dominance (
It will pose a challenge to identify a singular factor as the exclusive reason for the disruptions linked to bush encroachment, particularly considering the spatial correlation of all influencing environmental variables (Kgosikoma and Mogotsi et al. 2013;
However, many studies have shown that the spread of bushes in semi-arid areas can also have substantially effects (
The central issue in the woody plant encroachment is the role of the water and the landscape water balance. Several studies have shown that bush encroachment increases evapotranspiration and may lead to excessive water use contributing to more severe droughts (
Ecological Niche Modeling (ENM) has become a fundamental tool for understanding species’ ecological requirements and predicting their distribution patterns, particularly in light of current and future environmental changes (
Bioclimatic factors in the study area are projected to change in unprecedented ways. Temperatures in Namibia, Botswana and western South Africa are expected to experience significant warming, ranging from 0.2–0.5 °C per decade, leading to an increase in the number of hot days (
The encroachment of both native and non-indigenous invasive woody species, exacerbated by the impacts of climate change, is causing significant environmental, social, and economic harm (
The selection of three specific species—Terminalia sericea, Dichrostachys cinerea, and Senegalia mellifera—is based on their generalist behaviour, invasiveness, and biomass potential in semi-arid areas, aligning closely with the project’s objectives (Fig.
The study aims to elucidate the likely distribution of these three species, developing habitat suitability maps based on climate change scenarios proposed by the IPCC in 2021 following an accepted and well-known methodology (Fig.
Our case study (Fig.
Namibia has a total area of 825,418 km² and is the driest country in southern Africa, mainly in the southwest, and slightly wetter in the northeast region bordering Botswana, with over 600 mm of rainfall per year (
Botswana covers an area of over 600,370 km², with an average elevation of over 1100 m.a.s.l. (
Finally, South Africa is the largest and most rugged country with 1.22 million km² and an average elevation of 1200 m.a.s.l. (
The arid tropical climate of the Namib Desert and Kalahari Desert covers large parts of Namibia, Botswana, and certain parts of South Africa (
Different encroached farms of Namibia; (A) Dichrostachys cinerea and Terminalia prunoides encroached zone at Farm Deutsche Erde (Colin Lindeque, Carbon Capital, 8th May 2019), (B) Terminalia sericea encroached zone in Farm Lensrust (Colin Lindeque, Carbon Capital,19 November 2018) and (C) Senegalia mellifera encroached zone in Farm Seruka (Colin Lindeque, Carbon Capital, 24th April 2019).
The presence of species Senegalia mellifera, Terminalia sericea, and Dichrostachys cinerea predominantly occur in savanna ecosystems characterized by seasonal variability in precipitation and temperatures. These species are typically found in areas with annual precipitation ranging from 400 to 1000 mm and mean annual temperatures between 18 °C and 32 °C, depending on the species (
Distribution data for Senegalia mellifera, Dichrostachys cinerea and Terminalia sericea were obtained from literature and the Global Biodiversity Information Facility (GBIF), which compiles a lot of historical and current information on species for each country (Fig.
For instance, in the case of Terminalia sericea, there are a total of 1,017 occurrence points (120 in Namibia, 328 in Botswana, and 569 in South Africa). Senegalia mellifera has a total of 755 occurrence points (285 in South Africa, 280 in Botswana, and 190 in Namibia) and Dichrostachys cinerea has a total of 1,336 occurrence points (893 in South Africa, 320 in Botswana, and 123 in Namibia). After the rarefaction process for Terminalia sericea, Senegalia mellifera, and Dichrostachys cinerea, they were modeled with 286, 281, and 186 occurrence points respectively.
During the development of the ENMs for these species, the selection of environmental variables was grounded in an extensive review of the existing literature and empirical data on the factors influencing species occurrence (
Three types of variables: edaphic (pH, CEC, coarse fragments, clay, silt, sand and bulk density), physical (slope, elevation, roughness and aspect) and bioclimatic (19 bioclimatic variables) were selected based on research and literature on the habitat of woody encroachers (Table
Variables selected for testing in the MaxEnt model. We used environmental variables (clay, sand, silt, coarse fragments, bulk density, pH, cation exchange capacity, elevation, slope, roughness and aspect) and bioclimatic variables from WorldClim (bio1–bio19).
Bioclimatic variables | Code | Resolution | Source |
---|---|---|---|
Annual Mean Temperature | BIO1 | 30 s (~ 1 km) | WorldClim |
Mean Diurnal Range | BIO2 | 30 s (~ 1 km) | WorldClim |
Isothermality (BIO2/BIO7)*100 | BIO3 | 30 s (~ 1 km) | WorldClim |
Temperature seasonality | BIO4 | 30 s (~ 1 km) | WorldClim |
Max Temperature of the Warmest month | BIO5 | 30 s (~ 1 km) | WorldClim |
Min Temperature of Coldest Month | BIO6 | 30 s (~ 1 km) | WorldClim |
Temperature Annual Range | BIO7 | 30 s (~ 1 km) | WorldClim |
Mean Temperature of Wettest Quarter | BIO8 | 30 s (~ 1 km) | WorldClim |
Mean Temperature of Driest Quarter | BIO9 | 30 s (~ 1 km) | WorldClim |
Mean Temperature of Warmest Quarter | BIO10 | 30 s (~ 1 km) | WorldClim |
Mean Temperature of Coldest Quarter | BIO11 | 30 s (~ 1 km) | WorldClim |
Annual Precipitation | BIO12 | 30 s (~ 1 km) | WorldClim |
Precipitation of Driest Month | BIO13 | 30 s (~ 1 km) | WorldClim |
Precipitation of Wettest Month | BIO14 | 30 s (~ 1 km) | WorldClim |
Precipitation Seasonality | BIO15 | 30 s (~ 1 km) | WorldClim |
Precipitation of Wettest Quarter | BIO16 | 30 s (~ 1 km) | WorldClim |
Precipitation of Driest Quarter | BIO17 | 30 s (~ 1 km) | WorldClim |
Precipitation of Warmest Quarter | BIO18 | 30 s (~ 1 km) | WorldClim |
Precipitation of Coldest Quarter | BIO19 | 30 s (~ 1 km) | WorldClim |
Environmental variables | Code | Resolution | Source |
Clay content | clay | 250 m | SoilsGrid |
Sand | sand | 250 m | SoilsGrid |
Silt | silt | 250 m | SoilsGrid |
Coarse fragments | coarsefrg | 250 m | SoilsGrid |
Bulk density | bd | 250 m | SoilsGrid |
pH | pH | 250 m | SoilsGrid |
Cation Exchange Capacity | CEC | 250 m | SoilsGrid |
Elevation | DEM | 30 s (~ 1 km) | WorldClim |
Slope | slope | 30 s (~ 1 km) | Own source |
Roughness | roughness | 30 s (~ 1 km) | Own source |
Aspect | aspect | 30 s (~ 1 km) | Own source |
The elevation was obtained from WorldClim, a database of high spatial resolution global weather and climate data. This variable was excluded from the modelling due to the controversial use in future projections owing to correlations of elevation with temperature and precipitation (
Future projections were made using the shared socio-economic pathways (SSPs) for greenhouse gas concentrations, specifically SSP245-585 across two-time horizons: 2041–2060 and 2061–2180. We utilized WorldClim 1-km resolution climate data from three global Earth System Models (ESMs) from the sixth phase of the Coupled Model Intercomparison Project (CMIP6) that obtained better results than CMIP5 models (
There are four primary Shared Socio-economic Pathways (SSPs) as outlined by Meinshausen et al. (2022). Among them, SSP275-585, which represents medium-high emissions, was chosen due to its alignment with the current global climate change mitigation efforts. This selection reflects the existing policies and clean energy initiatives (
The UK-ESM1.0-LL, MPI-ESM1.2-HR and INM-CM5-0 models were chosen for this study because, as noted by
MaxEnt 3.4.4 software was used to create the species distribution models (SDM), which requires environmental variables, presence data and the selection of the various features to find the distribution which has maximum entropy (
In the feature selection, the data were split into two groups; 25% of the distribution points were randomly chosen as the test data set and the remaining 75% of the distribution were used as a training data set. The maximum number of iterations of the model was set to 5000 and the number of repetitions was set to 10. In this way, the model was iterated 10 each time using a different test and training subset. The method employed for repetition was “bootstrapping”, aimed at enhancing result variability and subsequently utilizing the average output, giving the possibility to set the training and test configuration to the models (
The contribution of the different variables was calculated marking the jackknife test analysis in the software (
High correlation and collinearity between variables can easily lead to the overfitting of the model, thus affecting the accuracy of the resulting predictions, so Pearson correlation matrix through Python was done, compared with jackknife test (leave-one-out strategy) in MaxEnt (
Model performance was assessed by the area under curve (AUC) of the receiver operator characteristic (
Finally, the species distribution and habitat suitability map were produced by averaging the 10 replicate runs. Moreover, the habitat suitability thresholds from the model were reclassified using the “10th percentile training minimum presence” (T) of each species model (
A total of 57 distribution maps were acquired, with three generated under current conditions and 54 under conditions of climate change. The AUC values spanned from 0.738 to 0.836, denoting their classification performance as favourable (0.8 < AUC < 0.9). Specifically, Terminalia sericea with a mean AUC of 0.836, Senegalia mellifera with 0.738 and Dichrostachys cinerea with 0.822.
The outputs from the model were reclassified according to the common threshold established. The next tables show the area of the habitat suitability in km² and % for each climate model, climate scenario and in the different periods.
Habitat suitability for Terminalia sericea is mainly associated with annual precipitation (bio12; 55.2%) and annual mean temperature (bio1; 31.2%) (Table
Prediction of habitat suitability for Senegalia mellifera was widespread and mainly associated with mean temperature of wettest quarter (bio8; 50.9%), annual mean temperature (bio1; 27.4%) and annual mean precipitation (bio12: 11.1%) (Table
Additionally, the categories of high suitability register their maxima in the SSP585 emission scenarios, with the UK-ESM1.0-LL climate model recording the highest value in the period 2061–2080, with up to 58.6% high suitability for the species. On the other hand, the INM-CM5-0 climate model shows lower values, reaching up to 34.9% for the same period and emission scenario, followed by the MPI-ESM1.2-HR with 45.7% under the same conditions (Table
For Dichrostachys cinerea, the most related variables are precipitation in the driest month (bio13; 41.3%), mean temperature in the coldest quartile (bio11; 21.8%) and seasonality of temperatures (bio4; 18.9%) (Table
Furthermore, high suitability categories reach their peaks in the SSP585 emission scenarios, with the UK-ESM1.0-LL climate model registering the highest value for the 2061–2080 period, showing up to 47.6% high suitability for the species. Conversely, the INM-CM5-0 climate model records lower values, with a maximum of 29.7% for the same period and emission scenario, followed by the MPI-ESM1.2-HR at 36.53% under the same conditions (Table
Habitat suitability in the current scenario. This map illustrates variations in habitat suitability for the species, featuring distinct colors to delineate areas deemed unsuitable or suitable for the species. The color spectrum is utilized to convey different levels of suitability: areas in dark green represent the highest suitability, while light green signifies low suitability. Unsuitable areas for the species are depicted in grey.
Mean model outputs from MaxEnt models to predict habitat suitability in southern Africa for three bush encroacher species.
Variables | Code | Terminalia sericea | Dichrostachys cinerea | Senegalia mellifera | |||
---|---|---|---|---|---|---|---|
Percent contribution | Permutation importance | Percent contribution | Permutation importance | Percent contribution | Permutation importance | ||
Annual Mean Temperature | BIO1 | 31.1 | 38.2 | 6.2 | 15.2 | 27.4 | 31.3 |
Mean Diurnal Range | BIO2 | na | na | na | na | na | na |
Isothermality (BIO2/BIO7)*100 | BIO3 | na | na | na | na | na | na |
Temperature seasonality | BIO4 | 2.6 | 5.9 | 18.9 | 12.4 | 6.6 | 10.9 |
Max Temperature of the Warmest month | BIO5 | na | na | na | na | na | na |
Min Temperature of Coldest Month | BIO6 | 9.6 | 3.5 | na | na | na | na |
Temperature Annual Range | BIO7 | 1.4 | 2.9 | 11.9 | 30.7 | na | na |
Mean Temperature of Wettest Quarter | BIO8 | na | na | na | na | 50.9 | 16.7 |
Mean Temperature of Driest Quarter | BIO9 | na | na | na | na | na | na |
Mean Temperature of Warmest Quarter | BIO10 | na | na | na | na | na | na |
Mean Temperature of Coldest Quarter | BIO11 | na | na | 21.8 | 11.3 | na | na |
Annual Precipitation | BIO12 | 55.2 | 49.5 | na | na | 11.1 | 24.6 |
Precipitation of Driest Month | BIO13 | na | na | 41.3 | 30.3 | na | na |
Precipitation of Wettest Month | BIO14 | na | na | na | na | na | na |
Precipitation Seasonality | BIO15 | na | na | na | na | na | na |
Precipitation of Wettest Quarter | BIO16 | na | na | na | na | na | na |
Precipitation of Driest Quarter | BIO17 | na | na | na | na | na | na |
Precipitation of Warmest Quarter | BIO18 | na | na | na | na | na | na |
Precipitation of Coldest Quarter | BIO19 | na | na | na | na | na | na |
Clay content | clay | na | na | na | na | na | na |
Sand | sand | na | na | na | na | na | na |
Silt | silt | na | na | na | na | na | na |
Coarse fragments | coarsfrg | na | na | na | na | 3.9 | 16.5 |
Bulk density | bd | na | na | na | na | na | na |
pH | pH | na | na | na | na | na | na |
Cation Exchange Capacity | CEC | na | na | na | na | na | na |
Elevation | DEM | na | na | na | na | na | na |
Slope | slope | na | na | na | na | na | na |
Roughness | roughness | na | na | na | na | na | na |
Aspect | aspect | na | na | na | na | na | na |
Habitat suitability | Senegalia mellifera | Terminalia sericea | Dichrostachys cinerea | |||
---|---|---|---|---|---|---|
Category | Area (km²) | (%) | Area (km²) | (%) | Area (km²) | (%) |
Unsuitable | 1147234.4 | 44.0 | 1634000.7 | 62.1 | 1475523.4 | 56.1 |
Suitable | 1460353.8 | 56.0 | 996168.2 | 37.9 | 1154645.5 | 43.9 |
Low suitability | 347455.6 | 13.3 | 418297.5 | 15.9 | 588943.6 | 22.4 |
Moderate suitability | 1033229.6 | 39.6 | 430823.1 | 16.4 | 428755.0 | 16.3 |
High suitability | 79668.6 | 3.1 | 147047.6 | 5.6 | 136946.9 | 5.2 |
Terminalia sericea change between scenarios. The model was built under three CMIP6 climate models (MPI-ESM1.2-HR, UK-ESM1.0-LL and INM-CM5-0) and two periods of time (2041–2060 and 2061–2080).
Time period | Habitat suitability | Senegalia mellifera climate models and emission scenarios | |||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | |||||||||||||||||
SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | ||
2041–2060 | Unsuitable | 928580.2 | 35.6 | 873144.8 | 33.5 | 849512.2 | 32.6 | 632117.3 | 24.2 | 539693.2 | 20.7 | 515546.7 | 19.8 | 856823.4 | 32.9 | 807744.1 | 31.0 | 769990.7 | 29.5288451 |
Suitable | 1679008.0 | 64.4 | 1734443.4 | 66.5 | 1758076.0 | 67.4 | 1975470.9 | 75.8 | 2067895.0 | 79.3 | 2092041.5 | 80.2 | 1750764.8 | 67.1 | 1799844.1 | 69.0 | 1837597.5 | 70.4711549 | |
Low suitability | 246713.3 | 9.5 | 236136.6 | 9.1 | 255231.1 | 9.8 | 211587.6 | 8.1 | 204602.3 | 7.8 | 199003.1 | 7.6 | 225638.8 | 8.7 | 239783.7 | 9.2 | 225397.3 | 8.64389991 | |
Moderate suitability | 974404.1 | 37.4 | 951222.0 | 36.5 | 913350.2 | 35.0 | 754758.4 | 28.9 | 641830.2 | 24.6 | 616790.5 | 23.7 | 952787.8 | 36.5 | 922776.7 | 35.4 | 925995.8 | 35.5115811 | |
High suitability | 457890.7 | 17.6 | 547084.9 | 21.0 | 589494.7 | 22.6 | 1009124.9 | 38.7 | 1221462.6 | 46.8 | 1276247.8 | 48.9 | 572338.2 | 21.9 | 637283.7 | 24.4 | 686204.4 | 26.3156739 | |
2061–2080 | Unsuitable | 869185.7 | 33.3 | 742393.7 | 28.5 | 683928.8 | 26.2 | 510980.9 | 19.6 | 416545.9 | 16.0 | 381876.1 | 14.6 | 767957.4 | 29.5 | 673543.3 | 25.8 | 610195.3 | 23.4007534 |
Suitable | 1738402.5 | 66.7 | 1865194.5 | 71.5 | 1923659.4 | 73.8 | 2096607.3 | 80.4 | 2191042.3 | 84.0 | 2225712.1 | 85.4 | 1839630.8 | 70.5 | 1934044.9 | 74.2 | 2019973.6 | 77.4652068 | |
Low suitability | 243989.6 | 9.4 | 218026.5 | 8.4 | 220400.4 | 8.5 | 191733.0 | 7.4 | 180873.7 | 6.9 | 167256.0 | 6.4 | 227678.3 | 8.7 | 212116.2 | 8.1 | 224935.2 | 8.6261794 | |
Moderate suitability | 980264.0 | 37.6 | 844356.6 | 32.4 | 792590.7 | 30.4 | 621628.0 | 23.8 | 579928.0 | 22.2 | 530999.6 | 20.4 | 914305.3 | 35.1 | 782880.9 | 30.0 | 601159.6 | 23.0542387 | |
High suitability | 514148.9 | 19.7 | 802811.4 | 30.8 | 910668.3 | 34.9 | 1283246.3 | 49.2 | 1430240.5 | 54.8 | 1527456.5 | 58.6 | 697647.2 | 26.8 | 939047.8 | 36.0 | 1193878.7 | 45.7847887 |
Senegalia mellifera change between scenarios. The model was built under three CMIP6 climate models (MPI-ESM1.2-HR, UK-ESM1.0-LL and INM-CM5-0) and two periods of time (2041–2060 and 2061–2080).
Time period | Habitat suitability | Terminalia sericea climate models and emission scenarios | |||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | |||||||||||||||||
SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | ||
2041–2060 | Unsuitable | 1322301.6 | 50.3 | 1311514.4 | 49.9 | 1222655.3 | 46.5 | 1112278.2 | 42.3 | 1071213.7 | 40.7 | 1064833.6 | 40.5 | 1560107.7 | 59.3 | 1332964.2 | 50.7 | 1357883.9 | 51.6272526 |
Suitable | 1307867.3 | 49.7 | 1318654.5 | 50.1 | 1407513.6 | 53.5 | 1517890.6 | 57.7 | 1558955.2 | 59.3 | 1565335.3 | 59.5 | 1070061.2 | 40.7 | 1297204.7 | 49.3 | 1272284.9 | 48.3727474 | |
Low suitability | 434230.3 | 16.5 | 380103.7 | 14.5 | 206358.5 | 7.8 | 494565.9 | 18.8 | 464141.5 | 17.6 | 463103.6 | 17.6 | 392721.4 | 14.9 | 489267.8 | 18.6 | 474838.9 | 18.0535532 | |
Moderate suitability | 440573.2 | 16.8 | 464829.6 | 17.7 | 509748.7 | 19.4 | 453154.6 | 17.2 | 456716.5 | 17.4 | 450509.8 | 17.1 | 379554.9 | 14.4 | 419518.1 | 16.0 | 390109.9 | 14.8321249 | |
High suitability | 433063.8 | 16.5 | 473721.3 | 18.0 | 691406.5 | 26.3 | 570170.2 | 21.7 | 638097.2 | 24.3 | 651721.9 | 24.8 | 297784.9 | 11.3 | 388418.7 | 14.8 | 407336.1 | 15.4870693 | |
2061–2080 | Unsuitable | 1325043.9 | 50.4 | 1229514.5 | 46.7 | 1189522.7 | 45.2 | 1051173.3 | 40.0 | 1043700.3 | 39.7 | 1001239.4 | 38.1 | 1344660.9 | 51.1 | 1232228.1 | 46.8 | 1023041.5 | 38.8964175 |
Suitable | 1305125.0 | 49.6 | 1345490.6 | 51.2 | 1440646.2 | 54.8 | 1578995.6 | 60.0 | 1586468.6 | 60.3 | 1628929.4 | 61.9 | 1285508.0 | 48.9 | 1397940.8 | 53.2 | 1607127.4 | 61.1035825 | |
Low suitability | 323162.8 | 12.3 | 323162.8 | 12.3 | 413203.8 | 15.7 | 458821.0 | 17.4 | 462040.1 | 17.6 | 412148.8 | 15.7 | 489205.9 | 18.6 | 497000.9 | 18.9 | 453741.2 | 17.2514111 | |
Moderate suitability | 446716.4 | 17.0 | 459912.3 | 17.5 | 422069.2 | 16.0 | 456385.2 | 17.4 | 431728.7 | 16.4 | 364602.8 | 13.9 | 427356.4 | 16.2 | 405230.8 | 15.4 | 482171.0 | 18.3323225 | |
High suitability | 535245.8 | 20.4 | 562415.5 | 21.4 | 605373.3 | 23.0 | 663789.4 | 25.2 | 692699.8 | 26.3 | 852177.9 | 32.4 | 368945.7 | 14.0 | 495709.1 | 18.8 | 671215.1 | 25.5198489 |
Dichrostachys cinerea change between scenarios. The model was built under three CMIP6 climate models (MPI-ESM1.2-HR, UK-ESM1.0-LL and INM-CM5-0) and two periods of time (2041–2060 and 2061–2080).
Time period | Habitat suitability | Dichrostachys cinerea climate models and emission scenarios | |||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | |||||||||||||||||
SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | SSP245 (km²) | % | SSP370 (km²) | % | SSP585 (km²) | % | ||
2041–2060 | Unsuitable | 1358133.2 | 51.6 | 1294573.8 | 49.2 | 1166255.5 | 44.3 | 1119215.6 | 42.6 | 1043170.1 | 39.7 | 1042242.9 | 39.6 | 1397773.6 | 53.1 | 1240426.3 | 47.2 | 1272302.0 | 48.3733948 |
Suitable | 1272035.7 | 48.4 | 1335595.1 | 50.8 | 1463913.4 | 55.7 | 1510953.3 | 57.4 | 1586998.8 | 60.3 | 1587926.0 | 60.4 | 1232395.3 | 46.9 | 1389742.5 | 52.8 | 1357866.9 | 51.6266052 | |
Low suitability | 459844.2 | 17.5 | 379808.8 | 14.4 | 349981.1 | 13.3 | 352005.1 | 13.4 | 320151.9 | 12.2 | 322766.5 | 12.3 | 437220.2 | 16.6 | 416900.4 | 15.9 | 408706.1 | 15.5391564 | |
Moderate suitability | 437526.7 | 16.6 | 401174.3 | 15.3 | 331330.8 | 12.6 | 379313.4 | 14.4 | 324476.3 | 12.3 | 346149.1 | 13.2 | 314569.1 | 12.0 | 327187.6 | 12.4 | 322029.7 | 12.2436878 | |
High suitability | 374664.8 | 14.2 | 554612.0 | 21.1 | 782601.5 | 29.8 | 779634.7 | 29.6 | 942370.5 | 35.8 | 919010.4 | 34.9 | 480606.0 | 18.3 | 645654.5 | 24.5 | 627131.2 | 23.8437609 | |
2061–2080 | Unsuitable | 1306886.6 | 49.7 | 1207524.4 | 45.9 | 1199494.9 | 45.6 | 995404.2 | 37.8 | 932851.1 | 35.5 | 839478.1 | 31.9 | 1246120.7 | 47.4 | 1127882.9 | 42.9 | 898520.3 | 34.1620781 |
Suitable | 1323282.3 | 50.3 | 1422644.5 | 54.1 | 1430674.0 | 54.4 | 1634764.6 | 62.2 | 1697317.8 | 64.5 | 1790690.8 | 68.1 | 1384048.2 | 52.6 | 1502286.0 | 57.1 | 1731648.5 | 65.8379219 | |
Low suitability | 364018.4 | 13.8 | 301512.5 | 11.5 | 303053.5 | 11.5 | 310556.7 | 11.8 | 286373.0 | 10.9 | 251913.0 | 9.6 | 380750.7 | 14.5 | 311232.4 | 11.8 | 348467.2 | 13.2488521 | |
Moderate suitability | 369174.0 | 14.0 | 343872.7 | 13.1 | 345311.6 | 13.1 | 351739.7 | 13.4 | 296800.4 | 11.3 | 287946.6 | 10.9 | 342163.0 | 13.0 | 315575.3 | 12.0 | 422426.8 | 16.0608225 | |
High suitability | 590089.9 | 22.4 | 777259.3 | 29.6 | 782308.9 | 29.7 | 972468.3 | 37.0 | 1114144.4 | 42.4 | 1250831.2 | 47.6 | 661134.5 | 25.1 | 875478.4 | 33.3 | 960754.6 | 36.5282473 |
Summary tables of percentage change between current and climate change scenarios in the two periods of time.
Senegalia mellifera area percentage change | |||||||||
Time period | INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | ||||||
SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | |
(2041–2060) | 8.0 | 10.5 | 11.4 | 19.8 | 23.3 | 24.2 | 11.1 | 13.0 | 14.5 |
(2061–2080) | 10.7 | 15.5 | 17.8 | 24.4 | 28.0 | 29.4 | 14.5 | 18.2 | 21.5 |
Terminalia sericea area percentage change | |||||||||
Time period | INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | ||||||
SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | |
(2041–2060) | 11.9 | 12.2 | 15.6 | 19.8 | 21.4 | 21.6 | 2.8 | 11.4 | 10.5 |
(2061–2080) | 11.7 | 15.4 | 16.9 | 22.1 | 22.4 | 24.0 | 11.0 | 15.3 | 23.2 |
Dichrostachys cinerea area percentage change | |||||||||
Time period | INM-CM5-0 | UK-ESM1.0-LL | MPI-ESM1.2-HR | ||||||
SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | SSP245 (%) | SSP370 (%) | SSP585 (%) | |
(2041–2060) | 4.5 | 6.2 | 11.8 | 13.5 | 16.4 | 16.5 | 3.0 | 8.9 | 7.7 |
(2061–2080) | 6.4 | 10.2 | 10.5 | 18.3 | 20.6 | 24.2 | 8.7 | 13.2 | 21.9 |
Habitat suitability area per scenario Senegalia mellifera. This map illustrates variations in habitat suitability for Senegalia mellifera, featuring distinct colors to delineate areas deemed unsuitable or suitable for the species. The color spectrum is utilized to convey different levels of suitability: areas in dark green represent the highest suitability, while light green signifies low suitability. Unsuitable areas for the species are depicted in grey. The map serves as a visual guide to understand how the suitability of habitat changes across different scenarios and climate models for Senegalia mellifera.
Habitat suitability area per scenario Terminalia sericea. This map illustrates variations in habitat suitability for Terminalia sericea featuring distinct colors to delineate areas deemed unsuitable or suitable for the species. The color spectrum is utilized to convey different levels of suitability: areas in dark green represent the highest suitability, while light green signifies low suitability. Unsuitable areas for the species are depicted in grey. The map serves as a visual guide to understand how the suitability of habitat changes across different scenarios and climate models for Terminalia sericea.
Habitat suitability area per scenario Dichrostachys cinerea. This map illustrates variations in habitat suitability for Dichrostachys cinerea, featuring distinct colors to delineate areas deemed unsuitable or suitable for the species. The color spectrum is utilized to convey different levels of suitability: areas in dark green represent the highest suitability, while light green signifies low suitability. Unsuitable areas for the species are depicted in grey. The map serves as a visual guide to understand how the suitability of habitat changes across different scenarios and climate models for Dichrostachys cinerea.
This study predicted habitat suitability for the species Senegalia mellifera, Dichrostachys cinerea, and Terminalia sericea in the southern African region. Distribution predictions were carried out under three different climate models (INM-CM5-0, UK-ESM1.0-LL and MPI-ESM1.2-HR), three emission scenarios (SSP245, SSP370 and SSP585) for the periods 2041–2060 and 2061–2080, highlighting changes relative from the current state. We used the MaxEnt methodology to predict areas at higher or lower risk of bush encroachment and to identify the key drivers influencing the distribution of these species. This approach provides valuable insights to support future tasks related to climate change and its underlying drivers that affect the distribution patterns of plant species. All models outperformed chance, achieving AUC values greater than 0.736 in all cases, with the Terminalia sericea model (mean AUC = 0.836) achieving the highest value.
In general terms, all three species show a pattern of increasing habitability in previously uninhabited areas according to climate scenarios and time periods. Habitability rises as the expected concentration of greenhouse gasses increases for each scenario. The SSP245 scenario (medium pathway of future greenhouse gas emissions) assumes that climate protection measures are being taken, while registering the lowest increase in habitat suitability among the climate change scenarios. Conversely, the SSP585 scenario (less optimistic), which involves drastic emissions increase, correlates with the highest expansion in habitat suitability for all three species.
The results confirm that variables related to rainfall and temperature are the most important for modeling the distribution of these bush encroacher species. Dichrostachys cinerea and Terminalia sericea showed similar changes in distribution between the current and climate change scenarios, with the species expanding their range limits mainly in South Africa and Botswana. Meanwhile, Terminalia sericea it does not have a big spread in Namibia, but it increases in its habitat suitability. However, Dichrostachys cinerea incresases the distribution along the north of the case study, incresing the habitat suitability in the future scenarios. In the other hand, Senegalia mellifera does not respond in the exact same way. The model results record an unsuitable area of 14.6% in the SSP585 scenario (2061–2080), which in comparison to the other species (Terminalia sericea: 38.1%; Dichrostachys cinerea: 31.9%) indicates that it will be the most generalist and widespread of the three species in scenarios with drastic emissions.
The results in the percentage change between the current scenario and future scenarios also support the idea that Senegalia mellifera will cover the most area, with a maximum increase in suitable distribution area of up to 29.4%, obtaining a maximum suitability of 85.4% of the total area. This is followed by Dichrostachys cinerea, with a 24.2% of maximum increase in habitat suitability, with a maximum suitability of 68.1% of the total area. Lastly, Terminalia sericea shows a 24% of maximum increase, with a suitability of 61.9% of the total area. All these results indicate that the UK-ESM1.0-LL climate model and the SSP585 emission scenario for the period 2061–2080 register the highest bush encroachment. The increase in habitat suitability for the species is highest in this climate model compared to INM-CM5-0 and MPI-ESM1.2-HR, which show lower average habitat suitability.
The figure that summarises changes in habitat suitability according to the proposed scenarios shows differences and similarities in the distribution patterns of the three species (Figs
According to the model outputs, these findings imply that Senegalia mellifera is the most potentially bush encroacher for the near future changes, with a larger distribution area and better habitat suitability in most of the scenarios.
This statement does not rule out or ignore the invasive potential of the remaining species due to the climate changes.
These results are similar to findings reported in other studies concerning various species. Based on studies involving growth rings and stem diameter (
These findings support our model outputs and underscore the importance of the variables considered. In the current distribution scenarios, the model outputs generate distribution maps highly resemblant to the existing maps describing species presence boundaries (
According to present and future Köppen-Geiger climate classification maps by
Certainly, socio-ecological processes and biotic interactions pose a significant challenge to quantification and modelling in scientific research. Numerous intricate interactions between species and their environments elude precise measurement, and even models exhibiting impeccable AUC values and statistical metrics cannot ensure absolute perfection. Furthermore, the broad geographical scope of our estimates introduces inherent uncertainty into the assessment of climatic and physical suitability.
Species distribution models have emerged as invaluable tools for implementing conservation strategies or conducting studies on the impacts of climate change, particularly in expansive regions characterized by notable physical and bioclimatic gradients. However, these models predominantly rely on interpolation methods and lack a robust foundation in ecological process theory. Thus, there remains substantial room for improvement, with the development and implementation of novel methodologies being imperative for enhancing our understanding of biotic interactions.
Additionally, it is crucial for biogeographers and ecologists to recognize that the pattern of species association in joint species distribution models is influenced not only by actual biotic interactions but also by shared habitat preferences, mutual migration history, phylogenetic background, and collective responses to absent environmental drivers (
The objectives of this study were to investigate the potential impacts of climate change on the spatial distribution of these three bush encroachers, important woody species in southern African rangelands, and compare these findings with the existing literature. This study used INM-CM5-0, UK-ESM1.0-LL and MPI-ESM1.2-HR climate models, three emission scenarios (SSP245, SSP370 and SSP585) for the periods 2041–2060 and 2061–2080, combined with georeferenced species occurrence data. The results of the study indicate that the bioclimatic habitat of these species is expected to change significantly in response to climate change in the southern African savannas. Thus, supporting other studies that suggest the greening trends in southern Africa due to woody plant cover change. The main patterns of expansion are related to expected changes in temperature and precipitation over the century. According to the results shown above, Terminalia sericea, Dichrostachys cinerea and Senegalia mellifera improve their habitat suitability in almost all scenarios and time periods compared to the current one. Current results showed in Table
Senegalia mellifera has the biggest increase in the area of high habitat suitability, with a 29.4% increase in the 2061–2080 time period for SSP585 scenario compared to the current scenario, making it the species with the best conditions for establishment throughout southern Africa in the future. Dichrostachys cinerea follows with a 24.2% and 24% for Terminalia sericea. In summary, Senegalia mellifera emerges as the most stable species covering the largest area across all scenarios. Conversely, Dichrostachys cinerea and Terminalia sericea exhibit a similar distribution pattern, with Senegalia mellifera being the most favoured species in future scenarios.
Finally, to contribute to the improvement of savanna rangelands management, we encourage developing new studies to enable innovative management and adaptation plans in line with ongoing climate change in the southern Africa regions. Finally, we recommend conducting validation studies at local scales, analyzing shrub regeneration and its growth in areas that were found to be highly suitable in future scenarios. This should be supported by localized studies measuring the differences between dense and sparse coverages in relation to the ecosystem. This would be very useful for future soil management plans and would support many issues addressed in the Africa 2063 agenda.
We want to thank Colin Lindeque and their team for graphical support of the Namibia study site and information related to bush encroached zones in the country. Also, thanks to Fernando Alonso-Martin, who read the manuscript a lot of times and gave statistics tips. SBA project , supported by the European Commission’s Horizon 2020 Programme. Grant agreement ID: 101036401.
Conceptualization, B.G.J.; methodology, B.G.J.; software, B.G.J.; validation, J.B.G., P.G.F and V.B.F.J; formal analysis, J.B.G., P.G.F and V.B.F.J.; investigation, B.G.J.; data curation, B.G.J.; writing—original draft preparation, J.B.G.; writing—review and editing, P.G.F., V.B.F.J., C.N.M.J; supervision, P.G.F., V.B.F.J and M.A.F.
The data used in this article are openly accessible through the following link on the Zenodo repository: [https://zenodo.org/records/13735837]. This ensures transparency and allows for further analysis or replication of the study by other researchers.