Skip to main content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Article
  • Published:

Limited increases in savanna carbon stocks over decades of fire suppression

Abstract

Savannas cover a fifth of the land surface and contribute a third of terrestrial net primary production, accounting for three-quarters of global area burned and more than half of global fire-driven carbon emissions1,2,3. Fire suppression and afforestation have been proposed as tools to increase carbon sequestration in these ecosystems2,4. A robust quantification of whole-ecosystem carbon storage in savannas is lacking however, especially under altered fire regimes. Here we provide one of the first direct estimates of whole-ecosystem carbon response to more than 60 years of fire exclusion in a mesic African savanna. We found that fire suppression increased whole-ecosystem carbon storage by only 35.4 ± 12% (mean ± standard error), even though tree cover increased by 78.9 ± 29.3%, corresponding to total gains of 23.0 ± 6.1 Mg C ha−1 at an average of about 0.35 ± 0.09 Mg C ha−1 year−1, more than an order of magnitude lower than previously assumed4. Frequently burned savannas had substantial belowground carbon, especially in biomass and deep soils. These belowground reservoirs are not fully considered in afforestation or fire-suppression schemes but may mean that the decadal sequestration potential of savannas is negligible, especially weighed against concomitant losses of biodiversity and function.

This is a preview of subscription content, access via your institution

Access options

Buy this article

Prices may be subject to local taxes which are calculated during checkout

Fig. 1: Vegetation characteristics of experimental burn plots in a mesic savanna in Kruger National Park, South Africa.
Fig. 2: Changes in whole-ecosystem carbon storage across fire treatments.
Fig. 3: Changes in SOC with tree cover and soil sand content.

Similar content being viewed by others

Data and code availability

Data and code are available in the Dryad Digital Repository: https://doi.org/10.5061/dryad.pg4f4qrr5Source data are provided with this paper.

References

  1. Giglio, L., Schroeder, W. & Justice, C. O. The collection 6 MODIS active fire detection algorithm and fire products. Remote Sens. Environ. 178, 31–41 (2016).

    Article  ADS  PubMed  PubMed Central  Google Scholar 

  2. Grace, J., José, J. S., Meir, P., Miranda, H. S. & Montes, R. A. Productivity and carbon fluxes of tropical savannas. J. Biogeogr. 33, 387–400 (2006).

    Article  Google Scholar 

  3. Van Der Werf, G. R. et al. Global fire emissions estimates during 1997–2016. Earth Syst. Sci. Data 9, 697–720 (2017).

    Article  ADS  Google Scholar 

  4. Bastin, J.-F. et al. The global tree restoration potential. Science 365, 76–79 (2019).

    Article  ADS  CAS  PubMed  Google Scholar 

  5. Russell-Smith, J. et al. Opportunities and challenges for savanna burning emissions abatement in southern Africa. J. Environ. Manage. 288, 112414 (2021).

    Article  CAS  PubMed  Google Scholar 

  6. Andela, N. et al. A human-driven decline in global burned area. Science 356, 1356–1362 (2017).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  7. Wu, C. et al. Historical and future global burned area with changing climate and human demography. One Earth 4, 517–530 (2021).

    Article  ADS  Google Scholar 

  8. Pellegrini, A. F. A. et al. Fire frequency drives decadal changes in soil carbon and nitrogen and ecosystem productivity. Nature 553, 194–198 (2018).

    Article  ADS  CAS  PubMed  Google Scholar 

  9. Higgins, S. I. et al. Effects of four decades of fire manipulation on woody vegetation structure in savanna. Ecology 88, 1119–1125 (2007).

    Article  PubMed  Google Scholar 

  10. Staver, A. C., Archibald, S. & Levin, S. A. The global extent and determinants of savanna and forest as alternative biome states. Science 334, 230–232 (2011).

    Article  ADS  CAS  PubMed  MATH  Google Scholar 

  11. Shi, Z. et al. The age distribution of global soil carbon inferred from radiocarbon measurements. Nat. Geosci. 13, 555–559 (2020).

    Article  ADS  CAS  Google Scholar 

  12. Pellegrini, A. F. A., Hedin, L. O., Staver, A. C. & Govender, N. Fire alters ecosystem carbon and nutrients but not plant nutrient stoichiometry or composition in tropical savanna. Ecology 96, 1275–1285 (2015).

    Article  PubMed  Google Scholar 

  13. Tilman, D. et al. Fire suppression and ecosystem carbon storage. Ecology 81, 2680–2685 (2000).

    Article  Google Scholar 

  14. Mokany, K., Raison, R. J. & Prokushkin, A. S. Critical analysis of root:shoot ratios in terrestrial biomes. Glob. Change Biol. 12, 84–96 (2006).

    Article  ADS  Google Scholar 

  15. de Miranda, S. D. C. et al. Regional variations in biomass distribution in Brazilian savanna woodland. Biotropica 46, 125–138 (2014).

    Article  Google Scholar 

  16. Wigley, B. J., Cramer, M. D. & Bond, W. J. Sapling survival in a frequently burnt savanna: mobilisation of carbon reserves in Acacia karroo. Plant Ecol. 203, 1 (2009).

    Article  Google Scholar 

  17. Sankaran, M. et al. Determinants of woody cover in African savannas. Nature 438, 846–849 (2005).

    Article  ADS  CAS  PubMed  Google Scholar 

  18. Staver, A. C., Botha, J. & Hedin, L. Soils and fire jointly determine vegetation structure in an African savanna. New Phytol. 216, 1151–1160 (2017).

    Article  CAS  PubMed  Google Scholar 

  19. Zhou, Y., Wigley, B. J., Case, M. F., Coetsee, C. & Staver, A. C. Rooting depth as a key woody functional trait in savannas. New Phytol. 227, 1350–1361 (2020).

    Article  PubMed  Google Scholar 

  20. Govender, N., Trollope, W. S. W., Van, & Wilgen, B. W. The effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in South Africa. J. Appl. Ecol. 43, 748–758 (2006).

    Article  Google Scholar 

  21. Colgan, M. S., Asner, G. P. & Swemmer, T. Harvesting tree biomass at the stand level to assess the accuracy of field and airborne biomass estimation in savannas. Ecol. Appl. 23, 1170–1184 (2013).

    Article  PubMed  Google Scholar 

  22. Davies, A. B. & Asner, G. P. Elephants limit aboveground carbon gains in African savannas. Glob. Change Biol. 25, 1368–1382 (2019).

    Article  ADS  Google Scholar 

  23. Butnor, J. R. et al. Utility of ground-penetrating radar as a root biomass survey tool in forest systems. Soil Sci. Soc. Am. J. 67, 1607–1615 (2003).

    Article  ADS  CAS  Google Scholar 

  24. Staver, A. C., Wigley-Coetsee, C. & Botha, J. Grazer movements exacerbate grass declines during drought in an African savanna. J. Ecol. 107, 1482–1491 (2019).

    Article  Google Scholar 

  25. Ryan, C. M., Williams, M. & Grace, J. Above- and belowground carbon stocks in a miombo woodland landscape of Mozambique. Biotropica 43, 423–432 (2011).

    Article  Google Scholar 

  26. Swezy, D. M. & Agee, J. K. Prescribed-fire effects on fine-root and tree mortality in old-growth ponderosa pine. Can. J. For. Res. 21, 626–634 (1991).

    Article  Google Scholar 

  27. Canadell, J. et al. Maximum rooting depth of vegetation types at the global scale. Oecologia 108, 583–595 (1996).

    Article  ADS  CAS  PubMed  Google Scholar 

  28. Coetsee, C., Bond, W. J. & February, E. C. Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover. Oecologia 162, 1027–1034 (2010).

    Article  ADS  PubMed  Google Scholar 

  29. Holdo, R. M., Mack, M. C. & Arnold, S. G. Tree canopies explain fire effects on soil nitrogen, phosphorus and carbon in a savanna ecosystem. J. Veg. Sci. 23, 352–360 (2012).

    Article  Google Scholar 

  30. Lloyd, J. et al. Contributions of woody and herbaceous vegetation to tropical savanna ecosystem productivity: a quasi-global estimate. Tree Physiol. 28, 451–468 (2008).

    Article  PubMed  Google Scholar 

  31. Wigley, B. J., Augustine, D. J., Coetsee, C., Ratnam, J. & Sankaran, M. Grasses continue to trump trees at soil carbon sequestration following herbivore exclusion in a semiarid African savanna. Ecology 101, e03008 (2020).

    Article  PubMed  Google Scholar 

  32. Khomo, L., Trumbore, S., Bern, C. R. & Chadwick, O. A. Timescales of carbon turnover in soils with mixed crystalline mineralogies. Soil 3, 17–30 (2017).

    Article  ADS  CAS  Google Scholar 

  33. Six, J., Conant, R. T., Paul, E. A. & Paustian, K. Stabilization mechanisms of soil organic matter: implications for C-saturation of soils. Plant Soil 241, 155–176 (2002).

    Article  CAS  Google Scholar 

  34. Abreu, R. C. R. et al. The biodiversity cost of carbon sequestration in tropical savanna. Sci. Adv. 3, e1701284 (2017).

    Article  ADS  PubMed  PubMed Central  Google Scholar 

  35. Bond, W. J., Stevens, N., Midgley, G. F. & Lehmann, C. E. The trouble with trees: afforestation plans for Africa. Trends Ecol. Evol. 34, 963–965 (2019).

    Article  PubMed  Google Scholar 

  36. West, T. A., Börner, J. & Fearnside, P. M. Climatic benefits from the 2006–2017 avoided deforestation in Amazonian Brazil. Front. For. Glob. Change 2, 52 (2019).

    Article  Google Scholar 

  37. Aleman, J. C., Blarquez, O. & Staver, C. A. Land-use change outweighs projected effects of changing rainfall on tree cover in sub-Saharan Africa. Glob. Change Biol. 22, 3013–3025 (2016).

    Article  ADS  Google Scholar 

  38. Huang, J., Yu, H., Guan, X., Wang, G. & Guo, R. Accelerated dryland expansion under climate change. Nat. Clim. Change 6, 166–171 (2016).

    Article  ADS  Google Scholar 

  39. Ratajczak, Z., Nippert, J. B. & Collins, S. L. Woody encroachment decreases diversity across North American grasslands and savannas. Ecology 93, 697–703 (2012).

    Article  PubMed  Google Scholar 

  40. Smit, I. P. & Prins, H. H. Predicting the effects of woody encroachment on mammal communities, grazing biomass and fire frequency in African savannas. PLoS One 10, e0137857 (2015).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  41. Huxman, T. E. et al. Ecohydrological implications of woody plant encroachment. Ecology 86, 308–319 (2005).

    Article  Google Scholar 

  42. Hermoso, V., Regos, A., Morán-Ordóñez, A., Duane, A. & Brotons, L. Tree planting: a double-edged sword to fight climate change in an era of megafires. Glob. Change Biol. 27, 3001–3003 (2021).

    Article  Google Scholar 

  43. Venter F. A. Classification of Land for Management Planning in the Kruger National Park. PhD thesis, Univ. South Africa (1990).

  44. Biggs, R., Biggs, H. C., Dunne, T. T., Govender, N. & Potgieter, A. L. F. Experimental burn plot trial in the Kruger National Park: history, experimental design and suggestions for data analysis. Koedoe 46, 15 (2003).

    Article  Google Scholar 

  45. Codron, J. et al. Taxonomic, anatomical, and spatio-temporal variations in the stable carbon and nitrogen isotopic compositions of plants from an African savanna. J. Archaeol. Sci. 32, 1757–1772 (2005).

    Article  Google Scholar 

  46. Zhou, Y., Boutton, T. W. & Ben Wu, X. Soil carbon response to woody plant encroachment: importance of spatial heterogeneity and deep soil storage. J. Ecol. 105, 1738–1749 (2017).

    Article  CAS  Google Scholar 

  47. Sheldrick B. & Wang C. In Soil Sampling and Methods of Analysis (ed. Carter, M. R.) 499–511 (CRC Press, 1993).

  48. Butnor, J. R. et al. Surface-based GPR underestimates below-stump root biomass. Plant Soil 402, 47–62 (2016).

    Article  CAS  Google Scholar 

  49. Pau, G., Fuchs, F., Sklyar, O., Boutros, M. & Huber, W. EBImage—an R package for image processing with applications to cellular phenotypes. Bioinformatics 26, 979–981 (2010).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  50. Hirano, Y. et al. Limiting factors in the detection of tree roots using ground-penetrating radar. Plant Soil 319, 15–24 (2009).

    Article  CAS  Google Scholar 

  51. Popescu, S. C. & Wynne, R. H. Seeing the trees in the forest. Photogramm. Eng. Remote Sensing 70, 589–604 (2004).

    Article  Google Scholar 

  52. Case, M. F., Wigley-Coetsee, C., Nzima, N., Scogings, P. F. & Staver, A. C. Severe drought limits trees in a semi-arid savanna. Ecology 100, e02842 (2019).

    Article  PubMed  Google Scholar 

  53. Beucher S. & Meyer F. In Mathematical Morphology in Image Processing (ed. Dougherty, E. R.) 433–481 (CRC Press, 1993).

  54. Nickless, A., Scholes, R. J. & Archibald, S. A method for calculating the variance and confidence intervals for tree biomass estimates obtained from allometric equations. S. Afr. J. Sci. 107, 1–10 (2011).

    Article  Google Scholar 

  55. Plowright A. & Roussel J.-R. ForestTools: analyzing remotely sensed forest data. R package version 0.2.1. https://CRAN.R-project.org/package=ForestTools (2020).

  56. Hijmans R. J. raster: geographic data analysis and modeling. R package version 3.3-7. https://CRAN.R-project.org/package=raster (2020).

  57. Penman J. et al. (eds) Good Practice Guidance for Land Use, Land-Use Change and Forestry (Intergovernmental Panel on Climate Change, 2003).

  58. Kuznetsova, A., Brockhoff, P. & Christensen, R. lmerTest package: tests in linear mixed effects models. J. Stat. Softw. 82, 1–26 (2017).

    Article  Google Scholar 

Download references

Acknowledgements

We gratefully acknowledge the logistical support provided by South African National Parks staff in Kruger National Park. Y.Z. was supported by a G. Evelyn Hutchinson Environmental Postdoctoral Fellowship from the Yale Institute for Biospheric Studies, A.C.S. was partially supported by a grant from the United States National Science Foundation (NSF MSB-1802453) and by funding from Yale University, J.S., P.B.B., E.G.H. and A.B.D. from Harvard University, and J.R.B. from the USDA Forest Service, Southern Research Station.

Author information

Authors and Affiliations

Authors

Contributions

Conceptualization: Y.Z. and A.C.S.; methodology: Y.Z., J.S., J.R.B., P.B.B., A.B.D. and A.C.S.; investigation: Y.Z., C.C., M.F.C., E.G.H., A.B.D. and A.C.S.; visualization: Y.Z. and A.C.S.; funding acquisition: A.B.D. and A.C.S.; project administration: A.B.D. and A.C.S.; supervision: A.B.D. and A.C.S.; writing — original draft: Y.Z. and A.C.S.; writing — review and editing: Y.Z., J.S., J.R.B., C.C., P.B.B., M.F.C., E.G.H., A.B.D. and A.C.S.

Corresponding authors

Correspondence to Yong Zhou or A. Carla Staver.

Ethics declarations

Competing interests

The authors declare no competing interests.

Peer review

Peer review information

Nature thanks Sebastian Dötterl, Niall Hanan and Douglas Morton for their contribution to the peer review of this work. Peer reviewer reports are available.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data figures and tables

Extended Data Fig. 1 An example showing belowground to aboveground biomass allocation for resprouting Terminalia sericea.

a, b, Five T. sericea individuals that have experienced annual burning were excavated in the Pretoriuskop landscape in Kruger National Park, South Africa. c, The difference between aboveground and belowground biomass and the ratio of belowground to aboveground biomass was 19.5. The box plots show medians (that is, 50th percentile), 25th and 75th percentiles, and the approximate 95% confidence interval for five replicates.

Extended Data Fig. 2 Maps showing the study site.

Maps showing the locations of different fire treatments (that is, annual, triennial and unburned) examined in this study and located in each string (Fayi, Kambeni, Numbi and Shabeni) across the Pretoriuskop landscape at Kruger National Park, South Africa. Base map for South Africa modified from Natural Earth.

Extended Data Fig. 3 Changes in SOC storage and soil δ13C across different fire treatments throughout the 60-cm soil column.

Effects of fire treatments on total SOC storage (Mg C ha−1) (a), soil δ13C (‰) (b), C3-derived SOC storage (that is, from woody plants) (Mg C ha−1) (c) and C4-derived SOC storage (that is, from grasses) (Mg C ha−1) (d). Values are mean ± standard errors (n = 4).

Extended Data Fig. 4 Long-term monitoring of grass fuel loads and their correlation to LiDAR-derived mean grass height.

ac, Grass fuel loads (kg ha−1) for annual (a), triennial (b) and April B2 (that is, burning in April for every two years, as a proxy for unburned) (c) treatments from 1982 to 2009 for different strings at the Pretoriuskop landscape in Kruger National Park, South Africa. Disconnected lines indicate missing data for specific years. d, The correlation between averaged grass fuel loads from 1982 to 2009 and LiDAR-derived mean grass heights (m) (R2 = 0.38, P = 0.03). The mean grass height was calculated by averaging heights of pixels that range from 0.05 to 0.5 m in the CHM derived from LiDAR. Please note especially that, in panel d, LiDAR-derived mean grass height was estimated from the unburned treatment itself, but that field-estimated grass fuel load was estimated from the April B2 treatment as a proxy (as grass fuel load is not routinely measured for the unburned treatment).

Extended Data Fig. 5 The uncertainty of coarse lateral and taproot biomass estimates.

a, The uncertainty of coarse lateral and taproot biomass for each treatment replicate. Error bars indicate the 95% confidence interval for coarse lateral and taproot biomass estimates derived from fitting regression lines (see Supplementary Figs. 5 and 10). Coarse-lateral-root biomass estimates were significantly correlated with taproot biomass estimates (R2 = 0.75, P < 0.001). Letters F, K, N and S indicate Fayi, Kambeni, Numbi and Shabeni strings at the Pretoriuskop landscape in Kruger National Park, South Africa; letters A, T and U indicate annual, triennial and unburned treatments. b, c The uncertainty of (that is, lower bound, mean and upper bound) coarse lateral and taproot biomass across different fire treatments. The box plots show medians (that is, 50th percentile), 25th and 75th percentiles, and the 95% confidence interval for four replicates. Points in b and c indicate outliers.

Extended Data Fig. 6 Depth distribution of coarse-lateral-root biomass across fire treatments and soil sand content.

a, Depth distribution of the GPR index (% in the number of pixels above the threshold for root detections) as an indicator of coarse-lateral-root biomass allocation throughout the soil column across different fire treatments at each string. Horizontal lines indicate the depth (cm) at which the GPR index reaches 50% of the total detections in the 60-cm soil column. b, Effects of fire treatment on the depth distribution of coarse-lateral-root biomass (P = 0.51). The box plots show medians (that is, 50th percentile), 25th and 75th percentiles, and the 95% confidence interval for four replicates. c, The correlation between soil sand content (%) and depth distribution of coarse-lateral-root biomass (R2 = 0.61, P = 0.003). The regression line indicates the significant linear fit and the shaded bands illustrate the 95% confidence interval of the linear fit.

Extended Data Fig. 7 The correlation between ratio of belowground to aboveground carbon storage and tree cover (%) (R2 = 0.83, P < 0.0001).

The regression line indicates the significant linear fit and the shaded bands illustrate the 95% confidence interval of the linear fit.

Extended Data Fig. 8 The validation of the object-based method to estimate aboveground woody biomass.

a, The correlation between LiDAR-derived stem density for trees with height > 5m (trees ha−1) and field-measured stem density (trees ha−1). The field-measured stem density was from ref. 52, which surveyed tree heights in eight 10-m-radius plots at each annual, triennial and unburned treatment in Kambeni, Numbi and Shabeni strings at the Pretoriuskop landscape in Kruger National Park, South Africa. The regression line indicates the significant linear fit and the shaded bands illustrate the 95% confidence interval of the linear fit. The dashed line indicates the 1:1 line. b, Differences in aboveground woody biomass between allometric-derived, object-based and plot-averaged estimates. The allometric-derived biomass estimation was on the basis of species-specific allometric equations developed in ref. 54, which predict aboveground woody biomass from DBH. This estimation was calculated for trees with DBH > 5 cm in each 10 × 10-m plot. The plot-averaged LiDAR biomass was estimated using an allometric equation derived from on-the-ground plot-level sampling relating aboveground woody biomass to LiDAR-derived canopy height and canopy area (aboveground woody biomass = −11.5 + 25.8 * canopy height * canopy area); please refer to ref. 21 for more details. The canopy height and canopy area were averaged across pixels with height > 0.5 m in each 30-m-radius plot. The box plots show medians (that is, 50th percentile), 25th and 75th percentiles, and the 95% confidence interval for four replicates. Points in b indicate outliers.

Extended Data Table 1 Results comparing the relative effect of different fire treatments on each component of ecosystem carbon storage from linear mixed-effects models using fire treatments as the fixed effect and string as a random effect

Supplementary information

Supplementary Information

This file contains Supplementary Figures 1–10 and Supplementary Tables 1–4.

Peer Review File

Source data

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Zhou, Y., Singh, J., Butnor, J.R. et al. Limited increases in savanna carbon stocks over decades of fire suppression. Nature 603, 445–449 (2022). https://doi.org/10.1038/s41586-022-04438-1

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1038/s41586-022-04438-1

This article is cited by

Comments

By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.

Search

Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing