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:

Negative effects of cattle on soil carbon and nutrient pools reversed by megaherbivores

Nature Sustainability volume 3, pages 360–366 (2020)Cite this article

Subjects

Abstract

Wild herbivore populations are declining in many African savannas, which is related to replacement by livestock (mainly cattle) and the loss of megaherbivores. Although some livestock management practices may be compatible with the conservation of native savanna biodiversity, the sustainability of these integrated wild herbivore/livestock management practices is unknown. For instance, how will these herbivore mixes influence key processes for the long-term functioning of savanna ecosystems, such as soil carbon, nitrogen and phosphorus pools and cycling? The Kenya Long-term Exclosure Experiment studies the ecosystem consequences of manipulating the presence and absence of wild herbivores and cattle at moderate densities in a ‘black cotton’ savanna. Here we show that after 20 years, cattle presence decreased total soil carbon and nitrogen pools, while the presence of megaherbivores (mainly elephants) increased these pools and even reversed the negative effects of cattle. Our results suggest that a mix of cattle at moderate densities and wild herbivores can be sustainable, provided that the assemblage of wild herbivores includes the largest species.

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

Access options

Buy this article

  • Purchase on Springer Link
  • Instant access to full article PDF
Buy now

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

Fig. 1: Herbivore effects on soil N.
Fig. 2: Herbivore treatment effects on soil C, N and P pools.
Fig. 3: Herbivore treatment effects on grass N and C/N ratio.
Fig. 4: Relationship between soil and grass.

Similar content being viewed by others

Data availability

The datasets collected and analyzed in this study are available in figshare at https://doi.org/10.6084/m9.figshare.11636595.v1, https://doi.org/10.6084/m9.figshare.11636577.v2 and https://doi.org/10.6084/m9.figshare.11636502.v1. Source data for Figs. 2–4 and Extended Data Figs. 1–4 and 6 are provided as Source Data files.

References

  1. du Toit, J. T. & Cumming, D. H. M. Functional significance of ungulate diversity in African savannas and the ecological implications of the spread of pastoralism. Biodivers. Conserv. 8, 1643–1661 (1999).

    Article  Google Scholar 

  2. Reid, R. Savannas of our Birth: People, Wildlife, and Change in East Africa (Univ. California Press, 2012).

  3. Veblen, K. E., Porensky, L. M., Riginos, C. & Young, T. P. Are cattle surrogate wildlife? Savanna plant community composition explained by total herbivory more than herbivore type. Ecol. Appl. 26, 1610–1623 (2016).

    Article  Google Scholar 

  4. du Toit, J. T., Kock, R. & Deutsch, J. Wild Rangelands: Conserving Wildlife while Maintaining Livestock in Semi-arid Ecosystems (Wiley-Blackwell, 2010).

  5. Ripple, W. J. et al. Collapse of the world’s largest herbivores. Sci. Adv. 1, e1400103 (2015).

    Article  Google Scholar 

  6. Malhi, Y. et al. Megafauna and ecosystem function from the Pleistocene to the Anthropocene. Proc. Natl Acad. Sci. USA 113, 838–846 (2016).

    Article  CAS  Google Scholar 

  7. Hempson, G. P., Archibald, S. & Bond, W. J. The consequences of replacing wildlife with livestock in Africa. Sci. Rep. 7, 17196 (2017).

    Article  CAS  Google Scholar 

  8. Young, T. P. et al. Relationships between cattle and biodiversity in multiuse landscape revealed by Kenya Long-term Exclosure Experiment. Rangel. Ecol. Manag. 71, 281–291 (2018).

    Article  Google Scholar 

  9. Keesing, F. et al. Consequences of integrating livestock and wildlife in an African savanna. Nat. Sustain. 1, 566–573 (2018).

    Article  Google Scholar 

  10. Asner, G. P., Vaughn, N., Smit, I. P. J. & Levick, S. Ecosystem-scale effects of megafauna in African savannas. Ecography 39, 240–252 (2016).

    Article  Google Scholar 

  11. Young, T. P., Palmer, T. M. & Gadd, M. E. Competition and compensation among cattle, zebras, and elephants in a semi-arid savanna in Laikipia, Kenya. Biol. Conserv. 122, 351–359 (2005).

    Article  Google Scholar 

  12. Kimuyu, D. M. et al. Influence of cattle on browsing and grazing wildlife varies with rainfall and presence of megaherbivores. Ecol. Appl. 27, 786–798 (2017).

    Article  Google Scholar 

  13. Augustine, D. J., McNaughton, S. J. & Frank, D. A. Feedbacks between soil nutrients and large herbivores in a managed savanna ecosystem. Ecol. Appl. 13, 1325–1337 (2003).

    Article  Google Scholar 

  14. Le Roux, E., Kerley, G. I. H. & Cromsigt, J. Megaherbivores modify trophic cascades triggered by fear of predation in an African savanna ecosystem. Curr. Biol. 28, 2493–2499.e3 (2018).

    Article  CAS  Google Scholar 

  15. Veldhuis, M. P., Gommers, M. I., Olff, H. & Berg, M. P. Spatial redistribution of nutrients by large herbivores and dung beetles in a savanna ecosystem. J. Ecol. 106, 422–433 (2018).

    Article  CAS  Google Scholar 

  16. van der Waal, C. et al. Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation. Oecologia 165, 1095–1107 (2011).

    Article  Google Scholar 

  17. Augustine, D. J. Long-term, livestock-mediated redistribution of nitrogen and phosphorus in an East African savanna. J. Appl. Ecol. 40, 137–149 (2003).

    Article  Google Scholar 

  18. Veblen, K. E. Savanna glade hotspots: plant community development and synergy with large herbivores. J. Arid Environ. 78, 119–127 (2012).

    Article  Google Scholar 

  19. Cech, P. G., Olde Venterink, H. & Edwards, P. J. N and P cycling in Tanzanian humid savanna: influence of herbivores, fire, and N2-fixation. Ecosystems 13, 1079–1096 (2010).

    Article  CAS  Google Scholar 

  20. Marshall, F. et al. Ancient herders enriched and restructured African grasslands. Nature 561, 387–390 (2018).

    Article  CAS  Google Scholar 

  21. Doughty, C. E. et al. Global nutrient transport in a world of giants. Proc. Natl Acad. Sci. USA 113, 868–873 (2016).

    Article  CAS  Google Scholar 

  22. Owen-Smith, N. Megaherbivores: The Influence of Very Large Body Size on Ecology (Cambridge Univ. Press, 1988).

  23. Kerley, G. et al. in Elephant Management: A Scientific Assessment for South Africa (eds Scholes, R. & Mennell, K.) 146–205 (Witwatersrand Univ. Press, 2008).

  24. Sitters, J., Edwards, P. J. & Olde Venterink, H. Increases of soil C, N, and P pools along an Acacia tree density gradient and their effects on trees and grasses. Ecosystems 16, 347–357 (2013).

    Article  CAS  Google Scholar 

  25. Blaser, W. J., Sitters, J., Hart, S. P., Edwards, P. J. & Olde Venterink, H. Facilitative or competitive effects of woody plants on understorey vegetation depend on N-fixation, canopy shape and rainfall. J. Ecol. 101, 1598–1603 (2013).

    Article  Google Scholar 

  26. Young, T., Okello, B., Kinyua, D. & Palmer, T. KLEE: A long-term multi-species herbivore exclusion experiment in Laikipia, Kenya. Afr. J. Range Forage Sci. 14, 94–102 (1997).

    Article  Google Scholar 

  27. Riginos, C. et al. Lessons on the relationship between livestock husbandry and biodiversity from the Kenya Long-term Exclosure Experiment (KLEE). Pastor. Res. Policy Pract. 2, 10 (2012).

    Article  Google Scholar 

  28. Charles, G. K., Porensky, L. M., Riginos, C., Veblen, K. E. & Young, T. P. Herbivore effects on productivity vary by guild: cattle increase mean productivity while wildlife reduce variability. Ecol. Appl. 27, 143–155 (2017).

    Article  Google Scholar 

  29. Riginos, C., Porensky, L. M., Veblen, K. E. & Young, T. P. Herbivory and drought generate short-term stochasticity and long-term stability in a savanna understory community. Ecol. Appl. 28, 323–335 (2018).

    Article  Google Scholar 

  30. Odadi, W. O., Okeyo-Owuor, J. B. & Young, T. P. Behavioural responses of cattle to shared foraging with wild herbivores in an East African rangeland. Appl. Anim. Behav. Sci. 116, 120–125 (2009).

    Article  Google Scholar 

  31. Kimuyu, D. M., Sensenig, R. L., Riginos, C., Veblen, K. E. & Young, T. P. Native and domestic browsers and grazers reduce fuels, fire temperatures, and acacia ant mortality in an African savanna. Ecol. Appl. 24, 741–749 (2014).

    Article  Google Scholar 

  32. Goheen, J. R. et al. Conservation lessons from large-mammal manipulations in East African savannas: the KLEE, UHURU, and GLADE experiments. Ann. N. Y. Acad. Sci. 1429, 31–49 (2018).

    Article  Google Scholar 

  33. Fox-Dobbs, K., Doak, D. F., Brody, A. K. & Palmer, T. M. Termites create spatial structure and govern ecosystem function by affecting N2 fixation in an East African savanna. Ecology 91, 1296–1307 (2010).

    Article  Google Scholar 

  34. Ritchie, M. E. & Raina, R. Effects of herbivores on nitrogen fixation by grass endophytes, legume symbionts and free-living soil surface bacteria in the Serengeti. Pedobiologia 59, 233–241 (2016).

    Article  Google Scholar 

  35. Sitters, J., Edwards, P. J., Suter, W. & Olde Venterink, H. O. Acacia tree density strongly affects N and P fluxes in savanna. Biogeochemistry 123, 285–297 (2015).

    Article  CAS  Google Scholar 

  36. Kelemu, S. et al. Detecting bacterial endophytes in tropical grasses of the Brachiaria genus and determining their role in improving plant growth. Afr. J. Biotechnol. 10, 965–976 (2011).

    Google Scholar 

  37. Hartnett, D. C., Potgieter, A. F. & Wilson, G. W. T. Fire effects on mycorrhizal symbiosis and root system architecture in southern African savanna grasses. Afr. J. Ecol. 42, 328–337 (2004).

    Article  Google Scholar 

  38. Craine, J. M. et al. Ecological interpretations of nitrogen isotope ratios of terrestrial plants and soils. Plant Soil 396, 1–26 (2015).

    Article  CAS  Google Scholar 

  39. Frank, D. A. & Evans, R. D. Effects of native grazers on grassland N cycling in Yellowstone National Park. Ecology 78, 2238–2248 (1997).

    Article  Google Scholar 

  40. Cech, P. G., Kuster, T., Edwards, P. J. & Olde Venterink, H. Effects of herbivory, fire and N2-fixation on nutrient limitation in a humid African savanna. Ecosystems 11, 991–1004 (2008).

    Article  CAS  Google Scholar 

  41. McNaughton, S. J., Banyikwa, F. F. & McNaughton, M. M. Promotion of the cycling of diet-enhancing nutrients by African grazers. Science 278, 1798–1800 (1997).

    Article  CAS  Google Scholar 

  42. Augustine, D. J. & McNaughton, S. J. Interactive effects of ungulate herbivores, soil fertility, and variable rainfall on ecosystem processes in a semi-arid savanna. Ecosystems 9, 1242–1256 (2006).

    Article  CAS  Google Scholar 

  43. Anderson, T. M., Ritchie, M. E. & McNaughton, S. J. Rainfall and soils modify plant community response to grazing in Serengeti National Park. Ecology 88, 1191–1201 (2007).

    Article  Google Scholar 

  44. Holdo, R. M. & Mack, M. C. Functional attributes of savanna soils: contrasting effects of tree canopies and herbivores on bulk density, nutrients and moisture dynamics. J. Ecol. 102, 1171–1182 (2014).

    Article  CAS  Google Scholar 

  45. van Langevelde, F. et al. Effects of fire and herbivory on the stability of savanna ecosystems. Ecology 84, 337–350 (2003).

    Article  Google Scholar 

  46. Sankaran, M., Ratnam, J. & Hanan, N. Woody cover in African savannas: the role of resources, fire and herbivory. Glob. Ecol. Biogeogr. 17, 236–245 (2008).

    Article  Google Scholar 

  47. Tobler, M. W., Cochard, R. & Edwards, P. J. The impact of cattle ranching on large-scale vegetation patterns in a coastal savanna in Tanzania. J. Appl. Ecol. 40, 430–444 (2003).

    Article  Google Scholar 

  48. Okigbo, B. N. in Ecology and Management of the World’s Savannas (eds Tothill, J. C. & Mott, J. J.) 95–113 (Australian Academy of Science, 1985).

  49. McClenachan, L., Cooper, A. B. & Dulvy, N. K. Rethinking trade-driven extinction risk in marine and terrestrial megafauna. Curr. Biol. 26, 1640–1646 (2016).

    Article  CAS  Google Scholar 

  50. Ahmad, N. in Vertisols and Technologies for their Management (eds Ahmad, N. & Mermut, A.) 1–41 (Elsevier, 1996).

  51. Bergstrom, B. J., Sensenig, R. L., Augustine, D. J. & Young, T. P. Searching for cover: soil enrichment and herbivore exclusion, not fire, enhance African savanna small-mammal abundance. Ecosphere 9, e02519 (2018).

    Article  Google Scholar 

  52. Schleppi, P., Conedera, M., Sedivy, I. & Thimonier, A. Correcting non-linearity and slope effects in the estimation of the leaf area index of forests from hemispherical photographs. Agric. For. Meteorol. 144, 236–242 (2007).

    Article  Google Scholar 

  53. Thimonier, A., Sedivy, I. & Schleppi, P. Estimating leaf area index in different types of mature forest stands in Switzerland: a comparison of methods. Eur. J. For. Res. 129, 543–562 (2010).

    Article  Google Scholar 

  54. R Core Team R: A Language and Environment for Statistical Computing Version 3.4.3 (2019); https://www.R-project.org/

  55. Pinheiro, J., Bates, D., DebRoy, S., Sarkar, D. & R Core Team. nlme: linear and nonlinear mixed effects models. R package version 3.1-144 (2020); https://CRAN.R-project.org/package=nlme

  56. Lenth, R. V. Least-Squares Means: the package. J. Stat. Softw. 69, 1–33 (2016).

    Article  Google Scholar 

Download references

Acknowledgements

This research was carried out under Government of Kenya research clearance permit no. NACOSTI/P/15/0830/4886. We thank F. Erii, J. Lochukuya, M. Namoni, J. Ekadeli, S. Ekuam and B. Kimiti for their invaluable assistance in the field. We also acknowledge the staff at Mpala Research Centre for their logistical support. The KLEE plots were built and maintained by grants from the Smithsonian Institution, The National Geographic Society (grants 4691-91 and 9106-12), the African Elephant Program of the US Fish and Wildlife Service (98210-0-G563) and the National Science Foundation (LTREB BSR-97-07477, 03-16402, 08-16453, 12-56004 and 12-56034). J.S. was funded by grants from the Research Foundation Flanders (FWO), grants 12N2615N and 12N2618N, and the Leopold III Fonds voor Natuuronderzoek en Natuurbehoud. Stable isotope facilities were supported by grants from FWO Hercules (HERC46) and VUB SRP2: Tracing and modelling of present and ancient global changes.

Author information

Authors and Affiliations

  1. Ecology and Biodiversity, Department Biology, Vrije Universiteit Brussel, Brussels, Belgium

    Judith Sitters & Harry Olde Venterink

  2. Mpala Research Centre, Nanyuki, Kenya

    Judith Sitters, Duncan M. Kimuyu & Truman P. Young

  3. Department of Natural Resources, Karatina University, Karatina, Kenya

    Duncan M. Kimuyu

  4. Department of Plant Sciences, University of California, Davis, CA, USA

    Truman P. Young

  5. Analytical, Environmental and Geo-Chemistry, Vrije Universiteit Brussel, Brussels, Belgium

    Philippe Claeys

Authors
  1. Judith Sitters
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Duncan M. Kimuyu
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Truman P. Young
    View author publications

    You can also search for this author in PubMed Google Scholar

  4. Philippe Claeys
    View author publications

    You can also search for this author in PubMed Google Scholar

  5. Harry Olde Venterink
    View author publications

    You can also search for this author in PubMed Google Scholar

Contributions

J.S., T.P.Y. and H.O.V. conceived the study. J.S., D.M.K. and T.P.Y. collected soil, plant and dung samples. D.M.K. and T.P.Y. provided additional data. J.S. and P.C. performed chemical analyses. J.S. and H.O.V. analysed the data and wrote the first draft of the manuscript. All authors contributed revisions.

Corresponding author

Correspondence to Judith Sitters.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

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

Extended data

Extended Data Fig. 1 Herbivore treatment effects on canopy cover.

a, Impact of herbivore treatments on canopy cover estimated with hemispherical photographs. Herbivore treatments (N = 3) include: no large herbivores (O), wild mesoherbivores (W), wild mega- and mesoherbivores (MW), only cattle (C), wild mesoherbivores and cattle (WC), wild mega- and mesoherbivores and cattle (MWC). Boxplots not sharing the same letter indicate significant differences between herbivore treatments. b, Linear regression showing the relationship between tree density and canopy cover. See ref. 31 for details on tree surveys.

Source data

Extended Data Fig. 2 Herbivore treatment effects on soil C, N and P pools.

Impact of herbivore treatments on plot-level soil C a, N b, and P pools c, calculated using the proportions of plot area located under and outside the canopy of A. drepanolobium trees. Herbivore treatments (N = 3) include: no large herbivores (O), wild mesoherbivores (W), wild mega- and mesoherbivores (MW), only cattle (C), wild mesoherbivores and cattle (WC), wild mega- and mesoherbivores and cattle (MWC). Boxplots not sharing the same letter indicate significant differences between herbivore treatments.

Source data

Extended Data Fig. 3 Herbivore treatment effects on grass P, C:P and N:P ratios.

Impact of herbivore treatments on live grass P concentrations a, C:P b, and N:P ratio c, outside (golden boxplots and points) and under (green boxplots and points) the canopy of A. drepanolobium trees. Herbivore treatments (N = 3) include: no large herbivores (O), wild mesoherbivores (W), wild mega- and mesoherbivores (MW), only cattle (C), wild mesoherbivores and cattle (WC), wild mega- and mesoherbivores and cattle (MWC).

Source data

Extended Data Fig. 4 Relationship between soil and grass.

Relationships between total soil P pool and live grass P concentrations a, and C:P ratio b, Herbivore treatments include: no large herbivores (O), wild mesoherbivores (W), wild mega- and mesoherbivores (MW), only cattle (C), wild mesoherbivores and cattle (WC), wild mega- and mesoherbivores and cattle (MWC). Sampled locations are outside (golden points) and under (green points) the canopy of A. drepanolobium trees.

Source data

Extended Data Fig. 5 The Kenya Long-term Exclosure Experiment (KLEE).

Schematic of the Kenya Long-term Exclosure Experiment (KLEE) plots. The letters inside each plot indicate the herbivore treatments.

Extended Data Fig. 6 Coverage of clipped grass species in each KLEE plot.

Coverage of different grass species in the two 25 x 25 cm subplots (one under the canopy and one outside the canopy of an A. drepanolobium tree) we clipped in each KLEE plot. Herbivore treatments include: no large herbivores (O), wild mesoherbivores (W), wild mega- and mesoherbivores (MW), only cattle (C), wild mesoherbivores and cattle (WC), wild mega- and mesoherbivores and cattle (MWC). N = 6 per treatment.

Source data

Supplementary information

Supplementary Information

Supplementary Notes 1–4, Tables 1 and 2, Figs. 1–6 and references.

Reporting Summary

Supplementary Data

Statistical source data soil and grass nutrients.

Source data

Source Data Fig. 2

Statistical source data soil C, N and P pools.

Source Data Fig. 3

Statistical source data grass N and C/N ratio.

Source Data Fig. 4

Statistical source data soil and grass N.

Source Data Extended Data Fig. 1

Statistical source data tree density and canopy cover.

Source Data Extended Data Fig. 2

Statistical source data plot-level soil C, N and P pools.

Source Data Extended Data Fig. 3

Statistical source data grass P, C/P and N/P ratios.

Source Data Extended Data Fig. 4

Statistical source data soil and grass P.

Source Data Extended Data Fig. 6

Statistical source data grass composition clipped sub-plots.

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Sitters, J., Kimuyu, D.M., Young, T.P. et al. Negative effects of cattle on soil carbon and nutrient pools reversed by megaherbivores. Nat Sustain 3, 360–366 (2020). https://doi.org/10.1038/s41893-020-0490-0

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1038/s41893-020-0490-0

This article is cited by

Search

Advanced 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