Skip to main content

Thank you for visiting 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:

Consequences of integrating livestock and wildlife in an African savanna


Globally, most wildlife lives outside of protected areas, creating potential conflicts between the needs of wildlife and the needs of humans. East African savannas epitomize this challenge, providing habitat for wildlife such as giraffes and elephants as well as for people and their livestock. Conflicts over land use are common, leading to the assumption of a necessary trade-off between wildlife and livestock management. Here, we show that the integration of livestock and wildlife in a large region of central Kenya can have ecological benefits, reducing the abundance of ticks and improving forage. These ecological benefits can be complemented by economic ones when property owners derive income both from wildlife through tourism and from livestock through meat and dairy production. Our results suggest that under specific ecological, economic and social conditions, integrating livestock with wildlife can provide benefits for the environment and for human well-being in African savannas.

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: Tick abundance in relation to management.
Fig. 2: Effects of ecological integration of livestock and wildlife on vegetation.
Fig. 3: Effects of livestock on wildlife abundance.
Fig. 4: Diversity of income sources based on economic classification (relative proportion of income derived from livestock versus wildlife on each property). See main text for details.
Fig. 5: Ecological classification (ln wildlife/livestock ratio) versus economic classification for each of the 23 properties surveyed.

Similar content being viewed by others

Data availability

A summary table containing data used in these analyses is provided in the Supplementary Information (Extended Data Table 7).


  1. Ogutu, J. O. et al. Extreme wildlife declines and concurrent increase in livestock numbers in Kenya: what are the causes? PLoS ONE 11, e0163249 (2016).

    Article  Google Scholar 

  2. Terrestrial Protected Areas (% of Total Land Area) (United Nations Environmental Program and the World Conservation Monitoring Centre, 2016);

  3. Western, D., Russell, S. & Cuthill, I. The status of wildlife in protected areas compared to non-protected areas of Kenya. PLoS ONE 4, e6140 (2009).

    Article  Google Scholar 

  4. World Development Indicators (World Bank, 2016);

  5. Herrero, M., Thornton, P. K., Gerber, P. & Reid, R. S. Livestock, livelihoods and the environment: understanding the trade-offs. Curr. Opin. Environ. Sustain. 1, 111–120 (2009).

    Article  Google Scholar 

  6. de Leeuw, J. et al. Distribution and diversity of wildlife in northern Kenya in relation to livestock and permanent water points. Biol. Conserv. 100, 297–306 (2001).

    Article  Google Scholar 

  7. Wildlife Conservation by Sustainable Use (eds H. H. T. Prins et al.) 51–80 (Springer, Dordrecht, 2000).

  8. Odadi, W. O., Karachi, M. K., Abdulrazak, S. A. & Young, T. P. African wild ungulates compete with or facilitate cattle depending on season. Science 333, 1753–1755 (2011).

    Article  CAS  Google Scholar 

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

    Article  Google Scholar 

  10. Reid, R. S. Savannas of Our Birth: People, Wildlife, and Change in East Africa (Univ. of California Press, Oakland, 2012).

  11. Allan, B. F. et al. Can integrating wildlife and livestock enhance ecosystem services in central Kenya? Front. Ecol. Environ. 15, 328–335 (2017).

    Article  Google Scholar 

  12. Butt, B. & Turner, M. D. Clarifying competition: the case of wildlife and pastoral livestock in East Africa. Pastoralism 2, 9 (2012).

    Article  Google Scholar 

  13. The Contribution of the Rural Economy of Laikipia as the Basis of a Model County (Laikipia Wildlife Forum, 2013);

  14. Economic Survey 2016 (Kenya National Bureau of Statistics, 2016);

  15. Jongejan, F. & Uilenberg, G. The global importance of ticks. Parisitology 129, S3–S14 (2004).

    Article  Google Scholar 

  16. Keesing, F., Allan, B. F., Young, T. P. & Ostfeld, R. S. Effects of wildlife and cattle on tick abundance in central Kenya. Ecol. Appl. 23, 1410–1418 (2013).

    Article  Google Scholar 

  17. Walker, J. B., Horak, I. G. & Keirans, J. E. The Genus Rhipicephalus (Acari, Ixodidae): a Guide to the Brown Ticks of the World (Cambridge Univ. Press, Cambridge, 2005).

  18. Porensky, L. M. & Veblen, K. E. Generation of ecosystem hotspots using short-term cattle corrals in an African savanna. Rangeland Ecol. Manag. 68, 131–141 (2015).

    Article  Google Scholar 

  19. Ryan, Z. Establishment and Evaluation of a Livestock Early Warning System for Laikipia, Kenya (Texas A & M University, College Station, 2005).

    Google Scholar 

  20. Lengoiboni, M., Bregt, A. K. & van der Molen, P. Pastoralism within land administration in Kenya: the missing link. Land Use Policy 27, 579–588 (2010).

    Article  Google Scholar 

  21. Systematic Aerial Sample Survey of Laikipia County (Laikipia Wildlife Forum, 2016);

  22. Augustine, D. J., Veblen, K. E., Goheen, J. R., Riginos, C. & Young, T. P. Pathways for positive cattle-wildlife interactions in semiarid rangelands. Smithson. Contr. Zool. 661, 55–71 (2011).

    Article  Google Scholar 

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

    Article  Google Scholar 

  24. Wesangula, D. Two rangers shot dead in Kenya’s Laikipia conservation area. The Guardian (6 June 2017);

  25. 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 

  26. Keesing, F. Ecological Interactions Between Small Mammals, Large Mammals, and Vegetation in a Tropical Savanna of Central Kenya. PhD thesis, Univ. of California, Berkeley (1997).

  27. Kinnaird, M. F., O’Brien, T. & Strindberg, S. Assessment of Aerial Sample Count Surveys as a Monitoring Method to Track Changes in Livestock and Wildlife Across Laikipia County (2012);

  28. Estes, R. D The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates (Univ. of California, Oakland, 1991).

    Google Scholar 

  29. Falco, R. C. & Fish, D. A comparison of methods for sampling the deer tick, Ixodes dammini, in a Lyme disease endemic area. Exp. Appl. Acarol. 14, 165–173 (1992).

    Article  CAS  Google Scholar 

  30. Xie, P. & Arkin, P. A. Global precipitation: a 17-year monthly analysis based on gauge observations, satellite estimates, and numerical model outputs. Bull. Am. Meteorol. Soc. 78, 2539–2558 (1997).

    Article  Google Scholar 

  31. ArcGIS Desktop, release 10 (ESRI, 2011);

  32. Walker, A. R. et al. Ticks of Domestic Animals in Africa: A Guide to Identification of Species. Bioscience Reports (2003).

  33. Okello-Onen, J. Taxonomy of African Ticks: an Identification Manual (ICIPE, Nairobi, Kenya, 1999).

    Google Scholar 

  34. Muturi, J. E., Kim, C.-H., Bara, J., Bach, E. M. & Siddappaji, M. Culex pipiens and Culex restuans mosquitoes harbor distinct microbiota dominated by few bacterial taxa. Parasit Vectors 9, 18 (2016).

    Article  Google Scholar 

  35. FastQC: a Quality Control Tool for High Throughput Sequence Data (Brabaham Bioinformatics, 2010);

  36. Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014).

    Article  CAS  Google Scholar 

  37. Zhang, J., Kobert, K., Flouri, T. & Stamatakis, A. PEAR: a fast and accurate Illumina Paired-End reAd mergeR. Bioinformatics 30, 614–620 (2014).

    Article  CAS  Google Scholar 

  38. Rognes, T., Flouri, T., Nichols, B., Quince, C. & Mahé, F.VSEARCH: a versatile open source tool for metagenomics. PeerJ 4, e2584 (2016).

    Article  Google Scholar 

  39. National Center for Biotechnology Information. Basic Local Alignment Search Tool (BLAST, accessed July 2016);

  40. R Development Core Team. R: A Language and Environment for Statistical Computing (2017);

  41. Bates, D., Mächler, M., Bolker, B. & Walker, S. C. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).

    Article  Google Scholar 

  42. Fox, J . & Weisberg, S. An R Companion to Applied Regression (SAGE Publications, Thousand Oaks, 2011).

  43. Hothorn, T., Bretz, F. & Westfall, P. Simultaneous inference in general parametric models. Biom. J. 50, 346–363 (2008).

    Article  Google Scholar 

  44. Wickham, H. Reshaping data with the reshape package. J. Stat. Softw. 21, 1–20 (2007).

    Article  Google Scholar 

  45. Oksanen, J. Vegan: An Introduction to Ordination (2017);

  46. Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer, Dordrecht, 2009).

  47. Wilke, C. O. cowplot: Streamlined Plot Theme and Plot Annotations for ‘ggplot2’ (2017);

  48. Stubben, C. & Milligan, B. Estimating and analyzing demographic models using the popbio package in R. J. Stat. Softw. 22, 1–23 (2007).

    Article  Google Scholar 

  49. Berggoetz, M. et al. Tick-borne pathogens in the blood of wild and domestic ungulates in South Africa: interplay of game and livestock. Ticks Tick Borne Dis. 5, 166–175 (2014).

    Article  CAS  Google Scholar 

  50. Jado, I. et al. Molecular method for identification of Rickettsia species in clinical and environmental samples. J. Clin. Microbiol. 44, 4572–4576 (2006).

    Article  CAS  Google Scholar 

Download references


We thank H. Kiai, K. Age, A. Mariki, J. Keesing Ostfeld and numerous enumerators for assistance with data collection, and V. Obiero for assistance with data entry. We extend our great appreciation to the owners, managers, staff members and residents of the properties we surveyed for allowing us to work on their land, and for welcoming and assisting us. We thank the management and staff of Ol Pejeta Conservancy, Lewa Wildlife Conservancy, Mpala Research Centre and Laikipia Wildlife Foundation for their hospitality and for their facilitation of this work. This research was supported by the United States National Science Foundation (Coupled Natural and Human Systems award 1313822).

Author information

Authors and Affiliations



B.F.A., F.K., R.S.O., R.C-K. and H.T. designed the study. S.O., B.F.A. and F.K. collected the ecological data. H.T. and S.A.W. developed the survey questions for property owners and managers and S.H. conducted the interviews. T.H. assisted in the field and identified ticks. L.P.F. conducted the pathogen identification using a protocol developed by B.F.A. and L.P.F. R.C-K., V.K., C.M.W., B.R.B. and S.A.W. assisted with data preparation and interpretation. F.K. analysed the data and F.K., B.F.A. and R.S.O. wrote the manuscript, which was edited and approved by all authors.

Corresponding author

Correspondence to Felicia Keesing.

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.

Supplementary information

Supplementary Information

Supplementary Figures 1–6, Supplementary Tables 1–7

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

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

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


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