Skip to main content
SpringerLink
Log in

The Enemy of My Enemy Hypothesis: Why Coexisting with Grasses May Be an Adaptive Strategy for Savanna Trees

  • Published:
Ecosystems Aims and scope Submit manuscript

Abstract

Savannas are characterized by the coexistence of trees and flammable grasses. Yet, tree–grass coexistence has been labeled as paradoxical—how do these two functional groups coexist over such an extensive area, despite being generally predisposed to excluding each other? For instance, many trees develop dense canopies that limit grass growth, and many grasses facilitate frequent/intense fires, increasing tree mortality. This study revisits tree–grass coexistence with a model of hierarchical competition between pyrogenic grasses, “forest trees” adapted to closed-canopy competition, and “savanna trees” that are inferior competitors in closed-canopy communities, but more resistant to fire. The assumptions of this model are supported by empirical observations, including a systematic review of savanna and forest tree community composition reported here. In general, the model simulations show that when savanna trees exert weaker competitive effects on grasses, a self-reinforcing grass community is maintained, which limits forest tree expansion while still allowing savanna trees to persist (albeit as a subdominant to grasses). When savanna trees exert strong competitive effects on grasses, savanna trees cover increases initially, but as grasses decline their inhibitory effect on forest trees weakens, allowing forest trees to expand and exclude grasses and savanna trees. Rather than paradoxical, these results suggest that having weaker competitive effects on grasses may be advantageous for savanna trees, leading to greater long-term abundance and stability. We label this the “enemy of my enemy hypothesis,” which might apply to species coexistence in communities defined by hierarchical competition or with species capable of generating strong ecological feedbacks.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Figure 1
Figure 2
Figure 3
Figure 4
Figure 5
Figure 6

Similar content being viewed by others

References

  • Adejuwon JO, Adesina FA. 1992. The nature and dynamics of the forest-savanna boundary in south-western Nigeria. Furley, PA, Proctor J, Ratter JA, editors. Nature and dynamics of forest–savanna boundaries. London: Chapman and Hall London. 21–33.

  • Attua EM, Pabi O. 2013. Tree species composition, richness and diversity in the northern forest-savanna ecotone of Ghana. Journal of Applied Bioscience 69:5437–48.

    Article  Google Scholar 

  • Azihou AF, Kakai RG, Bellefontaine R, Sinsin B. 2013. Distribution of tree species along a gallery forest-savanna gradient: patterns, overlaps and ecological thresholds. Journal of Tropical Ecology 29:25–37.

    Article  Google Scholar 

  • Banfai DS, Bowman DMJS. 2005. Dynamics of a savanna-forest mosaic in the Australian tropics inferred from stand structure and historical aerial photography. Australian Journal of Botany 53:185–94.

    Article  Google Scholar 

  • Banfai DS, Bowman DMJS. 2006. Forty years of lowland monsoon rainforest expansion in Kakadu National Park, Northern Australia. Biological Conservation 131:553–65.

    Article  Google Scholar 

  • Banfai DS, Bowman DMJS. 2007. Drivers of rain-forest boundary dynamics in Kakadu National Park, northern Australia: a field assessment. Journal of Tropical Ecology 23:73–86.

    Article  Google Scholar 

  • Beckage B, Platt WJ, Gross LJ. 2009. Vegetation, fire, and feedbacks: a disturbance-mediated model of savannas. The American Naturalist 174:805–18.

    PubMed  Google Scholar 

  • Bonan J. 2008. Ecological Climatology. Cambridge UK: Cambridge University Press.

    Book  Google Scholar 

  • Bond WJ. 2008. What limits trees in C4 grasslands and savannas. Annual Review of Ecology, Evolution, and Systematics 39:641–59.

    Article  Google Scholar 

  • Bond WJ, Keeley JE. 2005. Fire as a global ‘herbivore’: the ecology and evolution of flammable ecosystems. Trend in Ecology and Evolution 20:387–94.

    Article  Google Scholar 

  • Bouchenak-Khelladi J, Maurin O, Hurter J, van der Bank M. 2010. The evolutionary history and biogeography of Mimosoidea (Leguminosae): an emphasis on African acacias. Molecular Phylogenetics and Evolution 57:495–508.

    Article  PubMed  Google Scholar 

  • Bowman DMJS. 1992. Monsoon forests in North-western Australia. II. Forest-Savanna Transitions. Australian Journal of Botany 40:89–102.

    Article  Google Scholar 

  • Bowman DMJS, Wilson BA, Fensham RJ. 1990. Sandstone vegetation pattern in the Jim Jim Falls region, Northern Territory, Australia. Australian Journal of Ecology 15:163–74.

    Article  Google Scholar 

  • Bowman DMJS, Fensham RJ. 1991. Response of a monsoon forest-savanna boundary to fire protection, Weipa, northern Australia. Australian Journal of Ecology 16:111–18.

    Article  Google Scholar 

  • Bowman DMJS, Panton WJ. 1993. Factors that control monsoon-rainforest seedling establishment and growth north Australian Eucalyptus savanna. Journal of Ecology 81:297–304.

    Article  Google Scholar 

  • Brener AG, Silva SF. 1995. Leaf-cutting ants and forest groves in a tropical parkland savanna of Venezuela: facilitated succession? Journal of Tropical Ecology 11:651–69.

    Article  Google Scholar 

  • Charles-Dominique T, Staver AC, Midgley GF, Bond WJ. 2015. Functional differentiation of biomes in an African savanna/forest mosaic. South African Journal of Botany 101:82–90.

    Article  Google Scholar 

  • Cuni-Sanchez A, and others. 2016. African savanna-forest boundary dynamics: a 20-year study. PLoS One 11: e0156934.

  • Chesson P. 2000. Mechanisms of maintenance of species diversity. Annual Review of Ecology and Systematics 31:343–66.

    Article  Google Scholar 

  • D’Odorico P, Laio F, Ridolfi L. 2006. A Probabilistic Analysis of Fire-Induced Tree-Grass Coexistence in Savannas. The American Naturalist 3:E80.

    Google Scholar 

  • Dantas V, Batalha MA, Pausas JG. 2013. Fire drives functional thresholds on the savanna-forest transition. Ecology 94:2454–63.

    Article  Google Scholar 

  • Dezzeo N, Chacon N, Sanoja E, Picon G. 2004. Changes in soil properties and vegetation characteristics along a forest-savanna gradient in southern Venezuela. Forest Ecology and Management 200:183–93.

    Article  Google Scholar 

  • Dohn J, and others. 2013. Tree effects on grass growth in savannas: competition, facilitation and the stress gradient hypothesis. Journal of Ecology 101:202–209.

  • Edwards EJ, Osborne CP, Stromberg CAE, Smith SA, C4 grasses consortium. 2010. The origins of C4 grasslands: integrating evolutionary and ecosystem science. Science 328:587–91.

    Article  CAS  PubMed  Google Scholar 

  • Felfi JM, da Silva MC. 1992. Floristic composition, phytosociology and comparison of cerrado and gallery forests at Fazenda Água Limpa, Federal District, Brazil. Furley PA, Proctor J, Ratter JA, editors. Nature and dynamics of forest-savanna boundaries. London: Chapman and Hall.

  • Fernandez-Illescas CP, Rodriguez-Iturbe I. 2003. Hydrologically driven hierarchical competition-colonization models: The impact of interannual rainfall fluctuations. Ecological Monographs 73:207–22.

    Article  Google Scholar 

  • Frederiksen P, Lawesson JE. 1992. Vegetation Types and Patterns in Senegal Based on Multivariate Analysis of Field and NOAA-AVHRR Satellite Data. Journal of Vegetation Science 3:535–44.

    Article  Google Scholar 

  • Geiger EL, Gotsch SG, Damasco G, Haridasan M, Franco AC, Hoffmann WA. 2011. Distinct roles of savanna and forest tree species in regeneration under fire suppression in a Brazilian savanna. Journal of Vegetation Science 22:312–21.

    Article  Google Scholar 

  • Gignoux J, Lahoreau G, Julliard R, Barot S. 2009. Establishment and early persistence of tree seedlings in an annually burned savanna. Journal of Ecology 97:484–95.

    Article  Google Scholar 

  • Grime JP. 2006. Plant strategies, vegetation processes, and ecosystem properties. New York: Wiley and Sons.

    Google Scholar 

  • Hanan NP, Tredennick AT, Prihodko L, Bucini G, Dohn J. 2014. Analysis of stable states in global savannas: is the CART pulling the horse. Global Ecology and Biogeography 23:259–63.

    Article  PubMed  Google Scholar 

  • Hennenberg KJ, Goetze D, Kouame L, Orthmann B, Porembski S. 2005. Border and ecotone detection by vegetation composition along forest-savanna transects in Ivory Coast. Journal of Vegetation Science 16:301–10.

    Article  Google Scholar 

  • Higgins SI, Bond WJ, Trollope WSW. 2000. Fire, resprouting and variability: a recipe for grass-tree coexistence in savanna. Journal of Ecology 88:213–29.

    Article  Google Scholar 

  • Hirota M, and others. 2010. The climatic sensitivity of the forest, savanna and forest– savanna transition in tropical South America. New Phytologist 187:707–719.

  • Hirota M, Holmgren M, Van Nes E, Scheffer M. 2011. Global resilience of tropical forest and savanna to critical transitions. Science 334:232–5.

    Article  CAS  PubMed  Google Scholar 

  • Hoffmann WA. 1996. The effects of fire and cover on seedling establishment in neotropical savanna. The Journal of Ecology 84:383–93.

    Article  Google Scholar 

  • Hoffmann WA. 1999. Fire and population dynamics of woody plants in neotropical savanna: matrix model projections. Ecology 80:1354–69.

    Article  Google Scholar 

  • Hoffmann WA, Adasme R, Haridasan M, De Carvalho MT, Geiger EL, Pereira MAB, Gotsch SG, Franco AC. 2009. Tree topkill, not mortality, governs the dynamics of savanna-forest boundaries under frequent fire in central Brazil. Ecology 90:1326–37.

    Article  PubMed  Google Scholar 

  • Hoffmann WA, Geiger EK, Gotsch SG, Rossatto DR, Silva LCR, Lee Lau O, Haridasan M, Franco AC. 2012. Ecological thresholds at the savanna-forest boundary: how plant traits, resources and fire govern the distribution of tropical biomes. Ecology Letters 15:759–68.

    Article  PubMed  Google Scholar 

  • Holdo RM. 2013. Revisiting the Two-Layer Hypothesis: Coexistence of Alternative Functional Rooting Strategies in Savannas. PLoS One 8:e69625.

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Holdo RM, Holt RD, Fryxell JM. 2013. Herbivore-vegetation feedbacks can expand the range of savanna persistence: insights from a simple theoretical model. Oikos 122:441–53.

    Article  Google Scholar 

  • Hovestadt T, Yao P, Linsenmair E. 1999. Seed dispersal mechanisms and the vegetation of forest islands in a West African forest-savanna mosaic (Comoe National Park, Ivory Coast). Plant Ecology 144:1–25.

    Article  Google Scholar 

  • Ibanez T, Curt T, Hely C. 2013a. Low tolerance of New Caledonian secondary forest species to savanna fires. Journal of Vegetation Science 24:177–88.

    Article  Google Scholar 

  • Ibanez T, Hely C, Gaucherel C. 2013b. Sharp transitions in microclimate conditions between savanna and forest in New Caledonia: insights into the vulnerability of forest edges to fire. Austral Ecology 38:680–7.

    Article  Google Scholar 

  • Jeltsch F, Weber GE, Grimm V. 2000. Ecological buffering mechanisms in savannas: A unifying theory of long-term tree-grass coexistence. Plant Ecology 150:161–71.

    Article  Google Scholar 

  • Jensen M, Friis I. 2001. Fire regime, floristics, diversity, life forms and biomass in wooded grassland, woodland and dry forest at Gambella, Western Ethiopia. Biologiske Skrifter/Det Kongelige Danske Videnskabernes Selskab 54:349–87.

    Google Scholar 

  • Kerr B, Riely MA, Fledman MW, Bohannan JM. 2002. Local dispersal promotes biodiversity in a real-life game of rock-paper-scissors. Nature 418:171–4.

    Article  CAS  PubMed  Google Scholar 

  • Kikula IS. 1986. The influence of fire on the composition of Miombo woodland of SW Tanzania. Oikos 46:317–24.

    Article  Google Scholar 

  • Kulmatiski A, Beard KH, Verweij RJT, February EC. 2010. A depth-controlled tracer technique measures vertical, horizontal and temporal patterns of water use by trees and grasses in a subtropical savanna. New Phytologist 188:199–209.

    Article  PubMed  Google Scholar 

  • Lawes ML, Richards A, Dathe J, Midgley JJ. 2011. Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia. Plant Ecology 212:2057–69.

    Article  Google Scholar 

  • Lawrence D, D’Odorico P, Diekmann L, DeLonge M, Das R, Eaton J. 2008. Ecological feedbacks following deforestation create the potential for a catastrophic ecosystem shift in tropical dry forest. PNAS 104:20696–701.

    Article  Google Scholar 

  • Lawton RM. 1978. A study of the dynamic ecology of Zambian vegetation. Journal of Ecology 66:175–98.

    Article  Google Scholar 

  • Lehmann CER, and others 2014. Savanna vegetation-fire-climate relationships differ among continents. Science 343: 548–552.

  • Loschi RA, and others. 2013. Structural and environmental variations in the continuum of gallery forest/savanna stricto sensu in Itumirim, MG. Cerne 19: 213-227.

  • Macdermott HJ, Fensham RJ, Hua Q, Bowman DMJS. In press. Vegetation, fire and soil feedbacks of dynamic boundaries between rainforest, savanna and grassland. Austral Ecology.

  • Mapure I. 1997. A floristic classification of the vegetation of a forest-savanna boundary in southeastern Zimbabwae. Bothalia 27:185–93.

    Google Scholar 

  • Mapure I. 2012. Determinants of vegetation composition and diversity of a moist forest-savanna boundary in south-eastern Zimbabwae. International Journal of Biodiversity and Conservation 4:584–91.

    Google Scholar 

  • Marimon BS, Lima ES, Duarte TG, Chieregatto LC, Ratter JA. 2006. Observations on the vegetation of northeastern mato grosso, Brazil. IV. An analysis of the cerrado-amazonian forest ecotone. Edinburgh Journal of Botany 63:323–41.

    Article  Google Scholar 

  • Mitchard ETA, Saatchi SS, Gerard FF, Lewis SL, Meir P. 2009. Measuring woody encroachment along a forest-savanna boundary in central Africa. Earth Interactions 13:1–28.

    Article  Google Scholar 

  • Müller CS, Overbeck GE, Pfadenhauer J, Pillar VD. 2012a. Woody species patterns at forest-grassland boundaries in southern Brazil. Flora 207:586–98.

    Article  Google Scholar 

  • Müller JV, Sieglstetter R, Csontos P. 2012b. A multivariate approach to identify vegetation belts: gallery forest and its surrounding savanna along the river Kota in north Benin. Plant Biosystems 146:878–88.

    Article  Google Scholar 

  • Murphy BP, Russel-Smith J, Prior LD. 2010. Frequent fires reduce tree growth in northern Australian savannas: implications for tree demography and carbon sequestration. Global Change Biology 16:331–43.

    Article  Google Scholar 

  • Murphy BP, Bowman DMJS. 2012. What controls the distribution of tropical forest and savanna? Ecology Letters 15:748–58.

    Article  PubMed  Google Scholar 

  • Nangendo HT, Steege G, Bongers F. 2006. Composition of woody species in a dynamic forest-woodland-savannah mosaic in Uganda: implications for conservation and management. Biodiversity and Conservation 15:1467–95.

    Article  Google Scholar 

  • Pennec F, and others. 2016. Floristic and structural vegetation typology of bonobo habitats in a forest-savanna mosaic (Bolobo Territory, D.R.Congo). Plant Ecology and Evolution 149: 199-215.

  • Pérez-García EA, Meave JA, Villasenor JL, Gallardo-Cruz JA, Lebrija-Trejos EE. 2010. Vegetation heterogeneity and life-strategy diversity in the flora of the heterogeneous landscape of Nizanda, Oxacaca, Mexico. Folia Geotanica 45:143–61.

    Article  Google Scholar 

  • R Core Team (2015). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL http://www.R-project.org/.

  • Ratajczak Z, Nippert JB, Briggs JM, Blair JM. 2014. Fire dynamics distinguish grasslands, shrublands and woodlands as alternative attractors in the Central Great Plains of North America. Journal of Ecology 102:1374–85.

    Article  Google Scholar 

  • Ratnam J, Bond WJ, Fensham RJ, Hoffmann WA, Archibald S, Lehmann CER, Anderson MT, Higgins SI, Sankaran M. 2011. When is a ‘forest’ a savanna, and why does it matter? Global Ecology and Biogeography 20:653–60.

    Article  Google Scholar 

  • Ridolfi L, D’Odorico P, Laio F. 2011. Noise-induced Phenomena in the Environmental Sciences. New York: Cambridge University Press. p 314.

    Book  Google Scholar 

  • Riginos C, Grace JB. 2008. Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-town effects. Ecology 89:2228–38.

    Article  PubMed  Google Scholar 

  • Rodriques-Souza J, and others. 2015. Secondary forest expansion over a savanna domain at an ecological reserve in the Southeastern Brazil after 15 years of monitoring. Brazilian Journal of Botany 38: 311-322.

  • Rosell JA, Gleason S, Mendez-Alonzo R, Chang Y, Westoby M. 2014. Bark functional ecology: evidence for tradeoffs, functional coordination, and environment producing bark diversity. New Phytologist 201:486–97.

    Article  PubMed  Google Scholar 

  • Ruggiero PGC, Batalha MA, Pivello VR, Meirelles ST. 2002. Soil-vegetation relationships in cerrado (Brazilian savanna) and semideciduous forest, southeastern Brazil. Plant Ecology 160:1–16.

    Article  Google Scholar 

  • Sankaran M, Ratnam J, Hanan NP. 2004. Tree–grass coexistence in savannas revisited – insights from an examination of assumptions and mechanisms invoked in existing models. Ecology Letters 7:480–90.

    Article  Google Scholar 

  • Sankaran M, and others 2005. Determinants of woody cover in African savannas. Nature 438: 846-849.

  • Sarmiento G. 1984. The ecology of neotropical Savannas. Cambridge Massachusetts: Harvard University Press.

    Book  Google Scholar 

  • Schieter S, Higgins SI, Osborne CP, Bradshaw C, Lunt D, Ripley BS, Taylor LL, Beerling DJ. 2012. Fire and fire adapted vegetation promoted C4 expansion in the late Miocene. New Phytologist 195:653–66.

    Article  Google Scholar 

  • Scholes RJ, Archer SR. 1997. Tree-grass interactions in savannas. Annual Review of Ecology and Systematics 28:517–44.

    Article  Google Scholar 

  • Schwartz D, de Foresta H, Mariotti A, Balesdent J, Massimba JP, Girardin C. 1996. Present dynamics of the savanna-forest boundary in the Congolese Mayombe: a pedological, botanical and isotopic (13C and 14C) study. Oecologia 106:516–24.

    Article  CAS  PubMed  Google Scholar 

  • Silva JF, Zambrano A, Farinas MR. 2002. Increase in the woody component of seasonal savannas under different fire regime in Calbozo, Venezuela. Journal of Biogeography 28:977–83.

    Article  Google Scholar 

  • Simon MF, Pennington T. 2012. Evidence for adaptation to fire regimes in the tropical savannas of the Brazilian Cerrado. International Journal of Plant Sciences 173:711–23.

    Article  Google Scholar 

  • Smith PP, Fisher R. 2001. Chipya in Kasanka National Park, Zambia: Floristics, Soils and Dynamics. Systematics and Geography of Plants 71:923–34.

    Article  Google Scholar 

  • Staal A, Dekker SC, Hirota M, van Nes EH. 2015. Synergistic effects of drought and deforestation on the resilience of the south-eastern Amazon rainforest. Ecological Complexity 22:65–75.

    Article  Google Scholar 

  • Staver AC, Archibald S, Levin SA. 2011. The Global Extent and Determinants of Savanna and Forest as Alternative Biome States. Science 334:230–2.

    Article  CAS  PubMed  Google Scholar 

  • Staver AC, Levin SA. 2012. Integrating theoretical climate and fire effects on savanna and forest systems. The American Naturalist 180:211–24.

    Article  PubMed  Google Scholar 

  • Stromberg CAE. 2011. Evolution of grasses and grassland ecosystems. Annual Review of Earth and Planetary Sciences 39:517–44.

    Article  CAS  Google Scholar 

  • Tilman D, Isbell F, Cowles JM. 2014. Biodiversity and ecosystem functioning. Annual Review of Ecology, Evolution, and Systematics 45:471–93.

    Article  Google Scholar 

  • Trauernicht C, Murphy BP, Portner TA, Bowman DMJS. 2012. Tree cover–fire interactions promote the persistence of a fire-sensitive conifer in a highly flammable savanna. Journal of Ecology 100:958–68.

    Article  Google Scholar 

  • Trauernicht C, Murphy BP, Prior LD, Lawes MJ, Bowman DMJS. 2016. Human-Imposed, Fine-Grained Patch Burning Explains the Population Stability of a Fire- Sensitive Conifer in a Frequently Burnt Northern Australia Savanna. Ecosystems 19:869–909.

    Article  Google Scholar 

  • Turton SM, Duff GA. 1992. Light environments and floristic composition across an open forest-rainforest boundary in northeastern Queensland. Australian Journal of Ecology 17:415–23.

    Article  Google Scholar 

  • Unwin SL. 1989. Structure and composition of the abrupt rainforest boundary in the Herberton Highland, North Queensland. Australian Journal of Botany 37:413–28.

    Article  Google Scholar 

  • Van Langevelde F, and others 2003. Effects of fire and herbivory on the stability of savanna ecosystems. Ecology 84: 337-350.

  • Walpole MJ, Nabaala M, Matankory C. 2004. Status of the Mara Woodlands in Kenya. African Journal of Ecology 42:180–8.

    Article  Google Scholar 

  • Walter H. 1971. Ecology of tropical and subtropical vegetation. Edinburgh: Oliver and Boyd.

    Google Scholar 

  • Weltzin JF, McPherson GR. 1997. Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA. Oecologia 112:156–64.

    Article  PubMed  Google Scholar 

  • White L, Oslisly R, Abernethy K, Maley J. 1996. O’Kume (Aucoumea klaineana): expansion et declin d’un arbre pionnier en Afrique central atlantique au cours de l’Holocene. Servant M, Servant-Vildary S, editors. Dynamique a long terme des ecosystems forestiers intertropicaux. Paris: UNESCO.

  • Wiedemeier DB, Bloesch U, Hagedorn F. 2012. Stable forest-savanna mosaic in north-western Tanzania: local-scale evidence from delta 13C signatures and 14C ages of soil fractions. Journal of Biogeography 39:247–57.

    Article  Google Scholar 

  • Wolf J. 1998. Species composition and structure of the woody vegetation of the Middle Casamance region (Senegal). Forest Ecology and Management 111:249–64.

    Article  Google Scholar 

  • Xu C, Hanston S, Holmgren M, van Nes EH, Staal A, Scheffer M. 2016. Remotely sensed canopy height reveal three pantropical ecosystem states. Ecology 97:2518–21.

    Article  PubMed  Google Scholar 

  • Yu KL, D’Odorico P. 2015. Hydraulic lift as a determinant of tree-grass coexistence on savannas. New Phytologist 207:1038–51.

    Article  PubMed  Google Scholar 

Download references

Acknowledgements

ZR was supported by a National Science Foundation post-doctoral fellowship (DBI 1402033). We thank two anonymous reviewers for comments that greatly improved the literature review.

Author information

Authors and Affiliations

  1. Department of Environmental Sciences, University of Virginia, 291 McCormick Road, P.O. Box 400123, Charlottesville, Virginia, 22904, USA

    Zak Ratajczak, Paolo D’Odorico & Kailiang Yu

  2. Department of Environmental Science Policy and Management, University of California, Berkeley, Berkeley, California, 94720, USA

    Paolo D’Odorico

Authors
  1. Zak Ratajczak
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Paolo D’Odorico
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Kailiang Yu
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Zak Ratajczak.

Additional information

Author contributions

ZR, PD, and KY contributed to study design. ZR performed simulations and wrote the first draft. ZR, PD, and KY contributed to revisions.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (DOCX 2229 kb)

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Ratajczak, Z., D’Odorico, P. & Yu, K. The Enemy of My Enemy Hypothesis: Why Coexisting with Grasses May Be an Adaptive Strategy for Savanna Trees. Ecosystems 20, 1278–1295 (2017). https://doi.org/10.1007/s10021-017-0110-7

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10021-017-0110-7

Keywords

Search

Navigation