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Emeritus Professor William Bond

Phone: 021 650 2439 
Room No.: 3.19 (HW Pearson Building)






Key Research Interests

Processes most strongly influencing vegetation change in the past and present, including fire, vertebrate herbivory, climate extremes, atmospheric [CO2] and habitat fragmentation.

Plant-animal interactions
Plant form and function.
Systems studied include sub-tropical grasslands, savannas and winter rainfall shrublands.
A major research project (funded by the Mellon Foundation and the NRF) on biomes and their boundaries started in 2010.

UCT representative for MacArthur Foundation Project in Madagascar with Professor Joel Ratsirarson of University of Antananarivo and Professor John Silander of University of Connecticut and colleagues (2012-2014). 


BSc Hons (1970) University of Exeter ; MSc (1980) University of Cape Town ; PhD (1987) University of California , Los Angeles . Natural resource survey, Mozambique (1971-1974) Lake ecology, Cabora Bassa, Mozambique (1974-1976); Ecological research, Cape mountain catchments, Forestry Department (1976-1982; 1986-1987); Graduate studies, UCLA California Chaparral (1982-1986); Lecturer Botany Department, University of Cape Town (1988-1993), Professor, Botany Department, UCT (1993 - 2013), Emeritus Professor, Department of Biological Sciences UCT (2014- );

Chief Scientist, South African Environmental Observation Network (SAEON) (2014- ).

Professional Activities (1995-present)

Board member (Chairman of research committee), National Botanical Institute, 1995-1999

National Research Foundation Plant Scientists Rating Committee 1998-2000
Board Member, Cape Nature Conservation. 1999- 2006
Member of Editorial Board, Oecologia, 1994- 2001
Member of Editorial Board Global Ecology and Biogeography 2001 - present
Member of Editorial Board, Global Change Biology 2008-2010
Member of Editorial Board, Koedoe 2008-

Elected Foreign Associate of the National Academy of the USA (2013)

Post-Doctoral fellows

Tristan Charles-Dominique. Understanding the ecology of trees and shrubs in savanna: a developmental perspective. Plant architectural analysis across alternative biome states.

Gareth Hempson. Mammal assemblages in Africa and their functional organization. 

Current research students


Heath Beckett. The dynamics of forest-savanna boundaries.

Abraham Dabengwa (co-supervised with Lindsey Gillson) Reconstructing palaeovegetation sequences at grassland-savanna-woodland biome boundaries in KwaZulu-Natal and related landscape ecological processes in the late Holocene

Simon Power (co-supervised with Mike Cramer, Tony Verboom) The ecophysiological mechanisms that determine the afrotemperate forest-fynbos boundary in the Cape Floristic Region

Nikki Stevens (co-supervised with Sally Archibald, Barend Erasmus, Wits University). Change in the woody plant community in semi-arid savannas.

Masters by dissertation:

Cameron Wills. Factors influencing fire spread in an African savanna.

Recently graduated Doctoral:

Ben Wigley (co-supervised with Herve Fritz). Vertebrate browsing and plant functional traits.

Leonard Masubelele (co-supervised with Timm Hoffmann). Environmental change in the eastern Karoo Midlands over the last 100 years.

Recently graduated Masters:

Katherine Gordon (nee Bunney). Seed Dispersal in South African Trees: with a focus on the megafaunal fruit and their dispersal agents

Nick Zaloumis. Grassland diversity and its restoration.

Emma Gray. Thicket expansion and its effects on ecosystem goods and services.

Glenn Moncrieff. The demographic impacts of browsing on woody plants in savannas: from individual branches to whole populations.

Publications include:


Scott AC, Bowman DMJS, Bond WJ, Alexander ME. 2014. Fire on Earth: An Introduction. Wiley, Blackwell. 434 pages. ISBN: 978-1-119-95356-2

Keeley JE, Bond WJ, Bradstock RA, Pausas JG, Rundel PW. 2011. Fire Ecology, Evolution and Management in Mediterranean Climate Ecosystems. Cambridge University Press, Cambridge, UK.

Bond, W.J., van Wilgen, B.W. (1996) Fire and Plants. Chapman and Hall, London . (Volume 14 in the Population and Community biology series).

Selected Journal Articles


Staver AC, Bond WJ. 2014. Is there a ‘browse trap’? Dynamics of herbivore impacts on trees and grasses in an African savanna. J. Ecol 102: 595-602

Parr, CL, Lehmann CER, Bond WJ, Hoffmann WA, Andersen AN. 2014. Tropical grassy biomes: misunderstood, neglected, and under threat. Trends in Ecology and Evolution 29: 205-213

Bouchenak-Khelladi Y, Slingsby JA, Verboom GA, Bond WJ. 2014. Diversification of C4 grasses (Poaceae) does not coincide with their ecological dominance. American Journal of Botany 101:300-307.

Masubelele ML, Hoffman MT, Bond WJ, Gambiza J. 2014. A 50 year study shows grass cover has increased in shrublands of semi-arid South Africa. Journal of Arid Environments 104: 43-51

Lehmann, C. E., Anderson, T. M., Sankaran, M., Higgins, S. I., Archibald, S., Hoffmann, W. A., ... & Bond, W. J. (2014). Savanna Vegetation-Fire-Climate Relationships Differ Among Continents. Science, 343(6170), 548-552.

Moncrieff GR, Scheiter S, Bond WJ, Higgins SI. 2014. Increasing atmospheric CO2 overrides the historical legacy of multiple stable biome states in Africa. New Phytologist 201, 908-915.


Beale, C. M., Rensberg, S. V., Bond, W. J., Coughenour, M., Fynn, R., Gaylard, A., Grant R., Harris B., Jones T., Mduma S., Owen-Smith N., & Sinclair A.R.E (2013). Ten lessons for the conservation of African savannah ecosystems. Biological Conservation, 167, 224-232.

Gray, E.F. & Bond, W.J. 2013. Will woody plant encroachment impact the visitor experience and economy of conservation areas? Koedoe 55(1), Art. #1106, 9 pages. v55i1.1106

Wigley BJ, Coetsee C, Hartshorn AS, Bond WJ. 2013. What do ecologists miss by not digging deep enough? Insights and methodological guidelines for assessing soil fertility status in ecological studies. Acta Oecologica, 51, 17-27

February EC, Higgins SI, Bond WJ, Swemmer L. 2013. Influence of competition and rainfall manipulation on the growth responses of savanna trees and grasses. Ecology, 94(5), 1155–1164

Coetsee C, Gray EF, Wakeling J, Wigley BJ, Bond WJ. 2013. Low gains in ecosystem carbon with woody plant encroachment in a South African savanna. Journal of Tropical Ecology 29 (01), 49-60.

Cramer MD, Bond WJ. 2013. N-fertilization does not alleviate grass competition induced reduction of growth of African savanna species. Plant and Soil 366:563-574.


Bond WJ, Midgley JJ. 2012. Fire and the angiosperm revolutions. International J. Plant Sciences 173:569-583

Bond WJ, Cook G, Williams RJ. 2012. Which trees dominate in savannas? The escape hypothesis and eucalypts in northern Australia. Austral Ecology 37: 678-685.

West AG, Dawson TE, February EC, Midgley GF, Bond WJ, Aston TL. 2012. Diverse functional responses to drought in a Mediterranean-type shrubland in South Africa. New Phytologist 195: 396-407

Staver AC, Bond WJ, Cramer MD, Wakeling JL. 2012. Top-down determinants of niche structure and adaptation among African Acacias. Ecology Letters 15: 673-679.

Parr CL, Gray EF, Bond WJ. 2012. Cascading biodiversity and functional consequences of a global change–induced biome switch. Diversity and Distributions 18: 493–503

Wakeling JL, Cramer MD, Bond WJ. 2012. The savanna-grassland ‘treeline’: why don’t savanna trees occur in upland grasslands? J. Ecology 100: 381–391

Buitenwerf, R., Bond, W.J., Stevens, N., Trollope, W.S.W. 2012.  Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver. Global Change Biology 18,  675–684.   

Bond WJ, Midgley GF. 2012. CO2 and the uneasy interactions of trees and savanna grasses. Phil. Trans. R. Soc. B  367, 601-612

Vetter S., Bond W.J. 2012. Changing predictors of spatial and temporal variability in stocking rates in a severely degraded communal rangeland. Land Degradation & Development 23: 190-199.


Moncrieff GR, Chamaillé-Jammes S, Higgins SI, O’Hara RB, Bond WJ. (2011). Tree allometries reflect a lifetime of herbivory in an African savanna. Ecology 92(12) 2310-2315 [doi:10.1890/11-0230.1]

Wakeling JL, Staver AC, Bond WJ. 2011. Simply the best: the transition of savanna saplings to trees. Oikos 120: 1448–1451

Pauw A, Bond WJ. 2011. Mutualisms matter: pollination rate limits the distribution of oil-secreting orchids. Oikos 120: 1531–1538.

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: 653-660

Keeley JE, Pausas JG, Rundel PW, Bond WJ, Bradstock RA. 2011. Fire as an evolutionary pressure shaping plant traits. Trends in Plant Science 16:406-411

Estes JA, Terborgh J, Brashares JS, Power ME, Berger J, Bond WJ, Carpenter SR, Essington TE, Holt RD, Jackson JBC, Marquis RJ, Oksanen L, Oksanen T, Paine RT, Pikitch EK, Ripple WJ, Sandin SA, Scheffer M, Schoener TW, Shurin JB, Sinclair ARE, Soulé ME, Virtanen R, Wardle DA. 2011. Trophic Downgrading of Planet Earth. Science 333: 301-306.

Lehmann CER, Archibald S, Hoffmann W, Bond WJ. 2011. Deciphering the distribution of the savanna biome. New Phytologist 191: 197–209. 

Zaloumis NP, Bond WJ. (2011) Grassland restoration after afforestation: No direction home? Austral Ecology 36, 357–366.

Staver AC, Bond WJ, February EC. 2011. History matters: tree establishment variability and species turnover in an African savanna. Ecosphere 2:art49 pages 1-12. [doi:10.1890/ES11-00029.1]


Bond WJ, Scott AC. 2010. Fire and the spread of flowering plants in the Cretaceous. New Phytologist New Phytologist 188: 1137–1150

(See also Commentary: Belcher CM. 2010. From fiery beginnings: wildfires facilitated the spread of angiosperms in the Cretaceous. New Phytologist 188: 913-915)

Radloff FGT, Mucina L, Bond WJ, LeRoux PJ. 2010. Strontium isotope analyses of large herbivore habitat use in the Cape Fynbos region of South Africa. Oecologia 164: 567-578.

Bond WJ. 2010. Do nutrient-poor soils inhibit development of forests? a nutrient stock analysis. Plant Soil 334:47-60

Bond, WJ. 2010. Consumer control by megafauna and fire. In: J. Terborgh, J.A. Estes (eds), Trophic cascades: predators, prey and the changing dynamics of nature. Island Press, Washington DC. Pp. 275-285.

Bond WJ, Parr CL. 2010. Beyond the forest edge: Ecology, diversity and conservation of the grassy biomes. Biological Conservation 143: 2395-2404

Edwards EJ, Osborne CP, Strömberg CAE, Smith SA, and the C4 Grasses Consortium 2010. The Origins of C4 Grasslands: Integrating Evolutionary and Ecosystem Science. Science 328:587-591.

Coetsee C, Bond WJ, February EC. 2010. Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover. Oecologia 162:1027-1034

Wigley BJ, Bond WJ, Hoffman MT. 2010. Thicket expansion in a South African savanna under divergent land use: local vs. global drivers? Global Change Biology 16, 964-976

Kgope BS, Bond WJ, Midgley GF. 2010. Growth responses of African savanna trees implicate atmospheric [CO2] as a driver of past and current changes in savanna tree cover. Austral Ecology 35: 451-463

Stock WD, Bond WJ, van de Vijver CADM. 2010. Herbivore and nutrient control of lawn and bunch grass distributions in a southern African savanna. Plant Ecology 206:15-27

Cramer MD, van Cauter A, Bond WJ. 2010. Growth of N2-fixing African savanna Acacia spp. is constrained by below-ground competition with grass. Journal of Ecology 2010, 98, 156-167


Staver AC, Bond WJ, Stock WD, Van Rensburg SJ, Waldram MS. 2009. Browsing and fire interact to suppress tree density in an African savanna. Ecological Applications 19: 1909-1919

Bowman DMJS, Balch JK, Artaxo P, Bond WJ, Carlson JM, et al. 2009. Fire in the Earth System. Science 324: 481-484.

Schutz AEN, Bond WJ, Cramer MD. 2009. Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna. Oecologia 160:235-246

Wigley BJ, Cramer MD, Bond WJ. 2009. Sapling survival in a frequently burnt savanna: mobilisation of carbon reserves in Acacia karroo. Plant Ecology 2003:1-11


Bond, W.J. 2008. What limits trees in C4 grasslands and savannas? Annu. Rev. Ecol. Evol. Syst. 39:641–59

Waldram MS, Bond WJ, Stock WD. 2008. Ecological engineering by a mega-grazer: White Rhino impacts on a South African savanna. Ecosystems 11: 101-112.

Bond, W.J., Silander J.A., Ranaivonasy J, Ratsirarson J. 2008. The antiquity of Madagascar ’s grasslands and the rise of C4 grassy biomes. J. Biogeogr. 35: 1743-1758

(see also a commentary on this paper by K. J. Willis, L. Gillson, M. Virah-Sawmy 2008, Nature or nurture: the ambiguity of C4 grasslands in Madagascar. J. Biogeog 35: 1741-1742)

Coetsee C, February EC, Bond WJ. 2008. Nitrogen availability is not affected by frequent fire in a South African savanna. Journal of Tropical Ecology, 24: 647-654.


Bond WJ, Silander JA. 2007. Springs and wire plants: anachronistic defences against Madagascar's extinct elephant birds. Proc Roy Soc B, 274: 1985-1992. (see commentary with artist’s depiction: Dempewolf H., Rieseberg LH. 2007. Adaptive evolution: the legacy of past giants. Current Biology 17 (17): R773-R774.

Higgins SI, Bond WJ, February EC, Bronn A, Euston-Brown DIW, Enslin B, Govender N, Rademan L, O'Regan S, Potgieter ALF, Scheiter S, Sowry R, Trollope L, Trollope WSW. 2007. Effects of four decades of fire manipulation on woody vegetation structure in savanna. Ecology 88: 1119-1125.

Krook, K., Bond, W., & Hockey, P. (2007). The effect of grassland shifts on the avifauna of a South African savanna. Ostrich: Journal of African Ornithology, 78: 271-279

Hendricks HH, Bond WJ, Midgley JJ, Novellie PA. 2007. Biodiversity conservation and pastoralism-reducing herd size in a communal livestock production system in Richtersveld National Park. J. Arid Env. 70: 718-727


Bond, W.J., Keeley, J.E. 2005. Fire as global ‘herbivore’: the ecology and evolution of flammable ecosystems. Trends in Ecology and Evolution 20: 387-394

Bond, W.J. 2005. Large parts of the world are brown or black: a different view on the ‘Green World’ hypothesis. J. Veg. Sci. 16: 261-266 (Invited perspectives paper)

Archibald S, Bond W.J, Stock WD, Fairbanks DHK (2005) Shaping the landscape: Fire-grazer interactions in an African savanna. Ecological Applications 15: 96-109.

Bond, WJ, Woodward FI, Midgley GF. 2005. The global distribution of ecosystems in a world without fire. New Phytologist 165:525-538

(See commentary on this paper in same issue: Bowman D. 2005. Understanding a flammable planet – climate, fire and global vegetation patterns. New Phyt 165:341-345)


Hemborg, Å.M., Bond, W.J. 2005. Different rewards in female and male flowers can explain the evolution of sexual dimorphism in plants. Biological Journal of the Linnean Society 85: 97-109

Balmford, A., Bond, W.J. 2005. Trends in the state of nature and their implications for human well-being. Ecology Letters 8: 1218-1234.


Archibald S, Bond WJ (2004) Grazer movements: spatial and temporal responses to burning in a tall-grass savanna system. Intl Jnl Wildland Fire 13: 377-385.

Bond, W.J., Dickinson K.J.M, Mark, A.F. (2004). What limits the spread of fire-dependent vegetation? Evidence from geographic variation of serotiny in a New Zealand shrub. Global Ecology and Biogeography 13: 115-127.

Bond WJ, Lee WG, Craine JM. (2004). Plant structural defences against avian browsers: the legacy of New Zealand ’s extinct moas. Oikos 104: 500-508.

Uys, R., Bond, W.J., Everson, T.M. (2004) The effects of different fire regimes on plant diversity in southern African grasslands. Biological Conservation 118: 489-499.


Craine, J., W. Bond, W. Lee, P. Reich, and S. Ollinger (2003) The resource economics of chemical and structural defenses across nitrogen supply gradients. Oecologia 442:547-556.

Bond, W.J., Archibald, S. (2003) Confronting complexity: fire policy choices in South African savanna parks. International Journal of Wildland Fire 12: 1-9. pdf

Bond, W.J., Midgley, J.J. (2003) The evolutionary ecology of sprouting. International Journal of Plant Sciences 164: S103–S114 pdf

Archibald, S. Bond, W.J. (2003) Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments. Oikos 102: 3-14. pdf

Bond, W.J., Midgley, G.F., Woodward, F.I. (2003) The importance of low atmospheric CO 2 and fire in promoting the spread of grasslands and savannas. Global Change Biology 9:973-982 pdf

Bond, W.J., Midgley, G.F., Woodward, F.I. (2003) What controls South African vegetation - climate or fire? South African Journal of Botany 69:79-91. pdf

Pammenter, N.W., Midgley, G.F., Bond, W.J. 2003. Xylem hydraulics and angiosperm success: a test using broad-leafed species. In: A.R. Hemsley, I Poole (eds.), The Evolution of Plant Physiology, Academic Press, London . pp. 263-276.


Midgley, J.M., Bond, W.J. (2001). A synthesis of the demography of African acacias. Journal of Tropical Ecology 17:871-886

Bond, W.J., Loffell, D. (2001) Introduction of giraffe changes acacia distribution in a South African savanna. African Journal of Ecology 39 (3), 286-294. pdf

Bond, W.J., Midgley, JM. (2001) The persistence niche: ecology of sprouting in woody plants. Trends in Ecology and Evolution 16:45-51

Bond, W.J., Ladd, P.G. 2001. Dynamics of the overstorey and species richness in Australian heathlands. Journal of Mediterranean Ecology 2: 247-257

Bond WJ, Smythe, KA, Balfour DA (2001) Acacia species turnover in space and time in an African savanna. J. Biogeography 28: 117-128

Higgins, SI, Pickett, STA, Bond WJ (2000). Predicting extinction risks for plants: environmental stochasticity can save declining populations. Trends in Ecology and Evolution 15:516-520


West AG, Bond WJ, Midgley JJ. 2000. Soil carbon isotopes reveal ancient grassland under forest in Hluhluwe, Kwa-Zulu Natal. Sth Afr Jnl Sci 96:252-254.

Bond, WJ, Midgley, GF (2000) A proposed CO2-controlled mechanism of woody plant invasion in grasslands and savannas. Global Change Biology 6:865-869

Higgins, SI, Bond WJ, Trollope WSW (2000) Fire, resprouting and variability: a recipe for tree-grass coexistence in savanna. J. Ecol. 88, 213-229.

Gambiza, J., Bond, W., Frost, P.G.H., Higgins, S. (2000) A simulation model of miombo woodland dynamics under different management regimes. Ecological Economics 33: 353-368

Earlier (on request)

Other References

RESPONSES OF SAVANNAS TO STRESS AND DISTURBANCE. Eds: P. Frost, E. Medina, J.-C. Menaut, O. Solbrig, M. Swift and B. Walker. Biology International SPECIAL ISSUE lO[1986] PDF (1482 Kb)