Register      Login
Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE

Juvenile tree growth and demography in response to feral water buffalo in savannas of northern Australia: an experimental field study in Kakadu National Park

P. A. Werner A B C F , I. D. Cowie D and J. S. Cusack E
+ Author Affiliations
- Author Affiliations

A Centre for Resource and Environmental Studies, Australian National University, Canberra, ACT 0200, Australia.

B School for Environment Research, Charles Darwin University, Darwin, NT 0909, Australia.

C W. K. Kellogg Biological Station, Michigan State University, Hickory Corners, MI 49060, USA.

D Northern Territory Herbarium, NT Department of Infrastructure, Planning and Environment, PO Box 496, Palmerston, NT 0831, Australia.

E CSIRO Tropical Ecosystems Research Centre, PMB 44, Winnellie, NT 0821, Australia.

F Corresponding author. Email: wernerpa@ufl.edu

Australian Journal of Botany 54(3) 283-296 https://doi.org/10.1071/BT04135
Submitted: 7 September 2004  Accepted: 13 October 2005   Published: 15 May 2006

Abstract

Tree populations in the wooded savannas of northern Australia lack a well developed seed bank, but instead rely on a persistent ground layer of suppressed juvenile plants <1 m of indeterminate ages. The feral Asian water buffalo has been implicated as a factor in inhibiting the movement of juvenile trees to sapling stage. In a 6-year field study in Kakadu National Park, 656 juvenile plants were monitored for growth and survival, and in exploring the effects of buffalo, clipping surrounding vegetation and ambient fire. Buffalo were removed from one half of the sites across an environmental gradient in a landscape-scale experiment. Juvenile trees grew more slowly but fewer became dormant where buffalo were absent. Experimental clipping of surrounding vegetation produced significantly increased height at the buffalo-absent sites, but not at the buffalo-present sites, demonstrating that the positive effect of buffalo on growth was indirect, by reducing the competitive regime for juvenile trees. Within 3 years of removal of buffalo, ground-level green biomass increased 2–10 times and litter 1.3–2 times over a topographic gradient. Mortality of juvenile trees was not significantly different between buffalo-present and buffalo-absent sites when plots remained unburnt. By contrast, at buffalo-absent sites, three times more juveniles died after late dry-season fires than at unburnt sites, but there was no difference among buffalo-present sites. Overall, the total juvenile tree bank decreased by 7% where buffalo grazed and by 18% where buffalo had been removed. The study demonstrated a major mechanism(s) responsible for recorded changes in vegetation patterns of these savannas, whereby buffalo initiate a cascade of effects by changing ground-level biomass, which change competitive relationships and fuel loads, which then have an impact on tree growth and demography. The results are discussed with respect to dynamics of the juvenile tree bank and implications for long-term sustainability of these wooded savannas.


Acknowledgments

The field project was initiated with funding by the USA Man and Biosphere Program through the auspices of the USA Forest Service, Pacific Region (Grant No 59-PSW-82-005-G to P. A. Werner and P. G. Murphy). Dr M. G. Ridpath (CSIRO) conceived of, and set up, the original buffalo experiment; Mssrs P. Pan Quee, P. Brady, and D. Murphy (CSIRO, resident managers at Kapalga) maintained the buffalo-proof fence and protected the study plots from fires more than once with help from CSIRO and Kakadu National Park staff. Thanks go to W. Murray, J. Browne, K. Stagg and D .Murphy (all CSIRO) for field assistance; to J. S. Harrison (dec. MSU), K. Tidemann (CSIRO), S. Pidsley (CSIRO) and C. R. Dunlop (NT Conservation Commission) for technical assistance; M. Haynes (SIROMATH) for statistical analyses; L. D. Prior (Charles Darwin University (CDU)) for production of graphs and figures; and to Drs G. H. Lauff (MSU), R. J. Wasson (ANU) and B. M. Campbell, K. Gibb and G. J. E. Hill (all CDU) for infrastructure support. The paper was greatly improved by discussions with colleagues at Charles Darwin University and by constructive reviews of drafts by Drs R. J. Williams, L. D. Prior, I. J. McNaught, C. E. R. Lehmann and anonymous reviewers. We are very much indebted to all the above for their support, insights and considerable expertise. This is Contribution No. 1157 of the MSU Kellogg Biological Station and CSIRO TERC Publication No. 1312.


References


Andersen AN , Cook GD , Williams RJ (Eds) (2003) ‘Fire in tropical savannas: the Kapalga experiment.’ (Springer-Verlag: New York)

Anonymous (1999) ‘Kakadu National Park Plan of Management.’ Kakadu Board of Management and Parks Australia. (Commonwealth of Australia: Jabiru, NT)

Archer S (1996) Assessing and interpreting grass–woody plant dynamics. In ‘The ecology and management of grazing systems’. (Eds J Hodgson, AW Illius) pp. 101–134. (CAB International Press: Oxford, UK)

Bowman DMJS (1986) Stand characteristics, understorey associates and environmental correlates of Eucalyptus tetrodonta F.Muell. forests on Gunn Point, northern Australia. Vegetatio 65, 105–113.
Crossref | GoogleScholarGoogle Scholar | [verified on 17 February 2006].

East TJ (1996) Landform evolution. In ‘Landscape and vegetation ecology of the Kakadu region, northern Australia’. (Eds CM Finlayson, I von Oertzen) pp. 37–55. (Kluwer Academic Publishers: Dordrecht, The Netherlands)

Fensham RJ, Bowman DMJS (1992) Stand structure and the influence of overwood on regeneration in tropical eucalypt forest on Melville Island. Australian Journal of Botany 40, 335–352.
Crossref |
open url image1

Fowler J , Cohen L , Jarvis P (1998) ‘Practical statistics for field biology’. 2nd edn. (John Wiley and Sons: Chichester, UK)

Frost P (1996) The ecology of miombo woodlands. In ‘The miombo in transition: woodlands and welfare in Africa’. (Ed. B Campbell) pp. 11–57. (Centre for International Forestry Research: Bogor, Indonesia)

Hill RS, Truswell EM, McLoughlin S, Dettmann ME (1999) Evolution of the Australian flora: fossil evidence. Flora of Australia 1, 251–320. open url image1

Hoare JRL , Hooper RJ , Cheney NP , Jacobsen KLS (1980) A report on the effects of fire in tall open forests and woodland with particular reference to fire management in Kakadu National Park in the Northern Territory. Report to the Australian National Parks and Wildlife Service, Canberra, ACT.

Horn HS (1974) The ecology of secondary succession. Annual Review of Ecology and Systematics 5, 25–37.
Crossref | GoogleScholarGoogle Scholar | open url image1

Lacey CJ (1974) Rhizomes in tropical eucalypts and their role in recovery from fire damage. Australian Journal of Botany 22, 29–38.
Crossref |
open url image1

Lehmann CER (2006) Savanna dynamics in Kakadu National Park: 1964–2004. PhD Thesis, Charles Darwin University, Darwin, NT.

Letts GA , Bassingthwaighte A , De Vos WEL (1979) ‘Feral animals in the Northern Territory.’ (Northern Territory Government Printer: Darwin, NT)

McNaughton SJ (1985) Ecology of a grazing ecosystem: The Serengeti. Ecological Monographs 55, 259–294.
Crossref |
open url image1

McNaughton SJ (1993) Grasses and grazers, science and management. Ecological Applications 3, 17–20.
Crossref |
open url image1

McQuade CV , Arthur JT , Butterworth IJ (1996) Climate and hydrology. In ‘Landscape and vegetation ecology of the Kakadu region, northern Australia’. (Eds CM Finlayson, I von Oertzen) pp. 17–35. (Kluwer Academic Publishers: Dordrecht, The Netherlands)

Menaut J-C, Gignoux J, Prado C, Clobert J (1990) Tree community dynamics in a humid savanna of the Côte-d’Ivoire: modelling the effects of fire and competition with grass and neighbours. Journal of Biogeography 17, 471–481. open url image1

Minchin PR , Dunlop CR (1989) Impact of feral water buffalo on savanna ecosystems in the Australian monsoon tropics. In ‘Forests of the world: IAVS symposium, August 1989’. (IAVS: Uppsala, Sweden)

Norton-Griffiths M (1979) The influence of grazing, browsing, and fire on the vegetation dynamics of the Serengeti. In ‘Serengeti: dynamics of an ecosystem’. (Eds ARE Sinclair, M Norton-Griffiths) pp. 310–352. (University of Chicago Press: Chicago)

Northern Territory Herbarium (2004) ‘NTChecklist Jan 2003.’pdf. http://www.nt.gov.au/ipe/pwcnt/docs/

Press AJ , Lea D , Webb A , Graham A (Eds) (1995) ‘Kakadu: natural and cultural heritage and management.’ (Australian Nature Conservation Agency and NARU ANU: Darwin, NT)

Prior LD, Eamus D (1999) Seasonal changes in leaf water characteristics of Eucalyptus tetrodonta and Terminalia ferdinandiana saplings in a northern Australian savanna. Australian Journal of Botany 47, 587–599.
Crossref | GoogleScholarGoogle Scholar | open url image1

Prior LD, Eamus D (2000) Seasonal changes in hydraulic conductance, xylem embolism and leaf area in Eucalyptus tetrodonta and Eucalyptus miniata saplings in a north Australian savanna. Plant, Cell & Environment 23, 955–965.
Crossref | GoogleScholarGoogle Scholar | open url image1

Prior LD, Eamus D, Duff GA (1997a) Seasonal and diurnal patterns of carbon assimilation, stomatal conductance and leaf water potential in Eucalyptus tetrodonta saplings in a wet-dry savanna in northern Australia. Australian Journal of Botany 45, 241–258.
Crossref | GoogleScholarGoogle Scholar | open url image1

Prior LD, Eamus D, Duff GA (1997b) Seasonal trends in carbon assimilation, stomatal conductance and leaf water potential and growth in Terminalia ferdinandiana, a deciduous tree of northern Australian savannas. Australian Journal of Botany 45, 53–69.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ridpath MG, Waithman J (1988) Controlling feral Asian water buffalo in Australia. Wildlife Society Bulletin 16, 385–390. open url image1

Ridpath MG, Begg RJ, Dudzinski ML, Forbes MA, Graham A (1983) Counting the same populations of large tropical mammals from the ground and from the air. Australian Wildlife Research 10, 487–498.
Crossref | GoogleScholarGoogle Scholar | open url image1

Russell-Smith J , Needham S , Brock J (1995) The physical environment. In ‘Kakadu: natural and cultural heritage and management’. (Eds AJ Press, D Lea, A Webb, A Graham) pp. 94–196. (Australian Nature Conservation Agency and NARU ANU: Darwin, NT)

Russell-Smith J, Whitehead PJ, Cook GD, Hoare JL (2003) Response of Eucalyptus-dominated savanna to frequent fires: Lessons from Munmarlary, 1973–1996. Ecological Monographs 73, 349–375. open url image1

Scholes RJ , Walker BH (1993) ‘An African savanna: synthesis of the Nylsvley study.’ (Cambridge University Press: UK)

Schüle W (1990) Landscapes and climate in prehistory: interactions of wildlife, man, and fire. In ‘Fire in the tropical biota’. (Ed. JG Goldammer) pp. 271–318. (Springer-Verlag: Berlin)

Setterfield SA (2002) Seedling establishment in the Australian tropical savanna: effects of seed supply, soil disturbance and fire. Journal of Applied Ecology 39, 949–959.
Crossref | GoogleScholarGoogle Scholar | open url image1

Sharp BR, Whittaker RJ (2003) The irreversible cattle-driven transformation of a seasonally flooded Australian savanna. Journal of Biogeography 30, 783–802.
Crossref | GoogleScholarGoogle Scholar | open url image1

Silva JF (1996) Biodiversity and stability in tropical savannas. In ‘Ecological studies, Vol. 121. Biodiversity and savanna ecosystem processes: a global perspective’. (Eds OT Solbrig, E Medina, JF Silva) pp. 161–171. (Springer-Verlag: Berlin)

Sinclair ARE , Norton-Griffiths M (1979) ‘Serengeti: dynamics of an ecosystem.’ (University of Chicago Press: Chicago)

Skeat AJ , East TJ , Corbett LK (1996) Impact of feral water buffalo. In ‘Landscape and vegetation ecology of the Kakadu region, northern Australia’. (Eds CM Finlayson, I von Oertzen) pp. 155–177. (Kluwer Academic Publishers: Dordrecht, The Netherlands)

Taylor JA, Dunlop CR (1985) Plant communities of the wet–dry tropics of Australia: the Alligator Rivers Region, Northern Territory. Proceedings of the Ecological Society of Australia 13, 83–127. open url image1

Tulloch DG, Cellier KM (1986) Grazing behaviour of feral buffaloes on a native pasture in the northern portion of the Northern Territory. Australian Wildlife Research 13, 433–439.
Crossref | GoogleScholarGoogle Scholar | open url image1

Van Langevelde F, van de Vijer CADM, Kumar L, van de Koppel J, de Ridder N, van Andel J, Skidmore AK, Hearne JW, Stroosnijder L, Bond WJ, Prins HHT, Rietkerk M (2003) Effects of fire and herbivory on the stability of savanna ecosystems. Ecology 84, 337–350. open url image1

Werner PA (1986) Population dynamics and productivity of selected forest trees in Kakadu National Park. Final report to Australian National Parks and Wildlife Service, Canberra.

Werner PA (2005) Impact of feral water buffalo and fire on growth and survival of mature savanna trees: An experimental field study in Kakadu National Park, northern Australia. Austral Ecology 30, 625–647.
Crossref | GoogleScholarGoogle Scholar | open url image1

Werner PA , Murphy PG (1987) Tropical dry forests of northern Australia: population dynamics and productivity of selected forest trees. Final report to USA Man and Biosphere Program, USDA Forest Service, Pacific Division, Honolulu, HI.

van Wilgen BW , Trolloope WSW , Biggs HC , Potgieter ALF , Brocket BH (2003) Fire as a driver of ecosystem variability. In ‘The Kruger experience: ecology and management of savanna heterogeneity’. (Eds JT DuToit, KH Rogers, HC Biggs) pp. 149–170. (Island Press: Washington DC)

Williams PR, Congdon RA, Grice AC, Clarke PJ (2005) Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire. Austral Ecology 30, 79–90.
Crossref | GoogleScholarGoogle Scholar | open url image1

Williams RJ , Cook GD , Braithwaite RW , Anderson AN , Corbett LK (1995) Australia’s wet–dry tropics: identifying the sensitive zones. In ‘Impacts of climate change on ecosystems and species: terrestrial ecosystems’. (Eds J Pernetta, R Leemans, D Elder, S Humphrey) pp. 39–65. (IUCN: Gland, Switzerland)

Williams RJ, Myers BA, Muller WJ, Duff GA, Eamus D (1997) Leaf phenology of woody species in a northern Australian tropical savanna. Ecology 78, 2542–2558.
Crossref |
open url image1

Williams RJ, Cook GD, Gill AM, Moore PHR (1999) Fire regime, fire intensity and tree survival in a tropical savanna in northern Australia. Australian Journal of Ecology 24, 50–59.
Crossref | GoogleScholarGoogle Scholar | open url image1

Williams RJ , Müller WJ , Wahren CH , Setterfield SA , Cusack JS (2003) Vegetation. In ‘Fire in tropical savannas: the Kapalga experiment’. (Eds AN Andersen, GD Cook, RJ Williams) pp. 79–106. (Springer: New York)

Wilson BA , Russell-Smith J , Williams R (1996) Terrestrial vegetation. In ‘Landscape and vegetation ecology of the Kakadu region, northern Australia’. (Eds CM Finlayson, I von Oertzen) pp. 57–79. (Kluwer Academic Publishers: Dordrecht, The Netherlands)