RESPONSE OF EUCALYPTUS‐DOMINATED SAVANNA TO FREQUENT FIRES: LESSONS FROM MUNMARLARY, 1973–1996

We assess a replicated fire plot experiment undertaken between 1973 and 1996 in two Eucalyptus-dominated savanna vegetation formations (open forest, woodland), at Munmarlary, in monsoonal northern Australia. Four treatments, each with three replicates, were imposed on each vegetation type: annual early dry-season burning; annual late dry- season burning; biennial early dry-season burning; and unburned controls. Treatments were imposed faithfully, with noted exceptions, on 1-ha plots. Fire intensities were typically low (,1000 kW/m) to moderate (1000-2500 kW/m), varied significantly between treatments, and generally were greater in woodland. In both woodland and open forest, pH was sig- nificantly lower and NO 3-N was significantly higher in unburned plots. Organic C was not significantly greater in unburned treatments. Effects of fire regime on other soil chemical properties differed between open forest and woodland sites. Among the grasses, invariant frequent burning led to the dominance of a small number of annual species, notably re- gionally dominant Sorghum. In the absence of burning, annuals declined generally, whereas some perennials increased while most decreased. These responses usually were apparent within the first five years of the experiment. At the relatively small spatial scale of the grass sampling regime, there was high turnover of both annual and perennial grasses. Under low- to moderate-intensity, frequent burning regimes, woody vegetation dominated by ma- ture eucalypts is structurally stable. In the absence of burning for at least five years, there was release of the non-eucalypt, woody component into the midstory; this occurred more rapidly in open forest. Accession of rain forest species occurred on some woodland plots, especially the unburned treatment. In contrast, eucalypts were not released significantly from the understory. Rather, as suggested by other studies, recruitment of eucalypts into the canopy appears to involve significantly reduced root competition through death of dominant eucalypts. Although the Munmarlary experiment provides invaluable quantitative data for exploring relationships between fire regimes and the responses of north Australian savanna systems, it has been less successful in meeting the complex information require- ments of regional fire managers. Replicated experimental fire plot designs, no matter how elegant and rigorously implemented, may substantially fail the test of management rele- vance, given the fundamental requirement for savanna biodiversity managers to experience the integrated effects of fire regimes that vary idiosyncratically over multiple time and spatial scales. We suggest that such information requirements are better met through modest, targeted ''adaptive management'' studies, involving collaborative partnerships between managers and researchers.

[1]  Juan F. Silva,et al.  Increase in the woody component of seasonal savannas under different fire regimes in Calabozo, Venezuela , 2001 .

[2]  R. Fensham Regeneration in tropical eucalypt forest on Melville Island , 1990 .

[3]  P. Werner 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 , 2005 .

[4]  J. José,et al.  Temporal changes in the structure of a Trachypogon savanna protected for 25 years , 1991 .

[5]  M. Swaine,et al.  A re-assessment of a fire protection experiment in north-eastern Ghana savanna. , 1980 .

[6]  G. Friend Population Ecology of Mesembriomys gouldii (Rodentia : Muridae) in the Wet-Dry Tropics of the Northern Territory , 1987 .

[7]  Richard J. Williams,et al.  Fire regime, fire intensity and tree survival in a tropical savanna in northern Australia , 1999 .

[8]  L. Coutinho,et al.  Fire in the Ecology of the Brazilian Cerrado , 1990 .

[9]  G. Harrington,et al.  Recent contraction of wet sclerophyll forest in the wet tropics of Queensland due to invasion by rainforest , 1994 .

[10]  C. G. Trapnell,et al.  THE EFFECTS OF FIRE AND TERMITES ON A ZAMBIAN WOODLAND SOIL , 1976 .

[11]  P. Vitousek,et al.  Biological invasions by exotic grasses, the grass/fire cycle, and global change , 1992 .

[12]  E. O. Mclean Soil pH and Lime Requirement , 1982 .

[13]  P. Lavelle,et al.  Estimate of organic matter turnover rate in a savanna soil by 13C natural abundance measurements. , 1990 .

[14]  R. H. Groves,et al.  Fire and the Australian biota , 1981 .

[15]  S. Higgins,et al.  Fire, resprouting and variability: a recipe for grass–tree coexistence in savanna , 2000 .

[16]  Richard J. Williams,et al.  Patterns of Flowering and Seed Production in Eucalyptus miniata and E. tetrodonta in a Tropical Savanna Woodland, Northern Australia , 1996 .

[17]  Geoffrey J. Cary,et al.  Effects of fire frequency on plant species composition of sandstone communities in the Sydney region: Inter‐fire interval and time‐since‐fire , 1995 .

[18]  David M. J. S. Bowman,et al.  Response of a monsoon forest‐savanna boundary to fire protection, Weipa, northern Australia , 1991 .

[19]  Jeremy Russell-Smith,et al.  Conservation of monsoon rainforest isolates in the Northern Territory, Australia , 1992 .

[20]  W. Allee Ecological Monographs , 1930, Science.

[21]  W. Lonsdale Inviting trouble: Introduced pasture species in northern Australia , 1994 .

[22]  W. Hoffmann FIRE AND POPULATION DYNAMICS OF WOODY PLANTS IN A NEOTROPICAL SAVANNA: MATRIX MODEL PROJECTIONS , 1999 .

[23]  Carl J. Walters,et al.  Adaptive Management of Renewable Resources , 1986 .

[24]  K. Miyanishi,et al.  Nutrient Retention by Savanna Ecosystems: II. Retention After Fire , 1985 .

[25]  Richard J. Williams,et al.  Fire research for conservation management in tropical savannas: Introducing the Kapalga fire experiment , 1998 .

[26]  W. Trollope Ecological Effects of Fire in South African Savannas , 1982 .

[27]  A. Andersen Responses of Ground-Foraging Ant Communities to Three Experimental Fire Regimes in a Savanna Forest of , 1991 .

[28]  P. Whelan,et al.  Observations on the Ecological Significance of Vegetative Reproduction in the Katherine-Darwin Region of the Northern Territory , 1976 .

[29]  C. Lacey Rhizomes in Tropical Eucalypts and Their Role in Recovery From Fire Damage , 1974 .

[30]  A. Watkinson,et al.  Dynamics of shrub encroachment in an African savanna: relative influences of fire, herbivory, rainfall and density dependence , 2001 .

[31]  Emil Spjøtvoll,et al.  An Extension of the T-Method of Multiple Comparison to Include the Cases with Unequal Sample Sizes , 1973 .

[32]  T. A. Afolayan The effect of fires on the vegetation in Kainji Lake National Park, Nigeria , 1978 .

[33]  C. G. Trapnell Ecological Results of Woodland and Burning Experiments in Northern Rhodisia , 1959 .

[34]  M. Forbes,et al.  Counting the Same Populations of Large Tropical Mammals from the Ground and from the Air , 1983 .

[35]  Robert C. Maggio,et al.  Autogenic succession in a subtropical savanna: conversion of grassland to thorn woodland , 1988 .

[36]  R. Ostfeld,et al.  The Ecological Basis of Conservation: Heterogeneity, Ecosystems, and Biodiversity , 1999 .

[37]  M. Swaine,et al.  The effects of fire exclusion on savanna vegetation at Kpong, Ghana , 1992 .

[38]  C. Skarpe,et al.  Dynamics of savanna ecosystems , 1992 .

[39]  S. Archer,et al.  Tree-grass dynamics in a Prosopis-thornscrub savanna parkland: Reconstructing the past and predicting the future , 1995 .

[40]  J. Proctor,et al.  Nature and Dynamics of Forest-Savanna Boundaries. , 1994 .

[41]  J. Mistry Fire in the cerrado (savannas) of Brazil: an ecological review , 1998 .

[42]  D. Franklin Evidence of disarray amongst granivorous bird assemblages in the savannas of northern Australia, a region of sparse human settlement , 1999 .

[43]  R. Scholes,et al.  An African Savanna: Synthesis of the Nylsvley Study. , 1993 .

[44]  M. J. Savage,et al.  The Effect of Fire on Soil and Microclimate , 1984 .

[45]  W. M. Lonsdale,et al.  Assessing the effects of fire on vegetation in tropical savannas , 1991 .

[46]  E. Chidumayo A re-assessment of effects of fire on miombo regeneration in the Zambian Copperbelt , 1988, Journal of Tropical Ecology.

[47]  Jeremy Russell-Smith,et al.  Fire ecology and Aboriginal land management in central Arnhem Land, northern Australia: a tradition of ecosystem management , 2001 .

[48]  Richard J. Williams,et al.  Fire weather in the wet‐dry tropics of the World Heritage Kakadu National Park, Australia , 1996 .

[49]  A. Moreira Effects of fire protection on savanna structure in Central Brazil , 2000 .

[50]  A. Gill,et al.  Fire regimes and biodiversity of forested landscapes of southern Australia. , 2002 .

[51]  Colin R. Trainor,et al.  Responses of lizards to three experiments fires in the savanna forests of Kakadu National Park , 1994 .

[52]  J. Tester Effects of fire frequency on oak savanna in , 1989 .

[53]  G. Cook Effects of frequent fires and grazing on stable nitrogen isotope ratios of vegetation in northern Australia , 2001 .

[54]  A. Gill,et al.  Intervals between prescribed fires in Australia: what intrinsic variation should apply? , 1998 .

[55]  David M. J. S. Bowman,et al.  Factors that control monsoon-rainforest seedling establishment and growth in north Australian Eucalyptus savanna , 1993 .

[56]  J. Russell‐Smith,et al.  Fire regimes and the conservation of sandstone heath in monsoonal northern Australia: frequency, interval, patchiness , 2002 .

[57]  D. Franklin,et al.  Seasonal use of savanna landscapes by the Gouldian finch, Erythrura gouldiae, in the Yinberrie Hills area, Northern Territory , 2001 .

[58]  A. Ash,et al.  How season of grazing and herbivore selectivity influence monsoon tall-grass communities of northern Australia. , 1998 .

[59]  R. Braithwaite Biodiversity and Fire in the Savanna Landscape , 1996 .

[60]  Richard J. Williams,et al.  Seasonal Changes in Fire Behaviour in a Tropical Savanna in Northern Australia , 1998 .

[61]  R. Fensham Interactive effects of fire frequency and site factors in tropical Eucalyptus forest , 1990 .

[62]  David M. J. S. Bowman,et al.  Response of Eucalyptus forest and woodland to four fire regimes at Munmarlary, Northern Territory, Australia , 1988 .

[63]  G. Friend Ecological Studies of a Population of Antechinus bellus (Marsupialia : Dasyuridae) in Troprical Northern Australia , 1985 .

[64]  S. Materechera,et al.  Organic matter, pH and nutrient distribution in soil layers of a savanna Thornveld subjected to different burning frequencies at Alice in the Eastern Cape , 1998 .

[65]  W. J. Panton Changes in post world war ii distribution and status of monsoon rainforests in the Darwin area , 1993 .

[66]  A. W. Moore The influence of annual burning on a soil in the derived savanna zone of Nigeria. , 1960 .

[67]  C. Geldenhuys The Effect of Different Regimes of Annual Burning on Two Woodland Communities in Kavango / DIE INVLOED VAN VERSKILLENDE STELSELS VAN JAARLIKSE BRAND OP TWEE BOOMVELDGEMEENSKAPPE IN KAVANGO , 1977 .

[68]  D. Mueller‐Dombois,et al.  Ecology of Tropical and Subtropical Vegetation. , 1972 .

[69]  N. Cheney,et al.  The Influence of Fuel, Weather and Fire Shape Variables on Fire-Spread in Grasslands , 1993 .

[70]  A. Gill,et al.  Fire regimes and fire management of rainforest communities across northern Australia. , 2002 .

[71]  J. Woinarski Effects of fire on the bird communities of tropical woodlands and open forests in northern Australia , 1990 .

[72]  J. Sturtz,et al.  The use of fire to establish Townsville lucerne in the Northern Territory , 1966 .

[73]  N. M. Tainton,et al.  Ecological Effects of Fire in South African Ecosystems , 1984, Ecological Studies.

[74]  J. R. Charter,et al.  Assessment of the Olokemeji fire-control experiment (investigation 254) 28 years after institution. , 1960 .

[75]  John A. Taylor,et al.  Rainfall in the wet‐dry tropics: Extreme events at Darwin and similarities between years during the period 1870–1983 inclusive , 1985 .

[76]  C. Or Responses of Lizards to Three Experimental Fires in the Savanna Forests of Kakadu National Park , 1994 .

[77]  B. W. Wilgen,et al.  Fire and Plants , 1995, Population and Community Biology Series.

[78]  R. Scholes,et al.  Tree-grass interactions in Savannas , 1997 .

[79]  S. Setterfield The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia , 1997 .

[80]  Jeremy Russell-Smith,et al.  A LANDSAT MSS-derived fire history of Kakadu National Park, monsoonal northern Australia, 1980-94: seasonal extent, frequency and patchiness. , 1997 .

[81]  J. Kerle THE POPULATION DYNAMICS OF A TROPICAL POSSUM, TRICHOSURUS VULPECULA ARNHEMENSIS COLLETT , 1998 .

[82]  G. Cook Effects of fire regimen on two species of epiphytic orchids in tropical savannas of the Northern Territory , 1991 .

[83]  D. Bowman,et al.  Munmarlary revisited: Response of a north Australian Eucalyptus tetrodonta savanna protected from fire for 20 years , 1995 .

[84]  B. A. Wilson Vegetation survey of the Northern Territory, Australia 1990 - explanatory notes to accompany 1: 1000000 map sheets , 1990 .

[85]  G. Cook,et al.  The fate of nutrients during fires in a tropical savanna , 1994 .

[86]  J. Moss,et al.  An experimental investigation, using stomatal conductance and fluorescence, of the flood sensitivity of Boltonia decurrens and its competitors , 1998 .

[87]  R. Story,et al.  Lands of the Adelaide - Alligator area, Northern Territory , 1969 .

[88]  J. J. Mott,et al.  Ecology and management of the world's savannas , 1985 .

[89]  T. O’Connor Synthesis of field experiments concerning the grass layer in the savanna regions of southern Africa , 1985 .

[90]  P. J. Myerscough Flammable Australia. The Fire Regimes and Biodiversity of a Continent , 2003 .

[91]  E. Veenendaal,et al.  Effect of fire and soil texture on soil carbon in a sub-humid savanna (Matopos, Zimbabwe) , 2000 .

[92]  J. Woinarski,et al.  Wildlife of Lancewood [Acacia Shirleyi] Thickets and Woodlands in Northern Australia. 1. Variation in vertebrate species composition across the environmental range occupied by Lancewood vegetation in the Northern Territory , 1995 .