What Limits Trees in C4 Grasslands and Savannas

Though the distribution of global vegetation can generally be predicted from climate, grasslands are an exception. C4 grassy biomes cover vast areas that are warm enough and wet enough to support closed forests. The extent of this climate mismatch has been revealed by physiologically based global vegetation simulations and by large empirical data sets. Reasons for the existence of grassy biomes have long been debated, polarized into bottom-up (resources) or top-down (fire, herbivory) arguments. Recent studies indicate that both are important, especially in suppressing woody recruits. Grasses are formidable competitors belowground, create highly flammable fuels, and can support large herbivore densities. The net effect on trees is rare and episodic recruitment of adults in tree-fall gaps. The implication is that ecosystem structure and function depend on demographic transitions. Tree cover is increasing and grass/forest boundaries are changing. These changes can have large feedbacks to the earth-atmosphere...

[1]  Florian Jeltsch,et al.  Ecological buffering mechanisms in savannas: A unifying theory of long-term tree-grass coexistence , 2000, Plant Ecology.

[2]  M. Andreae,et al.  Smoking Rain Clouds over the Amazon , 2004, Science.

[3]  W. Trollope Controlling bush encroachment with fire in the savanna areas of South Africa , 1980 .

[4]  Sean C. Thomas,et al.  The worldwide leaf economics spectrum , 2004, Nature.

[5]  D. Louppe,et al.  The effects of brush fires on vegetation: the Aubréville fire plots after 60 years. , 1995 .

[6]  W. Bond,et al.  Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments , 2003 .

[7]  O. V. Auken Shrub Invasions of North American Semiarid Grasslands , 2000 .

[8]  L. Gillson Evidence of Hierarchical Patch Dynamics in an East African Savanna? , 2004, Landscape Ecology.

[9]  N. Ramankutty,et al.  Estimating historical changes in global land cover: Croplands from 1700 to 1992 , 1999 .

[10]  H. Prins,et al.  HERBIVORE POPULATION CRASHES AND WOODLAND STRUCTURE IN EAST-AFRICA , 1993 .

[11]  Bruce P. Hayden,et al.  Ecosystem feedbacks on climate at the landscape scale , 1998 .

[12]  Lindsay B. Hutley,et al.  Testing the grass‐fire cycle: alien grass invasion in the tropical savannas of northern Australia , 2003 .

[13]  A. Edwards,et al.  Rain forest invasion of eucalypt‐dominated woodland savanna, Iron Range, north‐eastern Australia: II. Rates of landscape change , 2004 .

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

[15]  Simon Scheiter,et al.  Effects of four decades of fire manipulation on woody vegetation structure in Savanna. , 2007, Ecology.

[16]  G. Asner,et al.  Net changes in regional woody vegetation cover and carbon storage in Texas Drylands, 1937–1999 , 2003 .

[17]  David M. J. S. Bowman,et al.  Australian Rainforests: Islands of Green in a Land of Fire , 2000 .

[18]  J. Leonard,et al.  The Vegetation of Africa , 1984 .

[19]  W. Hoffmann Post‐Establishment Seedling Success in the Brazilian Cerrado: A Comparison of Savanna and Forest Species 1 , 2000 .

[20]  Ilan Koren,et al.  Measurement of the Effect of Amazon Smoke on Inhibition of Cloud Formation , 2004, Science.

[21]  J. P. Grime,et al.  The plant traits that drive ecosystems: Evidence from three continents , 2004 .

[22]  Jon E. Keeley,et al.  Fire and the Miocene expansion of C4 grasslands , 2005 .

[23]  F. I. Woodward,et al.  The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas , 2003 .

[24]  L. Gillson A ‘large infrequent disturbance’ in an East African savanna , 2006 .

[25]  M. Cochrane,et al.  Fire as a large-scale edge effect in Amazonian forests , 2002, Journal of Tropical Ecology.

[26]  S. Higgins,et al.  Partitioning of Root and Shoot Competition and the Stability of Savannas , 2007, The American Naturalist.

[27]  F. Woodward,et al.  What controls South African vegetation — climate or fire? , 2003 .

[28]  W. Hoffmann,et al.  Constraints to seedling success of savanna and forest trees across the savanna-forest boundary , 2004, Oecologia.

[29]  A. Liedloff,et al.  Modelling the effects of rainfall variability and fire on tree populations in an Australian tropical savanna with the Flames simulation model , 2007 .

[30]  John C. Z. Woinarski,et al.  Response of vegetation and vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptus open forest, Northern Territory, Australia , 2004 .

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

[32]  J. Ehleringer,et al.  Global vegetation change through the Miocene/Pliocene boundary , 1997, Nature.

[33]  W. Bond,et al.  A synthesis of the demography of African acacias , 2001, Journal of Tropical Ecology.

[34]  R. Sage The evolution of C 4 photosynthesis , 2003 .

[35]  H. W. Polley,et al.  Viewpoint: atmospheric CO2, soil water, and shrub/grass ratios on rangelands. , 1997 .

[36]  K. Rogers,et al.  A Framework for Exploring the Determinants of Savanna and Grassland Distribution , 2006 .

[37]  R. B. Jackson,et al.  Regional feedbacks among fire, climate, and tropical deforestation , 2003 .

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

[39]  Robert B. Jackson,et al.  Positive feedbacks of fire, climate, and vegetation and the conversion of tropical savanna , 2002 .

[40]  R. B. Jackson,et al.  Elevated CO2 enhances resprouting of a tropical savanna tree , 2000, Oecologia.

[41]  R. J. Scholes,et al.  Leaf green-up in a semi-arid African savanna –separating tree and grass responses to environmental cues , 2007 .

[42]  David M. J. S. Bowman,et al.  Variation in the composition and structure of tropical savannas as a function of rainfall and soil texture along a large‐scale climatic gradient in the Northern Territory, Australia , 1996 .

[43]  S. Archer,et al.  WOODY PLANT ESTABLISHMENT AND SPATIAL HETEROGENEITY IN GRASSLANDS , 2003 .

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

[45]  G. Polis,et al.  Why Are Parts of the World Green? Multiple Factors Control Productivity and the Distribution of Biomass , 1999 .

[46]  T. Seastedt,et al.  Detritus Accumulation Limits Productivity of Tallgrass PrairieThe effects of its plant litter on ecosystem function make the tallgrass prairie unique among North American biomes , 1986 .

[47]  Kelly K. Caylor,et al.  Determinants of woody cover in African savannas , 2005, Nature.

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

[49]  N. Owen‐Smith,et al.  Pleistocene extinctions: the pivotal role of megaherbivores , 1987, Paleobiology.

[50]  J. Ehleringer The Influence of Atmospheric CO2, Temperature, and Water on the Abundance of C3/C4 Taxa , 2005 .

[51]  M. Kellman Synergistic relationships between fire and low soil fertility in neotropical savannas: a hypothesis. , 1984 .

[52]  D. Schimel,et al.  Mechanisms of shrubland expansion: land use, climate or CO2? , 1995 .

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

[54]  J. Gignoux,et al.  Biomass Cycle and Primary Production , 2006 .

[55]  Stephen R. Carpenter,et al.  The Trophic Cascade in Lakes , 1993 .

[56]  S. McNaughton,et al.  Regulation of shrub dynamics by native browsing ungulates on East African rangeland , 2004 .

[57]  Guy F. Midgley,et al.  A proposed CO2‐controlled mechanism of woody plant invasion in grasslands and savannas , 2000 .

[58]  F. Woodward,et al.  The global distribution of ecosystems in a world without fire. , 2004, The New phytologist.

[59]  W. Bond,et al.  Fire as a global 'herbivore': the ecology and evolution of flammable ecosystems. , 2005, Trends in ecology & evolution.

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

[61]  J. Ehleringer,et al.  C4 photosynthesis, atmospheric CO2, and climate , 1997, Oecologia.

[62]  K. Wiegand,et al.  Multi-scale patterns and bush encroachment in an arid savanna with a shallow soil layer , 2005 .

[63]  J. Carter,et al.  Carbon accounting, land management, science and policy uncertainty in Australian savanna landscapes: introduction and overview , 2005 .

[64]  P. Chesson,et al.  Short-term instabilities and long-term community dynamics. , 1989, Trends in ecology & evolution.

[65]  K. Wiegand,et al.  A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna , 2006 .

[66]  N. Hanan,et al.  A continental‐scale analysis of tree cover in African savannas , 2005 .

[67]  C. Field,et al.  Fire history and the global carbon budget: a 1°× 1° fire history reconstruction for the 20th century , 2005 .

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

[69]  L. Jacobs,et al.  The Origin of Grass-Dominated Ecosystems , 1999 .

[70]  Jean Clobert,et al.  Alternative fire resistance strategies in savanna trees , 1997, Oecologia.

[71]  C. Pendry,et al.  Neotropical seasonally dry forests and Quaternary vegetation changes , 2000 .

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

[73]  W. Trollope,et al.  Fire in Savanna , 1984 .

[74]  P. Pilewskie,et al.  Evolution of gases and particles from a savanna fire in South Africa , 2003 .

[75]  D. Beerling,et al.  Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[76]  M. Pärtel,et al.  ROOT DYNAMICS AND SPATIAL PATTERN IN PRAIRIE AND FOREST , 2002 .

[77]  F. Marret,et al.  Vegetation change in equatorial West Africa: time-slices for the last 150 ka , 2000 .

[78]  Jeremy Russell-Smith,et al.  RESPONSE OF EUCALYPTUS‐DOMINATED SAVANNA TO FREQUENT FIRES: LESSONS FROM MUNMARLARY, 1973–1996 , 2003 .

[79]  W. Bond Large parts of the world are brown or black: A different view on the ‘Green World’ hypothesis , 2005 .

[80]  S. Archer,et al.  SMALL‐MAMMAL REGULATION OF VEGETATION STRUCTURE IN A TEMPERATE SAVANNA , 1997 .

[81]  Vânia Regina Pivello,et al.  Soil-vegetation relationships in cerrado (Brazilian savanna) and semideciduous forest, Southeastern Brazil , 2002, Plant Ecology.

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

[83]  David M. J. S. Bowman,et al.  Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest , 2003 .

[84]  R. Monson,et al.  Evolutionary and Ecological Aspects of Photosynthetic Pathway Variation , 1993 .

[85]  Niall P. Hanan,et al.  Tree–grass coexistence in savannas revisited – insights from an examination of assumptions and mechanisms invoked in existing models , 2004 .

[86]  Houghton,et al.  The U.S. Carbon budget: contributions from land-Use change , 1999, Science.

[87]  P. Reich,et al.  PRESCRIBED FIRE IN OAK SAVANNA: FIRE FREQUENCY EFFECTS ON STAND STRUCTURE AND DYNAMICS , 2001 .

[88]  Barry W. Brook,et al.  Postcards from the past: charting the landscape-scale conversion of tropical Australian savanna to closed forest during the 20th century , 2006, Landscape Ecology.

[89]  O. Solbrig,et al.  The role of topkill in the differential response of savanna woody species to fire , 2003 .

[90]  M. Rietkerk,et al.  EFFECTS OF FIRE AND HERBIVORY ON THE STABILITY OF SAVANNA ECOSYSTEMS , 2003 .

[91]  F. Woodward,et al.  Vegetation dynamics – simulating responses to climatic change , 2004, Biological reviews of the Cambridge Philosophical Society.

[92]  J. Blair,et al.  An Ecosystem in Transition: Causes and Consequences of the Conversion of Mesic Grassland to Shrubland , 2005 .

[93]  R. Sage,et al.  The evolution of C4 photosynthesis. , 2004, The New phytologist.

[94]  Mark West,et al.  Carbon dioxide enrichment alters plant community structure and accelerates shrub growth in the shortgrass steppe , 2007, Proceedings of the National Academy of Sciences.

[95]  C. Lacey,et al.  Woody Clumps and Clumpwoods , 1990 .

[96]  I. Rodríguez‐Iturbe,et al.  Tree‐grass competition in space and time: Insights from a simple cellular automata model based on ecohydrological dynamics , 2002 .

[97]  B. Walker,et al.  The Savannas: Biogeography and Geobotany. , 1987 .

[98]  D. Beerling,et al.  The origin of the savanna biome , 2006 .

[99]  Steven R. Archer,et al.  Rainfall, land use and woody vegetation cover change in semi‐arid Australian savanna , 2005 .

[100]  I. Rodríguez‐Iturbe,et al.  HYDROLOGICALLY DRIVEN HIERARCHICAL COMPETITION–COLONIZATION MODELS: THE IMPACT OF INTERANNUAL CLIMATE FLUCTUATIONS , 2003 .