Edge fires drive the shape and stability of tropical forests.

In tropical regions, fires propagate readily in grasslands but typically consume only edges of forest patches. Thus, forest patches grow due to tree propagation and shrink by fires in surrounding grasslands. The interplay between these competing edge effects is unknown, but critical in determining the shape and stability of individual forest patches, as well the landscape-level spatial distribution and stability of forests. We analyze high-resolution remote-sensing data from protected Brazilian Cerrado areas and find that forest shapes obey a robust perimeter-area scaling relation across climatic zones. We explain this scaling by introducing a heterogeneous fire propagation model of tropical forest-grassland ecotones. Deviations from this perimeter-area relation determine the stability of individual forest patches. At a larger scale, our model predicts that the relative rates of tree growth due to propagative expansion and long-distance seed dispersal determine whether collapse of regional-scale tree cover is continuous or discontinuous as fire frequency changes.

[1]  W. Hoffmann,et al.  Differences in growth patterns between co-occurring forest and savanna trees affect the forest-savanna boundary , 2009 .

[2]  K. Mckinley,et al.  Fuels or microclimate? Understanding the drivers of fire feedbacks at savanna–forest boundaries , 2012 .

[3]  W. Romme,et al.  Historical Perspective on the Yellowstone Fires of 1988A reconstruction of prehistoric fire history reveals that comparable fires occurred in the early 1700s , 1989 .

[4]  Christie Allan,et al.  The Cerrados of Brazil: Ecology and Natural History of a Neotropical Savanna , 2003 .

[5]  R. B. Jackson,et al.  A Large and Persistent Carbon Sink in the World’s Forests , 2011, Science.

[6]  Janneke HilleRisLambers,et al.  Seed Dispersal Near and Far: Patterns Across Temperate and Tropical Forests , 1999 .

[7]  Holly T. Dublin,et al.  Elephants and Fire as Causes of Multiple Stable States in the Serengeti-Mara Woodlands , 1990 .

[8]  P. Bak,et al.  A forest-fire model and some thoughts on turbulence , 1990 .

[9]  E. Schertzer,et al.  Implications of the spatial dynamics of fire spread for the bistability of savanna and forest , 2015, Journal of mathematical biology.

[10]  Ty Kennedy-Bowdoin,et al.  Regional insight into savanna hydrogeomorphology from termite mounds. , 2010, Nature communications.

[11]  C. J. McGrath,et al.  Effect of exchange rate return on volatility spill-over across trading regions , 2012 .

[12]  P. Fearnside,et al.  Testing for criticality in ecosystem dynamics: the case of Amazonian rainforest and savanna fire. , 2010, Ecology letters.

[13]  W. Hoffmann,et al.  Comparative fire ecology of tropical savanna and forest trees , 2003 .

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

[15]  C. Potter,et al.  Large-scale impoverishment of Amazonian forests by logging and fire , 1999, Nature.

[16]  Ran Nathan Long-Distance Dispersal of Plants , 2006, Science.

[17]  M. Turner Disturbance and landscape dynamics in a changing world. , 2010, Ecology.

[18]  S. Carpenter,et al.  Catastrophic shifts in ecosystems , 2001, Nature.

[19]  A. Huth,et al.  High resolution analysis of tropical forest fragmentation and its impact on the global carbon cycle , 2017, Nature Communications.

[20]  C. Justice,et al.  High-Resolution Global Maps of 21st-Century Forest Cover Change , 2013, Science.

[21]  J. Fragoso,et al.  LONG‐DISTANCE SEED DISPERSAL BY TAPIRS INCREASES SEED SURVIVAL AND AGGREGATES TROPICAL TREES , 2003 .

[22]  Navashni Govender,et al.  The effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in South Africa , 2006 .

[23]  B. Drossel,et al.  Forest fires and other examples of self-organized criticality , 1996, cond-mat/9610201.

[24]  P. Grassberger Critical behaviour of the Drossel-Schwabl forest fire model , 2002, cond-mat/0202022.

[25]  J. Chave,et al.  Modelling forest–savanna mosaic dynamics in man-influenced environments: effects of fire, climate and soil heterogeneity , 2004 .

[26]  J. Kauffman,et al.  Deforestation, Fire Susceptibility, and Potential Tree Responses to Fire in the Eastern Amazon , 1990 .

[27]  M. Scheffer,et al.  Global Resilience of Tropical Forest and Savanna to Critical Transitions , 2011, Science.

[28]  S. Levin,et al.  The Global Extent and Determinants of Savanna and Forest as Alternative Biome States , 2011, Science.

[29]  S. Redner,et al.  Introduction To Percolation Theory , 2018 .

[30]  P. Bak,et al.  A deterministic critical forest fire model , 1990 .

[31]  A. Hastings,et al.  Thresholds and the resilience of Caribbean coral reefs , 2007, Nature.

[32]  Drossel,et al.  Self-organized critical forest-fire model. , 1992, Physical review letters.

[33]  D. Nepstad,et al.  Positive feedbacks in the fire dynamic of closed canopy tropical forests , 1999, Science.

[34]  J. Alroy Effects of habitat disturbance on tropical forest biodiversity , 2017, Proceedings of the National Academy of Sciences.

[35]  S. Gotsch,et al.  Tree topkill, not mortality, governs the dynamics of savanna-forest boundaries under frequent fire in central Brazil. , 2009, Ecology.

[36]  M. Turner Landscape ecology: what is the state of the science? , 2005 .

[37]  D. Nepstad,et al.  Road paving, fire regime feedbacks, and the future of Amazon forests , 2001 .

[38]  R. Didham,et al.  Ecosystem Decay of Amazonian Forest Fragments: a 22‐Year Investigation , 2002 .

[39]  I. Turner Species loss in fragments of tropical rain forest: a review of the evidence. , 1996 .

[40]  S. Higgins,et al.  Atmospheric CO2 forces abrupt vegetation shifts locally, but not globally , 2012, Nature.

[41]  Orr Spiegel,et al.  Mechanisms of long-distance seed dispersal. , 2008, Trends in ecology & evolution.

[42]  D. Schindler,et al.  Eutrophication science: where do we go from here? , 2009, Trends in ecology & evolution.

[43]  Marie-Josée Fortin,et al.  High-resolution global maps of 21st-century annual forest loss: Independent accuracy assessment and application in a temperate forest region of Atlantic Canada , 2017 .

[44]  G. Asner,et al.  Ecosystem‐scale effects of megafauna in African savannas , 2016 .

[45]  Roberta E. Martin,et al.  Large-scale impacts of herbivores on the structural diversity of African savannas , 2009, Proceedings of the National Academy of Sciences.

[46]  J. A. Ratter,et al.  SUCCESSIONAL CHANGES IN CERRADO AND CERRADO/FOREST ECOTONAL VEGETATION IN WESTERN SÃO PAULO STATE, BRAZIL, 1962–2000 , 2006 .

[47]  Glenn R. Moncrieff,et al.  Increasing atmospheric CO2 overrides the historical legacy of multiple stable biome states in Africa. , 2014, The New phytologist.

[48]  M. Cochrane Fire science for rainforests , 2003, Nature.

[49]  W. Laurance,et al.  Predicting the impacts of edge effects in fragmented habitats , 1991 .

[50]  S. Gotsch,et al.  Ecological thresholds at the savanna-forest boundary: how plant traits, resources and fire govern the distribution of tropical biomes. , 2012, Ecology letters.

[51]  J. Pausas,et al.  Disturbance maintains alternative biome states. , 2016, Ecology letters.

[52]  R. Durrett,et al.  Coexistence of grass, saplings and trees in the Staver–Levin forest model , 2014, 1401.5220.

[53]  M. Nowak,et al.  Habitat destruction and the extinction debt , 1994, Nature.

[54]  John W. Hearne,et al.  An improved cellular automaton model for simulating fire in a spatially heterogeneous Savanna system , 2002 .

[55]  W. Hoffmann,et al.  Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes , 2016, Global change biology.

[56]  T. H. Schubert,et al.  Nitrogen and Phosphorus , 1928, Nature.

[57]  H. Paerl,et al.  Controlling Eutrophication: Nitrogen and Phosphorus , 2009, Science.

[58]  Suzana Dragicevic,et al.  Design and implementation of an integrated GIS-based cellular automata model to characterize forest fire behaviour , 2008 .