The role of leaf traits in determining litter flammability of south-eastern Amazon tree species
Amoreena L. Parsons A , Jennifer K. Balch B F , Rafael B. de Andrade C and Paulo M. Brando D EA Pennsylvania State University, Department of Geography, 302 Walker Building, University Park, PA 16802, USA.
B University of Colorado-Boulder, Department of Geography, 110 Guggenheim Hall, Boulder, CO 80309, USA.
C Universidade Estadual de Campinas, Departamento de Biologia Animal, Campinas, SP 13083, Brazil.
D Instituto de Pesquisa Ambiental da Amazônia, 66035-170, Belém, Pará, Brazil.
E Woods Hole Research Center, 149 Woods Hole Road, Falmouth, MA 02450-1644, USA.
F Corresponding author. Email: jennifer.balch@colorado.edu
International Journal of Wildland Fire 24(8) 1143-1153 https://doi.org/10.1071/WF14182
Submitted: 7 October 2014 Accepted: 27 July 2015 Published: 13 October 2015
Abstract
Leaf traits can limit or promote flammability, but how these traits vary and influence forest flammability in humid tropical forests is unknown. Species within the south-eastern transitional forests of the Brazilian Amazon are experiencing fire, particularly surface fires, with greater frequency and severity than historically recorded. In this study, the leaf traits and consequent burning characteristics of the 17 most abundant species in a transitional forest in Mato Grosso, Brazil were analysed through controlled combustion experiments and leaf trait measurements. Mean maximum flame height (range 52–108 cm), flaming duration (range 21–71 s) and mass loss (range 82–97%), which relate to a fuel’s combustibility and consumability, varied substantially across species. Measured leaf traits, mainly surface area and volume, accounted for 78% of this variability. The most flammable species were those with thin, lightweight and loosely packed leaves, which produced rapid, intense fires that consumed larger fuel amounts. The least flammable species had thick, large and densely packed leaves. In diverse tropical forests, analysing the relationship between species-specific leaf traits and flammability will yield insights into fire behaviour and future forest composition in a frontier zone where exposure to anthropogenic fire is high.
Additional keywords: Brazilian Amazon, combustion experiments, experimental burns, fire ecology, forest fragmentation, tropical forests.
References
Afifi A, May S, Clark VA (Eds) (2003) ‘Computer-aided Multivariate Analysis’, 4th edn. (CRC Press: Boca Raton).Alencar AAC, Solórzano LA, Nepstad DC (2004) Modeling forest understory fires in an eastern Amazonian landscape. Ecological Applications 14, 139–149.
| Modeling forest understory fires in an eastern Amazonian landscape.Crossref | GoogleScholarGoogle Scholar |
Alessio GA, Peñuelas J, Llusià J, Ogaya R, Estiarte M, De Lillis M (2008) Influence of water and terpenes on flammability in some dominant Mediterranean species. International Journal of Wildland Fire 17, 274–286.
| Influence of water and terpenes on flammability in some dominant Mediterranean species.Crossref | GoogleScholarGoogle Scholar |
Alexander ME, Cruz MG (2012) Interdependencies between flame length and fireline intensity in predicting crown fire initiation and crown scorch height. International Journal of Wildland Fire 21, 95–113.
| Interdependencies between flame length and fireline intensity in predicting crown fire initiation and crown scorch height.Crossref | GoogleScholarGoogle Scholar |
Anderson HE (1970) Forest fuel ignitibility. Fire Technology 6, 312–319.
| Forest fuel ignitibility.Crossref | GoogleScholarGoogle Scholar |
Aragão LEOC, Shimabukuro YE (2010) The incidence of fire in Amazonian forests with implications for REDD. Science 328, 1275–1278.
| The incidence of fire in Amazonian forests with implications for REDD.Crossref | GoogleScholarGoogle Scholar |
Balch JK, Nepstad DC, Brando PM, Curran LM, Portela O, de Carvalho O, Lefebvre P (2008) Negative fire feedback in a transitional forest of southeastern Amazonia. Global Change Biology 14, 2276–2287.
| Negative fire feedback in a transitional forest of southeastern Amazonia.Crossref | GoogleScholarGoogle Scholar |
Balch JK, Nepstad DC, Curran LM, Brando PM, Portela O, Guilherme P, Reuning-Scherer JD, de Carvalho O (2011) Size, species, and fire behavior predict tree and liana mortality from experimental burns in the Brazilian Amazon. Forest Ecology and Management 261, 68–77.
| Size, species, and fire behavior predict tree and liana mortality from experimental burns in the Brazilian Amazon.Crossref | GoogleScholarGoogle Scholar |
Balch JK, Massad TJ, Brando PM, Nepstad DC, Curran LM (2013) Effects of high-frequency understorey fires on woody plant regeneration in southeastern Amazonian forests. Philosophical Transactions of the Royal Society B: Biological Sciences 368,
| Effects of high-frequency understorey fires on woody plant regeneration in southeastern Amazonian forests.Crossref | GoogleScholarGoogle Scholar |
Brando PM, Nepstad DC, Balch JK, Bolker B, Christman MC, Coe M, Putz FE (2012) Fire-induced tree mortality in a neotropical forest: the roles of bark traits, tree size, wood density and fire behavior. Global Change Biology 18, 630–641.
| Fire-induced tree mortality in a neotropical forest: the roles of bark traits, tree size, wood density and fire behavior.Crossref | GoogleScholarGoogle Scholar |
Brando PM, Balch JK, Nepstad DC, Morton DC, Putz FE, Coe MT, Silvério D, Macedo MN, Davidson EA, Nóbrega C, Alencar A, Soares-Filho B (2014) Abrupt increases in Amazonian tree mortality due to drought-fire interactions. Proceedings of the National Academy of Sciences of the United States of America 111, 6347–6352.
| Abrupt increases in Amazonian tree mortality due to drought-fire interactions.Crossref | GoogleScholarGoogle Scholar |
Clarke PJ, Prior LD, French BJ, Vincent B, Knox KJE, Bowman DMJS (2014) Using a rainforest–flame forest mosaic to test the hypothesis that leaf and litter fuel flammability is under natural selection. Oecologia 176, 1123–1133.
| Using a rainforest–flame forest mosaic to test the hypothesis that leaf and litter fuel flammability is under natural selection.Crossref | GoogleScholarGoogle Scholar |
Cochrane MA, Alencar A, Schulze MD, Souza CM, Nepstad DC, Lefebvre P, Davidson EA (1999) Positive feedbacks in the fire dynamic of closed canopy tropical forests (1832–1835). Science 284, 1832–1835.
| Positive feedbacks in the fire dynamic of closed canopy tropical forests (1832–1835).Crossref | GoogleScholarGoogle Scholar |
Coe MT, Marthews TR, Costa MH, Galbraith DR, Greenglass NL, Imbuzeiro HMA, Levine NM, Malhi Y, Moorcroft PR, Muza MN, Powell TL, Saleska SR, Solorzano LA, Wang J (2013) Deforestation and climate feedbacks threaten the ecological integrity of south–southeastern Amazonia. Philosophical Transactions of the Royal Society of London : Biological Sciences 368, 20120155
| Deforestation and climate feedbacks threaten the ecological integrity of south–southeastern Amazonia.Crossref | GoogleScholarGoogle Scholar |
Cornelissen JHC, Lavorel S, Garnier E, Díaz S, Buchmann N, Gurvich DE, Reich PB, ter Steege H, Morgan HD, van der Heijden MGA, Pausas JG, Poorter H (2003) A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany 51, 335–380.
| A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Crossref | GoogleScholarGoogle Scholar |
Costa MH, Pires GF (2010) Effects of Amazon and Central Brazil deforestation scenarios on the duration of the dry season in the arc of deforestation. International Journal of Climatology 30, 1970–1979.
| Effects of Amazon and Central Brazil deforestation scenarios on the duration of the dry season in the arc of deforestation.Crossref | GoogleScholarGoogle Scholar |
Coutinho LM (1990) Fire in the ecology of the Brazilian Cerrado. In ‘Fire in the Tropical Biota: Ecosystem Processes and Global Challenges’. (Ed. JG Goldammer) pp. 82–105. (Springer Verlag: Berlin)
Crawley MJ (2005) ‘Statistics: an Introduction using R.’ (J. Wiley & Sons: Chichester)
Davidson EA, de Araujo AC, Artaxo P, Balch JK, Brown IF, Bustamante MMC, Coe MT, DeFries RS, Keller M, Longo M, Munger JW, Schroeder W, Soares-Filho BS, Souza CM, Wofsy SC (2012) The Amazon basin in transition. Nature 481, 321–328.
| The Amazon basin in transition.Crossref | GoogleScholarGoogle Scholar |
De Lillis M, Bianco PM, Loreto F (2009) The influence of leaf water content and isoprenoids on flammability of some Mediterranean woody species. International Journal of Wildland Fire 18, 203–212.
| The influence of leaf water content and isoprenoids on flammability of some Mediterranean woody species.Crossref | GoogleScholarGoogle Scholar |
de Magalhães RMQ, Schwilk DW (2012) Leaf traits and litter flammability: evidence for non-additive mixture effects in a temperate forest. Journal of Ecology 100, 1153–1163.
| Leaf traits and litter flammability: evidence for non-additive mixture effects in a temperate forest.Crossref | GoogleScholarGoogle Scholar |
Dimitrakopoulos AP, Papaioannou KK (2001) Flammability assessment of Mediterranean forest fuels. Fire Technology 37, 143–152.
| Flammability assessment of Mediterranean forest fuels.Crossref | GoogleScholarGoogle Scholar |
Engber EA, Varner JM (2012) Patterns of flammability of the California oaks: the role of leaf traits. Canadian Journal of Forest Research 42, 1965–1975.
| Patterns of flammability of the California oaks: the role of leaf traits.Crossref | GoogleScholarGoogle Scholar |
Etlinger MG, Beall FC (2004) Development of a laboratory protocol for fire performance of landscape plants. International Journal of Wildland Fire 13, 479–488.
| Development of a laboratory protocol for fire performance of landscape plants.Crossref | GoogleScholarGoogle Scholar |
Fernandes PM, Cruz MG (2012) Plant flammability experiments offer limited insight into vegetation–fire dynamics interactions. New Phytologist 194, 606–609.
| Plant flammability experiments offer limited insight into vegetation–fire dynamics interactions.Crossref | GoogleScholarGoogle Scholar |
Fonda RW (2001) Burning characteristics of needles from eight pine species. Forest Science 47, 390–396.
Frost PHG, Robertson F (1987) The ecological effects of fire in savannas. In ‘Determinants of Tropical Savannas’. (Ed. BH Walker) pp. 93–141. (IRL Press Limited: Oxford, UK)
Gagnon PR, Passmore HA, Platt WJ, Myers JA, Paine CET, Harms KE (2010) Does pyrogenicity protect burning plants? Ecology 91, 3481–3486.
| Does pyrogenicity protect burning plants?Crossref | GoogleScholarGoogle Scholar |
Ganteaume A, Marielle J, Corinne LM, Thomas C, Laurent B (2011) Effects of vegetation type and fire regime on flammability of undisturbed litter in Southeastern France. Forest Ecology and Management 261, 2223–2231.
| Effects of vegetation type and fire regime on flammability of undisturbed litter in Southeastern France.Crossref | GoogleScholarGoogle Scholar |
Gentry AH (1993) ‘A Field Guide to the Families and Genera of Woody Plants of Northwest South America (Colombia, Ecuador, Peru), with Supplementary Notes on Herbaceous Taxa.’ (Conservation International: Washington, DC)
Goubitz S, Werger MJA, Ne’eman G (2003) Germination response to fire-related factors of seeds from non-serotinous and serotinous cones. Plant Ecology 169, 195–204.
| Germination response to fire-related factors of seeds from non-serotinous and serotinous cones.Crossref | GoogleScholarGoogle Scholar |
Hoffmann WA (1999) Fire and population dynamics of woody plants in a neotropical savanna: matrix model projections. Ecology 80, 1354–1369.
| Fire and population dynamics of woody plants in a neotropical savanna: matrix model projections.Crossref | GoogleScholarGoogle Scholar |
Instituto Nacional de Pesquisas Espaciais & National Institute for Space Research (2011) Projeto Prodes Monitoramento da Florsta Amazonica Brasileira por Satélite Prodes. Available at http://www.obt.inpe.br/prodes/ [Verified 25 August 2015].
Kane JM, Varner JM, Hiers JK (2008) The burning characteristics of southeastern oaks: discriminating fire facilitators from fire impeders. Forest Ecology and Management 256, 2039–2045.
| The burning characteristics of southeastern oaks: discriminating fire facilitators from fire impeders.Crossref | GoogleScholarGoogle Scholar |
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.
| Fire as an evolutionary pressure shaping plant traits.Crossref | GoogleScholarGoogle Scholar |
Keeley PJE, Bond WJ, Bradstock RA, Pausas JG, Rundel PW (2012) ‘Fire in Mediterranean Ecosystems: Ecology, Evolution and Management.’ (Cambridge University Press: Cambridge, NY)
Laurance WF, Williamson GB (2001) Positive feedbacks among forest fragmentation, drought, and climate change in the Amazon. Conservation Biology 15, 1529–1535.
| Positive feedbacks among forest fragmentation, drought, and climate change in the Amazon.Crossref | GoogleScholarGoogle Scholar |
Lewis SL, Brando PM, Phillips OL, van der Heijden GMF, Nepstad D (2011) The 2010 Amazon drought. Science 331, 554
| The 2010 Amazon drought.Crossref | GoogleScholarGoogle Scholar |
Martin RE, Gordon DA, Gurierrez ME, Lee DS, Molina DM, Schroeder RA, Sapsis DB, Stephens SL, Chambers M (1994) Assessing the flammability of domestic and wildland vegetation. In ‘Proceedings of the 12th Conference on Fire and Forest Meteorology’, 26–28 October 1993, Jekyll Island, GA. Society of American Foresters, Publ. 94–02, pp. 131–137. (Bethesda, MD)
Miranda HS, Bustamante MMC, Miranda AC (2002) The fire factor. In ‘The Cerrados of Brazil’. (Eds PS Oliveira, RJ Marquis) pp. 51–68. (Columbia University Press: New York, NY)
Montgomery KR, Cheo PC (1971) Notes: effect of leaf thickness on ignitibility. Forest Science 17, 475–478.
Morton DC, Page YL, DeFries R, Collatz GJ, Hurtt GC (2013) Understorey fire frequency and the fate of burned forests in southern Amazonia. Philosophical Transactions of the Royal Society B: Biological Sciences 368, 20120163
| Understorey fire frequency and the fate of burned forests in southern Amazonia.Crossref | GoogleScholarGoogle Scholar |
Nelson RM (1980) Flame characteristics for fires in southern fuels. USDA Forest Service, Southeast Forest Experiment Station, Research Paper SE-205. (Asheville, NC)
Nepstad DC, Stickler CM, Soares-Filho B, Merryl F (2008) Interactions among Amazon land use, forests and climate: prospects for a near-term forest tipping point. Philosophical Transactions of the Royal Society B: Biological Sciences 363, 1737–1746.
| Interactions among Amazon land use, forests and climate: prospects for a near-term forest tipping point.Crossref | GoogleScholarGoogle Scholar |
Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner H (2013) vegan: community ecology package (Version 2.0–10). Available at http://cran.r-project.org/web/packages/vegan/index.html [Verified 25 August 2015]
Ormeño E, Céspedes B, Sánchez IA, Velasco-García A, Moreno JM, Fernandez C, Baldy V (2009) The relationship between terpenes and flammability of leaf litter. Forest Ecology and Management 257, 471–482.
| The relationship between terpenes and flammability of leaf litter.Crossref | GoogleScholarGoogle Scholar |
Page WG, Jenkins MJ, Runyon JB (2012) Mountain pine beetle attack alters the chemistry and flammability of lodgepole pine foliage. Canadian Journal of Forest Research 42, 1631–1647.
| Mountain pine beetle attack alters the chemistry and flammability of lodgepole pine foliage.Crossref | GoogleScholarGoogle Scholar |
Papió C, Trabaud L (1990) Structural characteristics of fuel components of five Mediterranean shrubs. Forest Ecology and Management 35, 249–259.
| Structural characteristics of fuel components of five Mediterranean shrubs.Crossref | GoogleScholarGoogle Scholar |
Pausas JG (2015a) Bark thickness and fire regime. Functional Ecology 29, 315–327.
| Bark thickness and fire regime.Crossref | GoogleScholarGoogle Scholar |
Pausas JG (2015b) Evolutionary fire ecology: lessons learned from pines. Trends in Plant Science 20, 318–324.
| Evolutionary fire ecology: lessons learned from pines.Crossref | GoogleScholarGoogle Scholar |
Pausas JG, Schwilk D (2012) Fire and plant evolution. New Phytologist 193, 301–303.
| Fire and plant evolution.Crossref | GoogleScholarGoogle Scholar |
Pausas JG, Alessio GA, Moreira B, Corcobado G (2012) Fires enhance flammability in Ulex parviflorus. New Phytologist 193, 18–23.
| Fires enhance flammability in Ulex parviflorus.Crossref | GoogleScholarGoogle Scholar |
Pivello VR (2011) The use of fire in the Cerrado and Amazonian rainforests of Brazil: past and present. Fire Ecology 7, 24–39.
| The use of fire in the Cerrado and Amazonian rainforests of Brazil: past and present.Crossref | GoogleScholarGoogle Scholar |
R Development Core Team (2013) R: A language and environment for statistical computing. (R Foundation for Statistical Computing: Vienna). Available at http://www.R-project.org/ [Verified 25 August 2015]
Rasband WS (2012) ImageJ. (US National Institutes of Health: Bethesda, MD). Available at http://imagej.nih.gov/ij/ [Verified 25 August 2015]
Ray D, Nepstad D, Moutinho P (2005) Micrometeorological and canopy controls of fire susceptibility in a forested Amazon landscape. Ecological Applications 15, 1664–1678.
| Micrometeorological and canopy controls of fire susceptibility in a forested Amazon landscape.Crossref | GoogleScholarGoogle Scholar |
Rohlf FJ (2000) NTSYS-pc: numerical taxonomy and multivariate analysis system. Exeter Software Manual Version 2.0. (Setauket, NY)
Rothermel RC (1972) A mathematical model for predicting fire spread in wildland fuels. USDA Forest Service, Intermountain Forest & Range Experiment Station, Research paper INT-115. (Ogden, UT)
Sarmiento G, Monasterio M (1983) Life forms and phenology. In ‘Tropical Savannas’. (Ed. F Bourliére) pp. 79–108. (Elsevier Scientific Publishing: Amsterdam)
Saura-Mas S, Paula S, Pausas JG, Lloret F (2010) Fuel loading and flammability in the Mediterranean Basin woody species with different post-fire regenerative strategies. International Journal of Wildland Fire 19, 783–794.
| Fuel loading and flammability in the Mediterranean Basin woody species with different post-fire regenerative strategies.Crossref | GoogleScholarGoogle Scholar |
Scarff FR, Westoby M (2006) Leaf litter flammability in some semi-arid Australian woodlands. Functional Ecology 20, 745–752.
| Leaf litter flammability in some semi-arid Australian woodlands.Crossref | GoogleScholarGoogle Scholar |
Schwilk DW, Ackerly DD (2001) Flammability and serotiny as strategies: correlated evolution in pines. Oikos 94, 326–336.
| Flammability and serotiny as strategies: correlated evolution in pines.Crossref | GoogleScholarGoogle Scholar |
Schwilk DW, Caprio AC (2011) Scaling from leaf traits to fire behaviour: community composition predicts fire severity in a temperate forest. Journal of Ecology 99, 970–980.
| Scaling from leaf traits to fire behaviour: community composition predicts fire severity in a temperate forest.Crossref | GoogleScholarGoogle Scholar |
van Altena C, van Logtestijn R, Cornwell W, Cornelissen H (2012) Species composition and fire: non-additive mixture effects on ground fuel flammability. Functional Plant Ecology 3,
| Species composition and fire: non-additive mixture effects on ground fuel flammability.Crossref | GoogleScholarGoogle Scholar |
VanderWeide BL, Hartnett DC (2011) Fire resistance of tree species explains historical gallery forest community composition. Forest Ecology and Management 261, 1530–1538.
| Fire resistance of tree species explains historical gallery forest community composition.Crossref | GoogleScholarGoogle Scholar |