Non-additive effects on plant mixtures flammability in a tropical mountain ecosystem
Helena Roland
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Abstract
Consumability, a component of flammability, describes how well plants burn and may be influenced by species traits and climate change. However, knowledge gaps remain regarding how species mixtures interact and whether non-additive effects are mediated by functional traits and diversity.
This study examined the consumability of species mixtures in Brazil’s tropical montane grasslands (campos de altitude), focusing on traits and species interactions.
Laboratory tests measured remaining biomass (a proxy for consumability) in monospecific and three-species mixtures for seven species. Traits like specific leaf area (SLA), leaf dry matter content (LDMC), fuel moisture at ignition (FMCig) and maximum combustion rate (MCR) were assessed, along with functional diversity, to understand their influence on consumability.
Consumability in species mixtures differed from predictions based on individual species values, indicating non-additive effects. Leptostelma maximum, Pleroma hospita and Chusquea pinifolia, despite low or medium individual consumability, contributed most to mixture consumability. Higher SLA reduced consumability, whereas higher MCR, LDMC and functional index FRich increased it.
Our results suggest that low-consumability species like L. maximum may reduce fire severity, offering opportunities for fire management in the campos de altitude.
These findings highlight the importance of considering species interactions in mixtures and identifying traits that shape plant community flammability.
Keywords: Brazil, campos de altitude, climate change, consumability, ecosystem process, fire management, fire regimes, fire severity, species interactions, trait-based ecology.
References
Alam MA, Wyse SV, Buckley HL, Perry GLW, Sullivan JJ, Mason NWH, Buxton R, Richardson SJ, Curran TJ (2020) Shoot flammability is decoupled from leaf flammability, but controlled by leaf functional traits. Journal of Ecology 108, 641-653.
| Crossref | Google Scholar |
Alam MA, Wyse SV, Buckley HL, Perry GLW, Cui X, Sullivan JJ, Schwilk DW, Curran TJ (2024) Fuel architecture influences interspecific variation in shoot flammability, but not as much as leaf traits. Journal of Ecology 113, 322-338.
| Crossref | Google Scholar |
Anderson HE (1970) Forest fuel ignitibility. Fire Technology 6, 312-319.
| Crossref | Google Scholar |
Assis MV, Mattos EAD (2016) Vulnerabilidade da vegetação de campos de altitude às mudanças climáticas. Oecologia Australis 20, 162-174.
| Crossref | Google Scholar |
Aximoff I, Rodrigues RDC (2011) Histórico dos incêndios florestais no Parque Nacional do Itatiaia. Ciência Florestal 21, 83-92.
| Crossref | Google Scholar |
Bond W, Keeley J (2005) Fire as a global ‘herbivore’: the ecology and evolution of flammable ecosystems. Trends in Ecology & Evolution 20, 387-394.
| Crossref | Google Scholar | PubMed |
Boving I, Celebrezze J, Salladay R, Ramirez A, Anderegg LDL, Moritz M (2023) Live fuel moisture and water potential exhibit differing relationships with leaf‐level flammability thresholds. Functional Ecology 37(11), 2770-2785.
| Crossref | Google Scholar |
Bowman DMJS, Balch JK, Artaxo P, Bond WJ, Carlson JM, Cochrane MA, D’Antonio CM, DeFries RS, Doyle JC, Harrison SP, Johnston FH, Keeley JE, Krawchuk MA, Kull CA, Marston JB, Moritz MA, Prentice IC, Roos CI, Scott AC, Swetnam TW, van der Werf GR, Pyne SJ (2009) Fire in the arth system. Science 324, 481-484.
| Crossref | Google Scholar | PubMed |
Brade AC (1956) A flora do Parque Nacional do Itatiaia. Boletim do Parque Nacional do Itatiaia 5, 1-112 [In Portuguese].
| Google Scholar |
Burton JE, Cawson JG, Filkov AI, Penman TD (2021) Leaf traits predict global patterns in the structure and flammability of forest litter beds. Journal of Ecology 109, 1344-1355.
| Crossref | Google Scholar |
Cardoso AW, Oliveras I, Abernethy KA, Jeffery KJ, Lehmann D, Edzang Ndong J, McGregor I, Belcher CM, Bond WJ, Malhi YS (2018) Grass species flammability, not biomass, drives changes in fire behavior at tropical forest-savanna transitions. Frontiers in Forests and Global Change 1, 6.
| Crossref | Google Scholar |
Della Rocca G, Danti R, Hernando C, Guijarro M, Madrigal J (2018) Flammability of two Mediterranean mixed forests: study of the non-additive effect of fuel mixtures in laboratory. Frontiers in Plant Science 9, 825.
| Crossref | Google Scholar | PubMed |
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.
| Crossref | Google Scholar |
Dias ATC, Berg MP, De Bello F, van Oosten AR, Bílá K, Moretti M (2013) An experimental framework to identify community functional components driving ecosystem processes and services delivery. Journal of Ecology 101, 29-37.
| Crossref | Google Scholar |
Fernandes PM, Cruz MG (2012) Plant flammability experiments offer limited insight into vegetation–fire dynamics interactions. New Phytologist 194, 606-609.
| Crossref | Google Scholar | PubMed |
Finerty GE, De Bello F, Bílá K, Berg MP, Dias ATC, Pezzatti GB, Moretti M (2016) Exotic or not, leaf trait dissimilarity modulates the effect of dominant species on mixed litter decomposition. Journal of Ecology 104, 1400-1409.
| Crossref | Google Scholar |
Gill AM, Zylstra P (2005) Flammability of Australian forests. Australian Forestry 68, 87-93.
| Crossref | Google Scholar |
Grime JP (1977) Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. The American Naturalist 111, 1169-1194.
| Crossref | Google Scholar |
Grootemaat S, Wright IJ, van Bodegom PM, Cornelissen JHC, Cornwell WK (2015) Burn or rot: leaf traits explain why flammability and decomposability are decoupled across species. Functional Ecology 29, 1486-1497.
| Crossref | Google Scholar |
Grootemaat S, Wright IJ, van Bodegom PM, Cornelissen JHC (2017) Scaling up flammability from individual leaves to fuel beds. Oikos 126, 1428-1438.
| Crossref | Google Scholar |
Hantson S, Andela N, Goulden ML, Randerson JT (2022) Human-ignited fires result in more extreme fire behavior and ecosystem impacts. Nature Communications 13, 2717.
| Crossref | Google Scholar | PubMed |
Hardesty J, Myers R, Fulks W (2005) Fire, ecosystems, and people: a preliminary assessment of fire as a global conservation issue. The George Wright Forum 22(4), 78-87.
| Google Scholar |
Hooper DU, Chapin FS, Ewel JJ, Hector A, Inchausti P, Lavorel S, Lawton JH, Lodge DM, Loreau M, Naeem S, Schmid B, Setälä H, Symstad AJ, Vandermeer J, Wardle DA (2005) Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecological Monographs 75, 3-35.
| Crossref | Google Scholar |
Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) (2023) ‘Integrated Fire Management Plan of Itatiaia National Park.’ (Ministry of the Environment) Available at https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/planos-de-manejo-integrado-do-fogo/PMIFParnadoItatiaia_compressed.pdf [verified 25 September 2024]
Kataki R, Konwer D (2001) Fuelwood characteristics of some indigenous woody species of north-east India. Biomass and Bioenergy 20, 17-23.
| Crossref | Google Scholar |
Keeley JE (2009) Fire intensity, fire severity and burn severity: a brief review and suggested usage. International Journal of Wildland Fire 18, 116.
| Crossref | Google Scholar |
Krix DW, Phillips ML, Murray BR (2019) Relationships among leaf flammability attributes and identifying low-leaf-flammability species at the wildland–urban interface. International Journal of Wildland Fire 28(4), 295.
| Crossref | Google Scholar |
Laliberté E, Legendre P (2010) A distance‐based framework for measuring functional diversity from multiple traits. Ecology 91, 299-305.
| Crossref | Google Scholar | PubMed |
Lyra A, Tavares P, Chou SC, Sueiro G, Dereczynski C, Sondermann M, Silva A, Marengo J, Giarolla A (2018) Climate change projections over three metropolitan regions in southeast Brazil using the non-hydrostatic ETA regional climate model at 5-km resolution. Theoretical and Applied Climatology 132, 663-682.
| Crossref | Google Scholar |
Makkonen M, Berg MP, van Logtestijn RS, van Hal JR, Aerts R (2013) Do physical plant litter traits explain non‐additivity in litter mixtures? A test of the improved microenvironmental conditions theory. Oikos 122, 987-997.
| Crossref | Google Scholar |
Martin R, Gordon R, Gutierrez, MA, Lee D, Molina-Terrén DM, Schroeder RA, Sapsis DB, Stephens SL (1993) Assessing the flammability of domestic and wildland vegetation. In ‘Proceedings of the 12th Conference on Fire and Forest Meteorology’. pp. 26-28. (Society of American Foresters) 10.13140/RG.2.1.3999.3680
Mason NWH, Mouillot D, Lee WG, Wilson JB (2005) Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos 111, 112-118.
| Crossref | Google Scholar |
Matos IS, Eller CB, Oliveras I, Mantuano D, Rosado BHP (2021) Three eco‐physiological strategies of response to drought maintain the form and function of a tropical montane grassland. Journal of Ecology 109, 327-341.
| Crossref | Google Scholar |
Murray BR, Hardstaff LK, Phillips ML (2013) Differences in leaf flammability, leaf traits and flammability-trait relationships between native and exotic plant species of dry sclerophyll forest. PLoS One 8, e79205.
| Crossref | Google Scholar | PubMed |
Oliveras I, Malhi Y (2016) Many shades of green: the dynamic tropical forest–savannah transition zones. Philosophical Transactions of the Royal Society B: Biological Sciences 371, 20150308.
| Crossref | Google Scholar | PubMed |
Oliveras I, Meirelles ST, Hirakuri VL, Freitas CR, Miranda HS, Pivello VR (2013) Effects of fire regimes on herbaceous biomass and nutrient dynamics in the Brazilian savanna. International Journal of Wildland Fire 22, 368.
| Crossref | Google Scholar |
Pausas JG, Keeley JE, Schwilk DW (2017) Flammability as an ecological and evolutionary driver. Journal of Ecology 105, 289-297.
| Crossref | Google Scholar |
Parsons AL, Balch JK, de Andrade RB, Brando PM (2015) The role of leaf traits in determining litter flammability of south-eastern Amazon tree species. International Journal of Wildland Fire 24(8), 1143.
| Crossref | Google Scholar |
Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE , Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, De Vos AC, Buchmann N, Funes G, Quétier F, Hodgson JG, Thompson K, Morgan HD, Ter Steege H, Sack L, Blonder B, Poschlod P, Vaieretti MV, Conti G, Staver AC, Aquino S, Cornelissen JHC (2013) New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany 61, 167.
| Crossref | Google Scholar |
Petchey OL, Hector A, Gaston KJ (2004) How do different measures of functional diversity perform? Ecology 85, 847-857.
| Crossref | Google Scholar |
R Core Team (2023) ‘R: a language and environment for statistical computing’. (R Foundation for Statistical Computing: Vienna, Austria) Available at https://www.R-project.org/ [accessed 10 September 2024]
Rego CF, Morgan P, Fernandes P, Hoffman C (2021) Fuel and Fire Behavior Description. In ‘Fire Science: From Chemistry to Landscape Management’. Springer Textbooks in Earth Sciences, Geography and Environment. (Eds CF Rego, P Morgan, P Fernandes, C Hoffman) pp. 101–114. (Springer International Publishing: Cham, Switzerland) 10.1007/978-3-030-69815-7_6
Ritz C, Baty F, Streibig JC, Gerhard D (2015) Dose–response analysis using R. PLoS One 10, e0146021.
| Crossref | Google Scholar | PubMed |
Roderick ML, Berry SL, Noble IR, Farquhar GD (1999) A theoretical approach to linking the composition and morphology with the function of leaves. Functional Ecology 13, 683-695.
| Crossref | Google Scholar |
Rosado BH, Dias AT, de , Mattos EA (2013) Going back to basics: importance of ecophysiology when choosing functional traits for studying communities and ecosystems. Natureza & Conservação 11, 15-22.
| Crossref | Google Scholar |
Safford HD (1999) Brazilian Páramos I. An introduction to the physical environment and vegetation of the campos de altitude. Journal of Biogeography 26, 693-712.
| Crossref | Google Scholar |
Safford HD (2001) Brazilian Päramos. III. Patterns and rates of postfire regeneration in the campos de altitude. Biotropica 33, 282-302.
| Crossref | Google Scholar |
Segadas-Vianna F, Dau L (1965) Ecology of the Itatiaia range, southeastern Brazil. II. Climates and Altitudinal Climatic Zonation. Arquivos do Museu Nacional 53, 31–53. https://biostor.org/reference/248948
Simpson KJ, Ripley BS, Christin P, Belcher CM, Lehmann CER, Thomas GH, Osborne CP (2016) Determinants of flammability in savanna grass species. Journal of Ecology 104, 138-148.
| Crossref | Google Scholar |
Spehn EM, Hector A, Joshi J, Scherer-Lorenzen M, Schmid B, Bazeley-White E, Beierkuhnlein C, Caldeira MC, Diemer M, Dimitrakopoulos PG, Finn JA, Freitas H, Giller PS, Good J, Harris R, Högberg P, Huss-Danell K, Jumpponen A, Koricheva J, Leadley PW, Loreau M, Minns A, Mulder CPH, O’Donovan G, Otway SJ, Palmborg C, Pereira JS, Pfisterer AB, Prinz A, Read DJ, Schulze E-D, Siamantziouras A-SD, Terry AC, Troumbis AY, Woodward FI, Yachi S, Lawton JH (2005) Ecosystem effects of biodiversity manipulations in European grasslands. Ecological Monographs 75, 37-63.
| Crossref | Google Scholar |
Stevens N, Bond W, Feurdean A, Lehmann CER (2022) Grassy ecosystems in the Anthropocene. Annual Review of Environment and Resources 47, 261-289.
| Crossref | Google Scholar |
Tilman D, Knops J, Wedin D, Reich P, Ritchie M, Siemann E (1997) The influence of functional diversity and composition on ecosystem processes. Science 277, 1300-1302.
| Crossref | Google Scholar |
van Altena C, van Logtestijn RSP, Cornwell WK, Cornelissen JHC (2012) Species composition and fire: non-additive mixture effects on ground fuel flammability. Frontiers in Plant Science 3, 63.
| Crossref | Google Scholar | PubMed |
van der Werf GR, Randerson JT, Giglio L, van Leeuwen TT, Chen Y, Rogers BM, Mu M, van Marle MJE, Morton DC, Collatz GJ, Yokelson RJ, Kasibhatla PS (2017) Global fire emissions estimates during 1997–2016. Earth System Science Data 9, 697-720.
| Crossref | Google Scholar |