Costs and benefits of relative bark thickness in relation to fire damage: a savanna/forest contrast

In fire‐prone ecosystems, bark protects the stem bud bank from fire. Absolute bark thickness is a good indicator of this protective function, but it depends on stem size as well as inherent differences between species. Relative bark thickness (i.e. relative to stem diameter) takes the latter into account. We argue that relative bark thickness is an important functional trait offering insights to the evolution of species persistence in fire‐prone habitats. During growth ontogeny different species can acquire absolutely thick bark through having: (i) relatively thick bark (i.e. an early commitment to thick bark) or (ii) relatively thin bark but fast stem diameter growth rates. We test the hypothesis that the most effective way of protecting tree stems from frequent fire is by having relatively thick‐barked small stems. We predict that species with higher relative bark thickness are more common in fire‐prone habitats. In habitats with long fire‐free intervals such as rainforest, delayed investment in bark thickness results in thin bark. We examined the relative bark thickness of woody congeners from Australian non‐fire‐prone forest and fire‐prone savanna and in other tree‐dominated systems world‐wide. We determined the relative cost of acquiring absolute bark thickness of 0.5 cm for different rates of bark allocation. The insulating benefits of bark were considered a linear function of bark thickness. Synthesis. We suggest that relatively thick bark minimizes the costs of acquiring absolutely thick bark, and it confers greater protection to smaller stems. The cost of acquiring thick bark prevents small trees from merely accumulating bark as a consequence of fast height or stem diameter growth. Accordingly, our field survey indicated that forest species had relatively thin bark and acquired thick bark only as a consequence of very large size, while fire‐prone savanna species had relatively thick‐barked small stems. Based on this, relative bark thickness appears to be a good predictor of local fire regimes and is a useful plant functional trait.

[1]  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.

[2]  M. Lawes,et al.  Ecology of plant resprouting: populations to community responses in fire-prone ecosystems , 2011, Plant Ecology.

[3]  M. Lawes,et al.  Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia , 2011, Plant Ecology.

[4]  Juli G Pausas,et al.  Fire as an evolutionary pressure shaping plant traits. , 2011, Trends in plant science.

[5]  W. Bond,et al.  Pushing back in time: the role of fire in plant evolution. , 2011, New Phytologist.

[6]  D. Hartnett,et al.  Fire resistance of tree species explains historical gallery forest community composition , 2011 .

[7]  M. Lawes,et al.  How do small savanna trees avoid stem mortality by fire? The roles of stem diameter, height and bark thickness , 2011 .

[8]  David M J S Bowman,et al.  Flammable biomes dominated by eucalypts originated at the Cretaceous-Palaeogene boundary. , 2011, Nature communications.

[9]  S. Patiño,et al.  Functional explanations for variation in bark thickness in tropical rain forest trees , 2010 .

[10]  M. Lawes,et al.  History of the East Point Monsoon Forest , 2010, Northern Territory Naturalist.

[11]  M. Lawes,et al.  Resprouting as a key functional trait in woody plants--challenges to developing new organizing principles. Sprouting behaviour workshops, Working Group 67, ARC-NZ Research Network for Vegetation Function, Armidale, Australia, 2009-2010. , 2010, The New phytologist.

[12]  D. Bowman,et al.  A wide diversity of epicormic structures is present in Myrtaceae species in the northern Australian savanna biome – implications for adaptation to fire , 2010 .

[13]  M. Lawes,et al.  Savanna woody plant dynamics: the role of fire and herbivory, separately and synergistically , 2010 .

[14]  P. Whitehead,et al.  Culture, ecology and economy of fire management in North Australian Savannas : rekindling the Wurrk tradition , 2009 .

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

[16]  P. Whitehead,et al.  Culture, Ecology and Economy of Savanna Fire Management in Northern Australia: rekindling the Wurrk tradition , 2009 .

[17]  D. Bowman,et al.  Leaf Axil Anatomy and Bud Reserves in 21 Myrtaceae Species from Northern Australia , 2008, International Journal of Plant Sciences.

[18]  J. Gambiza,et al.  Fire‐tolerance mechanisms of common woody plant species in a semiarid savanna in south‐western Zimbabwe , 2007 .

[19]  E. Johnson,et al.  How forest fires kill trees: A review of the fundamental biophysical processes , 2007 .

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

[21]  Douglas Sheil,et al.  Drought, fire and tree survival in a Borneo rain forest, East Kalimantan, Indonesia , 2005 .

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

[23]  P. Reich,et al.  A handbook of protocols for standardised and easy measurement of plant functional traits worldwide , 2003 .

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

[25]  M. Schwartz,et al.  Bark heat resistance of small trees in Californian mixed conifer forests: testing some model assumptions , 2003 .

[26]  W. Hoffmann,et al.  Comparative growth analysis of tropical forest and savanna woody plants using phylogenetically independent contrasts , 2003 .

[27]  J. Barlow,et al.  Morphological correlates of fire-induced tree mortality in a central Amazonian forest , 2003, Journal of Tropical Ecology.

[28]  A. Michelsen,et al.  Post‐fire regeneration strategies and tree bark resistance to heating in frequently burning tropical savanna woodlands and grasslands in Ethiopia , 2002 .

[29]  G. Burrows Epicormic strand structure in Angophora, Eucalyptus and Lophostemon (Myrtaceae): implications for fire resistance and recovery , 2002 .

[30]  P. G. Murphy,et al.  Size-specific biomass allocation and water content of above- and below- ground components of three Eucalyptus species in a northern Australian savanna , 2001 .

[31]  James F. Jackson,et al.  Allometry of Constitutive Defense: A Model and a Comparative Test with Tree Bark and Fire Regime , 1999, The American Naturalist.

[32]  Madhav Gadgil,et al.  Variation in bark thickness in a tropical forest community of Western Ghats in India , 1998 .

[33]  J. Keeley,et al.  Evolution of life histories in Pinus , 1998 .

[34]  M. Pinard,et al.  Fire resistance and bark properties of trees in a seasonally dry forest in eastern Bolivia , 1997, Journal of Tropical Ecology.

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

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

[37]  Jeremy Russell-Smith,et al.  Classification, species richness, and environmental relations of monsoon rain forest in northern Australia , 1991 .

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

[39]  Kevin C. Ryan,et al.  Modeling postfire conifer mortality for long-range planning , 1986 .

[40]  A. Gill Toward an Understanding of Fire-Scar Formation: Field Observation and Laboratory Simulation , 1974 .