j.g. pausas' blog

La huella del fuego es un documental sobre incendios forestales en España realizado por el equipo del programa Crónica, de La 2 de TVE, y que se emitió el 28 Noviembre 2016. En él participaron algunas de las personas que recientemente realizaron el decálogo sobre incendios forestales (decálogo | blog). Podéis ver un  resumen del documental, o el programa entero en www.rtve.es/alacarta.

Nota: el documental no está relacionado con el libro que tiene el mismo título (de L. Otero 2006), que describe la historia de los bosques de Tierra del Fuego.

We live on a flammable planet [1,2] yet there is little consensus on the origin and evolution of flammability in our flora [3]. Part of the problem lies in the concept of flammability. In a recent paper [4] we suggest that flammability should not be viewed as a single quantitative trait or metric, rather we propose that flammability has three major dimensions that are not necessarily correlated: ignitability, heat release, and fire spread rate. These dimensions define three flammability strategies observed in fire-prone ecosystems: the non-flammable, the fast-flammable and the hot-flammable strategy (with low ignitability, high flame spread rate and high heat release, respectively). The non-flammable strategy refers to plants that do not burn (or rarely) in natural conditions despite living in fire-prone ecosystems: this is because they have biomass with very low ignitability (low flammability at the organ scale) or because their plant structure does not allow the ignition of the biomass (low flammability at the individual scale). The hot- and the fast-flammable strategies refer to flammable plants with contrasted heat release and spread rate. Flammability strategies increase the survival or reproduction under recurrent fires, and thus, plants in fire-prone ecosystems benefit from acquiring one of them; they represent different (alternative) ways to live under recurrent fires. This novel framework on different flammability strategies helps us to understand variability in flammability across scales [4].

Figure: Conceptual model describing the three plant flammability strategies in fire-prone ecosystems. While many plants fall at intermediate levels of these axes (i.e., the null model for flammability), plants in fire-prone ecosystems benefit from being at the extremes, forming the three flammability strategies considered here. From [4]

References
[1] The-fire-overview-effect, jgpausas.blogs.uv.es/2016/09/18/

[2]  A new global fire map, jgpausas.blogs.uv.es/2013/03/06/   [doi | pdf]

[3] Pausas J.G. & Moreira B. 2012. Flammability as a biological concept. New Phytol.  194: 610-613. [doi | wiley | pdf]

[4] Pausas J.G., Keeley J.E., Schwilk D.W. 2017. Flammability as an ecological and evolutionary driver. J. Ecol. 105: 289-297 [doi | wiley | pdf | brief for managers]

The first version of this paper was my talk at the University of Campinas, Unicamp: link

UPDATE: paper featured on the cover of J Ecol 105(2): cover | blog

There is a tendency to think that fires will increase in the near future due to global warming. This is because many fire risk prediction are based on climate only. However fire regime changes not only depend on climate [1]; there are other factors, like land-use changes, CO₂, plant invasion, fragmentation, etc. that are also important drivers of change in fire activity [1]. Even plant drought stress (and flammability) not only depends on climate [2,3].

A recent simulation study [4] suggests that global burned area is certainly predicted to increase in the following decades when simulations are based on climate only (blue line in the figure below). However, adding the effect increased CO₂ reduces the predicted burned area to no increase (green line below). Furthermore, when adding increased population density and urbanization (black and red lines), the model predicts much more area burnt in the last century (black lines 1900-2000) and a reduction of future burned area (red lines). The predicted reduction of fire during 1900-2000 is consistent with global charcoal records [5] and can be explained by increasing agriculture, land use and fragmentation. Overall, this study suggests that global area burned is unlikely to increase in the following decades.

Note that 1) this is a model, so take it with caution! 2) This model is at the global scale, but changes in different directions are expected in different regions, and this can have biodiversity consequences (even if the global balance is steady); for instance, in the Mediterranean Basin, fire are likely to keep increasing as land abandonment and fuels are increasing [6]. And 3) there is a high uncertainty in some fire drivers. For instance, temperature is likely to keep increasing, however, rainfall and wind changes are very uncertain, and landuse and emissions are subject to uncertain changes in environmental policies in different countries. In any case, this study gives us an idea of the possible sensitivity of different parameters.

Figure: Simulation of global area burned for 1900 to 2100 under different scenarios: a) climate only (blue line); b) climate + CO2 (green); c) climate + CO2 + population & urbanization (black lines; red area for the future predictions). From [4].

References
[1] Pausas J.G. & Keeley J.E., 2014. Abrupt climate-independent fire regime changes. Ecosystems 17: 1109-1120. [doi | pdf | blog]

[2] De Cáceres M, et al. 2015. Coupling a water balance model with forest inventory data to predict drought stress: the role of forest structural changes vs. climate changes. Agr. For. Meteorol. 213: 77–90. [doi | pdf | suppl. | blog]

[3] Luo, Y. & H. Y. H. Chen. 2015. Climate change-associated tree mortality increases without decreasing water availability. Ecol, Let. 18:1207-1215.

[4] Knorr W, Arneth A, & Jiang L, 2016. Demographic controls of future global fire risk. Nature Clim. Change 6:781-785.

[5] Marlon JR, et al. (2008). Climate and human influences on global biomass burning over the past two millennia. Nature Geosci, 1, 697-702.

[6] Pausas J.G. & Fernández-Muñoz S. 2012. Fire regime changes in the Western Mediterranean Basin: from fuel-limited to drought-driven fire regime. Climatic Change 110: 215-226. [doi | pdf | blog]

There are plants in which fire can breaks seed dormancy and stimulate germination. In some species, it is the heat of the fire that breaks seed dormancy and triggers germination (heat-stimulated germination, [1, 2]). In others, germination is stimulated by chemicals produced during the combustion of the organic matter (e.g., chemicals found in the smoke and charred wood) [1, 3]; we call this process, smoke-stimulated germination [5]. That is, in fire-prone ecosystems many plants have evolved seeds with sensitivity to heat and/or to chemicals produced by fire [1, 2, 3].

There are many species from a wide phylogenetic range with smoke-stimulated germination [5]; they appear in different regions worldwide and are stimulated by different combustion-related products, both organic and inorganic [4, 5]. All this suggest that smoke-stimulated germination is a trait that has appeared multiple times during the evolution, and thus is another example of convergent evolution [5].

In the Mediterranean Basin we currently know about 67 species (from 19 families) showing a significant increase in germination in response to smoke [6]. Families with many smoke-stimulated species in this region are Lamiaceae, Ericaceae and Asteraceae. However, there is still a lot of research to be done on smoke-stimulated germination in Mediterranean Basin flora, as many species have not yet been tested; in fact, very few annuals has been tested [6] despite there is evidence from field studies (3) and from other Mediterranean regions suggesting that smoke-stimulated germination is important in annuals.

But remember, plants are not the only organisms that have evolved in response to chemicals present in the smoke, humans too! [7].

Figure: Germination (proportion of seeds) in control conditions (light yellow) and after a smoke treatment (blue) for four Mediterranean species in which germination is strongly dependent on smoke: Coris monspeliensis (Primulaceae), Erica umbellata (Ericaceae), Onopordum caricum (Asteraceae) and Stachys cretica (Lamiaceae) See [6].

References
[1] Moreira B., Tormo J., Estrelles E., Pausas J.G. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Ann. Bot. 105: 627-635. [pdf | doi | blog]

[2] Moreira B and Pausas JG. 2012. Tanned or Burned: the role of fire in shaping physical seed dormancy. PLoS ONE 7:e51523. [doi | plos | pdf]

[3] Tormo, J., B. Moreira, and J. G. Pausas. 2014. Field evidence of smoke-stimulated seedling emergence and establishment in Mediterranean Basin flora. J. Veget. Sci. 25: 771-777. [doi | wiley | pdf | blog ]

[4] Smoke-stimulated germination, jgpausas.blogs.uv.es/2011/12/02/

[5] Keeley J.E. & Pausas J.G. 2018. Evolution of ‘smoke’ induced seed germination in pyroendemic plants. South African J. Bot. 115: 251-255 [doi | pdf] <- New

[6] Moreira B. & Pausas J.G. 2018. Shedding light through the smoke on the germination of Mediterranean Basin flora. South African J. Bot. 115: 244-250 [doi | pdf] <- New

[7] Smoke and human evolution, jgpausas.blogs.uv.es/2016/08/31/