Manual para la medición de caracteres funcionales de plantas

January 4th, 2017 No comments

En 2003 se publicó el  primer manual para la medición estandarizada de caracteres funcionales de plantas [1], y en 2013 se realizó una segunda versión más completa y actualizada [2]. Estas dos versiones se publicaron en inglés en la revista Australian Journal of Botany. Ahora, la misma revista, publica la traducción de la segunda versión (2013) en español y la pone disponible a todo el mundo.

Versión en inglés: Handbook  y  supplementay material  [2]

Versión en español: Manual  y  material suplementario  [3]

Original en:  Australian Journal of Botany


Emas2009Fotografía: Midiendo caracteres funcionales en plantas de la sabana brasileña (cerrdao) [4]



[1] Cornelissen, J.H.C., Lavorel, S., Garnier, E., Díaz, S., Buchmann, N., Gurvich, D.E., Reich, P.B., Ter Steege, H., Morgan, H.D., van der Heijden, M.G.A., Pausas, J.G. & Poorter, H. 2003. Handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot. 51: 335-380. [doipdf | CSIRO pub]

[2] 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, Quetier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, Sack L, Blonder B, Poschlod P, Vaieretti V, Conti G, Staver AC, Aquino S, Cornelissen JHC. 2013. New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany 61(3): 167-234. [doi | pdf | Suppl. Mat.]

[3] 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, Quetier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, Sack L, Blonder B, Poschlod P, Vaieretti V, Conti G, Staver AC, Aquino S, Cornelissen JHC. 2013. Nuevo manual para la medición estandarizada de caracteres funcionales de plantas. Australian Journal of Botany 61(3): 167-234. [doi | pdf | Mat. Supl.]

[4] Dantas V., Batalha MA & Pausas JG. 2013. Fire drives functional thresholds on the savanna-forest transition. Ecology 94:2454-2463. [doi | pdf | appendix]

Scale mismatch in ecology

January 2nd, 2017 No comments

A recent paper suggested that fire-vegetation feedback processes may be unnecessary to explain tree cover patterns in tropical ecosystems and that climate-fire determinism is an alternative possibility [1]. This conclusion was based on the fact that it is possible to reproduce observed broad scale patterns in tropical regions (e.g., a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire-vegetation feedback processes. We argue that this reasoning is misleading because these two mechanisms (feedbacks vs fire-climate control) operate at different spatial and temporal scales [2]. It is not possible to evaluate the role of a process acting at fine scales (e.g., fire-vegetation feedbacks) using a model designed for reproducing regional-scale pattern; i.e., there is a mismatch between the scale of the question and the scale of the approach for addressing the question. While the distribution of forest and savannas are partially determined by climate, the most parsimonious explanation for their environmental overlaps (as alternative states) is the existence of feedback processes [3,4], as has been shown in many ecosystems, not only tropical ones [4]. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire-vegetation feedbacks are complementary processes acting at different spatial and temporal scales [2].
Figure: Fire activity (based on remotely sensed data) for savannas and forests located in the range of environmental conditions where both occurs, for Africa and South America (Afrotropics and Neotropics, respectively). From [2,3].

[1] Good, P., Harper, A., Meesters, A., Robertson, E. & Betts, R. (2016) Are strong fire–vegetation feedbacks needed to explain the spatial distribution of tropical tree cover? Global Ecol. and Biogeogr. 25, 16-25.

[2] Pausas J.G. & Dantas V.L. 2017. Scale matters: Fire-vegetation feedbacks are needed to explain tropical tree cover at the local sacle. Global Ecol. and Biogeogr. [doiwiley | pdf]

[3] Dantas V.L., Hirota M., Oliveira R.S., Pausas J.G. 2016. Disturbance maintains alternative biome states. Ecology Letters 19: 12-19. [doi | wiley | pdf | suppl | blog]

[4] Pausas, J.G. 2015. Alternative fire-driven vegetation states. J. Veget. Sci. 26:4-6. [doi | pdf | suppl.] | blog]


Summary 2016

December 22nd, 2016 No comments

The ecology of bark thickness (2): another twist

December 3rd, 2016 No comments

Sometime ago we proposed that "at the global scale, a significant proportion of the variability in bark thickness is explained by the variability in fire regimes", and specifically predicted that frequent low intensity fires select for thick bark [1]. In addition, we suggested that differentiating between inner and outer bark thickness would help to better understand the functional role of bark, especially in non-fire prone ecosystems. The paucity of available data at a global scale limited an empirical demonstration of the proposed framework.

A recent paper has now provided evidence for the fire hypothesis of bark thickness at a global scale [2, 3]. Specifically, Rosell [2] regressed bark thickness against fire frequency and climate parameters and showed that the most sensitive part of the bark in relation to fire was the outer bark, while the inner bark was quite variable and slightly related to both fire and climate [2]. In the early paper [1] we also mentioned that little was known about the role of bark thickness in arid ecosystems. Recent research support the role of bark as a fire protection mechanisms in some arid ecosystems [4, 5].

To advance in the relationship between bark thickness and fire, it is necessarily to consider not only fire frequency, but also fire intensity, and to scale these fire characteristics with plant life-histories ([3], see figure below). This is because the relationship between fire regime and bark thickness is not expected to be simple and linear, but a bit more complex, including some threshold-type relationships (figure below).

Little by little we are improving our understanding on the role of bark as a fire-protection mechanism, and how fire regimes has shaped bark thickness in many ecosystems.


Figure: Bark thickness as a function of fire regime: flame height (an indicator of fire intensity) and mean fire return interval (fire frequency). Fire regime is scaled by the characteristics of the plant (height to the base of the crown and longevity, respectively). The shaded area represents the areas where thick bark is adaptive for fire protection, i.e., when return intervals are shorter than the lifespan of the plant and fires are of low intensity (flame height is shorter than the distance to the base of the crown, e.g., surface fires); the shade area is limited thresholds (values of 1 in the axes). The unshaded area represents the conditions where thick barks are not adaptive (thin bark is more likely), i.e., when fires are crown-fires or when the return interval is long (in relation to the longevity of the plant). From [3].


[1] Pausas, J.G. 2015. Bark thickness and fire regime. Functional Ecology 29:317-327. [doi | pdf | suppl. | blog]

[2] Rosell J.A. 2016. Bark thickness across the angiosperms: more than just fire. New Phytologist 211: 90–102

[3 ] Pausas J.G. 2017. Bark thickness and fire regime: another twist. New Phytologist 213: 13-15. [doi| pdf] <- New!

[4] Schubert, A. T., Nano, C. E. M., Clarke, P. J. & Lawes, M. J. 2016. Evidence for bark thickness as a fire-resistance trait from desert to savanna in fire-prone inland Australia. Plant Ecol. 217: 683-696.

[5] Cousins, S. R., Witkowski, E. T. F. & Pfab, M. F. 2016. Beating the blaze: Fire survival in the fan aloe (Kumara plicatilis), a succulent monocotyledonous tree endemic to the Cape fynbos, South Africa. Austral Ecol. 41:466-479.

La huella del fuego

November 30th, 2016 1 comment

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 aquí:

También en

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.

Flammability strategies

November 24th, 2016 No comments

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]

[1] The-fire-overview-effect,

[2]  A new global fire map,   [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. [doi | wiley | pdf]


Future fires

November 11th, 2016 No comments

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].

[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]


Smoke-stimulated germination (2): Shedding light through the smoke

November 1st, 2016 No comments

There are some plants with seeds that have a dormancy period and that fire can stimulate their 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].

smoke-germinationFigure: 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].


[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,

[5] Keeley J.E. & Pausas J.G. (in press). Evolution of 'smoke' induced seed germination in pyroendemic plants. South African J. Bot. [doi | pdf] <- New

[6] Moreira B. & Pausas J.G. (in press). Shedding light through the smoke on the germination of Mediterranean Basin flora. South African J. Bot. [doi | pdf] <- New

[7] Smoke and human evolution,

De incendios y cipreses (5)

October 11th, 2016 2 comments

Después de una serie de despropósitos sobre el posible uso de cipreses ignífugos [1-4], por fin parece que se encaucen las cosas: Los cipreses que estaban destinados para hacer de barrera cortafuegos en el monte, parece que finalmente se utilizarán en jardinería [5], y esperemos que para jardines urbanos, lejos del monte. En paisajes con incendios recurrentes, plantar cipreses en zonas semi-urbanas (en la interfaz urbano-forestal), no es recomendable, ya que si llega el fuego, o simplemente pavesas, pueden prender de manera intensa y actuar como antorchas. Por ello, los bomberos temen las casas rodeadas de cipreses, y de hecho, está prohibido plantarlos en jardines de diversas zonas de EEUU. Hay evidencias de que los cipreses pueden ejercer de captadores de pavesas (foto). La idea de utilizarlos como cortafuegos estaba fuera de toda lógica [4].

Cipreses-quemadosFoto: Valla de cipreses que prendió durante el incendio de La Granadella (4/Sep/2016, La Marina, Alicante). Nótese que el incendio no llegó directamente a la valla (los pinos y campos de cultivo  de los alrededores no se vieron afectados); es probable que el fuego llegase con una pavesa, como pasó con los distintos focos de este mismo incendio [6].


[1] De incendios y cipreses (1), 29/9/2012
[2] De incendios y cipreses (2), 7/10/2012
[3] De incendios y cipreses (3), 22/6/2013
[4] De incendios y cipreses (4), 31/8/2015

[5] La investigación española sobre cipreses cortafuegos acabará en plantas de jardín,

[6] El SEPRONA concluye que todos los focos del incendio de la Granadella fueron provocados por las pavesas (;  Una colilla mal apagada provocó el incendio de Xàbia (; El Seprona cree que una colilla originó el incendio y el viento causó los tres focos (

¿Será este el último post sobre el tema? ¿Se habrá ganado una pequeña batalla?
(podéis dejar vuestra opinión en los comentarios)


Postfire flowering: Lapiedra martinezii

October 8th, 2016 No comments

Lapiadra martinezzi
Lapiadra martinezzi (Amaryllidaceae) flowering after fire in eastern Spain. This is also an example of an hysteranthous geophyte (flowering before appearing the leaves).

Upper left: From La Granadella (Benitatxell, La Marina Alta, Alicante), one month after a high intensity wildfire that occurred the 5 Sept 2016.
All others: in a Pinus halepensis open woodland that was burned (at low intensity) in April 2016 (for firefighting training) near Valencia; photo taken the 29th Sept 2016. There were many individuals (hundreds to thousands) flowering and some with fruits. We did not find any flower in the surrounding unburned area.

For other species with fire-stimulated flowering, see:


Fire benefits plants by disrupting antagonistic interactions

October 2nd, 2016 2 comments

There are many plants that benefit from fire. Typical examples are those that despite they may be killed by fire, the germination of their seeds is stimulated by the fire (either by the heat or by the smoke; [1,2]), and thus they recruit very well (high offspring abundance) and often increase there population size postfire. Species with fire-stimulated flowering [3,4] also benefit from fire. In a recent paper [5] we propose that there may be another mechanisms by which fire may benefit plants: fire may remove seed predators, and thus create a window of opportunity for reproduction under a lower predation pressure (predator release hypothesis). This is specially applicable to specialist plant-insect interactions. We documented two cases: in Ulex parviflorus, a plant species with fire-stimulated germination [1,2], fire eliminated there specialist seed predator weevil (Exapion fasciolatum, Apioninae, Brentidae) and thus increased the available seed number for germination. Similarly, in Asphodelus ramosus, a fire-stimulated flowering species [3], fire reduced the specialist herbivore and seed predator (Horistus orientalis, Miridae, Hemiptera) and increased their fruit production. Thus, fire, by disrupting the antagonistic interactions, benefit plants; the temporal window of this predator release is likely to depend on fire size. For more information see reference [5].


Figure: Proportion of predated fruits of Ulex parviflorus in unburned sites (grey boxes) and at the edge and center of a recently burned area (white boxes), 2 and 3 years postfire. Data from two large wildfires in Valencia (2012) [5]; Edge and Center of the burned area refer to <1 km and >1.5 km from the fire perimeter, respectively. Photo of the seed predator (Exapion) from


[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. Annals of Botany 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] Postfire blooming of Asphodelous,

[4] Postfire flowering: Narcissus,

[5] García Y., Castellanos M.C. & Pausas J.G. 2016. Fires can benefit plants by disrupting antagonistic interactions. Oecologia 182: 1165–1173. [doi | pdf] <- New!!


The fire overview effect

September 18th, 2016 No comments

The overview effect is the feeling and awareness reported by some astronauts when viewing the entire Earth during space-flight. Fire ecologists have our own overview effect! When remote sensed fire information was available for the first time at the global scale, it provided a magnificent and unprecedented view of the importance of fires on the Earth, and fires become a global issue. This remotely sensed information was a very valuable data because, for the first time, it was possible study some fire ecology processes at the global scale (for example [1]). Here is an animation for a 10 years period (2000-2010). It shows that on our planet, fires are widespread and something is always burning; we live in a flammable planet.


MODIS Rapid Response System Global Fire Maps, NASA. Each colored dot indicates a location where MODIS detected at least one fire during a 10-day period.

More global fire animations: youtube | Earth Observatory |


[1] Pausas J.G. & Ribeiro E. 2013. The global fire-productivity relationship. Global Ecol. & Biogeogr. 22: 728-736. [doi | pdf | appendix | erratum | blog]


Smoke and human evolution

August 31st, 2016 1 comment

In this blog we have discussed that some plants have evolved seeds with sensitivity to chemicals produced by fire in such a way that these chemicals stimulate the germination of the plants after a fire; we call this process smoke-stimulated germination [1-3]. Well, plants are not the only organisms that have evolved in response to chemicals present in the smoke, humans too! A recent paper show that modern humans are the only primates (including early hominids as Nearthentals and Denisovans) that carry a mutation increasing tolerance to smoke chemicals produced by fires [4]. This mutation could have given an evolutionary advantage to modern humans in relation to other hominids as allowed them to use fire for many important activities (e.g., cooking, hunting, defense, heating, agriculture). This high exposure to smoke would have also increased the susceptibility to pulmonary infections, and even the evolution of some of them (tuberculosis [5]). The tolerance to smoke also allowed modern humans to have some tolerance to pollution and to smoke cigarettes! That is, the ability to smoke could be a side effect (an exaptation, if you'd like) of been adapted to use fire, and in fact, it currently acts as a secondary sexual character!

Smoking as a secondary sexual character (Woody Allen in Manhattan, 1979).

[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. Annals of Botany 105: 627-635. [doi | pdf | post]

[2] Tormo, J., B. Moreira, and J. G. Pausas. 2014. Field evidence of smoke-stimulated seedling emergence and establishment in Mediterranean Basin flora. Journal of Vegetation Science 25: 771-777. [doi | wiley | pdf | post]

[3] Smoke-stimulated germination,

[4] Hubbard, T.D., Murray, I.A., Bisson, W.H., Sullivan, A.P., Sebastian, A., Perry, G.H., Jablonski, N.G. & Perdew, G.H. (2016) Divergent Ah receptor ligand selectivity during hominin evolution. Mol. Biol. Evol., 33:2648-2658.

[5] Chisholm, R.H., Trauer, J.M., Curnoe, D. & Tanaka, M.M. (2016). Controlled fire use in early humans might have triggered the evolutionary emergence of tuberculosis. Proc. Natl. Acad. Sci. USA, 113, 9051-9056.

Fire ecology in Plant Ecology: homage to Peter Clarke

July 8th, 2016 No comments

The journal Plant Ecology has now published an special issue on Fire Ecology to homage Peter Clarke, who died in December 2014 after a long battle with cancer. Peter (University of New England, Australia) made a significant contribution to the fire ecology of Australia; many of his colleagues and collaborators have contributed to this issue (including myself). Link to the special issue.

Photo: Peter Clarke (center; arms crossed) and colleagues visiting the Otay Mountains (San Diego, southern California) in November 2006 after attending the 3rd International Fire Ecology and Management Congress in San Diego. From Left to right: Malcom Gill, Dylan Schwilk, Ross Bradstock, Peter Clarke, William Bond, and Juli Pausas. Photo by Jon Keeley.

Diversidad política (3)

July 2nd, 2016 No comments

Tras las Elecciones Generales del 20 de Diciembre de 2015, la diversidad política del Congreso de los Diputados de España incrementó considerablemente, y fue la más elevada de la historia de nuestra democracia; y ya explicamos las ventajas de tener un congreso con elevada diversidad (elecciones 2008,  elecciones 2015). La desventaja es que la elevada diversidad es más difícil de gestionar, especialmente en sociedades poco maduras dramáticamente (lo más simple de gestionar son las dictaduras, que tienen una diversidad muy baja). La composición del Congreso que salió de esas elecciones no supo gestionar esa diversidad, seguramente por la falta de experiencia democrática, es decir, falta de capacidad de diálogo, de capacidad para aceptar las diferencias, y de excesivo deseo de poder de los lideres. Ello nos llevó a repetir las elecciones 6 meses más tarde.

Las Elecciones Generales del 26 de Junio 2016 dieron un resultado en parte parecido a las anteriores, pero con un poco menos de diversidad (Figura). ¿Se ha reducido suficiente la diversidad para que los políticos sean capaces de ponerse de acuerdo y formar un gobierno?

Figura: Valores de diversidad (expresada por el índice de diversidad de Shannon-Weaver) según el número de votos (rojo) o de escaños (azul) en los diferentes partidos políticos, en las diferentes Elecciones Generales al Congreso de los Diputados de España realizadas durante la democracia (elaborado a partir de la base histórica de resultados electorales del Ministerio del Interior).

Dado que sólo han transcurrido 6 meses entre las dos elecciones, es tentador analizar las causas de las diferencias, al menos por lo que hace a los grandes partidos y los grandes números (Tabla):

  • Se redujo un poco el número de partidos. La ley electoral española perjudica a los partidos pequeños de ámbito estatal, y eso forzó ha hacer algunas uniones antes de las elecciones, en lugar de hacerlas después (cosa que permitiría que el programa conjunto se realizase según los votos de cada uno de los partidos y así reflejaría mejor el deseo de los votantes).
  • El PP aumentó en votos, probablemente a expensas de C's y de otros partidos menores, que disminuyeron en votos (Tabla, números en azul; este cambio aumenta la dominancia y disminuye la diversidad). Quizá los votantes de derechas castigaron el pacto Cs-PSOE y asumieron la corrupción como una mal menor (cosa bastante sorprendente en un país que desea ser moderno y democrático)
  • La abstención aumentó en más de 1 millón, y probablemente se concentró en las confluencias de izquierda, que perdieron aprox. 1 millón de votos, y en menor medida en el PSOE (Tabla, números en rojo). Parece que los que decidieron no votar en el 26J después de haber votado en 20D eran gente de izquierdas (desencantados, o votantes de IU que no aceptaron la coalición con Podemos). Si eso es cierto, no deja de ser curioso que haya gente de izquierdas que le de igual el partido que gobierne.

Tabla: Comparación (en número de votos y abstenciones) entre las elecciones generales al Congreso de los Diputados del 20D del 2015 y las del 26J del 2016.  Datos según el Ministerio del Interior (29 Junio 2016). [1] Incluye a Podemos, IU y las confluencias afines (Compromís, Mareas, Equo, Podem, …). [2] Opciones que no cuentan para la formación del Congreso.


Conclusiones 26J 2016:

1) A muchísimos españoles (unos 8 millones) les parece bien y reprueban la corrupción, los grandes recortes en sanidad y educación, la reforma laboral y la ley mordaza, entre otras. Consideran que es lo mejor para el país, o quizá, que más vale malo conocido, que bueno por conocer.

2) Una coalición muy reciente, dominada por un partido muy nuevo (Podemos) se mantiene con mas de 5 millones de votos, indicando las ansias de cambios importantes de una buena parte del electorado.

3) Muchísimos españoles (unos 11 millones) no opinaron acerca de cómo les gustaría que se organizase el país, y parce que les da igual cómo se repartan los impuestos que pagan.

4) El hecho de que los resultados sean bastante diferentes a lo previsto por las encuestas sugiere una cierta respuesta a estas encuestas, y por lo tanto, una cierta inmadurez política.

5) Aun no se sabe si se conseguirá formar gobierno, pero el Congreso salido de estas elecciones parece que representará peor a la población (menos votos) que el que se formó en el 2015, y especialmente a la población con ideas de izquierda (una parte no votó). En cualquier caso, será un Congreso mucho más diverso que el de las anteriores legislaturas (ver linea azul en la figura de arriba); cosa que es positiva para todos.

Todo ello asumiendo que no hay fraude en el recuento de votos; la falta de transparencia ha despertado algunas alarmas (p.e., lagranjahumana).

Diversidad política (2).
Diversidad política (1): pérdida de diversidad política en España.

Brazil 2016

June 27th, 2016 No comments

Summary of my June 2016 tour:

  • Brasilia: meeting point with S. Paula;  visit to the Jadim Botánico and the Chapada Imperial [photos A, E]
  • Pirenópolos: IAVS 2016; talk: “Plant strategies in fire-prone ecosystems: hidden buds” [photo B]; Pireneus
  • Rio Claro: Univesidade Estadual Paulista (UNESP); meeting with A. Fidelis & students (Luis, Talita, etc.). Visiting the Itirapina experimental fires [photo C]. Talk: “Fire, traits, and biodiversity: a global perspective”.
  • Piracicaba: Universidade de Sao Paulo (USP), meeting with B. Appezzato-da-Glória and collaborators to study underground resprouting structures [photo D]
  • Campinas: Universidade Estadual de Campinas (UNICAMP); meeting with V. Dantas and students (Paulo, André, etc.) [photo F]
  • Sao Paulo: meeting W. Delitti (USP)

Photos: A (top left):  Susana Paula, Ericaulaceae and myself; B (top right) : my talk in Pirenópolis; C (bottom left): A. Fidelis and her students in front of an 'underground tree'. D (center): Xylopodium with tuberous roots in Aldama (Appezzato-da-Glória lab); E (middle right): tree with a corky bark in the cerrado of the Jardim Botánico (Brasilia); E (bottom right): Having a drink with Vinicius Dantas in Campinas.

Related posts:

- Brazil 2015, 16 Mar 2015

- Fire shapes savanna-forest mosaics in the Brazilian cerrado, 14 May 2014

- Afrotropical and neotropical savannas are different, 29 Jul 2013

- Fire generates intraspecific trait variability in neotropical savannas, 28 Aug 2012

- Disturbance maintains alternative biome states, 9 Nov 2015


Fire behaviour by Vareschi

May 13th, 2016 No comments

Recently I came across this figure published in 1962 by Volkmar Vareschi [1] which nicely synthesize variations in temperatures in the flame and in the soil, as well as flame height and flame spread (time and distance) in a simple hand-drawing. It is not easy to see a figure on fire behaviour as simple and illustrative as this one; I only miss a bit of colour. It refers to a burn of a Trachypogon savanna in Los Llanos, Venezuela. Vareschi (1906-1991) was born in Austria and moved to Venezuela in 1950; he is considered a pioneer in tropical plant ecology; one of his papers was about savanna fires [1].


Vareschi-1962-burnFigure 2 from [1]



[1] Vareschi, V. (1962) La quema como factor ecológico en los Llanos. Boletin de la Sociedad Venezolana de Ciencias Naturales 23, 9-31.

Objeción a los gastos militares (renta 2015)

May 7th, 2016 No comments
ACTUALIZACIÓN: Hace unos días explicaba aquí la manera de hacer la objeción a los gastos militares en la declaración de la renta del 2015, basándome en la experiencia de los otros años. Pero como este año la Agencia Tributaria ha realizado algunas modificaciones en el procedimiento, ahora he modificado el texto. Disculpad las molestias.


Como cada año, en estas fechas los españoles realizamos la declaración de la renta. Y esta es una buena oportunidad para quejarse de los excesivos gastos militares que nuestro gobierno realiza; ¡hay muchas otras prioridades!. El año pasado ya expliqué algunos detalles, este voy a ir al grano y explicar los pasos a seguir:
  • Realizar la declaración de la renta de manera normal, ya sea con el programa PADRE o con la nueva aplicación Renta WEB.
  • Una vez finalizada la declaración, ir al subapartado “Retenciones y demás pagos a cuenta” en la página 16(I) (que está hacia el final; se llega fácilmente desde la página de resumen del programa PADRE). En una de las casillas que no utilices, se pone la cantidad a desviar; por ejemplo en la casilla 542 ("Cuotas del impuesto sobre la renta de no residentes") ponéis 100 euros (por ejemplo o lo que se quiera desviar).
  • Si vuestra declaración os salía a pagar (positiva), ahora os saldrá a pagar 100 euros menos (por ejemplo); si os salía a cobrar (negativa), ahora os saldrá a cobrar 100 euros más.
  • Finalizar la declaración de manera normal (si utilizáis PADRE, se finaliza imprimiendo y entregando la declaración a un banco).
  • Ingresar la cantidad desviada (por ejemplo, los 100 euros) a una ONG. Algunas asociaciones que promueven y apoyan la objeción, hacen anualmente sugerencias de posibles ONGs a ingresar, pero se puede realizar en cualquiera.
  • Enviar una carta al registro una oficina de Hacienda (por ejemplo, al Registro General del Ministerio de Hacienda, c/ Alcalá, 9. 28071, Madrid) en la que se explica los motivos  de vuestro desvío a una ONG y adjuntar el justificante del ingreso. Aquí hay un ejemplo de carta:  descargar doc. Se puede incluir también una copia impresa de la declaración donde se tacha el texto de la casilla 542 y se pone (a mano) "Por objeción a los gastos militares".
  • Para las estadísticas, es conveniente avisar que se ha realizado la objeción fiscal, por ejemplo, rellenado el formulario 2016, enviando un mensaje a, o contactando con tu asociación antimilitarista local (por ejemplo, en Valencia: moc-valencia ).

Más información: | | | (cat) | (cat) | jgpausas (Renta2014) | insumissia |


Odena: 9 meses posincendio

May 1st, 2016 No comments

El 27 de Julio de 2015 un incendio forestal afectó unas 1200 ha en Òdena (Anoia, Catalunya central), una zona dominada principalmente por pino carrasco (Pinus halepensis). Pocos días después ya se empezaba a ver un inicio de la regeneración del ecosistema [1, 2]. En una visita reciente (Abril 2016, 9 meses posincendio), vemos que en gran parte de la zona se han cortado y extraído los árboles quemados (y algunos no quemados). Antiguamente, cuando aun no se daba casi ningún valor a los ecosistemas naturales, y sí a la madera, se sacaban los árboles quemados para obtener algún beneficio económico; y algunas veces por motivos "estéticos". Hoy en día, no parece una acción muy apropiada [3], a no ser que haya una razón de peso, cosa que desconozco en el caso de este incendio.

Los árboles quemados benefician a la regeneración porque retienen un poco el suelo, disminuyen el impacto de las gotas de lluvia en el suelo, mantienen cierta humedad, captan agua de la niebla, sirven de posadero para aves que traen semillas (que contribuyen a la regeneración), y son hábitat para fauna diversa [4]. Cortar los árboles requiere entrar con maquinaría en la zona quemada (con suelos muy sensibles), generar caminos y arrastrar troncos. Esto conlleva la eliminación de todos los beneficios mencionados, ademas de la disminución de parte del suelo y mantillo, la mortalidad de las primeras germinaciones posincendio (por ejemplo del pino), la formación de surcos que pueden ser puntos de inicio de erosión (cárcavas), y disminución de la regeneración natural en general. En general, entrar en una zona recién quemada, y degradar el ecosistema disminuyendo la regeneración y aumentando la erosión, está poco justificado [3]; en algunos casos, estas intervenciones pueden ser más perjudiciales que el propio incendio.

Fotos: a) Pinar con rebrotes de madroño 4 meses después del incendio, antes de cortar los árboles; se aprecia un cierto ambiente forestal. b) surcos del arrastre de troncos durante la extracción de la madera quemada. c) Ambiente 9 meses después del incendio, una vez se han cortado los árboles. d) Germinación de pino 4 meses después del incendio; germinaciones susceptibles a ser eliminadas si se entra con maquinaria o se arrastran troncos. e) pinos vivos (no quemados) cortados y apilados (9 meses posincendio). f) Enebro rojo (Juniperus oxycedrus) que rebrota tras quema y corta (9 meses posincendio). Incendio de Odena, Abril 2016 (fotos: JG Pausas).


[1] Odena fire: first visitors, 10-08-2015

[2] Odena fire: 55 days postfire, 17-10-2015

[3] Lo que no se debe hacer después de un incendio, 13-08-2015

[4] Pausas, J.G., Ribeiro, E. & Vallejo, R. 2004. Post-fire regeneration variability of Pinus halepensis in the eastern Iberian Peninsula. Forest Ecology and Management 203: 251-259. [doi | pdf]

Flammable Mexico

April 13th, 2016 No comments

Mexico is a megadiverse North American country with a wide range of climates (e.g., wet tropical, warm temperate, mediterranean, and arid) and a diverse topography (from sea level up to 5700 m asl). These characteristics together with its location in the transition toward Central America make this land a global biodiversity hotspot with species belonging to northern (Neartic) lineages co-occurring with others from southern (Neotropical) lineages. An important factor contributing to this biodiversity are the frequent disturbances in this region where volcanoes, hurricanes, and wildfires are common, together with droughts and floods. Fires occur mainly in April-May (Figure below, [1]); the natural sources of ignition being lightning, especially in mountains, and volcanoes (with clear evidence of fires ignited by volcanoes, e.g. from the Popocatépetl volcano). However, currently most fires are caused by human activities, as in many other countries. Hurricanes add fuel and increase the intensity and probability of fire [2].

Mexico is a center of diversification of pines (Pinus) and oaks (Quercus), two species groups strongly related to fire [3,4]. Mexico harbors about 50 species of pines and these incorporate all the fire strategies and traits observed in this genus [4]. For instance, there are many fire tolerant pines with thick barks, self-pruning abilities, and in some cases, with basal or juvenile resprouting capabilities; fire embracers (postfire seeders) with thin bark and serotinous cones; and fire avoiders that lack these traits. The country also harbors some 160 Quercus species, ranging from strongly resprouting shrubby species to many tree oaks with relatively thick bark that live in surface fire ecosystems, and including evergreen and drought-deciduous species with a large range of leaf morphologies. I was surprised to see some oaks with very large, and very thick leathery leaves that are deciduous, certainly an outlier in the leaf economics spectrum. More details in [1].

Figure: Recent fire activity in Mexico (2001-2015) estimated from the monthly number of active fires recorded by the Terra satellite (MODIS hotspots). Top: temporal variability (x-axis ticks indicate the begining of the year). Bottom left: fire seasonality– the flammable season is concentrated into four months (March-June), with a peak in April-May (the end of the dry season). Bottom right: proportion of active fires in each biome (TrDry: tropical dry broadleaf forests; TrConif: tropical coniferous forests: TrMost: tropical moist broadleaf forests; Desert: deserts and xeric shrublands; Others). From [1]

[1] Pausas J.G. Flammable Mexico. Int. J. Wildland Fire [doi | pdf]

[2] Fire-wind interactions, 30 Oct 2015

[3] He T, Pausas JG, Belcher CM, Schwilk DW, Lamont BB. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytol. 194: 751-759. [doi | wiley | pdf | suppl.]

[4] Pausas, J.G. 2015. Evolutionary fire ecology: lessons learned from pines. Trends Plant Sci. 20: 318-324. [doi | sciencedirectpdf]

Postfire resprouting of Chamaerops humilis

March 18th, 2016 No comments

"A few, but only a few species of palms, are, like our Coniferae, Quercineae, and Betulineae, social plants : such are the Mauritia flexuosa, and two species of Chamaerops, one of which, the Chamaerops humilis, occupies extensive tracts of the ground near the Mouth of Ebro and in Valencia ..." -- Alexander von Humboldt (1848)

Chamaerops humilis (Mediterranean dwarf palm) is the only native palm in continental Europe, and the northernmost naturally occurring palm in the world. It is native to the western Mediterranean Basin, occurring along the Mediterranean cost of Spain (as mentioned by Humboldt), Portugal, France, Italy, Malta, Morocco, Algeria, and Tunisia. The other palm occurring in the Mediterranean Basin is Phoenix theophrasti, a rare palm growing in the Crete island and in the southern Turkey [MedTrees].

Humboldt probably did not know that Chamaerops humilis resprouts very quick after fire (at that time fire was not considered as part of the natural processes). The resprouting of this species does not necessary come from new dormant buds (as in most typical resprouters) but from the normal apical buds protected from the fire by the leaf bases in the stem. In fact, buds generate leaves that have the upper part affected by the fire, but not the lower part (as in all monocots, the meristem is at the base of the leaves, and thus more protected from the heat of the fire). Consequently the first leaves often show the typical burned-brown-green pattern of the photo below. In addition, it can generate basal suckers from an underground rhizome. C. humilis often flowers very quickly after fire, together with the first leaves (upper photo). Overall it is very resilient to recurrent fires.

Chamaerops humilis (one of the few 'social palms' following Humboldt) 2-3 months postfire in the Valencia region (eastern Spain; photos: JG Pausas)


Humboldt, A. von (1848). Aspects of nature (original title: Ansichten der Natur, 3rd ed).


Incendios forestales (de vegetación) en México

March 1st, 2016 1 comment

Recientemente se ha publicado un nuevo libro sobre incendios forestales. El libro describe la importancia y los regímenes de incendios en los diferentes ecosistemas de México, así como aspectos sobre historia y manejo del fuego [1]. Es un libro extenso, enciclopédico, con unas 1700 páginas publicadas en 2 volúmenes. El primer volumen (18 capítulos) representa un paseo por la gran variedad de paisajes de México (pinares, bosques de encinos, de oyamel, de galería, pastizales, matorrales, selvas, bosques mesófilos de montaña, sabanas, manglares, palmares, etc.), y en cada uno de ellos se explica el régimen de fuego y las respuesta de las especies y ecosistemas. El segundo volumen (11 capítulos) aborda cuestiones de comportamiento, prevención, y combate del fuego, así como una historia del fuego desde sus inicios (con los primeros ecosistemas terrestres [2]) hasta el uso del fuego en las culturas mesoamericanas y en el México actual. Si hay algo que se encuentra a faltar sería una visión evolutiva; quizá los lectores pueden encontrar esas visión en mi propio libro [3]. Más comentarios sobre el libro y sobre México en [6].

Dante-Rodriguez-Trejo_libroPortadas del volumen 1 (izquierda) y 2 (derecha)

A veces se ha considerado que el concepto de incendios forestales se refiere sólo a los incendios que ocurren en bosques, o incluso a incendios que ocurren en plantaciones forestales. Para evitar ese mal entendido, el autor titula el libro 'incendios de vegetación', ese decir, para enfatizar que el libro se refiere a incendios en cualquier tipo de vegetación. En España, actualmente el término 'forestal' se refiere a cualquier tipo de vegetación natural (también llamado monte), y en ese contexto es cómo a menudo usamos el concepto de incendios forestales [3]. Así es como lo explicaba yo en el prólogo del libro 'Incendios forestales':

"Los incendios son fuegos que se propagan sin control humano; cuando ocurren en la naturaleza se llaman incendios forestales. El término forestal está relacionado con una clasificación tradicional de los usos del suelo, donde el uso forestal incluye cualquier zona terrestre que no sea de uso urbano ni agrícola. De este modo, el término incendios forestales se refiere a los fuegos no controlados (sean de origen natural o antrópico) que ocurren en los ecosistemas terrestres, y que se propagan por la vegetación, sea ésta del tipo que sea (bosque, sabana, matorral, pastizal, humedal, turbera, etc.). Por lo tanto, incendios forestales, y por extensión este libro, no sólo hace referencia a bosques, como a veces se ha interpretado, sino a cualquier tipo de ecosistema terrestre" [3]

En inglés también hay una amplia nomenclatura para referirse a los incendios forestales, como por ejemplo, wildfires (principalmente utilizado en Norte América), forest fires (Europa), bush fires (Australia), vegetation fires, landscape fires, etc., todos ellos describen los incendios en cualquier tipo de vegetación.


[1] Rodríguez Trejo, D. A. 2015. Incendios de vegetación. Su ecología, manejo e historia. 2 vol. Ed. Colegio de Postgraduados, Biblioteca Básica de Agricultura, México.

[2] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doijstor | pdfpost]

[3] Pausas J.G. 2012. Incendios forestales. Una visión desde la ecología. Ed Catarata-CSIC. Madrid. [libro]

[4] He T., Pausas J.G., Belcher C.M., Schwilk D.W., Lamont B.B. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytol. 194: 751-759. [doi | wiley | pdf | suppl.]

[5] Pausas, J.G. 2015. Evolutionary fire ecology: lessons learned from pines. Trends Plant Sci. 20: 318-324. [doi | sciencedirect | cell | pdf]

[6] Pausas J.G. 2016. Flammable Mexico. Int. J. Wildland Fire [doi | pdf]


Olive trees resprouting

February 22nd, 2016 No comments

The typical image on a cultivated olive tree (Olea europaea) is a short squat tree with a thick gnarled trunk. Below are some olive trees with a slightly different shape, after being burned twice in different wildfires (1994 and 7/2015) in Montán (Castelló, eastern Spain). Before 1994 these trees were single-stemmed with the typical thick trunk; they were planted long ago for olive production. The 1994 fire killed the main stem and the tree produced many resprout from the base, around the trunk (it became multi-stemmed). In 2015 in burned again killing those 21 year-old resprouts and producing many new ones (the green ones in the pictures, 7 month-old resprouts). The 2015 fire also consumed the main stem that had died in the 1994 fire, including the base of the stem, and thus it produced a hole in the middle of the tree (second picture). This is quite common.

Olea resprouting 1
Olea resprouting 2
Photos: Olive trees (Olea europaea) resprouting after two fires (1994, 7/2015; JG Pausas 2/2016).

More on resprouting: Lignotubers | Resprouting at the global scaleEvolutionary ecology of resprouting and seedingPhysiological differences between resprouters and seedersTo resprout or not to resprout | Differences between resprouters and non-resprouters | Fire, drought, resprouting: leaf and root traits |


Convivir con el fuego: Decálogo de incendios forestales

February 3rd, 2016 No comments

Hace ya unos años escribimos un decálogo donde proponíamos unas bases ecológicas para convivir con los incendios forestales [1]. Ahora, la Fundación Pau Costa, en el marco de su 5º aniversario, ha compilado otro decálogo [2], este más amplio en temática y con muchos más autores, pero con un objetivo similar, aprender a convivir con el fuego. Los interesados en apoyar el decálogo tienen la posibilidad de hacerlo firmando el formulario que hay al final del mismo.

Foto: Quema experimental en Ayora (Valencia, 4/2009) realizada con la finalidad de entender el efecto de los incendios en los ecosistemas mediterráneos.


[1] Pausas J.G. & Vallejo R. 2008. Bases ecológicas para convivir con los incendios forestales en la Región Mediterránea - decálogo. Ecosistemas 17(2):128-129, 5/2008. [enlace | pdf]

[2] Decálogo de incendios forestales, Pau Costa Fondation, [enlace | pdf]

[3] Pausas, J.G. 2012. Incendios forestales. Catarata-CSIC. [Libro]

[4] 'Conviure amb el foc', entrevista en El Temps, 24 Julio 2012 [pdf]

[5] Otros textos de divulgación sobre incendios y ecología [divulgación]

[6] Towards prescribed fires,, 7 Oct 2013


Fire in the root of humans (2)

January 16th, 2016 No comments

Many people have the idea that fires scare animals and fled them in panic. However this is not always true, some species react still and calm and move away to safe sites. Some time ago I mentioned a study demonstrating that chimps in wild, when they see a wildfire, they react calmly, predict their behaviour and move accordantly without any stress or fear, suggesting that they have some understanding of fire behaviour [1]. Few days ago I came across other studies [2,3] suggesting that different species of primates not only react calmly to fire but after a fire, they increase their home range to include the area burned and used it for searching food, including 'cooked' fruits! So wildfires were very important in the history of humans [4], they could have contribute to the first step towards humanity from our ancestors ...

Figure: In captivity, some apes are able to light a fire a roast vegetables (see youtube1, youtube2). Photo from

[1] Fire in the root of humans, 19-1-2010.

[2] Jaffe KE, Isbell LA 2009. After the fire: benefits of reduced ground cover for vervet monkeys (Cercopithecus aethiops). Am. J. Primatol. 71:252-260.

[3] Herzog NM, Parker CH, Keefe ER, Coxworth J, Barrett A, Hawkes K 2014. Fire and home range expansion: A behavioral response to burning among savanna dwelling vervet monkeys (Chlorocebus aethiops). Am. J. Phys. Anthropol. 154:554-560.

[4] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doijstor | BioOne | pdfpost]


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