Posts Tagged ‘Mediterranean’

Fire-released seed dormancy

April 8th, 2022 No comments

Many plants concentrate their seedling recruitment after the passage of a fire. This is because postfire conditions are especially optimal for germination and establishment of many species as fires create extensive vegetation gaps that have high resource availability, minimal competition, and low pathogen load. Thus we propose that fireprone ecosystems create ideal conditions for the selection of seed dormancy as fire provides a mechanism for dormancy release and optimal conditions for germination [1]. We compiled data from a wide range of fire-related germination experiments for species in different ecosystems across the globe and identified four dormancy syndromes: heat-released (physical) dormancy, smoke-released (physiological) dormancy, non-fire-released dormancy, and non-dormancy. In fireprone ecosystems, fire, in the form of heat and/or chemical by-products (collectively termed ‘smoke’), are the predominant stimuli for dormancy release and subsequent germination, with climate (cold or warm stratification) and light sometimes playing important secondary roles. Fire (heat or smoke)-released dormancy is best expressed where woody vegetation is dense and fires are intense, i.e. in crown-fire ecosystems (e.g., mediterranean-type ecosystems). In grassy fireprone ecosystems (e.g. savannas), where fires are less intense but more frequent, seed dormancy is less common and dormancy release is often not directly related to fire (non-fire-released dormancy). Fire-released dormancy is rare to absent in arid ecosystems and rainforests. Heat-released dormancy can be traced back to fireprone floras in the ‘fiery’ mid-Cretaceous, followed by smoke-released dormancy, with loss of fire-related dormancy among recent events associated with the advent of open savannas and non-fireprone habitats. Anthropogenic influences are now modifying dormancy-release mechanisms, usually decreasing the role of fire. We conclude that contrasting fire regimes are a key driver of the evolution and maintenance of diverse seed dormancy types in many of the world’s natural ecosystems.

Fig. 1. Percentage germination of 68 populations or species subjected to simulated fire- (y axis) and summer-type (warm stratification) temperature (x-axis) (C., Cistus; F., Fumana; U., Ulex; A., Acacia; M., Mimosa). Points above the dotted line (1:1) have higher germination levels after fire heat than after summer heat. Note that all points at or below the line are for species in savannas [S], while the others are from mediterranean shrublands and other crown-fire ecosystems. That is, in crown-fire ecosystems, fire is the most likely selective agent for dormancy. From [1].
Fig. 2. Dated phylogeny for major clades in the New and Old World Cistaceae together with closely related ancestral clades. Pie charts at the tips show the fraction of species that occur in crown-fire ecosystems (red), surface-fire ecosystems (orange), those with physical dormancy – hard seeds (green), and those with heat-released dormancy (blue). Blank sectors mean that the trait is absent. Letters at the tips refer to growth forms in the clade (T, tree; S, shrub or subshrub; H, herb/annual). Black dots indicate the crown age of diversification of the corresponding clade. From [1].


[1] Pausas J.G. & Lamont B.B. 2022. Fire-released seed dormancy – a global synthesis. Biological Reviews  [doi | pdf | supp. mat. | data (figshare)]

Marmaris postfire regeneration

December 5th, 2021 No comments

During last summer, over 170,000 ha burnt in Turkey. One of these fires was the Marmaris fire, a fire of about 12,500 ha in SW of the country. The area includes part of the Marmaris National Park and an Special Environmental Protection Area. Most of the area were covered by Pinus brutia forests, a Mediterranean pine that has some serotinous cones.

Four months after the Marmaris fire, I visited the area together with Çagatay Tavsanoglu. Some plants were resprouting and some geophytes flowering (Cyclamen, Arisarum). Pine seedlings had started to germinate; there were also many other seedlings but still too small to identify them (e.g., Cistus species). Below are a few examples of plants that were regenerating after the fire (click on the photos to enlarge them).


Global change in the Mediterranean basin

January 9th, 2019 2 comments

The paleartic region with mediterranean climate (southern Europe and northern Africa; the Mediterranean Basin; Fig. 1) is a hotspot of biodiversity, a hotspot of climate change (warming of the region is above global average), and a hotspot of human population (a highly populated area and a top tourist and retirement destination). In addition, the Mediterranean Sea is the world’s largest inland sea, and climatic disruptions in the region have consequences in the large catchment area that includes central-eastern Europe (Fig. 1). That is, environmental changes and disruptions of the water cycling in the Mediterranean region have consequences affecting a large human population [1].

Fig. 1. Area with mediterranean climate (green) and limits of the Mediterranean catchment (red).  The European catchment limit based on Cortambert (1870). From [1].

The region, as all the planet, is subject to global warming. In addition there are three main local processes (not directly related to global warming) that are very important in understanding dynamic changes in the region [1]:

a) Rural abandonment in an environment depauperate of native herbivores; this increases wildlands (greening) but also the abundance and continuity of fuels that feed wildfires [2]

b) Increasing the wildland-urban interface; this increases biodiversity degradation (e.g., alien species), fire ignitions, and the vulnerability of the society to fires

c) Coastal degradation enhances drought (browning) through negative feedback processes; that is, the desiccation of coastal marshes, the deforestation for agriculture, and more recently, the explosive coastal urbanization, have drastically reduced the original ecosystems and thus the water available for the sea breeze that was once feeding the rain in the upper part of the mountains [1].

All these mechanisms act in different directions (greening, browning), and the current balance is still towards greening, as land abandonment is buffering the browning drivers; however, it is likely to switch with global warming. The challenge is to mitigate the browning processes. The good news is that the importance of small-scale drivers suggests that local policies and actions can make a difference in reducing overall impact on the landscape and society.

Mechanisms acting at a fine scale, together with global drivers (CO2 enrichment and climatic warming) interact and drive current vegetation changes in Mediterranean landscapes. Any model aiming to predict the future of our vegetation and climate must consider these local mechanisms; and failing to consider them at an appropriate scale is likely to produce inconclusive predictions.

Fig. 2. The disruption of the natural fire and drought regimes in Mediterranean landscapes is driven by global and local drivers. Increased fire activity is a response to the fuel amount and landscape homogeneity generated by rural abandonment (fire hazard) in an environment depauperated of herbivores and with increasing human ignitions (fire risk) and droughts (fire weather). The increased dry conditions are the consequence of global warming, but also of storm losses caused by the disruption of the water cycle generated by the coastal degradation. WUI: wildland-urban interface. From [1].


[1] Pausas J.G. & Millán M.M. 2019. Greening and browning in a climate change hotspot: the Mediterranean Basin. BioScience 69: 143-151 [doi | OUP | pdf]  

[2] 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 | springer | pdf]  

Every time you build anything, a highway, a school, whatever, you are altering the precipitation regime somewhere downwind” Millán M. Millán

Incendios forestales: encrucijada natural y social

December 28th, 2018 No comments

En febrero de 2018, poco después del incendio de Doñana (Las Peñuelas, 2017), la Academia de Ciencias Sociales y el Medio Ambiente de Andalucía organizó en Sevilla un seminario sobre regeneración después de incendios. A partir de ese seminario, se ha realizado un libro con algunas de las ponencias. Tuve la suerte de que me invitasen a dar la charla inaugural del seminario y a escribir un capitulo introductorio en el libro. Aquí abajo copio algunos fragmentos de mi capítulo. El texto entero del capítulo está disponible en PDF aquí. El libro entero también está disponible en PDF aquí y en en la Academia



Los incendios forestales constituyen procesos complejos, a muchas escalas. Los físicos aún tienen enormes dificultades para modelar y predecir el comportamiento del fuego a escalas temporales y espaciales pequeñas, debido a la complejidad de la vegetación y las interacciones con los parámetros meteorológicos. A escalas mayores, en paisajes heterogéneos, con historia, y con personas asentadas en él, la complejidad es enorme. Prueba de ello es que el llamado ‘problema’ de los incendios forestales no está resuelto ni en nuestro país ni en ningún otro. Parte de la dificultad probablemente resida en un enfoque poco apropiado para abordar el tema. Lo más probable es que se necesite un cambio de paradigma en cuanto a cómo entendemos los incendios forestales en nuestros paisajes. Para este cambio de paradigma se requiere acumular evidencias, cultivar el pensamiento crítico, y romper algunos mitos. Aquí estamos para contribuir a ello.

Régimen de incendios

El clima mediterráneo es propenso al fuego porque tiene una vegetación densa, una estación seca, y tormentas secas (que provocan igniciones). Por lo tanto, como mínimo, hay incendios desde que tenemos clima mediterráneos (hace unos pocos millones de años), aunque hay evidencias de carbones fósiles producto de incendios desde que las plantas colonizaron el medio terrestre. A lo largo de la historia geológica el régimen de incendios ha ido cambiando, es decir, han ido cambiando características tales como la frecuencia, la intensidad, la estacionalidad, el tamaño, o el tipo de propagación. Y estos cambios han sido debidos a las variaciones a escala geológica de la concentración de oxígeno en la atmósfera, a cambios en el clima y en la vegetación, y a variaciones en la abundancia y tipo de herbívoros.

A una escala temporal más reciente (humana), los regímenes de incendios se han ido modificando según el uso del fuego, los cambios antropogénicos del paisaje, de la ganadería/pastoreo, de las políticas de gestión y, finalmente, los cambios climáticos recientes (antropogénicos). Todos estos cambios han ido modificando el régimen de incendios en diferentes direcciones, a veces de manera abrupta. Por ejemplo, el aumento de la población está asociado a un mayor número de igniciones; el incremento de la agricultura está asociado a la disminución de los incendios; y el reciente abandono rural conllevó un incremento de estos, especialmente de su tamaño. De manera que la dinámica del régimen de incendios está fuertemente ligada a cambios socio-económicos.


La larga historia de incendios ha conllevado que muchas plantas mediterráneas tengan rasgos o estrategias que les confieren supervivencia y capacidad de reproducción en ambientes con incendios recurrentes (rasgos adaptativos a los incendios). Existe una variedad de rasgos de este tipo; por ejemplo, muchas plantas rebrotan muy bien aunque se quemen totalmente, gracias a tener yemas en estructuras subterráneas protegidas por el suelo, o protegidas debajo de cortezas gruesas. Otras especies germinan de manera abundante después de los incendios, y así aprovechan la elevada disponibilidad de recursos tras el paso del fuego; y en muchos casos, estas especies aumentan su tamaño poblacional respecto al tamaño previo al incendio. Otras especies florecen rápida y abundantemente después de un fuego, aprovechando las condiciones favorables. Todo ello genera una frenética actividad de plantas y animales durante la primavera después de los incendios. La gran mayoría de plantas mediterráneas presenta alguna de estas estrategias para vivir en zonas con incendios recurrentes (ver aquí), aunque cambios bruscos en el régimen pueden conllevar problemas ecológicos.

Los bosques no son la única alternativa

Aunque la vegetación potencial de muchos ecosistemas pueda ser el bosque cerrado, la larga historia de incendios ha producido la apertura en los bosques de manera tan recurrente que han evolucionado especies típicas de matorrales abiertos que ni siquiera pueden vivir dentro del bosque. De hecho no es raro encontrar matorrales más diversos que bosques. Por lo tanto, la sociedad debería poner más en valor los ecosistemas que no son bosques, tales como los matorrales, las dehesas y los pastizales; todos ellos con una elevada diversidad en nuestro territorio. El fuego genera espacios abiertos, y da oportunidad a especies heliófilas, tanto de plantas como de animales. Sin incendios recurrentes nuestro paisajes serían más pobres en especies.

Incendios en el antropoceno

A pesar del carácter natural y antiguo de los incendios, el incremento de igniciones (aumento de la población urbana en ambientes semi-rurales), la elevada continuidad de la vegetación (por abandono rural y falta de herbívoros), y el cambio climático, junto con la política de exclusión del máximo número de incendios, nos lleva en conjunto muy a menudo a regímenes de incendios fuera del rango histórico, donde la mayor frecuencia de incendios de grandes dimensiones podría generar problemas ecológicos y sociales. En un mundo con climas y paisajes cambiantes, la gestión de los incendios (que no la exclusión o extinción), resulta de suma importancia.

La gestión debe partir del conocimiento de que los incendios son propios de los ecosistemas mediterráneos. La política de tolerancia cero a los incendios no ha funcionado en ningún país del mundo. El reto de la gestión no debería ser eliminar los incendios, sino crear paisajes que generen regímenes de incendios sostenibles tanto ecológica como socialmente. Eliminarlos es imposible, antinatural y ecológicamente insostenible. Para generar esos paisajes resilientes se precisan acciones a distintos niveles, tales como aceptar abiertamente un cierto régimen de incendios, crear discontinuidades en paisajes forestales homogéneos (por ejemplo, mosaicos agrícola-forestales), o reducir el combustible (pastoreo y quemas prescritas) en zonas estratégicas o próximas a viviendas.  También implica decisiones tan conflictivas como limitar la interfaz urbano-forestal, es decir, limitar la expansión de urbanizaciones y polígonos industriales en zonas rurales y naturales. A los efectos ambientales que supone la expansión de estas zonas de interfaz (por ejemplo, en biodiversidad, especies invasoras, contaminación lumínica y visual, etc.), hay que añadir que constituyen una gran fuente de igniciones, y que convierten en catastróficos (socialmente) incluso a regímenes de incendios ecológicamente sostenibles.

En conclusión

Se están acumulando evidencias que sugieren que hasta ahora teníamos una visión muy incompleta y sesgada de los incendios forestales, relegándolos a un factor externo que destruye ecosistemas y genera problemas ecológicos y sociales. Los estudios realizados en los últimos años sugieren un cambio de visión, donde los incendios constituyen una característica interna de los sistemas socio-ecológicos, una perturbación natural en muchos ecosistemas y una herramienta de gestión para moldear los regímenes de incendios futuros. Por ello se requiere que aprendamos a convivir con los incendios. Este cambio de paradigma se hace más urgente con el cambio climático, ya que la actividad de incendios está aumentando, y solo se puede abordar si se integra los incendios y el fuego dentro de nuestros sistemas socio-ecológicos.


Lecturas relacionadas

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

Pausas J.G. & Paula S. 2012. Fuel shapes the fire-climate relationship: evidence from Mediterranean ecosystems. Global Ecol. & Biogeogr. 21: 1074-1082. [doi | pdf | supp]

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 | springer | pdf]

Chergui B., Fahd S., Santos X., Pausas J.G. 2018. Socioeconomic factors drive fire-regime variability in the Mediterranean Basin. Ecosystems 21: 619–628. [doi | pdf | post]

Pausas J.G. 2018. Incendios: cambios recientes y soluciones.

Pausas J.G. 2018. Incendios forestales, encrucijada natural y social. En: Ecología de la regeneración de zonas incendiadas (García Novo F., Casal M., Pausas J.G.). Academia de Ciencias Sociales y del Medio Ambiente de Andalucía. pp. 9-14. [pdf | libro]

Pausas J.G. & Millán M.M. 2019. Greening and browning in a climate change hotspot: the Mediterranean Basin. BioScience. [doi | pdf]


BROT 2 – a mediterranean plant trait database

July 10th, 2018 No comments

We have released a new version of the BROT database on plant functional traits for Mediterranean Basin species [1]. BROT 2.0 is an improved and expanded version of BROT 1.0 [2]; while the first version focused on fire related traits, the version 2.0 is more general and include a higher diversity plant functional traits. BROT 2 includes 25,764 individual records of 44 traits from 2,457 plant taxa distributed in 119 taxonomic families.

The structure of the BROT 2 database is very simple and include four files (or tables) linked by IDs: the Data file (the main file), the Taxa file, Sources file, and an additional Synonymous file. Tables 5, 6, 7, and 9 refer to the place in the paper [1] with the definitions of the corresponding field.Geographical scope of BROT in the Mediterranean Basin. Circles are the locations of the data (for records with geographical coordinates), with colors indicating the number of records. The blue region refers to the Mediterranean climate area following Quézel & Médail (2003).


[1] Tavsanoglu Ç. & Pausas J.G. 2018. A functional trait database for Mediterranean Basin plants. Scientific Data 5: 180135 [doi | SciDat | pdf | data | BROT web]

[2] Paula, S., M. Arianoutsou, D. Kazanis, Ç. Tavsanoglu, F. Lloret, C. Buhk, F. Ojeda, B. Luna, J. M. Moreno, A. Rodrigo, J. M. Espelta, S. Palacio, B. Fernández-Santos, P. M. Fernandes, and J. G. Pausas. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90:1420-1420. [doi | ESA journals | Ecological Archives | pdf | BROT web]


Incendios: cambios recientes y soluciones

June 19th, 2018 No comments

Recientemente, Greenpeace hizo un par de preguntas a varias personas que trabajan en diferentes ámbitos relacionados con incendios forestales (noticia | documento). Aquí copio mis respuestas.

¿En qué han cambiado los incendios?

En las últimas décadas se ha observado un cambio brusco en el régimen de incendios, aumentando la frecuencia y, especialmente, el tamaño de los incendios. Los incendios requieren de tres factores: combustible (vegetación densa y continua), igniciones, y condiciones propensas a la propagación del fuego (sequedad, viento). Estos tres factores se han visto favorecidos en los últimos años y de manera simultánea:

(1) ha aumentado la cobertura, continuidad y densidad de la vegetación en el paisaje, incrementando la biomasa y el combustible disponible para los incendios;

(2) ha cambiado el clima hacia veranos más cálidos, más secos, y más largos, por lo que se dan mejores condiciones para la propagación del fuego y durante más tiempo; y

(3) ha aumentado la población urbana en las zonas de interfaz urbano-forestal (ver foto), lo que conlleva un mayor número de igniciones, tanto accidentales como provocadas.

En definitiva, actualmente hay más incendios porque tenemos paisajes y climas más propensos a la propagación del fuego, y más igniciones.

De estos tres factores, el que más ha influido en el cambio del régimen de incendios es el aumento de vegetación (biomasa) en nuestros paisajes. Este aumento se debe principalmente al abandono de las actividades rurales (tales como la agricultura, el pastoreo, la extracción de leña, o la gestión de las plantaciones forestales), y a las políticas de exclusión total de los incendios, sin una sustitución por otros sistemas que controlen la vegetación, tales como los herbívoros silvestres, o las quemas y el pastoreo prescritos.

¿Como mitigar el impacto de los incendios?

La política de tolerancia cero a los incendios no ha funcionado en ningún país del mundo. El reto de la gestión no debería ser eliminar los incendios, sino crear paisajes que generen regímenes de incendios sostenibles tanto ecológica como socialmente. Para ello se precisan acciones a distintos niveles, tales como aceptar abiertamente un cierto régimen de incendios (especialmente en zonas poco pobladas y en ecosistemas con adaptaciones al fuego), crear discontinuidades en paisajes forestales homogéneos (por ejemplo, mosaicos agrícola-forestales), o reducir el combustible en zonas estratégicas o próximas a viviendas. También implica decisiones tan conflictivas como limitar la interfaz urbano-forestal, es decir, limitar la expansión de urbanizaciones y polígonos industriales en zonas rurales y naturales. A los efectos ambientales que supone la expansión de estas zonas de interfaz (por ejemplo, en biodiversidad, especies invasoras, contaminación lumínica y visual, etc.), hay que añadir que constituyen una gran fuente de igniciones, y que convierten en catastróficos (socialmente) incluso a regímenes de incendios ecológicamente sostenibles. Los mecanismos para limitar estas zonas pueden ser diversos, incluyendo la recalificación de terrenos (a no urbanizables), o la implementación de tasas (disuasorias) por construir en áreas con alto riesgo de incendios, entre otros. Además, y con carácter más general, incrementar las medidas que frenan el cambio climático contribuiría a reducir los cambios no deseados en el régimen de incendios.

Foto: Ejemplo de interfaz urbano-forestal (Port d’Andratx, Mallorca; foto: @xarxaforestal). Con este modelo de urbanismo, además de aumentar la probabilidad de igniciones, convertimos en catastróficos incluso a regímenes de incendios ecológicamente sostenibles.


Más información

Incendios forestales, una visión desde la ecología
Acabar con los incendios es antinatural e insostenible
Cinco cuestiones sobre inflamabilidad e incendios

Doñana postfire – Doñana posincendio

April 2nd, 2018 No comments

[English version]

Last summer, between Jun 24 and Jul 4 (2017), a wildfire burned ca. 10,000 ha of the Doñana Natural Park (Las Peñuelas fire, Moguer, Huelva, Spain); the fire did not affect the adjacent Doñana National Park (National P. + Natural P. = 108,000 ha). Most of the area burned was a shrubland in a fixed dune system that had been afforested with Pinus pinea during the early 20th C. Now, nine months after fire, there are many plants from the original shrubland that are resprouting and many seeds germinating (pictures below); most of the pines are dead with very poor or null regeneration, so part of the afforestation (which is much larger than this fire) is lost.

From the ecological point of view, this fire provides an opportunity to replace part of the afforestation with the natural shrubland, and thus the wildfire may help to restore the original ecosystems and their biodiversity. This ecosystem will also benefit from having more water available, as tree consume a lot of water. In addition, future fires occurring in the shrubland without the tree layer would be also less intense. Consequently, the regenerating shrubland will be more natural and more resilient to future fires than the prefire pine woodland.


[Versión en español]

El verano pasado, entre el 24 de junio y el 4 de julio (2017), un incendio afectó ca. 10.000 ha del Parque Natural de Doñana (incendio de Las Peñuelas, Moguer, Huelva, España); el fuego no afectó al Parque Nacional de Doñana adyacente al parque natural (P. Nacional + P. Natural = 108.000 ha). La mayor parte del área quemada era un matorral en un sistema de dunas fijas en el que se había plantado pinos (Pinus pinea) a principios del siglo XX. Ahora, nueve meses después del incendio, hay muchas plantas del matorral original que están rebrotando y muchas semillas germinando (ver fotos); la mayoría de los pinos están muertos y presentan una regeneración muy pobre o nula, por lo que una parte de la repoblación de pinos (que era mucho más grande que este incendio) ha desaparecido.

Desde el punto de vista ecológico, este incendio brinda la oportunidad de reemplazar la repoblación por el matorral natural y, por lo tanto, el incendio pueden ayudar a restaurar los ecosistemas originales y diversos de Doñana. Estos ecosistemas también se beneficiarán de una mayor disponibilidad de agua, ya que hasta ahora una parte era consumida por los pinos. Además, los incendios futuros que ocurran en estos matorrales sin pinos serán menos intensos. En consecuencia, el matorral en regeneración será más natural y más resiliente a los incendios futuros que el pinar anterior.

UPDATE (Jan 2019): this post has inspired the following article:

Leverkus AB, Murillo PG, Doña VJ, Pausas JG. 2019. Wildfires: opportunity for restoration? Science 363 (6423): 134-135. [doi | science | pdf]


Socioeconomics and fire regime in the Mediterranean

August 26th, 2017 No comments

In recent decades, fires in Mediterranean Europe have become larger and more frequent. This trend has been driven mainly by socioeconomic changes that have generated rural depopulation and changes in traditional land use. This has increased the amount and continuity of vegetation (fuel), and thus an increase in the fire size and area burnt [1-3]. In a recent paper [4] we compared fire statistics of the Western Rif (Morocco) with those form Valencia (eastern Spain) to show that current fire regimes in Mediterranean Africa resemble past fire regimes in the Mediterranean Europe when rural activities dominated the landscape. The temporal fire regime shift observed in different countries of the Mediterranean Europe (from small, fuel-limited fires to drought-driven fires) can be identified when moving from the southern to the northern rim of the Basin. That is, most spatial and temporal variability in fire regimes of the Mediterranean Basin is driven by shifts in the amounts of fuel and continuity imposed by changes in socioeconomic drivers (e.g., rural depopulation). In fact, we can use rural population density as an early warning for abrupt fire regime shift. Consequently we can predict future fire regimes in North Africa, based on the trends observed in southern Europe, and we can better understand past fire regimes in Europe based on the current situation in North Africa [4].

Figure 1. Western Rif (northern Morocco) and Valencia (eastern Spain).

Figure 2. Fire-size distribution in Valencia, for the period 1880-1970 (white boxes) and for the period 1975-2014 (grey boxes), and in the western Rif (red symbols, 2008-2015). For details see [4]


[1] Pausas, J.G. 2004. Changes in fire and climate in the eastern Iberian Peninsula (Mediterranean basin). Climatic Change 63: 337-350. [pdf | doi]

[2] 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 | springer | pdf]

[3] Pausas J.G. & Paula S. 2012. Fuel shapes the fire-climate relationship: evidence from Mediterranean ecosystems. Global Ecol. & Biogeogr. 21: 1074-1082. [doi | pdf | supp]  

[4] Chergui B., Fahd S., Santos X., Pausas J.G. 2018. Socioeconomic factors drive fire-regime variability in the Mediterranean Basin. Ecosystems 21: 619–628 [doi | pdf]


¿Hay incendios naturales?

July 14th, 2017 No comments

A menudo me preguntan, ¿pero tu realmente crees que los incendios forestales es un fenómeno natural? Aquí intento responder a esta pregunta. Este texto apareció primero en 20minutos (Ciencia para llevar); aquí incluyo la primera versión que escribí, un poco más larga que la publicada; la principal diferencia está en el último párrafo, que por razones de espacio se recortó en la versión final en 20minutos.


Para que se produzca un incendio forestal se requieren tres condiciones: una ignición que inicie el fuego, un combustible continuo e inflamable, y unas condiciones de propagación adecuadas. ¿Se dan estas tres circunstancias en nuestros ecosistemas?

Empecemos por el final, las condiciones de propagación. Una de las principales características del clima mediterráneo es que la estación más seca coincide con la más cálida (el verano), cosa que no se da en la mayoría de los climas del mundo. En verano se genera un periodo relativamente largo con unas condiciones de elevadas temperaturas y baja o nula precipitación, que son ideales para que, si hay un incendio, se propague fácilmente. Además no es raro tener días de viento relativamente fuerte, seco y cálido (por ejemplo, los ponientes en la costa valenciana) que aún facilitan más los grandes incendios.

La siguiente condición es la existencia de un combustible continuo e inflamable. En la mayoría de los ecosistemas ibéricos, la vegetación es suficiente densa y continua que permite, si hay un incendio en verano, que este se pueda extender a grandes superficies. Esto es aplicable tanto a los bosques como a la gran diversidad de matorrales que encontramos en nuestro territorio. De manera que la vegetación mediterránea forma lo que a menudo se llama el combustible de los incendios forestales. No hay que olvidar que este ‘combustible’ está compuesto por una gran diversidad de seres vivos que tienen detrás una larga historia evolutiva; son parte de nuestra biodiversidad. Esta continuidad en la vegetación era especialmente evidente antes de que los humanos realizará esa gran fragmentación que se observa actualmente en nuestros paisajes, principalmente debida a la agricultura, pero también a las abundantes vías y zonas urbanas y periurbanas.

Pero con una vegetación inflamable y unos veranos secos no es suficiente para que haya incendios, se requiere una ignición inicial. Hoy en día, la mayoría de igniciones son generadas por personas, ya sea de manera voluntaria o accidental. Pero, ¿Hay igniciones naturales? La respuesta es . A menudo tenemos tormentas secas en verano, cuando las condiciones de propagación son óptimas, de manera que los rayos generados por estas tormentas pueden actuar como fuente de ignición e iniciar un incendio forestal. Tenemos muchos ejemplos de incendios generados por rayos (la mayoría sofocados rápidamente por los bomberos); y en los meses de verano, la AEMET detecta miles de rayos potencialmente capaces de generar igniciones (Figura 1).

Figura 1. Imagen del 31 de Julio de 2015 donde se muestra la localización de 12835 rayos que se registraron durante 6 horas en la Península Ibérica. Los diferentes colores indican diferentes horas, entre las 12 y las 18h. Fuente: Agencia Estatal de Meteorología.

Por lo tanto, las tres condiciones arriba mencionadas se dan de manera natural en nuestros ecosistemas, y por lo tanto podemos afirmar que sí hay incendios naturales. Pero, ¿cuantos?

Las estadísticas de incendios actuales nos dicen que los incendios generados por rayos son una minoría, comparado con la gran cantidad de incendios generados por los humanos. ¿Podría esta minoría de incendios por rayo representar la cantidad de los incendios esperables en condiciones naturales? La respuesta es no. Una gran cantidad de rayos cae en suelo sin vegetación combustible (zonas agrícolas y urbanas) y por lo tanto no producen los incendios que producirían en unas condiciones más naturales. Además, de los rayos que sí generan igniciones en el monte, la mayoría son apagados por los bomberos forestales cuando aun son solo conatos o incendios muy pequeños. Cabe recordar que nuestros bomberos apagan la inmensa mayoría de las igniciones y sólo un porcentaje muy pequeño se escapa y se transforma en los incendios que aparecen en la prensa. Y además, de los incendios que realmente progresan, la mayoría son más pequeños de lo que serían esperable en condiciones más naturales, porque los apagan los bomberos, o porque se detienen en zonas no inflamables (zonas agrícolas, urbanas, cortafuegos, etc.). Como consecuencia, las estadísticas de incendios por rayos, ya sea en número de incendios como en área afectada, no reflejan la importancia que tendrían los incendios en condiciones naturales, sino que los subestima. Algunos de los incendios que actualmente se dan por actividad humana, en realidad están sustituyendo a incendios naturales.

Es decir, en unos paisajes más naturales (con menos presión humana) que los actuales, sería de esperar que hubiese menos incendios que en la actualidad porque habría muchas menos igniciones (la actual elevada población genera muchas igniciones), pero en muchos casos, esos incendios podrías ser más grande. En cualquier caso, el balance probablemente sería de menos área afectada por incendios que actualmente; pero sí habría incendios frecuentes. A todo esto hay que añadirle que actualmente estamos cambiando el clima, de manera que la estación con incendios tienden a ser más larga, y las olas de calor más frecuentes, y todo ello incrementa la actividad de los incendios; pero ahora no entraremos en detalle en ello.

Además, hablar de condiciones o paisajes ‘más naturales’ es complicado por varias razones. ¿Cuanto hacia atrás en el tiempo son esas condiciones ‘más naturales’? Los humanos han poblado la Península ibérica desde hace muchos años, modificando las igniciones, cambiando la estructura de la vegetación, así como la cantidad y tipo de herbívoros. Esto ha llevado a continuos cambios en la cantidad y continuidad del combustible y en el régimen (frecuencia, intensidad, y estacionalidad) de incendios. Y si nos vamos a periodos antes de los humanos, tanto el clima como la cantidad y tamaño de los herbívoros (también consumidores de biomasa, como el fuego) era bastante diferente. Por lo tanto, lo importante no es si el régimen de incendios actual es ‘natural’ o no. Lo importante es si el régimen de incendios actual y futuro es ecológica y socialmente sostenible, considerando el cambio climático. Eliminar los incendios es imposible, antinatural y ecológicamente insostenible. Nuestra sociedad ha de aceptar la existencia de incendios, aprender a convivir con ellos, adaptar las estructuras y los comportamientos, y gestionar las zonas semi-urbanas y los paisajes rurales para que el régimen de incendios sea ecológica y socialmente sostenible. Esto incluye gestionar y planear la zonas semi-urbanas, la plantaciones forestales, y los parques naturales, pensando que lo normal es que un día les llegue un incendio.

Todo esto y más en: ‘Incendios forestales’ Ed. CSIC-Catarata.

[Actualización 30/7/2017] Un ejemplo: La sierra de los rayos. El País, 30 Julio 2017

Homage to Louis Trabaud

June 6th, 2017 No comments

Louis Trabaud (born in Montpellier, 2nd Feb. 1937) has recently passed away (Collioure, 16th April 2017). He was a research on plant ecology at Centre d’Ecologie Fonctionnelle et Evolutive (CEFE) of the CNRS, France. He was a pioneer of fire ecology in the Mediterranean Basin and set the basis of this topic for the region; he was especially influential to the fire ecologist of Spain (including me), and was awarded Professor Honoris Causa by the Univeristy of León (Spain) [1]. He was also award in France as Chevalier du Mérite Agricole (Order of Agricultural Merit). As a person, Louis Trabaud was very kind and always happy to help any student.

His research was especially focused on mediterranean shrublands around Montpellier (garrigue), i.e., shublands dominated by Quercus coccifera, Cistus species, Rosmarinus officinalis, Fumana species, etc… sometimes with an overstory of Pinus halepensis. He performed the first fire experiments in the Mediterranean region to study the regeneration of these shrublands, where he recurrently burned them in different seasons to demonstrate their high resilience. He also performed the first studies in the Mediterranean Basin on heat-stimulated germination and on flammability traits. He produced many papers, some in French (the most earlier ones) and others in English, and also wrote or edited some books. The full Trabaud’s publications list (books and scientific papers) is available here.

Louis Trabaud, together with his friend Roger Prodon, organized the International Workshop on Fire Ecology in Banyuls-sur-Mer (south of France) in the years 1992, 1997, 2001; these workshops were key in building the knowledge on fire ecology for the region; they were the meeting point for all mediterranean fire ecologist; we all met there for the first time and we all have very good memories from those meetings.

Participants of the 2001 Banyuls meeting organized by Louis Trabaud (in the middle, with glasses and a pale sweater) and Roger Prodon (second from the right).


[1] Texto en memoria de Louis Trabaud, por la Universidad de León (in Spanish)


Incendios y biodiversidad

May 31st, 2017 No comments

El 12 de Mayo de 2017 impartí una charla titulada Incendios forestales y biodiversidad en el IVIA (Valencia), en la que expliqué las principales adaptaciones de las plantas mediterráneas a los incendios, y cómo estudiamos esas adaptaciones en el marco de la ecología del fuego. La conclusión es que el fuego explica una parte de la biodiversidad de nuestros ecosistemas. La charla tuvo cierto impacto en los medios (enlace). Aquí podéis ver la charla integra así como la discusión posterior:

También disponible en la web del IVIA, GV

Más información: | @jgpausas | Incendios forestalesFire and diversity at the global scale | Fire adaptations in Mediterranean basin plants | Evolutionary fire ecology in pines | Ulex born to burn (II) | Serotiny|

Pinus brutia

April 19th, 2017 No comments

Pinus halepensis is a strongly serotinous pine [1,2] occurring mainly in the western part of the Mediterranean Basin (especially in Spain; see map below). The most phylogenetically closely related species to P. halepensis is Pinus brutia that occurs in the eastern Mediterranean Basin (mainly in Turkey). P. brutia is called ‘red pine’ in Turkish (Kızıl çam) because sometimes the upper part of the trunk is reddish (as in P. sylvestris); the leaves are pale green as in P. halepensis. In relation to their fire response strategy [3], the main differences between the two species are that P. brutia is taller, the bark is thicker and the serotiny level is lower. Our observations suggest that P. brutia have relatively few serotinous cones, and most of then are less than 4 years old; P. halepensis have a higher proportion of serotinous cones, with many of them over 5 year old (and some more than 20 year old) [1].

Distribution maps of P. halepensis and P. brutia (from Wikipedia)

Pinus brutia forest, serotinous cones (2 serotinous and one non-serotinous (open)), and an example of a bark of more than 5 cm thick (the gaude was too short!). Photos from SW Turkey (by JG Pausas). For an example of serotinous cones in P. halepensis, see here.


[1] Hernandez-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. Am. J. Bot. 100: 2349-2356. [doi | pdf | supp.]

[2] Castellanos, M.C., González-Martínez, S. & Pausas, J.G. 2015. Field heritability of a plant adaptation to fire in heterogeneous landscapes. Mol. Ecol. 24, 5633-5642. [doi | pdf | suppl. | blog]

[3] Pausas, J.G. 2015. Evolutionary fire ecology: lessons learned from pines. Trends Pl. Sci. 20: 318-324. [doi | pdf | blog]

Eastern Mediterranean tour, 2017

April 18th, 2017 No comments

It is a pleasure to visit the different ecosystems of Lebanon and Turkey, and to meet old students and colleagues.

Cedar (Cedrus libani) forest, Al Shouf Biosphere Reserve, Lebanon

Anatolian steppe (bushes are Astragalus), central Turkey

Pinus brutia forest, SW Turkey


February 6th, 2017 1 comment

The XIV International Conference on Mediterranean Ecosystems (MEDECOS), has been successfully held on Sevilla, Spain, 31st Jan – 4th Feb, 2017, together with the conference of the Spanish Society of Terrestrial Ecology (AEET). For details, see the web page of the meeting and the post conference comments in the J. Ecol. blog. My contributions to this MEDECOS include two talks on fire and biotic interactions, two on resprouting, and one on fire hazard:

  • Pausas J.G – Fire and biotic interactions: the benefits of the disruption
  • García Y., Castellanos M.C., Pausas J.G. – Fires do not jeopardize reproduction of Chamaerops humilis despite disrupting its pollination
  • Tavsanoglu C. & Pausas J.G. – Resprouting ability encapsulates the most functional variability in the Mediterranean Basin flora
  • Paula, S. & Pausas J.G – Worldwide geographic and phylogenetic distribution of lignotubers
  • Cáceres M. de, Casals P., Álvarez D., Pausas J.G., Vayreda J., Beltrán M. – The role of understory fuel characteristics in the fire hazard of Mediterranean forests

The last day I attended to the field trip to Los Alcornocales Natural Park (a mosaic of cork oak forests and heathlands), where I enjoyed a long conversation on alternative vegetation states in non-tropical ecosystems (e.g., PDF) with William Bond (photo below).

Thank you very much to the organizers of MEDECOS, especially to Juan Arroyo (Universidad de Sevilla) and Montse Vilà (Doñana-CSIC) for this nice and smooth conference.


William Bond (left) and myself (right) in a Cork oak forest (photo: F. Ojeda)


November 17th, 2015 1 comment

Lignotubers are swollen woody structures located at the root-shoot transition zone of some plants; they contain numerous dormant buds and starch reserves [1]. They are ontogenetically programmed, that is, they are not the product of repeated disturbances; and thus they can be observed at very early stages of the plant development (other types of basal burls may be a response to multiple disturbances). Lignotubers enables the plant to resprout prolifically after severe disturbances that remove the aboveground biomass, thus they are considered adaptive in fire-prone ecosystems [2]. Lignotubers are not well-known in many floras because they are often below-ground (i.e., detected only after excavation) and because they are often confused by other non-ontogenetically determined basal burls; thus some reports of lignotubers in the literature are mistakes. In a recent review [1] we provide examples of species with a clear evidence of lignotubers in the Mediterranean basin, together with detailed morphological and anatomical description of lignotubers in saplings. The species with lignotuebers in the Mediterranean basin include many Erica species (e.g. E. arborea, E. scoparia, E. australis, E. lusitanica, E. multiflora), the two Arbutus species (A. unedo, A. andrachne), Rhododendron ponticum, Viburnum tinus, Phillyera angustifolia, Quercus suber (not obvious macroscopically!), Tetraclinis articulata and Juniperus oxycedrus (but not in all populations!). Please let me know (email address here) if you know of other Mediterranean basin species with lignotubers! Thanks

Figures: Examples of lignotubers for Mediterranean basin species. A Juniperus oxycedrus (resprouting after fire). B Viburnum tinus. C Arbutus unedo. D Quercus suber (not a clear basal swelling). E Olea europaea. F Phillyrea angustifolia (adult), G Phillyrea angustifolia (saplings). In many species (e.g., V. tinus, A. unedo and P. angustifolia) the lignotuber is only evident after excavating the root-shoot transition zone.


[1] Paula S., Naulin P.I., Arce C., Galaz C. & Pausas J.G. 2016. Lignotubers in Mediterranean basin plants. Plant Ecology  [doi | pdf | suppl.]

[2] Keeley J.E., Pausas J.G., Rundel P.W., Bond W.J., Bradstock R.A. 2011. Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci. 16: 406-411.  [doi | sciencedirect | pdf | For managers]


Flammable organic compounds: Rosmarinus officinalis

October 2nd, 2015 No comments

Given an ignition source and the right environmental conditions, all plants can potentially burn. However, some plants have characteristics that make them burn more easily. The capacity to store volatile organic compounds (VOCs) such as aromatic terpenes, can be considered one of these flammability-enhancing traits (flammable organic compounds, FOCs), as has now been demonstrated for Rosmarinus officinalis [1]: The more terpenes in the leaves, the more quickly they ignite (i.e., less time to ignition) (Figure below). Other species enhance flammability by having a very fine fuel, retaining dead fuel or having a flammable canopy structure [2-5]. There is growing evidence that flammability-enhancing traits are adaptive in Mediterranean fire-prone ecosystems [2-4]. To what extent the evolutionary pressure exerted by fire could have contributed to the abundance of aromatic plants in many fire-prone ecosystems (mints, rosemary, thyme, eucalypts, etc…) remains unknown. But certainly Mediterranean ecosystems are probably the most aromatic and among the most flammable ecosystems in the world.

Figure: relation between time to ignition (given a heat source, corrected by the differences in moisture) and the contents of terpenes (here the sum of camphene, para-cymene, borneol, limonene) in leaves of a wild population of rosmary (Rosmarinus officinalis), in Eastern Spain (from [1]). The top right corner shows the epiraditor, the device for  testing for time-to-ignition (see [2]).

[1] Pausas J.G., Alessio G.A., Moreira B., Segarra-Moragues J.G. (in press). Secondary compounds enhance flammability in a Mediterranean plant. Oecologia. [doi | pdf]

[2] Pausas J.G., Alessio G., Moreira B. & Corcobado G. 2012. Fires enhance flammability in Ulex parviflorus. New Phytologist 193: 18-23. [doi | wiley | pdf] [Ulex born to burn]

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

[4] Moreira B., Castellanos M.C., Pausas J.G. 2014. Genetic component of flammability variation in a Mediterranean shrub. Molecular Ecology 23: 1213-1223. [doi | pdf] [Ulex born to burn (II)]

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


Fire adaptations in Mediterranean Basin plants

September 7th, 2015 No comments

Few days ago a botanist colleague ask me whether there were some fire adaptations in the plants of the Mediterranean Basin, similar to those reported in other mediterraenan-climate regions. So I realised that researchers working on other topics may not be aware of the recent advances in this area. Here is my brief answer, i.e., some examples of species growing in Spain that show fire adaptations; this is by no means an exhaustive list, but a few examples of common species for illustrative purpose. You can find a description of these adaptations and further examples elsewhere [1, 2, 3, 4]. It is also important to note that plants are not adapted to fire per se, but to specific fire regimes, and thus some adaptations my provide persistence to some fire regimes but not to all [1]. That is, species that exhibit traits that are adaptive under a particular fire regime can be threatened when that regime changes.

  • Serotiny (canopy seed storage): Pinus halepensis, Pinus pinaster, with variability in serotiny driven by different fire regimes [5, 6]
  • Fire-stimulated germination: There are examples of heat-stimulated germination, like many Cistaceae (e.g., Cistus, Fumana [7, 8]) and many Fabaceae (e.g., Ulex parviflorus, Anthyllis cytisoides [7, 8]), as well as examples of smoke-stimulated germination like many Lamiaceae (e.g., Rosmarinus officinalis, Lavandula latifolia [7]) or Coris monspeliensis (Primulaceae [7]). There are also examples of species with smoke-stimulated seedling growth (Lavandula latifolia [7])
  • Resprouting from lignotubers: Arbutus unedo, Phillyrea angustifolia, Juniperus oxycedrus, many Erica species (e.g., E. multiflora, E. arborea, E. scoparia, E. australis) [4, 17]
  • Epicormic resprouting: Quercus suber [9, 10], Pinus canariensis [4]
  • Fire-stimulated flowering: Some monocots like species of Asphodelus, Iris, Narcissus [11, 12]
  • Enhanced flammability: Ulex parviflorus shows variability of flammability driven by different fire regimes [13] and under genetic control [14]. Many Lamiaceae species have volatile organic compounds that enhance flammability (e.g., Rosmarinus officinalis [16]).
  • Thick bark and self-pruning (in understory fires): Pinus nigra [3,15]




[1] Keeley et al. 2011. Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci 16:406-411. [doi | pdf]

[2] Keeley et al. 2012. Fire in Mediterranean Ecosystems. Cambridge University Press. [book]

[3] Pausas JG. 2012. Incendios forestales. Catarata-CSIC. [book]

[4] Paula et al. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90:1420-1420. [doi | pdf | BROT database]

[5] Hernández-Serrano et al. 2013. Fire structures pine serotiny at different scales. Am J Bot 100:2349-2356. [doi | pdf]

[6] Hernández-Serrano et al. 2014. Heritability and quantitative genetic divergence of serotiny, a fire persistence plant trait. Ann Bot 114:571-577. [doi | pdf]

[7] Moreira et al. 2010. Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Ann Bot 105:627-635. [doi | pdf]

[8] 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]

[9] Pausas JG. 1997. Resprouting of Quercus suber in NE Spain after fire. J Veget Sci 8:703-706. [doi | pdf]

[10] Catry et al. 2012. Cork oak vulnerability to fire: the role of bark harvesting, tree characteristics and abiotic factors. PLoS ONE 7:e39810. [doi | pdf ]

[11] Postfire flowering: 2 May 2015

[12] Postfire blooming of Asphodelous, 5 Apr 2014

[13] Pausas et al. 2012. Fires enhance flammability in Ulex parviflorus. New Phytol 193:18-23. [doi | pdf]

[14] Moreira et al. 2014. Genetic component of flammability variation in a Mediterranean shrub. Mol Ecol 23:1213-1223. [doi | pdf]

[15] He et al. 2012. Fire-adapted traits of Pinus arose in the fiery Cretaceous. New Phytol 194:751-759. [doi | pdf | picture]

[16] Flammable organic compounds: Rosmarinus officinalis, 2-Oct-2015

[17] Paula et al. 2016. Lignotubers in Mediterranean basin plants. Plant Ecology [doi | pdf | suppl. | blog]


How plants survive the harsh environment of Australia

June 1st, 2015 No comments

New book: Groom, P. K., and Lamont, B. B. (2015). Plant Life of Southwestern Australia. Adaptations for Survival. De Gruyter Open

Early explorers described Western Australia as ‘the most barren spot on the face of the earth’. In this book we learn that south-western Australia is one of the world’s biodiversity hotspots – not despite but because of its harsh environment. Nutrient-poor soils, frequent droughts, and recurrent fires, together with adverse fauna interactions (e.g., strong-billed cockatoos, voracious kangaroos, and the lack of efficient pollinating bees and hummingbirds) have made this region the perfect evolutionary scenario for developing a plethora of plant adaptations and assembling an hyperdiverse flora. The authors nicely describe this scenario and offer an impressive wealth of knowledge on the natural history of the region in an attractive book with abundant tables and quality full-colour pictures. One of the strengths of the book is that it brings together both biotic and abiotic factors to explain biodiversity, something uncommon in most specialised books.

Overall this is a must-read book for Australian naturalists but will also be a key reference for international ecologists interested in how plants thrive and evolve in dry, nutrient-poor, fire-prone environments. The lessons learned from this region help us understand evolutionary pathways in other dry regions worldwide.


Ecology and evolution in fire-prone ecosystems

February 28th, 2015 2 comments

During the last years I’ve been working in many topics related to fire ecology and plant evolution in ecosystems subject to recurrent fires (mainly mediterranean and savanna ecosystems). Because I believe knowledge should be spread around easily, I make my results available to the public in my web page (see publications list) and in this blog. However, having the cumulative list of paper published each year is not very convenient for people searching for a specific topic. For this reason, I’m rearranging most of my articles by topics as follows:

1. Fire history
2. Fire regime: climate & fuel
3. Fire traits (resprouting, postfire germination, serotiny, bark thickness, flammability, data & methods)
4. Fire & plant strategies (in Mediterranean ecosystems, in pines, in savannas, community assembly)
5. Fire & evolution
6. Some fire-adapted species (Pinus halepensis, Quercus suber, Ulex parviflorus)
7. Fire & vegetation modelling
8. Plant-animal interactions
9. Restoration & conservation

See: fire-ecology-evolution.html

Some papers may be repeated if they clearly fit in more than one topic; some papers, mainly old ones, do not fit well in any of these topics and have not been included (at least at the moment), they still can be found in the section of publications sorted by year. I’m still working on this rearrangement, so some modifications are possible; and any comment is welcome.
I hope this is useful for somebody!

Publications: by year | by topic | books


Cultural trees: Cupressus sempervirens

January 4th, 2015 3 comments


Cupressus sempervirens (Mediterranean cypress; ciprés) and Colosseum, Rome,
1 January 2015 (photo: J.G. Pausas)

Evolutionary ecology of resprouting and seeding

July 15th, 2014 No comments

There are two broad mechanisms by which plant populations persist under recurrent fires: resprouting from surviving tissues, and seedling recruitment [1]. Species that live in fire-prone ecosystems can have one of these mechanisms or both [1]. In a recent review paper [2], we propose a model suggesting that changes in evolutionary pressures that modify adult (P) and juvenile (C) survival in postfire conditions (Fig. 1 below) determine the long-term success of each of the two regeneration mechanisms, and thus the postfire regeneration strategy: obligate resprouters, facultative species and obligate seeders (Fig. 2). Specifically we propose the following three hypotheses: 1) resprouting appeared early in plant evolution as a response to disturbance, and fire was an important driver in many lineages; 2) postfire seeding evolved under conditions where fires were predictable within the life span of the dominant plants and created conditions unfavorable for resprouting; and 3) the intensification of conditions favoring juvenile survival (C) and adult mortality (P) drove the loss of resprouting ability with the consequence of obligate-seeding species becoming entirely dependent on fire to complete their life cycle, with one generation per fire interval (monopyric life cyle). This approach provides a framework for understanding temporal and spatial variation in resprouting and seeding under crown-fire regimes. It accounts for patterns of coexistence and environmental changes that contribute to the evolution of seeding from resprouting ancestors. In this review, we also provide definitions and details of the main concepts used in evolutionary fire ecology: postfire regeneration traits, postfire strategies, life cycle in relation to fire, fire regimes (Box 1), costs of resprouting (Box 2), postfire seeding mechanisms (Box 3), and the possible evolutionary transitions (Box 4).


Fig. 1 : Main factors affecting adult and offspring seedling survival (P and C, respectively), and thus the P/C ratio, in fire-prone ecosystems (from Pausas & Keeley 2014 [2]).



Fig. 2: The changes in the probability of resprouting along an adult-to-offspring survival (P/C) gradient are not linear but show two turning points related to the acquisition of key innovations: the capacity to store a fire-resistant seed bank (postfire seeding), and the loss of resprouting capacity. Changes in P/C ratio may be produced by different drivers (Fig. 1) which drove the rise of innovations during evolution, e.g., during the increasing aridity from the Tertiary to the Quaternary (from Pausas & Keeley 2014 [2]).



[1] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [doi | pdf | esa | jstor]

[2] Pausas J.G. & Keeley J.E. 2014. Evolutionary ecology of resprouting and seeding in fire-prone ecosystems. New Phytologist 204: 55-65 [doi | wiley | pdf]


Ulex born to burn (II): genetic basis of plant flammability

January 25th, 2014 No comments

In an previous study we found that Ulex parviflorus (Fabaceae) populations that inhabit in recurrently burn areas (HiFi populations) were more flammable than populations of this species growing in old-fields where the recruitment was independent of fire (NoFi populations) [1,2, 3]. That is, HiFi plants ignited quicker, burn slower, released more heat and had higher bulk density than NoFi plants. Thus, it appeared that repeated fires selected for individuals with higher flammability, and thus driving trait divergence among populations living in different fire regimes. These results were based on the study of plant flammability (phenotypic variability) without knowing whether plant flammability was genetically controlled. In a recent study using the same individuals [4], we show that phenotypic variability in flammability was correlated to genetic variability (estimated using AFLP loci) [figure below]. This result provide the first field evidence supporting that traits enhancing plant flammability have a genetic component and thus can be responding to natural selection driven by fire [5]. These results highlight the importance of flammability as an adaptive trait in fire-prone ecosystems.


Figure: Relationship between flammability and genotypic variability at individual level in Ulex parviflorus (red symbols: individuals in HiFi populations; green symbols: individuals in NoFi populations). Variations in flammability are described using the first axis of a Principal Component Analysis (PCA1) performed from different flammability traits, and genetic variability is described using the first axis of a Principal Coordinate Analysis (PCo1) from the set of AFPL loci that were significantly related to flammability. See details in [4].

[1] Ulex born to burn,, 9/Nov/2011

[2] Pausas J.G., Alessio G., Moreira B., Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193:18-23 [doi | wiley | pdf]

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

[4] Moreira B., Castellanos M.C., Pausas J.G. 2014. Genetic component of flammability variation in a Mediterranean shrub. Molecular Ecology 23: 1213-1223 [doi | pdf | data:dryad]

[5] Keeley J.E., Pausas J.G., Rundel P.W., Bond W.J., Bradstock R.A. 2011. Fire as an evolutionary pressure shaping plant traits. Trends in Plant Science 16: 406-411. [doi | trends | pdf]



November 16th, 2013 No comments

Serotiny is the delayed seed release for more than a year by retaining the seeds in a woody structure [1]. This implies an accumulation of a canopy seed bank. Serotiny confer fitness benefits in environments with frequent crown-fires, as the heat opens the cones and seeds are dispersed in the post-fire bed which is rich in resource and the competition and predation are low. It is typical of many Proteaceae and some conifers, like some pine species [1, 2; figure below].

Two recent papers analyse the serotiny of two mediterranean pines Pinus halepensis and Pinus pinaster [3, 4]. P. halepensis show higher proportion of serotinous cones than P. pinaster, but the latter retain the cones for longer [3]. The two species show high variability of serotiny within and between populations, but they show a clear pattern of higher serotiny in populations subject to high frequency of crown-fires than those living in areas where crown-fires are rare or absent. This is true either considering serotiny as the proportion of serotinous cones or as the age of the cones stored. Compared with other pines worldwide, the strength of the fire-serotiny relationship in P. pinaster is intermediate, and in P. halepensis is among the highest known [3]. For P. halepensis (the species with higher % serotiny), populations in high fire recurrence regimes have higher fine-scale spatial aggregation of serotiny than those inhabiting low fire recurrence systems. This phenotypic spatial structure generated by fire could be a consequence of the spatial genetic structure of the population. The second study used genomic tools to search for a genetic association for serotiny [4]. The analysis of 384 SNPs of 199 individuals of P. pinaster (in 3 populations included in the previous study [3])  shows that 17 loci were associated with serotiny and explain all together ca. 29% of the serotiny variation found in the field. All these results adds further evidence to the emerging view that fire shapes intraspecific variability of traits and generates phenotypic divergence between populations [5, 6, 7].

Figure: Serotinous cones of Pinus pinaster (Foto: K.B. Budde)


[1] Keeley J.E., Bond W.J., Bradstock R.A., Pausas J.G. & Rundel P.W. 2012. Fire in Mediterranean Ecosystems: Ecology, Evolution and Management. Cambridge University Press.  [The book]

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

[3] Hernández-Serrano A., Verdú M., González-Martínez S.C., Pausas J.G. 2013. Fire structures pine serotiny at different scales. American Journal of Botany 100: 2349-2356 [doi | amjbot | pdf | supp.]

[4] Budde, K. B., Heuertz, M., Hernández-Serrano, A., Pausas, J.G., Vendramin, G.G., Verdú, M. & González-Martínez, S.C. 2014. In situ genetic association for serotiny, a fire-related trait, in Mediterranean maritime pine (Pinus pinaster Aiton). New Phytologist  201: 230-241 [doi | pdf]

[5] Keeley J.E., Pausas J.G., Rundel P.W., Bond W.J., Bradstock R.A. 2011. Fire as an evolutionary pressure shaping plant traits. Trends in Plant Science 16(8): 406-411. [doi] [trends] [pdf]

[6] Pausas, J. G., Schwilk, D. W. 2012. Fire and plant evolution. New Phytologist, 193:301-303. [doi | wiley | pdf]

[7] Pausas J.G., Alessio G., Moreira B. & Corcobado G. 2012. Fires enhance flammability in Ulex parviflorusNew Phytologist 193: 18-23. [doi | wiley | pdf]


Physiological differences between resprouters and seeders

November 9th, 2013 No comments

The ability to resprout and to recruit after fire are two extremely important traits for the persistence in fire-prone ecosystems [1,2], and they define three life histories: obligate resprouters, obligate seeders (non-resprouters), and facultative seeders. After a fire, obligate seeders die and recruit profusely from the seeds stored in the seed bank [3-5]. In contrast, resprouters survive after fire and their above-ground tissues regenerate from protected (often below-ground) buds by using stored carbohydrates [6]. Facultative seeders not only recruit profusely after fire, but are also able to resprout. In fact, seeders and resprouters have different regeneration niches: seedling regeneration of obligate resprouters is not linked to fire, and they recruit during the inter-fire period under sheltered conditions (i.e., under vegetation cover), while seedling regeneration of seeders occurs in open postfire environments. Given the marked difference in water availability between microsites under vegetation and microsites open to the sun under Mediterranean conditions, seedlings of resprouters and seeders are subjected to different water-stress conditions, and thus they are expected to have different physiological attributes. Despite these differences, resprouters and seeders co-exist, are often well-mixed on local and landscape scales [7,8], and represent the two main types of post-fire regeneration strategies in Mediterranean ecosystems [2].

A recent study demonstrates marked differences in physiological attributes between seedlings of seeders and resprouters [9]: Seeders show a range of physiological traits that better deal with water-limited and highly variable conditions (e.g., higher resistance to xylem cavitation, earlier stomatal closure with drought, higher leaf dehydration tolerance), but they are also capable of taking full advantage of periods with high water availability (greater efficiency in conducting water through the xylem to to sustain high gas exchange rates when water is available). Conversely, resprouter species are adapted to more stable water availability conditions, favoured by their deeper root system, but they also display traits that help them resist water shortages during long summers.

Previous studies already showed marked differences between seeders and resprouters in a range of attributes: resprouters tend to exhibit a deeper root-system, while seedling root structure of seeders are more efficient in exploring the upper soil layer [10]. Leaves of seeders show higher water use efficiency (WUE) and higher leaf mass per area (LMA; i.e., higher sclerophylly, lower SLA) [11]. Seeds of seeder species are more tolerant to heat shocks and have greater heat-stimulated germination [3]. All these differences support the idea that they are distinct syndromes with different functioning characteristics at the whole plant level and suggest that they undertook different evolutionary pathways [12].

Figure: Coexistence of resprouters (R+) and seeders (R-) in postfire conditions near Valencia, Spain. (Foto: A. Vilagrosa).



[1] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [jstor |[pdf | Ecological Archives E085-029]

[2] Keeley J.E., Bond W.J., Bradstock R.A., Pausas J.G. & Rundel P.W. 2012. Fire in Mediterranean Ecosystems: Ecology, Evolution and Management. Cambridge University Press. [The book]

[3] Paula S. & Pausas J.G. 2008. Burning seeds: Germinative response to heat treatments in relation to resprouting ability. Journal of Ecology 96 (3): 543 – 552. [doi | pdf]

[4] 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 | blog]

[5] Moreira B. & Pausas J.G. 2012. Tanned or burned: The role of fire in shaping physical seed dormancy. PLoS ONE 7(12): e51523. [doi | plos | pdf | blog]

[6] Moreira B., Tormo J, Pausas J.G. 2012. To resprout or not to resprout: factors driving intraspecific variability in resprouting. Oikos 121: 1577-1584. [doi | pdf]

[7] Verdú M, & Pausas JG 2007. Fire drives phylogenetic clustering in Mediterranean Basin woody plant communities Journal of Ecology 95 (6), 1316-323 [doi | pdf]

[8] Ojeda, F., Pausas, J.G., Verdú, M. 2010. Soil shapes community structure through fire. Oecologia 163:729-735. [doi | pdf | blog]

[9] Vilagrosa A., Hernández E.I., Luis V.C., Cochard H., Pausas, J.G. 2014. Physiological differences explain the co-existence of different regeneration strategies in Mediterranean ecosystems. New Phytologist 201 : xx-xx [doi | pdf | suppl.] – NEW

[10] Paula S. & Pausas J.G. 2011. Root traits explain different foraging strategies between resprouting life histories. Oecologia 165:321-331. [doi | pdf | blog]

[11] Paula S. & Pausas J.G. 2006. Leaf traits and resprouting ability in the Mediterranean basin. Functional Ecology 20: 941-947. [doi | pdf | blog]

[12] Verdú M. & Pausas J.G. 2013. Syndrome-driven diversification in a Mediterranean ecosystem. Evolution 67: 1756-1766. [doi | pdf | blog]


Mediterranean diversification and plant syndromes

January 21st, 2013 No comments

Woody plants of the Mediterranean Basin can be classified in two contrasted morpho-functional syndromes [1]: a) plants with sclerophyllous, evergreen leaves and small, unisexual greenish or brownish flowers with a reduced perianth, and large seeds dispersed by vertebrates; and b) plants with alternative character states (non-sclerophyllous deciduous, semi-deciduous or summer deciduous species with large and conspicuous flowers pollinated by insects, and small seeds). The sclerophyllous syndrorme (a) occurs in clades whose characteristics pre-date the appearance of the mediterranean climate while the non-sclerophyllous syndrome (b) arose in clades that have evolved after the emergence of this distinctive climate (Tertiary – Quaternary transition).

A recent phylogenetic study [2] show that during the time with prevalent mediterranean climate, lineages with the non-sclefophyllous syndrome showed a higher speciation rate than the sclerophyllous lineages, suggesting that a syndrome-driven local diversification has occurred in shrublands under mediterranean conditions. The processes behind this result might be divers, but fire might had an important role. The rise of mediterranean climate increased fire activity [3] and traits defining these two syndromes are related to post-fire regeneration traits and to the age to maturity [4,5]. The non-sclerophyllous syndrome is associated with species considered post-fire seeders (i.e., killed by fire in which populations regenerate from a persistent seed bank; fire-stimulated germination [6,7]) and to species with early maturation. In fire-prone ecosystems, these characteristics reduce the generation time and the overlap between generations and thus they provide more opportunities for diversification.

Overall, the results provide an example of how the integration of the environmental filter in a dated phylogeny may recreate the local history of lineages and help to explain assembly processes in mediterranean ecosystems.

Figure: Frequency distribution of differences in local speciation rate (λ) between non-sclerophyllous (n) and sclerophyllous (s) syndromes in the Valencia woody flora for 3 different post cut temporal slices (cutoff of 10, 6, and 3.6 My) related to the increasing aridity associated with the rise of mediterranean climate. For all alternative phylogenies (i.e., accounting for the undertainity in node age), speciation rate of the non-sclerophyllous syndrome is greater than for the sclerophyllous one. See Verdú & Pausas (2013) for details [2].

[1] Herrera, CM. 1992. Historical effects and sorting processes as explanations for contemporary ecological patterns: character syndromes in Mediterranean woody plants. Am. Nat. 140:421-446.

[2] Verdú M. & Pausas J.G. 2013. Syndrome-driven diversification in a Mediterranean ecosystem. Evolution. [doi | pdf]

[3] Keeley JE., Bond WJ., Bradstock RA., Pausas JG. & Rundel PW. 2012. Fire in Mediterranean Ecosystems: Ecology, Evolution and Management. Cambridge University Press [the book]

[4] Pausas, J.G., Bradstock, R.A., Keith, D.A., Keeley, J.E. & GCTE Fire Network. 2004. Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85: 1085-1100. [pdf | jstor]

[5] Pausas J.G. & Verdú M. 2005. Plant persistence traits in fire-prone ecosystems of the Mediterranean Basin: A phylogenetic approach. Oikos 109: 196-202. [doi| pdf]

[6] 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]

[7] Moreira B. & Pausas J.G. 2012. Tanned or burned: The role of fire in shaping physical seed dormancy. PLoS ONE 7(12): e51523.  [doi | plos | pdf]