Posts Tagged ‘evolution’

Honoring Wallace

January 6th, 2024 No comments

Alfred Russel Wallace was a great naturalist who independently discovered natural selection; it is also recognized for his foundational work in evolutionary biogeography. While often overshadowed by Darwin’s popularity, Wallace’s scientific contributions remain significant and influential. James T. Costa’s recent book, ‘Radical by Nature’ (Princeton University Press, 2023), gives full credit and details on Wallace’s outstanding contribution to science.

Wallace laid out the first ideas of the evolution of species in 1855 with his Sarawak Law paper [1] when he was living (traveling) in the Malayan archipelago; the article likely inspired scientists of that time, including Darwin. Later, in 1858, Wallace produced another paper describing the mechanism of natural selection. But before sending it to the publisher, he sent the draft to Darwin for comments. Darwin was working on his book (On the origin of species) and realized that they both had independently arrived at the same conclusion (i.e., evolution by natural selection), so he was worried that Wallace was going to publish it first. Finally, with the help of Lyell & Hooker, they agreed to co-authored the famous paper that first described the mechanism of natural selection (Darwin & Wallace 1858 [2]). This paper, in fact, was not written together but included texts written independently from both to show that they arrived at the same idea (article edited by Lyell & Hooker). Wallace generously accepted Darwin’s priority. In fact, Wallace had started to write a book on the origin of species but put it down after knowing that Darwin was working on a similar book; instead, he dedicated his time to further exploring the Malayan archipelago (from 1854 to 1962). Darwin published his book On the origin of species in 1859, which described in great detail, and popularised the concept of natural selection.

Throughout his life, Wallace was not only a strong supporter of natural selection and Darwin’s ideas but also made significant contributions to other aspects of evolutionary thinking. For example, modern views on evolutionary biogeography, sexual selection, evolutionary mimicry, aposematism, and some aspects of the speciation process (such as allopatry, reinforcement and the Wallace effect) align more closely with Wallace’s ideas than with Darwin’s. Wallace also made numerous contributions to other fields, including anthropology, geology, physical geography, climatology, taxonomy, and systematics. He is often considered the first humanitarian scientist due to his strong support for social justice. He wrote papers on a variety of topics, including land reform, monetary reform, women’s rights, environmental conservation, and critiques of capitalism, militarism, and imperialism. One controversial aspect of his life was his belief in spiritualism, which made him believe that the human mind was not the product of natural selection!!. However, this should not be used to minimize the numerous contributions to evolutionary theory. And honoring Wallace does not need to detract from the recognition of Darwin’s major contributions.

So, Costa’s book is enjoyable and very welcome! 


[1] Wallace A.R. 1855. On the law which has regulated the introduction of new species. Annals and Magazine of Natural History 16, 184–196. [Sarawak Law paper]

[2] Darwin C.R. & Wallace A.R. 1858. On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. Proc. Linnean Soc: Zoo. 3: 45–62. [one of the most important papers in biology!]

The geological history of fire

December 19th, 2023 No comments

Fire is unique to Earth; it is a characteristic of our planet. As far as we know, no other planet has fire. Here is a video explaining the geological history of fire, including its relation to the evolution of pines and grasses. It concludes that “without fire, there would probably be no grasslands and the forest of the world would likely have a lot less diversity“. By PBS Eons.


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

The history of evolutionary fire ecology

August 9th, 2023 No comments

Evolutionary fire ecology is a relatively young discipline. The idea that fire acts as an evolutionary force contributing to shaping species traits started a century ago, but has not been widely recognized until very recently. Among the first to realize this force include E.B. Paulon, R.D. Guthrie, and E.V. Komarek in animals, and W.L. Jepson, W.W. Hough, T.M. Harris, P.V. Wells and R.W. Mutch in plants (all earlier than 1970). Our recent paper [1] is a tribute to these researchers that were ahead of their time in their evolutionary thinking.

Since them, evolutionary fire ecology has percolated very slowly into the mainstream ecology and evolutionary biology; in fact, this topic is still seldom mentioned in textbooks of ecology or evolution. Currently, there is plenty of evidence suggesting that we cannot understand the biodiversity of our planet without considering the key evolutionary role of fire [2]. We also provide thoughts on future direction of this discipline.

Traits mentioned in the article as potential adaptive to fire-prone environments are [1]:

  • Traits that enhance survival
    Resprouting traits (plant survival): Root crown, Lignotuber, Woody rhizomes, Epicormic resprouting, Sunken stem buds, Smoke-induced nutrient translocation
    Stem survival: Thick (outer) bark, Reduced flammability
  • Traits that enhance reproduction and recruitment
    Heat-released dormancy, Smoke-released dormancy, Seed traits enhancing seed survival, Serotiny, Fire-stimulated flowering, Increased flammability (chemically or structurally), Precocity (ie early reproduction), Elaiosomes (ant-dispersal)

In our search for early researchers with an evolutionary view of fire [1], we may have missed some women and non-English speaker researchers; if so, we would appreciate feedback on such omissions.

Figure 1. Two of the manzanitas from Jepson (1916, Madroño 1:3-12): Arctostaphylos glandulosa (left) and Arctostaphylos nummularia (right). The former is an obligate resprouter (note the basal burl) and the latter is an obligate seeder (note the even-aged cohort). Manzanitas were among the first plants that made scientists think about the role of fire in plant evolution.

[1] Pausas JG & Keeley JE. 2023. Evolutionary fire ecology: an historical account and future directions. BioScience. [doi | pdf]

[2] Keeley JE & Pausas JG. 2022. Evolutionary ecology of fire. Ann Rev Ecol, Evol, Syst 53: 203-225. [doi |eprint | pdf]

Evolutionary Ecology of Fire

November 4th, 2022 1 comment

Fire is an evolutionary pressure that shaped our biodiversity [1,2]. In a recent paper we summarized the current state of the art in this topic [3]. Fire has been an ecosystem process since plants colonized land over 400 million years ago [1]. Many diverse traits provide a fitness benefit following fires, and these adaptive traits vary with the fire regime [4]. Some of these traits enhance fire survival, while others promote recruitment in the postfire environment. Demonstrating that these traits are fire adaptations is challenging, since many arose early in the paleontological record, although increasingly better fossil records and phylogenetic analysis (figure below) make timing of these trait origins to fire more certain. Resprouting from the base of stems is the most widely distributed fire-adaptive trait, and it is likely to have evolved under a diversity of disturbance types. The origins of other traits like epicormic resprouting [5], lignotubers [6], serotiny [7], thick bark [8], fire-stimulated germination [9], and postfire flowering are more tightly linked to fire. Fire-adaptive traits occur in many environments: boreal and temperate forests, Mediterranean-type climate (MTC) shrublands, savannas, and grasslands. MTC ecosystems are distinct in that many taxa in different regions have lost the resprouting ability and depend solely on postfire recruitment for postfire recovery [10]. Overall, evolutionary fire ecology not only provides an understanding of the origin and history of our biota, it also sets the basis for the management of our ecosystems in a world undergoing fire-regime changes.

Time of origin (x-axis) of five different fire traits (different colors) for different lineages (y-axis) estimated from dated phylogenies. Bars expand the uncertainty of the time of origin (e.g., stem versus crown age). From [3]


[1] Pausas JG & Keeley JE 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doiOUP | pdf | post]

[2] He T, Lamont NB, Pausas JG 2019. Fire as s key driver of Earth’s biodiversity. Biol. Rev. 94:1983-2010. [doi | pdf

[3] Keeley JE & Pausas JG 2022. Evolutionary ecology of fire. Ann. Rev. Ecol. Evol. Syst. 53: 203-225. [doi | pdf] <- New paper

[4] Keeley JE, Pausas JG, Rundel PW, Bond WJ, Bradstock RA 2011. Fire as an evolutionary pressure shaping plant traits. Trends Pl. Sci. 16: 406-411. [doi | sciencedirect | trends | pdf | For managers]

[5] Pausas J.G. & Keeley J.E. 2017. Epicormic resprouting in fire-prone ecosystems. Trends Pl. Sci. 22: 1008-1015. [doi | sciencedirect | pdf] [post | Cover image]

[6] Pausas JG, Lamont BB, Paula S, Appezzato-da-Glória B & Fidelis A 2018. Unearthing belowground bud banks in fire-prone ecosystems. New Phyt. 217: 1435–1448. [doi | pdf | suppl. | BBB database]

[7] Lamont BB, Pausas JG, He T, Witkowski, ETF, Hanley ME. 2020. Fire as a selective agent for both serotiny and nonserotiny over space and time. Crit. Rev. Pl. Sci. 39:140-172. [doi | pdf | suppl.]

[8] Pausas JG 2015. Bark thickness and fire regime. Funct. Ecol. 29:317-327. [doi | pdf | suppl.] & Pausas JG 2017. Bark thickness and fire regime: another twist. New Phytol. 213: 13-15. [doi | wiley | pdf]

[9] Pausas JG & Lamont BB 2022. Fire-released seed dormancy – a global synthesis. Biol. Rev. 97: 1612-1639. [doi | pdf | supp. mat.]

[10] Pausas JG & Keeley JE 2014. Evolutionary ecology of resprouting and seeding in fire-prone ecosystems. New Phyt. 204: 55-65. [doi | wiley | pdf]

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)]

Serotiny: a review

June 9th, 2020 No comments

Here is a new review paper on serotiny in plants [1]. Serotiny refers to prolonged storage of seeds in woody structures (cones or fruits) on the mother plant for several growing seasons. This implies an accumulation of a canopy seed bank with seeds from different. Serotiny confer fitness benefits in environments with frequent crown-fires, as the heat of the fires opens the woody structures and thus seeds are dispersed in the post-fire bed (where resource are abundant and competition is low). There are other ways by which some cone/fruit can be opened (e.g., dry warm winds), but the number of seeds released and the chance of successful recruitment are much lower than in postfire conditions, and thus fire provides higher fitness benefits for serotinous plants than any other cue factor. This is why most serotinous plants occurs in ecosystems with frequent crown fires. Serotiny has been studied mainly in pines [2-4] and Proteaceae, but we know at least 12 families and more than 50 genera with serotinous species [1]. And there is a diversity of serotinous structures (cones and fruits) in different genera and families (Fig. 1).

Figure 1. A) Callitris (Actinostrobus) pyramidalis (Cupressaceae); B) Postfire Cupressus sempervirens (Cupressaceae); C) Cone of Pinus patula; D) Postfire cones of Pinus radiata; E) Folicles of Hakea cyclocarpa (Proteaceae); F) Hakea stenocarpa follicle; G) Follicles of H. platysperma; H) Xylomelum angustifolium (Proteaceae) follicle; I) Lambertia echinata (Proteaceae) follicle; J) Cone of Banksia lemanniana (Proteaceae); K) B. hookeriana burning; L) Postfire B. leptophylla cone; M) Cluster of capsules of Leptospermum spinescens (Myrtaceae), inset: seeds of Callistemon (Melaleuca) teretifolius; N) Elongated cluster of sessile capsules of Callistemon citrinus (Myrtaceae), + seeds of Callistemon (Melaleuca) teretifolius; O) Woody capsules of Eucalyptus todtiana (Myrtaceae), Inset: capsules of Angophora hispida (Myrtaceae); P) Spikes of Connomois parviflora (Restionaceae); Q) Protea burchellii (Proteaceae); R) fruits of Protea burchellii; S) Cone of Allocasuarina torulosus (Casuarinaceae); T) Cone of Isopogon trilobus (Proteaceae); U) Cones of Petrophile brevifolia; V) Seedlings from serotinous H. polyanthema, B. attenuata and B. hookeriana in litter microsite. For more details see [1].


[1] Lamont BB, Pausas JG, He T, Witkowski, ETF, Hanley ME. 2020. Fire as a selective agent for both serotiny and nonserotiny over space and time. Critical Rev. Plant Sci. [doi | pdf]

[2] Hernández-Serrano A, Verdú M, González-Martínez SC, Pausas JG. 2013. Fire structures pine serotiny at different scales. Am. J. Bot. 100: 2349-2356. [doi | amjbot | pdf | supp.]

[3] Hernández-Serrano A, Verdú M, Santos-Del-Blanco L, Climent J, González-Martínez SC, Pausas JG. 2014. Heritability and quantitative genetic divergence of serotiny, a fire-persistence plant trait. Ann. Bot. 114: 571-577. [doi | pdf | suppl.]

[4] Castellanos MC, González-Martínez S, Pausas JG. 2015. Field heritability of a plant adaptation to fire in heterogeneous landscapes. Mol. Ecol. 24, 5633-5642. [doi | pdf | suppl.]  


More on serotiny: Serotiny | Heritability of serotiny (1) | Heritability of serotiny (2): a molecular approach | Evolutionay fire ecology in pines 

Fire as a key driver of Earth’s biodiversity

July 12th, 2019 2 comments

Regions subject to regular fire have exceptionally high levels of species richness and endemism, and fire is likely a major driver of their diversity. In a recent paper [1] we reviewed the mechanisms that enable fire to act as a major ecological and evolutionary force that promotes and maintains biodiversity over different spatiotemporal scales. Specifically, we reviewed the different components of fire regime, the diversity through time (postfire), the intermediate disturbance hypothesis, the pyrodiversity-begets-biodiversity hypothesis, the fire-driven evolution and diversification, and the mutagenic effect of fire.

From an ecological perspective, the vegetation, topography and local weather conditions during a fire generate a landscape with spatial and temporal variation in fire-related patches (pyrodiversity), and these produce the biotic and environmental heterogeneity that drives biodiversity across local, regional and global scales scales [2]. We show that biodiversity should peak at moderately high levels of pyrodiversity. Overall species richness is typically greatest immediately after fire and declines monotonically over time, with postfire successional pathways dictated by animal habitat preferences and varying lifespans among resident plants.

From an evolutionary perspective, fire drives population turnover and diversification by promoting a wide range of adaptive responses to particular fire regimes [3,4]. In addition, fire and its byproducts may have direct mutagenic effects, contributing to the formation of novel genotypes that can lead to trait innovation. As a consequence of all these processes, the number of species in fire-prone lineages is often much higher than that in their non-fire-prone sister lineages.

Figure 1: The six components that define an individual fire event (in red the two core components). The fire regime arises from repeated patterns (means plus variance) over time of the properties of the components for each fire. For more details, see [1].

Figure 2. Relationship between species richness (S) of a reference area (community, landscape, region) and mean fire interval (a fire regime component). For a given fire regime, there is a mosaic of patches defined by different times since the last fire (represented by black circles) about the mean time interval (central circle). For more details, see [1].



[1] He T., Lamont B.B., Pausas J.G. (2019). Fire as s key driver of Earth’s biodiversity. Biological Reviews [doi | pdf]

[2] Pausas J.G. & Ribeiro E. 2017. Fire and plant diversity at the global scale. Global Ecol. Biogeogr. 26: 889–897. [doi | pdf | data & maps (figshare)]

[3] 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 | pdf]  

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

A diversity of Belowground Bud Banks (BBB) for resprouting

January 15th, 2018 No comments

Many plants are able to survive recurrent disturbance by resprouting from a bud bank. In fire prone ecosystems, plants must protect their buds from fire heat or perish. One way to protect them is by growing a thick insulating bark or sink the buds in the stem [1,2]. Another way is to locate the buds below ground, as soil is an excellent heat insulator (belowground bud bank or BBB). In fire-prone ecosystems, there is a diversity of ways by which plants successfully conceal their buds below ground that enable them to survive and resprout vigorously after fire [3]. There are at least six locations where belowground buds are stored [3]: roots, root crown, rhizomes, woody burls, fleshy swellings and belowground caudexes. These support many morphologically distinct organs (figure below). Considering their history and function, these organs may be divided into three groups:

(a) Those that originated in the early history of plants and that currently are widespread; they act as a resprouting source after a range of disturbances, not just fire. These include bud-bearing roots and root crowns.

(b) Those that also originated early and have spread mainly among ferns and monocots; they are often tied to seasonal stresses and have been highly successful under recurrent fire regimes. Theses include non-woody rhizomes and a wide range of fleshy underground swellings. They are characteristic of the geophyte growth forms occurring in many ecosystems, often tied to seasonal stresses; they have been highly successful under recurrent fire regimes.

(c) And those that originated later in history and are strongly tied to fire-prone ecosystems. These are woody rhizomes, lignotubers and xylopodia. They are characteristic of the geoxyle growth form.

Recognizing the diversity of BBBs is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.

Figure: Stylized diagrams of 16 belowground bud bank structures that enable plants to resprout postfire (highlighted in red). Broken horizontal line indicates position of soil surface. Structures characterized by woody tissues, in pink; fleshy tissues, in blue; and neither woody nor fleshy, in orange (usually highly sclerified primary tissues, fibrous or ‘wiry’). Shoots highlighted in apple green: stems with leaves, branched; leaves only, unbranched. Roots highlighted in olive green: triangular-shaped roots indicate a primary system, those arising directly from the bud-storing structures are adventitious. Drawings from [3]. From top left to bottom right:

· Xylopodium (in red) joined to tuberous root (in blue); Lignotuber; Root Crown; Woody Rhizome, here arising from a burl
· Bud-bearing lateral Root arising (here) from a burl (the root is not necessarily woody); Taproot Tuber; Bulb; Corm, with previous year’s corm still present
· Stem Tuber; Non-woody Fleshy Rhizome; Rhizophore (note buds are only supported by the oldest rhizophores); Adventitious Root Tuber
· Non-woody fibrous Rhizome with a monopodial arrangement leading to expansive clone; Non-woody fibrous Rhizome with sympodial arrangement leading to a caespitose habit; Stolons that produce new ramets postfire (note that it is not a BBB); Belowground Caudex

For details and a full description of each structure, see reference [3].

[1] Pausas J.G. 2017. Bark thickness and fire regime: another twist. New Phytologist 213: 13-15. [doi | wiley | pdf | post1, post2]

[2] Pausas J.G. & Keeley J.E. 2017. Epicormic resprouting in fire-prone ecosystems. Trends in Plant Science 22: 1008-1015. [doi | pdf | post ]

[3] Pausas J.G., Lamont B.B., Paula S., Appezzato-da-Glória B., Fidelis A. 2018. Unearthing belowground bud banks in fire-prone ecosystems. New Phytologist  [doi | pdf | suppl | BBB database]

More posts on resprouting


Postfire epicormic resprouting

September 22nd, 2017 No comments

Many plants resprout from basal buds after disturbance, and this is common in shrublands subjected to high-intensity fires [1]. However, resprouting after fire from epicormic (stem) buds is globally far less common. In a recent paper we review the ecology and evolution of this mechanism [2]. Many plants can generate epicormic shoots after light disturbances (e.g., browsing, drought, low intensity fires, insect defoliation, strong winds), but this does not mean they generally resprout epicormically after fire, as the heat of a fire may kill epicormic buds if they are not well protected (e.g., by a thick bark). The most well-known examples of epicormic resprouting are many species of eucalypts (Fig. 1A below), the cork oak (Quercus suber [3], Fig. 1B below), and Pinus canariensis ([4], Fig. 1C, D below). There are other pines and oaks that also resprout epicormically, and many species from savannas, especially those from the Brazilian savannas (cerrado) where many trees have a thick corky bark [5].

Epicormic resprouting has appeared in different lineages and on different continents and thus it is an example of convergent evolution in fire-prone ecosystems. It is an adaptation to a regime of frequent fires that affect tree crowns. It has probably been favoured where productivity is sufficient to maintain an arborescent growth form, fire intensity is sufficient to defoliate the tree canopy crown, and fire frequency is high (in conifers, too high for serotiny to be reliable) [2]. Given the high resilience of forest and woodlands dominated by epicormic resprouters, these species are good candidates for reforestation projects in fire-prone ecosystems [3].

Figure: Examples of postfire epicormic resprouting after a crown fire from very different lineages: (A) Eucalyptus diversicolor 18 months after fire in Western Australia. (B) Quercus suber woodland 1.5 years postfire in southern Portugal. (C) Pinus canariensis woodland a few years after fire; (D) epicormic resprouts of P. canariensis 3 months postfire. Photos by G. Wardell-Johnson (A); F.X. Catry (B) and J.G. Pausas (C, D), from [2].

[1] Pausas, J.G., Pratt, R.B., Keeley, J.E., Jacobsen, A.L., Ramirez, A.R., Vilagrosa, A., Paula, S., Kaneakua-Pia, I.N. & Davis, S.D. 2016. Towards understanding resprouting at the global scale. New Phytologist 209: 945-954. [doi | wiley | pdf | Notes S1-S4]

[2] Pausas J.G. & Keeley J.E. 2017. Epicormic resprouting in fire-prone ecosystems. Trends in Plant Science 22: xx-xx. [doi | pdf]

[3] Aronson J., Pereira J.S., Pausas J.G. (eds). 2009. Cork Oak Woodlands on the Edge: conservation, adaptive management, and restoration. Island Press, Washington DC. 315 pp. [The book]

[4] Pinus canariensis,

[5] Dantas V. & Pausas J.G. 2013. The lanky and the corky: fire-escape strategies in savanna woody species. Journal of Ecology 101 (5): 1265-1272. [doi | pdf | suppl.]

More information on: epicormic resprouting | cork oak | pines

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. 105: 289-297 [doi | wiley | pdf | brief for managers]

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

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


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

November 1st, 2016 No comments

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

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

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

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

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

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

[7] Smoke and human evolution,


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]


Evolutionary fire ecology in pines

April 1st, 2015 No comments

Fire is an ancient and recurrent disturbance factor in our planet and has been present since the origin of terrestrial plants [1]. However, demonstrating whether fire has acted as an evolutionary force is not an easy task [2]. In this context, the emerging discipline of evolutionary fire ecology aims to understand the role of wildfires in shaping biodiversity. In a recent review paper I summarize what we have learned on evolutionary fire ecology by studying the iconic genus Pinus [3]. I suggest that the study of pines has greatly increased our understanding of the role of fire as an evolutionary pressure on plants.

Macro-evolutionary studies of the genus Pinus provide the oldest current evidence of fire as an evolutionary pressure on plants and date back to ca. 125 Million years ago (Ma). Micro-evolutionary studies show that fire traits are variable within and among populations, and especially among populations subject to different fire regimes. In addition, there is increasing evidence of an inherited genetic basis to variability in fire traits. Added together, pines provide compelling evidence that fire can exert an evolutionary pressure on plants and thus shape our biodiversity. In addition, evolutionary fire ecology is providing insights to improve the management of our pine forests under changing conditions. The lessons learned from pines may guide research on the evolutionary ecology in other taxa.

Figure: Example of trait divergence among populations living under different fire regime. Serotiny (as % of closed cones) in populations living under frequent crown fires (red boxes) and in populations where crown-fires are rare (green boxes) for two pine species, Pinus halepensis (Allepo pine, left) and P. pinaster (maritime pine, right).

[1] Pausas, J.G. and Keeley, J.E. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doi | jstor | BioOne | pdf]

[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 in Plant Science 16: 406-411. [doi | sciencedirect | trends | pdf]

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


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


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]


Fire drives trait divergence: smoke-induced germination

April 3rd, 2014 No comments

There is an increasing evidence that recurrent fires are driving within species phenotypic variability, and that different fire regimes can generate trait divergence among populations [1]. For instance, populations of the annual species Helenium aromaticum (Asteraceae) growing under different fire histories in Chile have different seed traits in such a way that the anthropogenic increase in fire frequency selected for an increasing in seed pubescence [2]. In the Mediterranean Basin there is also evidence of phenotypic divergence among populations under different fire regimes: Ulex parviflorus (Fabaceae) plants living under high fire frequency are more flammable than those growing in sites that have not suffered fires [3-5]; Pinus halepensis and P. pinaster living under high crown-fire frequency have higher serotiny that those living in areas that rarely burn in crown fires [6]

A recent paper add further examples of this fire-driven trait divergence: Vandvik et al. show that smoke-induced germination is observed in populations of Calluna vulgaris (Ericaceae) from traditionally burnt coastal heathlands of Norway but it is lacking in populations from other habitats with infrequent fires [7]. The results are also consistent with the suggestion that smoke-induced germination is a fire adaptation [8-9].


Figure: Probability of germination of Calluna vulgaris in relation to time (days) since sowing for smoke-treated (pink) and control (grey) seeds, in coastal and inland heathlands of Norway. From Vandvik et al. 2014 [7].


[1] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193 (2). [doi | pdf | blog]

[2] Gómez-González S, Torres-Díaz C, Bustos-Schindler C, Gianoli E, 2011. Anthropogenic fire drives the evolution of seed traits. PNAS 108: 18743-18747. [doi blog]

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

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

[5] 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 | suppl. | data:dryad | blog]

[6] 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 (12): 2349-2356. [doi | amjbot | pdf | supp. | blog]

[7] Vandvik, V., J. P. Töpper, Z. Cook, M. I. Daws, E. Heegaard, I. E. Måren, and L. G. Velle. 2014. Management-driven evolution in a domesticated ecosystem. Biology Letters 10 (2): 20131082. [doi]

[8] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doi | jstor | BioOne | pdf | scribd | ppt slides | post]

[9] 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]


Proyecto VIRRA

February 28th, 2014 No comments

El proyecto “El papel del fuego en la Variabilidad Intraespecífica (fenotípica y genética) de plantas del matoRRAl mediterráneo (VIRRA)” finalizó hace unos meses. Aquí se puede ver un resumen y los principales productos de este proyecto: enlace.

Ulex parviforus_juli_sm

La aliaga (Ulex parviflorus) es una de las principales especies estudiadas en VIRRA [1, 2].

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

[2] Ulex born to burn (II): genetic basis of plant flammability,, 25/Jan/2014

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]


Fire and the evolution of pine life histories

August 15th, 2012 No comments

Many pines species are fire adapted. In 1998, JE Keeley & PH Zedler provided a seminal paper showing the various fire adaptations of pines, and the relation between the different adaptations and the different fire regimes [1]. Recent phylogenetic [2,3] and conceptual [4,5] advances in fire ecology have allowed to better understand the evolutionary role of fire in plants, and specifically in pines [2-6]. In a recent paper, JE Keeley provides a new review on the ecology and evolution of pine life histories [7]. Pinus originated ~150 Ma in the mid-Mesozoic Era and radiated across the northern continent of Laurasia during the Cretaceous period, when fire activity was high [3]. Pines have followed two evolutionary strategies interpreted as responses to competition by the newly emerging angiosperms: 1) The Strobus lineage mostly has radiated into stressful sites of low nutrient soils and extremes in cold or heat; ans 2) The Pinus (subgenus) lineage has radiated into fire-prone landscapes with diverse fire regimes. Based on the life history traits associated to fire, JE Keeley define four pine syndromes [7]: fire-avoiders (no fire-adapted; with thin bark), fire-toleraters (adapted to surface fires; with thick bark and self-pruning of dead branches; tall pines), fire-embracers (adapted to crown fires; with retention of dead branches and serotinous cones), and fire-refugia (with marked metapopulation dynamics) strategies.

Figure: Basal fire scar (a) and cross-section of pine with previous fires delineated (b) demonstrating fire survival after recurrent fires. Photos by JE Keeley from [7].

[1] Keeley J.E. & Zedler P.H. 1998. Evolution of life histories in Pinus. In: Ecology and biogeography of Pinus (ed. Richardson DM). Cambridge University Press Cambridge (UK), pp. 219-250.

[2] Schwilk D.W. & Ackerly D.D. 2001. Flammability and serotiny as strategies: correlated evolution in pines. Oikos, 94, 326-336. [doi]

[3] 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 ]

[4] Pausas, J. G. and J. E. Keeley. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doi | jstor | pdf]

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

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

[7] Keeley J.E. 2012. Ecology and evolution of pine life histories. Ann. For. Sci., 69, 445–453. [doi]


The age of fire adaptations

February 20th, 2012 No comments

How old are wildfires? Probably as old as terrestrial ecosystems [1]. The origin of fire is tied to the origin of land plants, which are responsible for two of the three elements essential to the existence of fire: oxygen and fuel. The third element, a heat source, has probably been available throughout the history of the planet (mainly through lightning). There is charcoal evidencs of fires already in the Silurian (440 Ma). However, the existence of fire does not necessarily mean that fire was playing an evolutionary role at that time. So when did fire start to play an evolutionary role generating fire adaptations [2, 3]? By mapping fire adaptation onto a dated phylogeny of Pinaceae, we recently demonstrated [4] that at least, and for this family, fire was an agent of natural selection since about 90-125 Ma! This is far back from what was known until now [4]. At this time, fire-protective thick barks were originated in Pinus species as response to surface fires. With increasing fire intensity, thicker barks and serotiny appeared by 70-90 Ma. These innovations appear at the same time as the Earth’s paleoatmosphere experienced elevated oxygen levels that led to high burn probabilities (mid-Cretaceous). That is, the fiery environments of the Cretaceous strongly influenced trait evolution in Pinus. Whether fire had an evolutionary role prior to this is a challance for future research.

Fotos: In many pines, the thick bark and the discontinuity between the canopy and the understory (self-pruning) allows survival after surface fires (left: Pinus nigra, eastern Spain). Serotinous cones allow a quick seed regeneation after crown fire (right: P. halepensis, eastern Spain). Photos: JG Pausas

[1] Pausas, J. G. and J. E. Keeley. 2009. A burning story: The role of fire in the history of life. Bioscience 59: 593-601. [doijstor | BioOne | pdf]

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

[3] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193:301-303. [doiwileypdf]

[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 Phytologist 194: 751-759 [doi | wileypdf]

Fire drive plant evolution

December 7th, 2011 No comments

Considering fire as evolutionary pressure driving evolution has traditional been neglected, and only now is becoming a topic of research [1-3]. The role of fire as an evolutionary pressure can be elucidated using both macro- and micro-evolutionary approaches. While the micro-evolutionary approach searches for trait divergences in different current selective environments, the macro-evolutionary approach uses dated phylogenies to trace the evolution of traits over long time scales (My). In a previous post [3] we mentioned an example of the macro-evolutionary approach. In a recent paper, S. Goméz-Gonzalez and collaborators [5] provided, for the first time, a clear example of the micro-evolutionary approach to demonstrate natural selection driven by fire.  They presented compelling evidence that the novel anthropogenic fires affecting the Chilean matorral shaped seed traits on a native annual plants (Helenium aromaticum). By studying populations growing on sites with different recent fire histories, they showed that increasing fire frequency selects for increasing seed pubescence (directional selection): a trait that was proven to be heritable and that increased fitness under experimental heat treatments. This paper was also presented in a special session at the MEDECOS Conference [3].

Figure: Habitat of Helenium aromaticum in central Chile [5]

[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doipdfpost]

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 [doitrends |pdf]

[3] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193: [pdf]

[4] Pausas J.G. 2011. Australia born to burn – phylogenetic evidences. URL:, 18/03/2011

[5] Gómez-González S, Torres-Díaz C, Bustos-Schindler C, Gianoli E, 2011. Anthropogenic fire drives the evolution of seed traits. PNAS 108: 18743-18747. [doi]

MEDECOS XII (2011): fire and evolution

November 26th, 2011 No comments

Mediterranean Ecosystem (MEDECOS) conferences are held every 3–5 yrs, rotating venues through all five Mediterranean-type climate (MTC) regions of the world. The first meeting was held in Valdivia (Chile) in 1971. The last MEDECOS was held in Los Angeles (University of California, September 6-9, 2011, [1]), and about 300 scientist from the different MTC regions got together and presented their research on the different aspects of the ecology in mediterranean ecosystems. In this MEDECOS, fire was an important topic, it was explicit in the title at least in the following 5 special sessions:

Fire as an evolutionary pressure shaping plant traits (6th Sept, Pausas & Schwilk)
– Fire management at the wildland-urban interface (7th Sept, Keeley)
– Global change and fire (7th Sept, van Mantgem)
– Fire ecology in Mediterranean woodlands ans shrublands (8th Sept, O’Leary)
– Fire management (9th Sept, Fotheringham)

Dylan Schwilk and I organised the first one which highlighted several key aspects on the role of fire in plant evolution: First, there is good evidence for vegetation-fire regime feedbacks at different spatial and temporal scales, in such a way that plant flammability is a major driver of plant evolution and vegetation distribution. Second, the evidence that fire acts as a selective force is apparent on both micro- and macro-evolutionary scales, suggesting that fire shapes plant traits and generates fire adaptations. And third, that fire is a complex selective pressure – plants adapt to (and, in turn, influence) particular fire regimes rather than fire in the abstract. This is an exciting time for fire ecologists, as fire is now recognized as fundamental for many ecological and evolutionary processes; the coming macro- and micro- evolutionary studies will certainly reinforce many of the ideas drawn during the meeting [2]. The details of this session, including slides of the talks and a summary of the session [2], are now available here .

[1 ] MEDECOS 2011:  web | program | abstracts | final resolution

[2] Pausas, J. G. and D. W. Schwilk. 2012. Fire and plant evolution. New Phytologist 193 (2). [doi | pdf]

Ulex born to burn

November 9th, 2011 No comments

Recurrent fires are a strong evolutionary pressure shaping plants [1,2]. It has been hypothesized that in fire prone-ecosystems, natural selection has favoured the development of traits that enhance flammability [3]. Consistent with this idea, in a recent study [4] we found that Ulex parviflorus (Fabaceae) populations that inhabit in recurrently burn areas (HiFi populations) are more flammable than populations of this species growing in old-fields where the recruitment was independent of fire (NoFi populations). That is, HiFi plants ignite quicker, burn slower, release more heat and have higher bulk density than NoFi plants. Thus, it appears that repeated fires select for individuals with higher flammability, and thus driving trait divergence among populations living in different fire regimes. These results provide some field support for the ‘kill thy neighbour’ hypothesis [3], but they also highlighted the need for heritability studies to unambiguously demonstrate natural selection driven by fire. This study together with other studies recently commented in this blog [5, 6] are placing flammability as a fundamental trait in plant evolution.

Figure: Flammability experiments using an epiradiator [4].


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

[2] Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience [doi | jstore | pdf]

[3] Bond, W. J. and J. J. Midgley. 1995. Kill thy neighbour: an individualistic argument for the evolution of flammability. Oikos 73:79-85.

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

[4′] Pausas J.G. & Moreira B. 2012. Flammability as a biological concept. New Phytologist 194: 610-613. [doipdf]

[5] Pausas JG. 2011. Australia born-to-burn: a phylogenetic approach., 18/March/2011 [link]

[6] Pausas JG. 2011. Fire and evolution: Cretaceous fires and the spread of angiosperms., 9/Sept/2011 [link]

Fire and plants: adaptations and exaptations

May 18th, 2011 No comments

There are still people believing that wildfires are a catastrophic disturbance to ecosystems, and that are the product of humans. However there is an increasing evidence from paleoecological records and from phylogenetic analyzes suggesting that fire is a very old process in the history of life, dating back to the origin of land plants [1, 2, 3]. As a consequence many plants have evolved in the presence of recurrent wildfires and acquired adaptive traits to persist and reproduce in those conditions. Examples of these traits are the resprouting ability, germination by head or smoke, and serotiny; all of these confer fitness advantage in fire-prone ecosystems. However, plants are not adapted to fire per se but to fire regimes. Species that exhibit traits adaptive under a particular fire regime can be threatened when that regime changes, like the recent human-induced fire regime changes (e.g., increasing or decreasing fire frequency or severity in relation to the historic fire regime).

In a recent paper, Keeley et al. [4] proposed five scenarios of change in a trait state (Figure 1). An adaptive trait might not change through time regardless of the selective environment (scenarios 1 and 2 in Figure 1). Such traits cannot be described as adaptations to the current selective (fire-prone) environment as there is no evidence that natural selection shaped this trait. Other adaptive traits that were shaped by natural selection under a previous evolutionary pressure, but not under the current (fire-prone) environment (scenario 3 in Figure 1) would be adaptations to previous evolutionary pressures and exaptations to the current (fire) environment [4, 5]. Fire adaptations are those adaptive traits in which natural selection is acting under the current fire-prone environment to shape the trait, and it is independent of how long this pressure has been present (scenarios 4 and 5 in Figure 1). For instance, there are clear examples of lineages that resprout after fire, but their origin and evolution is hardly liked to fire. However the most plausible scenario of lineages that resprouting from lignotubers is the number 4 in Fig. 1 (old origin of resprouting reshaped by current recurrent fires). Similarly serotiny and thick barks are traits that has been reshaped by natural selection under the framework of recurrent fires and thus they also fit under the concept of adaptation to fire (scenario 4 or 5 in Fig. 1).

[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doi pdfpostslides]

[2] Pausas J.G. 2011. Australia born to burn – phylogenetic evidences. URL:, 18/03/2011.

[3] Pausas J.G. 2010. Fire and evolution: Cretaceous fires and the spread of angiosperms. URL:, 9/Sep/2010.

[4] 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]

[5] Endler J.A. (1986) Natural selection in the wild. Princeton University Press.

Figure. 1. Five possible evolutionary scenarios of change in a trait state along the evolutionary time (simplified). For each scenario, different line types are periods under different dominant evolutionary pressures (e.g., the continuous line represents a period in which fire acted as an evolutionary pressure; dashed line the period with a different previous selective environment). The scenarios are:
1) and 2) No change along the time axis and no sign of natural selection (no adaptation to fire);
3) trait shaped during the first evolutionary pressure, but no change (with persistence of the state of the trait) during the second evolutionary pressure; natural selection acted during the first period only (no adaptation to fire, but exaptation);
4) and 5) Trait shaped during the whole period; natural selection acts during the whole period even if the dominant evolutionary pressure changed (adaptation to fire).
From Keeley et al. (2011, [4]).

Australia born to burn – phylogenetic evidence

March 18th, 2011 1 comment

Traditionally wildfires were considered a disturbance linked to the recent history of the Quaternary, and specially linked to the humans. However, evidence are accumulating about the ancient role of wildfires in terrestrial ecosystems [1]. In Australia, the flammable continent, the current believe is that fires started to be important during the onset of seasonal aridity in the Miocene (25 Ma). However, two recent and independent papers demonstrate, using phylogenetic techniques, that fire-dependent traits appeared about 60 Ma ago (early Paleocene), implying that fire was already an effective agent of selection by then. Crisp et al [2] studied the Myrtaceae family and showed that post-fire epicormic resprouting (typical of many eucalypts) is an ancient trait linked to the flammable sclerophyll biomes originated about 60-62 Ma. He et al. [3] studied the Banksia genus (Proteaceae) and showed that serotiny (fire dependent dispersal; figure below) and dead floret retention around the cones (enhanced flammability around serotinous cones) co-originate with the first appearance of Banksia 60.8 Ma ago. The coincidence of the two independent papers, using two different taxa (Myrtaceae and Banksia) is amazing, and clearly suggests that fire was a selective force in Australia during the Paleocene.  These papers are part of the accumulating research on the prominent and ancient role of fire in shaping plant species and biodiversity [1, 4 ].

[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doipdfpost slides]

[2] Crisp MD, Burrows GE, Cook LG, Thornhill AH, Bowman DMJS. 2011. Flammable biomes dominated by eucalypts originated at the Cretaceous-Palaeogene boundary. Nature Communications 2: 193. [doi]

[3] He T, Lamont BB, Downes KS. 2011. Banksia born to burn. New Phytol. [doi]

[4] Bond, W. J. and Scott, A. C. 2010. Fire and the spread of flowering plants in the Cretaceous. New Phytol. 188: 1137–1150 [post]

Figure: Banksia cone opened by the fire to release seeds (serotiny).

Fire and evolution: Cretaceous fires and the spread of angiosperms

September 9th, 2010 1 comment

Recently we have highlighted the importance of wildfires in the evolution of plants in many ecosystems worldwide [1 | previous post]. In this line, a recent paper by Bond & Scott suggest that the spread of angiosperms in the Cretaceous (145-65 Ma) was promoted by the development of novel fire regimes linked to the evolution of novel, highly productive (and flammable) plants. They suggest that Creatceous angiosperms were similar to current ruderal (weedy) species, i.e., short, with high maximum photosynthetic rates, rapid reproduction and small seeds. This fast-growing angiosperms would not only compete with regenerating gymnosperms, but would also rapidly accumulate fuel. More fuel would promote more frequent fires, which would help to maintain open habitats in which rapid growth traits of angiosperms would be most favoured, promoting rapid fuel accumulation. The authors emphasize the similitude of this “angiosperm–fire cycle” with  the grass fire-cycle that helped to spread C4 grasses in the Miocene (c. 8 Ma) [3] and with the grass fire-cycle replacing forests by invasive grasses in the modern world [4]. This would also imply that forest was slow to develop until the Eocene, when fire activity dropped to very low levels. This hypothesis could also help to explain the ancient origin of some fire traits like resprouting and the abundance and phylogenetically widespread examples of species with smoke-stimulated germination [1, 5]. In conclusion I think this is a nice and stimulating contribution to the evolution of angiosperms.


[1] Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doi | pdfpost | slides]

[2] Bond, W. J. and Scott, A. C. 2010. Fire and the spread of flowering plants in the Cretaceous. New Phytol. 188: 1137–1150 [doi]

[3] Keeley, J. E. and Rundel, P. W. 2005. Fire and the Miocene expansion of C4 grasslands. Ecol. Lett. 8: 1-8.

[4] D’Antonio, C. M. and Vitousek, P. M. 1992. Biological invasions by exotic grasses, the grass/fire cycle and global change. Annu. Rev. Ecol. Syst. 23: 63-87.

[5] 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]

Fuego y evolución en el Mediterráneo

August 1st, 2010 2 comments

En pleno verano, y como cada año, arden unas cuantas hectáreas de nuestros paisajes. En los ecosistemas mediterráneos esto lleva ocurriendo desde hace muchos años, incluso antes de la llegada de los humanos. El fuego y los incendios forestales no es ni mucho menos un invento de los pirómanos, sino que están en la naturaleza desde casi siempre, y por eso hay muchas plantas adaptadas a vivir en zonas con fuegos recurrentes (igual que hay plantas que pueden sobrevivir el pastoreo recurrente). Lo que ha hecho los humanos es modificar el régimen de incendios, aumentado o disminuyendo su frecuencia e intensidad.

A menudo los incendios se ven como una cosa “mala” para la naturaleza; en un artículo de divulgación en la revista “Investigación y Ciencia” (Agosto 2010) damos otra visión del fuego, y hacemos un repaso del papel de este en la evolución de las especies,  la otra cara del fuego. En nuestros ecosistemas, el fuego es una parte integral de los procesos ecológicos, y a lo largo de la historia ha ido moldeando las especies, las comunidades y los paisajes. Sin duda hay ciertos regímenes de incendios que son naturales y característicos de ciertos ecosistemas, y parte de la diversidad de nuestros ecosistemas se explica por la existencia reiterada y predecible de incendios. Sin embargo, también es cierto que hay zonas que están sufriendo regímenes de incendios fuera del rango natural y con graves consecuencias ecológicas. El objetivo de la gestión forestal no debería ser eliminar los incendios, ya que es prácticamente imposible, además de poco natural. Por el contrario, deberían asumirse ciertos regímenes sostenibles de incendios, y aprender a convivir con ellos.

Pausas, J.G. 2010. Fuego y evolución en el Mediterráneo. Investigación y Ciencia, 407 (Agosto): 56-63. [PDF: IyC | jgpausas]

Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience, 59: 593-601. [caliber] [pdf]

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). [link] [pdf]

Pausas J.G., Llovet J., Rodrigo A., Vallejo R 2008. Are wildfires a disaster in the Mediterranean basin? – A review. Intern. Journal of Wildland Fires, 17: 713-723. [pdf] [IJWF CSIRO] [doi]


Fire in the roots of humans

January 19th, 2010 No comments

One key difference between animals and humans is the use of fire; in fact, during the evolution, fire made us humans. For instance, cooking implied higher food energy, as well as an increased the diversity of available food (detoxifying effects of heating, etc…). Furthermore, cooking implied a delay in food consumption, which required the development of social abilities for the distribution of tasks within the group (e.g., collection, accumulation, cooking, defense, even stealing). These factors are thought to have prompted the evolution of large brains and bodies, small teeth, modern limb proportions, and other human traits, including many social aspects of human-associated behavior. However, the moment in which humans started to use fire is still debated. It is often believed that the rise of Homo erectus from its more primitive ancestors was fueled by the ability use fire.

Although the use and control of fire is a human trait, a recent study has demonstrated that chimpanzees have the ability to understand wildfires and predict their behavior (Pruetz & LaDuke 2010). Chimps calmly observed wildfires around them, predict their behaviour and move accordantly without any stress or fear. This suggest that the conceptualization of fire may be a old trait, in the hominids group.

To what extent current humans are losing this trait is another debate, but we may be better off at managing our fire-prone landscapes by learning from chimps!


  • Pausas J.G. & Keeley J.E. 2009. A burning story: The role of fire in the history of life. BioScience 59: 593-601 [doi] [pdf]
  • Pruetz JD & TC LaDuke 2010. Reaction to fire by savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal: Conceptualization of  fire behavior and the case for a chimpanzee model. Am J Phy Anthropol (in press) [doi]
  • Wrangham RW, Jones JH, Laden G, Pilbeam D, Conklin-Brittain NL. 1999. The raw and the stolen: Cooking and the ecology of human origins. Current Anthropol 40: 567–590.
  • Control of fire by early humans [Wikipedia]


Fires and megafauna: the answer is in the dung

November 20th, 2009 No comments

We recently proposed (Aug/2009) that “The spread of humans, perhaps concomitant with climatic changes, contributed to the mass disappearance of megafauna such as mammoths and other large herbivores (i.e., the Pleistocene-Holocene extinction); this extinction would also have resulted in fuel buildup and the consequent change in fire activity, as suggested by the contemporary effects of megaherbivores.” (Pausas & Keeley 2009; see also Flannery 1994).

Today (20/Nov/2009), in a paper in Science, Gill et al. demonstrate this link between megafauna extinction and fire activity by studying the fossils spores of a coprophilous fungi (Sporormiella) from a lake sediments in Indiana, North America, together with charcoal and pollen from the same sediments (ca. 14,000 years of history). Sporormiella produced spores in the dung of large herbivores, and the amount of spores can be considered an index of the abundance (or biomass) of herbivores vertebrates. The authors demonstrate that the decline in megafauna is associated to an increased fire activity. This also make us to think about the idea of Pleistocene rewilding (Donland et al. 2005) for fuel control and fire reduction.

These papers and other recent ones are putting fire ecology in the front-line of ecology as they demonstrate the strong influence of fire in shaping nature; they emphasize the importance of fire for understanding present, past and future ecosystems as well as global processes. For instance, Nevle & Bird (2008) recently demonstrated that the massive reduction of American natives by the European invasion of America, drastically reduced fire activity and the consequent increase in forest (carbon sequestration) contributed to the ca 2% global reduction in atmospheric CO2. Despite the importance of fire in the global context, they are still poorly represented in global models (Bowman et al. 2009).


Bowman D.M.J.S. et al. (2009). Fire in the Earth System. Science, 324, 481-484.

Donlan C.J. et al. (2006). Pleistocene Rewilding: an optimistic agenda for 21st century conservation. The American Naturalist 168: 660-681

Flannery T. (1994). The Future Eaters: An Ecological History of the Australasian Lands and People. Reed Press, Port Melbourne, Australia.

Gill JL et al. (2009). Pleistocene Megafaunal Collapse, Novel Plant Communities, and Enhanced Fire Regimes in North America. Science 326: 1100-1103.

Nevle R.J. & Bird D.K. (2008). Effects of syn-pandemic fire reduction and reforestation in the tropical Americas on atmospheric CO2 during European conquest. Palaeogeography, Palaeoclimatology, Palaeoecology 264: 25-38.

Pausas J.G. & Keeley J.E. (2009). A burning story: The role of fire in the history of life. BioScience 59: 593-601. [caliber] [BioOne] [doi] [pdf]

Convergent evolution

November 8th, 2009 No comments

Images from two different tree species (A and B), from different Families (and different Orders), taken in different continents…


The thick bark offers protection to fire and thus these species are both adapted to live in fire-prone ecosystems [1].

Can you guess the species name of A and B?    [ Answer: A | B ]


[1] See also: The ecology of bark thickness | The ecology of bark thickness (2): another twist


New paper: A burning story

October 11th, 2009 No comments

Ecologists, biogeographers, and paleobotanists have long thought that climate and soils controlled the distribution of ecosystems, with the role of fire getting only limited appreciation. Here we review evidence from different disciplines demonstrating that wildfire appeared concomitant with the origin of terrestrial plants and played an important role throughout the history of life. The importance of fire has waxed and waned in association with changes in climate and paleoatmospheric conditions. Well before the emergence of humans on Earth, fire played a key role in the origins of plant adaptations as well as in the distribution of ecosystems. Humans initiated a new stage in ecosystem fire, using it to make the Earth more suited to their lifestyle. However, as human populations have expanded their use of fire, their actions have come to dominate some ecosystems and change natural processes in ways that threaten the sustainability of some landscapes.

Pausas J.G. & Keeley J.E. 2009. A Burning Story: The role of fire in the history of life. BioScience 59: 593-601. [doicaliber | pdf | ppt slides | scribd]


Temporal position of key moments in the history of life (time in logarithmic scale) in relation to fire.