Opinion

Fire as an evolutionary pressure

shaping plant traits
Jon E. Keeley1,2, Juli G. Pausas3, Philip W. Rundel2,
William J. Bond4 and Ross A. Bradstock5
1

US Geological Survey, Sequoia-Kings Canyon Field Station, Three Rivers, CA 93271, USA
Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095, USA
3
Centro de Investigaciones sobre Desertificacion of the Spanish National Research Council (CIDE - CSIC), IVIA campus, 46113
Montcada, Valencia, Spain
4
Botany Department, University of Cape Town, Private Bag, Rondebosch 7701, South Africa
5
Centre for Environmental Risk Management of Bushfires, University of Wollongong, Wollongong, NSW 2522, Australia
2

Traits, such as resprouting, serotiny and germination by

heat and smoke, are adaptive in fire-prone environments. However, plants are not adapted to fire per se
but to fire regimes. Species can be threatened when
humans alter the regime, often by increasing or decreasing fire frequency. Fire-adaptive traits are potentially the
result of different evolutionary pathways. Distinguishing
between traits that are adaptations originating in response to fire or exaptations originating in response
to other factors might not always be possible. However,
fire has been a factor throughout the history of landplant evolution and is not strictly a Neogene phenomenon. Mesozoic fossils show evidence of fire-adaptive
traits and, in some lineages, these might have persisted
to the present as fire adaptations.
Adaptation to fire
In a recent publication [1], S. Don Bradshaw et al. raised
concerns about the widespread application of prescription
burning in hotspots of biodiversity, and we share those
concerns that too frequent prescription burning can negatively affect biodiversity and ecosystem processes. However, their paper used this issue inappropriately as a
springboard for speculating on the evolution of plant traits,
confusing issues of adaptive traits and adaptations, and
missing the fact that different fire regimes can select for
very different plant traits. No species is fire adapted but
rather is adapted to a particular fire regime, which, among
other things, includes fire frequency, fire intensity and
patterns of fuel consumption [2]. Species that exhibit traits
that are adaptive under a particular fire regime can be
threatened when that regime changes. For example, many
of the species-rich Mediterranean-type climate (MTC)
shrublands are resilient to periodic high-intensity crown
fires at intervals of several decades or more. However,
when the fire frequency increases, species can be rapidly
lost [3,4]. Bradshaw et al. [1] incorrectly equate adaptation
to fire as adaptation to frequent fires and are not justified
in inferring that frequent prescription burning is an adaptationist view. Here, we discuss those traits with apCorresponding author: Keeley, J.E. (jon_keeley@usgs.gov).

406

parent adaptive value in fire-prone environments and the

extent to which we can demonstrate that they are adaptations to fire. We also address the question of what fireadaptive traits and fire adaptations can tell us about fire
management.
Adaptive traits
Adaptive traits are those that provide a fitness advantage
in a given environment. There are many plant traits that
are of adaptive value in the face of recurrent fire and these
vary markedly with fire regime. For example, North American conifers (Coniferophyta) subject to frequent lightningignited fires have thick bark, which functions to protect the
living tissues from heat damage during surface fires; they
also self-prune lower dead branches, which ensures a gap
in fuels between the dead surface litter and live canopy.
Thick bark and self-pruning are adaptive in this surface
fire regime. At higher latitudes as productivity declines,
lodgepole and jackpine forests lack the potential for growth
sufficient to keep the canopy away from surface fuels,
leading to a crown fire regime; traits change to thin bark,
retention of lower dead branches, and serotinous cones
that synchronously release seeds following such standreplacing fires [5,6]. High-intensity crown fires are not
stand replacing in Australian eucalypt forests as most of
the trees, big and small, resprout epicormically along the
length of the bole, replacing the tree canopy within a year
of such fires. In MTC shrublands subject to crown fires,
many species restrict seedling recruitment to the immediate post-fire years, arising from previously dormant soilstored seed banks or synchronous seed release from serotinous fruits [2]. These traits are adaptive in fire-prone
environments and are key to providing resilience to specific
fire regimes.
Adaptive traits as adaptations to fire
A key evolutionary question about fire-adaptive traits is
whether they represent fire adaptations, defined as traits
that originated in response to fire. Fire-adaptive traits that
originated in response to some other environmental factor,
and then were appropriated for their value in fire-prone
landscapes, are exaptations [7]. Distinguishing between

1360-1385/$ see front matter . Published by Elsevier Ltd. doi:10.1016/j.tplants.2011.04.002 Trends in Plant Science, August 2011, Vol. 16, No. 8

Opinion

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Box 1. Adaptations and exaptations

adaptations and exaptations is a high bar that is difficult to

demonstrate as there is not a simple dichotomy between
paths leading to adaptations and exaptations (Box 1).
There are several flaws in Bradshaw et al.s analysis [1]
of fire adaptations. One is the misconception that fire is a
recent Neogene [23 million years ago (Ma) to the present]
phenomenon; however, fire has been part of Earth system
since the Silurian origin of land plants 443 Ma [8]. It was
globally important through much of the Mesozoic (25166
Ma) [911] and, by the Late Cretaceous (10066 Ma), some
taxa were already specialized for fire-prone environments
[12,13]. In the highly diverse Australian Eucalyptus, epicormic resprouting regenerates forests rapidly after highintensity crown fires and phylogenetic studies support an
early Tertiary origin (62 Ma) [14].
Resprouting
Another misconception in [1] is the belief that vegetative
resprouting cannot be an adaptation to fire because it is
found in non-fire-prone habitats. There are many forms of
resprouting and it is expected that different resprouting
types might have appeared in different lineages in response to different evolutionary pressures. No one has
ever argued that all instances of resprouting are adaptations to fire; however, it is still an open question as to
whether post-fire resprouting originated in response to fire
in some lineages [2]. For example, resprouting is coupled
with a lignotuber in various lineages (Figure 1), and the
tight coupling of this ontogenetic trait with fire-prone
environments [15] suggests that it is a fire adaptation.

ment owing to anthropogenic impacts, but this adaptive trait might

not have arisen in response to fire (scenarios 1, 2 or 3). By contrast, it
is plausible that some fern lineages have had a long association with
fire and resprouting might follow scenario 5 [2]. MTC shrub lineages
resprouting from lignotubers might best be interpreted in terms of
scenario 4a, an old origin of resprouting in response to diverse factors
but reshaped later by fire.[()TD$FIG]

1
2
3
4a
4b

Trait state

An adaptive trait enhances fitness in a defined habitat and adaptation

is the process of acquiring adaptive value by the natural selection of
new variants [28,29]. Exaptation defines an adaptive trait with a
particular contemporary function, but one that was previously shaped
by natural selection for another function [7,25]. However, exaptations
can become adaptations when natural selection acts and reshapes the
trait.
We distinguish five scenarios of change in a trait state (Figure I).
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
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) would be adaptations to
previous evolutionary pressures and exaptations to the current
environment.
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 some cases (scenario 4b) selective pressures
can cause traits to revert to a former state and, when conditions
change, fire might replace the former selective pressure.
It is crucial to note that our interpretation of adaptations and
exaptations depends on current understanding of the entire evolutionary scenario. If the knowledge base only extends back to the
second half of the time period in Figure I, it is not possible to
distinguish between different scenarios. This problem is inherent in
retracing the history of most traits.
Plants can have traits of adaptive value under specific fire regimes,
but their origin can follow different evolutionary scenarios. For
instance, one might find that resprouting is highly adaptive in some
tropical forest trees that are suddenly cast into a fire-prone environ-

Time (evolution)
TRENDS in Plant Science

Figure I. Evolutionary scenarios of change in a trait state (continuous lines can

be interpreted as representing a period in which fire acted as an evolutionary
pressure and dashed lines a period with a different selective environment).

Serotiny
Serotiny as an adaptation to fire has also been questioned
by [1] because it probably post-dates onset of Tertiary
drying and might be an evolutionary response to low soil
fertility. However, phylogenetic studies in the Australian
genus Banksia conclude that fire-adaptive traits, and serotiny in particular, originated much earlier and they place
their origins in the Paleocene (60 Ma) [16].Bradshaw et al.s
[1] linking of serotiny to infertile soils follows Stephen D.
Hoppers [17] contention that low fertility soils, such as
those widespread in Western Australia and South Africa,
are derived from ancient Cretaceous substrates. Hopper
argues that this ancient association with infertile soils is
the primary factor driving the evolution of plant traits in
these shrublands. However, plants persist in a multivariate environment stressed by not only soil fertility, but also
climate and potentially fire, not to mention biotic interactions. Indeed, low fertility soils select for traits such as
small sclerophyll leaves and provide well-drained substrates, both of which increase the likelihood of fires. In
our view, evolution on these substrates is driven by a
combination of geology, climate and fire [2]. Singling out
one of these factors as the only determinate of trait evolution handicaps our ability to understand plant evolution.
Serotiny is tied to crown fire regimes and, in the absence
of fire, there is relatively little successful recruitment, making it clearly of adaptive value in these systems. However,
demonstrating that serotiny is an adaptation that evolved in
response to fire, is another matter. Bradshaw et al. [1]
conclude that because serotiny is concentrated in the two
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()TD$FIG][ Opinion

Trends in Plant Science August 2011, Vol. 16, No. 8

TRENDS in Plant Science

Figure 1. Resprouting after fire from a lignotuber in the California chaparral shrub Adenostoma fasciculatum. It is one of a wide diversity of resprouting modes in the plant
kingdom. These basal burls are an ontogenetic trait that forms early in development and, in this respect, differ from swollen basal burls initiated in response to cutting or
burning observed in many woody plants.

southern hemisphere MTC regions characterized by low soil

fertility, it arose in response to soil infertility. However, they
conflate two separate issues: the reproductive strategy of a
single post-fire pulse of seedling recruitment and aerial
versus soil seed storage.
Sorting out the adaptive value of serotiny requires
consideration of ultimate versus proximal causation. Four
of the five MTC regions have species that restrict seedling
recruitment to a post-fire pulse; the reason for this is that
fires are a predictable ecosystem process and the removal
of the dense shrub canopy by fires provides superior
resources for the recruitment of some species. The proximal mechanism of how species do this varies with the
region; in two of the MTC regions, dormant seeds are
408

stored in the soil and serotiny is rare, whereas in two other

regions, many dominant shrubs store seeds in serotinous
fruits in the shrub canopy. We argue that low soil fertility
in South African fynbos and Western Australian heathlands has selected against soil storage in favor of aerial
storage because high-nutrient seeds are subject to more
intense predation when exposed on the soil surface
(Figure 2). Higher fertility soils in northern hemisphere
MTC shrublands of California and the Mediterranean
Basin allow shrub taxa to capitalize on the advantages
of soil-stored seed banks, which includes a capacity for
accumulating larger seed banks and a greater resilience to
unpredictably long fire-free intervals. The one MTC region
not included here is central Chile and the shrublands there

()TD$FIG][ Opinion

Trends in Plant Science August 2011, Vol. 16, No. 8

Infertile soils

Large &
high nutrient
seeds

Intense seed predation

Canopy seed
storage

(+) Reduces seed predation

(-) Limits seed bank size to amount that can be stored on plant
(-) Increases carbon costs for seed protection
(-) Not resilient to fire cycles longer than adult lifespan
TRENDS in Plant Science

Figure 2. Model of factors selecting for serotiny and costs and benefits of serotiny.
Low soil fertility is hypothesized to select for larger seeds with higher nutrient
stores to enhance seedling survival. Predation pressure is heightened owing to the
lower nutrient stores in most plant tissues in the community and this selects for
canopy seed storage. Modified, with permission, from [2].

lack both, a post-fire pulse of seedlings and serotiny, which

is interpreted as a result of the diminution of natural fires
since the late Miocene, owing to the rise of the Andes that
block summer lightning storms [2].
Daniel I. Axelrod [18] also expressed skepticism about
the role of fire in the evolution of serotiny in northern
hemisphere pines, in part because he understood fire to be
an anthropogenic factor and unknown throughout land
plant evolution. He proposed that drought was a more
probable factor selecting for serotiny. We argue that if
drought were a stronger driving force than fire, it would
be more likely to select for a bet-hedging strategy in which
seed dispersal was spread out over multiple years, rather
than all seeds being dumped immediately after fire, when
there is a significant probability that it could be a dry year.
Bradshaw et al. [1] suggest rainfall predictability is a
more probable selective force behind the evolution of serotiny than is fire. They contend that winter rainfall is less
predictable in the two northern hemisphere MTC regions
and, thus, soil seed storage, which could spread germination over multiple years, would be preferable to serotiny.
However, northern hemisphere MTC shrublands with soil
stored seed banks do not exhibit seed carryover after fire
and, just like their southern hemisphere serotinous cousins, they too have a single pulse of seedling recruitment
after fire. Another weakness of this theory is the likelihood
that subtle differences in rainfall predictability between
the northern and southern hemisphere MTCs have fluctuated greatly since the Tertiary origin of serotiny [2].
Heat-shock triggered germination
We agree with Bradshaw et al. [1] that the physical dormancy of seeds has had multiple origins; however, because
of this, Bradshaw et al. cannot use the presence of physical
dormancy in non-fire-prone landscapes as evidence that it
is always an exaptation in fire-prone environments. The
possibility remains that, in some plant lineages, physical
dormancy originated in response to fire and, in other
lineages, it was appropriated by species suddenly cast into

a fire-prone setting. Ultimately, however, sorting out adaptation from exaptation might be extremely challenging
in some lineages (Box 1). For example, the Fabaceae are
well known for being hard seeded, meaning that they
produce outer integuments of densely packed palisade cells
and waxy cuticles that make the seed impermeable to
water until scarified by heat or other abrasive agent.
However, not all Fabaceae seeds are water impermeable.
In California grasslands, there are species of Lupinus,
Astragalus and Trifolium that germinate every spring
without any apparent physical scarification, but many of
these same species also occur in adjacent chaparral and
have deeply dormant seeds that seldom germinate until
exposed to heat shock from a fire. Although hard seededness in the Fabaceae is widespread, it appears to wax and
wane with local conditions and selection for or against
physical dormancy depends on local conditions (scenario
4b in Figure I, Box 1). When derived from grassland
ecotypes, deeply dormant chaparral seeds clearly reflect
selection for dormancy, despite the fact that the integument structures are a feature of parallel evolution in the
Fabaceae. We hypothesize that, although the seed coat
structures per se might have originated in response to any
number of factors, their precise organization into a heatshock-dependent seed is an adaptation to fire in some
situations.
Germination triggered by combustion chemicals
Smoke-induced germination is a trait that, on the surface,
would seem to be a clear adaptation to fire, but it is more
complicated than appears at first glance. In most MTC
ecosystems, there are species with dormant seed banks that
only germinate in the first growing season after fire, and can
be shown experimentally to only germinate in response to
smoke or charred wood extracts. In these cases, smokestimulated germination is clearly an adaptive trait. Whether the origin of this adaptive trait represents an adaptation
evolved in response to fire remains an open question.
The relatively recent discovery of a butanolide compound (karrikinolide) in smoke that will trigger germination of a vast array of species, many that lack any ecological
connection with fire, has raised questions about the selective role of fire in MTC ecosystems [1]. In an apparent quest
for the unifying theory of seed germination, several investigators [19,20] have focused intensively on this compound
and ignored reports of other compounds in smoke that
trigger the germination of MTC species with fire-dependent seedling recruitment [21] and other negative evidence
for butanolide being a universal signal [22]. All of the work
to date on karrikinolides in germination involves lab studies and essentially nothing is known about its ecological
role in fire-prone environments. That this compound triggers germination at levels of parts per trillion [19] raises
important ecological questions. If it is the universal germination trigger, then how do MTC soil-stored seed banks
remain dormant for decades in the face of such extraordinary sensitivity?
Bradshaw et al. [1] do provide a refreshing new theory
about the Mesozoic origin of smoke-stimulated germination initially in response to organic matter decay. However,
this theory is subject to all of the same criticisms that they
409

Opinion
level against fire adaptations. The only potential evidence
they present is the observation that karrikinolide-stimulated germination is widespread throughout all major
clades of angiosperms and, thus, is a feature of basic seed
metabolism that possibly originated during the early evolution of angiosperms. In their view, only organic matter
decay and not fire has been present since early angiosperm
evolution, but this is not supported by the vast literature
on Paleozoic and Mesozoic fires [813].
However, assuming for arguments sake that karrikinolides evolved in response to some other environmental
factor and are the only compounds in smoke that play a role
in germination of species with strict post-fire recruitment,
the fact that karrikinolides might have played a role in the
evolution of seed metabolism of all angiosperms does not
preclude an adaptive role for species that currently use it to
cue germination to post-fire conditions. Given that most
species seem to be sensitive to karrikinolides, yet only a
small subset of MTC species have the capacity to remain
dormant and then cue germination to the immediate postfire environment, it seems feasible that these MTC species
might have evolved some level of gene regulation controlling the functioning of karrikinolides in germination. This
would constitute a fire adaptation.
Flammability as a fire adaptation
Evolution of flammability implies selection for traits that
increase the frequency or intensity of fires. The idea is
largely dismissed by Bradshaw et al. [1] because of their
contention that it is based only on theoretical models.
However, studies of trait origins are not readily amenable
to experimentation and evolutionary comparative methods
are among out best available tools. We believe it is a viable
hypothesis that species restricting reproduction to a single
post-fire pulse of recruitment might be under selective
pressure to affect fire activity in ways that enhance their
reproductive success [23]. In crown fire ecosystems, traits
that could enhance flammability include small leaves,
volatile compounds and retention of dead leaves and
branches, to name just a few. There are empirical data
showing that trait distribution in some genera and communities is consistent with this hypothesis [6,24,25]. Also
consistent with this hypothesis is the observation that,
within fire-prone communities, species with post-fire seedling recruitment have the most flammable canopies [2,24].
This could be selected owing either to greater inhibition of
neighbors [23] or to enhanced seedbank survival by limiting deposition of dead fuels on the soil surface [26]. We
acknowledge that there is limited evidence that characteristics contributing to enhanced flammability actually enhance fitness (i.e. are adaptive) in fire-prone environments,
let alone that they represent an adaptation evolved in
response to fire. However, evidence is likewise lacking
that, in species from crown fire regimes, traits such as
retention of dead branches serve another function, which
evolved in response to a non-fire-related pressure.
Conclusion
Bradshaw et al. [1] raise some legitimate questions about
traits that are interpreted to be adaptive in fire-prone
environments. However, pointing out gaps in the ability
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Trends in Plant Science August 2011, Vol. 16, No. 8

to trace the origin of many traits to a fire origin is not

equivalent to demonstrating these traits arose in response
to other environmental factors. Contrary to their assertion,
Bradshaw et al. have not demonstrated that any fireadaptive trait has a more complex origin or that any
single trait arose in response to some other factor. In
several cases, they have rejected a possible adaptive role
for traits by defining them in very broad terms so that,
across the plant kingdom, the traits appear in many different selective environments. More importantly, trait
evolution is shaped by many forces that act throughout
the history of the trait, and it is naive to think that each
trait should be related to a single evolutionary pressure.
One of the unfortunate aspects of their paper [1] is that it
uses evolutionary arguments to draw conclusions about
appropriate fire management decisions. The assertion is
made that plants with traits interpreted as exaptations
perform worse in the face of recurrent fire than do those
that represent adaptations originating in response to fire.
However, the concept of adaptation versus exaptations only
refers to the origin of the trait (Box 1), not to the current role.
Thus, there is no reason to believe that plants with exapted
fire traits perform worse than do plants with fire adaptations and the authors do not provide any support for such a
view. Resource managers and other conservationists are
interested in how plants perform under current fire regimes
and mostly regard the evolutionary origin of traits as irrelevant to management decisions. It is possible for managers to
predict the behavior and dynamics of plants under different
alternative fire regimes on the basis of their functional traits
[27], without regard to their origins.
Acknowledgments
JGP acknowledges VIRRA (CGL2009-12048/BOS) support from the
Spanish Government. Any use of trade, product, or firm names in this
publication is for descriptive purposes only and does not imply
endorsement by the US Government.

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Press

Plant Science Conferences in 2011

Plant Biology 2011
610 August, 2011
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14th Symposium on Insect-Plant Interactions
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Scandinavian Plant Physiology Society (SPPS) Meeting 2011
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Plant Organellar Signaling from Algae to Higher Plants
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