Bridging the Divide: Integrating Animal and Plant Paradigms to Secure the Future of Biodiversity in Fire-Prone Ecosystems

Abstract

Conserving animals and plants in fire-prone landscapes requires evidence of how fires affect modified ecosystems. Despite progress on this front, fire ecology is restricted by a dissonance between two dominant paradigms: ‘fire mosaics’ and ‘functional types’. The fire mosaic paradigm focuses on animal responses to fire events and spatial variation, whereas the functional type paradigm focuses on plant responses to recurrent fires and temporal variation. Fire management for biodiversity conservation requires input from each paradigm because animals and plants are interdependent and influenced by spatial and temporal dimensions of fire regimes. We propose that better integration of animal-based and plant-based approaches can be achieved by identifying common metrics that describe changes in multiple taxa; linking multiple components of the fire regime with animal and plant data; understanding plant-animal interactions; and incorporating spatial and temporal characteristics of fires into conservation management. Our vision for a more integrated fire ecology could be implemented via a collaborative and global network of research and monitoring sites, where measures of animals and plants are linked to real-time data on fire regimes.

Full text

fire
Perspective
Bridging the Divide: Integrating Animal and Plant
Paradigms to Secure the Future of Biodiversity in
Fire-Prone Ecosystems
Luke T. Kelly 1, *, Lluís Brotons 2,3,4 ID , Katherine M. Giljohann 5, Michael A. McCarthy 5ID ,
Juli G. Pausas 6and Annabel L. Smith 7ID
1School of Ecosystem and Forest Sciences, University of Melbourne, Parkville, VIC 30105, Australia
2
InForest Joint Research Unit (CREAF-CTFC), Crta. de Sant Llorenç de Morunys, Km. 2, 25280 Solsona, Spain;
lluis.brotons@gmail.com
3CREAF, Campus de Bellaterra (UAB) Edifici C 08193 Cerdanyola del Vallès, Spain
4CSIC, 08193 Cerdanyola del Vallès, Spain
5School of BioSciences, University of Melbourne, Parkville, VIC 3010, Australia;
kmgi@unimelb.edu.au (K.M.G.); mamcca@unimelb.edu.au (M.A.M.)
6CIDE-CSIC, Ctra. Naquera Km. 4.5, 46113 Montcada, Valencia, Spain; juli.g.pausas@ext.uv.es
7Department of Zoology, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland;
annabel@smithecology.org
*Correspondence: ltkelly@unimelb.edu.au
Received: 18 July 2018; Accepted: 8 August 2018; Published: 10 August 2018


Abstract:
Conserving animals and plants in fire-prone landscapes requires evidence of how fires
affect modified ecosystems. Despite progress on this front, fire ecology is restricted by a dissonance
between two dominant paradigms: ‘fire mosaics’ and ‘functional types’. The fire mosaic paradigm
focuses on animal responses to fire events and spatial variation, whereas the functional type paradigm
focuses on plant responses to recurrent fires and temporal variation. Fire management for biodiversity
conservation requires input from each paradigm because animals and plants are interdependent and
influenced by spatial and temporal dimensions of fire regimes. We propose that better integration
of animal-based and plant-based approaches can be achieved by identifying common metrics that
describe changes in multiple taxa; linking multiple components of the fire regime with animal
and plant data; understanding plant-animal interactions; and incorporating spatial and temporal
characteristics of fires into conservation management. Our vision for a more integrated fire ecology
could be implemented via a collaborative and global network of research and monitoring sites,
where measures of animals and plants are linked to real-time data on fire regimes.
Keywords:
biodiversity monitoring; fire ecology; fire management; fire mosaic; functional trait;
life-history; plant functional type; plant-animal interactions; pyrodiversity; wildfire
1. Introduction
Historical fire regimes have shaped biodiversity, but conserving animals and plants in
contemporary landscapes requires evidence of how fires affect ecosystems that are modified by
and subject to new threats [
1
]. Although progress has been made on this front, fire ecology is still
restricted by a dissonance between two dominant paradigms that differ in theory and application:
‘fire mosaics’ and ‘functional types’ [
2
]. The fire mosaic paradigm focuses on animal responses to
fire events. Fire management under this paradigm typically aims to create spatially diverse fire
mosaics to promote biodiversity or the persistence of a small number of focal species [
3
]. By contrast,
the functional type paradigm focuses on plant responses to recurrent fires. Fire management under
Fire 2018,1, 29; doi:10.3390/fire1020029 www.mdpi.com/journal/fire

Fire 2018,1, 29 2 of 8
this paradigm is guided by the life-history traits of plants and aims for temporal variation within
acceptable fire intervals [4].
Fire management for biodiversity conservation cannot be achieved by applying either paradigm
in isolation: it requires input from each because animals and plants are both of value, interdependent,
and influenced by the spatial and temporal dimensions of fire regimes. We propose that better
integration of animal-based and plant-based approaches can be achieved by identifying common
metrics that describe changes in multiple taxa, linking multiple components of the fire regime
with measures of animals and plants, understanding shared mechanisms underpinning animal and
plant associations with fires, and incorporating spatial and temporal characteristics of fires into
conservation management.
2. Animal and Plant Paradigms in Fire Ecology
Fire science and management vary greatly between continents and ecosystems but there is value
in condensing knowledge of both animals and plants to enhance application of fire ecology. To this end,
we identify four general characteristics of animal and plant paradigms and how they differ (Figure 1).
Fire 2018, 1, x FOR PEER REVIEW 2 of 8
recurrent fires. Fire management under this paradigm is guided by the life-history traits of plants and aims
for temporal variation within acceptable fire intervals [4].
Fire management for biodiversity conservation cannot be achieved by applying either paradigm in
isolation: it requires input from each because animals and plants are both of value, interdependent, and
influenced by the spatial and temporal dimensions of fire regimes. We propose that better integration of
animal-based and plant-based approaches can be achieved by identifying common metrics that describe
changes in multiple taxa, linking multiple components of the fire regime with measures of animals and
plants, understanding shared mechanisms underpinning animal and plant associations with fires, and
incorporating spatial and temporal characteristics of fires into conservation management.
2. Animal and Plant Paradigms in Fire Ecology
Fire science and management vary greatly between continents and ecosystems but there is value in
condensing knowledge of both animals and plants to enhance application of fire ecology. To this end, we
identify four general characteristics of animal and plant paradigms and how they differ (Figure 1).
Figure 1. Fire mosaic and functional type paradigms can be defined by the following characteristics: (i)
taxonomic focus (animal-based vs. plant-based); (ii) how they characterize fire regimes (fire events vs.
recurrent fires); (iii) the mode of generalization (habitat change vs. life-history traits); and (iv) implications
for fire management (spatial mosaics vs. temporal intervals). A more integrated approach would
emphasize: multiple taxonomic groups and their interactions (biodiversity); different components of the fire
regime acting in concert (fire regimes); comprehensive understanding of mechanisms underpinning biotic
associations with fire and interdependency between different taxa (shared mechanisms); and
decision-making that considers how both spatial and temporal dimensions of fire regimes influence
biodiversity (spatio-temporal management).
2.1. Taxonomic Focus
The different ecology and evolution of plants and animals in fire-prone ecosystems partly explains the
distinct management approaches and underpinning research [5]. Plants are immobile throughout much of
their life-cycle and many have endogenous means of reproduction from seedbanks or vegetative buds that
promote in situ persistence after a fire [6]. Individual animals are more mobile and some can survive fires
by moving to refuges or recolonize from surrounding areas after being extirpated from burnt areas [7].
Some differences in management and research are also likely to stem from the separate developments of
the animal and plant sciences. We reviewed the ten most recent papers on fire ecology published in each of
five exemplar journals (Biological Conservation, Ecology, Forest Ecology and Management, Global Ecology and
Biogeography and International Journal of Wildland Fire; n = 50 papers) and found that 78% were solely
animal-based or plant-based, and only 22% investigated both animal and plant associations with fire
[Supplementary Material]. This suggests research culture and training play a part in forming different
perspectives.
2.2. Measures of Fire
A central concept in fire ecology is the fire regime, which describes the repeated pattern of fires in a
given area and their characteristics [8,9]. For animals, commonly measured components of the fire regime
Figure 1.
Fire mosaic and functional type paradigms can be defined by the following characteristics:
(i) taxonomic focus (animal-based vs. plant-based); (ii) how they characterize fire regimes (fire events vs.
recurrent fires); (iii) the mode of generalization (habitat change vs. life-history traits); and (iv) implications
for fire management (spatial mosaics vs. temporal intervals). A more integrated approach would
emphasize: multiple taxonomic groups and their interactions (biodiversity); different components of the
fire regime acting in concert (fire regimes); comprehensive understanding of mechanisms underpinning
biotic associations with fire and interdependency between different taxa (shared mechanisms);
and decision-making that considers how both spatial and temporal dimensions of fire regimes influence
biodiversity (spatio-temporal management).
2.1. Taxonomic Focus
The different ecology and evolution of plants and animals in fire-prone ecosystems partly explains
the distinct management approaches and underpinning research [
5
]. Plants are immobile throughout
much of their life-cycle and many have endogenous means of reproduction from seedbanks or
vegetative buds that promote in situ persistence after a fire [
6
]. Individual animals are more mobile
and some can survive fires by moving to refuges or recolonize from surrounding areas after being
extirpated from burnt areas [
7
]. Some differences in management and research are also likely to stem
from the separate developments of the animal and plant sciences. We reviewed the ten most recent
papers on fire ecology published in each of five exemplar journals (Biological Conservation,Ecology,
Forest Ecology and Management,Global Ecology and Biogeography and International Journal of Wildland Fire;
n = 50 papers) and found that 78% were solely animal-based or plant-based, and only 22% investigated
both animal and plant associations with fire [Supplementary Material]. This suggests research culture
and training play a part in forming different perspectives.

Fire 2018,1, 29 3 of 8
2.2. Measures of Fire
A central concept in fire ecology is the fire regime, which describes the repeated pattern of fires in
a given area and their characteristics [
8
,
9
]. For animals, commonly measured components of the fire
regime include those representing fire events, including severity and patchiness, and time since the last
fire [
2
]. These components can affect animals’ capacity to survive fire and their subsequent ability to
find food, find shelter, and recolonize regenerating habitats. For plants, fire frequency and the interval
between successive fires have been of central importance to research [
10
]. The time between fires sets
the window for growth and reproduction in many plants. Studies of animals and plants that explore
combinations of fire regime characteristics are increasing [
11
] and represent an important development
in fire ecology.
2.3. Mode of Generalization
Generalization assists both ecological understanding and prediction across taxa, times, and places.
Theories of post-fire succession underpin much animal research in fire ecology. For example,
the ‘habitat accommodation model’ (Fox 1982) predicts that faunal species enter the post-fire succession
when vegetation structure and floristic resources becomes suitable for them. As the vegetation changes
and becomes less suitable for a species, it will be excluded from the succession or decline in abundance
from competition [
12
]. In search of unifying trends, fire response patterns have been linked to
animal life-history, including shelter, food, fecundity, and mobility; such approaches have provided
informative descriptions but have often met with little predictive success [13].
Life-history approaches have been widely used for generalization in plant ecology [
6
].
For example, Noble and Slatyer (1980) derived an important method for predicting plant dynamics
in relation to recurrent fires using three groups of traits or ‘vital attributes’: method of arrival or
persistence at a site during and after fire, the ability to establish and grow to maturity, and time taken
to reach critical life stages [
14
]. Grouping plants into functional types, i.e., sets of species that share
traits, has been used repeatedly to examine post-fire responses of plants with considerable predictive
success [6,15].
2.4. Management Emphasis
A focus on fire events, vegetation succession, and accompanying changes in habitat structure has
translated into management for animals that aims for spatially diverse fire mosaics [
2
]. A widespread
expectation is that ‘pyrodiversity promotes biodiversity’, which is a prediction based on linking
the persistence of individual species, and the coexistence of multiple species, to variation in the
composition and configuration of fire mosaics [
3
]. This perspective highlights threats to animal
conservation triggered by large, severe, and uniform fires, which, in some locations, have been linked
to a build-up of fuels under fire suppression. Management to achieve spatially diverse fire regimes
varies between continents and ecosystems. In Australia, South Africa, and increasingly in Europe and
South America, patch-mosaic burning or planned burning have been implemented [
3
,
6
]. In North
America, there has been a strong focus on restoration of fire regimes to historical conditions [
1
,
16
],
and, more recently, on encouraging natural patterns of ignitions that result in mixed-severity fires [
17
].
The fire mosaic paradigm and pyrodiversity hypothesis have been criticized for their simplistic
representation of fire regimes and animal habitat [2,3].
An emphasis on functional types has translated into management for plants that aims for
variation in fire intervals over time. An influential approach uses plant life-history traits, such as
time to reproductive maturity, life-spans of established plants, and seed longevity to determine
minimum and maximum fire intervals, beyond which a significant decline of species populations is
predicted [
8
]. Accordingly, domains of tolerable fire intervals are derived and used to guide vegetation
management [
4
]. This perspective highlights two main threats: immaturity risk, where fire intervals are
shorter than the time needed for plants to mature and accumulate seed; and senescence risk, where fire

Fire 2018,1, 29 4 of 8
intervals are longer than seed and plant life-spans [
6
]. Plant life-history and demographic data have
been used widely to guide fire management in Australia [
4
] and South Africa [
18
], with examples from
other locations, such as North America [
19
], growing. Useful approaches have also been developed
that consider multiple measures of fire regimes—‘thresholds of particular concern’ and ‘bounded
ranges of variation’ [16]—but in practice are rarely linked to requirements of plants.
3. Towards a More Integrated Fire Ecology
We propose practical ways to achieve better integration of animal and plant perspectives in
fire ecology and management (Figure 1: Integrated approach). Our aim is to identify a small set of
opportunities to advance fire ecology and management regardless of the focal ecosystem or taxa.
3.1. Common Measures of Biodiversity
A simple way to enhance management and learning is to directly measure plants, animals,
and habitat elements. While some measures of biodiversity will be context specific, recording
observations in a standardized way to allow comparisons between taxa and ecosystems is valuable.
Harmonization of global biodiversity monitoring using Essential Biodiversity Variables (EBVs) [
20
]
could be adapted to fire-prone ecosystems. Under the EBVs framework, six dimensions of biodiversity
have been identified—including genetic composition, species population, species traits, community
composition, ecosystem function, and ecosystem structure—to assess biodiversity change [
20
]. As a
first step, essential variables used to inform knowledge of species persistence in fire-prone landscapes
could include the occurrence and abundance of animals and plants at key life stages [
13
], traits that
mediate fire responses at different levels of ecological organization [
15
], and elements of habitat
structure, such as tree size and density [
17
]. Systematic measures of species occurrence or abundance
could be pooled to provide multi-species indicators of biodiversity, and models used to extrapolate
observations from sites to landscapes. Monitoring multiple species and taxa will help to avoid some
of the pitfalls of single-species management, such as when a focal or indicator species is incorrectly
assumed to represent biodiversity more broadly [
1
]. An ongoing challenge is the complexity in
measuring multiple taxonomic groups, from invertebrates and herbs to larger vertebrates and trees,
which show great variation in size, diversity, and detectability.
3.2. Multiple Components of Fire Regimes
Rapid progress in satellite remote sensing and aerial photography has already produced valuable
resources that aid choices about when and where to conduct fuel management and fire suppression [
2
].
One practical challenge is describing variation in fire regimes over space and time in a consistent way
so that studies can be compared at different times and places. Several metrics of pyrodiveristy have
been developed, and the synthesis and testing of these approaches would facilitate advances globally.
Another challenge is the coupling of growing spatial data on fire with the sampling of animals and
plants. New work on spatially explicit power analysis shows how the location and level of sampling
required to detect changes in animal and plant populations can be identified [
21
]. In combination with
common measures of plants, animals, and fire regimes, power analysis could be used to design the
assessment and monitoring of fire management and boost the understanding of multiple components
of fire regimes.
3.3. Shared Mechanisms
Approaches for generalization in animal and plant ecology usually concentrate on resource
change (e.g., food, light) or traits, but rarely both. One way to formally bridge resource change and
trait-based approaches is to incorporate functional traits into models of species-fire relationships.
For example, Thomas and Vesk (2017) built a hierarchical multi-species model to predict tree species
growth trajectories at different times since fire as a function of their traits (stem density, seed size,
and specific leaf area) [
22
]. This approach could be extended to animal data, and can be combined

Fire 2018,1, 29 5 of 8
with new measures of ecological, behavioral, and physiological traits [
5
], to provide a mechanistic
understanding of animal succession and better integration with plant functional trait frameworks.
Fire not only affects animals and plants but also their interactions. Interactions such as pollination
and seed dispersal play an essential role in the maintenance of biodiversity, and explicit consideration
of plant-animal interactions will provide new insights and benefit fire management and conservation
(Table 1). In addition to revealing shared mechanisms, a focus on interactions between trophic levels
will help to understand the evolution of life-history strategies and traits in fire-prone ecosystems.
For example, Talluto and Benkman (2014) showed that the frequency of serotiny in lodgepole pine
across Yellowstone National Park reflects selection pressure from fire and seed predation by the red
squirrel [
23
]. This highlights that ecosystem structure and function vary as a result of spatial and
temporal variation in fires and plant-animal interactions.
Table 1.
Consideration of both animals and plants provides new insights and benefits fire management
for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire
Management
Habitat development
Considering structural changes of plants provided new
understanding of habitat provision over century-long time
scales and showed that tolerable fire intervals based solely
on plant occurrence were too short for animal conservation
(Haslem et al. 2011) *.
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Pollination
Nest location and floral resource use mediate pollinator
responses to fire. Plant traits such as growth form,
phenology and bud location influence flowering responses
to fire. Pollinators most vulnerable to changing fire regimes
are predicted to be those that nest above-ground and have
one brood per year (Brown et al. 2017).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Decomposition
Fossorial mammals affected fire behavior via an increase in
the rate of organic matter breakdown. Reintroducing
fossorial mammal species to landscapes where they have
previously been extinct offers a new way to modify fire
regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Seed dispersal
Colonization of holm oak in burnt forest is mediated by
acorn dispersal by Eurasian jays. Post-fire salvage logging
reduced the strength of this key plant-animal interaction.
Management policies of non-intervention after forest fire are
likely to increase the resilience of the ecosystem (Castro et al.
2012).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Seed predation
Mediterranean gorse seeds are predated upon by the weevil
Exapion fasciolatum. Mediterranean gorse experienced lower
seed damage in burnt compared to unburnt areas. Plants can
benefit from fire through disruption of antagonistic
interactions with seed predators and this might be one
mechanism promoting success of fire-adapted plants (García
et al. 2016).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Grazing
In periods of low rainfall, grazing by kangaroos and feral
herbivores following prescribed fire had a large effect on the
survival of seedlings and resprouting plants. This indicates
that conservation of plants and animals will benefit from fire
management that considers how herbivore management
influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
Community assembly
Spatial and temporal variation in fires influences community
assembly globally (Kelly & Brotons 2017), and feedback
among fires, biodiversity and ecological processes are
central to understanding community-level changes
(Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a
desert shrubland to a low-diversity, invasive grassland. In
areas where they remained, rodents created biotic resistance
to invasive plants, with cascading effects on plant diversity
(St. Clair et al. 2016).
Fire 2018, 1, x FOR PEER REVIEW 5 of 8
and Benkman (2014) showed that the frequency of serotiny in lodgepole pine across Yellowstone National
Park reflects selection pressure from fire and seed predation by the red squirrel [23]. This highlights that
ecosystem structure and function vary as a result of spatial and temporal variation in fires and
plant-animal interactions.
Table 1. Consideration of both animals and plants provides new insights and benefits fire
management for biodiversity conservation.
Process Benefit to Ecological Understanding and Fire Management
Habitat
development
Considering structural changes of plants provided new understanding of
habitat provision over century-long time scales and showed that tolerable
fire intervals based solely on plant occurrence were too short for animal
conservation (Haslem et al. 2011) *.
Pollination
Nest location and floral resource use mediate pollinator responses to fire.
Plant traits such as growth form, phenology and bud location influence
flowering responses to fire. Pollinators most vulnerable to changing fire
regimes are predicted to be those that nest above-ground and have one
brood per year (Brown et al. 2017).
Decomposition
Fossorial mammals affected fire behavior via an increase in the rate of
organic matter breakdown. Reintroducing fossorial mammal species to
landscapes where they have previously been extinct offers a new way to
modify fire regimes for the benefit of plant and animal conservation
(Hayward et al. 2016).
Seed dispersal
Colonization of holm oak in burnt forest is mediated by acorn dispersal by
Eurasian jays. Post-fire salvage logging reduced the strength of this key
plant-animal interaction. Management policies of non-intervention after
forest fire are likely to increase the resilience of the ecosystem (Castro et al.
2012).
Seed predation
Mediterranean gorse seeds are predated upon by the weevil Exapion
fasciolatum. Mediterranean gorse experienced lower seed damage in burnt
compared to unburnt areas. Plants can benefit from fire through
disruption of antagonistic interactions with seed predators and this might
be one mechanism promoting success of fire-adapted plants (García et al.
2016).
Grazing
In periods of low rainfall, grazing by kangaroos and feral herbivores
following prescribed fire had a large effect on the survival of seedlings
and resprouting plants. This indicates that conservation of plants and
animals will benefit from fire management that considers how herbivore
management influences a foundation species of semi-arid Australia
(Giljohann et al. 2017).
Community
assembly
Spatial and temporal variation in fires influences community assembly
globally (Kelly & Brotons 2017), and feedback among fires, biodiversity
and ecological processes are central to understanding community-level
changes (Bowman et al. 2016). For example, fire in combination with
experimental exclusion of seed-eating rodents shifted a desert shrubland
to a low-diversity, invasive grassland. In areas where they remained,
rodents created biotic resistance to invasive plants, with cascading effects
on plant diversity (St. Clair et al. 2016).
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol. 2011, 48, 247–256; Brown, J. et al. J. Appl.
Ecol. 2017, 54, 313–322; Hayward, M.W. et al. Anim. Conserv. 2016, 19, 490–497; Castro, J. et al. Ecosphere 2012,
3, 1–12; García, Y. et al. Oecologia, 2016, 182, 1165–1173; Giljohann, K.M. J. Ecol. 2017, 105, 1562–1570; Kelly,
L.T. and Brotons, L. Science, 2017, 355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B. 2016, 371,
20150169; St. Clair, S.B. et al. Ecology 2016, 97, 1700–1711.
* Additional references in Table 1: Haslem, A. et al. J. Appl. Ecol.
2011
,48, 247–256; Brown, J. et al. J. Appl. Ecol.
2017
,54, 313–322; Hayward, M.W. et al. Anim. Conserv.
2016
,19, 490–497; Castro, J. et al. Ecosphere
2012
,3, 1–12;
García, Y. et al.
Oecologia,
2016
,182, 1165–1173; Giljohann, K.M. J. Ecol.
2017
,105, 1562–1570; Kelly, L.T. and Brotons,
L. Science,
2017
,355, 1264–1265; Bowman, D.M.J.S. et al. Phil. Trans. R. Soc. B.
2016
,371, 20150169; St. Clair, S.B. et al.
Ecology 2016,97, 1700–1711.

Fire 2018,1, 29 6 of 8
3.4. Spatio-Temporal Fire Management
There is broad agreement that spatial and temporal dimensions of fire regimes matter but it is rare
that interconnections between the two are explicitly incorporated into management for biodiversity.
One reason is that data on plants, animals, and spatio-temporal mosaics has been lacking [
11
]. In part,
this is being addressed by a surge of new studies on fire regimes and biodiversity [
3
]. A continuing issue
is that results of studies are rarely available as mapped outputs that can directly inform spatial-temporal
management. To complement maps of fire history, we recommend mapping outputs from models that
directly link biodiversity with measures of fires (e.g., species distribution and population models) [
24
].
This would facilitate comparisons of fire impacts across different taxa and in different places, help to
identify when and where key populations of plants and animals will be located, and strengthen efforts
to define management thresholds in different components of fire regimes. We are already using linked
fire and biodiversity data to inform decision making across large areas of southern Australia and
northern Spain ([
3
,
25
] and references therein). Useful extensions could include mapping interactions
and interdependencies between plants and animals, such as pollinator networks (Table 1); linking
spatial models to fire simulations and management strategies to predict future changes in plant and
animal populations; and producing real-time environmental data to aid the urgent decisions that affect
people and biodiversity during fire events.
We have only touched on some important debates that span animal and plant ecology such
as single- vs. multi-species management, historical range and variability vs. contemporary fire
management, and correlative vs. mechanistic approaches. Insights from these other approaches are
likely to produce ideas additional to those proposed in this paper.
4. Conclusions
Many animals and plants show different responses to fires but we argue that searching for
commonality and links will enhance learning about both groups. Our vision for a more integrated
fire ecology could be implemented via a collaborative and global network of research and monitoring
sites in fire-prone ecosystems, where measures of animals and plants are linked to real-time data on
the spatial and temporal dimensions of fires. Well-designed monitoring would provide the following:
opportunities to reveal new ecological patterns, processes, and interactions between animals and
plants; a more robust understanding of the effect of fire regimes on biodiversity; and a strong basis for
choosing between alternative fire management strategies. This proposal would be strengthened by
collaborations between animal and plant ecologists, using emerging approaches for generalization
such as trait-based models. Finally, bringing together animal and plant perspectives will benefit
from other recent developments in fire ecology, notably, frameworks that recognize the importance of
socio-ecological linkages, adaptive resilience, and decision making under uncertainty [3,16].
Supplementary Materials:
The following are available online at http://www.mdpi.com/2571-6255/1/2/29/s1.
Author Contributions:
L.K. conceived the idea for the paper and led the writing. All authors made substantial
contributions to the development of ideas and drafts.
Funding:
Kelly was funded by the Australian Research Council Centre of Excellence for Environmental Decisions
and a Victorian Postdoctoral Research Fellowship delivered by veski on behalf of the Victorian Government.
Brotons and Pausas were funded by the Government of Spain on Project CGL2017-89999-C2 and CGL2015-64086-P,
respectively. Smith was supported by Marie Skłodowska-Curie Individual Fellowship FIRESCAPE-746191 under
the EU H2020 Programme for Research and Innovation.
Acknowledgments:
Many thanks to Clare Kelly who co-designed Figure 1and Table 1. We appreciate the useful
comments and suggestions provided by the reviewers of this paper.
Conflicts of Interest: The authors declare no conflict of interest.

Fire 2018,1, 29 7 of 8
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2018 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access
article distributed under the terms and conditions of the Creative Commons Attribution
(CC BY) license (http://creativecommons.org/licenses/by/4.0/).