Journal of Biogeography (J. Biogeogr.

) (2016) 43, 863–873

PERSPECTIVE Revising the biome concept for

understanding and predicting global
change impacts
Glenn R. Moncrieff1,2*, William J. Bond1,3 and Steven I. Higgins4,5

1
Fynbos Node, South African Environmental
Observation Network (SAEON), Centre for
Biodiversity Conservation, Kirstenbosch
Gardens, Private Bag X7, Rhodes Drive,
Claremont 7735 Cape Town, South Africa,
2
Department of Botany and Zoology,
Stellenbosch University, Matieland 7602,
South Africa, 3Department of Biological
Sciences, University of Cape Town,
Rondebosch 7701, South Africa, 4Department
of Botany, University of Otago, PO Box 56,
Dunedin 9054, New Zealand, 5Biodiversity
and Climate Research Centre (BiK-F),
Senckenberg Gesellschaft f€
ur Naturforschung,
Senckenberganlage 25, 60325 Frankfurt am
Main, Germany
*Correspondence: Glenn R. Moncrieff. Fynbos
Node, South African Environmental
Observation Network (SAEON), Centre for
Biodiversity Conservation, Kirstenbosch
Gardens, Private Bag X7, Rhodes Drive,
Claremont 7735, Cape Town, South Africa.
E-mail: glenn.moncrieff@gmail.com
ABSTRACT
Biomes are globally distributed, structurally and functionally similar vegetation
units defined without reference to plant species composition. The boundaries
between biomes are presumed to correspond with species turnover and changes
in biogeochemical cycling. Determining the controls of biome distributions is
thus critical for anticipating global change impacts. Historically, climate and
soils have been understood to adequately explain the global distribution of
biomes. Convergent evolution and environmental filtering are assumed to be
pervasive, ultimately resulting in deterministic patterns of vegetation structure
and function in relation to prevailing environmental conditions. Recent studies
have highlighted significant problems with this view of biomes. Firstly, system-
atic structural and functional divergence within biomes has been identified
when comparing environmentally similar, yet floristically distinct regions. Sec-
ondly, climatic determinism is being further undermined by evidence suggest-
ing multiple stable biome states are possible for some combinations of climatic
drivers. We argue that biomes remain useful and necessary constructs for orga-
nizing our knowledge of how ecosystems function and for predicting how they
might respond to change. However, biome concepts should acknowledge the
limits to predictability from environmental conditions alone and the influence
of historical processes on modern vegetation patterns. We discuss how direct
mapping of plant structure and function, the incorporation of insights into
biome evolution and a new generation of vegetation models will lead to
improvements in the concept of what biomes are, where they occur, and efforts
to predict their distribution.
Keywords
alternate stable state, biome, DGVM, functional trait, global land cover, plant
functional type, potential natural vegetation, niche conservatism

Traditionally, it has been assumed that the occurrence of

INTRODUCTION
There is complex large-scale variation in vegetation across
the surface of the Earth, with important consequences for
the resources available to support human wellbeing,
biodiversity and biogeochemical cycling (Cramer et al., 2001;
Olson et al., 2001). Biomes are the basal unit used to orga-
nize and describe this variation. They refer to globally
convergent vegetation formations similar in structure and
function, explicitly ignoring floristic differences. The biome
concept is attractive because it can be applied globally with-
out reference to plant species composition, allowing general-
ity to be sought.
similar vegetation in areas distant both geographically and
evolutionarily is driven by two processes: environmental fil-
tering of organisms not well suited to the prevailing climate
and soils in the process of community assembly, and selec-
tion for strategies and forms that optimally use available
resources over evolutionary time resulting in convergence of
distantly related organisms (Schimper, 1903; Woodward
et al., 2004). Convergent evolution has been noted across a
range of plant and animal taxa. Striking examples include
the parallel forms of marsupials and placental mammals,
succulence and similar leaf morphology in distantly related
desert plants or alpine cushion plants (Aubert et al., 2014)

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doi:10.1111/jbi.12701
G. R. Moncrieff et al.
BOX 1: DYNAMIC GLOBAL VEGETATION MODELS
Dynamic Global Vegetation Models (DGVMs) integrate
plant physiological models into models of vegetation
biogeography and have allowed simulation of the temporal
dimension of vegetation change and scaling of leaf-level
processes to whole canopies and the globe (Prentice et al.,
2007). Gross primary productivity, respiration losses,
reproduction, mortality and interactions among plants are
all incorporated into models that simulate vegetation
dynamics in an effort to produce projections of the spatial
and temporal trajectory of vegetation change, the carbon
cycle and feedbacks to the climate system. Uncertainties in
the representation of various processes are large, as are
parameter estimates, but without this complexity projec-
tions of vegetation response to global change will be biased.
DGVMs descend from plant geography models which
initially attempted to predict the global distribution of
biomes based on climate correlates. Although no longer
purely correlative, the biome concept still underlies the
manner in which the complex reality of vegetation patterns
is simplified. Plant diversity in DGVMs is usually repre-
sented using Plant Functional Types (PFTs). PFTs in
DGVMs are defined based on similarity in photosynthetic
function (e.g. C3 vs. C4), response to disturbance, phenol-
ogy and physiognomy (Lavorel et al., 1997). Biomes are
delimited based on the relative dominance and combina-
tions of different PFTs (Bonan et al., 2002). PFTs provide a
useful conceptual link from biomes, through the plant types
of which they are composed, to ecosystem function, while
not necessitating the inclusion of species-specific informa-
tion. The traits that define PFTs can be related to specific
functions, allowing the mechanisms that control biome
distribution to be examined rather than merely producing
correlative models.
and the independent evolution of C4 and CAM photosynthe-
sis in multiple plant families (Keeley & Rundel, 2003).
The upshot of environmental filtering during community
assembly and convergent evolution in similar environments
is that the distribution of the plant forms that define biomes
should be predictable from the environment and not depend
on any regional or historical context. This assumption has
been at the foundation of efforts to understand plant bio-
geography at large scales. The equilibrium distribution of
biomes is often understood to be explained by only a few
variables that roughly describe the conditions relevant to
plants (Whittaker, 1975). The inclusion of additional vari-
ables and information on local topo-edaphic conditions can
improve predictions (Holdridge, 1947; Walter, 1973). This
thinking underpins most attempts to project the current,
past and future distribution of biomes using only climatic,
edaphic and atmospheric forcing variables as input (Box 1:
DGVMs). In a typical global vegetation model, globally
864
defined plant functional types (PFTs) or vegetation units
respond uniformly to environmental drivers, producing
biomes with consistent properties across continents.
The fundamental assumptions of interregional convergence
and the predictability of global vegetation patterns underly-
ing the biome concept and efforts to model global biome
distribution have been increasingly undermined. Here we
discuss how new directions from community ecology, vegeta-
tion modelling and phylogenetics motivate for an update of
how we define, map and understand the distribution of
global biomes. We explicitly ignore the substantial influence
of human activity and focus only on potential natural vegeta-
tion. We start by synthesizing the evidence that constraints
imposed by evolutionary and biogeographic history limit the
functional convergence observed within biomes. Next we
highlight how climate feedbacks, disturbance and ecosystem
engineering by plants alter conditions relevant for plant
functioning. These feedbacks result in non-deterministic
biome distribution patterns in which multiple biomes can
exist at the same location under the same macroclimatic and
geological template. In such instances this environmental
template will not necessarily prescribe the dominant biome.
We then look at new directions in mapping and modelling
spatial variation in plant form and function through func-
tional traits and recent analyses that show how historical
processes over evolutionary and ecological time-scales shape
modern biome distributions. We end by discussing how the
concept of biomes and their use to organize our knowledge
of global vegetation patterns and biogeochemistry can be
revised through the incorporation of these new directions.
CHALLENGES TO BIOMES
Interregional divergence in form and function
Biomes are defined and delimited using a variety of schemes
based on similarity in structure, function or climatic condi-
tions, but usually (at least when defining units at the global
scale) plant species composition and geographical location is
intentionally ignored (Box 2: Biome concepts). A comple-
mentary view of vegetation recognizes biogeographical realms
based on compositional similarity and shared evolutionary
history (Holt et al., 2013). Mapped boundaries between
realms represent significant barriers to current and historical
gene flow. Despite an important role for intercontinental and
trans-oceanic dispersal in the assembly of modern floras
(Pennington et al., 2004), past speciation, extinction and
vicariance have left an imprint on modern floras with geo-
graphically distant or isolated regions harbouring different
species assemblages. Historical differences in these processes
among regions are reflected in differences in contemporary
phylogenetic structure. Indeed it is possible for the same
biome to be composed of entirely different and distantly
related species in two different realms.
The evolutionary distinctiveness of different floras is likely
to have ecological implications. Closely related taxa are often
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 Biomes for understanding global change impacts

BOX 2: BIOME CONCEPTS

The occurrence of analogous vegetation formations in geographically disjunct although climatically similar regions was described
over 200 years ago by von Humboldt (1807). Schimper (1903) first defined and named the world’s biomes in an essentially
modern way. He argued that the dominant life-forms in each biome had convergent physiological attributes as a result of
selection to cope with the prevailing climate. Common to many early biome concepts is the explicit incorporation of climate
parameters into the definition of a biome (Table 1). Holdridge (1947) produced a systematic scheme of Life Zones describing
large vegetation formations that map onto axes of precipitation, temperature and potential evapotranspiration. Walter (1973)
similarly defined biomes in terms of the climate zone they occupy (Zonobiomes), with modifications caused by soils and
orographic features. The inclusion of climate to delimit a biome serves as a proxy for functional characteristics that are difficult to
quantify or map globally. For example, ‘tropical rain forests’ are, by definition, composed of plants sensitive to cold or frost and
non-deciduous. Remote sensing of land cover with satellites has allowed new global biome distribution maps to be created where
boundaries between vegetation units are ostensibly objectively defined (e.g. Arino et al., 2007). To allow the mapping of biomes
from remotely sensed data, most modern biome classification schemes explicitly ignore geographical context and climate when
defining biomes, rather emphasizing shared phenology and structure, and consistencies in the combination and relative
dominance of PFTs (Woodward et al., 2004). Focussing on characteristics observable from space can, however, lead to important
characteristics being ignored, such as whether plants are drought- or cold-deciduous and boundaries delimited based on arbitrary
cut-offs in order to produce agreement with training data. Both early and contemporary biome concepts and maps recognize
lower hierarchical levels of vegetation formations below global biomes which are defined with reference to geographical context
(e.g. Ecozones (Olson et al., 2001) or Biomes sensu strictu (Walter, 1973)). While these lower hierarchical levels are useful for
purposes such as conservation planning and regional analyses, we concern ourselves with biomes defined globally without
reference to geographical context or environmental conditions. This approach allows the distribution of PFTs and biomes in
relation to the environment to be analysed without circularity, and this biome concept is implicit in global vegetation models and
meta-analyses of global biogeochemical cycling or species richness.

Table 1 An overview of some well known biome concepts used for vegetation modelling, mapping and understanding the
determinants of global vegetation patterns.

Actual or
Biome potential
concept Term Defined based on Mapping variables vegetation Hierarchical Reference

Walter Zonobiome Climate Climate Potential Yes Walter, 1973

Schultz Ecozone Climate Climate Potential Yes Schultz, 2002
WWF Biome Climate, Structure, Climate, Soils, Expert Potential Yes Olson et al., 2001
Orography knowledge, Structure
MODIS- Land cover Structure, Function, Spectral reflectance, Actual No Friedl et al., 2002
IGBP Human influence Temperature, Topography,
Snow/Ice
DGVMs Biome Structure, Function Structure, Function Actual and No Sitch et al., 2003;
(Combination of PFTs) Potential Prentice et al., 2007

more similar ecologically than would be expected if traits
evolved randomly, resulting from the tendency of species to
retain their ancestral ecology (Wiens et al., 2010). This phy-
logenetic conservatism is evident in species traits (Chazdon
et al., 2003), environmental tolerances (Petitpierre et al.,
2012) and the biomes in which species occur (Crisp et al.,
2009). If regional floras are phylogenetically structured, and
phylogeny constrains trait evolution, it follows that there
ought to be divergence in plants traits among floristically
distinct regions, potentially affecting emergent ecosystem
properties. In contradiction, convergence of floras to similar
vegetation structure and function under similar environmen-
tal conditions is a key assumption implicit in the concept
of global biomes. While many convincing examples of
convergence in plants and animals exist, when systematically
comparing ostensibly convergent plant groups among realms
functional differences masked by superficial structural simi-
larity are often revealed (Alvarado-C
ardenas et al., 2013).
When using PFTs to model biome distributions, optimal-
ity assumptions are used that imply convergence (Fisher,
2013). For example, the perfect plasticity assumption (PPA)
– used when simulating tree canopy interactions and light
availability – assumes that tree crowns perfectly explore and
fill available canopy space (Purves et al., 2007). This greatly
simplifies the task of modelling light competition among
trees and can improve model performance (Purves et al.,
2008). Representing the wide array of plant architecture
observed naturally is another major challenge for modelling

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G. R. Moncrieff et al.

vegetation structure. However, rather than quantifying and
modelling the entire range of architectures observed across
biomes and environments, the assumption that plant
resource supply networks are designed to optimize the trans-
port of nutrients and water has been used to estimate plant
allometries that are ostensibly globally applicable (Enquist &
Niklas, 2002; Price et al., 2007).
Intercontinental comparisons are beginning to reveal bio-
geographical biases across scales of plant form and function
that suggest convergence upon optimal strategies is seldom
realized. In forests and savannas, plant architectural traits are
starkly different among continents (Banin et al., 2012; Dan-
tas & Pausas, 2013; Moncrieff et al., 2014a). These differ-
ences remain even after controlling for environmental
differences, and are explained by the divergent architecture
of a few regionally dominant taxa. The remarkably tall, thin-
stemmed and narrow canopied eucalypts of Australian savan-
nas diverge from the architectures of savanna trees of South
America which diverge from those from Africa (Moncrieff
et al., 2014a,b; Fig. 1). Similarly, the dipterocarp-dominated
forests of south-east Asia are composed of trees relatively tal-
ler and thinner than those found in other tropical forests
(Banin et al., 2012). In both examples cited above, the lack
of systematic divergence among all species in the regions
compared suggests that observed differences are not the
result of any unaccounted-for systematic abiotic or biotic
factor, but rather the product of the idiosyncrasies of a few
dominant groups. Scaling upwards from species-level trait
divergence to emergent ecosystem patterns, the structural
responses of savannas to variation in climate, soils and dis-
turbance also vary among continents, leading to differences
in the environmental limits of savannas (Lehmann et al.,
2011, 2014). Although not explicitly linked, it is likely that
this pattern is partially underpinned by the trait divergence
described by Moncrieff et al. (2014a) and Dantas & Pausas
(2013)

Figure 1 Comparative architecture of savanna trees from three continents. Canopy diameter, tree height and bark thickness are shown
at 20 cm stem diameter summarizing data from Moncrieff et al. (2014a), Dantas & Pausas (2013) and Lawes et al. (2011). Allometries
are drawn to scale except bark thickness, which is exaggerated by a factor of 2 relative to stem diameter in the diagram. The extensive
savannas now found in South America, Africa and Australia evolved in the late Miocene (c. 5–8 Ma, Beerling & Osborne, 2006) long
after Gondwana began to break-up; circa 184 Ma. The tree flora of each continent has thus been independently assembled. Savanna
trees evolved in situ on each continent from ancestors in closed-canopy forests and sclerophyll biomes (Simon et al., 2009; Bouchenak-
Khelladi et al., 2010; Crisp et al., 2011; Maurin et al., 2014). Thus, different clades characterize different continents, with Mimosoideae,
Melastomataceae and Malpighiaceae and the genus Qualea common in South American Cerrado, Myrtaceae (particularly Eucalyptus and
Corymbia) often dominant in Australian savannas and Mimosoideae, Caesalpinioideae and Combretaceae common throughout African
savannas.

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Divergence in structure and function among multiple
instances of a single biome has important consequences for
ecological forecasting. Although savannas share multiple
characteristics (including a discontinuous tree layer, continu-
ous understory of C4 grass, frequent fire), instances of the
savanna biome differ enough in structure and function that
each is expected to respond differently to global change
(Lehmann et al., 2014). This observation makes modelling all
savannas as a single cohesive entity indefensible. When it
comes to tropical forests there is an imperative to provide
reliable ecological forecasting, as they are critical carbon
reservoirs and are extremely biodiverse. But is the concept of
a single ‘tropical forest’ biome helpful when biodiversity,
architecture and biogeochemistry vary to the extent that
insights from one region are not relevant to another (Corlett
& Primack, 2006; Banin et al., 2014)?
The potential for multiple stable states
Despite the widespread notion that biome distributions are cli-
matically predetermined, it is well established that there exist
regions of climate space in which the dominant biome cannot
be accurately predicted (Whittaker, 1975; Bond, 2005). Vegeta-
tion is not simply a product of the environmental template but
simultaneously alters that template across scales, from local to
global. At large scales, the earth’s vegetation feeds back to cli-
mate by altering surface albedo, hydrology and the partitioning
of latent and sensible heat (Bonan, 2008). Furthermore, vege-
tation affects both directly and indirectly the fluxes and storage
of a range of greenhouses gases (IPCC, 2013). At smaller scales,
the presence of plants creates local micro-climates, altering
wind, temperature and moisture availability (Hoffmann et al.,
2012). Either actively or passively, plants change the physical,
chemical and biological properties of soils (Hawkes et al.,
2005). Beyond impacting climate and soils, vegetation itself
constructs ecosystem attributes relevant to plant functioning.
The amount of light that passes through canopies and is avail-
able to the lowest vegetation layers is determined by the den-
sity and arrangement of leaves. The ignition and spread of
wildfires is a function of the arrangement, production and
moisture content of the plant growth forms fuelling the fires
(Burgan & Rothermel, 1984). These feedbacks create the
potential for alternate biome states with profoundly different
biodiversity, ecological function and evolutionary history
given the same macroclimatic and geological templates that
are not simply transitional to a predictable climax vegetation
state (sensu Clements (1916)) but stable over long time series
(Charles-Dominique et al., 2015).
Fire is a good example of a non-climatic factor altering
vegetation at large scales. Many studies have shown that
when fire regimes are artificially altered vegetation change
often follows, sometimes leading to a biome switch (Bond
et al., 2005; Higgins et al., 2007). However, this does not
imply that vegetation is not predetermined by climate. If cli-
mate ultimately predetermines the prevailing fire regime
unless subject to external forcing, then vegetation will still be
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Biomes for understanding global change impacts
predictable based on the climate. If multiple biomes are truly
possible within a set of climatic constraints, after a transition
occurs, vegetation (and the fire regime) must not return to
the previous state once the external forcing is removed. Mul-
tiple stable biome states have been suggested over vast areas
of the tropics and subtropics as a result of feedbacks among
fire, flammable, shade-intolerant grasses and fire-sensitive
trees (Staver et al., 2011a). Through promoting recurring
natural fires, grasses can prevent the establishment of
fire-sensitive forest trees and maintain a grassland or savanna
ecosystem. Alternately, low light availability below forest tree
canopies reduces C4 grass growth and fuel bed continuity,
ultimately reducing the occurrence of fires and maintaining
an ecosystem with an entirely different suite of species and
drastically altered structure (Scheiter & Higgins, 2007; Hoff-
mann et al., 2012; Parr et al., 2012). These alternative states
can be internally stable – forests maintaining a fire-free state
without the need for external forcing and resistant to the
incursion of fires and flammable species; savannas persisting
in a frequently burned state unless perturbed by anthro-
pogenic action or a rare climatic event (Staver et al., 2012).
Herbivores can also play an important role along with fire in
facilitating stabilizing feedbacks and transitions between
alternative stable states (Dublin et al., 1990).
Perceptions of the factors controlling vegetation distribu-
tion are built upon the assumption that universal bioclimatic
constraints acting on convergent plant functional types pro-
duce predictable patterns of biome distribution (Whittaker,
1975). This thinking is reflected in the design of the current
generation of global vegetation models (Prentice et al.,
2007), and in maps of potential natural vegetation that give
the impression of deterministic biome distribution patterns
(Olson et al., 2001). These maps are used for benchmarking
model performance, and hence further select for models that
represent biomes as deterministic constructs. Some models
now incorporate the processes responsible for non-determi-
nistic biome distribution patterns (Higgins & Scheiter, 2012;
Moncrieff et al., 2014b). If a model allows for multiple
potential biome states it is the initialization and maintenance
of feedbacks that determines which of the suite of potential
biomes is realized (Fig. 2). The legacy of past climates is
indeed visible in current species and biome distributions,
some of which may not yet have reached equilibrium with
prevailing conditions (Svenning & Skov, 2007; Moncrieff
et al., 2014b). Unfortunately many DGVM analyses do not
routinely report on the extent to which predictions are influ-
enced by initial conditions, making it difficult to assess how
widespread this problem is. Even a perfect model could sim-
ulate incorrect vegetation patterns with a mis-specified ini-
tialization. Separating these two sources of uncertainty
(model versus initial condition) would require vegetation
maps used for validation to reflect not only observed vegeta-
tion patterns, or a single climax community defined by cli-
mate, but rather the full range of possible states given the
environmental template used to force models (e.g. Staver
et al. (2011a), Fig. 2).
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(a) Historical biomes (b) Potential biomes (c) Predicted biomes

alternate past climates actual current climate actual current climate

Low tree cover

Both states possible
High tree cover

Figure 2 Differences in historical vegetation and climate can result in divergent contemporary biome distribution patterns even under a
fixed climate if multiple stable biomes are possible. In (a) two different hypothetical historical states over ecological time-scales (0.1–
10 ka) show a wetter Africa with more extensive forests (top panel) and a drier Africa with more extensive deserts, shrublands,
grasslands and savannas (bottom panel). In (b) the relationship between climate and vegetation is shown in climate space (bottom
panel) and projected in geographical space (top panel) based on contemporary climate data from New et al. (2000). The plotted
relationship between climate and biome state shows deterministically high tree cover in wet environments with aseasonal rainfall,
deterministically low tree cover when mean annual precipitation is low and the potential for multiple stable biomes states with
intermediate, seasonal rainfall (Staver et al., 2011b). Contemporary biome distributions (c) will depend on historical conditions in areas
where multiple stable biome are possible and not be predictable without the inclusion of accurate information on initial conditions.

trade-offs between species traits these analyses suggest traits

NEW DIRECTIONS
Mapping and modelling functional traits
Once we accept that history may often overwhelm conver-
gence as an evolutionary force it becomes tempting to aban-
don global plant biomes for more regional vegetation
formations as the highest unit of vegetation organization
(e.g. ecozones sensu Olson et al. (2001), or biomes crossed
with biogeographical realms e.g. Neotropical savannas). It
may be beneficial to avoid extrapolation of findings among
other regions when studies are concerned with only a single
continent or region. However, we do not deem this necessary
or prudent as a general rule. Biomes are widely used in
global conservation planning, meta-analyses, and Earth sys-
tem modelling. This necessitates a construct for organizing
our knowledge and understanding of the world’s major
ecosystem types. Recent developments have linked plant
traits and ecosystem functions such as nitrogen cycling or
carbon sequestration, thereby implying new ways of defining
global ecosystems (Reich et al., 1997). By revealing universal
that have a globally consistent interpretation (Wright et al.,
2004). The increasing availability of global data sets on plant
traits means that definitions of biomes that are driven by
data on plant functional traits is an realizable prospect
(Kattge et al., 2011).
Rather than producing biome classifications and delimiting
boundaries based on artificially imposed groupings and dis-
junctions that create the illusion of sharp boundaries or
mask intercontinental differences, new maps based on multi-
variate analyses of trait distributions reflect where change in
vegetation structure is gradual or abrupt. Such trait maps
would not need to group distinct vegetation types in differ-
ent regions based only on coarse structural similarity. Maps
obtained from direct measurements exist for well studied
regions (Dahlin et al., 2013). Van Bodegom et al. (2014)
produced global maps for three plant functional traits (leaf
mass per area, stem-specific density and seed mass) by pro-
jecting trait-environment relationships geographically. An
advantage of mapping plant traits and defining biomes based
on trait clustering is the necessity to simultaneously map

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trait variability at any location and the uncertainty in vegeta-
tion classification.
Another approach is to directly examine and map the
functional response of vegetation as revealed by remotely
sensed metrics of vegetation function. For example, Buiten-
werf et al. (2015) examined NDVI time series, calculated
important phenological parameters for each pixel, and classi-
fied them into ‘phenomes’. This differs from the approach
previously used to map biomes or land cover types from
remotely sensed reflectance data – where regression or
machine learning approaches are used to train classifiers –
because biome boundaries reflect interpretable functional
turnover. Biomes defined and delimited based on such func-
tional similarities are more likely to represent units that will
respond coherently to global change than units delimited
using artificial thresholds chosen to match training data.
Moving away from a PFT-based approach to dynamic veg-
etation modelling in favour of trait-based models is already a
research theme. Much criticism has been levelled at the con-
cept of PFTs, and the potential biases they introduce when
attempting to summarize all plant variation into these crude
groupings (Scheiter et al., 2013). New models of global vege-
tation patterns base projections on the simulated distribution
and functioning of plant types or individuals with continu-
ous variation in the values of multiple traits (Pavlick et al.,
2013; Scheiter et al., 2013; Sakschewski et al., 2015). Plant
performance in such a trait-based model is the outcome of
traits influencing plant physiology and ecological interac-
tions, and the integration of numerous trade-offs which con-
strain the possible realm of trait combinations. Yet even if
the grand task of linking plant performance to traits and
defining trade-off surfaces is achieved there is no guarantee
that trait distributions will be predictable based on environ-
mental conditions alone. Weak correlations between environ-
mental factors and community-weighted traits and
differences among separate evolutionary lineages and floras
suggest that trait convergence under similar environmental
conditions is unlikely (Heberling & Fridley, 2012; Mitchell
et al., 2015). In addition, the presence and intensity of top-
down controls on vegetation – such as fire and herbivores –
will profoundly alter traits (Staver et al., 2012; Wigley et al.,
2015), but may not be predictable from climate and soils.
Constraining the potential trait space available to modelled
plants according to regional trait pools, and adjusting the
parameterisation of trade-offs for different regions may
increase agreement between simulated and observed trait dis-
tributions. These constraints could be viewed as a means of
optimizing traits and trade-offs to match observations. We
view them rather as the necessary acknowledgement of the
importance of the unique evolutionary context of different
regions for ecosystem functioning.
Insights from evolutionary and ecological history
Modern biomes have been assembled over millions of years
with their distributions constantly changing in response to
Journal of Biogeography 43, 863–873
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Biomes for understanding global change impacts
past changes in the earth system and evolutionary innova-
tion. We believe elucidating the historical assembly of mod-
ern biomes can help identify convergence and divergence
among floristically distinct regions and move explanations of
modern difference in traits and vegetation patterns beyond
the simple invocation of ‘biogeographic idiosyncrasies’. For
example, the divergence already described in the savanna tree
flora (Fig. 1) contrasts with the relative similarity of the
savanna grass flora. While the globally widespread savanna
biome and endemic savanna trees are relatively young
(c. 5–8 Ma), C4 savanna grasses have existed for over 30 Ma
(Beerling & Osborne, 2006). Does the difference in age of
savanna trees and grasses explain why there is divergence in
the properties of the tree layer but not the grass layer among
Africa, Australia and South America? Relative convergence/
divergence as a function of age of a biome is becoming testa-
ble across biomes as information on historical biome assem-
bly becomes available.
Another useful example is provided by fire-prone biomes
of Australia and the eucalypt trees (Eucalyptus, Corymbia,
Angophora), which often dominate these vegetation types.
Australian biome distribution patterns and the variation of
basal area in relation to fire and climate differ markedly
from other continents (Lehmann et al., 2011; Murphy &
Bowman, 2012; Moncrieff et al., 2015). While the non-euca-
lypt tree component of these biomes responds to fire accord-
ing to expectations derived from studies on the fire ecology
of tree species from other continents, eucalypts appear to be
remarkably insensitive to variation in fire regimes (Bond
et al., 2012; Murphy et al., 2015). A unique suite of traits in
eucalypts related to growth rates and the abundance and dis-
tribution of bud-forming structures in stems and branches
enables fire escape and post-fire epicormic resprouting in
stems otherwise vulnerable to the total loss of aboveground
biomass after intense fires (Burrows, 2013). Modern euca-
lypts evolved from ancestors in fire-prone sclerophyllous
biomes (Crisp et al., 2011). The ancestral condition of fire-
adapted traits in eucalypts (c. 60 Ma) may explain their
remarkable success in fire-prone areas of Australia now
(Burrows, 2013). In contrast, the savanna tree flora of the
Brazilian Cerrado evolved from wet forest ancestors and cor-
respondingly these species differ from eucalypts in their fire
adaptations (Simon et al., 2009; Fig. 1). These differences
may partially explain why the distribution of South Ameri-
can savannas is restricted in environmental space relative to
African and Australian savannas (Lehmann et al., 2011,
2014). However, other factors such as differences in
megafaunal associations and their extinction rates may also
be involved (Doughty et al., 2015). Regardless, it is evident
that the historical context from which the fire-adapted flora
of Australia and South America evolved underpins modern
biome-level divergence.
We are also beginning to appreciate the importance of his-
tory over shorter (ecological) time spans for predicting the dis-
tribution of global plant biomes. The importance of boundary
conditions for modelling climate and weather has long been
869
G. R. Moncrieff et al.
recognized. This behaviour is not viewed as an artefact of
model structure, but a realistic representation of the chaotic
dynamics of atmospheric circulation (Lorenz, 1963). In global
vegetation models there is seldom direct lateral transfer of
mass or energy among pixels. Thus, the only relevant bound-
ary conditions relate to historical vegetation states. Progress in
modelling global vegetation depends on the acknowledgement
that vegetation patterns are not deterministic in relation to cli-
mate, the inclusion of processes that produce the feedbacks
responsible for multiple stable equilibria and explicit explo-
ration of vegetation history – especially after the end of the last
glacial. DGVM initializations should consider realistic histori-
cal vegetation states rather than uniform initial states. When
data on vegetation history are unavailable multiple initializa-
tions should be explored (Moncrieff et al., 2014b).
CONCLUSIONS
What we thought
The biome concept as developed by Schimper (1903) is
based on the idea that similar climates will select for similar
plant forms independent of differences in history. Underlying
this is the assumption that given the constraints common to
all plants optimal solutions to environmental conditions have
evolved. Deviations from global vegetation-climate patterns
at local scales have always been acknowledged (Walter,
1973). Many landscapes are a complex mosaic of vegetation
types often very different from the macroclimate-predicted
biome. At regional and continental scales, however, it was
presumed that aggregate properties would converge to the
macroclimate-derived expectation as fundamental biophysical
constraints limit the domain of possibilities and evolutionary
change drives global convergence. Accordingly, biome distri-
bution patterns have been thought of as deterministic, and
thus predictable given knowledge of the prevailing climate
and soils. Hence most efforts to model current and future
vegetation patterns work on the implicit assumption that
only one biome should be possible given prescribed environ-
mental conditions.
What we have learnt
Biomes defined by coarse structural similarity do not occur
within a consistent region of environmental space even at
the largest scales nor will they respond uniformly to global
change. The legacy of evolutionary history constrains con-
temporary plant functioning, resulting in divergence among
floristically distinct regions. Furthermore, many regions can
support multiple stable biome states rather than having
deterministic climate-vegetation associations. In these
regions, present-day vegetation patterns will depend on his-
torical conditions and maintenance of feedbacks that create
the potential for alternate stable states. A biome concept that
does not account for role of evolutionary history and multi-
ple stable biomes states will hinder, rather than help, efforts
870
to predict how global change will reorganize vegetation pat-
terns at large scales.
Moving forward
At the boundaries between biomes there is often rapid turn-
over of species, nutrient cycling and ecosystem services (Pay-
ette et al., 2001; Hoffmann et al., 2012). Biomes themselves
are the theatre of evolution, shaping contemporary patterns
of diversity, leaving an imprint on species evolutionary tra-
jectories (Pennington et al., 2004; Crisp et al., 2009). They
are thus not fictional constructs, and can indeed represent
important ecological and evolutionary units (Crisp et al.,
2009). We should therefore not do away with the biome
concept, but rather recognize that it requires updating from
its original conception over a century ago. Functional simi-
larities, which can now be observed globally – either directly
across the entire globe using satellites or as plant traits syn-
thesized in databases that are increasingly formidable – form
a better basis for defining and delimiting vegetation units.
Indicating uncertainty and multiple potential biomes on dis-
tribution maps will help to move away from the misleading
notion of biomes as climatically predetermined. Including
insights into historical biome assembly and historic vegeta-
tion states will help explain why differences in the behaviour
of different instances of a biome exist for no apparent envi-
ronmental reason. The result of this update will be an
improved understanding of the drivers of spatial and tempo-
ral vegetation change and an improved ability to predict how
global vegetation patterns will respond to global environ-
mental change.
ACKNOWLEDGEMENTS
We benefitted from discussions with Jasper Slingsby, Tho-
mas Hickler and Robert Buitenwerf. Funding was provided
by the South African Environmental Observation Network
and the National Research Foundation of South Africa. We
are grateful for the helpful feedback from three anonymous
referees.
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BIOSKETCH
Glenn Moncrieff is a postdoctoral researcher at the Fynbos Node of the South African Environmental Observation Network
(SAEON). He is interested in how evolutionary history affects present-day vegetation functioning and response to disturbance at
very large scales, and applies a range of tools from dynamic global vegetation models to phylogenetic methods to investigate
these processes.
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